FAO Species Identification Guide for Fishery Purposes Western

click for previous page 

iv 

Table of Contents 

Page 

CEPHALOPODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .687 

Introduction and General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .688 

Remarks on Major Groups of Commercial Importance . . . . . . . . . . . . . . . . . . . . . . . . .688 

Principal Measurements and Methods Useful for Identification . . . . . . . . . . . . . . . . . . . . .690 

Glossary of Technical Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .692 

Key to Families of Cephalopods Encountered in Fishing Activities in the Area . . . . . . . . . . . .699 

Annotated List of Families Encountered in Fishing Activities in the Area . . . . . . . . . . . . . . .705 

List of Families Treated in this Contribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .708 

Nautilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .709 

Sepiolidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .712 

Sepiadariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .719 

Idiosepiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .721 

Spirulidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .722 

Sepiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .723 

Loliginidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .764 

Enoploteuthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .781 

Onychoteuthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .784 

Histioteuthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .787 

Ommastrephidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .788 

Thysanoteuthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .797 

Chiroteuthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .798 

Mastigoteuthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .799 

Octopodidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .800 

STOMATOPODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .827 

Technical Terms and Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .828 

General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .829 

Guide to Families of Interest to Fisheries Occurring in the Area . . . . . . . . . . . . . . . . . . . .829 

Key to Major Families of Stomatopods Occurring in the Area . . . . . . . . . . . . . . . . . . . . 830 

List of Families Occurring in the Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .831 

Odontodactylidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .832 

Lysiosquillidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .835 

Harpiosquillidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .838 

Squillidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .842 

SHRIMPS AND PRAWNS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .851 



Guide to the Major Groups of Shrimps and Prawns Occurring in the Area . . . . . . . . . . . . . .856 

List of Families Occurring in the Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .857 

Infraorder Penaeidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .858 

Superfamily Sergestoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .858 

Sergestidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .860 

Superfamily Penaeoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .866 

Aristeidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .868 

Solenoceridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .875 

Penaeidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .889 

Sicyoniidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .952 

Infraorder Stenopodidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .955 

Stenopodidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .955 

Infraorder Caridea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .957 

Atyidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .960 

Hippolytidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .961 

Hymenoceridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .963 

Palaemonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .964 

Pandalidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .969 

Rhynchocinetidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .971

Page 

LOBSTERS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .973 



Guide to Families Occurring in the Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .977 

List of Families and Species Occurring in the Area . . . . . . . . . . . . . . . . . . . . . . . . . . .980 

Nephropidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .982 

Enoplometopidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .995 

Synaxidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1001 

Palinuridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1005 

Scyllaridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1028 

CRABS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1045 

Technical Terms and Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1046 

General Remarks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1048 

Imported Crabs of Commercial Importance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1055 

Guide to Families of Interest to Fisheries Occurring in the Area . . . . . . . . . . . . . . . . . . . 1056 

Key to the Families of Brachyuran Crabs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1061 

Key to the Families of Crab-like Anomura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1077 

List of Families of Marine Brachyura and Crab-like Anomura Presently Recognized . . . . . . . . 1080 

Infraorder Brachyura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1083 

Homolidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1983 

Dromiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1085 

Raninidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1089 

Calappidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1091 

Xanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1098 

Eriphiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1103 

Carpiliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1110 

Pilumnidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1112 

Goneplacidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1114 

Portunidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1115 

Geryonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1132 

Majidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1136 

Grapsidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1138 

Gecarcinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1147 

Ocypodidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1152 

Infraorder Anomura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1154 

Coenobitidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1154 

HOLOTHURIANS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1157 


Glossary of Technical Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1163 

Key to the Shallow-water Orders of the Class Holothuroidea . . . . . . . . . . . . . . . . . . . . . 1164 

Order Aspidochirotida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1164 

Holothuriidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1165 

Stichopodidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1185 

HAGFISHES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1191 

Myxinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1192 

SHARKS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1193 

Technical Terms and Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1194 


Key to Families Occurring in the Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1198 

List of Families and Species Occurring in the Area . . . . . . . . . . . . . . . . . . . . . . . . . . 1203 

Hexanchidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1208 

Echinorhinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1211 

Squalidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1213 

Pristiophoridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1233 

Squatinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1235 

Heterodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1238 

Parascylliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1241 

v

vi 

Page 

Brachaeluridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1243 

Orectolobidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1245 

Hemiscylliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1249 

Ginglymostomatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1260 

Stegostomatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1262 

Rhinocodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1263 

Odontaspididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1264 

Pseudocarchariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1268 

Alopiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1269 

Lamnidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1274 

Scyliorhinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1279 

Proscylliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1293 

Pseudotriakidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1296 

Triakidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1297 

Hemigaleidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1305 

Carcharhinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1312 

Sphyrnidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1361 

INDEX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1367 

click for next page


CEPHALOPODS

688 Cephalopods 

Introduction and General INTRODUCTION Remarks AND GENERAL REMARKS 

by M.C. Dunning, M.D. Norman, and A.L. Reid 

Living cephalopods include nautiluses, bobtail and bottle squids, pygmy cuttlefishes, cuttlefishes, 

squids, and octopuses. While they may not be as diverse a group as other molluscs or as the bony 

fishes in terms of number of species (about 600 cephalopod species described worldwide), they are very 

abundant and some reach large sizes. Hence they are of considerable ecological and commercial fisheries 

importance globally and in the Western Central Pacific. 

Remarks on Major REMARKS Groups of Commercial ON MAJOR Importance GROUPS OF COMMERCIAL IMPORTANCE 

Nautiluses (Family Nautilidae) 

Nautiluses are the only living cephalopods with an external shell throughout their life cycle. This shell is 

divided into chambers by a large number of septae and provides buoyancy to the animal. The animal is 

housed in the newest chamber. A muscular hood on the dorsal side helps close the aperture when the 

animal is withdrawn into the shell. Nautiluses have primitive eyes filled with seawater and without lenses. 

They have arms that are whip-like tentacles arranged in a double crown surrounding the mouth. Although 

they have no suckers on these arms, mucus associated with them is adherent. Nautiluses are restricted to 

deeper continental shelf and slope waters of the Indo-West Pacific and are caught by artisanal fishers using 

baited traps set on the bottom. The flesh is used for food and the shell for the souvenir trade. Specimens 

are also caught for live export for use in home aquaria and for research purposes. 

Squids (Order Teuthida) 

The 2 suborders, Myopsida, “covered-eyed”, nearshore (neritic) squids, and Oegopsida, “open-eyed”, oceanic 

(pelagic) squids, occur in the oceans and seas of the world and the species reaching larger sizes form the basis 

of major fisheries. Some squids are demersal or epibenthic at some period of their life cycle, but most species 

are pelagic, living off the bottom in the water column where they are caught using a variety of fishing gear such 

as trawls, lift nets, and jigs. FAO’s Yearbook of Fishery Statistics records about 2 169 000 t of squids taken 

worldwide in 1995, with around 159 000 t taken from the Western Central Pacific. 

The suborder Myopsida is represented in the Western Central Pacific by the very speciose Family 

Loliginidae which includes 4 genera and is important in many small- and large-scale fisheries. Like all 

myopsids, the loliginids are demersal, predominantly near-shore or shelf species, frequently feeding near 

or on the bottom. These squid occur in schools and are often caught in large spawning aggregations. Some 

species tolerate reduced salinities and more turbid estuarine situations (e.g. Loliolus spp.) while others 

occur in clear waters around coral reefs (e.g. Sepioteuthis lessoniana). In some species, the spawning 

season is extended with peaks in early summer and autumn. Many small- to medium-sized eggs are 

encapsulated in gelatinous strings attached to shells, corals, and other substrates. Loliginid squids are 

opportunistic carnivores and grow rapidly. All species so far studied have life spans of less than one year; 

for the small tropical species, only a few months. 

major ocean habitats showing indicative distributions of abundant cephalopods 

(after Packard et al., 1972)

Remarks on Major Groups of Commercial Importance 689 

Our knowledge of the taxonomy of the Indo-West Pacific loliginids remains poor. This is especially true for 

members of the genus Photololigo which includes the majority of the large commercially important species. 

The present set of diagnostic morphological characters (fin shape and relative length, sucker dentition, 

hectocotylus structure) may be highly variable, differ between the sexes, change with growth and do not 

always ensure a reliable identification of species. In many cases, these characters have been inadequately 

defined in the type descriptions (many from the mid 1800s), type reference material is poorly preserved 

and had poor geographic locality information. Several poorly known and new, unnamed species are referred 

to in the recent literature and “seasonal forms” with different life history characteristics and of questionable 

taxonomic status and distribution have also been described. This poor state of taxonomic knowledge has 

been highlighted previously by various researchers but little progress has been made. Current and future 

fisheries assessments of the loliginid resource and subsequent decisions concerning the management of 

the stocks are dependent on accurate identification of species. Therefore, there is an urgent need for a 

substantial cooperative regionwide taxonomic study of the genus Photololigo using classical morphology 

supported by modern techniques including allozyme electrophoresis and DNA analysis. 

Oceanic squids of various families of the suborder Oegopsida occur in the tropical Western Central Pacific 

but do not form a significant component of current reported fisheries catches. However, in some areas, 

arrow squid of the family Ommastrephidae are taken in localized artisanal fisheries (i.e. Sthenoteuthis 

oualaniensis in Melanesia and the Philippines) and their fisheries resource potential has been considered 

as large by some authors. In contrast to the loliginids, the taxonomy of this group is relatively well known. 

Cuttlefishes (Family Sepiidae) 

Cuttlefishes occur on the continental shelf and upper continental slope of tropical and temperate areas in 

all oceans. All are demersal and are believed to be more active at night. Many of the larger species of 

cuttlefish are important to fisheries in the Western Central Pacific. Fishing activity ranges from local, or 

subsistence fisheries to major export industries. Cuttlefish are targetted using a variety of gear including 

jigs and lures, baited and unbaited traps (sometimes with mangrove branches as attractants to spawning 

females), and spears. They are also an important component of finfish and prawn trawl bycatch in the area. 

They are used primarily for human consumption, but also as bait and are marketed fresh, frozen or dried. 

In 1995, FAO’s Yearbook of Fishery Statistics reports 96 198 t of cuttlefish (and bobtail squids) from the 

Western Central Pacific (about 44% of the total world catch of cuttlefish for that year). This figure comprises 

42 700 t caught off Thailand, 37 000 t from Viet Nam and 2 836 t caught in the Philippines. The taxonomy 

and biology of the cuttlefish in the area is generally poorly known and in need of review. While some, 

particularly commercial species, can be easily recognized, others which may occur in catches are not well 

defined by simple external morphological characters. 

Octopuses (Order Octopoda) 

The order Octopoda contains 2 suborders: the finned “cirrate” octopuses (suborder Cirrata) and the finless 

familiar “incirrate” octopuses (suborder Incirrata). The “cirrate” octopuses are soft and semigelatinous. All 

occur in deep water, possess paired fins on the mantle, deep webs, and rows of sensory papillae (“cirri”) 

adjacent to the suckers. These octopuses are rarely captured and, due to the soft flesh, are of no economic 

value. The “incirrate” octopuses include the familiar bottom-living octopuses and a range of pelagic species. 

All lack the fins and sensory cirri of the cirrate octopuses. All incirrate octopuses brood their young, either 

in lairs, within their webs, within the mantle, or using an egg case (as in the argonauts). Incirrate octopuses 

are found in all marine waters of the world from intertidal reefs to the deepest ocean trenches. 

The benthic octopuses (family Octopodidae) of the continental shelf are the primary targets of commercial 

fisheries. FAO’s Yearbook of Fishery Statistics reports about 247 600 t of octopods taken worldwide in 1995 

(about 10% of the total world catch of cephalopods) for which reported catches from the Western Central 

Pacific accounted for about 8% (24 487 t). The majority are harvested for human consumption as the 

bycatch of demersal trawl fisheries. They are also caught in artisanal fisheries by trapping, spearing, and 

using baited hooks or lures with certain species collected primarily as bait for finfish fisheries. They are 

marketed fresh, frozen, or dried. The taxonomy of this family is very poor. There are a large number of 

undescribed or poorly-defined species occurring in the Western Central Pacific, a number of which form 

the basis of local and commercial fisheries. Inappropriate species names are frequently used including 

many European names, species which only occur in the Atlantic Ocean (e.g. Octopus macropus, O. 

vulgaris). Much of this confusion has originated from poorly preserved reference material and use of 

limited, ill-defined distinguishing characters. Recent research working with live animal attributes and biology 

has clarified some the taxonomic problems. However, the majority of species in the Western Central Pacific 

(more than 40 species) still lack formal descriptions or any detailed information on biology, distribution, or 

importance to fisheries.


Principal PRINCIPAL Measurements MEASUREMENTS and Methods AND METHODS USEFUL FOR IDENTIFICATION 

External characteristics and measurements used 

Orientation, arm numbering, and external morphological terms are illustrated in Figure 1. Orientation is 

relative to the resting animal, the arms and arm/tentacle crown being anterior. The body or mantle is 

considered posterior. The upper surface of the resting animal is considered dorsal and the underside 

ventral. Arms are numbered as left or right, commencing from the dorsal arms. 

The mantle length is the standard length measurement for all cephalopods (except Nautilus where shell 

diameter is used). In squids and cuttlefishes, the measurement is made along the dorsal surface from the 

posteriormost point to the anteriormost point of the mantle. In octopuses, mantle length is measured from 

the midpoint between the eyes to the posterior tip of the mantle along the dorsal surface. 

I II 

tentacle 

III 

anterior 

left arms right arms 

III 

IV 

II 

IV 

arm 

I tentacle 

total 

length 

length 

head 

mantle 

length 

fin 

length 

tail 

stalk 

club 

total 

length 

mantle 

length 

a) squid 

posterior 

b) octopus 

c) cuttlefish 

Fig. 1 measurements, arm numbering, and major external features of cephalopods in dorsal view 

Determining sex 

In most squids and some cuttlefishes, the sex of specimens can be determined externally by examining the 

arms to find modifications in maturing and mature males (hectocotylization). Modifications include change in 

normal sucker arrangement or loss of suckers, increased sucker size, or thickening or lengthening of sucker 

stalks, protective membranes, and their supports. For many squids and cuttlefishes, one or both of the ventral 

arms are modified although in some species, no external modification is apparent. In some groups such as the 

bobtail squids, these modifications may occur to dorsal rather than (or as well as) ventral arms. The modified 

arms are used to gather spermatophores from the mantle cavity of the males and transfer them to the mouth 

region or sometimes in squids, inside the dorsal or ventral mantle of the female. 

The sex of benthic octopuses can be determined externally by examining the third arms to find the modified 

arm developed in maturing and mature males. This is typically the third arm on the right-hand side (left in 

some genera). It consists of a modified tip with a channel or gutter (the spermatophore groove) running 

along the edge of the arm. In copulation, the spermatophores are shunted along this groove to the modified 

tip. This tip usually consists of a spoon or club-like structure (ligula) and a short tongue-like flap (calamus). 

This tip inserts spermatophores directly into the oviducts of the female octopus. 

In poor material, for immature specimens or for species which lack hectocotylized arms, dissection of the 

mantle cavity is necessary to determine sex (see below). 

head 

I IV 

II 

III 

mantle 

length

Principal Measurements and Methods 691 

Internal structures in the mantle cavity 

In identifying certain species or determining the sex of damaged or immature animals, it is necessary to 

dissect open the mantle cavity, exposing the funnel-mantle locking apparatus, gills, and reproductive 

structures. 

Figure 2a shows the mantle cavity of typical squids (oegopsids) of both sexes, opened with a mid-ventral 

longitudinal cut along the length of the mantle. The funnel and mantle elements of the locking apparatus 

are visible just inside the mantle immediately posterior to the funnel opening itself. Males are recognized 

by the spermatophoric complex [including Needham’s sac (also called the spermatophoric sac) which 

stores fully formed spermatophores in mature squid and the coiled spermatophoric organ] and penis-like 

structure on the left side (right in ventral view) of the midline. Females possess paired nidamental glands 

(white elongate structures) and paired white oviducal glands and oviducts in the anterior part of the mantle. 

In myopsid squids (such as Photololigo), only a single oviducal gland and oviduct is present on the left-hand 

side of the animal and paired accessory nidamental glands are present anterior to the nidamental glands. 

seminal 

duct 

testes 

funnel locking 

cartilage 

anus 

penis 

spermatophore 

sac 

funnel 

mantle locking 

cartilage 

nidamental 

gland 

oviducal 

gland 

oviduct 

male female 

terminal organ 

(“penis” ) 

male 

testes spermatophore 

storage 

a) squid 

funnel 

funnel organ 

(W-shaped in male, 

UU-shaped in female) 

anal flaps 

anus 

septum 

gills 

branchial 

hearts 

renal papillae 

b) octopus 

mature ovary as in 

“large-egg” species 

Fig. 2 major features of the mantle cavity of a typical squid and octopus 

gills 

ovary 

distal 

oviducts 

mature ovary as in 

“small-egg” species 

female


To examine the contents of the mantle cavity of a cuttlefish, a median 

longitudinal incision needs to be made through the mantle on the ventral 

side of the animal. Mature females can readily be distinguished from 

males by the presence of a pair of leaf-shaped creamy yellow nidamental 

glands (Fig. 4). Eggs may also be seen in the ovary, below and 

posterior to the nidamental glands. In immature females, the nidamental 

glands may be greatly reduced in size or visible only as two short slits. 

The shape of the male and female genital openings on the left side of 

the mantle cavity also differs slightly between the sexes. 

The open mantle cavity of an octopod is illustrated in Figure 2b for both 

sexes. There are 2 gills, each consisting of distinct leaves (lamellae)in 

an inner and outer series. Males are recognized by the penis-like 

structure (terminal organ) on the left side (right in ventral view) of the 

central septum, while females possess paired oviducts on either side 

of the septum. 

Removing internal shells (gladii and cuttlebones) 

Squid - In fresh and preserved specimens, the gladius (Fig. 9) may be 

removed by making a mid longitudinal incision along the length of the 

dorsal mantle and peeling away the skin laterally. Care must be taken 

at the posterior end to ensure that all of the skin and muscle tissue is 

removed from around the cone. 

Cuttlefish - The cuttlebone (Fig. 10) can easily be removed from a fresh 

animal by making a median longitudinal incision along the length of the 

dorsal mantle, and 2 shorter incisions at the anterior end of the mantle 

(Fig. 3). The skin can then be peeled open to reveal the cuttlebone. 

Fig. 3 cuttlefish in dorsal view 

(broken line indicates suggested 

incision for dissection) 

Glossary of Technical Terms GLOSSARY OF TECHNICAL TERMS 

Accessory nidamental glands - glands of unknown function consisting of tubules containing symbiotic 

bacteria. Found in cuttlefishes and loliginid squids. Occur in both sexes, anterior to the nidamental glands 

in females; rudimentary in males (Fig. 4). 

Afferent blood vessels - vessels leading towards the gills. 

Anal flaps - pair of fleshy papillae that arise on either side of the anus (Fig. 5). 

Anal pad - ovoid pads of unknown function, appear glandular, lie either side of the rectum, just behind the 

anal opening (Fig. 5). 

Anterior - toward the head end or toward the arm tips of cephalopods. 

Anus - opening of the alimentary canal, or gut, through which undigested remains of food are expelled. 

Arm formula - the relative order of arm lengths from longest to shortest, e.g. “I II III IV” is arms decreasing in 

length from the dorsal pair (arm pair I) to the ventral pair (pair IV), “IV= III=II I” is dorsal pair shorter than all 

other, equal length, arms (note: arms are numbered in Arabic numerals by some authors, i.e. 2 or II, 3orIII). 

Arm - one of the 8 fixed appendages surrounding the mouth of squids, octopuses, and cuttlefishes (see 

also tentacles). 

Armature - refers to the presence and arrangement of suckers and/or hooks on the arms and tentacular 

clubs of cephalopods. 

Benthic - bottom dwelling, living on or near the bottom of sea (= demersal). 

Branchial - pertaining to the gills. 

Branchial canal - canal between afferent and efferent blood vessels. 

Buccal membrane - thin web of tissue that encircles the mouth, reinforced by 6 to 8 buccal supports (Fig. 6). 

Buccal membrane connectives - muscular bands that connect the supports of the buccal membrane to 

the bases of the arms (Fig. 6). 

Calamus - tongue-like projection at base of ligula on hectocotylized arm of male octopuses (Fig. 7). 

Calcareous - chalky, calcified by deposition of calcium salts (calcium carbonate). 

Carpus - the proximal zone of (small) suckers (and knobs) on the tentacular club (Fig. 8).

Glossary of Technical Terms 693 

accessory 

nidamental 

gland 

nidamental gland 

buccal 

membrane 

stalk of 

tentacle 

buccal connective 

(dorsally attached) 

Fig. 4 mantle cavity of female cuttlefish 

(opened ventrally) 

arm I (dorsal) 

arm IV 

(ventral) 

arm II 

Fig. 6 diagram of oral surface of brachial crown 

and buccal membrane of a squid 

ligula calamus sucker 

arm III 

Fig. 7 tip of hectocotylized arm in a male octopus 

buccal connective 

(ventrally attached) 

anal 

flaps 

epirenal 

body 

renal 

papilla 

dactylus 

manus 

(hand) 

carpus 

(wrist) 

anus 

anal 

pad 

penis 

Fig. 5 anterior end of 

mantle cavity of a 

sepiolid 

club 

Fig. 8 distal end of tentacle 

of a squid 

stalk


Chitin(ous) - a horny polysaccharide substance (fingernail-like) that forms the sucker rings, hooks and 

beaks of cephalopods. 

Chromatophores - pigment-filled, generally flat muscular sacs in the skin under individual nervous control 

that collectively provide the background colour, colour patterns, and colour play of cephalopods (never 

ovoid or embedded in muscle tissue as light organs may be). 

Club-fixing apparatus - the mechanism of suckers and knobs on the carpal region of the tentacular club 

that permits the 2 clubs to be locked together during capture of prey (see also carpus). 

Cone, conus - the spoon-like or cup-like conical posterior terminus of the gladius or cuttlebone (Figs 9 and 10). 

Cone flag - lateral extensions of the gladius developed from the cone (Fig. 9). 

Corneal membrane - the very thin, transparent skin that covers the eyes of myopsid squids and cuttlefish 

(Fig. 11b). 

Cuttlebone - calcareous supporting plate in the dorsal part of the mantle of cuttlefishes. Organ used to 

maintain buoyancy. Consists of many thin plates, or septae, arranged in a thick bundle. The system of 

plates is called the phragmocone (= sepion). The septae are interconnected by supporting poles and pillars 

which are visible on the ventral side of the cuttlebone as striae) (Fig. 10). 

Dactylus - the distal, terminal section of the tentacular club in squids and cuttlefishes, often characterized 

by suckers of reduced size (Fig. 8). 

Distal - away from the body or point of origin; toward the peripheral parts (opposite of proximal). 

Dorsal shield - hard calcareous dorsal surface of the cuttlebone. 

Efferent blood vessels - vessels leading away from the gills. 

Epipelagic - living in the surface waters of the ocean. 

Epirenal bodies - glandular structures of unknown function which lie on either side of the renal papillae 

(Fig. 5). 

Fins - the pair of muscular flaps that arise along the lateral or dorsolateral surface of the mantle of sepioids, 

teuthoids, and cirrate octopods; used for locomotion, steering and stabilization. 

Foveola - transverse membranous fold of skin that forms a pocket in the funnel groove of some oegopsid 

squids (e.g. some ommastrephids) (Fig. 12). 

Funnel - the ventral, subconical tube through which water is expelled from the mantle cavity during locomotion 

and respiration (Fig. 13) (reproductive and waste products and the ink also pass through the funnel). 

Funnel locking apparatus - the combination of the funnel locking cartilage (or component) and the mantle 

locking cartilage (or component); also called the funnel-mantle locking apparatus. It is found laterally at 

the ventral mantle opening joining the posterior extension of the funnel to the mantle in squids and 

cuttlefishes (Fig. 13). The cartilages may be very simple in structure such as in loliginid squids or highly 

complex such as in the ommastrephids (illustrated in Fig. 13). The apparatus may also involve a partial or 

complete muscular fusion between the funnel and mantle elements such as in the ommastrephid Sthenoteuthis 

oualaniensis, the bobtail squid Sepiadarium kochii, and all cranchiid squids. 

Funnel locking cartilage - the cartilaginous groove, pit, pocket, or depression on each ventrolateral side 

of the posterior part of the funnel that joins with the mantle locking cartilage to lock the funnel and mantle 

together during locomotion, so water is expelled only through the funnel and not around the mantle opening 

(Figs 13 and 14; see also mantle locking cartilage). 

Genital opening - exit duct for products formed in the reproductive tract; through which pass eggs and 

spermatophores. 

Gills - paired structures each consisting of many lamellae through which gas exchange occurs. How to 

count gill lamellae: to identify some octopuses, it is necessary to count the number of gill lamellae on 

each side of each gill (= per demibranch, an inner and outer demibranch on each gill). Count per 

demibranch excludes the central terminal (anterior) lamella, e.g. gill count of 10 refers to 10 lamellae on 

each side of each gill. The animal illustrated in Fig. 15 has a gill count of 10. 

Gladius (pl. = gladii) - the feather or rod-shaped chitinous supporting structure (or shell) in the dorsal 

midline of squids and sepioids other than cuttlefish (= pen) (Fig. 9). 

Hectocotylus - the part of 1 (or more) arm(s) of male cephalopods modified for transferring spermatophores 

to the female; modifications may involve suckers, sucker stalks, protective membranes, trabeculae in 

squids and cuttlefishes or a distinct ligula/calamus on tip of the modified arm in octopods. 

Hooks - chitinous, claw-like structures derived from the suckers on the arms and/or clubs of some 

oegopsids (Fig. 16).


rachis 

vane 

anterior 

lateral plates 

wings 

lateral 

asymptotes 

cone 

posterior 

cone flags 

Fig. 9 examples of gladii 

anterior 

last loculus 

(anterior smooth 

zone of cuttlebone) 

striae 

outer cone 

inner cone 

ventral view 

spine 

(or rostrum) 

posterior 

lateral view 

Fig. 10 cuttlebone 

a) myopsid squid 

(e.g. Loliginidae) 

eye 

naked 

eye covered 

by corneal 

membrane 

b) oegopsid squid 

(e.g. Ommastrephidae) 

Fig. 11 head, eyes, and arms in ventral view 

mantle locking 

cartilage 

foveola 

Fig. 12 funnel groove of squids 

anterior 

posterior 

(illustration: K.Hollis/ABRS) 

Fig. 13 internal view of anterior mantle 

(Ornithoteuthis) 

side 

pockets 

funnel 


cartilage 

mantle


Ink sac - the structure that stores the ink of cephalopods; it lies ventrally along the intestine or is embedded 

in the digestive gland (hepatopancreas) and empties anteriorly via a duct into the rectum. 

Inner cone - forked limbs on the ventral side of the cuttlebone, between the inner side of the outer cone 

and the phragmocone; usually extends to the anterior end of the striated zone (Fig. 10). 

Keel - a flattened, muscular extension along the aboral surface of some arms and tentacular clubs to render 

them more hydrodynamic (= swimming membrane) (Fig. 16); also a narrow longitudinal ridge on the keel 

of some cuttlebones. 

Last loculus - anterior part of the cuttlebone that is smooth, not striated below (Fig. 10). 

Lateral ridge - muscular keel along the lateral mantle of some octopuses. 

Light organ - a simple or complex structure that produces bioluminescence by intrinsic (self-generated) 

or extrinsic (bacterial) means. May be present in the skin, on the eyes, embedded in muscle in the mantle, 

head, arms, tentacles or on the viscera (= photophore) (Fig. 17). 

Ligula - spoon- or club-like tip of the modified (hectocotylized) arm of octopuses (Fig. 7). 

Mantle - the fleshy (muscular) tubular or sac-like body of cephalopods; provides propulsion through jet-like 

expulsion of water; contains the viscera. 

Mantle locking cartilage - the cartilaginous ridge, knob or swelling on each side of the ventrolateral, 

internal surface of mantle that locks into the funnel component of the funnel-mantle locking apparatus 

(Figs 13 and 14, see also funnel locking cartilage). 

Manus - central or “hand” portion of club between the dactylus distally and the carpus proximally (Fig. 8). 

Medial (median) - pertaining to a structure located towards, on, or along the midline. 

Mesopelagic - living in the middle layers of the water column in oceanic waters. 

Neritic - inhabiting the sea over the continental shelf; arbitrarily taken to be the sea where it is shallower 

than 200 m. 

Nidamental glands - glands inside the mantle of females that secrete a fourth envelope for the eggs. 

Present in squids (except Enoploteuthidae), sepiolids, sepiadariids, and cuttlefishes. Glands tongue-like 

and bifurcated in squids, and oval in cuttlefishes (Fig. 4). 

Nuchal cartilages - a pair of cartilaginous structures (a mantle element and a neck element) connecting 

the mantle to the neck dorsally in many cephalopods. 

Ocellus - dark false-eye spot; found in “ocellate” octopuses as a pair of ocelli, one spot below each eye on 

the lateral arm crown between the bases of arms II and III (as in Octopus cyanea). 

Olfactory organ - organs of smell; in squids, cuttlefish, and vampyromorphs represented by olfactory 

papillae, while in octopus there are olfactory pits. Positioned on sides of the head near the neck. 

Orbit - cavity, or depression housing the eyeball. 

Outer cone - rim surrounding the phragmocone in cuttlebones. 

Pedicel - a short, tubular stalk that supports a sucker in sepioids and squids (Fig. 18). 

Phragmocone - system of plates comprising the cuttlebone. 

Pocket - an open depression in the anteroventral surface of the head between the bases of arms III and 

IV of cuttlefish into which the tentacles are retracted when not in use. 

Posterior - away from the arms and tentacles, towards the tail or rear end of mantle. 

Protective membrane - thin web-like integument along the lateral angles of the oral surface of the arms 

and clubs lateral to the suckers, supported by muscular rods called trabeculae (Fig. 19). 

Proximal - toward the body or nearest or next to the point of origin or attachment (opposite of distal). 

Rachis - the thickened central axis of the gladius that generally extends its entire length. Free rachis is the 

portion that does not support vanes (Fig. 9; see also vanes, gladius). 

Renal papilla - kidney opening. 

Renal sac -kidney. 

Secondary fold - on the eyelid, conspicuous in cuttlefishes. 

Secondary sexual character - a characteristic of animals which differs between the 2 sexes, but which is 

not a primary component of the reproductive system, e.g. enlarges suckers, hectocotylized arms. 

Spermatophores - encapsulated packets of sperm. Tubular structures manufactured by male cephalopods 

capable of holding millions of sperm, being transferred intact by a modified arm of the male and attaching 

to the female (or being inserted into the oviducts of octopuses) until fertilization occurs.


a b 

a b b 

simple (Loliginidae) complex (Sepiidae) other complex funnel cartilages 

light organs 

on eye 

light organs 

on ink sac 

Fig. 14 basic types of mantle locking cartilage (a) and funnel locking cartilage (b) 

funnel 

Fig. 15 gills 

anterior 

posterior 

gill 

lamella 

Fig. 17 examples of light organs 

light 

organs 

on 

ventral 

mantle 

keel 

hooks 

Fig. 16 tentacular club 

of an onychoteuthid 

squid 

pedicels 

trabeculae 

Fig. 18 detail of arm 

Fig. 19 detail of arm 

protective 

membrane 

sucker


Spermatophore groove - channel-like groove along the edge of the hectocotylized arm of mature male 

octopuses along which spermatophores are shunted to the tip for transfer to the female. 

Spine - the sharp extension on the posterior tip of the gladius or cuttlebone, dorsal to the outer cone 

(Fig. 10; also called the rostrum). 

Suckers - muscular, suction-cup structures on the arms and tentacles (rarely on the buccal membrane) of 

cephalopods; stalked and placed on muscular rods that contract (squids and cuttlefishes) (Fig. 20a); sessile 

and embedded without stalks on the oral surface of the arms of benthic octopuses (Fig. 20b). They are 

counted in either longitudinal rows (sometimes called “series”) or in oblique, transverse rows (Fig. 20c). 

Sucker ring - chitinous, often serrated or toothed ring that encircles the opening of suckers of squids and 

cuttlefishes. 

Sulcus - a median longitudinal groove, sometimes flanked by 2 low ridges on the ventral side of the 

cuttlebone. 

Swimming membrane - an elongate, muscular vane along the aboral surface of arms of cephalopods that 

functions to streamline and support the arms during swimming (= keel). 

Tail - the posterior extension of the mantle in some squids, frequently elongate. Fins or tapered terminations 

of fins may extend posteriorly along the tail. 

Tentacles - the 2 elongate, stalked appendages used for prey capture; distal ends contain clubs with 

suckers (or hooks); stalks usually devoid of suckers. Tentacles in squids can only contract rather than retract 

into pockets as in cuttlefish. 

Tentacular club - terminal portion of a tentacle; armed with suckers (or suckers and/or hooks), used for 

capturing prey. 

Terminal organ - penis-like muscular process of the male reproductive tract in octopuses which passes 

spermatophores into the base and spermatophore groove of the hectocotylized arm. 

Total length - length from posterior tip of mantle to tip of longest arm in octopods or tentacles in squids 

and cuttlefishes. (Because of the various degrees of contraction of the highly extensible tentacles during 

capture or fixation/preservation, this is not generally a useful comparative measurement in squids and 

cuttlefishes). 

Trabeculae - muscular rods that support the protective membranes on the arms and clubs of cephalopods; 

occasionally membranes are reduced and/or trabeculae are elongated, so they extend beyond the edge 

of the membrane, papilla-like (Fig. 19). 

Umbilicus - the central core of the chambered Nautilus shell, representing the juvenile shell with its initial coils. 

Vane - thin lateral expansion of the gladius that arises from the rachis (Fig. 9; see also rachis). 

Ventral - the lowermost or belly surface of a cephalopod; the surface on which the funnel is located; opposite 

the dorsal surface. 

Web - a membranous sheet that extends between the arms of many octopuses and some squids and 

cuttlefish, giving an umbrella-like appearance when the arms are spread out (Fig. 21). 

Web depth - distance from mouth to shallowest point of web between adjacent arms in octopods (Fig. 22). 

distal margin suckers in 

longitudinal 

web 

rows 

web depth 

(”series”) 

a) sucker of squid 

proximal 

margin 

b) sucker of octopus c) tentacular club 

Fig. 20 suckers and how they are counted 

8 suckers in 

(oblique) 

transverse 

rows 

Fig. 21 octopus of the genus 

Tremoctopus, with expanded web 

Fig. 22 octopus in oral view

Key to Families of Cephalopods 699 

Key to Families of Cephalopods KEY TO FAMILIES OF CEPHALOPODS ENCOUNTERED 

IN FISHING ACTIVITIES IN THE AREA 

(compiled from Nesis, 1987, and Roper et al., 1984) 

Note: families and higher taxa which are not treated further in this guide because they are unlikely to be 

regularly encountered in fishing or research activities are indicated by an asterisk (*). The reader is referred 

to the general references listed for further information on these groups. 

1a. Animal with hard, chambered external shell; numerous (more than 50) slender arms 

without suckers or hooks (Fig. 23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . Subclass Nautiloidea: Order Nautilida (monotypic order): Family Nautilidae (p. 709) 

1b. Shell absent or internal, external shell only present in female argonauts as thin brittle 

egg case; 8 arms or 8 arms and 2 tentacles always armed with suckers or hooks . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Subclass Coleoidea → 2 

2a. Eight arms and 2 tentacles (except in adult Octopoteuthidae which lack tentacles but 

are otherwise squid-like); suckers with chitinous rings, sometimes modified into hooks; 

fins always present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2b. Eight arms only; suckers without chitinous rings or hooks; fins absent or present only 

as short paddles on the sides of the mantle in certain deep-water gelatinous forms . . . . . . → 19 

3a. Internal shell (if present) either straight and laminate, coiled and chambered, or 

rudimentary and straight; pockets present which house the tentacles between arms III 

and IV. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Order Sepiida (= Sepioidea)) → 4 

3b. Internal shell straight, feather- or rod-shaped; no pockets present between arms III and 

IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Order Teuthida (= Teuthoidea)) → 8 

4a. Internal shell calcified, as flat laminate cuttlebone or coiled chambered shell . . . . . . . . . . → 5 

4b. Internal shell chitinous (thin and transparent) or absent . . . . . . . . . . . . . . . . . . . . . → 6 

5a. Internal shell coiled, chambered, embedded in posterior mantle (Fig.24) . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Spirulidae (p. 722) 

5b. Internal shell a thick, oval to elongate calcareous plate (the cuttlebone) embedded in 

the dorsal mantle (Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . . Family Sepiidae (p. 723) 

lateral view 

Fig. 23 Nautilidae (Nautilus) 

coiled 

internal 

shell 

dorsal view 

Fig. 24 Spirulidae (Spirula) 


dorsal view 

Fig. 25 Sepiidae (Sepia)


6a. Fins small and restricted to posterior end of mantle; large adhesive gland on dorsal 

surface of mantle (Fig. 26). . . . . . . . . . . . . . . . . . . . . . . Family Idiosepiidae (p. 721) 

6b. Fins semicircular to kidney-shaped on lateral mantle, never restricted to posterior 

mantle; adhesive gland absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

7a. Shell absent; both or only left ventral arm hectocotylized; dorsal border of mantle always 

fused to head (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . Family Sepiadariidae (p. 719) 

7b. Internal thin chitinous shell (gladius) present (except in Euprymna); hectocotylus 

developed on 1 dorsal arm, both dorsal arms (I) or dorsolateral arm (II); dorsal border 

of mantle free from, or fused to, head (Fig. 28) . . . . . . . . . . . . . Family Sepiolidae (p. 712) 

dorsal view 

large 

adhesive 

gland 

Fig. 26 Idiosepiidae (Idiosepius) 

mantle 

fused to 

head 

dorsally 

8a. Eye covered by transparent membrane (cornea); arms with suckers always in 2 rows; 

hooks absent; left or both ventral arms (IV) hectocotylized; paired light organs either 

side of the intestine in many species but no external light organs (Fig. 29) . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . (Suborder Myopsida): Family Loliginidae (p. 764) 

8b. Eye without cornea and in open contact with seawater; arm sucker arrangement variable 

(dependent on family); many species possessing hooks; hectocotylization present or 

absent; light organs may be present external on the mantle, head, arms and tentacles 

and ventral surface of the eyes as well as internally . . . . . . . . . . (Suborder Oegopsida) → 9 

9a. Mantle fused to head dorsally 

and to the funnel; funnel-mantle 

locking apparatus absent 

(Fig. 30) . . . . . . Family Cranchiidae * 

9b. Mantle not fused to head dorsally; 

funnel-mantle locking apparatus 

present (although 

funnel and mantle cartilages 

are fused in some species) . . . . . . → 10 

10a. Mantle-funnel locking apparatus 

a simple, straight groove 

and ridge . . . . . . . . . . . . . . . → 11 

10b. Mantle-funnel locking apparatus 

not a simple, straight 

groove and ridge . . . . . . . . . . . → 16 

dorsal view 

Fig. 27 Sepiadariidae (Sepiadarium) 

ventral view 

Fig. 29 Loliginidae 

(Loligo) 

dorsal view 

Fig. 28 Sepiolidae (Rossia) 

ventral view 

Fig. 30 Cranchiidae 

(Cranchia)


11a. Arms with hooks rather than 

suckers in adults; in juveniles 

where hooks are absent, suckers 

are in 4 rows . . . . . . . . . . . → 12 

11b. Arms without hooks, suckers in 

2 rows . . . . . . . . . . . . . . . . . → 13 

12a. Tentacles with fully developed 

clubs present; buccal membrane 

connectives attach to 

dorsal sides of ventral arms 

(IV) (Fig. 31) . . . . . . . . . 

. . . . . Family Enoploteuthidae (p.781) 

12b. Tentacles and clubs absent in 

adults although rudimentary 

clubs present in larvae or occasionally 

in juveniles; buccal 

membrane connectives attach 

to ventral sides of ventral arms 

(IV) (Fig. 32) . . Family Octopoteuthidae * 

13a. Buccal membrane connectives 

attach to ventral sides of ventral 

arms (IV) . . . . . . . . . . . . . → 14 

13b. Buccal membrane connectives 

attach to dorsal sides of ventral 

arms (IV) . . . . . . . . . . . . . . . → 15 

14a. Hooks present on tentacular clubs (Fig. 33) . . . . . . . . . . . Family Onychoteuthidae (p.784) 

14b. No hooks on tentacular clubs (Fig. 34) . . . . . . . . . . . . . . . . . Family Brachioteuthidae * 

hooks 

tentacular 

club 

dorsal view 

Fig. 33 Oncychoteuthidae (Onychoteuthis) 

ventral view 

Fig. 31 Enoploteuthidae 

(Pterygioteuthis) 

no 

hooks 

ventral view 

Fig. 32 Octopoteuthidae 

(Octopoteuthis) 

light 

organ 

tentacular 

club 

dorsal view 

Fig. 34 Brachioteuthidae (Brachioteuthis)


15a. Surface of mantle, head and arms covered with many light organs (usually large and 

distinct); a few small suckers at the proximal end of the manus; left eye considerably 

larger than right in adults (Fig. 35) . . . . . . . . . . . . . . . . . Family Histioteuthidae (p. 787) 

15b. Surface of mantle and head without light organs; many small to minute suckers (or suckers 

and knobs) at proximal end of manus and along tentacle shaft; eyes equal sized (Fig. 36) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Architeuthidae * 

light 

organs 

ventral view 

Fig. 35 Histioteuthidae (Histioteuthis) 

club 

Fig. 36 Architeuthidae (Architeuthis) 

16a. Funnel locking cartilage with a longitudinal and a transverse groove, ⊥-shaped or 

–| -shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 17 

16b. Funnel locking cartilage oval with 1 or 2 knobs directed toward centre of concavity . . . . . . → 18 

17a. Funnel locking cartilage with a longitudinal groove crossed by a transverse groove at its 

posterior end, ⊥-shaped; fins less than 60% of mantle length (Fig. 37) ........... 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Ommastrephidae (p. 788) 

17b. Funnel locking cartilage with a longitudinal groove from which a shorter groove 

branches medially, –|-shaped; fin length equal to mantle length (Fig. 38) ......... 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Thysanoteuthidae (p. 797) 


cartilage 

ventral view 

Fig. 37 Ommastrephidae (Ommastrephes) 

dorsal view 

cluster of 

suckers 

knobs alternating 

with suckers 

tentacular 


cartilage 

dorsal view 

Fig. 38 Thysanoteuthidae (Thysanoteuthis)


18a. Club with 4 longitudinal rows of suckers (Fig. 39) . . . . . . . . . Family Chiroteuthidae (p. 798) 

18b. Club with more than 15 longitudinal rows of minute suckers (Fig. 40) . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Mastigoteuthidae (p. 799) 

4 rows of suckers 

ventral view 

tentacular 

club 

Fig. 39 Chiroteuthidae (Chiroteuthis) 

19a. Suckers stalked, with chitinous rings; fins on posterior mantle, 1 pair in adults, 2 pairs 

in juveniles; internal shell as a thin broad chitinous plate; pair of small filamentous 

“tentacles” in pouch between bases of arms I and II; whole animal typically black; 2 

transverse pairs of light organs present (Fig. 41) . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . Order Vampyromorpha (monotypic order): Family Vampyroteuthidae 

19b. Suckers sessile (not stalked), without chitinous rings; fins present or absent; internal 

shell vestige either a U-shaped fin support, a pair of small rods (stylets), or absent; 

secondary filamentous “tentacles” absent; light organs absent (except around the mouth 

in some boliteanids); never completely black . . . . . . . . . . . . . . . (Order Octopoda) → 20 

20a. Fins present; body gelatinous; 

rows of sensory digits 

of skin (cirri) adjacent 

to single row of suckers 

. . . . . . (Suborder Cirrata) * 

20b. Fins absent; body gelatinous 

to muscular; sensory 

cirri absent . . . . . . . 

. . . (Suborder Incirrata) → 21 

filaments 

21a. Body jelly-like; often semitransparent 

. . . . . . . . . → 22 

21b. Body firm and muscular; 

opaque . . . . . . . . . . → 25 

22a. Suckers on arms in 2 rows 

(Fig. 42) . . Family Alloposidae * 

22b. Suckers on arms in 1 row . .→ 23 

light organs 

suckers 

minute, more 

than 4 rows 

dorsal view 

Fig. 40 Mastigoteuthidae (Mastigoteuthis) 

dorsal view 

Fig. 41 Vampyroteuthidae 

(Vampyroteuthis) 

ventral view 

Fig. 42 Alloposidae 

(Haliphron) 

2 rows of 

suckers


23a. Arms short, typically less than mantle length; webs shallow (less than 50% arm length); 

eyes moderate size and not telescopic (Fig. 43) . . . . . . . . . . . . . . . Family Bolitaenidae * 

23b. Arms longer than mantle length; webs deep (more than 50% of arm length); eyes 

telescopic or small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 24 

24a. Eyes elongate, tubular; mantle opening reduced to 2 separate small lateral slits; mantle 

and arm musculature enclosed in gelatinous outer coat (Fig. 44) . . . . . Family Amphitretidae * 

24b Eyes small and normal-shaped; single wide mantle opening; body gelatinous; mantle 

and arm musculature not enclosed in gelatinous outer coat (Fig. 45). . Family Vitreledonellidae * 

lateral view 

Fig. 43 Bolitaenidae 

(Japetella) 

1 row of 

suckers 

lateral view 

Fig. 44 Amphitretidae 

(Amphitretus) 

wide 

mantle 

opening 

lateral view 

Fig. 45 Vitreledonellidae 

(Vitreledonella) 

25a. Funnel-mantle locking apparatus absent; suckers in 1 or 2 rows (Fig. 46) . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Octopodidae (p. 800) 

25b. Funnel-mantle locking apparatus present; suckers in 2 rows . . . . . . . . . . . . . . . . . . → 26 

26a. Female housed in thin calcareous shell (“paper nautilus”);thin flared webs on tips of modified 

first (dorsal) arms in females (in live animal, webs of each modified dorsal arm can cover 

each entire face of the shell); third left arm modified in male (Fig. 47) . . . . . Family Argonautidae * 

26b. Shell absent; web (as in Argonauta) on tips of dorsal arm absent in females; third right 

arm modified in males . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 27 

dorsal view 

Fig. 46 Octopodidae (Octopus) 

 

lateral view 

 

Fig. 47 Argonautidae (Argonauta)

Annotated List of Families 705 

27a. Arms I and II of female joined by very deep thin web; arms I and II longer than other 

arms; ventral mantle smooth; 2 pairs of open holes on head (cephalic water pores), 

adjacent to bases of arms I and IV (Fig. 48) . . . . . . . . . . . . . . Family Tremoctopodidae * 

27b. Web absent; arms I and IV longer than other arms; ventral mantle of female sculptured 

with keratin-like inclusions in the skin forming a reticulate pattern; 1 pair of cephalic 

water pores on dorsal surface adjacent to bases of fourth arms (Fig. 49) . . . Family Ocythoidae * 

 

dorsal view 

expanded web 

water pores 

Fig. 48 Tremoctopodidae (Tremoctopus) 

reticulated 

pattern 

lateral view 

Fig. 49 Ocythoidae (Ocythoe) 

Annotated ANNOTATED List of Families LIST OF FAMILIES ENCOUNTERED IN FISHING ACTIVITIES IN THE AREA 

Classification modified after Nesis (1987), and Clarke and Trueman (1988). Importance to fisheries listed 

in increasing order as “non-commercial”, “minor commercial”, “commercial”, or “major commercial”. 

Class CEPHALOPODA Cuvier, 1798 

Subclass NAUTILOIDEA Agassiz, 1847 

Order NAUTILIDA, Monotypic Family Nautilidae Blainville, 1825 (minor commercial) - 

Chambered nautiluses. Characterized by: COILED PEARLY EXTERNAL CHAMBERED 

SHELL WITH ANIMAL LIVING IN THE OUTERMOST CHAMBER; 2 PAIRS OF GILLS; MORE 

THAN 50 SMOOTH “ARMS” WITHOUT SUCKERS; EYES SIMPLE WITHOUT LENSES. 

Medium-sized cephalopods with shell diameters reaching 250 mm; occur adjacent 

to coral reefs at the edge of the continental shelf and upper continental slope; 6 

species. 

Subclass COLEOIDEA Bather, 1888 

Order SEPIOLIDA - Dumpling or bobtail squid and pygmy cuttlefishes. Characterized by: 

FINS ROUNDED AND TYPICALLY WIDE; CHITINOUS GLADIUS RUDIMENTARY (Sepiolidae) 

OR GLADIUS ABSENT (Idiosepiidae); ONE OR BOTH DORSAL (Sepiolidae) OR BOTH 

VENTRAL ARMS (Idiosepiidae) HECTOCOTYLIZED IN MALES; LIGHT ORGANS PRE- 

SENT ON THE INK SAC IN SOME SPECIES. 

Family SEPIOLIDAE Leach, 1817 (minor commercial) - Dumpling or bobtail squid. 

Small rounded animals up to 80 mm mantle length; associated with sandy or rubbly 

substrates and seagrass beds in coastal waters and deeper continental shelf to 

60 m; 11 species. 

Family IDIOSEPIIDAE Appelöf, 1898 (non-commercial) - Pygmy cuttlefishes. 

Small animals maturing at less than 20 mm mantle length; occur in shallow seagrass 

and other inshore habitats; 3 species.


Order SEPIIDA - Cuttlefishes. Characterized by: CALCAREOUS SHELL INTERNAL IN 

THE MANTLE (except Sepiadariidae); TENTACLES RETRACTILE INTO POCKETS; ARM 

AND TENTACULAR SUCKERS WITH CHITINOUS RINGS; ONE PAIR OF GILLS WITHOUT 

BRANCHIAL CANAL BETWEEN AFFERENT AND EFFERENT BLOOD VESSELS; LIVER 

DIVIDED OR BILOBED; POSTERIOR FIN LOBES FREE. 

Family SPIRULIDAE Owen, 1836 (minor commercial) - Ram’s horn squid. Small 

(up to 45 mm mantle length); mesopelagic in warm oceanic waters; a single species. 

Family SEPIIDAE Keferstein, 1866 (major commercial) - Cuttlefishes. Mediumsized 

cephalopods to 500 mm mantle length; demersal species of the continental shelf 

and upper slope; more than 35 species. 

Family SEPIADARIIDAE Naef, 1912 (non-commercial) - Bottle squids. Small cephalopods 

(less than 50 mm mantle length); nektobenthic in coastal waters to depths of 

60 m; 2 species. 

Order TEUTHIDA - Squids. Characterized by: CHITINOUS GLADIUS INTERNAL IN THE 

MANTLE, SIMPLE, ROD- OR FEATHER-LIKE; TENTACLES CONTRACTILE BUT NOT 

RETRACTILE INTO POCKETS; ARM AND TENTACULAR SUCKERS WITH CHITINOUS 

RINGS AND/OR HOOKS; ONE PAIR OF GILLS WITH BRANCHIAL CANAL BETWEEN 

AFFERENT AND EFFERENT BLOOD VESSELS; LIVER UNDIVIDED AND SINGLE 

LOBED; POSTERIOR FIN LOBES MAY BE JOINED. 

Suborder MYOPSIDA Orbigny, 1845 - “covered-eyed” squids 

Family LOLIGINIDAE Steenstrup, 1861 (major commercial) - Inshore or pencil 

squids. Medium-sized cephalopods to 500 mm mantle length; nektonic species of the 

continental shelf; more than 20 nominal species. 

Suborder OEGOPSIDA Orbigny, 1845 - “open-eyed” squids 

Family ENOPLOTEUTHIDAE Pfeffer, 1900 (minor commercial) (includes M.R. 

Clarke’s families Enoploteuthidae, Ancistrocheirinae and Pyroteuthinae) - Firefly or 

enope squids. Small to medium squids to 400 mm mantle length; pelagic species of 

the continental slope and oceanic waters; 17 species. 

Family OCTOPOTEUTHIDAE Berry, 1912 (non-commercial) - Octopus squids. 

Medium to large squids to 1 700 mm mantle length; meso- to bathypelagic and benthic 

in oceanic waters; at least 4 species. 

Family ONYCHOTEUTHIDAE Gray, 1849 (minor commercial?) - Hooked squids. 

Medium to large squids to 400 mm mantle length; epi- to mesopelagic in oceanic 

waters and benthic in continental slope waters; at least 4 species. 

Family LEPIDOTEUTHIDAE Naef, 1912 (non-commercial) (includes M.R. Clarke’s 

family Pholidoteuthidae) - Scaled squids. Medium to large squids to 970 mm mantle 

length; nektobenthic in continental slope and oceanic waters; 3 species. 

Family ARCHITEUTHIDAE Pfeffer, 1900 (non-commercial) - Giant squids. Reach 

maturity at large sizes to 2 000 mm mantle length; mesopelagic in oceanic waters; 

number of species uncertain. 

Family HISTIOTEUTHIDAE Verrill, 1881 (non-commercial) - Jewel squids. Small to 

medium squids to 200 mm mantle length; mesopelagic to nektobenthic in continental 

slope and oceanic waters; 4 species. 

Family CTENOPTERYGIIDAE Grimpe, 1922 (non-commercial) - Ribbed finned 

squids. Small to medium squids to 90 mm mantle length; meso-to bathypelagic in 

oceanic waters; 2 species. 

Family BRACHIOTEUTHIDAE Pfeffer, 1908 (non-commercial) - Arm squids. Small 

to medium squids to 90 mm mantle length; meso- to bathypelagic in oceanic waters; 

2 species. 

Family OMMASTREPHIDAE Steenstrup, 1857 (commercial) - Arrow squids. Medium 

to large squids to 600 mm mantle length; nektobenthic in deeper continental 

shelf and slope waters and epi-mesopelagic in oceanic waters; 9 species. 

Family THYSANOTEUTHIDAE Keferstein, 1866 (commercial) - Diamond squids. 

Large-sized squids reaching 1 000 mm mantle length; epi-mesopelagic in warm 

oceanic waters; a single species.

Annotated List of Families 707 

Family CHIROTEUTHIDAE Gray, 1849 (non-commercial) - Chiroteuthid squids. 

Medium to large squids to 800 mm mantle length; mesopelagic to nekto-benthic in 

oceanic and continental slope waters; 2 species. 

Family MASTIGOTEUTHIDAE Verrill, 1881 (non-commercial) - Mastigoteuthid 

squids. Medium to large squids reaching 1000 mm mantle length; mesopelagic to 

nekto-benthic in oceanic and continental slope waters; several species. 

Family CRANCHIDAE Prosch, 1849 (non-commercial) - Cranch squids. Small to 

large squids to 800 mm mantle length; epi- to bathypelagic in oceanic waters; many 

species. 

Order VAMPYROMORPHA Pickford, 1939, Monotypic Family VAMPYROTEUTHIDAE 

Thiele, 1915 (non-commercial) - Vampire squids. Characterized by: EIGHT ARMS 

ONLY, TENTACLES ABSENT; SINGLE ROW OF SUCKERS STALKED WITH CHITINOUS 

RINGS; DORSAL MANTLE JOINED TO HEAD; PAIRED, ROUNDED FINS PRESENT; 

INTERNAL SHELL A CHITINOUS THIN BROAD PLATE; A PAIR OF THIN FILAMENTOUS 

FILAMENTS IN POUCHES BETWEEN ARMS I AND II; LIGHT ORGANS PRESENT 

IN THE MANTLE AT THE BASE OF EACH FIN AND MEDIAL TO THE EYE DORSALLY; 

COLOUR BLACK. 

Medium-sized octopuses reaching 130 mm mantle length; mesopelagic in slope and 

deeper waters; a single species. 

Order OCTOPODA Leach, 1818 - Octopuses. Characterized by: EIGHT ARMS ONLY, 

TENTACLES ABSENT; SINGLE OR PAIRED ROWS OF SUCKERS SESSILE WITHOUT 

SUCKER RINGS OR HOOKS; DORSAL MANTLE JOINED TO HEAD; FINS PRESENT 

OR ABSENT; CHITINOUS SHELL VESTIGE EITHER SMALL CARTILAGINOUS RODS 

OR A U-SHAPED SUPPORT; LIGHT ORGANS GENERALLY ABSENT (PRESENT 

AROUND THE MOUTH IN BOLITAENIDS). 

Suborder INCIRRATA Grimpe, 1916 

Superfamily BOLITAENOIDEA 

Family BOLITAENIDAE Chun, 1911 (non-commercial) - Small to medium-sized 

octopuses reaching 150 mm mantle length; meso- to bathypelagic (juveniles 

epipelagic); 2 species. 

Family AMPHITRETIDAE Hoyle, 1886 (non-commercial) - Medium-sized octopuses 

reaching 90 mm mantle length; meso- to bathypelagic (juveniles epipelagic); 

a single species. 

Superfamily OCTOPODOIDEA 

Family OCTOPODIDAE Orbigny, 1845 (major commercial) - Benthic octopuses. 

Small to medium-sized octopuses reaching 200 mm mantle length; demersal species 

of the continental shelf and upper slope; more than 30 species. 

Superfamily ARGONAUTOIDEA 

Family TREMOCTOPODIDAE Brock, 1882 (non-commercial) - Banket octopuses. 

Medium-sized octopuses reaching 200 mm mantle length (females), 15 mm mantle 

length (males); epi- to mesopelagic in oceanic waters; 2 species. 

Family OCYTHOIDAE Gray, 1849 (non-commercial) - Medium-sized octopuses 

reaching 200 mm mantle length; epi- to mesopelagic in oceanic waters; a single 

species. 

Family ARGONAUTIDAE Naef, 1912 (non-commercial) - Argonauts or paper 

nautiluses. Small to medium-sized octopuses reaching 120 mm mantle length 

(females), less than 20 mm mantle length (males); epi- to mesopelagic in deeper 

shelf and oceanic waters although occasionally encountered on the shelf; several 

species. 

Family ALLOPOSIDAE Verrill, 1882 (non-commercial) - Jelly octopuses. Mediumsized 

octopuses reaching 150 mm mantle length; epi- to mesopelagic oceanic 

octopods; a single species.


List of Familes LIST OF FAMILIES TREATED IN THIS CONTRIBUTION 

Order NAUTILIDA 

NAUTILIDAE - Chambered nautiluses 

Order SEPIOLIDA 

SEPIOLIDAE - Bobtail squids 

IDIOSEPIIDAE - Pygmy cuttlefishes 

Order SEPIIDA 

SPIRULIDAE - Ram’s horn squid 

SEPIIDAE - Cuttlefishes 

SEPIADARIIDAE - Bottle squids 

Order TEUTHIDA 

Suborder MYOPSIDA 

LOLIGINIDAE - Inshore squids, pencil squids 

Suborder OEGOPSIDA 

ENOPLOTEUTHIDAE - Firefly or enope squids 

ONYCHOTEUTHIDAE - Hooked squids 

HISTIOTEUTHIDAE - Jewel squids 

OMMASTREPHIDAE - Arrow squids 

THYSANOTEUTHIDAE - Diamond squids 

CHIROTEUTHIDAE - Chiroteuthid squids 

MASTIGOTEUTHIDAE - Mastigoteuthid squids 

Order OCTOPODA 

OCTOPODIDAE - Benthic octopuses 

References 

Chikuni, S. 1983. Cephalopod resources in the Indo-Pacific region. FAO Fish. Tech. Pap., 231:264-305. 

Chotiyaputta, C. 1993. Cephalopod resources of Thailand. In Recent advances in cephalopod fisheries biology. Tokyo, 

Tokai University Press, pp. 71-80. 

Clarke, M.R. 1966. A review of the systematics and ecology of oceanic squids. Adv. Mar. Biol., 4:91-300 

Clarke, M.R. and E.R. Trueman (eds). 1988. The Mollusca, Volume 12. Palaeontology and Neontology of cephalopods. 

London: Academic Press. 

Dunning, M.C. 1982. Squid and cuttlefish resources of Australian waters. FAO Fish. Rep., 275:103-111. 

Lu, C.C. and M.C. Dunning. 1998. Subclass Coleoidea Bather, 1888. Fauna of Australia. Volume 5. Mollusca. Canberra, 

Australian Government Publishing Service. 

Lu, C.C. and J.U. Phillips. 1985. An annotated checklist of the Cephalopoda from Australian waters. Occas. Pap. Mus. 

Vict., 2:21-36. 

Natsukari, Y. and M. Tashiro. 1991. Neritic squid resources and cuttlefish resources in Japan. Mar. Behaviour Physiol., 

8:149-226. 

Nesis. K.N. 1987. Cephalopods of the world. Neptune City, T.F.H. Publications, 351 p. 

Norman, M.D. and M.J. Sweeney. 1997. The shallow-water octopuses (Cephalopoda: Octopodinae) of the Philippine 

Islands. Invert. Taxonomy, 11:89-140. 

Okutani, T. 1995. Cuttlefishes and squids of the world in color. Tokyo, National Cooperative Association of Squid 

Processors, 185 p. 

Roper, C.F.E., C.C. Lu, and F.G. Hochberg (eds). 1983. Proceedings of the workshop on the biology and resource 

potential of cephalopods, Melbourne, Australia, 9-13 March 1981. Mem. Natl. Mus. Vict., (44):311 p. 

Roper, C.F.E., M.J. Sweeney, and C.E. Nauen. 1984. FAO species catalogue. Volume 3. Cephalopods of the world. An 

annotated and illustrated catalogue of species of interest to fisheries. FAO Fish. Synop., 125(3):277 p. 

Roper, C.F.E., R.E. Young, and G.L. Voss. 1969. An illustrated key to the families of the order Teuthoidea (Cephalopoda). 

Smithson. Contrib. Zool., (13):32 p. 

Sasaki, M. 1929. A monograph of the dibranchiate cephalopods of the Japanese and adjacent waters. J. Faculty Agric., 

Hokkaido Univ., Japan, 20 (Supplementary Number), 357 p. 

Sweeney, M.J., C.F.E Roper, K.M. Mangold, M.R. Clarke, and S.V. Boletzky (eds). 1992. “Larval” and juvenile cephalopods: 

A manual for their identification. Smithson. Contrib. Zool., (513):282 p. 

Voss, G.L. 1963. Cephalopods of the Philippine Islands. U.S. Natl. Mus. Bull., 234:1-180. 

Voss, G.L. and G.R. Williamson. 1971. Cephalopods of Hong Kong. Hong Kong, Hong Kong Government Press, 138 p. 

Voss, N.A. 1980. A generic revision of the Cranchiidae (Cephalopoda; Oegopsida). Bull. Mar. Sci., 30:365-412. 



Nautilidae 709 

Nautilidae NAUTILIDAE 

Chambered nautiluses 

by M.C. Dunning 

Diagnostic characters: Coiled, pearly, external shell punctuated with chambers with the animal 

living in the outermost chamber; 2 pairs of gills; up to 47 pairs of arm-like appendages around mouth; 

suckers and hooks lacking; eyes simple, without lenses; funnel (or infundibulum) consisting of 2 lobes 

which fold together to form a tube-like structure that serves for locomotion. Chromotophores and ink sac 

absent. 

septa separating chambers 

arm-like 

appendages 

R 

S| 

T| 

mouth 

obturating 

muscle of funnel 

radula 

(tooth-plate) 

hood 

funnel 

dorsal region 

of mantle 

umbilicus 

ventral region of 

mantle gills 

anus 

schematic cross-section of a Nautilus 

Habitat, biology, and fisheries: The nautiloids are represented by 6 living species of Nautilus. All of these 

are found in the tropical Indo-West Pacific. Nautiluses live in association with the bottom, primarily coral 

reefs, from depths of about 50 to 500 m. They are slow foragers of the deep nektobenthos and are generally 

found in deeper water during the day and at shallower depths at night. Nautiluses are the longest lived of 

the extant cephalopods, recent mark-recapture studies indicating they may live for more than 20 years. 

They exhibit determinate growth, i.e. after reaching maturity they show no more somatic growth, but may 

live several years after reaching maturity. Egg capsules in natural 

habitats are unknown but in captivity are laid singly attached to hard 

substrates and take up to 14 months to hatch in warm water (22° to 

25°C). It is hypothesized that they are laid in nature in relatively 

shallow water (80 to 100 m). At least 2 of these species are of 

commercial value as food (largely at the artisanal and subsistence 

levels) and in the specimen shell trade (e.g. Indonesia, Fiji, New 

Caledonia, and the Philippines). Nautiluses are also collected alive 

for public display and home aquaria, and for research. They are 

caught using baited fish traps. 

Similar families occurring in the area 

Argonautidae: a kind of octopus, the female argonaut produces a 

white calcareous “shell” in which she resides and eggs are laid and 

incubated. This “shell” has a single chamber only. Female argonauts 

have 8 true arms with biserial suckers compared to the many 

sucker-less arm-like appendages of nautiluses. 

digestive gland 

chambers 

shell 

Argonautidae 

ovary 

 

siphonal 

tube 

stomach


Key to the species of Nautilidae occurring in the area 

1a. Umbilicus small, up to 5% of shell diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Umbilicus larger, at least 10% of shell diameter . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2a. Umbilicus covered by a calcareous deposit, the umbilical callus . . . . . . . . . . . . . . . . . → 4 

2b. Umbilicus without umbilical callus (northeastern Australia). . . . . . . . . . Nautilus stenomphalus 

3a. Umbilicus moderate with sloping umbilical walls and an evenly rounded umbilical 

shoulder, approximately 16% of shell diameter (New Caledonia, Loyalty Islands) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nautilus macromphalus 

3b. Umbilicus large (approximately 20% shell diameter) with subangular shoulders and 

vertical walls (Solomon Islands, Papua New Guinea) . . . . . . . . . . . . . Nautilus scrobiculatus 

4a. Umbilicus approximately 5% of shell diameter, brown to reddish brown colour banding 

from the shell margin to the umbilicus or at least half way . . . . . . . . . . . . . . . . . . . . → 5 

4b. Umbilicus small, fine yellowish brown colour banding, sometimes greatly reduced 

(northwestern Australia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nautilus repertus 

5a. Fine raised longitudinal growth lines on the shell (found only around Palau, Western 

Caroline Islands) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nautilus belauensis 

5b. Shell generally smooth, without growth lines (widespread distribution, Indo-West Pacific) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nautilus pompilius 

a 

(after Swan and Saunders, 1986) 

schematic lateral view of Nautilus shells showing the umbilicus width (a) and the shell diameter (d) 

List of species occurring in the area 

The symbol is given when species accounts are included. 

Nautilus belauensis Saunders, 1981 

Nautilus macromphalus Sowerby, 1849 

Nautilus pompilius Linnaeus, 1758 

Nautilus repertus Iredale, 1944 

Nautilus scrobiculatus (Lightfoot, 1786) 

Nautilus stenomphalus Sowerby, 1849 

Reference 

Saunders, W.B., and N.H. Landman. 1986. Nautilus, the biology and paleobiology of a living fossil. New York, Plenum 

Press. 

d

Nautilidae 711 

Nautilus macromphalus Sowerby, 1849 

Frequent synonyms / misidentifications: None / None. 

FAO names: En - Bellybutton nautilus; 

Fr - Nautile bouton; Sp - Nautilo ombligo. 

Diagnostic characters: Umbilicus a deep, 

round shouldered concavity, approximately 

16% of shell diameter. 

Size: Maximum shell diameter about 

160 mm. 

Habitat, biology, and fisheries: Inhabits 

continental shelf and slope waters associated 

with coral reefs, from the surface to a 

depth of about 500 m. Consumed by artisanal 

fishers; also supports a small fishery 

for public and private aquarium and research 

trade. Collected alive at a depth of 

about 65 m on the outer slope of the barrier 

reef in New Caledonia; in the Coral Sea, 

trapped at depths between 300 and 400 m. 

Distribution: Restricted to New Caledonia 

and Loyalty Islands. 

Nautilus pompilius Linnaeus, 1758 


FAO names: En - Emperor nautilus; 

Fr - Nautile flammé; Sp - Nautilo común. 

Diagnostic characters: Umbilicus small, 

filled in with a concretion; brown to reddish 

brown striped colour pattern, extending to 

the umbilicus in some specimens or only 

half way across the shell in others. 

Size: Populations of this species reach 

shell diameters typically between 170 and 

180 mm around Fiji and the Philippines. 

Habitat, biology, and fisheries: Inhabits 

deeper continental shelf and slope waters 

around coral reefs, from near the surface to 

a depth of about 750 m. Supports shell 

trade, mostly from beach-drift specimens, 

and subsistence and artisanal fisheries in 

the Philippines. Captured in bamboo fish 

traps at depths from 60 to 240 m. 

Distribution: Indo-West Pacific; Andaman 

Islands, Ambon, the Philippines, 

New Guinea to Fiji; northeastern Australia. 

Absent from around New Caledonia where 

it is replaced by N. macromphalus. Sympatric 

with N. scrobiculatus off New Guinea, N. 

repertus off northwestern Australia, and N. 

stenomphalus off northeastern Australia. 

Replaced by N. belauensis around Palau.


Sepiolidae SEPIOLIDAE 

Bobtail squids 

by A.L. Reid and M.D. Norman 

Diagnostic characters: Small rounded 

squids (mantle length typically less than 

80 mm) with 8 arms and 2 functional retractile 

tentacles with well-developed clubs. Dorsal 

mantle free from, or fused to, head. 

Ventral mantle attached to funnel by funnel 

locking apparatus, mantle edge may 

cover funnel base. Fins present, rounded 

and typically wide. Maximum fin length 

distinctly longer than length of attachment 

to mantle. Suckers spherical, usually 

larger in males than females. Internal shell 

chitinous, rudimentary or absent. Frequently 

with light organ on ink sac. One or both 

dorsal arms hectocotylized in males. 

Habitat, biology, and fisheries: Benthic or 

mesopelagic squids. Mesopelagic species 

live in midwater over or near the continental 

slope. Benthic species associate with soft 

substrates and seagrass beds, typically remaining 

submerged in soft sediments during 

the day. Bury using fins and funnel to cover 

the entire animal with sand or shell, using the 

arms to rake grains onto the head and mantle. 

Animals typically emerge at night to forage 

for benthic and free-swimming 

crustaceans. A number of sepiolid species 

are of minor commercial importance 

throughout the area, harvested primarily as 

bycatch in trawl fisheries. 

Remarks: The taxonomy of many genera 

within this family is poorly known as most 

species are identified solely on sexual characters 

of mature males. 


Sepiadariidae (bottle squids): no internal shell; no light organs 

within the mantle cavity; dorsal mantle fused to head 

in all species; fins elongate, much longer than wide; ventral 

arms hectocotylized in males. 

Key to the genera of Sepiolidae occurring in the area 

1a. Only third and fourth arms united by a 

broad web; anterior edge of ventral mantle 

not covering funnel base; light organ 

present or absent; internal shell present 

or absent; benthic species . . . . . . . . . . . . → 2 

1b. All arms except the fourth pair united by 

a broad web; anterior edge of ventral 

mantle extends to cover base of funnel, 

reaching level of eye in certain species; 

light organ on ventral ink sac; internal 

shell absent; pelagic or deeper benthic 

species . . . . . . (subfamily Heteroteuthinae) → 3 

1 or both dorsal arms 

hectocotylized in males 

dorsal view 

both or only left ventral 

arms hectocotylized 

dorsal view 

Sepiadariidae 

fins wide and 

rounded 

mantle 

always 

fused to 

head 

dorsally 

fins distinctly 

longer than 

wide

Sepiolidae 713 

2a. Dorsal mantle fused to head; nuchal cartilage absent; left dorsal arm hectocotylized; 

internal shell rudimentary or absent . . . . . . . . . . . . . . . . . . . (subfamily Sepiolinae) → 4 

2b. Dorsal mantle not fused to head; nuchal cartilage present; left or both dorsal arms 

hectocotylized; shell present . . . . . . . . . . . . . . . . . . . . . . . .(subfamily Rossinae) → 6 

3a. Dorsal mantle fused to head by narrow strip . . . . . . . . . . . . . . . . . . . . . . . . Sepiolina 

(a single species, S. nipponensis, in this genus) 

3b. Dorsal mantle not fused to head . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroteuthis 

(a single species, H. weberi, in the area) 

4a. In male, only distal half of left dorsal arm modified: distal suckers modified into a row of 

column-like structures with tiny, fleshy, slit openings; base of arm with normal suckers 

and 1 to 3 elongate papillae with or without tiny sucker on tip; arm suckers in 4 or more 

longitudinal rows; tentacular clubs with many tiny suckers in more than 10 rows; light 

organ saddle-shaped, a lobe visible on each side of the septum; third arms not bent 

inwards towards mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euprymna 

4b. In male, entire hectocotylized arm or basal part modified with recesses and projections; 

arm suckers in 2 longitudinal rows (sometimes more on tips of ventral arms; tentacular 

clubs with 10 or less rows of small suckers; third arms of male usually greatly bent inward 

towards mouth; light organ present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

5a. Light organ present on ink sac, saddle-like or in form of 2 “ears”; junction of dorsal mantle 

and head wide, 33 to 50% of head width; suckers on ends of ventral arms sometimes 

arranged in 4 to 8 rows; tentacular club suckers usually in 4 to 8 rows . . . . . . . . . . . . Sepiola 

5b. Light organ absent; junction of dorsal mantle and head narrow, less than 33% of head 

width; hectocotylized arm widened in basal half in area of copulatory structure, distal 

part of arm normal; tentacular club suckers in 8 to 10 rows . . . . . . . . . . . . . . . . Inioteuthis 

6a. Anal flaps present; ink sac well developed; vane extends entire length of shell . . . . . . . . Rossia 

6b. Anal flaps reduced or absent; ink sac greatly reduced; vane present on posterior half of 

shell only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neorossia 1/ 



Euprymna morsei (Verrill, 1881) 

Euprymna tasmanica (Pfeffer, 1884) 

Heteroteuthis weberi Joubin, 1902 

Inioteuthis maculosa Berry, 1918 

? Neorossia sp. 1/ 

Rossia australis Berry, 1918 

Rossia bipapillata Sasaki, 1920 

Sepiola birostrata Sasaki, 1918 

Sepiola parva Sasaki, 1913 

Sepiola trirostrata Voss, 1962 

Sepiolina nipponensis (Berry, 1911) 

References 

Okutani, T. 1995. Cuttlefish and squids of the world in colour. Tokyo, Japan, Okumura Pringing Co. Ltd., 185 p. 

Reid, A. 1991. Taxonomic review of the Australian Rossinae (Cephalopoda: Sepiolidae), with a description of a new 

species, Neorossia leptodons and redescription of N. caroli (Joubin, 1902). Bull. Mar. Sci., 19(3):748-831. 

1/ Not yet recorded from the area. Representatives of this genus have been collected off the northwest shelf of western 

Australia in depths of 690 to 1 277 m, and 1 species, Neorossia leptodons, is known from southeastern Australia. 

Another representative of the genus is known from Taiwan Province of China, so the genus may extend throughout 

the area in deep water.


Eupryma morsei (Verrill, 1881) 

En - Double-ear bobtail squid. 

Maximum mantle length 40 mm. Dorsal mantle fused to head. 

Fins large and round. Arms with 4 longitudinal rows of suckers. 

Enlarged suckers of males restricted to ventral rows of 

arms II to IV, approximately 10 on each arm. Left arm I 

hectocotylized in distal half as rows of columnar suckers with 

tiny openings. Colour: large black chromatophores over iridescent 

gold to purple base colour. Taxonomy of genus poorly 

known, several undescribed or unresolved species in the area. 

Harvested on a minor scale, primarily as trawl bycatch. Southern 

Japan and Philippines to at least Indonesia. 

q¬ 

r¸do 

Eupryma tasmanica (Pfeffer, 1884) 

En - Southern bobtail squid. 

Maximum mantle length 40 mm. Dorsal mantle 

fused to head. Fins large and round. Arms with 4 

longitudinal rows of suckers. Enlarged suckers of 

males restricted to ventral rows of arms II and 

III, 1 to 3 on each arm. Left arm I hectocotylized 

in distal half as rows of columnar suckers with tiny 

openings. Colour: large black chromatophores over 

iridescent green to gold iridescent base colour. Taxonomy 

of genus poorly known, several undescribed 

or unresolved species in the area. No current exploitation. 

Eastern and southeastern Australia. 

dorsal view 

dorsal view


Heteroteuthis weberi Joubin, 1902 

En - Stumpy bobtail squid. 

Arms of mature male subequal in length, arms II not 

longer than arms I and IV; suckers reaching to distal 

tips of arms, 3 enlarged suckers on arms II, twice 

as large as normal suckers. Depth of web between 

arms I in males 33 to 50% arm length. Three enlarged 

suckers present on arms II, approximately twice as 

large as normal suckers. Anterior edge of fin extends 

well beyond anterior edge of mantle. Ink 

ejected with a luminous liquid. Females unknown. 

Bathyl. Known from central Indonesia. 

? 

? 

Rossia australis Berry, 1918 

En - Big bottom bobtail squid. 

Maximum mantle length 34 mm (males) and 63 mm (females). 

Anterior edge of mantle not fused to head dorsally. 

Fins large, ovate; attached within anterior 2/3 of mantle. 

Nuchal locking cartilage elongate oval, rounded anteriorly; 

tapering, slightly narrower posteriorly. Arm suckers 

biserial, largest suckers of arms II and III larger than those 

of arms I and IV in both sexes. Dorsal arm pair of males 

hectocotylized: ventrolateral edge of oral surface bordered 

by a swollen glandular crest, the inner edge of which forms 

a deep furrow extending from sucker rows 4 to 6, to sucker 

rows 8 to 11 (usually 4 to 9); proximal 8 to 10 series of 

suckers small, next 4 to 8 series enlarged; remaining suckers 

gradually diminish in size. Tentacular club slender, uniform 

in diameter; suckers minute; males with 18 to 26 

suckers in transverse rows, females with 25 to 

33 suckers in transverse rows. A pair of epirenal 

bodies present in males only, near the renal 

papillae. Anal pads present in both sexes, on 

either side of the rectum. Vane extends entire 

length of gladius. Colour: uniform pinkish. 

Sandy and muddy substrates to a depth of 

670 m. No fisheries importance at present. 

Found in eastern Australia. A second Rossia 

species is found on the northwest shelf of western 

Australia. 

tentacular 

club 

nuchal locking 

cartilage 

dorsal view 

dorsal view of female 

(illustration: K.Hollis/ABRS)


Rossia bipapillata Sasaki, 1920 

En - Big-eyed bobtail. 

Maximum mantle length 50 mm. Mantle short, saccular, 

width 70 to 80% of mantle length. Fins circular. Nuchal 

cartilage oval. Arm suckers globular; biserial.Tentacular 

club slender, with more than 25 minute suckers in 

transverse rows. Epirenal bodies present in both 

sexes, near the renal papillae. Anal pads present. 

Differs from R. pacifica Berry, 1911, in having an oval, 

rather than elongate, parallel-sided, nuchal locking cartilage, 

and in the tentacle club suckers, which in R. 

pacifica are arranged in 8 to 10 rows, rather than 25 or 

more as in R. bipapillata.R. pacifica does not possess 

papillae on either side of the anus. Minor fisheries 

importance at present. Found in Japan and the Philippines. 

? 

Sepiola birostrata Sasaki, 1918 

En - Butterfly bobtail; Fr - Sépiole papillon; Sp - Sepiola mariposa. 

Maximum mantle length approximately 12.5 mm. Arm III in both 

sexes stout and strongly curved inward (more obviously so in 

males); ventral suckers of right arm I and arms II to III of males 

larger than dorsal suckers, those on arms III more markedly 

enlarged. Left arm I hectocotylized: 2 pairs of normal suckers 

proximally, followed by 2 long, slender papillae arising ventrolaterally 

to blunt papilla; anterior of 2 papillae longer, and 

thicker; a smooth region distal to modified papillae; remaining 

distal suckers reduced, borne on swollen, closely packed, prismlike 

pedicels, together forming a fleshy mound; distal portion of 

arm twisted and strongly recurved.Tentacular club suckers large; 

4 suckers in transverse rows; dorsal suckers larger than ventral 

suckers. Light organ on each side of ink sac. Colour (preserved 

specimens): mantle and head with many minute brown or 

black chromatophores. Arms III deep pink; arms I to III with 

single rows of large chromatophores, arms IV with double row 

of small chromatophores. Differs from Sepiola trirostrata in 

lacking the blunt, fleshy papilla in addition to the 2 slender 

hectocotylus papillae. Fisheries significance unknown. Philippines, 

Singapore. 

? 

dorsal view 

(after Okutani, 1995) 

dorsal view 

(after Okutani, 1995)


Sepiola parva Sasaki, 1913 

En - Spotty bobtail squid. 

Maximum mantle length 10 mm. 

Mantle short, dome-shaped; anterior 

margin fused dorsally with large 

head. Fins large, ear-shaped. Arm 

suckers biserial.Male left arm I hectocotylized 

with a hook-like 

fleshy projection proximally following 

a few normal suckers;distal 

sucker pedicles swollen, forming 

a fleshy mound.Tentacular club with 

8 suckers in transverse rows, with 

median suckers larger than marginal 

ones. Subtidal zone in hard 

substrates. Southern Japan to 

northern Philippines. 

hectocotylus 

Sepiola trirostrata Voss, 1962 

En - Knobby bobtail squid. 

Maximum mantle length approximately 12.5 mm. Arm III in both 

sexes stout and strongly curved inward (more obviously so in 

males); ventral suckers of right arms I and arms II to III of males 

larger than dorsal suckers, those on arm III more markedly enlarged. 

Left arm I hectocotylized: 2 pairs of normal suckers proximally, 

followed by a large, blunt, fleshy papilla with 2 long, 

slender papillae arising ventrolaterally to blunt papilla; anterior 

of 2 papillae longer, and thicker; a smooth region distally to modified 

papillae; remaining distal suckers reduced, borne on swollen, 

closely packed, prism-like pedicels, together forming a fleshy 

mound; distal portion of arm twisted and strongly recurved. Tentacular 

club suckers large; 4 suckers in transverse rows; dorsal 

suckers larger than ventral suckers. Light organ on either side of 

ink sac. Colour (preserved specimens): mantle and head with 

many minute brown or black chromatophores. Arms III deep pink; 

arms I to III with single rows of large chromatophores, arms IV 

with double row of small chromatophores. Differs from Sepiola 

birostrata in having the blunt, fleshy papilla in addition to the 2 

slender hectocotylus papillae. Fisheries significance unknown. 

Philippines, Singapore. 

II left 

? 

I right Ileft 

male female 

dorsal views 


III left 

arm arrangement (male) 

dorsal view 

IV left 

tentacle 

(after Voss, 1963)


Sepiolina nipponensis (Berry, 1911) 

En - Japanese bobtail; Fr - Sépiole gros yeux; 

Sp - Sepiolina. 

Maximum mantle length 25 mm. Mantle short and 

dome-shaped; anteriorly fused to dorsal side of 

head by a narrow (approximately 3 mm) band. 

Fins oval. Arm suckers biserial; in females 

suckers numerous, small throughout; males with 

fewer suckers, enlarged on arms II to III and 

to a lesser extent, on arms IV. Both dorsal 

arms hectocotylized, thickened, with no 

special structure; suckers small, widely spaced, 

oral surface with transversely grooved ridges. 

Club same diameter as tentacle stalk; suckers 

minute, 13 to 16 suckers arranged in 

transverse rows; swimming keel extending for a 

short distance along stalk. Light organ roundish, 

on ink sac in mantle cavity; in fresh specimens, 

visible through mantle. Colour: with numerous 

chromatophores; ventral mantle margin dark and 

encircled by a silvery iridescent band, 

approximately 5 mm wide, chromatophores 

small, evenly peppered over this region. A neritic 

species occurring on the continental shelf to a 

depth of 200 m. Light organ excretes a luminous 

cloud, instead of ink. Fisheries interest 

undetermined. Found off southern Japan, the 

Philippines, and the Great Australian Bight. 

? 

arms of female (oral view) 

I 

IV 

dorsal view 


II 

III 

I 

IV 

arms of male (oral view) 

II 

III

Sepiadariidae 719 

Sepiadariidae SEPIADARIIDAE 

Bottle squids, bottletail squids 

by M.D. Norman and A.L. Reid 

Diagnostic characters: Small rounded squids (mantle 

length typically less than 40 mm) with 8 arms and 2 

functional retractile tentacles. Internal shell absent. 

Light organ absent. Dorsal mantle fused to head. 

Ventral mantle attached to funnel by fixed ligament or 

by plug-and-socket type funnel locking apparatus. III 

Fins present, relatively narrow and kidney-shaped (longer 

than wide). Ventral arms hectocotylized in males. 

Habitat, biology, and fisheries: Benthic squids, typically mantle 

associated with soft substrates and seagrass beds. Bottle always 

fused to 

squids remain submerged in soft sediments during the head 

day, bury using the fins and funnel to cover the entire dorsally 

animal with sand or shell, using the arms to rake grains 

onto the head and mantle. Animals emerge at night to 

forage for benthic and free-swimming crustaceans. There 

are no records for fisheries harvests of bottle squids in the 

area. They are likely to be taken infrequently and may be 

sold as bycatch in trawl fisheries along with bobtail squids 

(sepiolids). 


Sepiolidae (bobtail squids): fins wide and rounded; dorsal 

arms hectocotylized in males; an internal shell in most 

species; light organs within mantle cavity in many species; 

dorsal mantle not fused to head in many species. 

Key to the species of Sepiariidae occurring in the area 

1a. Ventral mantle permanently connected with 

funnel by muscular band in place of funnel 

locking apparatus; anterior edge of dorsal 

mantle smooth; fins situated in posterior half 

of mantle; colour pattern of longitudinal lines 

absent . . . . . . . . . . . . . . . . Sepiadarium kochii 

1b. Funnel locking apparatus present, consisting 

of 2 projections fitting into corresponding 

sockets; anterior edge of dorsal mantle 

fringed with finger-like projections at border 

of mantle aperture; fins narrow and elongate, 

extending along majority of mantle length; 

colour pattern of narrow longitudinal lines 

over white to pink base colour on dorsal 

surfaces . . . . . . . . . . . . . . Sepioloidea lineolata 



Sepiadarium kochii Steenstrup, 1881 

Sepioloidea lineolata (Quoy and Gaimard, 1832) 

mantle 

not fused 

to head 

dorsally 

in several 

species 

both or only left ventral arms 


dorsal view 

1 or both dorsal arms 


fins distinctly 

longer than wide 

References 

Berry, S.S. 1921. A review of the cephalopod genera Sepioloidea, Sepiadarium and Idiosepius. Rec. South Aust. Mus., 

1:347-364. 

Berry, S.S. 1932. Cephalopods of the genera Sepioloidea, Sepiardium and Idiosepius. Philipp. J. Sci., 47(1):39-53. 

Voss, G.L. 1964. Cephalopods of the Philippines. Bull. U.S. Natl. Mus., (234):180 p. 

II 

IV I 

dorsal view 

Sepiolidae 

fins wide 

and rounded


Sepiadarium kochii Steenstrup, 1881 

Frequent synonyms / misidentifications: Sepiadarium 

malayense Robson, 1932 / None. 

En - Koch’s bottle squid. 

Maximum mantle length 20 mm. Mantle fused to 

head dorsally and to base of both sides of 

funnel ventrally. Fins small and in posterior 

half of mantle. Internal shell absent. Tentacular 

clubs with 8 rows of minute suckers. Arms 

with 2 longitudinal rows of suckers near base, 

changing to 4 longitudinal rows in last 20 transverse 

rows. Left arm IV modified to form hectocotylus 

of 18 to 20 fleshy low lamellae over 

distal 60% of arm. Colour: spotted with large 

white to orange spots over dorsal surfaces. 

Found in coastal waters on soft sediments to 

depths of at least 60 m. Found throughout Indo- 

Malayan waters from India to Japan. 

? 

? 

Sepioloidea lineolata (Quoy and Gaimard, 1832) 


En - Striped (or Tiger) dumpling squid. 

Maximum mantle length 40 mm. Mantle fused to head 

dorsally. Ventral mantle connected to funnel by cartilaginous 

locking apparatus of dash-and-dot projections 

fitting into corresponding double sockets. Fins 

elongate and narrow, extending along majority of mantle 

length. Anterior margin of mantle aperture on dorsal 

mantle fringed with finger-like projections. Internal 

shell absent. Tentacular clubs with minute suckers in 20 

rows. Sides and ventral surfaces of head and mantle with 

rounded raised bumps. Colour: white with many narrow 

longitudinal stripes of pink to black on dorsal and 

lateral surfaces. In shallow waters on sand or mud 

substrates. Found in coastal waters of eastern, southern, 

and western Australia.

Idiosepiidae 721 

Idiosepiidae IDIOSEPIIDAE 

Pygmy cuttlefish 


Diagnostic characters: Very small cephalopods 

(less than 30 mm mantle length). Mantle elongated 

and slightly pointed at the posterior end; anterior 

edge of mantle not fused to head; head prominent with 

large, bulbous eyes covered by a cornea; nuchal cartilage 

absent; funnel locking cartilage oval-shaped in 

all species in the Western Central Pacific. Glandular, 

oval attachment organ located posteriorly on dorsal 

surface of mantle. Gladius a vestige only. Fins small, 

kidney-shaped, attached laterally to the posterior end 

of the mantle and slightly oblique to the longitudinal body 

axis. Arms short, with 2 rows of suckers; both ventral 

arms (IV) of mature males hectocotylized by loss of 

suckers on most parts of arms and by tips of left ventral 

arm becoming bilobed. Tentacles short and slender, with 

tentacular clubs not expanded, supporting 2 to 4 rows of 

suckers. 

Habitat, biology, and fisheries: Very abundant in 

shallow tropical coastal environments, particularly 

inshore seagrass beds and mangrove areas. Species 

are variously nektonic and planktonic and Idiosepius 

pygmaeus has a life span of less than 3 months. Animals 

appear to be solitary. Over a few days, each female lays 

up to 65 solitary eggs attached to hard substrates and 

then dies; eggs hatch after about 15 days. Of no value 

to fisheries because of their small size, but may be tentacular 

confused on cursory examination with hatchling and club 

juvenile loliginid squids which also occur in the same 

inshore habitats. 


Loliginidae: Idiosepiids are readily separated from 

juvenile loliginids on close examination by the presence 

of the dorsal attachment organ and by the lack of a true 

gladius. Idiosepius has fully developed reproductive 

organs and hectocotylization in males at less than 

15 mm mantle length. In Western Central Pacific 

species, the funnel locking cartilage is deep and 

oval-shaped compared to the simple straight cartilage in 

loliginids. 

left arm 

dorsal view 

(after Voss, 1963) 

ventral arms hectocotylized (oral view) 


Idiosepius pygmaeus Steenstrup, 1881 (southern Japan to northern Australia) 

Idiosepius paradoxus (Ortmann, 1888) (southern Japan to northern Australia) 

Idiosepius thailandicus Chotiyaputta, Okutani, and Chaitiamvong, 1991 (Gulf of Thailand) 

right arm 

attachment 

organ 

References 

Chotiyaputta, C., T. Okutani, and S. Chaitiamvong. 1991. A new pygmy cuttlefish from the Gulf of Thailand Idiosepius 

thailandicus n. sp. (Cephalopoda: Idiosepiidae). Venus (Jap. J. Malacology), 50:165-174. 

Jackson, G.D. 1988. The use of statolith microstructures to analyze life-history events in the small tropical cephalopod 

Idiosepius pygmaeus. Fish. Bull. NOAA, 87:265-272. 

Sasaki, M. 1923. On an adhering habit of a pygmy cuttlefish Idiosepius pygmaeus Steenstrup. Annot. Zool. Jap., 

10:209-213.


Spirulidae SPIRULIDAE 

A single species in the family. 

Ram’s horn squid 


Spirula spirula (Linnaeus, 1758) 

Diagnostic characters: A small squid. Characterized by the 

spirally-coiled internal shell located in the posterior end of 

the animal; shell contains over 30 chambers in adults. Mantle 

cylindrical, thin and muscular externally; a pair of small, round 

fins attached transversely to posterior end of mantle. A large 

photophore is located between the fins. Anterior margin of mantle 

with 3 pronounced projections on the dorsal midline and ventrolaterally 

on each side of the funnel-mantle locking cartilages. 

Funnel-mantle locking cartilage simple and straight. Eyes large, 

equipped with muscular eyelids. Length of arms increases from 

dorsal to ventral arms; each arm with 4 to 6 rows of small suckers. 

Non-expanded club on each long tentacle with 16 rows of numerous 

small suckers. All arms (except between ventral arms), connected 

with a web. Both ventral arms (IV) of males hectocotylized; 

left arm tip modified into a very complex organ of unknown 

function. 


None. Ram’s horn squid is easily 

distinguished by its spirally-coiled 

internal shell. 

Size: Maximum mantle length 45 mm, 

rarely larger. 

Habitat, biology, and fisheries: 

During the day the animals concentrate 

at depths between 600 and 

700 m. During darkness, the majority 

of the population occurs at depths 

less than 300 m. Able to control 

buoyancy by regulating the pressure 

internal shell 

dorsal view of male 


of gas contained in the shell. The eggs are small. The capture of young at depths of about 1 000 to 1 750 m 

suggests that females possibly lay eggs on the bottom on the continental slope. The smallest young known, 

presumably newly hatched, have a mantle length of about 1.5 mm with 2 shell chambers. Attains sexual 

maturity at about 30 mm mantle length. No fisheries are based on this species because of its relative 

scarcity and the small size as well as the tough and thin mantle. The beach collected shells are sold in the 

shell trade. 

Distribution: Found in tropical and subtropical oceanic waters worldwide, where water temperature at 

400 m is 10°C or more. 

References 

Bruun, A. F. 1943. The biology of Spirula spirula (L.). Dana Report, Carlsberg Foundation, (24):44 p. 

Bruun, A. F. 1955. New light on the biology of Spirula, a mesopelagic cephalopod. In Essays in the Natural Sciences 

in Honour of Captain Allan Hancock. Los Angeles, University of California Press, pp. 61-72. 

Clarke, M.R. 1970. Growth and development of Spirula spirula. J. Mar. Biol. Assoc. U.K., 50:53-64. 

Denton, E.J. and J.B. Gilpin-Brown. 1971. Further observations on the buoyancy of Spirula. J. Mar. Biol. Assoc. U.K., 

51:363-373. 

Lu, C.C., A. Guerra, F. Palumbo, and W.C. Summers. 1992. Order Sepioidea Naef, 1916. In “Larval” and juvenile 

cephalopods: A manual for their identification, edited by M.J. Sweeney, C.F.E. Roper, K.M. Mangold, M.R. Clarke, 

and S.V. Boletzky. Smithson. Contrib. Zool., 513:21-36. 



Sepiidae 723 

Sepiidae SEPIIDAE 

buccal 

membrane 

fin 

mantle 

biserial suckers 

tetraserial 

suckers 

ventral view 

tentacular 

club 

stalk 

pocket for 

retractile 

tentacle 

hectocotylus 

funnel 

spine 

Fig. 1 diagram of basic cuttlefish features 

5 suckers in 

(oblique) 

transverse 

row 

club suckers 

differ in size, 

medial suckers 

enlarged 

a) Sepia apama 

(after Lu, in press) 

dorsal and 

ventral 

protective 

membranes 

joined 

club 

suckers 

uniform 

in size 

ventral 

protective 

membrane 

Cuttlefishes 

normal 

suckers at 

distal end 

of arm 

modified region 

with reduced 

suckers 

normal suckers 

at proximal end 

of arm 

by A.L. Reid 

b) Sepia esculenta 

a) 

(after Sasaki, 1929) 

(after Okutani et al., 1987) 

Fig. 3 tentacular clubs 

swollen dorsal 

protective 

membrane 

transverse 

folds 

b) 

(after Nesis, 1982) 

Fig. 2 examples of male hectocotylized arms 

swimming 

keel 

dorsal 

protective 

membrane 

dorsal and 

ventral 

protective 

membranes 

not joined 

membrane 

separating 

sucker-bearing 

face of club 

from stalk 

c) Sepia latimanus 

(after Roper et al., 1984) 

proximal 

half of arm 

modified: 

suckers 

reduced, 

widely 

spaced, 

and arm 

swollen 

dorsal 

protective 

membrane 

swimming 

keel


anterior 

last loculus (anterior 

smooth zone of 

cuttlebone) 

anterior striae 

inverted 

U-shape 

outer cone 

inner cone 

median 

sulcus 

(furrow) 

spine with keel 

posterior 

spine 

(or rostrum) 

(ridge) 

ventral view a) 

lateral view 



cartilage 

mantle 

locking 

cartilage 

gill 

fin 

a) male 

I II 

IV 

III 

buccal 

membrane 

funnel 

renal 

sac 

ink 

sac 

anus 

renal 

papilla 

genital 

opening 

keel 

mantle 

locking 

cartilage 

gill 

accessory 

nidamental 

gland 

nidamental 

gland 

Fig. 5 mantle cavity (Sepia officinalis) 

ink sac 

b) 

I 

b) female 

II 

anterior 

striae 

inverted 

V-shape 

inner cone 

forming a 

thickened ledge 

posteriorly 

IV 

ovary 

(containing 

eggs) 

III 

funnel 

valve 

funnel 

organ 

anus 

genital 

opening

Sepiidae 725 

Diagnostic characters: Small to medium-sized cephalopods. Mantle robust, slightly flattened dorsoventrally, 

may be broad or slender; oval, oblong or nearly circular in outline; anterior dorsal mantle 

margin projected forward, not fused with head. Fins narrow, located dorsolaterally on mantle, approximately 

equal to mantle length; posterior fin lobes free, not connected to each other (Fig. 1). Head robust, slightly 

narrower than mantle; eyes prominent, covered by a transparent membrane and a conspicuous secondary 

fold on the eyelid. Mouth surrounded by 10 appendages (8 arms, 2 tentacles). Arms with 2 to 4 suckers in 

transverse rows (Fig. 1). Males of some species with hectocotylized ventral arm(s) IV for holding spermatophores; 

when present, usually consists of a modified region of reduced suckers (Fig. 2); hectocotylized 

region may also be swollen and crenulated by transverse folds (Fig. 2b). Tentacular clubs (Fig. 3) with 4 or 

more suckers in transverse rows; tentacles retractile into pockets on the ventrolateral sides of the 

head (Fig. 1). Arm and club suckers with chitinous rings. Mantle locking apparatus angular (Fig. 10a) or 

curved (Fig. 10b) in shape. Internal calcareous cuttlebone (Fig. 4) located dorsally underneath the skin, 

cuttlebone length usually equal to mantle length; cuttlebone shape ranges from lanceolate, oval, to 

diamond-shaped; dorsal side a calcareous plate (dorsal shield); ventrally, finely laminate, porous and 

comprised of thin, transverse septa supported by transverse calcareous rods. One pair of gills (Fig. 5); no 

branchial canal between afferent and efferent branchial blood vessels. Liver divided or bilobed. Buccal 

membrane present (Figs 1 and 5), with or without suckers; each radula tooth unicuspid (with a single 

projection). Olfactory organ a ciliated pit. 

Habitat, biology, and fisheries: On the continental shelf and upper slope to a maximum depth of 

approximately 600 m. Primarily bottom dwellers over a range of habitats, including rocky, sandy and muddy 

bottoms to seagrass, seaweed and coral reefs. Slower swimmers than the more streamlined squids. Able 

to attain neutral buoyancy by regulating the relative amounts of gas and fluid in the chambers of the 

cuttlebone; able to hover in midwater, with fins acting as stabilisers. Some species migrate seasonally in 

response to temperature changes and aggregate, usually in shallow water, at the time of spawning. Within 

a species, individuals may attain sexual maturity at very different sizes, depending upon the combined 

effects of temperature and light. Eggs, relatively few in number, are individually attached to various 

substrates in clusters; length of development varies with temperature. Life span (studied for Sepia 

officinalis) between 18 months and 2 years, though males may live longer; post-spawning mortality is high 

in females. Prey on a wide range of invertebrates and fish. 

Many species of cuttlefish are important to fisheries in the area. 

Fishing activity ranges from local, or subsistence fisheries, to 

major export industries. Sepiids are also an important 

component of finfish and prawn trawl bycatch in the area. They 

are used primarily for human consumption, but also as bait and 

are marketed fresh, frozen or dried. In 1995, FAO’s Yearbook 

of Fishery Statistics reports 96 198 t of cuttlefish (and bobtail 

squids) from the Western Central Pacific (about 44% of the 

total world catch of cuttlefish for that year). This figure 

comprises 42 700 t caught off Thailand, 37 000 t from Viet 

Nam, 2 836 t from the Philippines, and 3 t from Australia. 


Some species are superficially similar in appearance to the 

loliginid squid genus Sepioteuthis, but can be readily 

distinguished due to the presence of a calcareous cuttlebone 

in the sepiids. 

Loliginidae 

(Sepioteuthis) 

Sepiidae 

Identification note 

The taxonomy and biology of the cuttlefish from the area is generally poorly known and in need of review. 

While some, particularly commercial species, can be easily recognized, others which may occur in catches 

are not well defined. This presents difficulties in attempting to construct a key to all species occurring in 

the area, as particular character states for many species have been incompletely documented. For this 

reason, the key below should only be used in combination with the species accounts to follow to confirm 

identification. 

Many distinguishing features are seen only in males, so animals of this sex are generally required for 

identification. Where possible, however, features of the cuttlebone and tentacular clubs have been included 

in the key to enable females to be identified. Males can often be recognized by the presence of a 

hectocotylus. One (usually the left), or both ventral arms (IV) may be modified. Not all species have a 

hectocotylus. In addition, in juvenile males this secondary sexual character may not be fully developed, so


it may be necessary to check internal anatomical features to determine sex. To examine the contents of 

the mantle cavity, a median longitudinal incision needs to be made through the mantle on the ventral side 

of the animal. The internal features of males and females are shown in Figure 5. Mature females can readily 

be distinguished from males by the presence of a pair of leaf-shaped creamy yellow nidamental glands 

(Fig. 5b). Eggs may also be seen in the ovary, below and posterior to the nidamental glands. In immature 

females, the nidamental glands may be greatly reduced in size or visible only as two short slits. The shape 

of the male and female genital openings on the left side of the mantle cavity also differs slightly between 

the sexes (Fig. 5a, b). Any body patterning, such as transverse wavy lines or bands on the dorsal side of 

the mantle, is usually more pronounced in males and may be faint or absent in females. 

The number and size of the tentacular club suckers are important traits. If the tentacles are retracted, 

they can readily be extracted in fresh animals by gently probing inside the pouches between arms III and 

IV on either side of the buccal mass (Fig. 1). Extraction of the club may be more difficult in preserved 

specimens and dissection may be necessary. The number of club suckers given for each species, and used 

in the keys, refers to the number of suckers in transverse rows. This is determined by counting the number 

of suckers that are intersected in an oblique line midway along the tentacular club as shown in Fig. 3a. 

Other important features of the tentacular club include: 

1. The relative sizes of the suckers: they may vary in size (Fig. 3a, c), or be of similar size (Fig. 3b); 

2. The protective membranes may be joined at the posterior end of the sucker-bearing face of the club 

(Fig. 3a, c), or not joined (Fig. 3b); 

3. The swimming keel may extend beyond the sucker-bearing face of the club (Fig. 3c), or may be 

equivalent in length to the sucker-bearing face; 

4. The sucker-bearing face may be joined to the tentacle stalk (Fig. 3b), or may be separated from it 

by a membrane (Fig. 3c). 

Suckers are described as normal or normal-sized if they do not differ obviously in size from other suckers. 

As for club suckers, arm sucker “rows” refer to suckers positioned in transverse rows, that is, positioned 

in oblique lines approximately perpendicular to the longitudinal axis of the arm. “Series” refers to suckers 

positioned approximately parallel to the longitudinal axis of the arm. Biserial suckers are those arranged 

in 2 series, tetraserial are those arranged in 4 series. 

The cuttlebone (Fig. 4) can easily be removed from a fresh animal by 

making a median longitudinal incision along the length of the mantle, 

and two shorter incisions at the anterior end of the mantle as shown in 

Fig. 6. The skin can then be peeled open to reveal the cuttlebone below. 

Aside from its shape, important features of the cuttlebone include: 

1. The width of the outer cone: it may broaden posteriorly 

(Fig. 4a, b), or be of approximately uniform width along the length 

of the cuttlebone (Fig. 7); 

2. The inner cone may be narrow throughout (Fig. 7), or broaden 

posteriorly (Fig. 4a, b); 

3. The inner cone may be thickened posteriorly, and/or form a raised 

ledge (Fig. 4b); 

4. The median sulcus (or furrow) may be wide, or narrow; 

5. The shape of the striae: while the striae may vary in shape within 

species along the length of the cuttlebone, the shape of the 

anteriormost striae (where the striated zone joins the last loculus) 

is rather uniform and used for diagnoses. However, in some 

cases, the distinction between “anterior striae inverted U-shape”, 

and “anterior striae inverted V-shape” is not clear-cut, so identification 

should be confirmed following examination of other characters. 

The anterior extent of the striated zone may vary with the 

size of the animal, so may differ in some cases from that shown 

in illustrations. 

No other internal features, requiring dissection of animals, are included 

in the following key. Reference should be made to Figures 1 to 4 and the 

“Glossary of Technical Terms” (pages 692 to 698). 

suggested 

incision for 

dissection 

Fig. 6 cuttlefish in dorsal view

Sepiidae 727 

Key to the species of Sepiidae occurring in the area 

1a. Cuttlebone diamond-shaped in outline (Fig. 7); cuttlebone much shorter than mantle, 

located in the anterior 1/2 to 2/3 of mantle; dorsal anterior edge of mantle without 

tongue-like projection (Fig. 8a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metasepia 

(a single species, M. pfefferi, occurring in the area; unconfirmed records of M. tullbergi from the Philippines and 

Viet Nam) 

1b. Cuttlebone outline elliptical to lanceolate; cuttlebone length approximately equal to 

mantle length; dorsal anterior edge of mantle usually with tongue-like projection 

(Fig. 8b). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

cuttlebone (ventral view) 

Fig. 7 Metasepia pfefferi 

(illustration: K. Hollis/ABRS) 

2a. A gland and gland pore located 

on the ventral side of 

the posterior end of the mantle 

(Fig. 9); mantle locking 

apparatus (position shown in 

Fig. 5) with triangular projection 

(Fig. 10a); cuttlebone in- gland pore 

ner cone with very short 

limbs; outer cone a wide, 

flared, chitinized border 

around posterior end of cuttlebone 

(Fig. 11a) . . . . (Sepiella) → 3 

2b. Gland and gland pore absent; 

mantle locking apparatus 

(position shown in 

Fig. 5) semicircular, without 

triangular projection (Fig. 

10b); cuttlebone inner cone 

with relatively long limbs; 

outer cone usually calcareous, 

not obviously flared posteriorly 

(Fig. 11b) .......(Sepia) → 6 

3a. Tentacular club with 7 to 10 

suckers in transverse rows . . . . . → 4 

3b. Tentacular club with 12 to 

24 suckers in transverse 

rows . . . . . . . . . . . . . . . . . → 5 

tongue-like 

projection 

a) Metasepia pfefferi b) Sepia, Sepiella 

Fig. 8 head and anterior mantle (dorsal view) 


triangular 

projection 

posterior end of mantle 

(ventral view) 

Fig. 9 Sepiella 


outer cone 

chitinous, 

flared 

posteriorly 

a) Sepiella b) Sepia 

Fig. 10 mantle locking 

apparatus 

a) Sepiella b) Sepia 

Fig. 11 cuttlebone (ventral view) 


outer cone 

calcareous, 

not flared 

posteriorly


4a. Five to 6 wine-coloured spots at base of each fin; spots increasing in size posteriorly 

(Fig. 12a); cuttlebone width about 30% the length; cuttlebone broadens posteriorly, 

tapers anteriorly (Fig. 13a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepiella weberi 

4b. Six to 7 wine-coloured spots along fins, close to fin margin; spots all similar in size 

(Fig. 12b); cuttlebone width narrow, 20 to 25% the length; cuttlebone of approximately 

uniform width throughout its length, sides parallel (Fig. 13b) . . . . . . . . . . . . Sepiella ocellata 

5-6 winecoloured 

spots 

a) Sepiella weberi b) Sepia ocellata 

Fig. 12 dorsal view 

6-7 winecoloured 

spots 

broadening 

posteriorly 

a) Sepiella weberi b) Sepia ocellata 


5a. Mantle colour greyish brown with 8 or 9 reddish patches along base of each fin (Fig. 14); 

cuttlebone width approximately 40% the length; outer cone indented towards posterior 

end of striated zone (Fig. 15a), flared, spoon-shaped posteriorly; anterior striae 

m-shape, following the deep median sulcus (Fig. 15a) . . . . . . . . . . . . . . . Sepiella inermis 

5b. Mantle dark greyish brown with white spots and a pale reflective line along base of fins; 

cuttlebone width approximately 30% the length; outer cone tapers gradually posteriorly, 

not indented towards posterior end of striated zone (Fig. 15b); anterior striae inverted 

U-shape, median sulcus indistinct (Fig. 15b) (unconfirmed records from the Philippines) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepiella japonica 

Sepiella inermis 


8-9 

reddish 

blotches 

anterior 

striae 

m-shape 

deep 

sulcus 

outer cone 

indented 

a) Sepiella inermis b) Sepiella japonica 


(after Sasaki, 1929)

Sepiidae 729 

6a. Cuttlebone outline narrow, lanceolate (Fig. 16a) . . . . . → 7 

[includes Sepia mira, until recently known only from cuttlebone; see 

species account to follow] 

6b. Cuttlebone outline moderate to broad, oval (Fig. 16b) . . → 11 

[includes Sepia bartletti, status uncertain, known only from the type and 

6 other specimens reported by Iredale (1954) from Misima and the 

Conflict Group of Islands, Louisiade Archipelago, southeast of Papua 

New Guinea; see species account to follow] 

7a. Tentacular club with 5 or 6 suckers in transverse rows . . → 8 

7b. Tentacular club with 8 suckers in transverse rows . . .→10 8a. Arms I to IV of females with biserial suckers proximally 

and at extreme distal tips, remaining suckers 

tetraserial; male arms I to IV with biserial suckers 

on the proximal third and at distal tips, remaining 

suckers enlarged, tetraserial; male arms III protective 

membranes thickened with alternating transverse 

ridges . . . . . . . . . . . . . . . . . . . Sepia cottoni 

8b. Arms I to III of females with tetraserial suckers proximally 

and biserial suckers distally, female arms IV with 

tetraserial suckers; male arms I to III with tetraserial 

suckers proximally and biserial suckers distally, 

male arms IV with tetraserial suckers; male arms III 

protective membranes not thickened . . . . . . . . . . . → 9 

9a. Arms III of males greatly elongate; hectocotylus absent; 

club suckers differ markedly in size (Fig. 17a) . . Sepia braggi 1/ 

9b. Arms III of males not greatly elongate; hectocotylus 

present; club suckers differ only slightly in size 

(Fig. 17b) . . . . . . . . . . . . . . . . . . Sepia vietnamica 

10a. Arms II of males greatly elongate, 

3 times longer than other 

arms (Fig. 18); male arms II 

with tetraserial suckers proximally, 

biserial suckers at distal 

tips; left ventral hectocotylized 

arm (IV) with approximately 10 

rows of normal suckers proximally, 

then remaining suckers 

reduced to tip of arm; club suckers 

differ markedly in size 

male 

arms II 

greatly 

elongate 

(Fig. 19a) . . . . . . . Sepia andreana 

10b. Male arms all of similar length; 

male arm suckers all tetraserial; 

left ventral hectocotylized arm 

(IV) with approximately 9 to 12 

rows of normal suckers proximally, 

followed by 8 to 10 rows 

of reduced suckers, then rest 

normal to tip of arm; club suckers 

differ only slightly in size 

(Fig. 19b). . . . . . . . Sepia kobiensis 

I 

II 

head and anterior mantle 

of male (dorsal view) 

Fig. 18 Sepia 

andreana 

III 

IV 

a) lanceolate 

(Sepia braggi) 

b) oval 

(Sepia apama) 



a) Sepia braggi 

b) Sepia vietnamica 


a) Sepia andreana b) Sepia kobiensis 


1/ Until recently believed to occur in the Western Central Pacific. See footnote on species account to follow (p. 739).


11a. Tentacular club suckers all similar in size . . . . . . . . . . . . . → 12 

11b. Tentacular club suckers differ markedly in size . . . . . . . . . . → 24 

12a. Tentacular club with 8 or fewer suckers in transverse rows . . . . . .→ 13 

12b. Tentacular club with 10 or more suckers in transverse rows . . . . .→ 18 

13a. Arms I to IV of both sexes with biserial suckers proximally, 

tetraserial distally . . . . . . . . . . . . . . . . . . . . . . Sepia sulcata 

13b. Arms I to IV of both sexes with tetraserial suckers throughtout . . . . → 14 

14a. Cuttlebone inner cone forming a raised ledge posteriorly 

(Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . Sepia brevimana 

14b. Cuttlebone inner cone may be thickened, but does not form a 

raised ledge posteriorly . . . . . . . . . . . . . . . . . . . . . . . .→15 raised ledge 

15a. Hectocotylus absent; cuttlebone inner cone limbs not thickened 

posteriorly (Fig. 21) . . . . . . . . . . . . . . . . . . . . Sepia mestus 

15b. Hectocotylus present; cuttlebone inner 

cone limbs thickened posteriorly . . . . . . .→16 16a. Tentacular club with 5 or 6 suckers in 

transverse rows; anterior margin of cuttlebone 

triangular, posterior margin bluntpointed; 

cuttlebone striated zone flat, or 

slightly convex; cuttlebone outer cone of 

approximately uniform width along its 

length (Fig. 22) . . . . . . . . . . . Sepia cultrata 

16b. Tentacular club with 8 suckers in transverse 

rows; anterior margin of cuttlebone 

blunt-pointed or rounded, posterior margin 

rounded; cuttlebone striated zone concave; 

cuttlebone outer cone broadens 

posteriorly . . . . . . . . . . . . . . . . . .→ 17 

17a. Dorsal mantle with wrinkled 

striped pattern and distinctive 

ocellate patches, and a 

pale reflective line along 

base of fins (Fig. 23); cuttlebone 

anterior striae inverted 

V-shape; cuttlebone inner 

cone limbs thick, broadens 

posteriorly (Fig. 24a) . Sepia lycidas 

17b. Dorsal mantle brownish, 

sometimes with darker 

brown blotches and scattered 

white spots; cuttlebone 

anterior striae inverted 

U-shape; cuttlebone inner 

cone limbs thick, of uniform 

width (Fig. 24b) . . . Sepia madokai 

ocellate 

patches 

dorsal view 

Fig. 22 Sepia lycidas 

Sepia mestus 




inverted V-shape 

Sepia brevimana 


anterior 

margin 

triangular 

Sepia cultrata 




inverted U-shape 

a) Sepia lycidas b) Sepia madokai 

Fig. 23 cuttlebone (ventral view)

Sepiidae 731 

18a. Tentacular club with approximately 20 suckers in 

transverse rows . . . . . . . . . . . . . . . . . . . . . → 19 

18b. Tentacular club with 16 or less suckers in transverse 

rows . . . . . . . . . . . . . . . . . . . . . . . . → 20 

19a. Left ventral arm (IV) hectocotylized: 8 rows of 

normal arm suckers proximally, followed by 8 

rows of reduced suckers; 2 ventral series of reduced 

suckers smaller than 2 dorsal series; club 

swimming keel extends beyond sucker-bearing 

face of club; cuttlebone inner cone forming a 

raised ledge posteriorly (Fig. 25a) . . . . . . . . Sepia smithi 

19b. Both ventral arms (IV) hectocotylized: left ventral 

arm with 7 or 8 rows of normal suckers proximally, 

followed by 5 or 6 rows of reduced suckers, 

and right ventral arm with 4 or 5 rows of reduced 

suckers distally; reduced suckers equal-sized 

across series; club swimming keel equal to or 

shorter than sucker-bearing face of club; cuttlebone 

inner cone a narrow rim, does not form a 

inner cone 

forming a 

raised ledge 

posteriorly 

raised ledge posteriorly (Fig. 25b) . . . . . Sepia whitleyana 

a) Sepia smithi b) Sepia whitleyana 


20a. Cuttlebone spine with dorsal and ventral keel (Fig. 26) . . . . . . . . . . . . . . . . Sepia stellifera 

20b. Cuttlebone spine without keel(s) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .→ 21 

21a. Hectocotylus with 9 or 10 rows of reduced suckers; cuttlebone inner cone limbs uniform 

in width, not forming a ledge posteriorly (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . Sepia rex 

21b. Hectocotylus with between 5 and 7 rows of reduced suckers; cuttlebone inner cone limbs 

broaden and form a ledge posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .→ 22 

22a. Hectocotylus (left ventral arm IV) with 7 or 8 rows of normal suckers proximally, followed 

by 7 rows of slightly reduced suckers; dorsal and ventral protective membranes not fused 

at base of tentacular club in small specimens, fused in large specimens; cuttlebone inner 

cone with posterior ledge not thickened, flat, thin; dull white, not shiny (Fig. 28) . . . . . Sepia elliptica 

22b. Hectocotylus (left ventral arm IV) with 6 or less rows of normal suckers proximally, 

followed by 5 or 6 rows of markedly reduced suckers; dorsal and ventral protective 

membranes not fused at base of club; cuttlebone inner cone with posterior ledge 

thickened, rounded, not flat; ledge shiny, yellowish . . . . . . . . . . . . . . . . . . . . . . . .→ 23 

spine with 

dorsal and 

ventral keel 

Sepia stellifera 



inner cone 

limbs uniform 

in width 

Sepia rex 



inner cone not 

thickened 

posteriorly 

Sepia elliptica 


(ventral view)


23a. Hectocotylus with 3 rows of normal suckers proximally; cuttlebone anterior striae 

inverted U-shape (Fig. 29a). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepia aculeata 

23b. Hectocotylus with 5 to 6 rows of normal suckers proximally; cuttlebone anterior striae 

inverted V-shape (Fig. 29b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepia esculenta 

24a. Tentacular club with 6 or less suckers in transverse rows . . . . . . . . . . . . . . . . . . . . . .→25 [Ghofar (1989) reports Sepia bandensis as having 7 to 9 suckers in transverse rows on the tentacular club. The 

number of suckers in transverse rows cited in the original description of Adam (1939), and by subsequent workers 

e.g. Okutani (1995), and followed here is 5] 

24b. Tentacular club with 8 suckers in transverse rows . . . . . . . . . . . . . . . . . . . . . . . . . → 31 

25a. Tentacular club with 3 or 4 suckers in transverse rows . . . . . . . . . . . . . . . . Sepia opipara 

25b. Tentacular club with 5 or 6 suckers in transverse rows . . . . . . . . . . . . . . . . . . . . . .→ 26 

26a. Cuttlebone without spine, or spine reduced to a tiny tubercle . . . . . . . . . . . . . . . . . .→ 27 

26b. Cuttlebone with spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .→ 28 

27a. Maximum size 500 mm mantle length; anterior edge of inner cone with a V-shaped 

calcareous callus, or rough thickening posteriorly (Fig. 30a) . . . . . . . . . . . . . . Sepia apama 

27b. Maximum size 50 mm mantle length; cuttlebone inner cone without calcareous callus 

posteriorly (Fig. 30b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepia bandensis 


inverted 

U-shape 

a) Sepia aculeata b) Sepia esculenta 


anterior 

striae 

inverted 

V-shape 

28a. Male arms I to IV with tetraserial 

suckers cuttlebone inner cone limbs 

of uniform width; cuttlebone sulcus 

distinct, flanked by rounded ribs 

(Fig. 31a, b) . . . . . . . . . . . . . . . . .→ 29 

28b. Male arms I to III with tetraserial 

suckers proximally, biserial suckers 

distally; cuttlebone inner cone limbs 

broaden posteriorly; cuttlebone sulcus 

indistinct, not flanked by 

rounded ribs (Fig. 32a, b) . . . . . . . . . .→ 30 

calcareous 

callus 

a) Sepia apama 

b) Sepia bandensis 



anterior 

striae 

inverted 

U-shape 

sulcus 

shallow 

but distinct 

a) Sepia latimanus b) Sepia plangon 


anterior 

striae 

inverted 

V-shape 

sulcus 

deep

Sepiidae 733 

29a. Hectocotylus absent; tentacular club separated from stalk by a membrane; cuttlebone 

spine without keel; cuttlebone sulcus shallow (Fig. 31a); cuttlebone anterior striae blunt 

inverted V-shape (Fig. 31a); males dark purple-brown with white spots and white 

transverse stripes on fins, extending to mantle . . . . . . . . . . . . . . . . . . . . Sepia latimanus 

29b. Hectocotylus present; tentacular club joined to stalk, not separated from stalk by a 

membrane; cuttlebone spine with keel; cuttlebone sulcus deep (Fig. 31b); cuttlebone 

anterior striae inverted V-shape (Fig. 31b); males with pale, transverse stripes . . . . Sepia plangon 

30a. Female arms I to III with tetraserial 

suckers throughout length; hectocotylus 

absent; cuttlebone inner 

cone limbs not thickened posteriorly, 

do not form a rough chitinous callus 

(Fig. 32a) . . . . . . . . . . . . Sepia papuensis 

30b. Female arms I to III with tetraserial 

suckers proximally, biserial suckers 

on distal tips; hectocotylus present; 

cuttlebone inner cone limbs with a 

thick, rough, chitinous callus posteriorly 

(Fig. 32b) . . . . . . . . Sepia recurvirostra 

31a. Dorsal and ventral protective membranes 

fused at base of club; club 

separated from stalk by a membrane; 

cuttlebone inner cone limbs broaden 

posteriorly, forming a thick pink rim 

(Fig. 33) . . . . . . . . . . . . . . . Sepia rozella 

31b. Dorsal and ventral protective membranes 

not fused at base of club; club 

joined to stalk, not separated from 

stalk by a membrane; cuttlebone inner 

cone limbs of uniform width, or 

form a distinctive rounded swelling, 

not pink . . . . . . . . . . . . . . . . . . . → 32 

32a. Cuttlebone inner cone limbs 

broaden, thicken posteriorly to form 

a distinct bulbous, shiny swelling 

(Fig. 34); cuttlebone sulcus deep, 

wide, flanked by rounded ribs; hectocotylus 

(left ventral arm IV)with10 

rows of normal suckers at base followed 

by reduced suckers . . . . Sepia pharaonis 

32b. Cuttlebone inner cone limbs of uniform 

width, form a thickened rim, do 

not form a bulbous swelling; cuttlebone 

sulcus shallow, narrow, not 

flanked by rounded ribs; hectocotylus 

(left ventral arm IV) with 3 rows 

of normal suckers at base, followed 

by reduced suckers . . . . . . . . . . Sepia vossi 

inner cone 

limbs not 

thickened 

thick 

calcareous 

callus 

posteriorly 

a) Sepia papuensis b) Sepia recurvirostra 


thick 

pink rim 

Sepia rozella 



bulbous 

swelling 

Sepia pharaonis 


(ventral view)




Metasepia pfefferi (Hoyle, 1885) 

Sepia aculeata Férussac and d’Orbigny, 1848 

Sepia andreana Steenstrup, 1875 

Sepia apama Gray, 1849 

Sepia bandensis Adam, 1939 

Sepia bartletti (Iredale, 1954) 1/ 

[Sepia braggi Verco, 1907] 2/ 

Sepia brevimana Steenstrup, 1875 

? Sepia carinata Sasaki, 19203/ Sepia cottoni Adam, 1979 

Sepia cultrata Hoyle, 1885 

Sepia elliptica Hoyle, 1885 

Sepia esculenta Hoyle, 1885 

Sepia kiensis Hoyle, 1885 

Sepia kobiensis Hoyle, 1885 

Sepia latimanus Quoy and Gaimard, 1832 

? Sepia lorigera Wülker, 19103/ Sepia lycidas Gray, 1849 

Sepia madokai Adam, 1939 

Sepia mestus Gray, 1849 

Sepia mira (Cotton, 1932) 1/ 

Sepia opipara (Iredale, 1926) 

Sepia papuensis Hoyle, 1885 

Sepia pharaonis Ehrenberg, 1831 

Sepia plangon Gray, 1849 

Sepia recurvirostra Steenstrup, 1875 

Sepia rex (Iredale, 1926) 

Sepia rozella (Iredale, 1926) 

Sepia smithi Hoyle, 1885 

Sepia stellifera Homenko and Khromov, 1984 

Sepia sulcata Hoyle, 1885 

Sepia vietnamica Khromov, 1987 

Sepia vossi Khromov, 1996 

Sepia whitleyana (Iredale, 1926) 

Sepiella inermis Van Hasselt, 1835 

? Sepiella japonica Sasaki, 19294/ Sepiella ocellata Pfeffer, 1884 

Sepiella weberi Adam, 1939 

Sepiella sp. 5/ 

1/ Uncertain status. 

2/ Until recently believed to occur in the area. See footnote on species account to follow (p. 739). 

3/ Unconfirmed records from Viet Nam. 

4/ Species known as Sepiella maindroni in China. It is important in the fisheries of Japan, South Korea, and China. 

Unconfirmed records from the Philippines. 

5/ New species from northern Australia (A. Reid and C.C. Lu, submitted manuscript).

Sepiidae 735 

References 

Adam, W. 1939. Cephalopoda, II. Révision des espèces Indo-Malaises du genre Sepia Linné, 1758. III. Revision du 

genre Sepiella (Gray) Steenstrup, 1880. Siboga Exped., Leiden, 55b(135):35-122. 

Adam, W. 1979. The Sepiidae (Cephalopoda, Decapoda) in the collections of the Western Australian Museum. Rec. 

West. Aust. Mus., 7(2):111-212. 

Adam, W. and W.J. Rees. 1966. A review of the cephalopod family Sepiidae. Scientific reports of the John Murray 

Expedition 1933-1934. 11(1):165 p. 

Hoyle, W.E. 1886. Report on the Cephalopoda collected by H.M.S. “Challenger” during the years 1873-1876. Rep. Sci. 

Res. Voy ’Challenger’1873’76(Zool.), (16)44:245 p. 

Khromov, D.N. 1987. A new species of Sepia (Cephalopoda:Sepiidae) from the north-western South China Sea. Asian 

Mar. Biol., 4:35-40. 

Khromov, D.N. 1988. Cuttlefishes of the famly Sepiidae (Cephalopoda) of the Zoological Institute of the USSR Academy 

of Sciences. Tr. Zool. Inst. Akad. Nauk SSSR, 171:174-195. [in Russian] 

Khromov, D.N. 1996. Some notes on the shelf and slope cephalopod fauna of Vietnam, and a new species of Sepia 

(Cephalopoda, Sepiidae) from this region. Ruthenica, 5(2):139-145. 

Khromov, D.N., C.C. Lu, A. Guerra, Z. Dong, and S.V. Boletzky. (in press). A synopsis of Sepiidae outside Australian 

waters (Cephalopoda: Sepioidea). Smithson. Contrib. Zool. 

Lu, C.C. (in press). A synopsis of Sepiidae in Australian waters. Smithson. Contrib. Zool. 

Lu, C.C. and M.C. Dunning. (in press). Subclass Coleoidea Bather, 1888. Fauna of Australia. Volume 5. Mollusca. 

Canberra, Australian Government Publishing Service. 

Lu, C.C. and A.L. Reid. 1997. Two new cuttlefishes (Cephalopoda: Sepiidae) from the North West Shelf, and a 

redescription of Sepia sulcata Hoyle, 1885. Rec. W.A. Mus., 18:277-310. 

Nesis, K.N. 1987. Cephalopods of the world: squid, cuttlefish, octopuses and their allies. Neptune City, New Jersey, 

T.F.H. Publications Inc. Ltd., 351 p. [Levitov, B.S. trans. from Russian, edited by L.A. Burgess, 1987]. 

Okutani, T. 1995. Cuttlefish and squids of the world in colour. Tokyo, Japan, Okumura Printing Co. Ltd., 186 p. 

Reid, A.L. (in press) A complete description of Sepia mira (Cotton, 1932) (Cephalopoda: Sepiidae) from eastern 

Australia. Proc. Linn. Soc. N.S.W., 119. 

Voss, G.L. 1963. Cephalopods of the Philippine Islands. Bull. U.S. Natl. Mus., (234):180 p. 

Voss, G.L. and G.R. Williamson. 1971. Cephalopods of Hong Kong. Hong Kong, Hong Kong Government Press, 138 p.


Sepiidae Sepia aculeata Férussac and d’Orbigny, 1848 

Frequent synonyms / misidentifications: Sepia indica Férussac and d’Orbigny, 1848 / None. 

FAO names: En - Needle cuttlefish; Fr - Seiche aiguille; Sp - Sepia con punta. 

dorsal view 

ventral view 

cuttlebone 

10 mm 

tentacular club dorsal view 

Diagnostic characters: Left ventral arm (IV) of males hectocotylized: 3 rows of normal suckers proximally, 

followed by 5 or 6 rows of reduced suckers, remaining suckers normal to tip of arm; reduction marked, 

suckers in 2 dorsal series smaller than rest; 2 dorsal and 2 ventral series widely spaced. Tentacular club 

long, with 10 to 12 suckers in transverse rows (males), or13 or 14 suckers in transverse rows 

(females); suckers all of similar size, minute; dorsal and ventral protective membranes not fused at base 

of club, extending beyond sucker-bearing face along stalk. Buccal membrane with few, minute suckers. 

Cuttlebone outline oval; dorsal and lateral ribs present, distinct; spine present, keel(s) absent; anterior 

striae inverted U-shape; inner cone limbs broaden posteriorly, raised to form a thickened, rounded, 

ledge; outer cone narrow anteriorly, broadens posteriorly. Colour: variable; dorsal mantle with fine 

transverse reticulated colour pattern in the spawning season, or may be pale brownish with, or without 

white blotches or spots, sometimes with transverse saddle mark; fins with pale reflective line along base. 

Size: Maximum mantle length 230 mm; maximum weight 1.3 kg. Commercially caught animals from the 

Gulf of Thailand and the Andaman Sea range between 60 and 130 mm mantle length. 

Habitat, biology, and fisheries: A demersal neritic species; at depths to 60 m. Spawning occurs all year in the 

Gulf of Thailand, with peak months from March to April and July to September. Males in this region mature at 

70 mm, and females at 810 mm mantle length, with the sex ratio of males to females 1:1.3. An object of fisheries 

in southern China, Taiwan Province of China, and also commercially important in Thailand; mostly caught using 

otter trawl, smaller catches are 

made using pair trawl, and to a 

lesser extent, squid light-lures, 

traps, and push nets; bottom 

otter and pair trawls are used 

offshore, and push nets and lift 

nets in inshore and coastal 

waters. 

? 

Distribution: Southern India 

to Andaman Sea, Indonesia, 

Thailand, Malaysia, the 

Philippines, China, and north 

to central Japan.

Sepiidae 737 

Sepia andreana Steenstrup, 1875 

Frequent synonyms / misidentifications: 

None / Sepia kobiensis Hoyle, 1885. 

FAO names: En - Andrea cuttlefish; 

Fr - Seiche andreana; Sp - Sepia andreana. 

Diagnostic characters: Fins end in small 

auriculate lobes posteriorly, with a small Vshaped 

interstice between lobes where the 

cuttlebone spine is located. Male arms II 

greatly elongated, 3 times longer than the 

other arms (except in very young animals); 

rounded distally, not tapered.Male arms II with 

4 suckers in transverse rows proximally and 2 

suckers in transverse rows distally, with suckers 

becoming rudimentary and sparse; male arms I, 

III and IV with 4 suckers in transverse rows. 

Female arms all with 4 suckers in 

transverse rows. Left ventral arm 

(IV) of males hectocotylized: 

approximately 10 rows of normal 

suckers proximally, remaining 

suckers greatly reduced, 

rudimentary, on swollen peduncular 

bases; reduced sucker rows 

evenly spaced on arm. Tentacular 

club with 8 suckers in transverse 

rows; differing markedly in size, 

with 4 or 5 large median suckers; 

swimming keel extending slightly 

beyond sucker-bearing face; dorsal 

and ventral protective membranes 

not fused at base of club, 

terminating at posterior end of sucker- 

bearing face, dorsal membrane 

forming deep cleft at junction with 

the stalk. Cuttlebone lanceolate; 

without dorsal median rib present; 

striated zone and last loculus 

dorsal 

view 

tentacular 

club 


ventral 

view 

cuttlebone 


arms II elongate 

dorsal view 

head of male 

dorsal view 


convex; sulcus shallow, narrow; anterior striae M-shape; inner cone limbs of uniform width. Colour: dorsal 

mantle pale brownish with yellow spots; arms I to III with orange pigmented stripe along aboral surface. 

Size: Maximum mantle length 120 mm. 

Habitat, biology, and fisheries: A demersal species occurring in coastal waters to a depth of 50 m. Taken 

as bycatch in trawl and set-net fisheries. 

Distribution: Western Pacific from northern Philippines, along south China coast to central Japan. 

Remarks: May be confused with Sepia kobiensis. Mature males of S. andreana can be distinguished by the 

greatly elongate second arm 

pair and the hectocotylus 

? 

sucker arrangement: in S. 

kobiensis, 9 to 12 rows of 

normal suckers are followed 

by 8 to 10 rows of reduced 

suckers, then remaining 

suckers normal to tip of arm. 

Also, S. kobiensis is smaller, 

reaching a maximum mantle 

length of 70 mm. 

 

 




Sepia apama Gray, 1849 

Frequent synonyms / misidentifications: Sepia 

palmata Owen, 1881; Amplisepia verreauxi 

Iredale, 1926; A. parysatis Iredale, 1954 / None. 

FAO names: En - Giant Australian cuttlefish; 

Fr - Seiche géante; Sp - Sepia gigante. 

Diagnostic characters: Three large flap-like 

papillae posterior to each eye; sometimes 

additional smaller papillae anterior to these large 

papillae. Left ventral (IV) arm of males 

hectocotylized, proximal 6 to 10 sucker rows 

slightly reduced; reduced suckers equal in size 

across rows. Tentacular club crescent-shaped; 

sucker-bearing face flattened, with 5 or 6 suckers 

in transverse rows, differing markedly in size 

(median suckers enlarged); swimming keel 

extending slightly beyond sucker-bearing face; 

dorsal and ventral protective membranes fused at 

base of club, separated from stalk by a membrane. 

Cuttlebone outline broadly oval in juveniles, wider 

in anterior half, becoming elongate in adults; 

cuttlebone rounded anteriorly and posteriorly in 

juveniles, acuminate in adults; dorsal surface with 

faint median and lateral ribs; spine present in 

juveniles, lost in adults; last loculus flat, striated 

zone convex, with shallow sulcus, anterior striae 

inverted U-shape; inner cone limbs broadening 

posteriorly, anterior margin forms a rough 

calcareous callus posteriorly; outer cone narrow 

anteriorly, broadens posteriorly, extends well 

beyond inner cone in adults. Colour: during 

breeding season, dorsal mantle with bold, 

transverse stripes. 

Size: Maximum mantle length 500 mm; maximum 

weight over 5 kg. One of the largest cuttlefishes. 

Habitat, biology, and fisheries: A neritic 

demersal species occurring in rocky areas, 

seagrass beds and on open trawl grounds to a 

depth of 100 m. Spawning extends from May to 

September. Taken as bycatch of prawn and mixed 

species trawl fisheries and also caught by 

hook-and-line, or speared by divers. Commonly 

seen in fish markets along the southern coast of 

Australia. Sold for human consumption and as bait. 

Distribution: Southern Australia. 

20 mm 

tentacular 

club 


dorsal view 


ventral view 

30 mm dorsal view 

lateral view 

cuttlebone (subadult) 

(illustration: K. Hollis/ABRS)

Sepiidae 739 

Sepia braggi Verco, 1907 

Frequent synonyms 1/ / misidentifications: Arctosepia 

limata Iredale, 1926; A. versuta Iredale, 1926; A. rhoda 

Iredale 1954; A. treba Iredale 1954; A. braggi xera Iredale, 

1954 / None. 

FAO names: En - Slender cuttlefish; Fr - Seiche gracile; 

Sp - Sepia grácil. 

tentacular 

club 

ventral 

view 

10 mm dorsal 

lateral view view 

cuttlebone 


dorsal view 

Diagnostic characters: Mantle shape oblong. Both sexes with longitudinal row of 5 or 6 ridges along each 

side, close to fins. Male arms III greatly elongated; female arms all similar in length. Male and female 

arm suckers: arms I to III with tetraserial suckers proximally, biserial suckers at distal tips; arms IV 

with tetraserial suckers throughout. Hectocotylus absent. Tentacular club short, with 5 or 6 suckers 

in transverse rows, differing markedly in size, 5 or 6 median suckers twice the diameter of rest; dorsal and 

ventral protective membranes not fused at base of club, joined to stalk at base of club, extending beyond 

sucker-bearing face along stalk; dorsal protective membrane much wider than ventral membrane. Cuttlebone 

outline lanceolate; dorsal surface pinkish; evenly convex; faint median rib present; spine present, long, curving 

upward, keel(s) absent; ventral surface convex; sulcus deep, narrow, flanked by rounded ribs; anterior striae 

m-shaped; inner cone limbs of uniform width, thickened posteriorly forming a shiny rounded ridge; outer cone 

limbs expanded forming 2 short wings. Colour: pinkish purple dorsally, pale ventrally; arms II to IV with 

median longitudinal dark pink strips 

and transverse bars and spots. 

Size: Maximum mantle length 

65 mm. 

Habitat, biology, and fisheries: A 

demersal species, at depths between 

30 and 146 m.Taken as bycatch with 

other cuttlefish species in southern 

and southeastern Australia. 

Distribution: Southern Australia 

(see footnote!). 

1/ Since the time of writing, some additional information has been become available. Lu (in press) included 5 species 

in synonymy with Sepia braggi (see section on “Frequent synonyms” above). Work in progress by the author [A.L. 

Reid] and currently being prepared for publication has shown S. limata (Iredale, 1926) and S. rhoda (Iredale, 1954) 

to be valid species. In addition, the distribution of S. braggi is now known to be restricted to the southeastern corner 

of Australia (outside the WCP area). 

10 mm


Sepia brevimana Steenstrup, 1875 

Frequent synonyms / misidentifications: Sepia rostrata 

Férussac and d’Orbigny, 1848 / Sepia stellifera Homenko 

and Khromov, 1984. 

FAO names: En - Shortclub cuttlefish; Fr - Seiche petites 

mains; Sp - Sepia mazicorta. 

Diagnostic characters: Mantle blunt-pointed posteriorly. 

Tentacular club small; with 5 to 8 minute suckers in 

transverse rows; swimming keel extending well beyond 

sucker-bearing face; dorsal and ventral protective 

membranes not fused at base of club, dorsal membrane 

much wider than ventral membrane. Cuttlebone very 

angular, V-shaped anteriorly; dorsal surface flat, texture 

uniformly pustulose; spine present, long, keels present 

dorsally and ventrally, also with nose-like protuberance in the 

anterior part of spine; a shallow, narrow sulcus extending 

along striated zone, flanked by rounded ribs; anterior striae 

inverted V-shape; inner cone limbs broaden posteriorly, 

thickened, rose-coloured or yellowish orange, outer 

margin of inner cone forming a raised, flattened ledge 

posteriorly; outer cone narrow anteriorly, broadens 

posteriorly. Colour: buff-brown. 

Size: Maximum mantle length 100 mm. In the Gulf of 

Thailand, commonly caught at 40 to 60 mm mantle length, 

with a maximum mantle length of 90 mm. 

Habitat, biology, and fisheries: A demersal species, 

ranging to a depth of 100 m. The vertical range is 10 to 40 m 

10 mm 

and the peak occurrence 30 to 40 m in the Gulf of Thailand 

ventral view 

and the Andaman Sea, with the sex ratio of males to females 

caught 1:2.2. Important to the commercial squid fishery in 

cuttlebone 

Thailand. Most are caught using otter trawl, some using pair 

trawl, and small catches are made using squid light-lures, traps and push nets, with bottom otter and pair 

trawls used offshore, and push nets and lift nets used in inshore and coastal waters. It is also fished in 

South China Sea. 

Distribution: Southern India to the Andaman Sea, Gulf of Tonkin, including Java, Sulu, and Celebes Seas. 

Remarks: The number of tentacular club suckers in transverse rows has been recorded as 5, 5 to 8, and 

6 to 8 by various workers. The exact number needs to be confirmed. Can be confused with Sepia stellifera, 

but differs in having up to 8, 

rather than 10 suckers in 

transverse rows on the club, 

and the shallow and narrow 

sulcus of the cuttlebone, 

which is deep and wide in S. 

stellifera. Similar to S. 

esculenta, but differs in having 

fewer club suckers (10 to 16 

suckers in transverse rows in 

S. esculenta), in having no 

fleshy papillae along the base 

of the fins, and in having a 

coloured inner cone.

Sepiidae 741 

Sepia elliptica Hoyle, 1885 

Frequent synonyms / misidentifications: None / 

Sepia esculenta Hoyle, 1885; S. stellifera Homenko 

and Khromov, 1984. 

FAO names: En - Ovalbone cuttlefish. 

Diagnostic characters: Left ventral arm (IV) of 

males hectocotylized: 7 or 8 rows of normal 

suckers proximally, followed by 7 rows of 

reduced suckers, then remainder normal to arm 

tip; reduction slight; 2 dorsal series smaller than 

remaining suckers.Tentacular club sucker-bearing 

face flattened; with 10 to 12 minute suckers in 

transverse rows; dorsal and ventral protective 

membranes not fused in small specimens, fused at 

base of club in large specimens, separated from 

stalk by a membrane. Cuttlebone outline oval; 

rounded posteriorly; median ribs indistinct; spine 

present, keel(s) absent; sulcus deep, wide; anterior 

striae inverted U-shape; inner cone limbs 

broadening posteriorly, not thickened, outer 

margin of inner cone forms a raised, flattened 

anteriorly directed ledge; outer cone narrow 

anteriorly, broadens posteriorly. Colour: pale 

pinkish purple. 

ventral view 


cuttlebone 

Habitat, biology, and fisheries: Coastal waters at 

depths from 10 to 142 m. In the Gulf of Carpentaria, 

both sexes of Sepia elliptica reach maturity at 

36 mm mantle length, with broad size ranges of both 

sexes over most of the year indicative of extended spawning season. 

Distribution: Northern Australia and New Guinea, South China Sea, possibly 

Philippines. 

Remarks: May be confused with Sepia esculenta which can be distinguished by the 

following characters: hectocotylus with 5 or 6 rows of normal suckers proximally, 

followed by 6 rows of reduced suckers; dorsal and ventral protective membranes not 

fused at base of club; cuttlebone anterior striae inverted V-shape, and inner cone 

limbs thickened posteriorly. 

? 

? 

dorsal ventral 

hectocotylus 

(after Hoyle, 1886) 

tentacular club 

(subadult)


Sepia esculenta Hoyle, 1885 


Sepia elliptica Hoyle, 1885. 

FAO names: En - Golden cuttlefish; Fr - Seiche dorée; 

Sp - Sepia dorada. 

tentacular 

club 


10 mm 

ventral view 

cuttlebone 

dorsal view 


Diagnostic characters: Mantle with a row of 6 or 7 yellowish fleshy tubercles along silvery stripe at 

base of each fin. Left ventral arm (IV) of males hectocotylized: 5 or 6 rows of normal suckers proximally, 

followed by 6 rows of reduced suckers, then remaining suckers normal to tip of arm. Tentacular club 

long; sucker-bearing face flattened, with 10 to 16 similar-sized, minute suckers in transverse rows; 

dorsal and ventral protective membranes not fused at base of club. Buccal membrane without suckers. 

Cuttlebone rounded anteriorly and posteriorly; spine present; sulcus deep, wide; anterior striae inverted 

V-shape; inner cone limbs thickened posteriorly, forming a raised, rounded ledge. Colour: dorsal 

mantle pale with transverse stripes, sometimes studded with yellow tubercles; fins with pale, golden 

reflective line along base, visible dorsally, and ventrally; arms with pinkish bands along the aboral 

surface. 

Size: Maximum mantle length 175 mm; maximum weight 0.6 kg. 

Habitat, biology, and fisheries: A demersal inner shelf species found on sandy bottoms between depths 

of 10 and 100 m, sometimes burrowing into the substrate. After overwintering in deeper water, animals 

migrate into shallower coastal waters where they spawn when the water temperature increases in spring 

and summer.Eggs are deposited on macrophytes and other substrates.Supports localized and subsistence 

fisheries in the Philippines and Hong Kong. Caught using otter trawls, pound nets, hoop nets and on 

hook-and-line. The flesh is highly thought of as food. 

Distribution: South and East 

China seas. Japan, Hong Kong 

(China), Viet Nam, northern 

Philippines, and Indonesia. 

Remarks: This species has 

been confused with Sepia 

elliptica. InS. elliptica,theinner 

cone ledge of the cuttlebone is 

thinner, flatter and directed 

anteriorly, and the anterior striae 

are inverted U-shaped. The 

hectocotylus sucker arrangement 

also differs.

Sepiidae 743 

Sepia kobiensis Hoyle, 1885 


andreanoides Hoyle, 1885; S. kobiensis var. 

andreanoides Sasaki, 1929; S. kobiensis var. 

toyamensis Sasaki, 1929; S. kobiensis var. beppauna 

Sasaki, 1929; S. kobiensis var. crassa Sasaki, 1929; S. 

kobiensis var. albatrossi Sasaki, 1920 / Sepia 

andreana Steenstrup, 1875. 

FAO names: En - Kobi cuttlefish; 

Fr - Seiche kobi; Sp - Sepia kobí. 

Diagnostic characters: Body 

narrow. All arms similar in length, 

with tetraserial suckers. Left 

ventral arm (IV) of males 

hectocotylized: 9 to 12 rows of 

normal suckers proximally, 

followed by 8 to 10 rows of 

reduced suckers, then remaining 

suckers normal to tip of arm; 

reduction marked, 2 dorsal and 2 

ventral series widely spaced; oral 

surface wide, fleshy, with 

transversely grooved ridges and 

adeepmedianfurrow. Tentacular 

club short, with 8 suckers in 

transverse rows, differing only 

slightly in size; swimming keel 

extends slightly beyond club. 

Cuttlebone lanceolate, slightly 

broader in females; dorsal surface 

10 mm 

ventral view 

cuttlebone 

dorsal view 

pinkish, or yellowish; spine present, long; sulcus shallow, narrow; anterior striae 

inverted m-shape; inner cone limbs of uniform width, thickened, U-shaped posteriorly; 

outer cone limbs narrow throughout most of length, limbs expanded posteriorly 

forming 2 short wings.Colour: dark reddish purple; head with V-shaped reddish stripe 

on dorsal margin of orbit; dorsal mantle brownish with darkish central area and reddish 

orange spots; reddish stripes on aboral surface arms I to III. 


Habitat, biology, and fisheries: Subtidal to a depth of 150 m or more. Taken in small 

quantities in the Hong Kong, China area, also caught as bycatch in small scale fisheries 

off southern Japan and China, mainly with fixed nets, trawls, and beach seines. 

Distribution: South and East China Seas, Yellow Sea to southern and central Japan. 

Indo-Malayan waters, Philippines, Gulf of Tonkin and the northern part of the Indian 

Ocean from the Arabian Sea and the Persian Gulf to Myanmar. 

tentacular 

club 

Remarks: Probably a complex of related species. Sepia kobiensis has been confused with S. andreana.In 

adult male S. andreana, the second pair of arms is greatly elongate and bears biserial suckers, rather than 

tetraserial suckers; the hectocotylus has about 10 rows of normal suckers at the base, and the remaining 

suckers are reduced and very 

rudimentary. Also, the club 

suckers differ markedly in size 

in S. andreana, while those in ? 

S. kobiensis differ only slightly 

? 

in size.


Sepia latimanus Quoy and Gaimard, 1832 


Ponderisepia eclogaria Iredale, 1926; Sepia rappiana 

Férussac, 1835; S. mozambica Rochebrune, 1884; S. 

hercules Pilsbry, 1894 / None. 

FAO names: En - Broadclub cuttlefish; Fr -Seiche 

grandes mains; Sp - Sepia mazuda. 


(from Roper et al., 1984) 

10 mm 

ventral view 

cuttlebone 

dorsal view 


Diagnostic characters: Head tubercles present, papilliform. Hectocotylus absent. Tentacular club 

crescent-shaped, sucker-bearing face flattened; with 5 or 6 suckers in transverse rows, differing 

markedly in size with a few median suckers enlarged; swimming keel extends beyond sucker-bearing face; 

dorsal and ventral protective membranes fused at base of club, separated from stalk by membrane; 

dorsal protective membrane forms a deep cleft at junction with stalk. Cuttlebone rounded anteriorly and 

posteriorly; dorsal surface granulose, loosely following growth lines; spine present, short; keel(s) absent; 

sulcus shallow, narrow, extending entire length of cuttlebone, flanked by rounded ribs; anterior 

striae blunt inverted V-shape; inner cone limbs thickened posteriorly, form a rounded, shiny ridge; outer 

cone deep, cup-like posteriorly. Colour: dorsal mantle dark purple/brown with white spots; fins with 

white transverse stripes extending onto mantle; arms vivid with transverse stripes. 

Size: Maximum mantle length 500 mm. Males up to 170 mm (0.5 kg) and females to 240 mm (1.3 kg) 

mantle length in Alas Strait, Indonesia. 

Habitat, biology, and fisheries: A shallow-water species inhabiting tropical coral reefs to a depth of 30 m. 

Mating occurs on the west coast of Guam and off Okinawa in shallow waters from January to May and the 

eggs hatch after 38 to 40 days. This species supports local fisheries in western Japan and the Philippines. 

Caught with jigs, handlines, set nets, and spears and commonly taken as bycatch in Southeast Asian trawl 

fisheries. Fished in small quantities in the Ryukyu Islands, China, near Taiwan Province of China, and in 

the waters of Indochina. In the Philippines, the large cuttlefish, Sepia pharaonis and S. latimanus, are split 

open, the cuttlebone and viscera removed and are dried in the sun without salt. 

Distribution: Widespread in 

the tropical Indo-West Pacific 

from the Andaman Sea, South 

and East China Seas, Japan, 

the Philippines, Malaysia, 

Indonesia to New Guinea, 

northern Australia and east to 

Palau, Guam, New Caledonia, 

and Fiji. Westernmost record 

Madagascar (doubtful).

Sepiidae 745 

Sepia lycidas Gray, 1849 

Frequent synonyms / misidentifications: Sepia subaculeata Sasaki, 1914 / None. 

FAO names: En - Kisslip cuttlefish; Fr - Seiche baisers; Sp - Sepia labiada. 

10 mm 

ventral view 

cuttlebone 



dorsal view 

Diagnostic characters: Left ventral arm (IV) of males hectocotylized: 6 rows of normal suckers proximally, 

then 4 rows of reduced suckers. Tentacular club with 8 similar-sized suckers in transverse rows; 

swimming keel extending nearly to base of club; dorsal and ventral protective membranes not fused 

at base of club. Buccal membrane with a few, minute suckers, single sucker on most lobes. Cuttlebone 

outline elliptical, rounded anteriorly and posteriorly, dorsal median rib absent; spine present, short; sulcus 

deep, wide; anterior striae inverted V-shape; inner cone limbs thickened posteriorly broadens to 

form a rounded ridge, outer cone narrow anteriorly, broadens posteriorly. Colour: head, mantle and fins 

with wrinkled zebra-stripe pattern; dorsal mantle with scattered ocellate patches (pattern more obvious 

in males than females). Fins with wide stripe at base. 

Size: Maximum mantle length about 200 mm. In the Gulf of Thailand, most animals caught are between 

100 and 200 mm mantle length, with maximum mantle length 380 mm and maximum weight 5 kg. 

Habitat,biology,and fisheries: A neritic demersal species.The depth range in the Gulf of Thailand and Andaman 

Sea is 10 to 100 m, with most animals caught between 20 to 40 m.In the South China Sea, it is abundant between 

depths of 60 and 100 m in the pre-spawning period (November to February), and migrates inshore to spawn in 

depths of 15 to 30 m from March to May. Sepia lycidas is important to the commercial cuttlefish fishery in Thailand 

and Viet Nam. Most cuttlefish are caught off Thailand using otter trawl, with smaller catches made using pair trawl 

and to a lesser extent, squid light-lures, traps and push nets, with bottom otter and pair trawls used offshore, and 

push nets and lift nets used in 

inshore and coastal waters. 

The mantle flesh is thick and 

tasty and therefore highly 

esteemed. 

Distribution: East and South 

? 

China Seas, the Philippines, Viet 

Nam, and Borneo. Commonly 

distributed in the Andaman Sea. 

In the Gulf of Thailand it occurs 

south of 10°N, and never 

appears in the inner and eastern 

coast of the Gulf.


Sepia opipara (Iredale, 1926) 

Frequent synonyms / misidentifications: Glyptosepia opipara Iredale, 1926 / None. 

FAO names: En -Staregaze cuttlefish. 

Diagnostic characters: Both sexes with tetraserial 

arm suckers. Left ventral arm (IV) of males 

hectocotylized: 5 or 6 rows of normal suckers 

proximally, followed by 6 or 7 rows of reduced 

suckers, then remainder normal to arm tip.Tentacular 

club sucker-bearing face flattened; 

with 3 or 4 suckers in transverse 

rows; suckers differ markedly in 

size, middle of club with 4 or 5 big 

suckers; swimming keel extending 

well beyond club; dorsal and ventral 

protective membranes not fused at 

base of club, joined to stalk at base 

of club, not separated from stalk by a 

membrane; protective membranes 

extending beyond sucker-bearing 

face along stalk, dorsal membrane 

forming a deep cleft at junction with 

stalk. Cuttlebone dorsal surface 

pinkish; flat medially and laterally; 

median rib present, distinct, sides 

approximately parallel; lateral ribs 

distinct; spine with ventral keel; sulcus 

shallow, narrow; anterior striae 

inverted U-shape; inner cone limbs 

of uniform width, thickened 

posteriorly forming a rounded ridge; tentacular club 

dorsal view 

ventral view 

outer cone narrow anteriorly, 

cuttlebone 

broadens posteriorly, lateral limbs 

wide. Colour: pale pinkish brown. 

(after Iredale, 1926) 

Size: Maximum mantle length about 150 mm. 

Habitat, biology, and fisheries: At depths from 83 to 184 m. Species taken as bycatch of prawn and mixed 

species trawl fisheries. 

Distribution: Northern Australia. 

? 

?

Sepiidae 747 

Sepia papuensis Hoyle, 1885 


Sepia galei Meyer, 1909; S. prionota Voss, 1962; 

Solitosepia submestus Iredale, 1926; S. occidua 

Cotton, 1929; S. genista Iredale, 1954; S. lana 

Iredale, 1954 / None. 

FAO names: En - Papuan cuttlefish. 

Diagnostic characters: Papillae on head, 

mantle and arms. Dorsal mantle with a row of 

elongated tubercles along silvery stripe at base 

of fins, or dorsal and ventral mantle covered with 

numerous small papillae. Head with numerous 

scattered small papillae and 2 pairs of large 

papillae located over each eye and 1 on each 

eyelid. Tubercles present on dorsal surface of 

arms I to III. Arm protective membranes (both 

sexes) wide, well developed. Male arms I to III 

with tetraserial suckers proximally, biserial 

suckers at distal tips; male arms IV with 

tetraserial suckers. Female arms all tetraserial. 

Hectocotylus absent. Tentacular club 

sucker-bearing face flattened; with 5 or 6 

suckers in transverse rows, differing 

markedly in size; swimming keel extending well 

beyond club; dorsal and ventral protective 

membranes not fused in small specimens, fused 

at base of club in large specimens, extending 

beyond sucker-bearing face along stalk; dorsal 

protective membrane much longer than 

10 mm 

ventral view 

cuttlebone 


(subadult) 


ventral membrane, dorsal membrane forming deep cleft at junction with stalk. Cuttlebone rounded 

anteriorly and posteriorly; median rib distinct, broadens anteriorly; spine with ventral keel; striated 

zone and last loculus concave; sulcus shallow, wide; anterior striae inverted U-shape; inner cone limbs 

broaden posteriorly, not thickened; outer cone narrow anteriorly, broadens posteriorly. Colour: dorsal 

mantle pale brownish with white blotches or spots; paired dorsal eye spots present. 


Habitat, biology, and fisheries: A shelf species found on silt, sand or mud bottoms, at depths from 17 to 

155 m. Species taken as bycatch of prawn and mixed species trawl fisheries. In the Gulf of Carpentaria, 

Australia, probably forms part of bycatch of Taiwanese trawl fisheries in the region. 

Distribution: Philippines, 

Indonesia, Arafura and Coral 

seas, northern Australia.


Sepia pharaonis Ehrenberg, 1831 

Frequent synonyms / misidentifications: Crumenasepia 

hulliana Iredale, 1926;C. ursulae Adam, 1939;Sepia rouxii 

Férussac and d’Orbigny, 1841; S. formosana Berry, 1912; 

S. tigris Sasaki, 1929 / None. 

FAO names: En - Pharaoh cuttlefish; Fr - Seiche 

pharaon; Sp - Sepia faraónica. 

Diagnostic characters: Mantle with a 

row of elongated tubercles along 

silvery stripe at base of fins, or 

covered with numerous small 

papillae. Head tubercles absent. Left 

ventral arm (IV) of males hectocotylized: 

10 rows of normal suckers 

proximally, followed by 6 rows of 

reduced suckers, then suckers 

normal to arm tip; reduction marked, 

2 dorsal series, smaller than 2 

ventral serries, oral surface wide, 

fleshy, with transversely grooved 

ridges. Tentacular club long; sucker- 

bearing face flattened; with 8suckers 

in transverse rows, differing 

markedly in size (5 or 6 median 

suckers enlarged); dorsal and 

ventral view 

cuttlebone 

dorsal view 

ventral protective membranes not fused at base of club, joined to stalk at base of club, 

not separated from stalk by a membrane. Buccal membrane with few, minute suckers. 

Cuttlebone median rib present, distinct, wider anteriorly; lateral ribs indistinct; sulcus 

extending entire length, flanked by rounded ribs; anterior striae inverted U-shape; 

inner cone limbs thickened posteriorly, form a shiny swelling; outer cone narrow 

anteriorly, broadens posteriorly; spine without keel. Colour: head with a vivid, 

transverse striped pattern; dorsal mantle pale brownish with white blotches or spots, 

or pale brown with transverse saddle mark (in females), or with vivid transverse striped 

pattern (especially in males); fins with pale reflective line along base (sometimes 

broken), line is blue in females; arms vivid with transverse stripes. 

Size: Maximum mantle length 420 mm; maximum weight 5 kg. In the Gulf of 

Carpentaria (Australia), the largest male and female collected in 1990 and 1991 

surveys were 192 mm and 173 mm mantle length, respectively. In the Gulf of Thailand, 

the maximum size of animals caught is for males 350 mm mantle length, and females 

300 mm mantle length. 

Habitat,biology,and fisheries: A neritic demersal species, at depths from 10 to 110 m. 

In the Gulf of Thailand and the Andaman Seas, found from the coastal shallows to a 

tentacular 

club 


depth of 100 m, with most caught between 10 and 40 m. In the Gulf of Thailand, spawning occurs all year 

round, with peak months of January-February and July-September. Males mature at 13.7 mm mantle 

length, females at 142 mm mantle length. Supports industrial or artisanal fisheries throughout its range. 

With Sepia esculenta, the most abundant cuttlefish species caught in the Philippines. Also important to the 

commercial squid fishery of Thailand. Contributes about 90% of the cuttlefish catch off Australia by Chinese 

pair trawlers. This amounted to 

some 1 000 t in 1979. Taken by 

domestic fisheries as bycatch of 

prawn and mixed species trawl 

fisheries. The flesh is thick, tender 

and excellent for human consumption. 

Distribution: South and East 

China seas, Indonesia, Malaysia, 

Philippines, northern Australia, Gulf 

of Suez, Zanzibar, Madagascar to 

the Arabian Sea. 

10 mm

Sepiidae 749 

Sepia plangon Gray, 1849 

Frequent synonyms / misidentifications: Solitosepia plangon 

adhaesa Iredale, 1926 / None. 

FAO names: En - Striking cuttlefish. 

Diagnostic characters: Left ventral arm (IV) of males hectocotylized: 

5 rows of normal suckers proximally, followed by 5 

rows of reduced suckers then remainder normal to arm tip; 

reduction slight, suckers equal sized across series; oral surface 

as for remaining arms, not wide, fleshy. Tentacular club suckerbearing 

face flattened; with 5 suckers in transverse rows, differing 

markedly in size; swimming keel extends well beyond suckerbearing 

face; dorsal and ventral protective membranes fused 

at base of club; joined to stalk at base of club, not separated from 

stalk by a membrane; protective membranes extend beyond 

sucker-bearing face along stalk, dorsal membrane forming deep 

cleft at junction with stalk. Cuttlebone blunt-pointed anteriorly; 

rounded posteriorly; dorsal median rib distinct, sides approximately 

parallel; lateral ribs indistinct; spine present, with ventral keel; 

striated zone concave, last loculus flat; sulcus extending length of 

cuttlebone, deep, narrow, flanked by rounded ribs (giving striae 

a wavy appearance); anterior striae inverted V-shape; inner cone 

limbs of uniform width; outer cone narrow anteriorly, broadens 

posteriorly. Colour: deep purplish brown. 


Habitat, biology, and fisheries: Intertidal to a depth of 83 m. Taken 

as bycatch of prawn and mixed species trawl fisheries. 

Distribution: Eastern Australia. 

? ? 

10 mm 

ventral view 

cuttlebone 




Sepiidae Sepia recurvirostra Steenstrup, 1875 


singaporensis Pfeffer, 1884 / S. pagenstecheri Pfeffer, 

1884. 

FAO names: En - Curvespine cuttlefish; 

Fr - Seiche hameçon; Sp - Sepia 

ganchuda. 

Diagnostic characters: Arm suckers 

(both sexes): arms I to III with tetraserial 

suckers proximally, biserial suckers at 

distal tips; arms IV with tetraserial 

suckers. Hectocotylus present: suckers 

normal proximally, then reduced, followed 

by normal suckers to arm tip; reduction 

marked, rows evenly spaced on arm; 2 

dorsal series, smaller than 2 ventral 

series. Tentacular club sucker-bearing 

face flattened; with 5 or 6 suckers in 

transverse rows, differing markedly in 

size, 5 or 6 median suckers enlarged; 

swimming keel extends slightly beyond 

sucker-bearing face; dorsal and ventral 

protective membranes not fused at 

base of club; dorsal protective membrane 

forming deep cleft at junction with 

stalk. Cuttlebone outline oblong; very 

angular, V-shaped anteriorly; rounded 

ventral view 

cuttlebone 

posteriorly; dorsal median and lateral ribs present; spine 

present, curving ventrally (giving species its name); last 

loculus concave; sulcus extending along striated zone only; 

sulcus shallow, narrow; anterior striae inverted U-shape; 

inner cone limbs broadening posteriorly, thickened 

posteriorly forming a rough chitinous callus; outer cone 

narrow anteriorly, broadens posteriorly. Colour: dorsal 

mantle pale with transverse stripes; fins with pale 

opalescent blue reflective line along base. 

Size: Maximum mantle length 170 mm; maximum weight 0.4 kg. In the Gulf of 

Thailand, most animals caught are between 40 and 130 mm mantle length, with 

a maximum size of 170 mm mantle length. 

Habitat, biology, and fisheries: A demersal species inhabiting the continental 

shelf to depths of around 160 m. In the Gulf of Thailand and Andaman Seas, it is 

found from depths of 10 to 100 m, with peak of occurrence at 20 to 40 m. In this 

region, spawning occurs all year, with peak times in November-February and 

July-September. A commercial species in Gulf of Thailand, South and East China 

Seas, Japan; and Hong Kong, China. In Thailand, most are caught using otter 

trawl, with smaller catches made using pair trawl, squid light-lures, traps and push 

nets, with bottom otter and 

pair trawls used offshore, and 

push nets and lift nets used 

in inshore and coastal waters. 

A few are caught using purse 

seine and hook-and-line. 

Distribution: Andaman Sea 

to Celebes Sea and from Java 

Sea to the Yellow Sea. 

dorsal view 

10 mm 

tentacular club

Sepiidae 751 

Sepia rex (Iredale, 1926) 


Decorisepia rex Iredale, 1926; D. 

cottlesloensis Cotton, 1929; D. jaenschi 

Cotton, 1931 / Sepia madokai Adam, 1939. 

FAO names: En - King cuttlefish. 

Diagnostic characters: Arms in both sexes 

with tetraserial suckers. Left ventral arm (IV) 

of males hectocotylized: 6 to 8 rows of 

normal suckers proximally, followed by 9 

or 10 rows of reduced suckers, then rest 

normal to arm tip; reduction marked, 2 

dorsal and 2 ventral series widely spaced; 

suckers equal sized across series; oral 

surface wide, fleshy, with transversely 

grooved ridges and shallow median furrow. 

Tentacular club sucker-bearing face 

flattened; with 10 to 12 similar-sized, 

minute suckers in transverse rows; 

swimming keel extends well beyond club; 

dorsal and ventral protective membranes 

not fused at base of club, joined to stalk at 

base of club and terminating at posterior end 

of sucker-bearing face; dorsal protective 

membrane forming deep cleft at junction 

with stalk. Cuttlebone outline oblong, 

narrowed anteriorly, tending to diamondshaped, 

maximum width in middle of 

cuttlebone; dorsal surface pinkish; median 

dorsal view 

ventral view 

cuttlebone 

(after Iredale, 1926) 

rib present, distinct; spine without keel(s); ventral sulcus extending entire length of cuttlebone, shallow, 

narrow; anterior striae inverted U-shape; inner cone limbs of uniform width, thickened posteriorly 

forming a rounded ridge; outer cone narrow anteriorly, broadens posteriorly. Colour: pale pinkish purple. 


Habitat, biology, and fisheries: At depths from 55 to 400 m. Taken as bycatch of prawn and mixed species 

trawl fisheries. 

Distribution: Southern Australia. 

Remarks: Sepia rex has been 

confused with S. madokai. It 

differs in having all reduced 

suckers of the hectocotylus 

similar in size, while in S. 

madokai, the 2 dorsal series 

are much smaller than the 2 

ventral series. The club 

suckers in S. madokai (8 

suckers in transverse rows), 

are fewer than those (10 to 

12) seen in S. rex. The 

cuttlebone in this species has 

distinct median ribs and the 

inner cone limbs are of 

uniform width, while in S. 

madokai it has faint median 

ribs, and the inner cone limbs 

broaden posteriorly. S. rex 

may be synonymous with S. 

hedleyi Berry, 1918.


Sepia rozella (Iredale, 1926) 

Frequent synonyms / misidentifications: Solitosepia rozella 

Iredale, 1926; Solitosepia rozella peregrina / None. 

FAO names: En - Rosecone cuttlefish. 

Diagnostic characters: Arm suckers tetraserial in both sexes. 

Hectocotylus present, left ventral arms (IV) modified: suckers 

reduced proximally for 2/5 of arm length; oral surface wide, 

fleshy, with transversely grooved ridges and shallow median 

furrow, thicker than corresponding portion on right ventral arm. 

Tentacular club sucker-bearing face flattened, with 8 suckers in 

transverse rows, differing markedly in size (third series from 

dorsal margin larger than other suckers); swimming keel extending 

well beyond sucker-bearing face; dorsal and ventral protective 

membranes fused at base of club, separated from stalk by 

membrane; dorsal protective membrane forms a deep cleft at 

junction with stalk. Cuttlebone outline oblong (widest in middle); 

blunt-pointed anteriorly and posteriorly; dorsal surface pinkish; 

median rib present, distinct, wider anteriorly; lateral ribs indistinct; 

spine with ventral keel; sulcus deep, wide, flanked by 2 

strongly convex rounded ribs; anterior striae inverted V-shape; 

inner cone limbs broaden posteriorly (rose-coloured), thickened 

forming a slightly raised, rounded ledge from outer margin of inner 

cone; outer cone narrow anteriorly, broadens posteriorly. Colour: 

pinkish purple. 


Habitat, biology, and fisheries: At depths from 27 to 183 m. 

Species taken as bycatch of prawn and mixed species trawl 

fisheries. 

10 mm 

Distribution: Eastern Australia. ventral view 

cuttlebone

Sepiidae 753 

Sepia smithi Hoyle, 1885 


Acanthosepion pageorum Iredale, 1954 / 

Sepia whitleyana (Iredale, 1926). 

FAO names: En - Smiths cuttlefish. 

Diagnostic characters: Arms in both sexes 

with tetraserial suckers. Left ventral arm (IV) 

of males hectocotylized: 8rowsofnormal 

suckers proximally, followed by 8 rows of 

reduced suckers, then remainder normal to 

tip of arm; reduction marked, 2 dorsal and 2 

ventral series widely spaced; 2 ventral series, 

smaller than dorsal series; oral surface 

wide, fleshy, with transversely grooved ridges 

and a shallow median furrow. Tentacular club 

sucker-bearing face convex; with 20 

similar-sized, minute suckers in transverse 

rows; swimming keel extends beyond 

sucker-bearing face of club; dorsal and ventral 

protective membranes not fused at base of 

club, joined to stalk at base of club, not 

separated from stalk by a membrane. Cuttlebone 

outline oblong, V-shaped anteriorly, 

rounded posteriorly; dorsal surface convex 

medially, flattened laterally, granulose; 

median rib faint, broadens anteriorly; lateral 

ribs indistinct; spine present, curving upward; 

keel absent; striated zone concave; sulcus 

deep, wide; anterior striae inverted U-shape; inner cone limbs broaden posteriorly, thickened 

posteriorly forming a raised, flattened ledge from outer margin of inner cone; outer cone narrow 

anteriorly, broadens posteriorly. Colour: pale pinkish purple. 


Habitat, biology, and fisheries: At depths of 33 to 138 m. Taken as bycatch of prawn and mixed species 

trawl fisheries. 

Distribution: Northern Australia, Timor, Arafura, and Coral Seas. 

Remarks: Often confused with 

Sepia whitleyana. In S. 

whitleyana, both ventral arms 

are modified in males, with all 

reduced suckers equal in size, 

rather than differing in size as in 

S. smithi. The swimming keel 

does not extend beyond the 

sucker-bearing face of the club 

in S. whitleyana. The inner 

cone limbs form a small ridge 

but do not form a distinct 

ledge posteriorly in S. 

whitleyana. 

? 

10 mm 

ventral view 

cuttlebone 

? 

tentacular club


Sepia stellifera Homenko and Khromov, 1984 

Frequent synonyms / misidentifications: None / Sepia brevimana Steenstrup, 1875. 

FAO names: En - Starry cuttlefish. 

ventral view 

cuttlebone 

hectocotylus 

(after Homenko and Khromov, 1984) 


dorsal view 

(after Homenko and Khromov, 1984) 

Diagnostic characters: Tentacular club with 10 similar-sized suckers in transverse rows. Cuttlebone 

outline oval; very angular, V-shaped anteriorly; spine present, long; keels present dorsally and ventrally; 

inner cone forms a raised ridge posteriorly; sulcus extending along striated zone only; deep, wide, flanked 

by rounded ribs. 


Habitat, biology, and fisheries: At depths to 200 m. Fished commercially in India and the Gulf of Thailand 

and may figure in statistical data with S. brevimana. 

Distribution: Arabian Sea, 

Bay of Bengal, Andaman Sea, 

Gulf of Thailand. 

Remarks: This species is 

? 

similar to Sepia brevimana. It 

differs in having 10 club 

suckers in transverse rows, ? 

rather than 6 to 8 in S. 

brevimana. Also, the sulcus is 

deep and wide, rather than 

narrow and shallow, as seen in 

S. brevimana.

Sepiidae 755 

Sepia whitleyana (Iredale, 1926) 

Frequent synonyms / misidentifications: Acanthosepion 

whitleyanum Iredale, 1926 / Sepia smithi Hoyle, 1885. 

FAO names: En - Whitleys cuttlefish. 

Diagnostic characters: Arms in both sexes with tetraserial 

suckers. Hectocotylus present, both male ventral arms (IV) 

modified: left ventral arm with 7 or 8 rows of normal suckers 

proximally; 5 or 6 rows reduced suckers medially, then 

remainder normal to arm tip; right ventral arm with 4 or 5 rows of 

minute suckers distally, remaining suckers normal; reduction 

marked, left arm IV with 2 dorsal and 2 ventral series widely 

spaced; suckers equal size across series; oral surface wide, 

fleshy, with transversely grooved ridges and shallow median furrow. 

Tentacular club sucker-bearing face flattened; with 20 

similar-sized, minute suckers in transverse rows; swimming 

keel does not extend beyond sucker-bearing faced of the club; 

dorsal and ventral protective membranes not fused at base of club, 

joined to stalk at base of club, not separated from stalk by a 

membrane and terminating at posterior end of sucker-bearing face. 

Cuttlebone outline oblong; texture uniformly pustulose; median rib 

faint; lateral ribs indistinct; spine without keel; striated zone deeply 

concave; last loculus slightly convex; sulcus deep, wide; anterior 

striae inverted U-shape; inner cone limbs broadening posteriorly, 

thickened forming a rounded ridge; outer cone narrow anteriorly, 

wider posteriorly. Colour: pale pinkish purple. 

Size: Cuttlebone length of holotype 168 mm (excluding spine). 

Habitat, biology, and fisheries: At depths of 23 to 160 m. Taken 

as bycatch of prawn and mixed-species trawl fisheries. 

Distribution: Eastern Australia 

from the Gulf of Carpentaria to 

New South Wales. 

Remarks: Often confused with 

Sepia smithi, but differs in 

having both ventral arms 

modified in males (only the left 

ventral arm hectocotylized in S. 

smithi). The swimming keel 

extends beyond the suckerbearing 

face of the club in S. 

smithi, but is equal to the club 

length in S. whitleyana. The 

distinct inner cone limbs do not 

form a ledge posteriorly in S. 

whitleyana but form a narrow 

rim. 

? 

? 

? 

ventral view 

cuttlebone


Sepiella inermis Férussac and dOrbigny, 1835 


(Sepiella) microcheirus Gray, 1849; Sepia affinis 

Eydoux and Souleyet, 1852; Sepiella mandroni 

Rochebrune, 1884 / None. 

FAO names: En - Spineless cuttlefish; Fr -Sepia 

inerme; Sp - Sepia inerme. 

dorsal view lateral view 

cuttlebone 

(after Adam, 1939) 

ventral view 

dorsal view 

Diagnostic characters: Arms in both sexes with tetraserial 

suckers. Hectocotylus present, suckers reduced and widely 

spaced on proximal half of club. Tentacular club with 12 to 24 

equal-sized suckers in transverse rows; swimming keel 

shorter than sucker-bearing face of club; dorsal and ventral 

protective membranes not fused at base of club, extending 

proximally along stalk as low ridges. Cuttlebone width 

approximately 33 to 43% the length; spine absent; sulcus 

extending entire length of cuttlebone; anterior striae m-shape; 

outer cone chitinous, flared, spoon-shaped, indented at posterior 

end of striated zone. Colour: dorsal mantle greyish brown with 

8 or 9 reddish patches along base of each fin. 


hectocotylus 


Size: Maximum mantle length 125 mm. In the Gulf of Thailand, most animals caught range between 20 

and 80 mm mantle length, with the maximum size recorded at 105 mm mantle length. 

Habitat, biology, and fisheries: A demersal, shallow-water species, depth range to 40 m. Probably a species 

complex. Highly abundant in the Gulf of Thailand, and Andaman Sea, it occurs from the shallows to a depth of 

40 m, with most caught between 10 and 20 m. Sepiella inermis is an object of several small fisheries in the waters 

of Indochina. It is important to the commercial cuttlefish fishery in Thailand. Cuttlefish in this region are usually 

caught using squid light lures, traps, push nets, purse seines and hook-and-line. This species has been reared 

successfully in culture 

experiments conducted by the 

Thai Coastal Culture Division, 

Department of Fisheries and 

there is hope to convert this 

work to a commercial basis. 

Distribution: Mouth of 

Zambezi River (Mozambique), 

Indian Ocean, southern Red 

Sea, Gulf of Aden to Andaman 

Sea, eastern Indonesia, the 

? 

Gulf of Tonkin and the 

southern South China Sea.

Sepiidae 757 

Metasepia pfefferi (Hoyle, 1885) 

En - Pfeffers flamboyant cuttlefish. 

Maximum mantle length 80 mm. Mantle very broad. 

Dorsal mantle with 3 pairs of large, flat, flap-like 

papillae; head with papillae over eyes. Arms broad, 

blade-like; male and female arms I reduced. Fins 

broad, transparent positioned dorsolaterally. Left ventral 

arm (IV) of males hectocotylized. Tentacular club short, 

sucker- bearing face flattened; with 5 or 6 suckers in 

transverse rows, differing markedly in size, 4 median 

suckers greatly enlarged; dorsal and ventral protective 

membranes not fused at base of club; swimming keel 

twice as long as sucker-bearing face. Cuttlebone much 

shorter than mantle; located in anterior 2/3 to 3/4; 

outline diamond-shaped; blunt-pointed anteriorly and 

posteriorly; median rib absent; spine absent; chitin 

covers entire dorsal surface; striated zone concave; 

last loculus convex (thick in anterior third); sulcus deep, 

wide; anterior striae inverted V-shape; inner cone limbs 

of uniform width; outer cone narrow throughout. 

Colour: a pair of prominent white bars on 

mantle; striking flowery body pattern; dorsal 

surface of cuttlebone yellowish. From depths of 

3to86m.NorthernAustralia. 

Sepia bandensis Adam, 1939 

En - Stumpy spined cuttlefish. 

Maximum mantle length 48 mm in males, 50 mm in 

females. Tentacular club short, with 5 suckers in 

transverse rows, central 3 suckers enlarged; dorsal and 

ventral protective membranes fused at base of club. 

Cuttlebone outline broad, oval; rounded anteriorly and 

posteriorly. Dorsal surface evenly convex with reticulate 

sculpture; median and lateral ribs absent; spine reduced 

to tiny tubercle; sulcus shallow, narrow; anterior striae 

inverted U-shape; inner cone limbs broadening 

posteriorly, not thickened. Colour: dark brown with 

scattered yellowish spots. Coastal shallow waters. 

Importance to fisheries unknown. Java, eastern 

Indonesia, Philippines, New Guinea, and Marshall 

Islands. 

? 

? 

10 mm 

lateral view 

10 mm 

dorsal view 


10 mm 10 mm 

10 mm 


cuttlebone 


ventral view 

10 mm 

dorsal view 

cuttlebone 

10 mm 

ventral view 

tentacular 

club


Sepia bartletti (Iredale, 1954) 

En - Bartletts cuttlefish. 

Known only from the cuttlebone. Status uncertain. 

Cuttlebone length 73 mm. Dorsal surface of cuttlebone 

without sculpturing; spine absent; a shallow, wide 

sulcus present on the last loculus only, absent from 

striated zone; inner cone limbs extending to last loculus. 

Fisheries importance unknown. Known only from the 

Misima and the Conflict Group of islands, Louisiade 

Archipelago (southeast of Papua New Guinea). 

Sepia cottoni Adam, 1979 

dorsal view 

En - Cottons cuttlefish. 

Maximum mantle length 65 mm. Male arms all elongate, twice as long as in 

females, tips of arms I, II, andIV thread-like; arms III with protective 

membrane in middle part widened and thickened with transverse 

ridges; ridges alternate, uniting inner margin of one membrane and outer 

margin of opposite membrane, each ridge with 2 small suckers. Male arms 

proximally with a few normal-sized biserial suckers, followed by 7 or 8 

tetraserial rows of large suckers; distally, suckers biserial, reduced, 

disappearing on thread-like tips of arms I, II, and IV. Female arms with 

biserial suckers at the base, tetraserial suckers medially, biserial suckers at 

distal tips. Tentacular club with 5 suckers in transverse rows; suckers 

differing markedly in size, about 6 in the middle of club twice as wide as 

others. Dorsal and ventral protective membranes not fused at base of club. 

Cuttlebone lanceolate; spine present, keel(s) absent; sulcus, shallow, 

narrow; striae slightly convex-straight; inner cone limbs thickened posteriorly 

forming a rounded ridge; outer cone limbs expanded forming 2 short 

wings. At depths from 83 to 164 m. Importance to fisheries unknown. 

Western Australia and the Gulf of Carpentaria; possibly Viet Nam 

(unconfirmed records). One cuttlebone collected from North Stradbroke 

Island Queensland, Australia implying a northern Australian distribution. 

? 

? 

? 

cuttlebone 

ventral view 

ventral view 

cuttlebone

Sepiidae 759 

Sepia cultrata Hoyle, 1885 

En - Knifebone cuttlefish. 

Left ventral arm of male hectocotylized: suckers normal 

proximally, followed by reduced suckers, then 

remaining suckers normal to tip of arm; reduction 

marked, 2 dorsal and 2 ventral series widely spaced, 2 

dorsal series smaller than 2 ventral series. 

Tentacular club with 5 or 6 suckers in transverse rows; 

suckers approximately of similar size, small; swimming 

keel extending well beyond club sucker-bearing face; 

dorsal and ventral protective membranes not fused at 

base of club. Cuttlebone triangular anteriorly; median rib 

distinct; spine present; sulcus indistinct; striae slightly 

convex-straight; inner cone limbs of uniform width, very 

narrow, thickened posteriorly forming a rounded ridge. 

Outer shelf and upper bathyl species, at depths from 

132 to 803 m (majority of catches from 300 to 500 m). 

Species taken as bycatch of prawn and mixed species 

trawl fisheries. Southern Australia. 

Sepia madokai Adam, 1939 

En - Madokais cuttlefish; Fr - Seiche madokai; Sp - Sepia 

madokai. 

Maximum mantle length 80 mm. Left ventral arm of 

males hectocotylized: suckers normal proximally, 

followed by 10 rows of reduced suckers, then rest 

normal to arm tip; 2 dorsal series of reduced 

suckers smaller than 2 ventral series. Tentacular 

club sucker-bearing face flattened, with 8 equal-sized, 

small suckers in transverse rows. Cuttlebone dorsal 

surface pinkish; spine present without keel(s); anterior 

striae inverted U-shape; inner cone limbs 

approximately of uniform width, very slender, 

thickened, forming a rounded ridge posteriorly. Colour: 

dorsal mantle without distinct markings, brownish with 

some whitish blotches, or spots. A demersal species, 

most common in bays. Commonly confused with Sepia 

acuminata and S. rex. A minor object of fisheries in 

Japan and China. East and South China Sea. 

? 

10 mm 

dorsal view 

cuttlebone 

10 mm 

ventral view 

ventral view 

cuttlebone tentacular club 

(after Okutani et al., 1987)


Sepia mestus Gray, 1849 

En - Reaper cuttlefish; Fr - Seiche moisson; Sp - Sepia segadora. 

Maximum mantle length 140 mm. Arms all of similar length in both 

sexes, with tetraserial suckers. Hectocotylus absent. Tentacular club 

sucker-bearing face flattened; with 6 small suckers in transverse 

rows, those in middle of third dorso-longitudinal row slightly enlarged; 

swimming keel extending well beyond sucker-bearing face of club; 

dorsal and ventral protective membranes fused at base of club, 

joined to stalk at base of club, not separated from stalk by a membrane. 

Cuttlebone oval, rounded anteriorly and posteriorly; dorsal median 

rib indistinct; spine with ventral keel; sulcus shallow, narrow; anterior 

striae inverted U-shape; inner cone limbs broaden posteriorly, not 

thickened. A neritic demersal species, at depths to 146 m. Cuttlebones 

have been found in eastern Australia. Whole animals have been 

collected only from the Sydney region. Also reported from northern 

China and Viet Nam though it is uncertain whether either of these 

records refer to species conspecific with S. mestus from Australia. 

? 

? 

Sepia mira (Cotton, 1932) 

? 

En - Little cuttlefish. 

Cuttlebone length 55 mm. Cuttlebone outline 

lanceolate, narrowed between middle and 

posterior third, widened near posterior end; 

dorsal surface white, evenly convex; median 

and lateral ribs absent; spine present, straight, 

without keels; striated zone and last loculus 

convex; sulcus absent; striae slightly 

convex-straight; inner cone limbs of uniform 

width; thickened posteriorly forming a rounded, 

shiny ridge; outer cone narrow anteriorly, wider 

posteriorly. Until recently, known only from 

cuttlebones found in Queensland, at Lizard 

Island and North-West Islet, as well as Trial 

Bay (New South Wales). On the basis of 

specimens trawled betwen 20 and 72 m off 

northern New South Wales, Sepia mira has 

now been fully described (see Reid, in press). 

? 


cuttlebone 

(after Cotton, 1952) 

10 mm 

ventral view 

cuttlebone 

ventral view 



Sepiidae 761 

Sepia sulcata Hoyle, 1885 

En - Grooved cuttlefish. 

Maximum mantle length 68 mm. Mantle broadest at 

mantle opening, blunt-pointed posteriorly. Suckers 

biserial proximally, tetraserial distally on arms I to IV 

in both sexes. Left ventral arm of males hectocotylized: 14 

rows of reduced suckers proximally; reduction marked, 

2 dorsal and 2 ventral series widely spaced; oral surface 

wide, fleshy, with transversely grooved ridges and deep 

median furrow. Tentacular club with 5 to 7 similar-sized, 

minute suckers in transverse rows; swimming keel 

extending well beyond club sucker-bearing face; dorsal 

and ventral protective membranes not fused at base of 

club, dorsal membrane much wider than ventral 

membrane (as in Sepia brevimana). Cuttlebone outline 

broad, oval; rounded anteriorly, blunt-pointed posteriorly; 

dorsal median rib and lateral ribs present, distinct; sulcus 

shallow, narrow, flanked by rounded ribs; inner cone 

raised to form a thin, flat ledge posteriorly, outer cone 

recurved ventral to spine. Ki Island, Arafura Sea to 

Australia. 

? 

Sepia vietnamica Khromov, 1987 

En - Viet Nam cuttlefish. 

Maximum mantle length 56 mm. Body papillae present, 

mantle with row of elongated tubercles along silvery stripe 

at base of fins. Male and female arms all of similar 

length. In both sexes arms I to III with tetraserial 

suckers proximally, biserial suckers at distal tips 

of arm. Left ventral arm of males hectocotylized: 8 to 11 

rows of normal suckers proximally, suckers reduced 

medially, then rest normal to distal tip of arm; reduction 

marked, 2 dorsal and 2 ventral series widely spaced; 2 

dorsal series smaller than 2 ventral series;oralsurface 

with deep median furrow. Tentacular club sucker-bearing 

face flattened, with 5 or 6 suckers in transverse rows, 

differing only slightly in size; dorsal and ventral 

protective membranes not fused at base of club. 

Cuttlebone outline lanceolate; convex ventrally; sulcus 

deep, narrow; anterior striae M-shape; inner cone limbs 

of uniform width; outer cone limbs expanded forming 

2 short wings. Colour: head with 2 small, 

crescent-shaped orange spots near dorsal projection 

of mantle margin and above eyes; dorsal mantle dark 

brown; fins pale, with transverse rows of large 

wine-coloured spots; arms I to III with spots on aboral 

surface same as for mantle (sometimes visible, males 

only); spots may not be visible in preserved animals. At 

depths to 105 m. Fisheries importance unknown. 

Northwestern South China Sea, off Viet Nam. 

ventral view 

cuttlebone 

? 

10 mm 

ventral view 

cuttlebone 

(after Khromov, 1987) 


tentacular 

club


Sepia vossi Khromov, 1996 

En - Voss cuttlefish. 

Maximum mantle length 100 mm. Arms in both sexes 

with tetraserial suckers. Hectocotylus present: 3rows 

of normal suckers proximally, followed by reduced 

suckers, then rest normal to arm tip; reduction 

pronounced, 2 dorsal and 2 ventral series widely 

spaced, oral surface wide, fleshy, with transversely 

grooved ridges. Tentacular club with 8 suckers in 

transverse rows; suckers differing markedly in size: 

3 to 5 suckers in middle of third longitudinal row 

extremely enlarged; swimming keel extending slightly 

beyond club; dorsal and ventral protective 

membranes not fused at base of club. Cuttlebone 

blunt-pointed anteriorly and posteriorly; spine with 

keels; sulcus shallow, narrow; inner cone limbs of 

uniform width, form a ledge posteriorly, outer cone 

recurved, cup-like. Colour: dorsal mantle light brown, 

with dark brown transverse stripes. A neritic demersal 

species, at depths to 200 m. Until recently, this 

Indo-Pacific cuttlefish was considered to be 

conspecific with Sepia omani Adam and Rees, 1966. 

It differs from S. omani in the cup-like formation of the 

outer cone and inner cone ledge. Known from 

Pakistan, western India, and Hong Kong, China to 

southern Viet Nam. 

Sepiella ocellata Pfeffer, 1884 

En - Spotty cuttlefish. 

Maximum mantle length 50 mm. Known 

only from a single male specimen. 

Tentacular club with 8 to 10 suckers in 

transverse rows. Cuttlebone narrow, 

width 20 to 25% of length, oval, of uniform 

width along its length, sides approximately 

parallel. Colour: 6 or 7 winecoloured, 

similar-sized spots along fins, 

close to fin margin. Very similar to 

Sepiella ornata from the west coast of 

Africa, which differs in having a broader 

cuttlebone and 10 to 14 arm suckers in 

transverse rows. The status of this 

species is questionable. Fisheries 

importance unknown. Known only from 

the type locality, Java. 

? 

? 

ventral view 

cuttlebone 

dorsal view ventral view 

cuttlebone 

lateral view 


tentacular 

club 

hectocotylus 

(after Voss and Williamson, 1971) 

dorsal view 

(after Adam, 1939)

Sepiidae 763 

Sepiella weberi Adam, 1939 

En - Webs cuttlefish. 

Left ventral arm of males hectocotylized: 10 to 12 rows of 

reduced suckers proximally; reduction marked, 2 dorsal and 

2 ventral series widely spaced; oral surface wide, fleshy, with 

transversely grooved ridges and a deep median furrow. 

Tentacular club sucker-bearing face convex; with 7 to10 

suckers in transverse rows; dorsal and ventral protective 

membranes not fused at base of club, joined to stalk at base of 

club, not separated from stalk by a membrane; swimming keel 

equal in length to sucker-bearing face of club. Cuttlebone 

median rib present, faint, wider anteriorly; lateral ribs absent; 

sulcus shallow, narrow; striae wavy; inner cone limbs of uniform 

width; outer cone chitinous, spoon-shaped, expanded. Colour: 

5 or 6 large wine-coloured spots at base of each fin. Depth range 

1 to 88 m. Eastern Indonesia and northern Australia; 

unconfirmed records from Viet Nam. In contrast to Sepiella 

weberi, the ventral side of the cuttlebone is strongly convex 

medially in all other Sepiella species. 

dorsal view 

lateral view ventral view 

cuttlebone 


schematic presentation 

in dorsal view 

? 

wine-coloured 

spots 

? 

10 mm 

dorsal view 





Loliginidae LOLIGINIDAE 

Inshore squids, pencil squids 


Diagnostic characters: Shape variable from short and stout to long and slender. Fins terminal or 

marginal, but always reaching posterior end of body (present in Uroteuthis as very thin membranes 

around the “tail”). Funnel locking apparatus a simple, straight groove. Eyes covered with transparent 

membrane (cornea). Buccal connectives attached to ventral borders of arms (IV); 7 buccal lappets 

supplied with small suckers (can be readily lost after capture). Mouth surrounded by 10 appendages 

(8 arms, 2 tentacles). Two longitudinal rows of suckers on arms and 4 rows on tentacular clubs; hooks 

never present. Usually, left ventral arm (IV) hectocotylized in males; hectocotylus with suckers reduced 

in size or number, and/or modified into fleshy papillae or flaps, or lost completely. Colour: translucent to 

dense colour, bright scarlet (in some species with yellow and pink chromatophores) to dark brown (nearly 

black), darker dorsally, but highly variable depending on the behavioural situation and degree of expansion 

of chromatophores. 

eye covered by funnel 

ventral 

membrane groove 

row of 

suckers 

ventral view 

ventral (4 th ) 

pair of arms 

corneal 

membrane 

fin angle 

funnel 

funnel 

locking 

cartilage 

4 longitudinal 

rows of 

suckers 


tentacle 

mantle 

locking 

cartilage 

dorsal 

row of 

suckers 

pedicels 

gladius 

(internal skeletal mantle 

support) feather-shaped 

modified 

portion 

with 

reduced 

suckers 

arms I beak (jaws) 

arms IV 

(left arm hectocotylized in males) 

hectocotylized 

arm of males 

arm II 

diagram of oral surface of 

brachial crown and buccal area 

buccal 

membrane 

arm III 

buccal lappets 

often with 

suckers 

buccal 

connectives 

ventrally 

attached

Loliginidae 765 

Habitat, biology, and fisheries: Demersal or semipelagic inhabitants of coastal and continental shelf areas 

to a maximum depth of about 400 m. Several species are restricted to extremely shallow waters and some 

of these penetrate into brackish waters (Loliolus). Typically, they undertake diel movements, aggregating 

near the bottom during the day, but dispersing into the water column at night. Many species are positively 

phototactic, and hence often are captured with fishing techniques using light attraction. Some species 

undergo seasonal onshore-offshore migrations in response to temperature changes. The eggs (which 

range from about 2 mm diameter in Photololigo to about 10 mm diameter in Sepioteuthis) are encapsulated 

in gelatinous, finger-like strings and attached in clusters to various substrates. Hatchlings resemble the 

adults. Recent studies of growth using direct measurements from statoliths reveal that some tropical 

loliginids may live only for a few months. Inshore squids are predators on crustaceans and small fishes. 

Loliginids are the dominant component of the Western Central Pacific cephalopod catch, but separate 

statistics on total catches are not reported for this family. They are a significant component of major trawl 

fisheries in the Gulf of Thailand and South China Sea areas; they also are taken as bycatch in many 

nearshore trawl fisheries for crustaceans and demersal finfish. Numerous artisanal and subsistance 

fisheries take inshore squids either in multispecies catches or as prime target species; gear used includes 

purse seines, dip nets, lift nets, cast nets, encircling nets including fixed or tunnel nets in intertidal areas, 

baited and unbaited jigs sometimes trolled rather than used vertically, often used at night in association 

with light attraction. Loliginids are highly valued for human consumption. They are marketed either fresh, 

frozen, dried or processed into cleaned mantles (whole hoods, rings). Small squid are also used as bait 

for both commercial and recreational fisheries. 

Remarks: A recent study using modern biochemical genetic techniques has highlighted the incomplete 

state of knowledge of the taxonomy of the Indo-West Pacific loliginids. This is especially true for members 

of the genus Photololigo which includes the majority of the large commercially important species. The 

present set of diagnostic morphological characters (fin shape, sucker dentition, hectocotylus structure) 

does not always ensure a reliable identification of species. They do not take into account variability with 

growth and sex in some of these characters, particularly when these have been poorly and inadequately 

defined in the type descriptions (many from the mid 1800s). In many cases, type reference material is 

poorly preserved and had poor geographic locality information (e.g. Yokohama market for Loligo edulis, 

“Australian Seas (?S.E.)” for Loligo etheridgei. Generic level revisions have recently been undertaken for 

Sepioteuthis, Loliolus, and “Loligo”. However, additional studies at the species level over broad geographic 

areas are required to clarify the “species complexes” represented by what are currently known as 

“Photololigo edulis”, “P. chinensis“ and ”Sepioteuthis lessoniana“. Several poorly known and new unnamed 

species are referred to in the literature and ”seasonal forms" with different life history characteristics and of 

questionable taxonomic status and distribution are known (e.g. Sepioteuthis lessoniana around Okinawa). This 

poor state of taxonomic knowledge has been highlighted previously in workshops and published papers but 

little progress has been made.Current and future fisheries assessments of the loliginid resource and subsequent 

decisions concerning the management of the stocks are dependent on accurate identification of species. 

Therefore, there is an urgent need for a substantial cooperative 

region-wide taxonomic study of the genus Photololigo 

using classical morphology supported by modern techniques 

including allozyme electrophoresis and DNA analysis. 


Sepiidae (cuttlefishes): the loliginid genus Sepioteuthis 

shows some external similarities to cuttlefish but can be 

easily distinguished from them by the presence of a gladius 

in the dorsal mantle rather than a chalky cuttlebone with a 

posterior calcareous spine; the presence of the midline connection 

of the fin lobes posteriorly in Sepioteuthis and the 

possession of contractile rather than retractile tentacles only 

without pockets. 


The key and descriptions provided below are based on the 

existing, inadequate adult character sets and users should 

recognize that in some cases, they may not readily assist 

with identifying the animals at hand. It should also be noted 

calcareous 

spine 

Sepiidae Loliginidae (Sepioteuthis) 

that in some cases, known or suspected species complexes are referred to rather than single species 

(“Photololigo chinensis”, “Photololigo edulis” and “Sepioteuthis lessoniana“). Some nominal species 

described in the literature and listed below are not included in the key because detailed descriptions have 

yet to be published, they remain poorly characterized from the type descriptions or are described from limited 

material and their status as valid species remains questionable. These include:


Loligo reesi (Voss, 1962), a form that matures at a small size (less than 80 mm mantle length) known only 

from the Philippines which has long square-tipped arm sucker teeth, both ventral arms hectocotylized in 

males (left with papillae along more than 50% arm length and right with much reduced suckers on the distal 

half) and the the tips of both ventral arms in males devoid of suckers and papillae. The existing descriptions 

do not mention the presence of light organs; 

Loligo vossi Nesis, 1982 [= species A of Voss (1963)], reaching 140 mm mantle length and reported from 

around the Philippines and northern Indian Ocean perifery, with 15 to 20 triangular teeth on the large 

tentacular suckers, 2 to 4 on the distal edge much larger, sharp and hook-like in Philippine specimens. 

Only the left ventral arm is hectocotylized in males by the development of long fleshy papillae along 40 to 

50% of the arm. The existing descriptions do not mention the presence of light organs; 

Loligo n.sp. of Chotiyaputta (1993), a form that matures at a small size (less than 60 mm mantle length) 

known only from the Gulf of Thailand which has long square-tipped arm sucker teeth and sharp conical 

widely spaced subequal teeth on the large tentacular sucker rings. The existing incomplete description 

does not mention the presence of light organs; 

Photololigo spp. 1, 2, 3 of Yeatman and Benzie, 1994, are “species” which have been distinguished using 

allozyme electrophoresis. Morphological characteristics of species 1 and 2 from northern Australia (which 

mature at less than 120 mm mantle length) fall within the range of the “Photololigo edulis” complex with 

square tipped teeth on the arm sucker rings, sharp conical teeth on the large tentacular sucker rings and 

hectocotylization by development of long fleshy papillae along more than 50% of the left ventral arm in 

males. Species 3 is a larger maturing form (more than 150 mm mantle length) which falls within the 

“Photololigo chinensis” complex with a broad, robust mantle and large head and arms; sharp conical 

toothed sucker rings on the arms and large sharp conical teeth interspersed with smaller ones on the large 

tentacular sucker rings. All species have paired light organs adjacent to the rectum on the ink sac. 

It is recommended that samples of animals (including mature males and females where available) which 

cannot be identified readily from the key should be preserved in formalin or alcohol, labelled with location, 

date of capture, depth and means of capture, and name of collector, sealed in a plastic bag and sent to an 

expert for identification. Major regional cephalopod collections and cephalopod specialists are at the 

Museum of Victoria (Invertebrate Zoology), Swanston Street, Melbourne, Victoria 3000 Australia and at the 

National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, Tokyo 169, Japan. Material marked “squid 

for scientific study” may also be sent for identification to Dr Malcolm Dunning, Fisheries Group, Queensland, 

Department of Primary Industries, GPO Box 3129, Brisbane, Queensland 4001, Australia. 

Key to the species of Loliginidae occurring in the area 

1a. Mantle very long, narrow, its posterior end drawn out into a long, pointed tail, especially 

in males; posterior border of fins strongly concave, extend part way or entirely along tail 

as narrow membranes (Fig. 1a) . . . . . . . . . . . . . . . . . . . . . . . . . Uroteuthis bartschi 

1b. Mantle elongate or short, relatively robust, its posterior end pointed or rounded, but 

never produced into an elongate, pointed tail; posterior border of fins straight, slightly 

concave, or rounded (Fig. 1b-d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

tail long, 

pointed 

heart-shaped 

fins 

rhombic 

fins 

a) Uroteuthis bartschi b) Loliolus c) Photololigo d) Sepioteuthis 

Fig. 1 characteristic fin shapes of loliginid genera (dorsal view) 

(after Adam, 1984; Voss, 1963; Okutani, 1995) 

fins 

long, 

broad


2a. Fins very long, more than 90% of mantle length, broad; 

Sepia-like but much wider and more muscular; mantle 

very robust (Fig. 2) . . . . . . . . . . . . “Sepioteuthis lessoniana” 

2b. Fins short to moderately long, less than 75% of mantle 

length; mantle elongate and narrow to short and stout . . . . . . → 3 

3a. Fins heart-shaped, rounded (Fig. 1b); mantle stout; mature 

at small sizes (less than 50 mm mantle length) . . . . . . . → 4 

3b. Fins rhomboidal, with posterior borders straight or 

slightly concave (Fig. 1c); mantle elongate, bluntly to 

sharply pointed; generally mature at more than 50 mm 

mantle length . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

4a. Pair of light organs present on either side of the rectum 

on ink sac; large tentacular sucker rings smooth 

(Fig. 3a); sucker stalks on hectocotylus (left ventral arm 

IV) of males modified as papillae along entire length 

(Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . Loliolus noctiluca 

dorsal view 

4b. No light organs on ink sac; large tentacular suckers with 

small blunt truncate teeth around entire margin (Fig. 3b); 

no conspicuous papillae on hectocotylus of males . . . . . . . 

Fig. 2 “Sepioteuthis lessoniana” 

→ 5 (illustration: K.Hollis/ABRS) 

a) smooth 

b) small blunt 

truncate teeth 

c) subequal-sized regularly 

spaced sharp conical teeth 

Fig. 3 large tentacular sucker 

d) unequal-sized irregularly 

spaced sharp conical teeth 

5a. Large tentacular suckers with 20 to 29 small truncate teeth on the entire margin; suckers 

and sucker stalks on ventral column of hectocotylus (left ventral arm IV) absent in 

mature males, ventral protective membrane broad, thickened, forming a large crest 

present proximally and becoming narrower towards the arm tip . . . . . . . . . Loliolus hardwickei 

5b. Large tentacular suckers with 15 to 20 small truncate teeth on the entire margin; 

hectocotylized arm in males without a proximal crest on the modified ventral protective 

membrane (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loliolus affinis 

a) dorsal view b) hectocotylus 

Fig. 4 Loliolus noctiluca 


papillae 

dorsal view 

Fig. 5 Loliolus affinis 

(after Lu, Roper, and Tait, 1985)


6a. Gladius relatively broad without 

thickened lateral asymptotes, 

cone flags developed (Fig. 6a) . . . . . . → 7 

6b. Gladius narrow with thickened 

lateral asymptotes and much reduced 

cone flags (Fig. 6b) . . . . . . . . → 8 

7a. Large suckers of arms III with 

low, broad plate-like square 

teeth in the distal margin, proximal 

margin smooth (Fig. 7a) . . . . . . . → 9 

7b. Large suckers of arms III with 

slender teeth on the distal margin, 

either square-tipped (Fig. 

7b) or sharp conical (Fig. 7c), 

proximal margin smooth or with 

low serrations . . . . . . . . . . . . . . → 10 

8a. Suckers of lateral arms (II and 

III) with 7 to 11 wide teeth distally 

(Fig. 8) . . . . Photololigo singhalensis 

8b. Suckers of lateral arms (II and 

III) with wide rough teeth around 

entire ring; in males arms II and 

III suckers greatly enlarged to 

be more than twice the diameter 

of the median club suckers . . . 

. . . . . . . . . . . . . Photololigo pickfordae 

cone 

flags 

reduced 

cone 

flags 

developed 

a) broad b) narrow 

Fig. 6 gladius 

(after Alexyev, 1989) 

a) broad plate-like 

teeth 

b) long, square-tipped 

truncate teeth 

c) sharp conical 

teeth 

Fig. 7 arm III 

sucker rings 

9a. Medial tentacular sucker rings smooth (Fig. 3a) or with only low serrations distally . . . . . . → 11 

9b. Medial tentacular sucker rings with subequal-sized, regularly spaced sharp conical or 

triangular teeth around their entire diameter (Fig. 3c) . . . . . . . . . . . . . . . . . . . . . . → 12 

10a. Large suckers of arms III with 6 to 12 long, square-tipped teeth on the distal margin 

(Fig. 7b); posterior fin margins distinctly concave; in males, dorsal and ventral sucker 

stalks of the left ventral arm modified into papillae for more than 50% of its length (Fig. 9) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . “Photololigo edulis” complex 

10b. Large suckers of arms III with 10 to 18 sharp, conical teeth on the distal margin (Fig. 7c); 

posterior fin margins almost straight; in males, papillate hectocotylus involving less than 

40% of the left ventral arm (Fig. 10) . . . . . . . . . . . . . . . . “Photololigo chinensis” complex 

a) dorsal view 

7-11 wide 

teeth distally 

b) arm III 

sucker ting 

Fig. 8 Photololigo singhalensis 

ventral view 

Fig. 9 “Photololigo edulis” 

dorsal view 

Fig. 10 “Photololigo chinensis” 

(illustration: K.Hollis/ABRS)


11a. Arm III sucker rings with 5 to 10 low plate-like teeth; diameters less than those of the 

largest tentacular sucker rings (Fig. 11a) which are smooth or have low serrations 

around only part of their diameter (Fig. 12) . . . . . . . . . . . . . . . . Nipponololigo sumatrensis 

11b. Arm III sucker rings with 3 to 6 low plate-like teeth; diameters approximately equal to 

those of the largest tentacular sucker rings (Fig. 11b) which are smooth . . . . . Nipponololigo uyii 

91.5 mm 

mantle length 

110 mm 

mantle length 

arm III sucker tentacular sucker 

ring 

ring 

a) Nipponololigo sumatrensis 

68.5 mm 

mantle length 

Fig. 11 relative sizes of sucker rings of arms III and tentacles 

(after Natsukari, 1984) 

92 mm 

mantle length 

arm III sucker 

tentacular sucker 

ring 

ring 

b) Nipponololigo uyi 

12a. Suckers on arms II and III equal to or greater than 

diameter of largest tentacular sucker rings (Fig. 13a); 

suckers of arm II in males greatly enlarged (more than 

ventral view 

2 times larger than large tentacular sucker diameter, 

Fig. 13a); arm suckers with 3 to 7 large low plate-like 

Fig. 12 Nipponololigo 

sumatrensis 

teeth; no paired light organs either side of the rectum on 

ink sac (Fig. 14) . . . . . . . . . . . . . . . . . Nipponololigo beka 

12b. Suckers on arms II and III generally less than or equal to diameter of largest tentacular 

sucker rings (Fig. 13b); suckers of arm III in males somewhat enlarged (about 1.5 times 

larger than large tentacular sucker diameter, Fig. 13b); arm suckers with 5 to 13 low 

plate-like teeth; paired light organs either side of the rectum on ink sac (Fig. 15) . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Photololigo duvaucelii 

II 

6.5 mm mantle length 

II 

III 

III 

104 mm mantle length 


sucker rings of arms 

large tentacular 

sucker ring 


sucker rings of arms 

large tentacular 

sucker ring 

a) Nipponololigo beka b) Photololigo duvaucelii 



Fig. 13 relative sizes of sucker rings of arms II and III and the large tentacular sucker rings 

II 

II 

III 

III


ventral view 

Fig. 14 Nippololigo beka 



? Loligo reesi (Voss, 1962) 

? Loligo vossi Nesis, 1982 

Loligo sp. nov. [Chotiyaputta, 1993] 

Loliolus affinis Steenstrup, 1856 

Loliolus hardwickei Gray, 1849 

Loliolus noctiluca Lu, Roper, and Tait, 1985 

Nipponololigo beka (Sasaki, 1929) 

Nipponololigo sumatrensis (d’Orbigny, 1835) 

Nipponololigo uyii Wakiya and Ishikawa, 1921 

“Photololigo chinensis“ (Gray, 1849) 1/ 

Photololigo duvaucelii (Orbigny, 1848) 

“Photololigo edulis“ (Hoyle, 1885) 2/ 

Photololigo pickfordae Adam, 1954 

? Photololigo robsoni Alexeyev, 1992 

Photololigo singhalensis (Ortmann, 1891) 

Photololigo sp. 1 [Yeatman and Benzie, 1994] 



“Sepioteuthis lessoniana“ Lesson, 1830 3/ 

Uroteuthis bartschi Rehder, 1945 

dorsal view 

Fig. 15 Photololigo duvaucelii 

1/ Probably a species complex including Loligo etheridgei Berry, 1918, L. formosana Sasaki, 1929, and Photololigo 

sp. 3 of Yeatman and Benzie, 1994. 

2/ Probably a species complex including Loligo edulis, L. budo, and Photololigo spp. 1 and 2 of Yeatman and Benzie, 

1994. 

3/ Probably a species complex.


References 

Adam, W. 1954. Cephalopoda. Partie III,IV. Cephalopodes a’ l’exclusion des genres Sepia, Sepiella et Sepioteuthis. 

Siboga Expéd., Leiden, 55c:123-193. 

Chotiyaputta, C. 1993. A survey on diversity and distribution of juvenile squids in the inner and western Gulf of Thailand. 

Thai Mar. Fish. Res. Bull., 4:19-36. 

Ghofar, A. 1989. Population biology and fishery of squid and cuttlefish in the Alas Strait, Indonesia. Unpublished Ph.D. 

Thesis, Gwynedd, UK, University College of North Wales, 254 p. 

Jackson, G.D. 1990a. Age and growth of the tropical nearshore loliginid squid Sepioteuthis lessoniana determined from 

statolith growth ring analysis. Fish. Bull. NOAA, 88:113-118 

Jackson, G.D. 1990b. The use of tetracycline staining techniques to determine statolith growth ring periodicity in the 

tropical loliginid squids Loliolus noctiluca and Loligo chinensis. Veliger, 33:389-393 

Lu, C.C. and R.W. Tait. 1983. Taxonomic studies on Sepioteuthis Blainville (Cephalopoda: Loliginidae) from the Australian 

region. Proc. R. Soc. Vict., 95:181-204. 

Lu, C.C., C.F.E. Roper, and R.W. Tait. 1985. A revision of Loliolus (Cephalopoda: Loliginidae) including L. noctiluca, a 

new species of squid from Australian waters. Proc. R. Soc. Vict., 97:59-85. 

Nateewathana, A. 1992. Taxonomic studies on loliginid squids (Cephalopoda: Loliginidae) from the Andaman Sea coast 

of Thailand. Phuket Mar. Biol.Centre Res. Bull., 57:1-40. 

Natsukari, Y. 1983a. Taxonomical and morphological studies on the loliginid squids. V. Redescription on the type 

specimen of Loligo sumatrensis d’Orbigny, 1835. Venus, 42:259-263. 

Natsukari, Y. 1983b. Taxonomical and morphological studies on the loliginid squid. III. Nipponololigo, a new subgenus 

of the genus Loligo. Venus, 42:313-318. 

Natsukari, Y. 1984. Taxonomical and morphological studies on the loliginid squids. IV. Two new genera of the family 

Loliginidae. Venus, 43:229-239. 

Natsukari, Y. and T. Okutani. 1975. Taxonomic and morphological studies on the loliginid squids - I. Identity of Loligo 

chinensis Gray, 1849, redescription of the type specimen and taxonomic review (Cephalopoda: Loliginidae). Jap. 

J. Malacology (Venus), 34:85-91. 

Natsukari, Y. and M. Tashiro. 1991. Neritic squid resources and cuttlefish resources in Japan. Mar. Behav. Physiol., 

18:149-226. 

Nesis, K.N. 1982. Kratkiy opredelitel’ golovonogikh molluskov Mirovogo okeana. (Abridged key to the cephalopod 

mollusks of the World Ocean). Light and Food Industry Publishing House, Moscow, 358 p. (Also translated and 

revised as Nesis, K.N. 1987. Cephalopods of the world. Neptune City, TFH Publications Inc., 351 p.) 

Segawa, S., S. Horayama, and T. Okutani. 1993. Is Sepioteuthis lessoniana in Okinawa a single species? In Recent 

advances in cephalopod fisheries biology, edited by T. Okutani, R.K. O’Dor, and T. Kubodera. Tokyo, Tokai 

University Press, pp. 513-521. 

Vecchione, M., T.F. Brakoniecki, Y. Natsukari, and R.T. Hanlon. (in press). A provisional generic classification of the Family 

Loliginidae. Smithson. Contrib. Zool. 

Voss, G.L. 1962. Six new species and two new subspecies of cephalopods from the Philippine Islands. Proc. Biol. Soc. 

Wash., 75:169-176. 

Voss, G.L. 1963. Cephalopods of the Philippine Islands. Bull.U.S. Natl. Mus., 234:1-180. 

Yeatman, J. and J.H. Benzie. 1994. Genetic structure and distribution of Photololigo spp. in Australia. Mar. Biol., 

118:79-87.


Nipponololigo beka (Sasaki, 1929) 

Frequent synonyms / misidentifications: None / Nipponololigo uyii 

Wakiya and Ishikawa, 1921. 

FAO names: En - Beka squid; Fr - Calmar cracheur; Sp - Calamar beka. 

Diagnostic characters: Mantle short, slender; fins rhomboidal with 

round lateral angles, length more than 50% of mantle length. Tentacular 

clubs expanded, lanceolate; medial manus suckers twice the diameter 

of lateral suckers and with 18 to 30 sharp teeth on the sucker rings. 

Arm sucker rings with 3 to 7 very low broad squared teeth; left 

ventral arm modified in males with palisade-like fleshy papillae on 

the distal 60%. 

Size: Maximum dorsal mantle length 70 mm. 

Habitat, biology, and fisheries: Found in coastal 

waters, particularly bays. Occurs in trawl and lightluring 

net catches in the Gulf of Thailand. 

Distribution: Southeast Asian coastal waters to 

southern Japan. 

? 


sucker ring 

arm III sucker ring 

ventral view


Nipponololigo sumatrensis (Orbigny, 1835) 

Frequent synonyms / misidentifications: Loligo kobiensis Hoyle, 1885; 

L. rhomboidalis Burgess, 1967;L. yokoyae Ishikawa, 1926 / Nipponololigo 

uyii Wakiya and Ishikawa, 1921. 

FAO names: En - Kobi squid; Fr - Calmar kobi; Sp - Calamar kobí. 

Diagnostic characters: Mantle short, slender; fins rhomboidal with 

round lateral angles, length approximately 65% of mantle length. Tentacular 

clubs expanded, lanceolate; club suckers in 4 rows, 6to8medial 

manus suckers up to 4 to 5 times the diameter of lateral suckers and 

with smooth sucker rings (occasionally with low serrations distally); 

other larger tentacular suckers with 6 to 15 sharp teeth. Arm sucker rings 

with 5 to 10 low broad squared teeth, left and right ventral arms (IV) 

modified in males; beyond the first 3 rows of normal suckers, sucker 

stalks of the left arm modified as 

low stump-like papillae in the dorsal 

column and broad thick palisade-like 

papillae in the ventral 

column, reducing in size distally; 

right arm IV with 3 or 4 rows of 

much enlarged suckers proxi- 

mally. 

Size: Maximum dorsal mantle length 

100 mm, commonly to 50 mm mantle 

length. 


Found in coastal waters, common in 

upper 10 m of the water column. 

Abundant in Gulf of Thailand trawl 

and light-luring net catches. 

Distribution: Southeast Asian 

coastal waters to central Japan. 


sucker ring 


tentacular 

club 


ventral view


“Photololigo chinensis” Gray, 1849 species complex 

Frequent synonyms (named species of uncertain status included 

in the complex) / misidentifications: Loligo chinensis Gray, 1849; L. 

australis Gray, 1849; L. formosana Sasaki, 1929; L. etheridgei Berry, 

1918; L. indica Hoyle, 1886; ?L. vossi Nesis, 1982; Photololigo sp. 3 of 

Yeatman and Benzie, 1994 / ?Doryteuthis singhalensis of Voss, 1963. 

FAO names: En - Mitre squid; Fr - Calmar mitre; Sp - Calamar mitrado. 

Diagnostic characters: Mantle 

elongate, slender, bluntly pointed 

posteriorly; fins rhombic, long, 

over 60% of mantle length in 

adults. About 12 medial manal 

suckers of tentacular clubs enlarged 

to 1.5 times the diameter of 

lateral suckers and twice the diameter 

of largest arm sucker; large 

rings with 20 to 30 sharp, conical 

separate teeth, the 6 to 12 larger 

ones interspersed with 1 to 4 

smaller ones. Larger sucker rings 

of arms II and III with 10 to 18 

sharp conical teeth distally, with 

degenerate teeth or smooth proximally; 

left arm IV hectocotylized 

at distal 33 to 40% by modification 

of more than 30 suckers and stalks 

in each row into slender, conical 

papillae that remain larger in the 

ventral row. 


sucker ring 


tentacular 

club 


dorsal view 

Size: Maximum 400 mm mantle length, commonly to 200 mm mantle (illustration: K. Hollis/ABRS) 

length. 

Habitat, biology, and fisheries: Neritic species ranging from intertidal areas to a depth of 170 m; positively 

phototactic and periodically forming large aggregations. Spawning occurs throughout the year, with peaks 

in Spring and Autumn. This species complex is targetted or a welcome bycatch of numerous commercial 

and small-scale fisheries throughout its range; major squid species together with Photololigo duvaucelii 

in the Gulf of Thailand, where it is taken in waters from depths between 15 and 30 m and amounts to up 

to 71% of the catch (more than 40 000 t in 1984) from light-luring vessels. Bycatch (together with 

“Photololigo edulis”) in prawn trawling off northern Australia and occurs in minor quantities in Indonesian, 

Malaysian, and Philippine catches. Taken with a variety of gears, including demersal trawls, purse seines, 

cast, lift, dip and box nets, hook-and-lines including jigs, scoop nets, and stake nets, often involving light 

attraction. Marketed fresh, frozen, or dried and is also processed into cleaned “hoods” and rings. 

Distribution: Western Pacific: 

South and East China Seas to 

Japan, Gulf of Thailand, 

Arafura and Timor seas, northern 

Australia.


Photololigo duvaucelii (d’Orbigny, 1835) 

Frequent synonyms / misidentifications: Loligo 

duvauceli Orbigny, 1848, L. oshimai Sasaki, 1929; 

L. indica Pfeffer, 1884 / None. 

FAO names: En - Indian squid; Fr - Calmar indien; 

Sp - Calamar índico. 

Diagnostic characters: Mantle relatively short, stout. 

Fins rhombic, broad, short, just over 50% of mantle 

length. Tentacular clubs expanded; large median 

manal suckers 1.5 times larger than marginals, with 

14 to 17 short, sharp teeth around ring. Arm suckers 

of female of about equal size on arms II and 

III, rings smooth proximally, toothed with about 7 

broad, blunt teeth distally; in males, suckers of 

arm II and especially III greatly enlarged, with 5 to 13 

low, squared to rounded, truncate teeth in the 

distal 2/3 of ring, proximal 1/3 smooth; leftarmIV 

of male hectocotylized for 

more than 1/2 its length, with 2 

rows of large papillae, some 

with minute suckers on tip, 

ventral rows larger, turned outward, 

comb-like; an oval photophore 

on each side of rectum 

on the ink sac. 

Size: Maximum mantle length 

300 mm, commonly to 150 mm 

mantle length. 


A neritic, inshore species 

occurring in depths between 

30 and 170 m, forming large 

aggregations during the 

spawning season. Spawning 

occurs throughout the year. 

Feeds on crustaceans (such 

as mysids, euphausids and 

ostracods), fishes and squids; 

cannibalism is common. Exploited 

throughout its range, 

mainly by artisanal subsistance 

fisheries; in the Gulf of 

Thailand it is one of the target 

species of the demersal trawl 

and light-luring fisheries (with 

“Photololigo chinensis”) with 

over 15 000 t taken in 1984. 

Distribution: Indo-Pacific: Indian 

Ocean periphery, including 

the Red Sea and the 

Arabian Sea, extending eastwards 

from Mozambique to the 

South China Sea and the Philippines 

Sea, northward to Taiwan 

Province of China. 


sucker ring 

arm III sucker ring dorsal view


“Photololigo edulis” (Hoyle, 1885) species complex 

Frequent synonyms / misidentifications: Doryteuthis kensak (Wakiya 

and Ishikawa, 1921); Loligo budo Wakiya and Ishikawa, 1921; Photololigo 

sp. 1 and 2 of Yeatman and Benzie, 1994 / ?Doryteuthis sibogae Adam, 

1954. 

FAO names: En - Swordtip squid; Fr - Calmar épée; Sp - Calamar espada. 

Diagnostic characters: Mantle moderately stout to elongate, slender in 

mature males. Fins rhombic, attaining 70% of mantle length in adults, 

their posterior margin slightly concave. Tentacular clubs expanded, lanceolate; 

about 16 medial manal suckers 1.2 times larger than the marginals, 

approximately equal to largest arm suckers, with 30 to 40 sharp conical 

teeth, 20 to 30 small ones interspersed between 

10 larger ones. Arm sucker rings with 6 to 12 

distinct, squared, truncate teeth in distal 2/3, 

smooth or with rudimentary denticles proximally; 

slightly more than 50% of left arm IV hectocotylized 

by enlargement of about 50 pairs of 

sucker stalks into swollen papillae, each with a 

minute rudimentary sucker on the tip; papillae 

slightly larger in ventral row; a fusiform photophore 

on each side of rectum. Seasonal forms present 

around Japan (“budo” and “kensaki”) varying in 

the robustness of the mantle and number of normal 

sucker rows at the base of the hectocotylized 

arm; small maturing (less than 120 mm mantle 

length) unnamed species morphologically similar 

but genetically distinct occurring off northern Australia. 

Size: Maximum mantle length 400 mm; common 

size in commercial catches in South China Sea 

between 150 and 250 mm. 

Habitat, biology, and fisheries: A neritic species 

occurring in depths from 30 to 170 m. May undergo 


sucker ring 


ventral view 

seasonal inshore-offshore migrations, forming large aggregations. Spawning of the kensaki form has been 

reported on sandy bottoms in depths from 30 to 40 m off southern Japan. Supports local fisheries 

throughout its range, in western Japan, the Philippines, and in Indonesia. At least 2 species in this complex 

are taken as bycatch in prawn trawling in northern Australian waters. Other gears include hand jigs and 

set nets. The flesh is of good quality and sold at high prices both fresh and frozen, processed into a dried 

product and also used for 

sashimi in Japan. 

Distribution: Western Pacific 

from northern Australia, Philippine 

Islands, South China 

Sea to central Japan.


Photololigo singhalensis (Ortmann, 1891) 

Frequent synonyms / misidentifications: Doryteuthis sibogae Adam, 

1954; Loligo sibogae (Adam, 1954) / Photololigo chinensis (Gray, 1849). 

FAO names: En - Long barrel squid; Fr - Calmar baril; Sp - Calamar buril. 

Diagnostic characters: Mantle long and slender. Fins long, reaching 

approximately 50% of mantle length. Tentacular clubs short, slightly 

expanded; suckers in medial rows of manus only about 25% larger 

than those on lateral rows with 20 to 22 sharply pointed, curved 

teeth, some of which are quite reduced in size. Arms relatively short; 

sucker rings with 7 to 11 long, plate-like squared teeth distally, 

smooth proximally.LeftarmIV hectocotylized in distal half with slender 

papillae. Paired, bean-shaped light organs present 

adjacent to the rectum on the ink sac. 

Size: Maximum mantle length 500 mm (males) 

and 310 mm (females), commonly caught at 150 

to 200 mm mantle length. 

Habitat, biology, and fisheries: A neritic, 

semipelagic species occurring at depths from 30 

to 120 m. It is positively phototactic, a feature that 

is utilized in the fishery by attracting it by light prior 

to capture. It aggregates in large schools in summer, 

probably for mating and spawning. Males 

grow larger than females. In the Philippines, Indonesia 

and Taiwan Province of China (Penghu), it 

supports localized and subsistence fisheries and 

is taken by jigs, purse seines, and dip-nets using 

light attraction together with Photololigo chinensis 

and other large loliginids. Seasonally abundant, 

indicating some migration. 

Distribution: Indo-Pacific from Eastern Arabian 

Sea, Bay of Bengal to South China Sea and Philippines 

Sea, Indonesian waters, Solomon Sea, 

and Taiwan Province of China (Penghu). 

? 

? 

? 

? 

? 

largest tentacular 

club sucker ring 

largest arm III 

sucker ring 

dorsal view


“Sepioteuthis lessoniana” Lesson, 1830 ? species complex 

Frequent synonyms / misidentifications: Sepioteuthis 

arctipinnis Gould, 1852 / None. 

FAO names: En - Bigfin reef squid; Fr - Calmar tonnelet; 

Sp - Calamar manopla. 

Diagnostic characters: Mantle 

long, robust, its width about 40% 

of length. Fins very large, length 

over 90% mantle length, width up 

to 75% mantle length; greatest 

width occurs posterior to the 

midpoint of the fins. Tentacular 

clubs long, expanded; median manus 

suckers enlarged; rings with 14 

to 23 sharp conical teeth. Arm 

sucker rings with 18 to 29 long 

sharp, regularly spaced, conical 

teeth around entire margin; tentacles 

long, robust; left arm IV hectocotylized 

along distal 25 to 30%. 

Colour: dark to light brown to pale 

translucent; iridescent pale transverse 

bars or spots are present dorsally 

in all colour phases and are 

distinctive of this species. 

Size: Maximum dorsal mantle length 

422 mm in males, smaller in females; 

commonly 200 to 300 mm mantle 

length. 


Sepioteuthis lessoniana is a neritic 

species occurring from the surface 

down to a depth of at least 100 m. The 

spawning season may be quite extended. 

Egg capsules containing 3 to 

7 eggs are finger-shaped and attached 

in clusters to seaweeds, submerged 

mangrove roots, twigs, stones and corals 

in coastal waters. Around Okinawa 

andPalau,aformlaying2eggsper 

capsule attached underneath coral 

boulders has recently been reported. 

Preys primarily on prawns and fishes, 

occasionally on stomatopods and crabs. Grows to 200 mm in about 150 days in northern Australia. Males 

reach larger sizes than females. Of major commercial and artisanal value throughout Southeast Asia, 

captured using a variety of gear from demersal trawls to jigs and hooks, spears, set nets, and traps. 


the Indo-Pacific: Red Sea 

eastward to the Hawaiian Islands, 

northern Australia to 

central Japan. 


sucker ring 


hectocotylus 

distal end of 

hectocotylus 

? 

? 

? 

dorsal view 


? 

? 

? 

?


Uroteuthis bartschi Rehder, 1945 


FAO names: En - Bartsch’s squid; Fr - Calmar tépo; 

Sp - Calamarete. 

Diagnostic characters: Mantle very narrow, elongate, 

with very long, pointed tail, more pronounced in males. 

Fins rhomboidal, their lateral angles rounded, posterior borders 

concave, generally extending the entire length of tail 

but sometimes as only a minute membrane. Head relatively 

small, narrow. Arm suckers with broad, plate-like teeth in the 

distal margin, smooth proximally; large medial tentacular 

suckers with long square-tipped teeth. Left arm IV hectocotylized 

in distal half by abrupt transformation of suckers 

into long, stout papillae. 


Habitat, biology, and fisheries: A neritic species; upper 

and lower limit of depth range undetermined. Taken as 

bycatch in local trawl and net fisheries in Indonesia and the 

Philippines. Utilized mostly fresh. 

Distribution: Western Pacific Ocean: in Philippine and Indonesian 

waters. 

? 

? 

? 

? 

dorsal view 

of female 


dorsal view 

of male


Loliolus affinis Steenstrup, 1856 

Frequent synonyms / misidentifications: None / ?Photololigo spp.; 

Nipponololigo spp. 

En - Steenstrup’s bay squid. 

Maximum mantle length to 39 mm (females 

larger than males), occur in shallow coastal 

habitats to depths of at least 13 m. Prob- 

ably a schooling species. Caught occasionally 

in demersal trawl catches in the Gulf of 

Thailand among larger loliginids, may be 

confused with juveniles of other larger 

more commercially important species, mature 

males readily identified by the absence 

of any normal suckers from the 

hectocotylized arm. 

Loliolus noctiluca Lu, Roper, and Tait, 1985 

Frequent synonyms / misidentifications: None / Photololigo spp.; 

Nipponololigo spp. 

En - Luminous bay squid. 

Maximum mantle length to 90 mm 

(females larger than males). Occur 

in shallow coastal habitats including 

seagrass beds to depths of about 

50 m. Caught incidentally in inshore 

prawn trawls along the northeastern 

Australian coast, may be confused 

with juveniles of other larger more 

commercially important species 

from which they can be separated 

by fin shape and, in freshly caught 

specimens, by the presence of yellow 

and pink chromatophores on 

the fin margins and mantle. Short 

lived, reaching 60 mm mantle 

length in tropical Australian waters 

in approximately 70 days. 


sucker ring 

arm III 

sucker ring 


sucker ring 

arm III 

sucker ring 

hectocotylus 

hectocotylus 

dorsal view 

(after Lu, Roper, and Tait, 1985) 

dorsal view 


Enoploteuthidae 781 

Enoploteuthidae ENOPLOTEUTHIDAE 

Firefly squids, enope squids 


Diagnostic characters: Small to medium-sized squids (generally less than 150 mm mantle length) with 

only moderately muscular or gelatinous mantles. Fin shape varies. Light organs present in adults 

of all genera; arrangement, size and number of light organs is a generic and species specific characteristic. 

Characterized by a simple, straight funnel locking apparatus. Sharp toothed suckers (Pterygioteuthis) 

or suckers and hooks (remaining genera) arranged in biserial rows on arms and in 4 columns on 

tentacular clubs. Buccal connectives attached to dorsal border of ventral arms (IV). One or both 

ventral arms hectocotylized in males, with considerable variation between species. Hectocotylized arm of 

Pyroteuthis with hooks modified by the development of a secondary cusp. All hooks replaced in Pterygioteuthis 

by a single chitinous plate housed in a fleshy pocket midway along the oral surface of the ventral 

arm. 

ventral view 

Pterygioteuthis 

fin shape varies 

light 

organ 

ventral view 

Ancistocheirus 

Habitat, biology, and fisheries: Epi- to mesopelagic, tropical and subtropical oceanic squids, some 

species spawning in continental slope and deeper continental shelf waters. Female reproductive organs 

are modified from the typical oegopsid squid form by the loss of the nidamental glands and strong 

development of the oviducal glands in the genera Abralia, Abraliopsis, and Ancistrocheirus. In Pyroteuthis 

and Pterygioteuthis, only the right or left oviducal glands are present. Though some oegopsid squid spawn 

eggs in masses surrounded by jelly, some female enoploteuthids spawn individual eggs into the plankton. 

Females of enoploteuthid squid such as Abralia and Abraliopsis which mature at about 50 mm mantle 

length are capable of producing between 10 000 and 20 000 eggs, typically with a maximum diameter of 

1.2 mm. Ripe eggs of mature Ancistrocheirus are large for oegopsids, reaching 3 mm over the long axis. 

Enoploteuthids are among the most commonly caught squids in midwater trawls and migrate to deeper 

waters during the daytime. Several genera are caught in demersal trawls in deeper slope waters. Their 

abundance in the epipelagic and mesopelagic zones is reflected by their prevalence in the diets of tunas 

and lancetfish, sharks, and marine mammals. Enoploteuthids are not the subject of target fisheries in 

tropical waters of the Western Central Pacific, although they do appear in some markets as bycatch from 

trawling activities. In the Sea of Japan, up to several thousand tonnes of a temperate species closely related 

to Abralia and Abraliopsis, Watasenia scintillans (Berry, 1911), are taken annually using set nets to 

capture spawning aggregations. Watasenia are processed by boiling whole.



Octopoteuthidae: larger species may appear superficially similar to 

Ancistrocheirus (long rhomboid fins, hooks on arms) but are distinguished 

by the following characters: buccal connectives attached to 

ventral border (rather than to dorsal border) of ventral arms (IV); loss 

of tentacles in adult stage; no light organs on ventral mantle (although 

present on tips on some of the arms, internally on the viscera, 

and in the tail). Species of Ancistrocheirus have several light organs 

on the ventral mantle and the ventral surface of the head. 

Key to the genera of Enoploteuthidae occurring in the area 

(after Nesis, 1987) 

1a. Light organs present on the surface of mantle, 

head, and arms; none or only a single light organ 

present inside the mantle cavity; no light organs 

or a single row only on ventral side of eyeball; 

posterior end of mantle conical, not produced into 

an acute tail, terminal cone of gladius slightly 

ventral view 

developed; posterior end of fins concave or 

straight; both oviducts developed . . . . . . . . . . . . . → 2 Octopoteuthidae 

1b. No light organs on the surface of mantle, head, or arms; 8 to 10 light organs present 

inside the mantle cavity, embedded in the tentacular stalk; 12 to 15 light organs on 

ventral side of eyeball; posterior end of mantle extending into an acute tail with a 

needle-like terminal cone of the gladius; fins rounded, their posterior edges convex, not 

reaching end of body; only 1 oviduct developed . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

2a. Twenty-two large semicircular light organs present on ventral surface of mantle; large 

light organs also present on head and tentacular stalk; many minute light organs 

scattered over mantle, head, and ventral arms; no light organs inside mantle or on 

ventral side of eyeball; anterior end of mantle elongated in adults to form a tail extending 

beyond posterior edge of fins; hooks in 2 rows on tentacular clubs; in adults, no marginal 

suckers on central part of clubs; nidamental glands developed; adults reaching 400 mm 

mantle length (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Ancistrocheirus 

2b. Hundreds of small, approximately equal-sized light organs present on ventral surface 

of mantle, head, and arms; light organs also present on ventral side of eyeball; no light 

organs present in the tentacular stalk; nidamental glands absent and replaced by 

oviducal glands that are greatly enlarged and divided into 2 halves; adults rarely 

exceeding 200 mm mantle length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Eight to 12 light organs on ventral 

side of eyeball, the 2 marginal 

ones large, the others 

small and abutting one another; 

posterior end of mantle in 

adults usually developed into a 

soft conical tail extending beyond 

edge of fins (Fig. 2) . . Enoploteuthis 

3b. Five to 8 unequal-sized, separate 

light organs on ventral side 

of eyeball; fins reaching posterior 

end of mantle; central part 

of tentacular clubs with 1 row of 

hooks and 2 rows of suckers, or 

with 2 rows of hooks and several 

small suckers between 

them (suckers only present in 

larvae and juveniles) . . . . . . . . . . → 4 

ventral view 

Fig. 1 Ancistrocheirus 

dorsal view 

Fig. 2 Enoploteuthis 

light 

organs 

no tentacles 

in adults 


Enoploteuthidae 783 

4a. No large black globular light organs on tips of ventral arms; only silvery light organs may 

be present, smaller than diameter of arms; hooks in a single row on clubs; 5 to 8 light 

organs of different size and structure (posterior one usually differing from the others) 

on ventral side of eyeball; buccal membrane pink; left (sometimes right) ventral arm (IV) 

hectocotylized (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Abralia 

4b. Three large light organs on tip of each ventral arm (appear like small black balls in 

preserved specimens, but are green in life); hooks in 2 rows on clubs; 5 round light 

organs of similar structure on ventral side of eyeball, both marginal light organs larger 

than middle ones, which are equal-sized; buccal membrane dark violet; right ventral arm 

(IV) hectocotylized . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Abraliopsis 

5a. Hooks present on tentacular clubs and on ventral arms (IV); hooks on club in a single 

row and in 2 rows along entire length of ventral arms; 12 light organs on ventral side of 

eyeball; 10 light organs present inside the mantle cavity; 6 to 8 light organs present in 

the tentacular stalks; the right ventral arm hectocotylized in males; only the left oviduct 

developed in females . . . . . . . . . . Pyroteuthis 

5b. No hooks on tentacular clubs and not more 

than 2 hooks on ventral arms (IV); a few 

hooks in 1 or 2 rows present in the middle 

of arms I to III; 14 to 15 light organs on 

ventral side of eyeball; 8 light organs present 

inside the mantle cavity; 4 light organs present 

in the tentacular stalks; left ventral arm 

hectocotylized in males; only the right oviduct 

developed in females (Fig. 4) . . . . . 

. . . . . . . . . . . . . . . . . . . Pterygioteuthis 


Abralia andamanica Goodrich, 1896 

Abralia armata (Quoy and Gaimard, 1832) 

Abralia astrolineata Berry, 1914 

Abralia astrostricta Berry, 1909 

Abralia lucens Voss, 1963 

Abralia renschi Grimpe, 1931 

Abralia similis Okutani and Tsuchiya, 1982 

Abralia spaercki Grimpe, 1931 

Abraliopsis chuni Nesis, 1982 

Abraliopsis lineata (Goodrich, 1896) 

Abraliopsis pfefferi Joubin, 1896 

Ancistrocheirus lesueuri Férussac and d’Orbigny, 1839 

Enoploteuthis galaxias Berry, 1912 

Enoploteuthis jonesi Burgess, 1982 

Enoploteuthis leptura magnoceani Nesis, 1982 

Enoploteuthis reticulata Rancurel, 1970 

Pterygioteuthis giardi Fischer, 1896 

Pyroteuthis margaritifera (Rüppell, 1844) 

dorsal view 

Fig. 3 Abralia 

dorsal view 

Fig. 4 Pterygioteuthis 


References 

Kubota, T., K. Iizuka, and T. Okutani. 1982. Some biological aspects of Abralia andamanica from Suruga Bay, Japan 

(Cephalopoda: Enoploteuthidae). J. Fac. Mar. Sci. Tech., Tokai Univ., 15:333-343. 

Okutani, T. 1976. Rare and interesting squid from Japan V. A gravid female of Ancistrocheirus lesueuri (d’Orbigny, 1839) 

collected in the Kuroshio Area (Oegopsida, Enoploteuthidae). Venus, 35:73-81. 

Okutani, T. and S. Tsukada. 1988. Squids eaten by lancetfish and tunas in the tropical Indo-Pacific oceans. J. Tokyo 

Univ. Fish., 75:1-44. 

Rancurel, P. 1976. Note sur les Cephalopodes des contenus stomacaux de Thunnus albacares (Bonnaterre) dans le 

Sud-ouest Pacifique. Cah. ORSTOM (Ser. Océanog), 14:71-80.


Onychoteuthidae ONYCHOTEUTHIDAE 

Hooked squids 


Diagnostic characters: Medium-sized to large oceanic squids, 

with a strong, muscular torpedo-shaped mantle and large 

muscular rhomboidal fins attenuated posteriorly in some species. 

Funnel locking cartilage simple, straight. Buccal connectives 

attached to ventral border of ventral arms. Arms with 

biserial and sometimes smooth suckers; tentacular clubs with 

biserial medial hooks and 2 rows of marginal suckers in the 

immature stages. Club-fixing apparatus consisting of smoothringed 

suckers and corresponding knobs present at the base of the 

tentacular clubs. Hectocotylisation not evident in males. Numerous 

distinctive pleats or folds are present in the neck region in Onychoteuthis 

and Ancistoteuthis (see Fig. 2a), and the skin of Moroteuthis 

often contains subcutaneous papillae and ridges. Lateral 

arms have swimming keels in some of the larger species. Visceral 

light organs are present in the genus Onychoteuthis. 

Habitat, biology, and fisheries: Onychoteuthids occur in tropical 

to polar waters in all oceans. They are oceanic squid, occasionally 

encountered in continental slope waters where they may be caught 

in demersal trawls. They feed on a broad range of fish, crustaceans 

including krill, squid, and other pelagic molluscs, and are at least 

partially cannibalistic. Known predators include lancetfish, tunas, 

whales, dolphins, seals, and albatrosses. Onychoteuthids form 

schools. Juvenile Onychoteuthis are able to leap above the surface 

to escape predators, as do ommastrephids, and sometimes are 

found on the decks of vessels. Immature Moroteuthis species 

share with other epipelagic cephalopods the deep blue dorsal day 

coloration characteristic of the neuston generally. Onychoteuthids 

are not fished commercially in the Western Central Pacific, although 

sometimes Moroteuthis are caught incidentally in demersal 

trawls in continental slope waters (300 to 600 m) off northern 

Australia and in the South China Sea. 

Remarks: Recent taxonomic studies have indicated that at least 

some tropical oceanic onychoteuthids previously assigned to the 

genus Onykia may be juveniles of various species of Moroteuthis. 


Ommastrephidae: also with an arrow-shaped fin, but can be 

distinguished by the following characters: funnel-mantle locking 

apparatus ⊥-shaped; sucker rings toothed; no hooks on tentacular 

clubs; buccal connectives attached to dorsal border (rather 

than to ventral border) of ventral arms. 

4 rows of 

suckers, 

no hooks 


Ommastrephidae 

dorsal view 

2 rows of 

hooks 


Onychoteuthidae

Onychoteuthidae 785 

Key to the genera of Onychoteuthidae occurring in the area 

1a. Skin smooth; posterior end of gladius a short chitinous needle (Fig. 1a); light organs 

present: 1 elongated light organ on ventral side of eyeball, 2 light organs on intestine 

and ink sac (anterior one small, posterior one large) (Fig. 2a) . . . . . . . . . . . . . Onychoteuthis 

1b. Skin smooth or rough; posterior end of gladius a long opaque cartilaginous rostrum 

(Fig. 1b); no light organs present (Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

a) Onychoteuthis 

short 

needle 

Fig. 1 gladius 

long 

rostrum 

b) Ancistroteuthis, 

Moroteuthis 

nuchal 

folds or 

pleats 

intestine 

2a. Six to 10 well-developed nuchal folds; skin smooth (Fig. 3) . . . . . . . . . . . . . Ancistroteuthis 

2b. No nuchal folds (Fig. 4a); skin thick, rough, or tuberculate (Fig. 4b) . . . . . . . . . . . Moroteuthis 

dorsal view 

Fig. 3 Ancistroteuthis 

ink 

sac 

light 

organs 

a) head and anterior b) light organs on 

part of mantle 

intestinal tract 

(dorsal view) (ventral view) 

c) dorsal view 

Fig. 2 Onychoteuthis 

nuchal 

folds 

absent 

a) head and anterior 

part of mantle 

b) dorsal view 

(dorsal view) 


Fig. 4 Moroteuthis



Ancistroteuthis lichtensteini (Orbigny, 1839) 

Moroteuthis lönnbergi Ishikawa and Wakiya, 1914 

Onychoteuthis “banksii” Leach, 1817 - species complex 

Onychoteuthis compacta (Berry, 1913) 

Onychoteuthis meridiopacifica Rancurel and Okutani, 1990 

References 

Kubodera, T., U. Piatkowski, T. Okutani, and M.R. Clareke. 1996. Taxonomy and Zoogeography of the Family Onychoteuthidae 

(Cephalopoda: Oegopsida). Smithson. Contrib. Zool. 

Rancurel, P. 1976. Note sur les Cephalopodes des contenus stomacaux de Thunnus albacares (Bonnaterre) dans le 

Sud-ouest Pacifique. Cah. ORSTOM (Ser. Océanog.), 14:71-80. 

Tsuchiya, K and T. Okutani. 1991. Growth stages of Moroteuthis robusta (Verrill, 1881) with a re-evaluation of the genus. 

Bull. Mar. Sci., 49:137-147.

Histioteuthidae 787 

Histioteuthidae HISTIOTEUTHIDAE 

Jewel squids, umbrella squids 


Diagnostic characters: Medium-sized to large squids. 

Characterized by a simple, straight funnel-mantle locking 

apparatus. Many large, anteriorly-directed light organs 

over the surface of mantle, head, and arms, more concentrated 

on the ventral surface. Unmodified toothed suckers arranged 

biserially on arms, and in 4 and 8 rows on tentacular 

clubs. Buccal connectives attached to dorsal border of ventral 

arms.Bothdorsalarms(I) usually hectocotylized in males. 

Head large in adults, with left eye considerably larger than 

right eye. Most species with mantle relatively short, broad, 

conical, and typically somewhat gelatinous (like the rest of the 

body). Terminal fins consist of separate oval lobes, free anteriorly 

but united posteriorly with a distinct median notch on the 

posterior border; the convex posterior margin of the fins may 

extend posteriorly beyond the mantle. Internal light organs absent. 

The light organ pattern on the mantle, head, and arms and 

the presence of enlarged terminal light organs on the arms are 

specific characters and may be used in ventral countershading. 

The species show considerable variation in the development of 

a web connecting the inner margins of the arms, extending up 

to 50% of the arm length in some species. 

Habitat, biology, and fisheries: The only genus of the family, 

Histioteuthis, comprises oceanic species, also occurring on the 

continental slope. No comprehensive life history studies of any 

histioteuthid in tropical waters have been conducted. Mature 

eggs of H. miranda have a maximum diameter of 0.8 mm. The 

number and size of beaks in sperm whale stomachs indicate 

that histioteuthids are schooling species. Although the family is 

considered to be primarily mesopelagic, H. celetaria pacifica 

and H. miranda have been caught frequently in deep-water 

lobster trawls at depths of 300 to 600 m off northern Australia, 

Histioteuthis celetaria pacifica 

and the latter species in demersal trawls in slope waters of the 

ventral view 

South China Sea, suggesting that they are associated with the 

bottom during part of their life cycle. They are important in the 


diets of sperm whales and also form part of the diets of yellowfin and albacore tuna, scabbard fish, 

lancetfish, spotted dolphin, and perhaps albatrosses elsewhere. Fish predominates in the diet of adult 

histioteuthids. Off Hawaii, H. dofleini shows both diel vertical migration and ontogenetic descent. Small 

jewel squids occur at depths from 200 to 300 m during the night and descend to 400 to 700 m during the 

day. Larger adults have been caught at depths of up to 800 m and only occasionally deeper. Other species 

are more common in waters of 2 000 m and are captured only rarely in the upper 200 m. Histioteuthids are 

not presently of commercial interest. Although some histioteuthid species reach in excess of 200 mm mantle 

length and are frequently, if not abundantly, encountered in continental slope waters, the gelatinous nature 

of their bodies and high ammonium content would detract from their market acceptance. 


None. Jewel squids are readily distinguished from other cephalopods by the anteriorly-directed light organs 

which cover the surface of the mantle, head, and arms. 


Histioteuthis celetaria pacifica (Voss, 1962) 

Histioteuthis dofleini (Pfeffer, 1912) 

Histioteuthis miranda (Berry, 1918) 

Histioteuthis meleagroteuthis (Chun, 1910) 

Reference 

Voss, N.A. 1969. A monograph of the Cephalopoda of the North Atlantic. The family Histioteuthidae. Bull. Mar. Sci., 

19:713-867. 




Ommastrephidae OMMASTREPHIDAE 

Arrow squids, flying squids 


Diagnostic characters: Medium to large oegopsid squid (to 

400 mm mantle length) with a strong, muscular torpedoshaped 

mantle. Large muscular fins generally rhomboidal but may 

be attenuated posteriorly to varying degrees. Lateral arms (II, III) 

strongly keeled in many species. Funnel-mantle locking apparatus 

⊥-shaped, sometimes fused with mantle and funnel elements. 

Suckers arranged biserially on arms and tetra-serially on tentacular 

clubs; chitinous sucker rings with sharp conical teeth.Buccal 

connectives attached to 

dorsal border of ventral 

arms (IV). Light organs 

present in some species 

on dorsal and ventral mantle, 

viscera, and ventrally 

on eyes and head. Mouth 

surrounded by 10 appendages 

(8 arms, 2 tentacles). mantle locking funnel locking 

One or both ventral arms cartilage 

cartilage 

hectocotylized in males, 

modifications vary from 

funnelmantle 

elaborate development and locking 

sculpturing of sucker bases apparatus 

and trabeculae to simple ⊥-shaped 

head 

loss of suckers and stalks 

funnel 

and development of protective 

membranes. Unlike 

most teuthoids which hatch 

as miniatures of the adults, 

ommastrephids have a distinct 

larval form, the rhynchoteuthion, 

in which the 

precursors to the tentacles 

mantle 

remain fused along their 

internal view of anterior 

length into a proboscis with 

mantle 

a terminal disc of suckers 

ventral view 

until the larva reaches (illustration: K. Hollis/ABRS) 

about 5 mm mantle length. 

Habitat, biology, and fisheries: In the Western Central Pacific, adult members of this family occur in deeper 

continental shelf, slope and throughout oceanic waters. Larvae and juveniles may also be found in shallow 

coastal waters. Ommastrephids are nektonic squid, associated with all depth strata from the surface to near the 

bottom.They are capable of extensive vertical and horizontal migrations.Ommastrephids feed on a broad range 

of crustaceans, fish, squid and other pelagic molluscs, and are at least partially cannibalistic. Known predators 

include seabirds, teleosts and sharks, whales, and dolphins. Ommastrephids form schools, which decrease in 

size as the animal grows. Evidence from commercial fishing operations suggests that neritic species congregate 

close to the bottom during the day, and move up through the water column at night. Oceanic species are often 

seen feeding at the surface at night. Juveniles of several species are able to glide like exocoetid flying fish 

to escape predators for distances in excess of 10 m. There is evidence for long-distance migration 

associated with spawning in some species. Ommastrephid squid are caught incidentally in demersal trawls 

in shelf and slope waters in the area. Also caught using baited or artificial lures or jigs, generally operated 

by hand. Exploratory use of jigging machines has been attempted for several species in the area with limited 

success. No target fisheries for these species currently exist in the area and regional catch statistics do 

not separate ommastrephids from other squids in most cases. The shelf and slope species in the area 

(Todarodes pacificus, Nototodarus hawaiiensis) are unlikely to represent large resources with the exception 

of the broader slope areas off northwestern and northeastern Australia and in the South China Sea. 

The resource of the oceanic ommastrephids (especially Sthenoteuthis oualaniensis) may be large and is 

probably underexploited.

Ommastrephidae 789 


Onychoteuthidae: also relatively large muscular squids with an 

arrow-shaped terminal fin, but can be distinguished by the following 

characters: funnel-mantle locking cartilage simple, straight; tentacular 

clubs with biserially arranged hooks and marginal suckers; 

buccal connectives attached to ventral border (rather than to dorsal 

border) of ventral arms. 

Key to the species of Ommastrephidae occurring in the area 

Note: most ommastrephid species are easily identified in the field 

as adults. However, ontogenetic development or loss of light organs 

and variability in sucker dentition with growth may make identification 

of specimens smaller than 80 mm mantle length more difficult 

and specimens should be preserved and referred to regional 

teuthologists for confirmation. 


Onychoteuthidae 

2 rows of 

hooks 

4 rows of 

suckers, 

no hooks 


Ommastrephidae 

1a. Light organs apparent externally on head and/or mantle (skin may need to be removed 

to observe dorsal mantle light organs) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. No light organs visible externally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

2a. A large oval light-organ patch present on dorsal mantle just posterior to head (Fig. 1a); 

funnel and mantle cartilages of locking apparatus fused (Fig. 1b) . . . Sthenoteuthis oualaniensis 1/ 

(form with a dorsal light-organ patch) 

2b. Light organs clearly visible on ventral mantle and head, no light organs evident on dorsal 

mantle; funnel and mantle cartilages of locking apparatus not generally fused . . . . . . . . . . → 3 

3a. Nineteen spherical light organs arranged in a distinctive pattern on ventral mantle 

(Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyaloteuthis pelagica 

3b. Light organs form 2 longitudinal stripes along the length of ventral mantle (Fig. 3) . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eucleoteuthis luminosa 


light 

organ 

patch 

funnel 

cartilage 

b) funnel cartilage fused 

with mantle cartilage 

Fig. 1 Stenoteuthis oualaniensis 

mantle 

cartilage 

ventral view 

spherical 

light 

organs 

Fig. 2 Hyaloteuthis pelagica 

(after Wormuth, 1976) 

ventral view 

light 

organs in 2 

longitudinal 

stripes 

Fig. 3 Eucleoteuthis luminosa 

1/ Two forms of Sthenoteuthis oualaniensis, the larger more abundant form with, and the smaller without, a dorsal 

light-organ patch, are known from the area. The former reaches in excess of 300 mm mantle length while females 

of the latter form reach maturity at less than 200 mm mantle length. It remains unresolved as to whether these 

represent separate species or whether onset of reproductive maturation at a small size triggered by unknown factors 

inhibits the formation of the light organ patch in the smaller form.


4a. Mantle very slender, with elongated posterior tail 

(Fig. 4) . . . . . . . . . . . . . . . . . . . .Ornithoteuthis volatilis 

4b. Mantle moderately stout, without an elongated tail 

(Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

5a. Large medial tentacular sucker rings with a large tooth 

in each quadrant, separated by smaller subequal teeth 

(Fig. 5a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

5b. Large medial tentacular sucker rings with equal-sized 

or subequal teeth or with a single larger tooth present 

distally (Fig. 5b, c) . . . . . . . . . . . . . . . . . . . . . . . . → 7 

6a. Two small round light organs on the intestine, one 

anteriorly near the anus and the other posteriorly 

(Fig. 6); ventral mantle uniformly coloured . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . Sthenoteuthis oualaniensis 2/ 

(form without a dorsal light-organ patch) 

6b. No light organs on the intestine, but small spherical 

sub-cutaneous light organs distributed broadly in the 

ventral mantle in larger specimens (visible in the mantle 

wall when cut midventrally); mantle with a midventral 

bronze stripe (Fig. 7) . . . . . . . . . . . . Ommastrephesbartramii 

7a. Medial tentacular sucker rings 

with approximately 30 subequal 

teeth (Figs 8a and 9) . . Todaropsis eblanae 

7b. Medial tentacular sucker rings 

with less than 25 teeth 

(Fig. 8a, c) . . . . . . . . . . . . . . . . → 8 

8a. Medial tentacular sucker rings 

with a single larger tooth distally 

(Fig. 8c) . . . . . . . . . . . . . . . . . → 9 

8b. Medial tentacular sucker rings 

with 16 to 20 subequal teeth 

(Figs 8b and 10) ........ 

. . . . . . . .Todarodes pacificus subspecies 

intestine 

funnel 

mantle 

internal view of anterior 

ventral mantle 

2 light 

organs 

Fig. 6 Sthenoteuthis oualaniensis 

2/ See footnote on previous page. 

a) with large tooth 

in each quadrant 

ventral view 

bronze 

stripe 

Fig. 7 Ommastrephes bartramii 

about 30 

teeth 

16-20 

subequal 

teeth 

ventral view 

tail 

enlongated, 

pointed 

Fig. 4 Ornithoteuthis volatilis 

b) with subequal 

teeth 

large tooth 

Fig. 5 tentacular sucker rings 

c) with single large 

tooth distally 

a) Todaropsis eblanae 

b) Todarodes pacificus 

large tooth 

distally 

c) Nototodarus hawaiiensis 

Fig. 8 medial tentacular 

sucker rings


9a. Arms with less than 30 transverse rows of biserial suckers; skin appears rough (Fig. 11) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nototodarus hawaiiensis 

9b. Arms with more than 35 transverse rows of biserial suckers; skin appears smooth 

(Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nototodarus gouldi 

dorsal view 

Fig. 9 Todaropsis 

eblanae 



Eucleoteuthis luminosa (Sasaki, 1915) 

Hyaloteuthis pelagica (Bosc, 1802) 

Nototodarus gouldi (McCoy, 1888) 

Nototodarus hawaiiensis (Berry, 1912) 

Ommastrephes bartramii (Lesueur, 1821) 

Ornithoteuthis volatilis (Sasaki, 1915) 

Sthenoteuthis oualaniensis (Lesson, 1830) 

Todarodes pacificus Steenstrup, 1880 

Todaropsis eblanae (Ball, 1841) 

dorsal view 

Fig. 10 Todarodes 

pacificus 


dorsal view 

Fig. 11 Nototodarus 

hawaiiensis 

ventral view 

Fig. 12 Nototodarus 

gouldi 

References 

Dunning, M.C. 1988a. Todarodes pacificus pusillus new subspecies (Cephalopoda, Ommastrephidae) from northern 

Australian waters. Mem. Natl. Mus. Vict., 49:149-157. 

Dunning, M.C. 1988b. First record of Nototodarus hawaiiensis (Berry, 1912) from northern Australian waters with a 

reconsideration of the identity of N. sloani philippinensis Voss, 1962. Mem. Natl. Mus. Vict., 49:159-168. 

Dunning, M.C. (in press). A review of the systematics, distribution and biology of the arrow squid genera Ommastrephes 

Orbigny, 1835, Sthenoteuthis Verrill, 1880, and Ornithoteuthis Okada, 1927 (Cephalopoda, Ommastrephidae). 

Smithson. Contrib. Zool. 

Dunning, M.C. and S.B. Brandt. 1985. Distribution and life history of deep-water squid of commercial interest from 

Australia. Aust. J. Mar. Freshwat. Res., 36:343-359. 

Dunning, M.C. and J.H. Wormuth. (in press). The ommastrephid squid genus Todarodes: a review of systematics, 

distribution and biology. Smithson. Contrib. Zool. 

Lu, C.C. 1982. First record of Todaropsis eblanae (Ball, 1841) (Cephalopoda: Oegopsida) in the Pacific Ocean. Venus, 

41:67-70. 

Lu, C.C. and M.C. Dunning. 1982. Identification guide to Australian arrow squid (Family Ommastrephidae). Tech. Rep. 

Vict. Inst. Mar. Sci., 2:1-30. 

Nesis, K.N. 1979. Squid of the family Ommastrephidae in the Australian - New Zealand Region. Tr. Inst. Okeanol., 

109:140-146 (in Russian). 

Wormuth, J.H. 1976. The biogeography and numerical taxonomy of the oegopsid squid family Ommastrephidae in the 

Pacific Ocean. Bull. Scripps Inst. Oceanogr., 23:1-90.


Nototodarus hawaiiensis (Berry, 1912) 

Frequent synonyms / misidentifications: Nototodarus philippinensis 

Voss, 1962; N. nipponicus Okutani and Uemura, 1973 / 

N. gouldi (McCoy, 1888). 

FAO names: En - Hawaiian flying squid; Fr - Encornet bouquet; 

Sp - Pota hawaiana. 

Diagnostic characters: Mantle cylindrical, moderately muscular, 

with abrupt caudal taper; fins short, rhombic. Head large, slightly 

wider than mantle. Mantle element of T-shaped locking apparatus 

with straight ridge, no muscular fusion to funnel element; 

funnel groove with foveola but no side 

pockets. Arms subequal and large, the 

longest slightly less than half the mantle 

length; swimming keels show greatest development 

on arms III; largest arm suckers 

with 9 to 16 conical teeth interspersed with 

low flat truncated teeth, distal tooth sig- 

nificantly larger; arms with 19 to 28 

transverse rows of suckers. Protective 

membranes and their supports of uniform 

height, not higher than suckers. Hectocotylization 

in males involving both ventral 

arms (IV); left ventral arm modified 

proximally by the enlargement of the first 4 

or 5 pairs of sucker bases as sculptured 

cushions; right ventral arm mirrors its partner 

proximally and shows modification of 

the distal ventral protective membrane supports 

and the sucker bases to form the 

equivalent of three columns of papillae; 

right ventral arm slightly shorter than its 

partner. Suckers of tentacular manus in 9 

to 11 tetraserial rows; largest medial manus sucker rings with 14 to 18 moderately large conical teeth 

interspersed with low horny plates, one distal conical tooth significantly larger. Fixing apparatus 

absent. Light organs absent in larvae, juveniles and adults. Colour: mantle and arms chestnut brown 

with a distinctly darker brown dorsal stripe along the mantle midline broadening over head and extending 

anteriorly as thin stripes along the arms. 

Size: Maximum mantle length 250 mm, commonly to 150 mm. 

Habitat, biology, and fisheries: Occurs predominantly in continental slope waters from depths of more 

than 200 m to at least 500 m although small juveniles and larvae have also been caught in adjacent shelf 

waters. Only occasionally taken on jigs in the area near the surface suggesting it is predominantly demersal 

rather than pelagic; alternatively, may not be as positively phototactic as most ommastrephids. Size at 

reproductive maturity seasonally variable in northern Australian populations; males generally mature from 

130 mm and females from 150 mm mantle length; occurrence of mature females and larvae indicates that 

spawning occurs throughout the year. Caught incidentally in demersal trawling in deep water and occasionally 

abundant in catches. No targeted fisheries. Suitable for human consumption and used as bait. 


deeper shelf and slope waters 

throughout the Indo-Pacific 

region from east Africa to Sri 

Lanka, northern Australia, 

Philippines, around Hong 

Kong (China), southern Japan, 

Hawaii, seamounts of the 

southeastern Pacific off Peru. 

? 


sucker ring 

arm III 

sucker ring 

(from Voss and Williamson, 1971) 

? 

dorsal view 

(after Voss, 1963)


Ommastrephes bartramii (Lesueur, 1821) 


Ommastrephes caroli stenodactyla Rancurel, 

1976 / Dosidicus gigas of Brazier, 1892. 

FAO names: En - Neon flying squid;Fr - Encornet 

volant; Sp - Pota saltadora. 

Diagnostic characters: Funnel groove with 

foveola and side pockets. Tetraserial suckers 

on dactylus of tentacular clubs; medial manus 

sucker rings with 1 tooth 

in each quadrant greatly 

enlarged; carpal fixing apparatus 

consisting of 

smooth-ringed suckers and 

knobs on the tentacular 

stalk. Small, irregularlyshaped, 

subcutaneous 

light organs present in 

adults, embedded in the 

ventral mantle and ventrally 

in the head; no light 

organs in larvae. Either left 

or right ventral arm (IV) hectocotylized 

by complete loss 

of suckers and sucker bases 

distally in mature males. 

Mantle-funnel locking apparatus 

not fused. 


sucker ring 

arm III 

sucker ring 

ventral view 

dorsal view 

Size: Maximum mantle length 700 mm (females) and 400 mm (males), commonly between 300 and 

450 mm mantle length. 

Habitat, biology, and fisheries: Ommastrephes bartramii occurs as mature adults and larvae around 

the northern Hawaiian Islands but is very rare between 25°N and the equator in the western North Pacific. 

In the southwest Pacific, adult O. bartramii have been caught between 23°42’ S and 45°45’ S where 

surface water temperatures varied from 14.2° to 25.7°C and larvae as far north as 24°S in East Australian 

Current waters. Adult O. bartramii are known to occupy a broad depth range both day and night from the 

surface to at least 1 500 m. An adult female in the northwest Pacific carrying an ultrasonic tag remained 

in the upper 100 m (mostly 40 to 70 m) during the night but stayed below 400 m depth during the day. 

The vertical distribution of larvae and juveniles remains poorly known. Male O. bartramii first reach 

maturity from 320 mm mantle length in the southwest Pacific. Considerable variation was evident in size 

of maturity of females in summer catches in this region. The smallest female observed with oviducal eggs 

measured 420 mm mantle length while the largest female with no eggs in the oviducts was 550 mm mantle 

length. An examination of the degree of development of the web on the ventrolateral arms in O. bartramii 

of both sexes suggests that growth of this structure is correlated with development of reproductive organs. 

Mature males and a female (520 mm mantle length) washed ashore near New Caledonia in October 1973, 

indicating that, in addition to off the central eastern Australian coast, spawning may be occurring in this 

region in spring. Its rarity in the area makes it of little importance as a target fisheries species. However, 

the northern and southern border regions of the Western Central Pacific may be important as a spawning 

ground and larval habitat for this species. 

Distribution: Ommastrephes bartramii is 

the dominant surface-dwelling oceanic 

ommastrephid circumglobally in subtropical 

and temperate oceanic waters 

and is only rarely encountered in continental 

slope waters and tropical latitudes. 

This bisubtropical species is the 

most widespread of all the ommastrephids. 

It is replaced as the dominant om- 

? 

mastrephid in tropical Indo-Pacific 

? 

waters by Sthenoteuthis oualaniensis.


Sthenoteuthis oualaniensis (Lesson, 1830) 


Symplectoteuthis oualaniensis (Lesson, 1830) / 

None. 

FAO names: En - Purpleback flying squid; 

Fr - Encornet bande violette; Sp - Pota cárdena. 

Diagnostic characters: Mantle cylindrical, very 

muscular, with abrupt caudal taper; fins large 

rhombic, with fin angle 60 to 70°. Head large, only 

slightly narrower than mantle. Mantle element of 

T-shaped locking apparatus curved with an 

anterior bifurcation, fused to funnel element 

along the posterior third of the longitudinal groove. 

Funnel groove with foveola and side pockets; 

tetraserial suckers on the dactylus of the tentacular 

club; medial manus sucker rings with 1 tooth in 

each quadrant greatly enlarged, 5 to 7 subequal 

teeth in between; carpal fixing apparatus 

consisting of smooth-ringed suckers and knobs on 

the tentacular stalk; 2 approximately equal-sized 

light organs present between the intestine and ink 

sac in larvae and juveniles, one anteriorly near the 

anus and the other posteriorly; single, oval light 

organ also present ventrally on each eye in larvae 

and juveniles; large dorsal light organ may be 

present in larger individuals anteriorly on the 

mantle. Either left or right ventral arm (IV) hectocotylized 

in mature males, 6 to 8 pairs of normal 

suckers proximally, sucker stalks and bases lost 

distally and pores developed in the thickened dorsal 

and ventral protective membranes basally and 

medially. 

Size: Maximum mantle length for the large form 

to about 300 mm in the area, commonly to about 200 mm mantle length (another larger form from the 

northwestern Indian Ocean reaches 650 mm mantle length). 

Habitat, biology, and fisheries: Adults generally occur only where bottom depth is greater than 200 m, larvae 

and juveniles occur also in clear, shallower water including around coral reefs. The more abundant large form 

with the dorsal light organ patch occurs in oceanic waters throughout the area, capable of extensive vertical 

migrations but also found in surface waters day and night, occurs in small schools (about 30 individuals) of like 

size. Males of the large form reach maturity at 110 mm mantle length, females at 180 mm mantle length but with 

seasonal and geographic variability; spawning appears to be geographically widespread in continental slope 

waters and occurring year-round;spawning migrations have been reported from around Taiwan Province of China. 

Males of the small maturing form reach maturity at 90 mm mantle length and females at 130 mm mantle length, 

spawning times and locations for this form unknown. Preys primarily upon small pelagic fishes and crustaceans, 

cooperatively hunting with other school members. Supports local fisheries around Nansei Islands (Okinawa) and 

Taiwan Province of China from Spring to Autumn, caught by hand jigging, mechanized jigging has been tried 

around Okinawa and Fiji without success, used for human consumption and for bait for tuna. 

Distribution: Tropical waters throughout the Indo-Pacific region, from Okinawa in the north to northern Australia and 

from East Africa to the eastern central Pacific. 

Remarks: S. oualaniensis as it is currently rec- 

ognized includes 2 forms in the Western Central 

Pacific, one with and the other without a large 

dorsal light organ patch on the anterior mantle 

in adults. The former reaches in excess of 

300 mm mantle length while females of the 

latter form reach maturity at less than 200 mm 

mantle length. The dorsal light organ patch develops 

in specimens from 65 mm mantle length 

and is readily visible at 100 mm mantle length. 

funnel 

2 light 

organs 

mantle 

intestine 

internal view of 

anterior ventral mantle 

(after Wormuth, 1976) 

funnel 

cartilage 

mantle 

funnel cartilage fused 

with mantle cartilage 

? 

? 

? 

arm III 

sucker ring 

dorsal view 

dorsal 

lightorgan 

patch 

(not 

always 

present) 


sucker ring


Todarodes pacificus Steenstrup, 1880 SQJ 

Frequent synonyms / misidentifications: Ommastrephes 

sloani pacificus Sasaki, 1929 / Nototodarus 

philippinensis Voss, 1962 (= N. hawaiiensis (Berry, 

1912)). 

FAO names: En - Japanese flying squid; Fr - Toutenon 

japonais; Sp - Pota japonesa. 

Diagnostic characters: Mantle cylindrical, slender, only 

moderately muscular, with abrupt caudal taper; fins 

short, rhombic. Head large, only slightly narrower than 

mantle. Mantle element of T-shaped locking apparatus 

with straight ridge, no muscular fusion to funnel element;funnel 

groove with foveola but no side pockets. 

Arms subequal and large, the longest slightly less than 

half the mantle length; swimming keels well developed; 

largest arm sucker rings with 9 to 11 sharp subequal 

somewhat flattened teeth; protective membranes and 

their supports of uniform height, not higher than suckers. 

Only right ventral arm (IV) hectocotylize in males; 

arm slightly thicker and shorter than its partner; 44 to 48 

normal arm suckers present proximally in T. pacificus 

pacificus, 11 to 13 in T. pacificus pusillus Dunning, 1988; 

approximately 20 pairs of trabeculae present in modified 

distal section forming an undulating spatula on the dorsal 

edge in both subspecies; in T. p. pacificus the hectocotylus 

represents about 30% of the arm length, in T. p. pusillus, it 

reaches 45 to 57%. Suckers of tentacular manus in 6 to 8 tetraserial 

rows; largest medial manus sucker rings with 16 to 18 moderately 

large subequal conical teeth interspersed with low horny plates. 

Light organs absent.Colour: dorsal mantle brown with a distinct deep 

blue-black dorsal stripe along the mantle midline broadening over head 

and extending anteriorly as thin stripes along aboral edges of dorsal and 

dorsolateral arms. 

Size: Maximum mantle length T. p. pacificus 500 mm, commonly 

300 mm in temped Japan, less than 200 mm off Hong Kong and in 

the South China Sea; T. p. pusillus from northern Australia reaches a 

maximum mantle length of less than 100 mm. 

Habitat, biology, and fisheries: Two subspecies are presently 

recognized: Todarodes pacificus pacificus Steenstrup, 1880, and 

Todarodes pacificus pusillus Dunning, 1988. Occurs in mid-shelf to 

slope waters in tropical latitudes, in shallow coastal to near oceanic 

waters in temperate latitudes. Populations around Japan are highly 

migratory and occur in large aggregations around oceanic fronts, seamounts and gyres where food is 

abundant. Feed primarily on small pelagic fishes (including anchovies, myctophids) and crustaceans; 

cannibalism is common. Predators include tunas, dolphins, and whales. Major mechanized jig fishery in 

the northwest Pacific, catches there have exceeded 600 000 t; also caught in coastal set nets and demersal 

trawls. Incidental trawl 

catches only in the Western 

Central Pacific. Large established 

markets in Japan, Korea, 

and Taiwan Province of 

China as fresh, frozen, processed, 

dried, and canned 

product. 


continental shelf and upper 

slope waters of the northwestern 

Pacific, from north of 

Japan to northern Australia. 

? 

? 

? ? 

right arm IV 

of male 


? 

? 

dorsal view 

Todarodes pacificus pusillus 


arm III tentacular club 

sucker ring sucker ring 

(after Voss and Williamson, 1971)


Todaropsis eblanae (Ball, 1841) 


FAO names: En - Lesser flying squid; Fr - Toutenon souffleur; 

Sp - Pota costera. 

Diagnostic characters: Mantle cylindrical, moderately muscular, 

relatively short with abrupt caudal taper; fins less than 50% mantle 

length but broad (about 90% mantle length), rhombic. Head large, 

slightly wider than mantle. Mantle element of T-shaped locking 

apparatus with straight ridge, no muscular fusion to funnel 

element; funnel groove without foveola or side pockets.Arms 

subequal and large, the longest more 

than half the mantle length; swimming 

keels show greatest development on 

arms III; largest arm suckers with about 

10 conical teeth on the distal margin, 

distal tooth only marginally larger. 

Protective membranes and their supports 

of uniform height, not higher than 

suckers. Hectocotylization in males 

involving both ventral arms (IV); 

suckers on proximal 20 to 30% of both 

arms lost and trabeculae modified into 

hard bracket-like structures; right ventral 

arm mirrors its partner proximally 

and shows modification of the protective 

membrane supports and the sucker 

bases to broad fan-like brackets as on 

the proximal portion; right ventral arm 

slightly shorter than its partner.Suckers 

of tentacular manus in 6 tetraserial 

rows; largest medial manus sucker 

rings with 30 or more subequal, regularly 

spaced small conical teeth. Fix- 

ing apparatus absent. Light organs 

absent. Colour: mantle and arms golden-brown 

with a darker brown dorsal 

stripe along the mantle midline. 

Size: Maximum mantle length to about 160 mm. 

Habitat, biology, and fisheries: Occurs predominantly in continental 

slope waters from depths of more than 200 m to at least 

800 m (temperatures 9 to 17.5°C) although small juveniles have 

also been caught in adjacent deeper shelf waters. Demersal 

rather than pelagic as adults. Males generally mature from 90 mm 

and females from 150 mm mantle length in northern Australian waters; spawning season is apparently 

protracted from midsummer to winter. Caught incidentally in demersal trawling in deep water; sometimes 

caught together with Nototodarus hawaiiensis and Todarodes pacificus spp. No targeted fisheries. Suitable 

for human consumption. 

Distribution: Occurs in the 

eastern Atlantic and Mediterranean, 

western Indian 

Ocean and Timor Sea, South 

China Sea, and off northeastern 

Australia in slope waters. 

? 

? 

left arm right arm 

arms IV of male 


? 

dorsal view 


arm III 

sucker ring 


sucker ring

Thysanoteuthidae 797 

Thysanoteuthidae THYSANOTEUTHIDAE 

A single species occurring in the area. 

Thysanoteuthis rhombus Troschel, 1857 

Rhomboid squids, diamondback squids 


Frequent synonyms / misidentifications: Thysanoteuthis 

nuchalis Pfeffer, 1912 / None. 

FAO name: En - Diamondback squid; Fr - Chipliloua commun; 

Sp - Chipirón volantín. 

Diagnostic characters: A large squid. Mantle thick, muscular, 

tapering to a blunt tip posteriorly. Fins rhombic, extending the 

entire length of mantle in adults. Funnel locking cartilage 

–|-shaped, with a long narrow longitudinal groove and a short 

broad transverse groove. Sharp toothed suckers arranged in 2 

rows on arms, and in 4 rows on tentacular clubs. Lateral arms 

(II, III) strongly keeled in large specimens. Long, cirrate trabeculae 

on arms support a well-developed web; buccal connectives 

attached to ventral border of ventral arms (IV). Left 

ventral arm hectocotylized in males. Light organs absent. 

Size: Maximum mantle length 850 mm, commonly to 600 mm 

mantle length; maximum weight 24 kg. 

Habitat, biology, and fisheries: The monotypic Thysanoteuthis 

rhombus is an oceanic species generally caught in the upper 50 m 

of the water column. Juveniles are apparently capable of leaping 

out of the water but do not “fly” (glide) in the same manner as 

funnel 

locking 

cartilage 

tentacular 

club 

dorsal view 

ommastrephids and onychoteuthids. Adults are slow swimmers and are often observed in monogomous, pairs 

(male and female) although groups of up to 20 have been observed elsewhere. It is preyed upon by yellowfin 

tuna, lancetfish, spotted dolphin, and blue marlin. Females spawn gelatinous, sausage-shaped egg masses, 

150 to 200 mm in diameter and up to 1 m long, which have been found near the surface. A spiral, double row 

of eggs of up to 2 mm diameter is contained in the external layers. Females are thought to be multiple spawners, 

the spawning season lasting perhaps a few months. In warm waters around Japan, spawning occurs during 

summer although in warmer tropical waters, it is thought to occur year-round. The smallest larvae hatch at 

1.1 mm mantle length as near replicas of the adults. Recent studies of age and growth of this species in the 

tropical Atlantic have shown them to be fast growers, reaching maturity (450 mm mantle length for males, 

600 mm mantle length for females) at less than 8 months of age; the oldest specimen examined (a 770 mm 

mantle length male) was 309 days old. T. rhombus is caught using drifting, sometimes baited jigs and set nets 

and is renowned for its tender flesh which is sometimes eaten as sashimi. 

Distribution: Occurs circumglobally in 

tropical and warm subtropical oceanic waters 

but nowhere abundant. 


Octopoteuthidae (Octopoteuthis, Taningia), 

Enoploteuthidae (Ancistrocheirus lesueuri): 

also have long rhomboid fins, but biserial 

hooks instead of suckers on the arms and 

tentacular clubs; in addition, all have obvious 

light organs on the ventral mantle, arm tips 

or internally. Unlike Thysanoteuthis, these other oceanic squid are rarely encountered near the surface. 

Loliginidae (Sepioteuthis lessoniana): has long fins extending the full length of the mantle but these are 

oval-shaped; also, the funnel-mantle locking apparatus is simple and straight, not –| -shaped. Sepioteuthis 

occurs only in continental shelf waters. 

Reference 

Nigmatullin, Ch.M., A.I. Arkhipkin, and R.M. Sabirov. 1995. Age, growth and reproductive biology of diamond-shaped 

squid Thysanoteuthis rhombus (Oegopsida: Thysanoteuthidae). Mar. Ecol. Prog. Ser., 124:73-87.


Chiroteuthidae CHIROTEUTHIDAE 

Chiroteuthid squids 


Diagnostic characters: Small to mediumsized 

squids (less than 400 mm mantle 

length) with somewhat gelatinous bodies. Characterized 

by an oval mantle-funnel locking apparatus, 

the funnel cartilage with 1 or 2 knobs 

projecting towards the centre of the cavity. 

Arms with biserial, toothed suckers; ventral 

pair (IV) greatly enlarged. Some species possess 

very elongate tentacles (up to 5 times the 

mantle length) and distinctive clubs with 

tetraserial suckers on long stalks. Buccal 

connectives attached to ventral border of ventral 

arms. Abundant light organs present along 

tentacle stalks and ventral arms; large light organs 

also present ventrally on the eyes, at the tips 

of the tentacular clubs, and embedded in the ink 

sac on both sides of the intestine. 

Habitat, biology, and fisheries: Although considered 

to be predominantly oceanic, Chiroteuthis 

imperator was taken recently in demersal 

trawls in continental slope waters at depths from 

300 to 600 m off northern Australia and is also 

caught in “jala-oras”, light-lured surround nets, in tentacular 

Indonesian waters. In deeper oceanic waters, club 

adult Chiroteuthis apparently occur below 500 m 

during the day but are distributed throughout the 

water column during the night. Diel vertical migration 

is also evident in some species. Nothing is 

known of the life history biology of these species. 

Chiroteuthids form part of the diet of lancetfish 

and yellowfin tuna in the tropical Indo-West Pacific 

waters. Off Japan, C. imperator feeds on 

micronektonic crustaceans, molluscs, and fish. 

Chiroteuthids have no commercial fisheries potential, 

due to their soft gelatinous bodies. funnel locking 

ventral view 

cartilage 


Mastigoteuthidae: also with enlarged ventral arms, but distinguished by the following characters: funnel 

locking cartilage with posterior knob and (occasionally) medial knob poorly developed; tentacles with many 

hundreds of small suckers, not tetraserially arranged. 


Asperoteuthis acanthoderma (Lu, 1977) 

Chiroteuthis imperator Chun, 1913 

Reference 

Kubota, T., M. Koshiga, and T. Okutani. 1981. Rare and interesting squid from Japan VII. Some biological data on 

Chiroteuthis imperator from Suruga Bay, Japan. Venus, 40:150-159.

Mastiogoteuthidae 799 

Mastiogoteuthidae MASTIGOTEUTHIDAE 

Mastigoteuthid squids 


Diagnostic characters: Medium-sized to large 

squids (500 to 1 000 mm mantle length), with gelatinous 

bodies. Funnel locking cartilage oval, with 

inward projecting knobs. Generally, posterior knob 

and (occasionally) medial knob poorly developed. 

Arms with biserial, toothed suckers; ventral arms 

(IV) enlarged. Males without hectocotylized arm. 

Characterized by long, whip-like tentacles, bearing 

many hundreds of minute suckers. Buccal connectives 

attached to ventral border of ventral arms. 

Fins large. Many species with light organs on surface 

of mantle, ventral surfaces of head, ventral arms, and 

eyeball. Some species possess minute dermal tubercles 

covering the body surface. Colour: body often 

pink or brick red coloured. 

Habitat, biology, and fisheries: Mastigoteuthids are 

deep living, oceanic squids occurring from tropical 

waters to the polar regions. Closing net data indicate 

that all species live at depths of 500 to 1 000 m during 

the day and may ascend to shallower water at night, 

even to as shallow as 50 to 100 m. The known predators 

of various species of Mastigoteuthis include 

Alepisaurus ferox, pilot whales, and sperm whales. 

Increased fishing effort using bottom trawls on the 

continental slopes of northern Australia in recent years 

has captured many mastigoteuthids including large 

specimens of Mastigoteuthis cordiformis up to 

700 mm mantle length. Of no commercial fisheries 

potential because of their gelatinous body consistency 

and the ammonia content in the mantle and arms. 

Remarks: The taxonomy of this family is in need of 

major revision. It is likely that several species occur in 

the area, but only adults of M. cordiformis have been 

many rows 

of suckers 

reported. 

dorsal view 

(Mastigoteuthis cordiformis) 



Chiroteuthidae: also with enlarged ventral arms, but distinguished by the following characters: funnel 

locking cartilage with posterior knob and medial knob well developed; tentacular suckers with long stalks, 

tetraserially arranged. 


Mastigoteuthis cordiformis Chun, 1908 

References 

Lu, C.C. and J.U. Phillips. 1985. An annotated checklist of the Cephalopoda from Australian waters. Occas. Pap. Mus. 

Vict., 2:21-36. 

Salcedo-Vargas, M.A. and T. Okutani. 1994. New classification of the squid family Mastigoteuthidae. Venus (Jap. J. 

Malacology), 53(2):119-127. 




Octopodidae OCTOPODIDAE 

Benthic octopuses 

by M.D. Norman 

Diagnostic characters: Bottom-living muscular octopuses 

with 8 arms, lacking tentacles. Eacharm 

possesses 1 or 2 rows of suckers. Fins absent. Rows of 

cirri adjacent to suckers absent. Internal shell absent or 

reduced to small rod-like stylets. One of third pair of arms 

8 arms, no tentacles 

IV 

II 

III 

(typically right-hand side) modified in mature males (the 

hectocotylized arm), consisting of a gutter along the 

margin of this arm (spermatophore groove) and a modified 

I 

tip (ligula) used to grip and pass spermatophores into the 

oviducts of the female. Funnel locking apparatus absent. 

Habitat, biology, and fisheries: Octopuses of the family 

Octopodidae are all bottom-dwelling and are found from 

intertidal waters down to abyssal depths (more than 

5 000 m). They occur on a wide range of substrates from 

coral and rocky reefs, to seagrass beds, sand, and mud. 

All brood their young, the female tending the eggs until 

hatching. The egg size in different species dictates the 

behaviour of the hatchlings. Species with small eggs 

(approximately 1 to 2 mm long) produce many tiny 

planktonic young which spend at least some time 

transported in the water column. Species with large eggs 

(10 to 30 mm long) produce few, large “crawl-away” young. 

Many octopus species have high fisheries profiles in the 

area, important in local and subsistence fisheries, as well 

as forming major export industries. They are collected in 

subtidal habitats by trawl, lure, and spear, and on intertidal 

reefs by hand or spear. The majority are harvested for 

human consumption with certain species collected 

right arm III 

mostly 


in males 

primarily as bait for finfish fisheries. They are marketed 

fresh, frozen, or dried. The reported yearly production of 

all octopods in the Western Central Pacific from 1990 to 

no fins 

1995 ranged from 20 023 t to 25 567 t (FAO Yearbook of 

Fishery Statistics). The actual annual total harvest in the 

area is likely to exceed 50 000 t. 

Notes on the taxonomy of Octopodidae 

dorsal view 

The taxonomy of this family is in a very poor state. There are a large number of undescribed or 

poorly-defined species (more than 80) occurring in the area, many of which form the basis of local and 

commercial fisheries. The majority of named species are placed in the catch-all genus Octopus, which 

currently contains over 200 nominal species. The limited existing literature frequently uses inappropriate 

species names, including several European names for species restricted to the Atlantic Ocean (e.g. 

Octopus macropus, O. vulgaris). 

There is negligible information available on biology, distribution or importance to fisheries for all but a 

handful of species. The list for the Western Central Pacific region, presented below is preliminary, including 

better known species and only undescribed species of high profile. It excludes species names coined from 

the area for which there is inadequate original descriptions or insufficient reference material to enable 

identification (i.e. dubious taxa). 

A further complicating factor in the taxonomy of benthic octopuses is the number of species groups within 

the broad genus Octopus (as it currently stands). Each of these groups contains similar species, often 

difficult to distinguish in the field. The knowledge of members of 3 groups in particular is very poor. The 

least resolved species group is the “drop-arm” octopuses, the Octopus horridus group. Members are found 

throughout the tropical Indo-West Pacific region and are characterized by small size, long arms (4 to 10 

times mantle length), the capacity to sever (“autotomize”) arms at the base (used as a wriggling decoy to 

predators) and complex skin sculpture producing accurate camouflage against coral, rubble, or algae. They 

typically occur on intertidal reefs where they are active during daytime low tides. Named species in the 

area include Octopus abaculus and O. aculeatus (treated below in species accounts). More than 12 species 

occur throughout the area and the limits to their distributions are unknown. Western Central Pacific species

Octopodidae 801 

are frequently misidentified under the names Octopus horridus (a large-egg species restricted to the Red 

Sea and western Indian Ocean) and O. defillipi (a species of a different group restricted to the 

Mediterranean Sea and Atlantic Ocean). See Norman and Sweeney (1997) for discussion of this species 

group. 

The second group of octopuses is the Octopus macropus group. These animals are all moderate to large 

octopuses (up to 3 kg and more than 1 m in total length) characterized by longer and more robust dorsal 

arms, high gill counts (10 or more lamellae per demibranch), simple colour patterns of red and white, and 

nocturnal foraging behaviour. Members occur throughout tropical and temperate waters of the world. 

Named species include Octopus alpheus, O. aspilosomatis, O. dierythraeus, O. graptus, O. luteus, O. 

nocturnus, and O. ornatus (all treated below). More than 5 undescribed species also occur in the area 

covered here and the limits to their distributions are unknown. Members of this group are frequently treated 

incorrectly under the name Octopus macropus, an European member of this group restricted to the 

Mediterranean Sea and Atlantic Ocean. See Norman (1993a) and Norman and Sweeney (1997) for 

discussion of this species group. 

A suite of pygmy species also occur in the area (more than 10 species), mature at under 1 g and with a 

mantle length less than 30 mm. These tiny species have low gill counts (5 to 7 lamellae per demibranch), 

short arms (1.5 to 3 times mantle length) and relatively few suckers (less than 150 suckers on each normal 

arm). Named species include Octopus bocki and O. wolfi (not treated here). Many pygmy species 

throughout the Indo-West Pacific region are undescribed. See Norman and Sweeney (1997) for treatment 

of pygmy octopuses from the Philippines. 

Major revisions of octopus taxonomy have been carried out by Robson (1929), Sasaki (1929) and Nesis 

(1987). 

Octopuses of deeper waters 

When trawling in deeper waters (more than 200 m), some genera of octopuses may be collected which 

are not normally encountered in shallower waters. These include species of the poorly-known genera 

Scaeurgus (characterized by a left hand hectocotylized arm in males and a lateral skin ridge around the 

mantle), Benthoctopus (smooth-skinned pale octopuses with large eyes and no ink sac) and Berrya 

(muscular octopuses with loose soft skin, a narrow opening to the mantle cavity and 2 long digit-like papillae 

over each eye). Eledone palari may also be encountered, and is easily distinguished in that it possesses 

a lateral skin ridge on the mantle and a single row of suckers on each arm (compared with 2 rows in all 

other members of the family Octopodidae in the area). 


Note: the order Octopoda, as it currently stands, contains 2 distinct groups of octopuses: finned “cirrate” 

octopuses (suborder Cirrata) and the more familiar “incirrate” octopuses (suborder Incirrata). The latter 

includes the family Octopodidae. 

Opisthoteuthidae and Cirroteuthidae 

(suborder Cirrata): cirrate octopuses are 

soft, semi-gelatinous animals of deeper 

waters easily distinguished from incirrate 

octopuses (including the family 

Octopodidae) by paired fins on the mantle, 

deep webs and rows of sensory papillae 

(“cirri”) adjacent to the suckers. Cirrate 

octopuses are rarely captured and, due to 

their soft flesh, are of no economic value. 

Argonautidae (suborder Incirrata): the 

“argonauts” are muscular pelagic octopuses 

(genus Argonauta) easily distinguished from 

members of the Octopodidae. Female 

argonauts have expanded webs on the ends 

of the dorsal arms which secrete a brittle 

white shell, the “paper nautilus” shell. This 

shell is used as an egg case in which the 

eggs are brooded. The tiny male argonaut 

lacks a shell and has a detachable third left 

arm in a pouch. A funnel locking apparatus 

paired fins 

 

is present as a lug-and-socket type. 

Ophisthoteuthidae 

Argonautidae


Ocythoidae (suborder Incirrata): contains a single species, the muscular pelagic octopus, Ocythoe 

tuberculata, in which the large female possesses a network of collagen-like ridges in the ventral mantle. 

The tiny male lacks the collagen sculpture and has a detachable third right arm in a pouch. A funnel locking 

apparatus is present as a lug-and-socket type. 

Tremoctopodidae (suborder Incirrata): contains a single genus and at least 2 species of muscular pelagic 

“blanket octopuses”, the best known being Tremoctopus violaceus. Females possess greatly expanded 

webs along length of dorsal arms, capable of being shed in segments from the tips. The tiny male lacks 

the dorsal webs and has a detachable third right arm in a pouch. A funnel locking apparatus is present as 

a transverse flap and shelf. 

Alloposidae (suborder Incirrata): contains a single species, the deep-water semi-gelatinous octopus, 

Haliphron atlanticus (frequently treated in the literature under the junior synonym, Alloposus pacificus). 

Little is known of this octopus other than the male possesses a detachable third right arm in a pouch. A 

funnel locking apparatus is present as a transverse flap and shelf. 

reticulate 

collagen 

sculpture 

Ocythoidae 

 

expanded web 

Alloposidae 

Families Amphitretidae, Bolitaenidae and Vitreledonellidae (suborder Incirrata): These 3 related families 

of rarely encountered open-water pelagic octopuses are distinguished from members of the family 

Octopodidae by their gelatinous consistency and a range of internal characters including distinctive radula 

morphology, the nature of the mantle aperture (double opening in Amphitretidae) and distinctive floorplans 

to the digestive and reproductive tracts. 

 

Tremoctopodidae 

Amphitretidae Bolitaenidae Vitreledonellidae


Key to the species of Octopodidae occurring in the area 

Remarks on key characters: measurements and sucker counts used below refer to submature and mature 

animals, not juvenile material. In interpreting order of arm lengths, partial arm regeneration can produce 

confusing combinations. A distinct decrease in sucker diameter along the arms is an indication of partial 

regeneration from that point. 

1a. Dorsal arms distinctly longer and frequently more robust than lateral and ventral arms 

(Fig. 1a); dorsal webs deeper than ventral webs . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Arms subequal (Fig. 1b) or lateral arms longer and more robust than dorsal arms 

(Fig. 1c); webs subequal in depth or dorsal webs shallower than other webs . . . . . . . . . . → 9 

dorsal arms longer 

ventral 

arms 

2a. Water pouches present between 

bases of arms on oral surface of webs 

(Fig. 2), visible as pores open to exterior 

at level of 3 rd to 4 th proximal sucker 

(pores difficult to find in contracted or 

smaller preserved specimens); ligula 

tiny (less than 1% of arm length), calamus 

absent . . . . . . . . . . . . Cistopus indicus 

2b. Water pouches and pores absent; 

ligula in submature and mature males 

well developed, cylindrical with deep 

groove and a distinct calamus . . . . 

. . . . . . . . . . .(Octopus macropus group) → 3 

(only 7 common members treated here of more than 12 

species in the area) 

3a. Colour pattern of pale cream to pink 

base with irregular short scribbles on 

dorsal mantle and arm crown (Fig. 3a) 

. . . . . . . . . . . . . . . . . . . Octopus graptus 

3b. Colour pattern of numerous white or 

red spots; dark scribbles absent . . . . . . . . → 4 

4a. Colour pattern on dorsal mantle of distinctive 

arrangement of cream to white 

longitudinal stripes over pink to red 

base colour (Fig. 3b) . . . . . . . Octopus ornatus 

4b. Dorsal mantle plain or with spots, longitudinal 

stripes absent . . . . . . . . . . . . → 5 

all arms subequal in length lateral arms longer 

a) b) c) 


dorsal 

arms 

water pouch 

Fig. 2 Cistopus indicus (oral view) 

pore 

a) Octopus graptus b) Octopus ornatus 

Fig. 3 dorsal mantle and arm crown


5a. Distinctly elongate species with almost tubular arms and short webs (Fig. 4a); mantle 

greatly elongated in many specimens (Fig. 4b); deepest web always less than 15% of 

length of longest arm, typically 10% . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

5b. Moderately elongate species with robust arms and deep webs (Fig. 5); mantle never greatly 

elongated; deepest web always more than 15% of length of longest arm, typically 20% . . . . . . → 7 

6a. Colour pattern on dorsal mantle plain red or white, lacking distinctive spots (Fig. 6a) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Octopus aspilosomatis 

6b. Colour pattern on dorsal mantle of white spots over orange to deep red base colour 

(Fig. 6b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Octopus nocturnus 

a) dorsal view b) elongate mantle 

Fig. 4 Octopus aspilosomatis 

dorsal view 

Fig. 5 Octopus dierythraeus 

a) Octopus 

aspilosomatis 

7a. More than 230 suckers on normal arms of submature and mature animals; more than 100 

suckers on hectocotylized arm of males; alarm display in live animals of large red spots over 

white base colour on dorsal mantle, arm crown, and arms (Fig. 5) . . . . . . . . .Octopus dierythraeus 

7b. Typically less than 230 suckers on normal arms of submature and mature animals; less 

than 100 suckers on hectocotylized arm of males; alarm display in live animals of white 

spots over red base colour on dorsal mantle, arm crown, and arms (as in O. nocturnus, 

Fig. 6b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

8a. Eggs large (more than 8 mm) and produced in low 

numbers (less than 500) (known only from intertidal reef 

flats of the Capricorn Bunker Islands of the southern 

Great Barrier Reef, Australia) . . . . . . . . . . . . Octopus alpheus 

8b. Eggs small (less than 4 mm) and produced in high 

numbers (more than 10 000) (known at this stage only 

from the Philippine Islands) . . . . . . . . . . . . Octopus cf. luteus 

9a. Arms greatly elongated (typically 7 to 9 times mantle 

length), subequal in length when intact; regular alternating 

white and pink-brown wide bands present on arms 

(Fig. 7); mantle walls very thin, almost transparent, branchial 

hearts visible through mantle walls (Fig. 7); arm 

autotomy present; ink sac absent . . . . . . . . Ameloctopus litoralis 

9b. Arms short to long, dorsal arms always slightly shorter 

than other arms; regular alternating white and pink-brown 

bands absent; mantle walls muscular, never thin and 

transparent; arm autotomy present or absent; ink sac 

present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

b) Octopus 

nocturnus 

Fig. 6 dorsal mantle and arm crown 

white and 

pink-brown 

bands 

branchial 

heart 

dorsal view 

Fig. 7 Ameloctopus litoralis


10a. Small animals with mantle and arm crown covered in rings or lines (Fig. 7), iridescent 

blue in live animals; arms short, 1.5 to 3 times mantle length . . . . . . . . (Hapalochlaena) → 11 

10b. Iridescent tissue absent or, if present, restricted to a pair of false eye-spots (“ocelli”) on 

the arm crown over bases of arms II and III, one below each eye (as in Fig. 13a); arms 

short to long (more than 2 times mantle length) . . . . . . . . . . . . . . . . . . . . . . . . . → 13 

11a. Iridescent blue lines on dorsal and lateral mantle; rings on arm crown and arms (Fig. 8a) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hapalochlaena fasciata 

11b. Iridescent blue lines never present; small or large iridescent blue rings on all dorsal 

surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 12 

12a. Large rings (up to 12 mm in diameter, up to 40% of mantle length) over dorsal and lateral 

surfaces of mantle, head, arm crown, and arms (Fig. 8b) . . . . . . . . . Hapalochlaena lunulata 

12b. Small rings (up to 2 mm in diameter, approximately 5% of mantle length) over dorsal and 

lateral surfaces of mantle, head, arm crown, and arms (Fig. 8c) . . . . . Hapalochlaena cf. maculosa 

a) Hapalochlaena fasciata b) Hapalochlaena lunulata 


c) Hapalochlaena cf. maculosa 

(after Roper and Hochberg, 1988) 

13a. Raised keel (lateral ridge) around lateral mantle (Fig. 9); swollen club-like ligula in 

mature males (Fig. 10a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Octopus australis 

(species of the deep-water genus Scaeurgus appear similar to this species but are distinguished by a left-hand 

hectocotylized arm in males) 

13b. Lateral ridge absent; ligula small to spear-like (Fig. 10b), never as a swollen club . . . . . . . → 14 

lateral 

ridge 

lateral mantle 

Fig. 9 Octopus australis 

(from Stranks and Norman, 1993) 

ligula club-like 

a) Octopus australis b) other Octopus species 

Fig. 10 tip of hectocotylized arm of male 

(from Stranks and Norman, 1993) 

ligula 

small 

spearlike


14a. Small, elongate octopuses (mantle length to 70 mm, 

weight to 150 g); arms long, more than 4 times mantle 

length; arms sever at set level near arm base (arm autotomy), 

as a decoy to predators; skin typically highly sculptured 

including raised branched papillae (Fig. 11) . . . . 

. . . . . . . . . . . . . . . . . . . . . (Octopus horridus group) → 15 

(only 2 distinct members treated here of more than 10 species in the area) 

14a. Small to large robust octopuses; arms typically less than 

4 times mantle length; arms not capable of autotomy at set 

level near arm base; skin smooth to sculptured . . . . . . . . . . → 16 

15a. Colour pattern of dark brown to black circular reticulations 

which define a mosaic of distinct round cream spots 

(Fig. 12a); sucker counts approximately 90 to 120 on hectocotylized 

arm; ligula of moderate size (approximately 5% 

of arm length) . . . . . . . . . . . . . . . . . . . . . Octopus abaculus 

15b. Colour mottled grey-brown to dark grey on dorsal and 

lateral surfaces (Fig. 12b); sucker counts approximately 

140 to 175 on hectocotylized arm; ligula small (less than 

2% of arm length) . . . . . . . . . . . . . . . . . . Octopus aculeatus 

16a. Large robust species (mantle length more than 140 mm, 

weight more than 1 kg), moderate to long arms (more than 

3 times mantle length), more than 200 suckers on normal 

Fig. 11 typical representative 

of the Octopus horridus group 

arms, more than 140 on hectocotylized arm of males . . . . . . . . . . . . . . . . . . . . . . → 17 

16b. Small to moderate-sized species (mantle length less than 120 mm, weight less than 

500 g), short to moderate length arms (2 to 3 times mantle length), less than 200 suckers 

on normal arms, less than 110 on hectocotylized arm of males . . . . . . . . . . . . . . . . . → 18 

17a. False eye-spot (ocellus) present (Fig. 13a), as a dark oval spot surrounded by pale band 

of skin (not iridescent) and narrow dark outer band (Fig. 13b); dark zebra bars on 

lateroventral surfaces of all arms in submature and mature animals (Fig. 13c); sucker 

counts around 400 to 500 on normal arms, 180 to 230 on hectocotylized arm . . . Octopus cyanea 

17b. Ocellus and zebra bars on ventral faces of arms absent; sucker counts 200 to 300 on 

normal arms, 140 to 160 on hectocotylized arm . . . . . . . . . . . . . . . . . . . Octopus tetricus 

a) Octopus abaculus b) Octopus aculeatus 

Fig. 12 dorsal mantle and arm crown 

a) whole animal 

2 ocelli 

Fig. 13 Octopus cyanea 

b) detail of ocellus 

dark zebra 

bars 

c) lateroventral 

surface of arm


18a. Ocelli (“false eye-spots”) present, 1 on each lateral face of arm crown between bases 

of arms II and III (position shown in Fig. 13a) . . . . . . . . . . . . . . . . . . . . . . . . . . → 19 

18b. Ocelli absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 23 

19a. Ocellus plain, without iridescent ring, as plain black spot (Fig. 14a) . . . . . . Octopus exannulatus 

19b. Ocellus with iridescent ring within black spot (Fig. 14b) . . . . . . . . . . . . . . . . . . . . . → 20 

20a. Widely-spaced dark transverse bars on all arms separated by approximately 3 to 5 

suckers (Fig. 15a); mature males with 1 to 3 enlarged suckers (12 th to 14 th )onallarms 

atlevelofedgeofweb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Octopus polyzenia 

20b. Dark longitudinal lines down leading edge of arms I to III (Fig. 15b), never transverse 

bars; enlarged suckers of mature males (if present) on arms II and III (never I) . . . . . . . . → 21 

a) Octopus exannulatus 

21a. Circular cluster of dark spots above 

each eye forming petals of “flower” pattern 

(Fig. 16a); 4 longitudinal rows of 

dark spots (as solid stripes in some 

colour patterns) on dorsal body; large 

digit of skin in white spot on dorsal arm 

crown anterior to eyes; approximately 

100 suckers on hectocotylized arm of 

b) 

Fig. 14 detail of ocellus 

iridescent 

ring 

males . . . . . . . . . . . . . . . . Octopus mototi 

21b. Base colour of whole animal creambrown 

with short black longitudinal bar 

lines 

through each eye (Fig. 16b); pedal of 

of 

dark 

black spots on head and longitudinal 

spots 

stripes on dorsal mantle absent; large 

digit of skin anterior to eyes absent; 65 

to 80 suckers on hectocotylized arm of 

males . . . . . . . . . . . . . . . . . Octopussp. A 

IV 

I 

II III 

dark 

longitudinal 

lines 

dark 

transverse 

bars 

a) Octopus polyzenia b) Octopus sp. A 

Fig. 15 right arms I to IV 

circular 

cluster 

of dark 

spots 

I 

black bar 

through eye 

digit of 

skin in 

white 

spot 

IV II III 

a) Octopus mototi b) Octopus sp. A 

Fig. 16 dorsal view


22a. Pale longitudinal stripe present 

along midline of dorsal mantle 

(Fig. 17); pattern of circular 

cream to green spots bound by 

dark boundaries over all dorsal 

and lateral mantle and arm 

crown (Fig. 17), distinct on lateral 

faces of arms I to III . .Octopus aegina 

22b. Longitudinal stripe along dorsal 

midline of mantle absent; dark 

purple/black longitudinal stripe 

along lateral faces of arms I to 

III, in marked contrast with white 

or pink suckers (Fig. 18a); body 

colour variable from pale purplebrown 

to deep purple-black; purple 

black grooves between 

patches on lateral arm crown 

form distinctive branching reticulations 

(Fig. 18a); distinct 

white triangle under eye 

(Fig. 18b) . . . . Octopus marginatus 


The symbol is given when species accounts are included. Distribution Harvest 

Ameloctopus litoralis Norman, 1992b Northern Australia none 

Cistopus indicus (Rapp, 1835) SE Asia to India high 

Hapalochlaena fasciata (Hoyle, 1886) Australia, southern Qld and NSW 

Hapalochlaena lunulata (Quoy and Gaimard, 1832) Indo-Malayan Arch. to Vanuatu 

Hapalochlaena cf. maculosa (from Roper and Hochberg, 1988) Great Barrier Reef 

none 

none 

none 

Octopus abaculus Norman and Sweeney, 1997 

Octopus aculeatus d’Orbigny, 1835 

Octopus aegina Gray, 1849 

Philippines 

Philippines 

none known 

low 

1/ 

Octopus alpheus Norman, 1993a 

Octopus aspilosomatis Norman, 1993a 

Octopus australis Hoyle, 1885 

Octopus bocki Adam, 1945 

Octopus cyanea Gray, 1849 

Octopus dierythraeus Norman, 1993a 

Octopus exannulatus Norman, 1993b 

Octopus graptus Norman, 1993a 

Octopus cf. luteus (from Norman and Sweeney, 1997) 

Octopus marginatus Taki, 1964 

Indian Ocean and SE Asia 

Southern Great Barrier Reef 

Northern Great Barrier Reef 

Australia, southern Qld and NSW 

Tropical Pacific 

Tropical Indo-West Pacific 

Northern Australia 

Indo-Malayan Archipelago 


Philippines 

high 

none 

none 

low 

none 

moderate 

none 

low 

low 

moderate 

2/ 

Octopus mototi Norman, 1993b 

Octopus nocturnus Norman and Sweeney, 1997 

Octopus ornatus Gould, 1852 

Octopus polyzenia Gray, 1849 

Octopus pumilus Norman and Sweeney, 1997 

Octopus tetricus Gould, 1852 

Indian Ocean and SE Asia 

Tropical Pacific 

Philippines 



Philippines 

low 

none 

low 

low 

none 

none known 

3/ 

Octopus tonganus Hoyle, 1885 

Australia, southern Qld and NSW 

Tonga 

low 

none known 

Octopus vitiensis Hoyle, 1885 Tropical Pacific none known 

Octopus wolfi Wülker, 1913 

Octopus sp. A 


Indo-Malayan Archipelago 

none known 

very high 

pale 

stripe 

Fig. 17 Octopus aegina 


b) lateral view 

of head 

Fig. 18 Octopus marginatus 

1/ The genuine O. aegina is treated in Roper et al. (1984) under the name of a junior synonym, O. dollfusi. 

2/ O. marginatus is treated in Roper et al. (1984) incorrectly under the name of the related O. aegina. 

3/ The species treated under the name O. tetricus in Roper et al. (1984) is not identical with O. tetricus described originally 

from eastern Australia.


References 

Chotiyaputta, C. 1993. Cephalopod resources of Thailand. In Recent advances in fisheries biology, edited by T. Okutani, 

R.K. O’Dor, and T. Kubodera. Tokyo, Tokai University Press, pp. 71-80. 

Joll, L. M. 1983. Octopus tetricus. In Cephalopod life cycles. Volume 1. Species accounts, edited by P.R. 

Boyle, pp. 325-334. New York, Academic Press, 475 p. 

Lu, C. C. and T.N. Stranks. 1992. Eledone palari, a new species of octopus (Cephalopoda: Octopodidae) from Australia. 

Bull. Mar. Sci., 49(1-2):73-87. 

Nesis, K. N. 1987. Cephalopods of the world: Squids, cuttlefish, octopuses and allies. Neptune City, New Jersey, TFH 

Publications, 351 p. 

Norman, M. D. 1992a. Octopus cyanea Gray, 1849 (Mollusca: Cephalopoda) in Australian waters: Description, distribution 

and taxonomy. Bull.Mar.Sci., 49(1-2):20-38. 

Norman, M. D. 1992b. Ameloctopus litoralis gen. et sp. nov. (Cephalopoda: Octopodidae), a new shallow-water octopus 

from tropical Australian waters. Invert. Taxonomy, 6:567-582. 

Norman, M. D. 1993a. Four new species of the Octopus macropus group (Cephalopoda: Octopodidae) from the Great 

Barrier Reef, Australia. Mem. Natl. Mus. Vict., 53(2):267-308. 

Norman, M. D. 1993b. Ocellate octopuses (Cephalopoda: Octopodidae) of the Great Barrier Reef, Australia: description 

of two new species and redescription of Octopus polyzenia Gray, 1849. Mem. Natl. Mus. Vict., 53(2):309-44. 

Norman, M. D. 1993c. Octopus ornatus Gould, 1852 (Cephalopoda: Octopodidae) in Australian waters: morphology, 

distribution and life history. Proc. Biol. Soc. Wash., 106(4):645-60. 

Norman, M. D. and F.G. Hochberg. 1994. Shallow-water octopuses (Cephalopoda: Octopodidae) from Hong Kong’s 

territorial waters. In The Malacofauna of Hong Kong and Southern China III, edited by B. Morton. Proceedings 

of the third international workshop on the malacofauna of Hong Kong and southern China. Hong Kong, Hong 

Kong University Press, pp. 141-59. 

Norman, M. D. and M.J. Sweeney. 1997. The shallow-water octopuses (Cephalopoda: Octopodidae) of the Philippines. 

Invert. Taxonomy, 11:89-140. 

Pickford, G. E. 1974. Cistopus indicus (Orbigny): a common Indo-Malayan species of octopus. J. Mar. Biol. Assoc. Ind., 

16(1):43-8. 

Robson, G. C. 1929. A monograph of the recent Cephalopoda. Part I. Octopodinae. London, British Museum, 236 p. 

Roper, C. F. E. and F.G. Hochberg. 1988. Behavior and systematics of cephalopods from Lizard Island, Australia, based 

on colour and body patterns. Malacologia, 29(1):153-93. 

Roper, C. F. E., M.J. Sweeney, and C.E. Nauen. 1984. FAO species catalogue. Volume 3. Cephalopods of the world: An 

annotated and illustrated catalogue of species of interest to fisheries. FAO Fish. Synop., 125(3):277 p. 

Sasaki, M. 1929. A monograph of the dibranchiate cephalopods of the Japanese and adjacent waters. J. Coll. Agric., 

Hokkaido Imp. Univ., (20) (supplement):357 p. 

Stranks, T. N. and M.D. Norman. 1993. Review of the Octopus australis complex from Australia and New Zealand, with 

description of a new species (Mollusca: Cephalopoda). Mem. Natl. Mus. Vict., 53(2):345-373 (1992). 

Van Heukelem, W. F. 1983. Octopus cyanea. In Cephalopod life cycles. Volume 1. Species accounts, edited by P.R. 

Boyle, pp. 267-276. New York, Academic Press, 475 p. 

Voss, G. L. 1963. Cephalopods of the Philippine Islands. Bull. U.S. Natl. Mus., 234:1-180. 

Voss, G. L. 1981. A redescription of Octopus ornatus Gould, 1852 (Octopoda: Cephalopoda) and the status of 

Callistoctopus Taki, 1964. Proc. Biol. Soc. Wash., 94(2):525-534. 

Voss, G. L. and G.R. Williamson. 1972. Cephalopods of Hong Kong. Hong Kong, Hong Kong Government Press, 138 p.


Cistopus indicus (Rapp, 1835 in Férussac and d’Orbigny, 1834-48) 

Frequent synonyms / misidentifications: None / Octopus macropus 

Risso, 1826. 

FAO names: En - Old woman octopus (from Chinese “Laai Por”); Fr -Poulpe 

vieille femme; Sp - Pulpo perforado. 

Diagnostic characters: Large species. Water pouches present, embedded 

in the web around the mouth, one in each sector between arms. 

Pouch opens through a “water pore” situated at the level of third or fourth 

sucker from the mouth. Arms long, 6 to 7 times mantle length. Dorsal arms 

longer than ventral arms (arm formula I.II.III.IV). Right third arm of 

males hectocotylized with tiny blunt ligula (less than 1% of arm length) 

and no apparent calamus. Gills with 9 to 11 lamellae per demibranch. In 

larger animals, around 120 to 190 suckers on each normal arm, 60 to 130 

on hectocotylized arm of male (see comments below). One or 2 enlarged 

suckers (11 th to 13 th ) sometimes visible in mature males on arms I, II, and 

IV, but never on arm III. Colour: pink or cream, to slate grey with iridescent 

purple sheen on lateral mantle in fresh specimens; skin sculpture consisting 

of a few scattered low papillae on the dorsal mantle; no large primary 

papillae. 

Size: Maximum mantle length 180 mm, total length to over 1 m; weight to 

2kg. 

Habitat, biology, and fisheries: Known from muddy coastal waters, living 

subtidally on soft substrates to depths of at least 80 m. Nothing known of 

diet or foraging behaviour. No mature females found. The function of the 

unique water pores is not known. There may be more than 1 species 

treated under this species name. Specimens from Thailand and Singapore, 

and west to India appear to show lower sucker counts (around 110 

to 140 on normal arms, less than 80 on hectocotylized arm of males) and 

distinctly enlarged suckers in males, compared with animals from further 

north and east (180 to 200 suckers on normal arms, 110 to 130 suckers 

on hectocotylized arm of males, and enlargement of suckers in males 

slight or absent). These octopuses form major fisheries, 

important in subsistence, local small-scale 

and larger commercial fisheries in many coastal 

Asian countries, harvested primarily by trawlers. 

Distribution: Cistopus indicus, as it currently 

stands, occurs in tropical and subtropical coastal 

waters of the Asian mainland from China, the Philippines, 

and northern Indonesia, south to Malaysia 

and west to Pakistan. 

tip of hectocotylized 

arm of male 

pore 

dorsal view 

oral view 

water 

pouch


Octopus aegina Gray, 1849 

Frequent synonyms / misidentifications: Octopus dollfusi 

Robson, 1828; O. hardwickei Gray, 1849 / None. 

FAO names: En - Marbled octopus; Fr - Poulpe nain; Sp -Pulpo 

marmóreo. 

Diagnostic characters: Moderate-sized robust species. 

Arms relatively short, 2 to 3 times mantle length. Lateral and 

ventral arms longest (arm formula IV=III.II.I). Web deep on 

lateral arms, very shallow between dorsal arms. Right third 

arm of males hectocotylized with moderate length sharp ligula 

(4 to 6% of arm length) with distinct calamus. Gills with 8 or 9 

lamellae per demibranch. In larger animals, around 110 to 130 

suckers on each normal arm, around 60 to 70 on hectocotylized 

arm of male. Mature males with 2 or 3 enlarged 

suckers (6 th to 8 th )onarmsII and III (slightly enlarged on arms 

IV). Colour: pattern of reticulations formed by dark 

grooves defining large, and smaller intermediate, circular 

patches; reticulated pattern most distinctive on dorsal 

arm faces; cream coloured longitudinal stripe along dorsal 

midline of mantle; cream transverse head bar visible 

in many specimens; skin sculpture of regular round patches 

and grooves matching colour pattern; diamond of longitudinal 

skin ridges on dorsal mantle. 

Size: Maximum mantle length 90 mm, total length to around 

300 mm; weight to around 100 g. 

Habitat, biology, and fisheries: Known from muddy coastal 

waters found subtidally on soft substrates to depths of at least 

40 m. Nothing known of diet or foraging behaviour. This octopus 

is a major fisheries species throughout coastal mainland 

Asia, important in commercial trawl fisheries, particularly from 

the Gulf of Thailand and South China Sea. Exported throughout 

the world on a large scale along with Octopus sp. A (tens 

of thousands of tonnes annually). Both species are sold and 

prepared under the name “baby octopus” (at least in Australia 

and the United States). Octopus aegina has frequently been 

treated under the name of a junior synonym, Octopus dollfusi. 

Large catches of this species reported from Gulf of Thailand 

under the name Octopus “dollfusi”. 

Distribution: Found in coastal waters of continental Asia, 

from China south to Malaysia and west to at least Madras, 

India. Also reported from the Philippines. 

dorsal view


Octopus cyanea Gray, 1849 

Frequent synonyms / misidentifications: Octopus magnocellatus 

(Taki, 1964); O. marmoratus Hoyle, 1885 / O. bimaculatus Verrill, 1883. 

FAO names: En - Day octopus (formerly reported as big blue octopus). 

Diagnostic characters: Large and robust ocellate 

octopus. Arms robust and long, 4 to 6 times mantle 

length. Dorsal arms slightly shorter than other arms 

(arm formula IV=III=II.I). Right third arm of males 

hectocotylized with tiny ligula (1 to 2% of arm 

length). Gills with 9 to 11 lamellae per demibranch. In 

larger animals, 400 to 500 suckers on each normal 

arm, 180 to 230 on hectocotylized arm of male. Mature 

males with 2 to 4 enlarged suckers (12 th to 14 th )on 

arms II and III (slightly enlarged on arm IV). Colour: 

variable from dark chocolate brown through mottled 

patterns to pale grey; ocellus present as plain black 

spot surrounded by pale and dark rings; dark zebra 

bars on ventrolateral faces of all arms in submature 

and mature animals; skin sculpture of 

irregular patches separated by distinct grooves inter- 

spersed by large primary papillae, including diamond 

of 4 papillae on dorsal mantle and large papilla over 

each eye. 

Size: Maximum mantle length 160 mm, total length to 

over 1 m; weight to 6 kg. 

Habitat, biology, and fisheries: Known from clear 

tropical waters, from intertidal reefs to depths of at 

least 25 m, on coral reefs amongst both live corals and 

dead coral rubble. Preys primarily on crabs and other 

crustaceans, foraging throughout the day with peak 

activity at dusk and dawn. Occupies lairs in coral or 

rock, often visible by midden of large crab carapaces. 

Females lay up to 600 000 eggs in festoons, each egg 

around 1 mm long. Important fisheries species collected 

in large numbers throughout its range in local 

subsistence and small-scale fisheries. Frequently sold 

in fish markets, particularly through the central and 

southern tropical Pacific, in countries such as Fiji, 

Tonga, Solomons, New Caledonia, Papua New 

Guinea, and Philippines. Harvested using spear, lures, 

traps, and chemical irritants to flush animals from lairs. 

Distribution: Found widely 

throughout shallow waters of 

the tropical Indo-West Pacific 

region, from Hawaii in the east 

to the east African coast in the 

west. Reported as far north as 

southern Japan and as far 

south as New South Wales, 

Australia. 

lateroventral 

surface of arm 

dorsal view 


arm ocellus


Octopus graptus Norman, 1993a 


FAO names: En - Scribbled night octopus. 

Diagnostic characters: Large robust and muscular species. Arms 

long, 4.5 to 7 times mantle length. Dorsal arms longer than ventral 

arms (arm formula I.II.III.IV). Webs moderately deep, deepest 16 

to 22% of longest arm. Right third arm of males hectocotylized with 

large cylindrical ligula (around 6% of arm length), with deep groove. 

Gills with 13 or 14 lamellae per demibranch. In larger animals, 

around 200 to 280 suckers on each normal arm, around 80 to 90 on 

hectocotylized arm of male. Suckers on longer 

dorsal arms proportionally larger than other 

arms, but none markedly enlarged. Eggs large 

(to 40 mm) and produced in low numbers (to 

700). Colour: white to pink base colour with dark 

irregular spots and short lines (“scribbles”) 

over dorsal surfaces; distal tips of all arms 

grading to crimson purple; skin sculpture simple, 

consisting of scattered low papillae evenly distributed 

over dorsal surfaces. 

Size: Maximum mantle length 200 mm, total 

length to over 1.3 m; weight to 5 kg. 

Habitat, biology, and fisheries: Known from 

muddy coastal waters, living subtidally on soft 

substrates to depths of at least 40 m. Trawl capture 

data shows this species is caught at night, 

presumably when it is emerged from lairs and 

foraging. Captive animals emerge at night to forage, 

taking a range of prey including crustaceans, 

bivalves and fish. Females lay large eggs, 

attached singly to the roof of a lair, not in fes- 

toons. This large octopus is commercially harvested 

as bycatch in prawn trawl fisheries, sold 

for both human consumption and as bait. 

Distribution: Restricted to 

tropical muddy coastal waters 

of northern Australia from the 

Great Barrier Reef to northern 

Western Australia. 


arm 

dorsal view


Octopus cf. luteus (from Norman and Sweeney, 1997) 

Frequent synonyms / misidentifications: None / Octopus 

macropus Risso, 1826. 

FAO names: En - Small-spot night octopus. 

Diagnostic characters: Large elongate species. Arms long, 4.5 

to 5.5 times mantle length. Dorsal arms longer than ventral arms 

(arm formula I.II.III.IV). Webs moderately deep, deepest around 

15 to 20% of longest arm. Right third arm of males hectocotylized 

with large cylindrical ligula with deep groove. Gills with 12 or 13 

lamellae per demibranch. In larger animals, greater than 200 

suckers on each normal arm, approximately 80 to 90 on hectocotylized 

arm of male. Suckers on longer dorsal arms proportionally 

larger than other arms, but none markedly enlarged. Eggs 

small type in submature females examined. Colour: pink to 

bright red base colour with many small white spots over dorsal 

mantle, arm crown, webs, and arms; skin sculpture of raised 

papillae in centres of white spots, interspersed by smaller low 

papillae over all dorsal and lateral surfaces. 

Size: Maximum mantle length at least 90 mm, total length to over 

0.5 m; weight to at least 500 g. 

Habitat, biology, and fisheries: Little known of the biology and 

distribution of this species. Material examined was collected from 

coral rubble and rocky reefs at depths between 1 and 82 m. 

Females lay small type eggs, no mature females examined. This 

octopus is taken in local subsistence harvests and in line and trawl 

fisheries as bycatch. This species shows similarities with Octopus 

luteus Sasaki, 1929, from Taiwan Province of China and coastal 

waters of mainland China. Resolution of the identity of this species 

is not possible until more mature material is available. 

Distribution: At this stage, the species treated here is only known 

from the Philippines. 

dorsal view 




Octopus marginatus Taki, 1964 

Frequent synonyms / misidentifications: Octopus striolatus 

Dong, 1976 / Octopus aegina Gray, 1849. 

FAO names: En - Sand bird octopus (from Chinese “Saa liu”); 

Fr - Poulpe des sables; Sp - Pulpo reticulado. 

Diagnostic characters: Moderate-sized octopus. Arms short, 2 

to 3 times mantle length. Dorsal arms slightly shorter than other 

arms (arm formula IV=III=II.I or III.IV=II.I). Right third arm of 

males hectocotylized with small ligula (1.5 to 3.5% of arm 

length). Gills with 9 to 11 lamellae per demibranch. Up to 150 

suckers on each normal arm, 60 to 85 on hectocotylized arm of 

male. Mature males possess 4 to 5 slightly enlarged suckers (7th to 11th )onarmsII and III. Colour: pattern typically orange-brown 

to purple with dark reticulations defining distinct patches in 

irregular longitudinal rows; suckers white to pink contrasting 

against dark brown to black along leading edge of arms I to 

III; narrow transverse “head bar” visible in live animals; white 

triangle below each eye; dark reticulations distinctive on 

lateral arm crown in same position as false eye-spots in ocellate 

species; skin sculpture of regular patches separated by distinct 

grooves; diamond of 4 longitudinal skin ridges on dorsal mantle 

and large papilla over each eye. 

Size: Maximum mantle length around 

100 mm, total length to around 

300 mm; weight to 400 g. 

Habitat, biology, and fisheries: Known 

from coastal muddy waters on mud 

and sand substrates, subtidal to 

depths of at least 190 m. Little known 

of biology or behaviour.Females lay up 

to 100 000 small eggs, up to 3 mm 

long. Important fisheries species collected 

by trawlers, pots and lines. Taxonomy 

confused with Octopus aegina 

(treated above) and O. kagoshimensis 

from Japan. 

Distribution: Found in tropical continental 

waters of the Indian Ocean, 

from the Red Sea and East Africa to lateral view of head 

Southeast Asia and eastern Australia. 

? ? 

? 

? 

? 

dorsal view


Octopus nocturnus Norman and Sweeney, 1997 


macropus Risso, 1826. 

FAO names: En - Philippine night octopus. 

Diagnostic characters: Moderate-sized octopus. Arms long, 5 to 

6.5 times mantle length. Dorsal arms distinctly longer than other 

arms (arm formula I.II.III.IV). Webs shallow, deepest 10 to 15% of 

longest arm. Right third arm of males hectocotylized with moderate 

length ligula (3 to 5% of arm length). Gills with 10 or 11 lamellae per 

demibranch. In larger specimens, 180 to 220 suckers on each normal 

arm, 80 to 90 on hectocotylized arm of male. No enlarged suckers 

in either sex. Eggs small type and numerous (more than 1 000) in 

submature females examined. Colour: pattern of red-brown base 

with irregular darker blotches and white spots over dorsal surfaces 

including mantle; white spots paired along length of arms; 

skin sculpture of small low round papillae; no primary papillae visible. 

Size: Maximum mantle length around 60 mm, 

total length to around 350 mm; weight to at 

least 100 g. 

Habitat, biology, and fisheries: Known primarily 

from intertidal coral and rocky reefs. 

Deepest record a poison station affecting between 

a depth of 1.5 and 4.5 m. Members of 

this species emerge at night to forage during 

low tides on intertidal reefs. Females produce 

small eggs in large numbers. Collected in local 

subsistence harvest, speared on night low 

tides, historically using burning bamboo 

torches to find active octopuses. 

Distribution: At this stage, known only from 

throughout the Philippines. 


arm of male 

dorsal view


Octopus ornatus Gould, 1852 

Frequent synonyms / misidentifications: Octopus (or Callistoctopus) 

arakawai (Taki, 1964) / None. 

FAO names: En - White-striped octopus. 

Diagnostic characters: Large and elongate octopus. Arms long, 6 

to 8 times mantle length. Dorsal arms much longer than ventral arms 

(arm formula I.II.III.IV). Webs shallow, deepest 5 to 11% of longest 

arm. Right third arm of males hectocotylized with large cylindrical 

ligula (4 to 6% of arm length), with deep groove. Gills with 13 or 14 

lamellae per demibranch. In larger animals, 300 to 400 suckers on 

each normal arm, 150 to 170 on hectocotylized arm of male. Suckers 

on longer dorsal arms proportionally larger than other arms, but none 

markedly enlarged. Colour: pattern of brown to deep red base colour 

with white markings; distinctive pattern of longitudinal white bars 

on dorsal mantle, visible in live, fresh dead and preserved material; 

large paired white spots along arms; skin sculpture of low irregular 

patches separated by distinct grooves; elongate skin flaps can be 

raised within longitudinal white bars on mantle. 

Size: Maximum mantle length 130 mm, total length to 

over 1 m; weight to at least 1 kg. 

Habitat, biology, and fisheries: Known from clear 

tropical waters, from intertidal shallows to a depth of at 

least 10 m, on coral reefs amongst both live corals and 

dead coral rubble. Forages exclusively at night, preying 

primarily on crustaceans and other octopus species. 

Typically encountered foraging along the edges of intertidal 

reefs adjacent to deeper water. Sometimes 

encountered swimming at the surface in or near lagoons. 

Occupies deep lairs during the day, the entrance 

of which is blocked at several levels. Females 

lay up to 35 000 eggs in festoons, each egg around 3 

to 4 mm long. Harvested on a small scale throughout 

its range, primarily in local subsistence fisheries. It is 

sold in fish markets in the central and southern tropical 

Pacific, but less frequently than Octopus cyanea. Har- 

vested at night using torches and spears in at least 

Hawaii. 

Distribution: Found widely throughout shallow waters 

of the tropical Indo-West Pacific region, from Hawaii in 

the east to the east African coast in the west. Reported 

as far north as southern Japan (as Callistoctopus arakawai) 

and as far south as New South Wales, Australia. 


arm III 

dorsal view


Octopus tetricus Gould, 1852 


cyanea Gray, 1849; O. cyaneus Gray, 1849. 

FAO names: En - Common Sydney octopus. 

Diagnostic characters: Large and robust species. Arms 

moderate to long, 3 to 4.5 times mantle length. Dorsal arms 

slightly shorter and less robust than other arms (arm formula 

IV=III=II.I). Right third arm of males hectocotylized with tiny 

ligula (1 to 2% of arm length). Gills with 8 or 9 lamellae per 

demibranch. In larger animals, around 220 to 260 suckers on 

each normal arm, around 140 to 160 on hectocotylized arm 

of male. Mature males with around 3 to 5 enlarged suckers 

(13 th to 17 th )onarmsII and III. Colour: active animals cream 

to mottled orange and dark brown; transverse narrow dark 

bands along arms in some colour patterns; resting animals 

within lairs show grey dorsal surfaces, orange arm faces 

and eyes with a white iris; skin sculptured in rounded 

patches separated by distinct grooves; capable of raising 

large papillae over dorsal surfaces, including 4 in diamond 

pattern on dorsal mantle and 1 above each eye. 

Size: Maximum mantle length at least 140 mm, total length to 

over 0.6 m; weight to 1 kg. 

Habitat, biology, and fisheries: Known from shallow coastal 

waters, living subtidally on and adjacent to rocky reefs, to a 

depth of at least 60 m. Active primarily at night, although alert 

in the mouth of lairs throughout the day. Preys primarily on 

crabs, but will also take shellfish and finfish (at least in 

captivity). Occupies lairs in rock crevices or excavated under 

rocks on sand or mud. Females lay over 150 000 eggs in 

festoons, each egg around 2 to 3 mm long. Moderate scale 

harvest as bycatch in prawn and finfish trawl fisheries. Frequently 

sold in fish markets in New South Wales and southern 

Queensland. A distinct undescribed species from Western 

Australia has been incorrectly treated under this name. 

Distribution: Found in warm temperate waters of coastal 

New South Wales and southern Queensland, Australia. 

dorsal view


Octopus sp. A 


membranaceus Quoy and Gaimard, 1832. 

FAO names: En - Eye-bar ocellate octopus. 

Diagnostic characters: Small to moderate-sized ocellate octopus. 

Arms short to moderately long, 2.5 to 3 times mantle length. 

Dorsal arms slightly shorter than other arms (arm formula 

IV=III.II.I). Right third arm of males hectocotylized with moderate 

length, elongate ligula (4 to 10% of arm length) with a shallow 

groove. Gills with 8 to 10 lamellae per demibranch. Up to 150 

suckers on each normal arm, 65 to 80 on hectocotylized arm of 

male. Mature males possess 2 to 4 moderately enlarged suckers 

(4 th to 8 th )onarmsII and III. Colour: pattern typically cream to 

light green colour on all surfaces; short longitudinal bar 

through each eye; dark brown or black lines along edges of 

arms; ocellus present as black spot containing a fine iridescent 

blue to purple ring (4 to 6 mm in diameter in adults); 

head bar and longitudinal stripes on mantle absent; skin sculpture 

of small low rounded papillae, interspersed by slightly larger 

pink papillae; diamond of four longitudinal skin ridges on dorsal 

mantle and large papilla over each eye. 

Size: Maximum mantle length around 60 mm, total length to 

around 250 mm. 

Habitat, biology, and fisheries: Known from coastal waters on 

mud and sand substrates, from intertidal shallows to a depth of 

at least 60 m. Females lay small eggs, up to 3 mm long, often in 

shells or bottles. Very important fisheries species collected primarily 

from the Gulf of Thailand by trawlers. Exported throughout 

the world on a large scale along with Octopus aegina (tens of 

thousands of tonnes annually). Both species are sold and prepared 

under the name “baby octopus” (at least in Australia and 

the United States). The ocellate octopuses of Southeast Asia 

require extensive revision. A number of undescribed taxa occur 

throughout the area, several of which are treated under the name 

O. membranaceus. Fisheries statistics for Octopus sp. A were 

reported from Gulf of Thailand, under the name Octopus “membranaceus”. 

Distribution: Found in tropical continental waters of Southeast 

Asia, from at least Gulf of Thailand, through Indonesia to northern 

Australia. dorsal view


Ameloctopus litoralis Norman, 1992b 

En - Banded long-arm octopus. 

Small octopus (maximum mantle length 30 mm) with greatly elongated arms 

(5 to 10 times mantle length). Arms frequently sever at base as decoy to 

predators, around 10 th sucker. more than 180 suckers on intact normal arms 

of larger animals, 20 to 40 on hectocotylized arm of male. No ink sac. Right 

third arm long in submature males, lacking ligula. Mature males sever arm 

around 40 th sucker and develop ligula from stump. Eggs large, to 10 mm. Skin 

smooth, no papillae. Colour: pink to brown, hearts visible through thinwalled 

mantle, arms with regular purple-brown bands. Intertidal mud, 

sand, and rubble reefs. No fisheries value but may be poisonous. Restricted 

to tropical coastal waters of northern Australia. 

Hapalochlaena fasciata (Hoyle, 1886) 

tip of 


arm 

En - Blue-lined octopus. 

Small octopus (maximum mantle length 40 mm) with short arms (2 to 3 times 

mantle length). Colour: cream to orange base colour with iridescent blue 

lines (not rings) on dorsal mantle and single or linked blue rings on arm 

crown and arms. Intertidal and shallow rocky reefs to depths of at least 20 m. 

No fisheries value but extremely venomous, tetrodotoxin venom produced in 

the salivary glands and responsible for a number of human deaths. Subtropical 

waters of eastern Australia from southern Queensland to southern New 

South Wales. 


dorsal view 

dorsal view


Hapalochlaena lunulata (Quoy and Gaimard, 1832) 

En - Greater blue-ringed octopus. 

Small octopus (maximum mantle length 50 mm) with short arms (1.5 to 2 times 

mantle length). Colour: cream to orange base colour with large iridescent 

blue rings (to 12 mm in diameter) on dorsal mantle, arm crown and arms. 

Intertidal and shallow coral reefs. No fisheries value but extremely venomous, 

tetrodotoxin venom produced in the salivary glands and responsible for a 

number of human deaths. Tropical waters of Indo-Malayan Archipelago from 

at least the Philippines to northern Australia and east to Vanuatu. 

Hapalochlaena cf. maculosa (from Roper and Hochberg, 1988) 


En - Lesser blue-ringed octopus. 

Small octopus (maximum mantle length 40 mm) with short arms (1.5 to 2.5 

times mantle length). Colour: cream to orange base colour with small 

iridescent blue rings (approximately 2 mm in diameter) on dorsal mantle 

and single or linked blue rings on arm crown and arms. Intertidal and shallow 

coral reefs to depths of at least 55 m. No fisheries value but potentially 

venomous, as in other members of the genus. Tropical waters of northeastern 

Australia from southern Great Barrier Reef to southern Gulf of Carpentaria. 


dorsal view 

dorsal view


Octopus abaculus Norman and Sweeney, 1997 

En - Mosaic drop-arm octopus. 

Small octopus (maximum mantle length 33 mm) with long arms (5 to 6 times mantle 

length). Arms frequently sever at base as decoy to predators, between 5 th and 

8 th sucker. In adults, 170 to 210 suckers on normal arms, 90 to 120 on hectocotylized 

arm of male. Mature males with 8 to 12 enlarged suckers on arms III 

to IV. Gills with 6 lamellae per demibranch. Colour: dark grey to purple-black 

reticulations define large circular cream spots on dorsal surfaces, producing 

a mosaic pattern. Intertidal and shallow coral, rubble, or rocky reefs to depths of 

at least 6 m. No known fisheries value, but may be taken in local subsistence 

harvest. Only known from the Philippines. 

Octopus aculeatus d’Orbigny, 1835 

tip of 


arm 

En - Greater drop-arm octopus. 

Small to moderate octopus (maximum mantle length 65 mm) with long arms (5 to 

6 times mantle length). Arms frequently sever at base as decoy to predators, 

between 5 th and 8 th sucker. In adults, 190 to 250 suckers on normal arms, 140 

to 175 on hectocotylized arm of male. Mature males with 5 to 12 enlarged suckers 

on arms II to IV. Gills with 6 or 7 lamellae per demibranch. Colour: mottled 

grey-brown to dark grey with small cream spots forming fine mosaic pattern 

towards arm tips. Intertidal and shallow mangroves, coral rubble, and rocky reefs 

to depths of at least 10 m. Small-scale local subsistence harvest. Only known from 

the Philippines. 

tip of 


arm 

dorsal view 

dorsal view


Octopus alpheus Norman, 1993a 

En - Capricorn night octopus. 

Moderate to large robust octopus (maximum mantle length 90 mm) with 

moderately long arms (3 to 5 times mantle length). Dorsal arms longer 

than lateral and ventral arms. Webs moderately deep, deepest 16 to 

25% of longest arm. In adults, 190 to 230 suckers on normal arms, 80 

to 100 on hectocotylized arm of male. No enlarged suckers. Large eggs 

(more than 8 mm) produced in low numbers (less than 500). Colour: 

orange-brown to red base colour with large white spots over dorsal 

mantle, arm crown, webs and arms. Intertidal coral 

reef flats, emerging to forage during night low tides. No 

commercial harvest. Tropical species restricted to the 

Capricorn Bunker Islands, southern Great Barrier Reef, 

Australia. 

Octopus aspilosomatis Norman, 1993a 

tip of 


arm 

En - Plain-body night octopus. 

Moderate-sized, elongate octopus (maximum mantle length 80 mm) 

with long arms (4.5 to 6 times mantle length). Dorsal arms longer than 

lateral and ventral arms. Webs shallow, deepest 9 to 15% of longest 

arm. In adults, 200 to 270 suckers on normal arms, 75 to 100 on 

hectocotylized arm of male. No enlarged suckers. Small eggs (less 

than 3 mm) produced in large numbers (more than 

10 000). Colour: orange-brown to deep maroon base 

colour with white spots on arm crown, webs, and paired 

down arms; no white spots ever expressed on mantle. 

Intertidal coral reef flats, emerging to forage during 

night low tides. No commercial harvest. Tropical species 

known only from coastal reefs and offshore islands of 

northern Great Barrier Reef, Australia. 

tip of 


arm 

dorsal view 

dorsal view


Octopus australis Hoyle, 1885 

En - Hammer octopus. 

Moderate-sized octopus (maximum 

mantle length 70 mm) with 

moderately long arms (2.5 to 4 

times mantle length). Up to 220 

suckers on normal arms, 60 to 80 

on hectocotylized arm of male. 

Ligula of large swollen club 

(“hammer”). Eggs large, to 

12 mm. Raised ridge (“keel”) of 

skin around lateral mantle. Skin 

covered in regular rounded papil- tip of hectocotylized arm 

lae. Colour: cream to purplebrown 

dorsally, cream on ventral 

surfaces. Sand and mud substrates 

to a depth of 134 m. Small-scale trawl 

harvest for human consumption and as bait. 

Restricted to subtropical waters of southern 

Queensland and New South Wales, Australia. 

Octopus dierythraeus Norman, 1993a 

lateral view 

En - Red-spot night octopus. 

Large robust octopus (maximum mantle length 140 mm) with moderately 

long arms (4 to 5 times mantle length). Dorsal arms longer than 

lateral and ventral arms. Webs moderately deep, deepest 18 to 28% 

of longest arm. In adults, 230 to 280 suckers on normal arms, 100 to 

130 on hectocotylized arm of male. No enlarged suckers. Large eggs 

(more than 14 mm) produced in low numbers (less than 500). 

Colour: foraging animals orange brown to red with white spots over 

dorsal surfaces, alarm display of white base colour with red spots 

over mantle, arm crown, webs and arms. Shallow coastal waters on 

rubble, sand or mud from intertidal reefs to depths of at 

least 80 m. Potential bycatch harvest. Tropical species 

restricted to coastal waters of northern Australia. 


arm 

dorsal view 

(after Stranks and Norman, 1993) 

dorsal view


Octopus exannulatus Norman, 1993b 

En - Plain-spot ocellate octopus. 

Small octopus (maximum mantle length 50 mm) with short arms (2 to 3 

times mantle length). 120 to 190 suckers on normal arms of adults, 60 

to 80 on hectocotylized arm of male. Gills with 7 or 8 lamellae per 

demibranch. In mature males, 2 or 3 very large suckers on arms II and 

III (10 th to 12 th ). Eggs small, to 4 mm. Skin covered in regular oval to 

round papillae. Colour: cream with 4 black stripes along dorsal 

mantle and arm crown; plain ocellus without iridescent ring; black 

lines along leading edge of arms. Shallow sand and mud substrates 

to depths of at least 84 m. Small harvest in Australia, primarily as bait. 

Indo-Malayan area from Philippines, to Thailand, and northern Australia. 

Octopus mototi Norman, 1993b 

En - Poison ocellate octopus (from Rapa, Rapa Is., “Fe’e motot”). 

Moderate-sized octopus (maximum mantle length 70 mm) with short to 

moderate arms (2.5 to 3 times mantle length). In adults, 140 to 170 

suckers on normal arms, 90 to 110 on hectocotylized arm of male. Gills 

with 9 to 11 lamellae per demibranch. No distinct enlargement of suckers 

in mature males. Eggs small, to 6 mm. Skin covered in regular polygonal 

patches. Colour: resting colour of orange with ring of 5 large black 

spots above each eye; 6 longitudinal rows of spots on dorsal mantle 

which become solid maroon stripes in live alarm displays; ocellus 

with large iridescent blue ring. Sand and coral rubble substrates to 

depths of 54 m. No fisheries value, potentially venomous as for blueringed 

octopuses. Reported as poisonous on Rapa Island. Occurs in at 

least the western and southern tropical Pacific Ocean. 

ocellus dorsal view 

ocellus 

dorsal view


Octopus polyzenia Gray, 1849 

En - Arm-band ocellate octopus. 

Small octopus (maximum mantle length 40 mm) with short to moderate 

arms (2 to 3 times mantle length). 80 to 140 suckers on normal arms of 

adults, 45 to 55 on hectocotylized arm of male. Gills with 6 or 7 lamellae 

per demibranch. In mature males, 1 to 3 enlarged suckers (12 th to 14 th ) 

on all arms. Eggs large relative to mantle length, to 7.5 mm.Skin covered 

in low small papillae. Colour: mottled cream to dark brown; ocellus with 

small iridescent blue ring; transverse dark bars widely spaced on 

arms, one every 3 to 5 sucker pairs. Shallow coastal waters on rubble, 

sand or mud to depths of at least 20 m. No fisheries value but may be 

harvested with other northern Australian ocellate octopuses. Coastal 

waters of northern Australia. 

ocellus 

dorsal view 



STOMATOPODS 

by R.B. Manning

828 Stomatopods 

Technical Terms and Measurements TECHNICAL TERMS AND MEASUREMENTS 

eye 

carapace 

tail 

thoracic somites abdominal somites 

antennule 

5 6 7 8 I II III IV 

V 

antenna 

raptorial claw 

total 

length 

antennule cornea 

peduncle of eye 

median carina 

of carapace 

antenna 

length of 

carapace 

length of thorax 

(free somites 5-8) 

length of 

abdomen 

(somites II-IV 

omitted) 

ocular scales 

antennular process 

anterolateral angle 

of carapace 

anterior part of body (dorsal view) 

rostral plate 

walking legs 

lateral view of a stomatopod (mantis shrimp) 

5 

6 

7 

8 

I 

V 

VI 

dorsal view (without head and extremities) 

antennal scale 

endopod 

gill 

pleopods 

uropod 

VI 

CARAPACE AND ROSTRAL PLATE 

rostral plate 

anterolateral angle of carapace 

anterior bifurcation of median carina 

marginal 

lateral } CARINAE 

intermediate 

gastric groove 

cervical groove 

reflected portion of marginal carina 

EXPOSED THORACIC SOMITES 

lateral process of 5 th thoracic somite 

median 

submedian } CARINAE 

intermediate 

FIRST ABDOMINAL SOMITE 

lateral 

marginal } CARINAE 

intermediate 

TELSON 

marginal carina 

prelateral lobe 

median carina 

lateral 

intermediate} CARINAE 

submedian 

lateral 

intermediate} DENTICLES 

5 

submedian 

TH AND 6TH ABDOMINAL SOMITES 

median 

submedian 

intermediate}CARINAE 

lateral 

marginal 

propodus 

(basal segment) 

proximal 

segment 

left uropod (ventral view) 

distal 

segment 

telson 

exopod 

basal 

prolongation 

of uropod

General Remarks 829 

General Remarks GENERAL REMARKS 

Stomatopods, also called mantis shrimps, are elongate, flattened, shrimp-like or lobster-like crustaceans, 

which are characterized by the following features: large, often T-shaped, movable eyes, often with a 

bilobed cornea; a very short carapace, not more than about 1/3 the total length and not covering the 

eyes; only 3 pairs of walking legs; 5 pairs of pleopods under the anterior 5 abdominal somites (I to V); 

a long, flattened tail (which includes part of the thorax, the abdomen, and the terminal telson); 1 pair 

of lateral uropods on the abdominal somite VI which includes a strongly spined ventral process; a telson 

that is often spined posteriorly. The most conspicuous characteristic of mantis shrimps is a pair of massive, 

conspicuous, praying mantis-like “raptorial” claws which are folded under the sides of the carapace. 

In many of the larger stomatopods, the terminal 2 segments of the claws usually are lined with sharp, 

serrated teeth. The claws are adapted for crushing or spearing. Those species with crushing claws have 

the terminal segment of the claw broadened and heavily buttressed basally. Spearing claws are more 

elongate, more conspicuously toothed, and much more slender, not inflated basally. 

carpus 

merus 

propodus 

crushing claw 

These are burrowing animals that hunt from the burrow or leave it to forage for food. Few, if any, mantis 

shrimps are fished commercially in the Western Central Pacific. However, very large, up to 38 cm long, 

representatives of 4 families can be found in markets or may be fished artisanally. They are often caught 

in trawls and fish traps, at night lights, and by hand. 

Banded mantis shrimps 

Body smooth, lacking longitudinal ridges or 

carinae; telson without median carina; eyes 

T-shaped, with large, bilobed cornea; raptorial 

claw large and slender. Four species found in the 

area, 2 of Lysiosquilla and 2 of Lysiosquillina, 

only 1 of which, Lysiosquillina maculata, is 

known to be fished, at least artisanally. 

dactylus 

inflated 

basally 

basic types of raptorial claw 

spearing claw 

Guide GUIDE to Families TO FAMILIES OF INTEREST TO FISHERIES OCCURRING IN THE AREA 

ODONTODACTYLIDAE 

eye globular, not 

T-shaped 

Page 832 

telson with 

median carina 

Odontodactylid mantis shrimps 

Body smooth, lacking longitudinal ridges or 

carinae anterior to terminal abdominal somite 

(= telson); telson with distinct median 

longitudinal carina; eyes subglobular, cornea not 

bilobed; raptorial claw short, with dactylus 

heavily buttressed, swollen basally. Four species 

of Odontodactylus occur in the area, only 1 of 

which, O. scyllarus, is large and common 

enough to occasionally be found in markets. 

claw slender, 

dactylus 

swollen basally 

surface of 

body smooth 

(except telson) 

Odontodactylidae 

claw large and slender, 

with many teeth 

body conspicuously 

banded 

no median 

carina on telson 

LYSIOSQUILLIDAE Page 835 

Lysiosquillidae 

body smooth 

teeth


propodus of 

HARPIOSQUILLIDAE Page 838 claw with 

small spines 

Harpiosquillid mantis shirmps 

Body with longitudinal carinae or ridges; 

posterolateral angles of carapace with 

conspicuous excavation; telson with distinct 

median longitudinal carina; eyes T-shaped, with 

large bilobed cornea; raptorial claw large and 

slender; opposable margin of propodus of claw 

with erect spines. The larger, more conspicuous 

harpiosquillids belong to a single genus, 

Harpiosquilla. Nine species of Harpiosquilla 

found in the area, 2 of which are common 

enough to be found in markets. 

propodus of claw with 

SQUILLIDAE Page 842 

blunt pectinations 

Squillid mantis shrimps 

Body with longitudinal carinae or ridges; telson 

with distinct median longitudinal carina; eyes 

T-shaped, with bilobed cornea; raptorial claw 

large and slender; opposable margin of 

propodus of claw lined with low, blunt 

pectinations; posterolateral angles of carapace 

rounded, convex, rather than excavate. The 

family Squillidae comprises some 40 genera, 20 

of which are represented in the Indo-West 

Pacific, but members of only 5 genera are large 

enough or abundant enough to be used for food. 

Key to Families KEY TO MAJOR FAMILIES OF STOMATOPODS OCCURRING IN THE AREA 

1a. Telson without median carina 

(Fig. 1a) . . . . . . . . . . . . . . . . . . . → 2 

1b. Telson with distinct median carina 

(Fig. 1b) . . . . . . . . . . . . . . . . . . . → 3 

2a. Distal segment of endopod of walking 

legs circular or nearly so 

(Fig. 2a); eyes not T-shaped, with 

cornea rounded or oval, not bilobed 

(Fig. 3a); total length not exceeding 

7cm. . . . . . . . . . . . . . Nannosquillidae 

posterolateral corner 

of carapace excavate 

Harpiosquillidae 

evenly rounded 

Squillidae 

longitudinal 

ridges on body 

longitudinal 


1 2 

a) Nannosquillidae 

3 

1 2 

3 

b) Lysiosquillidae a) Nannosquillidae b) Lysiosquillidae 

Fig. 2 walking legs 1-3 

Fig. 3 eye 

3a. Dactylus of raptorial claw inflated basally, strongly buttressed (Fig. 4a, b) . . . . . . . . . . . . → 4 

3b. Dactylus of raptorial claw slender, not inflated or buttressed basally (Fig. 4c, d) . . . . . . . . . → 5 

a) 

no 

median 

carina 

Fig. 1 telson (dorsal view) 

b) 

median 

carina 

2b. Distal segment of endopod of walking legs strap-shaped, elongate (Fig. 2b); eyes 

T-shaped, cornea strongly bilobed (Fig. 3b); total length to at least 30 cm . . . . . . Lysiosquillidae 

circular 

strap-shaped 

cornea 

cornea 

bilobed 

rounded

List of Families and Species 831 

4a. Dactylus of raptorial claw with teeth on inner margin (Fig. 4a); total length up to 17 cm 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontodactylidae 

4b. Dactylus of raptorial claw unarmed on inner margin (Fig. 4b); total length less than 10 

to 11 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gonodactylidae 

inner margin 

with teeth 

dactylus 

inflated 

basally 

a) Odontodactylidae b) Gonodactylidae c) Harpiosquillidae 

Fig. 4 right claw 

5a. Carapace with posterolateral excavation 

(Fig. 5a); propodus of 

raptorial claw lined with erect 

spines (Fig. 4c) . . . . . . . Harpiosquillidae 

5b. Carapace rounded posterolaterally, 

not excavate (Fig. 5b); 

propodus of raptorial claw lined 

with blunt pectinations (Fig. 4d) . . . Squillidae 

Fig. 5 posterolateral carapace and lateral processes of 

thoracic somites (dorsal view) 

List of Families and SpeciesLIST OF FAMILIES OCCURRING IN THE AREA 

The symbol is given for those families which are treated further in this contribution. 

Superfamily BATHYSQUILLOIDEA Manning, 1967 

BATHYSQUILLIDAE Manning, 1967 

INDOSQUILLIDAE Manning, 1995 

Superfamily ERYTHROSQUILLOIDEA Manning and Bruce, 1984 

ERYTHROSQUILLIDAE Manning and Bruce, 1984 

Superfamily GONODACTYLOIDEA Giesbrecht, 1910 

ALAINOSQUILLIDAE Moosa, 1991 

EURYSQUILLIDAE Manning, 1977 

GONODACTYLIDAE Giesbrecht, 1910 

HEMISQUILLIDAE Manning, 1980 

ODONTODACTYLIDAE Manning, 1980 

PARASQUILLIDAE Manning, 1995 

PROTOSQUILLIDAE Manning, 1980 

PSEUDOSQUILLIDAE Manning, 1977 

TAKUIDAE Manning, 1995 

Superfamily LYSIOSQUILLOIDEA Giesbrecht, 1910 

CORONIDIDAE Manning, 1980 

HETEROSQUILLIDAE Manning, 1995 

LYSIOSQUILLIDAE Giesbrecht, 1910 

NANNOSQUILLIDAE Manning, 1980 

TETRASQUILLIDAE Manning and Camp, 1993 

Superfamily SQUILLOIDEA Latreille, 1803 

HARPIOSQUILLIDAE Manning, 1980 

SQUILLIDAE Latreille, 1803 

erect 

spines 

emarginate 

d) Squillidae 

a) Harpiosquillidae b) Squillidae 

blunt 

pectinations 

rounded 

Reference 

Manning, R.B. 1995. Stomatopod Crustacea of Vietnam: the legacy of Raoul Serène. Tokyo, Crustacean Research, The 

Carcinological Society of Japan, Special Number 4:339 p.


ODONTODACTYLIDAE 

Odontodactylid mantis shrimps 

Diagnostic characters: Moderate-sized mantis shrimps 

(maximum total length more than 17 to 18 cm). Eyes 

globular, not T-shaped, cornea not bilobed. Carapace, thorax, 

and abdomen smooth, not ornamented with any longitudinal 

ridges or carinae anterior to last abdominal somite 

(= telson). Telson with median longitudinal carina. Raptorial 

claw short and heavily buttressed at base of terminal segment, 

adapted for smashing prey; inner margin of dactylus 

toothed with no more than 5 short teeth. 

Habitat, biology, and fisheries: In coarse-bottom or 

level-bottom habitats. Essentially nothing is known about the 

biology of odontodactylids and no organized fisheries are 

known to exist for them. 


Gonodactylidae: share the buttressed, inflated claw, but have 

the inner margin of the claw unarmed and are much smaller. 

right claw 

Gonodactylidae 

Key to the species of Odontodactylidae occurring in the area 

1a. Proximal segment of uropodal exopod shorter than distal (Fig. 1a) . . . . . . . . Raoulius cultrifer 

1b. Proximal segment of uropodal exopod longer than distal (Fig. 1b-d) . . . . . . . . . . . . . . . → 2 

proximal segment 

of exopod 

distal segment 

of exopod 

a) Raoulius cultrifer 

longitudinal carina 

2 pairs of 

carinae 

intermediate 

no teeth 

right claw 

inner 

margin of 

dactylus 

toothed 


1 pair of 

carinae 

surface of 

body 

smooth 

(except 

telson) 

b) Odontodactylus brevirostris 

2 pairs of 

carinae 

dorsal view 

denticle intermediate denticle 

c) Odontodactylus japonicus 

d) Odontodactylus scyllarus 

Fig. 1 sixth abdominal somite, telson, and right uropod 

(dorsal view) (from Manning, 1967) 

eye globular, 

not T-shaped 

claw 

slender, 

dactylus 

swollen 

basally 

telson with 

median carina 

(after Manning, 1995)

Odontodactylidae 833 


2a. Ocular scales separate in midline (Fig. 2a); telson with 1 pair of carinae converging 

under apex of median carina (Fig. 1b) . . . . . . . . . . . . . . . . . . Odontodactylus brevirostris 

2b. Ocular scales fused in midline (Fig. 2b); telson with 2 pairs of carinae converging under 

apex of median carina (Fig. 1c, d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Fifth abdominal somite unarmed 

posterolaterally 

III 

(Fig. 3a); telson with longitudinal 

carina extending 

anteriorly from inner intermediate 

denticle (Fig. 1c) 

. . . . . . . .Odontodactylus japonicus 

3b. Fifth abdominal somite 

with posterolateral spine 

(Fig. 3b); telson lacking 

longitudinal carina extending 

anteriorly from inner intermediate 

denticle 

(Fig. 1d) . . . Odontodactylus scyllarus 



Odontodactylus brevirostris (Miers, 1884) 

Odontodactylus japonicus (De Haan, 1844) 

Odontodactylus scyllarus (Linnaeus, 1758) 

Raoulius cultrifer (White, 1850) 

Reference 

a) Odontodactylus brevirostris b) Odontodactylus scyllarus 

(from Manning, 1967) 

Fig. 2 anterior part of body (dorsal view) 

IV V 

ocular scales 

spine 

a) Odontodactylus japonicus b) Odontodactylus scyllarus 

Fig. 3 abdominal somites III-V (lateral view) 

Manning, R.B. 1967. Review of the genus Odontodactylus (Crustacea: Stomatopoda). Proc. U.S. Natl. Mus., 

123(3606):1-33. 

III 

IV 

V


Odontodactylus scyllarus (Linnaeus, 1758) 

En - Reef odontodactylid mantis shrimp. 

Maximum total length about 17 cm, the largest “smasher”. 

One of the most brightly coloured stomatopods, with deep 

blue uropods and those and other appendages lined with 

bright red setae. Lives in existing burrows in shallow rough 

bottom habitats, often on or near coral reefs. Active during 

the day and uses its buttressed raptorial claws to smash 

hard-bodied prey like other scyllarids. Collected primarily 

by hand. Used in the aquarium trade because of its bright 

coloration. Widely distributed from Japan to the western 

Indian Ocean. 

Raoulius cultrifer (White, 1850) 

En - Pastel odontodactylid mantis shrimp. 

Maximum total length about 12 cm. Colour primarily in 

pastels, with pink or purple uropods and antennal scales. 

Burrows in level bottoms near shore, in depths to about 

25 m. May be taken together with lysiosquillids and squillids 

in trawling operations and at night lights. Known from 

southern China to Australia. 

dorsal view 

(after Manning, 1995) 

dorsal view 

anterior part of body 

dorsal view 

6 


th abdominal somite, telson, and right uropod

Lysioquilidae 835 

Lysioquilidae LYSIOSQUILLIDAE 

Banded mantis shrimps 

Diagnostic characters: Eye T-shaped, cornea bilobed. 

Carapace, thorax, and abdomen smooth, lacking 

longitudinal ridges or carinae. Telson lacking distinct 

median carina; marginal teeth or spines of telson inconspicuous. 

Raptorial claw slender and elongate, adapted 

for spearing prey, with toothed edge of dactylus bearing 

numerous, large, serrated teeth or spines. Lysiosquillids 

usually are clearly banded with alternate light and 

darkly pigmented bands. Members of the 2 genera likely 

to be encountered in markets are readily distinguished by 

2 features: in Lysiosquilla, (1) the antennal scale is slender, 

elongate, more than 3 times longer than wide and it 

is outlined by dark pigment, and (2) there is a spine-like 

projection on the anterior margin of the antennal peduncle; 

in Lysiosquillina, the antennal scale is oval, about 2 times 

longer than wide, and it bears a central spot or blotch of 

dark pigment; the antennal peduncle is smooth anteriorly, 

lacking a distinct spine-like projection. 

body 

Habitat, biology, and fisheries: Almost nothing is known smooth 

about the biology of lysiosquillids. They form simple 

burrows with 2 entrances, one at each end, in level-bottom 

habitats in shallow water, from shore to a depth of about 

25 m.The burrow openings may be as much as 10 m apart. 

Although they generally hunt from the mouth of their 

burrow, they occasionally leave their burrows and may be 

caught at night lights or in trawls. Most fisheries for 

lysiosquillids are artisanal. Five species of lysiosquillids 

occur within the area, but information on distribution is 

limited and contradictory. Only 1 species, Lysiosquillina 

maculata (Fabricius, 1793), is particularly common and 


known to be fished artisanally. 


Nannosquillidae: also smooth-bodied and also may be conspicuously 

banded with alternate dark and light bands of pigment, but much smaller, 

rarely exceeding 7 cm in length, and of no commercial importance. 

Key to the species of Lysiosquillidae occurring in the area 

1a. Dorsal processes of antennular somite unarmed 

(Fig. 1a); telson with movable submedian teeth (Fig. 2a) 

. . . . . . . . . . . . . . . . . . . . . . . Lysiosquilloides siamensis 

1b. Dorsal processes of antennular somite produced into 

spines (Fig. 1b); telson lacking movable submedian teeth 

(Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

unarmed 

spined 

a) Lysiosquilloides siamensis b) Lysiosquilla, Lysiosquillina 


dorsal view 

Nannosquillidae 

claw large and 

slender, with 

many teeth 

body 

conspicuously 

banded 

no median 

carina on 

telson 

movable tooth no movable tooth 

a) Lysiosquilloides siamensis b) Lysiosquilla, Lysiosquillina 

Fig. 2 telson (dorsal view)


2a. Antennal protopod with 

angled dorsal projection 

(Fig. 3a); antennal scale 

slender, length more 

than 3 times width 

(Fig. 3a) . . . . . (Lysiosquilla) → 3 

2b. Antennal protopod lacking 

angled dorsal projection 

(Fig. 3b); antennal 

scale broad, length less 

than 3 times width 

(Fig. 3b) . . . . (Lysiosquillina) → 4 

antennal 

scale 

slender 

angled dorsal 

projection 

antennal 

scale broad b) Lysiosquillina 


a) Lysiosquilla 

3a. Median carina on rostral plate flanked by longitudinal grooves (Fig. 4a); dactylus of 

raptorial claw with 7 or 8 teeth (Fig. 5a) . . . . . . . . . . . . . . . . . . . Lysiosquilla sulcirostris 

3b. Median carina on rostral plate not flanked by longitudinal grooves (Fig. 4b);dactylus ofraptorial 

claw with 10 to 13 teeth (Fig. 5b) . . . . . . . . . . . . . . . . . . . . . . Lysiosquilla tredecimdentata 

longitudinal 

groove 

a) Lysiosquilla sulcirostris b) Lysiosquilla tredecimdentata 


4a. Distal end of uropodal endopod light (Fig. 6a); dactylus of raptorial claw with 7 to 9 teeth 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lysiosquillina sulcata 

4b. Distal end of uropodal endopod dark (Fig. 6b); dactylus of raptorial claw with 10 or 11 

teeth (usually less in adult females) . . . . . . . . . . . . . . . . . . . . . Lysiosquillina maculata 

7-8 

teeth 

10-13 

teeth 

a) Lysiosquilla sulcirostris b) Lysiosquilla tredecimdentata 

Fig. 5 right claw 



Lysiosquilla sulcirostris Kemp, 1913 

Lysiosquilla tredecimdentata Holthuis, 1941 

Lysiosquillina maculata (Fabricius, 1793) 

Lysiosquillina sulcata (Manning, 1978) 

Lysiosquilloides siamensis Naiyanetr, 1980 (a single record from Thailand) 

no dorsal 

projection 

light 

tip 

dark 

tip 

a) Lysiosquillina sulcata b) Lysiosquillina maculata 

Fig. 6 left uropod (ventral view) 

Reference 

Manning, R.B. 1978. Synopses of the Indo-West-Pacific species of Lysiosquilla Dana, 1852 (Crustacea: Stomatopoda: 

Lysiosquillidae). Smithson. Contrib. Zool., (259):16 p.

Lysioquilidae 837 

Lysiosquillina maculata (Fabricius, 1793) 

En - Common banded mantis shrimp. 

Maximum total length about 38 cm; the characteristic raptorial claw may be 4 cm long. Distinctly 

banded with alternate light and dark bands. Burrows in level bottoms near shore. Collected with spears, 

snares and bait, or at night lights. Widely distributed from Japan and Hawaii to the western Indian 

Ocean. 

male 



female


Harpiosquillidae HARPIOSQUILLIDAE 

Harpiosquillid mantis shrimps 

Diagnostic characters: Very large mantis shrimps 

(maximum total length at least 30 cm). Eye very large, 

T-shaped, cornea strongly bilobed. Carapace, thorax, 

and abdomen with longitudinal ridges. Telson with 

median longitudinal carina and conspicuous posterior 

spines. Posterolateral corners of carapace deeply and 

conspicuously excavate. Raptorial claw very large and 

conspicuous, adapted for spearing. In males outer edge 

of the claw forming an obtuse angle; in females it is evenly 

rounded. Propodus of raptorial claw with irregularly 

spaced, large and small erect spines on margin opposite 

terminal segment (= dactylus). Dactylus of claw 

usually with 7 or 8 large, serrated teeth. 

Habitat, biology, and fisheries: Harpiosquillids are 

burrowers in level bottoms. They use the burrow as a 

refuge while waiting for prey and they also leave the 

burrow to hunt prey. Their large raptorial claws are well 

adapted for capturing fishes. There is no organized fishery 

for these species which reach markets as bycatch. At least 

9 species occur within the area, but only 2 are abundant 

enough to be sold in markets. 


Squillidae: carapace rounded posterolaterally; propodus 

of raptorial claw lined with blunt pectinations. 

dorsal view 

propodus 

of claw 

with small 

spines 

posterolateral 

corner of 

carapace 

excavate 

longitudinal 


Key to the species of Harpiosquillidae occurring in the area 

1a. Carapace lacking median carina (Fig. 1a); distal segment of uropodal exopod entirely 

black (Fig. 2a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harpiosquilla melanoura 

1b. Carapace with median carina (Fig. 1b); distal segment of uropodal exopod with white 

midline, or inner half dark (Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

median 

no median 

carina 

a) 

carina 

b) 



distal 

segment 

black 

distal segment 

not black (inner 

half may be dark) 

a) Harpiosquilla melanoura b) other species 

Fig. 2 left uropod (ventral view) 

(from Manning, 1969)

Harpiosquillidae 839 

2a. Intermediate carinae of thoracic somites with posterior spine (Fig. 3a) . . . . . . . . . . . . . .→ 3 

2b. Intermediate carinae of thoracic somites unarmed (Fig. 3b, c) . . . . . . . . . . . . . . . . . .→ 4 

5 

6 

7 

posterior 

spines 

a) Harpiosquilla annandalei b) Harpiosquilla stephensoni 

Fig. 3 lateral processes of thoracic somites 5-7 


3a. Submedian carinae of fifth abdominal somite with 

5 

posterior spine (Fig. 4); distal segment of uropodal 

exopod black with white midline (Fig, 4); total length 

to 15 cm or less . . . . . . . . . . . . Harpiosquilla annandalei 

3b. Submedian carina of fifth abdominal somite unarmed; 

distal segment of uropodal exopod with inner half 

dark; total length to at least 30 cm . . . . Harpiosquilla raphidea 

th 

abdominal 

somite 

5 

6 

7 

acute 5 

rounded 

c) Harpiosquilla japonica 

4a. Rostral plate with anterior projection (Fig. 5a) . . . . . . . . → 5 

4b. Rostral plate without anterior projection (Fig. 5b) . . . . . . . → 7 Harpiosquilla annandalei 

Fig. 4 posterior part of body 

5a. Marginal carina of telson more than twice as long as 

carina of lateral tooth (Fig. 6a); dactylus of raptorial 

(dorsal view) 

claw with 8 teeth . . . . . . . . . . . . . . Harpiosquilla harpax 

5b. Marginal carina of telson twice or less than twice as long as carina of lateral tooth 

(Fig. 6b); dactylus of raptorial claw with 9 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

anterior 

projection 

of rostral 

plate 

a) Harpiosquilla 

b) Harpiosquilla 

harpax 

stephensoni 



a) Harpiosquilla harpax 

6a. Intermediate carinae of second abdominal somite with posterior spine . . . . Harpiosquilla indica 

6b. Intermediate carinae of second abdominal somite unarmed . . . . . . . . Harpiosquilla philippina 

7a. Dactylus of raptorial claw with 7 teeth; fifth thoracic somite acute laterally (Fig. 3b) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harpiosquilla stephensoni 

7b. Dactylus of raptorial claw with 8 teeth; fifth thoracic somite rounded laterally (Fig. 3c) . . . . . . → 8 

6 

7 

marginal 

carina of 

telson 

carina of 

lateral 

tooth 

b) other species 


posterior 

spine 

white 

midline 

marginal 

carina of 

telson 

carina of 

lateral 

tooth


8a. Rostral plate longer than broad 

(Fig. 7a); marginal carina of telson 

twice as long as carina of lateral tooth 

. . . . . . . . . . . . . Harpiosquilla intermedia 

8b. Rostral plate length and width subequal 

(Fig. 7b); marginal carina of telson 

less than twice as long as carina 

of lateral tooth . . . . . . Harpiosquilla japonica 



Harpiosquilla annandalei (Kemp, 1911) 

Harpiosquilla harpax (De Haan, 1844) 

Harpiosquilla indica Manning, 1969 

Harpiosquilla intermedia Manning and Michel, 1973 

Harpiosquilla japonica Manning, 1969 

Harpiosquilla melanoura Manning, 1968 

Harpiosquilla philippina Garcia, 1978 

Harpiosquilla raphidea (Fabricius, 1798) 

Harpiosquilla stephensoni Manning, 1969 

Reference 

a) Harpiosquilla intermedia b) Harpiosquilla japonica 


Manning, R.B. 1969. A review of the genus Harpiosquilla (Crustacea: Stomatopoda), with descriptions of three new 

species. Smithson. Contrib. Zool., (36):41 p. 

rostral 

plate

Harpiosquillidae 841 

Harpiosquilla harpax (De Haan, 1844) 

En - Robber harpiosquillid mantis shrimp. 

Maximum total length about 25 cm; usually 17 cm or less. Inhabits level bottom habitats, near shore 

to depths of about 100 m. Collected by trawl, trapping, or hook-and-line. Japan to the Red Sea. 

anterior 

projection 


Harpiosquilla raphidea (Fabricius, 1798) 

lateral processes of 

thoracic somites 5-7 

no spine on 

5 th somite of 

thorax 

En - Giant harpiosquillid mantis shrimp. 

The largest known squilloid; maximum total length more than 33 cm, although most specimens range 

from 16 to 29 cm. Inhabits level bottoms in shallow water and can be found in estuaries. Collected in 

traps, by trawls, and by hand. Indo-Malaya and Indonesia to East Africa. 

long anterior 

projection 




sharp 

spine 



5 

6 

5 

6 

7 

8 

7


Squillidae SQUILLIDAE 

Squillid mantis shrimps 

Diagnostic characters: Large mantis shrimps (maximum 

total length more than 20 cm). Eye T-shaped, 

cornea bilobed. Carapace, thorax, and abdomen with longitudinal 

ridges or carinae. Telson with median longitudinal 

ridge and conspicuous posterior spines. 

Posterolateral corners of carapace evenly rounded, not 

excavate. Raptorial claw large and conspicuous, slender, 

adapted for spearing. Dactylus of raptorial claw usually with 

5 or 6 serrated teeth on inner margin. Propodus of raptorial 

claw lined with short, blunt pectinations on margin opposite 

toothed margin of distal segment (= dactylus). 

Habitat, biology, and fisheries: Squillids are burrowers in 

level bottoms and they seek prey at night. They are often 

collected by commercial trawls fishing for penaeid shrimps. 

Some of the larger and more abundant species have been 

reported to be edible and may be found in markets, but there 

is no organized fishery for them in the area. One species of 

Oratosquilla is fished commercially in Japan. 

evenly 

rounded 

Similar families ocurring in the area 

Harpiosquillidae: carapace with posterolateral excavation; 

propodus of raptorial claw lined with erect spines. 

Notes on genera of interest to fisheries 

Species of at least 5 genera, Cloridopsis, Erugosquilla, 

Miyakea, Oratosquilla, andOratosquillina, can be expected 

to be found in markets in the Philippines and in the continental 

parts of the area.These genera can be distinguished as follows: 

Species of Cloridopsis differ from other squillids that might 

dorsal view 


propodus of 

claw with 

blunt 

pectinations 

longitudinal 

ridges on 

body 

be found in markets in having but one broad, anteriorly-curved lateral spine on the fifth thoracic somite, 

the first free somite behind the carapace. In all of the other squillids large enough to be found in markets 

(except Anchisquilla fasciata), the lateral process on the fifth thoracic somite is distinctly bilobed, with an 

anteriorly-directed acute lobe and a shorter, laterally-directed acute lobe. The terminal segment of the 

raptorial claw usually is armed with 5 serrated spines. Adults rarely exceed 10 cm in length. Six species of 

Cloridopsis are known from the Indo-West Pacific, but only C. scorpio is relatively common. 

Members of Erugosquilla can be distinguished by their broad, smooth, carapace, with its anterior margin 

more than half as long as its median length. The carapace is characteristically smooth and shiny, lacking 

any hint of roughness or pits. The median carina of the carapace completely lacks the anterior bifurcation, 

characteristic of members of Miyakea, Oratosquilla, andOratosquillina. The raptorial claw is armed with 

6 serrated teeth on the opposable margin of its terminal segment. Bright blue color on the uropods is 

characteristic of members of this genus, and in at least 1 species the usual rounded lobe between the 

spines of the basal prolongation of the uropod may be replaced by a sharp spine.Erugosquilla woodmasoni 

(Kemp, 1911) is the most common member of this genus, which contains 4 species. 

A single species of Miyakea is abundant enough to be found in markets, and it occurs on level-bottom habitats 

throughout the area. Miyakea nepa has relatively small eyes and a distinctive median carina on the carapace 

that is uninterrupted, splits posterior to the dorsal pit on the midline of the carapace, and extends almost to the 

anterior margin of the carapace. The raptorial claw is armed with 6 teeth on its margin that folds. This is one of 

the most common species of squillids in the western part of the area. Miyakea nepa (Latreille, 1828) is the only 

abundant and widespread member of this genus which contains 1 other species. 

Oratosquilla and Oratosquillina have much larger eyes than species of Miyakea, and in both genera the 

median carina of the carapace splits or bifurcates anterior to the pit on the midline of the carapace. In 

Oratosquilla the median carina of the carapace is entire, uninterrupted, from its base to its bifurcate anterior 

part near the anterior margin. In members of Oratosquillina, the median carina of the carapace is 

interrupted at its bifurcation, and the anterior branches of the bifurcation may be indistinct or absent. In 

both genera the distal segment of the claw usually is armed with 6 teeth, but some species of Oratosquillina 

have but 5 teeth on the claw.

Squillidae 843 

Oratosquilla includes only 4 species, 1 of which, O. oratoria, occurs in the western part of the area, in 

the northern part of Viet Nam. A second species is known from a single record from New Caledonia, and 

a third from a single record in the Philippines. 

Oratosquillina includes 22 species, 3 of which, O. gravieri, O. interrupta, andO. perpensa are common 

enough in the area to be found in markets. Oratosquillina interrupta is one of the most characteristic 

species of the genus, as it can be recognized at once by the convex lobe between the spines of the ventral 

prolongation of the uropod. 

Key to species of interest to fisheries occurring in the area 

1a. Lateral process of fifth thoracic somite a single, anteriorly- or anterolaterally-directed 

spine (Fig. 1a-c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Lateral process of fifth thoracic somite double, with an anteriorly-directed spine and a 

shorter, laterally-directed lobe (Fig. 1d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

2a. Lateral process of fifth thoracic somite a slender, straight spine, directed anterolaterally 

(Fig. 1a); telson with distinct longitudinal carinae on surface . . . . . . . . . . Anchisquilla fasciata 

2b. Lateral process of fifth thoracic somite a broad, anteriorly-curved spine (Fig. 1b, c); 

telson lacking longitudinal carinae on surface . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Rostral plate longer than broad (Fig. 1b); lateral process of fifth thoracic somite with 

black spot (Fig. 1b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cloridopsis scorpio 

3b. Rostral plate broader than long (Fig. 1c); lateral process of fifth thoracic somite lacking 

any dark color (Fig. 1c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cloridopsis immaculata 

5 

rostral plate 

longer than 

broad 

lateral process a 

single spine 

rostral plate 

broader than 

long 

lateral 

process 

double 

6 

5 

single 

spine 

6 

7 

black 

spot 

6 

7 

7 

a) Anchisquilla fasciata b) Chloridopsis scorpio c) Chloridopsis immaculata d) Oratosquilla oratoria 

Fig. 1 anterior part of body and lateral thoracic somites 5-7 (dorsal view) 

4a. Thorax and abdomen completely covered with longitudinal carinae . . . . . . . . . . . . . . . → 5 

4b. Thorax and abdomen with no more than 8 longitudinal carinae . . . . . . . . . . . . . . . . . . → 6 

5a. Submedian carinae of sixth to eighth thoracic somites and fourth to sixth abdominal 

somites ending in spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Keijia lirata 

5b. Most carinae on thorax and abdomen terminating posteriorly in spines . .Carinosquilla multicarinata 

6a. Dorsal surface of carapace smooth, shiny, lacking any trace of an anterior bifurcation 

on the median carina (Fig. 2a, b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

6b. Dorsal surface of carapace pitted or eroded, at least part of an anterior bifurcation of 

the median carina present (Fig. 4a, b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

5


7a. Dorsal surface of telson lacking a line of tubercles on each side of median carina 

(Fig. 3a); rostral plate broader than long (Fig. 2a) . . . . . . . . . . . . . Erugosquilla woodmasoni 

7b. Dorsal surface of telson with a line of tubercles on each side of median carina, the 2 

lines converging posteriorly under apical spine of median carina (Fig. 3b); rostral plate 

longer than broad.(Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . Erugosquilla hesperia 

rostral plate 

broader 

than long 

mediana 

carina 

a) Erugosquilla woodmasoni b) Erugosqilla hesperia 


rostral plate 

longer than 

broad 

median carina 

line of 

tubercles 

median carina 

a) Erugosquilla woodmasoni b) Erugosquilla hesperia 


8a. Median carina of carapace bifurcates posterior to dorsal pit (Fig. 4a) . . . . . . . . . Miyakea nepa 

8b. Median carina of carapace bifurcates anterior to dorsal pit (Fig. 4b, c) . . . . . . . . . . . . . . → 9 

9a. Median carina of carapace entire, not interrupted at base of bifurcation (Fig. 4b) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oratosquilla oratoria 

9b. Median carina of carapace interrupted at base of bifurcation (Fig. 4c) . . . . . . . . . . . . . → 10 

base of 

bifurcation 

of median 

carina 

a) Miyakea nepa 

dorsal pit 

dorsal pit 

base of 

bifurcation 

of median 

b) Oratosquilla oratoria 


10a. Dactylus of raptorial claw with 5 teeth . . . . . . . . . . . . . . . . Oratosquillina quinquedentata 

10b. Dactylus of raptorial claw with 6 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11 

11a. Dorsal ridge on carpus of raptorial claw tuberculate (Fig. 5a); lobe between spines of 

basal prolongation of uropod convex (Fig. 6a) . . . . . . . . . . . . . . . Oratosquillina interrupta 

11b. Dorsal ridge on carpus of raptorial claw smooth, undivided (Fig. 5b); lobe between 

spines of basal prolongation of uropod concave (Fig. 6b) . . . . . . . . . . . . . . . . . . . . → 12 

smooth 

tuberculate 

a) Oratosquillina interrupta b) Oratosquillina perpensa 

Fig. 5 carpus of right raptorial claw (dorsal view) 

carina 

median 

carina 

interrupted 

convex 

lobe 

a) Oratosquillina interrupta 

c) Oratosquillina gravieri 

concave 

lobe 

b) Oratosquillina perpensa 

Fig. 6 basal prolongation of left uropod (ventral view)


12a. Rostral plate broader than long (Fig. 7a); anterolateral spines of carapace extending to 

or overreaching base of rostral plate (Fig. 7a) . . . . . . . . . . . . . . . .Oratosquillina perpensa 

12b. Rostral plate longer than broad (Fig. 7b); anterolateral spines of carapace not overreaching 

base of rostral plate (Fig. 7b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 13 

13a. Lateral process of sixth thoracic somite with broad, rectangular anterior lobe (Fig. 8a) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oratosquillina solicitans 

13b. Lateral process of sixth thoracic somite with slender, triangular anterior lobe, acute 

apically (Fig. 8b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oratosquillina gravieri 

rostral plate 

broader 

than long 

a) Oratosquillina perpensa 

anterolateral 

spine 

b) Oratosquillina gravieri 


rostral 

plate 

longer 

than broad 



Anchisquilla fasciata (De Haan, 1844) 

Carinosquilla multicarinata (White, 1848) 

Cloridopsis immaculata (Kemp, 1913) 

Cloridopsis scorpio (Latreille, 1828) 

Erugosquilla hesperia (Manning, 1968) 

Erugosquilla woodmasoni (Kemp, 1911) 

Keijia lirata (Kemp and Chopra, 1921) 

Miyakea holoschista (Kemp, 1911) 

Miyakea nepa (Latreille, 1828) 

Oratosquilla calumnia (Townsley, 1953) 

Oratosquilla mauritiana (Kemp, 1913) 

Oratosquilla oratoria (De Haan, 1844) 

Oratosquillina anomala (Tweedie, 1935) 

Oratosquillina asiatica (Manning, 1978) 

Oratosquillina fossulata (Moosa, 1986) 

Oratosquillina gravieri (Manning, 1978) 

Oratosquillina inornata (Tate, 1883) 

Oratosquillina interrupta (Kemp, 1911) 

Oratosquillina pentadactyla (Manning, 1978) 

Oratosquillina perpensa (Kemp, 1911) 

Oratosquillina quinquedentata (Brooks, 1886) 

Oratosquillina solicitans (Manning, 1978) 

Oratosquillina stephensoni (Manning, 1978) 

Oratosquillina subtilis (Manning, 1978) 

References 

a) Oratosquillina solicitans b) Oratosquillina gravieri 

Fig. 8 lateral processess of thoracic somites 5-7 


Manning, R.B. 1971. Keys to the species of Oratosquilla, (Crustacea: Stomatopoda), with descriptions of two new 

species. Smithson. Contrib. Zool., (71):16 p. 

Manning, R.B. 1978. Further observations on Oratosquilla, with accounts of two new genera and nine new species 

(Crustacea: Stomatopoda: Squillidae). Smithson. Contrib. Zool., (272):44 p. 

5 

6 

7 

broad 

lobe 

6 

7 

5 

triangular, 

acute lobe


Cloridopsis scorpio (Latreille, 1828) 

En - Spotted squillid mantis shrimp. 

Maximum total length 10 cm or less. A shore species not known to occur in brackish water. Taken by 

traps. The most common species among the 6 Cloridopsis known from the Indo-West Pacific. Known 

from Viet Nam, Indonesia, and Malaysia to Pakistan. 

anterior 

curved 

spine 


Erugosquilla woodmasoni (Kemp, 1911) 

En - Smooth squillid mantis shrimp. 

Maximum total length about 15 cm. A common shallow water species inhabiting burrows on level 

bottoms. Commonly taken by trawlers. Widely distributed from Japan to the western Indian Ocean. 



black 

spot 

5 

6 

7 



bilobed


Miyakea nepa (Latreille, 1828) 

En - Smalleyed squillid mantis shrimp. 

Maximum total length about 17 cm. A very common shore species that burrows in level-bottom habitats 

and one of the most common squillids in the western part of the area. Usually taken by trawls. Known 

from Taiwan Province of China to the Red Sea. 


Oratosquilla oratoria (De Haan, 1844) 

En - Japanese squillid mantis shrimp. 

Maximum total length more than 18 cm. The common commercial species in Japan. Burrows in 

near-shore level-bottom habitats. Taken by trawls and traps. Known from Japan to Hong Kong (China) 

and Viet Nam, rare in southern part of its range. 



median carina bifurcates 

posterior to dorsal pit 

5 

6 

7 


thoracic somites 5-7


Oratosquillina gravieri (Manning, 1978) 

En - Vietnamese squillid mantis shrimp. 

Maximum total length about 11 cm. Burrows in level bottoms in depths of 15 to 25 to more than 100 m. 

Taken by trawls. Known only from Viet Nam, where it is a common species, and the Philippines. 



Oratosquillina perpensa (Kemp, 1911) 

5 

6 

7 



En - Common squillid mantis shrimp. 

Maximum total length about 10 cm. Occurs in sublittoral, level-bottom habitats, in depths of 100 m or 

less. Usually taken in trawls. Known from localities between southern Taiwan Province of China and 

Myanmar. 



5 

6 

7 


thoracic somites 5-7


Oratosquillina quinquedentata (Brooks, 1886) 

En - Fivespined squillid mantis shrimp. 

Maximum total length 14 cm. Occurs in sublittoral, level-bottom habitats in depths of at least 50 m. 

Usually taken in trawls. Known from the Arafura Sea, Gulf of Thailand, and Bombay, India. 



Oratosquillina solicitans (Manning, 1978) 

En - Variable squillid mantis shrimp. 

Maximum total length less than 10 cm. Occurs in level-bottom habitats near shore. Usually taken in 

trawls. Known from the western Pacific, Taiwan Province of China, the Gulf of Thailand, Malaysia, and 

Indonesia. 



5 

6 

7 



5 

6 

7 





SHRIMPS AND PRAWNS 

by T.Y. Chan

852 Shrimps and Prawns 


antennule 

antennal 

scale 

antenna 

antennal 

flagellum 

gastrofrontal 

groove 

gastrofrontal 

crest 

postorbital spine 

antennal spine 

postantennal spine 

pterygostomian 

spine 

length of 

rostrum 

length of 

carapace 

antennular 

flagella 

3 rd 

maxilliped 

branchiostegal 

spine 

rostrum 

gastro-orbital crest 

antennal 

crest 

exopod 

cervical 

groove 

hepatic crest 

epigastric (or last 

rostral) tooth 

adrostral 

crest 

suprahepatic spine 

hepatic spine 

transverse or 

vertical suture 

carapace (lateral view) 

rostrum 

non-grooved 

carapace 

carapace (dorsal view) 

carapace 

length 

grooved 

carapace 

pereiopods or legs 

apical 

portion 

(tip) 

orbital margin 

1 

branchiocardiac 

crest 

longitudinal 

suture 

gastrofrontal crest 

gastrofrontal groove 

gastro-orbital crest 

median groove 

postrostral crest 

adrostral groove 

adrostral crest 

1-6 abdominal segments 

(or somites) 

endopod 

2 

pleopods 

(or abdominal 

appendages) 

lateral view 

stridulating 

organ 

(or ridges) 

body length 

3 

dactylus 

antennule 

basis 

coxa 

epipod 

dorsal 

ventral 

4 

uropod 

pincer 

podobranch 

ischium 

flagella 

3 rd (or distal) 

article 

2 nd article 

distolateral 

spine 

1 st (or basal) 

article 

stylocerite 

5 

length of 

abdomen 

6 

endopod 

exopod 

propodus 

exopod 

arthrobranch 

pleurobranch 

carpus 

merus 

pereiopod with branchiae 

exopod 

podobranch 

epipod 

dorsal 

crest 

arthrobranch 

pleurobranch 

telson 

body 

ischium 

basis 

coxa 

proximal part of pereiopod 

(schematic) 

fixed spines 

movable 

spines 

types of spined telsons

Technical Terms and Measurements 853 

penultimate 

thoracic 

segment 

(sternite XIII) 

last thoracic 

segment 

(sternite XIV) 

anterior 

plate 

posterior 

plate 

anterior 

plate 

posterior 

transverse 

ridge 

coxal plate 4 th 

leg 

intermediate 

plate 

anterior sternal 

plate 

posterior 

sternal plate 

anterior process 

RST 

RST 

lateral 

plates 

thelycal plate 


basic types of thelycum of 

female penaeid shrimps 


4 

4 

5 

left pereiopod of 

4 th pair 

5 

posterior 

process 

left pereiopod of 

5 th pair 

Penaeus 

proximolateral 

lobes 

Parapenaeopsis 

Metapenaeus 

right 

distolateral 

projection 

Metapenaeopsis 

distomedian projections 

distolateral 

projections 


distolateral 

projections 

left 

distomedian 

projection 

outer 

intermediate 

strip 

distomedian 

projection 

ventral costa 

median lobe 

lateral lobe 

inner 

intermediate 

strip 

distal part 

(dorsal view) 

distomedian 

lobule 

basic types of petasma 

(joined endopods of first pair of pleopods) 

of male penaeid shrimps 

(ventral view - except otherwise stated)



Shrimps and prawns constitute a large group of crustaceans with an extended abdomen (or “tail”), varying 

in size from microscopic to about 35 cm body length (measured dorsally from the posterior orbital 

margin to the end of the tail, excluding the rostrum and the appendages). Taxonomically, shrimps and 

prawns belong to the “swimming group” of decapod crustaceans in the suborder Macrura Natantia. They 

differ from the lobsters (suborder Macrura Reptantia) by having the body generally more laterally compressed, 

the pleopods (abdominal appendages) well developed, the thoracic sternum (i.e. ventral part of 

thoracic body segments, between the legs) often narrow and not easy to observe, the first abdominal 

pleuron (or lateral plate) well developed (about as large as the pleura of following segments, see figure 

below), and the telson usually tapering distally. 

1 st abdominal pleuron 

well developed 

pleopods 

long 

shrimps and prawns 

lobsters 

(after Chan and Yu, 1993) 

pleopods short 

1 st abdominal 

pleuron 

reduced 

conspicuous morphological differences between shrimps and lobsters 

body laterally 

compressed 

telson usually 

pointed 

body dorsoventrally 

depressed 


broadly convex


The terms “shrimp” and “prawn” have no definite reference to any known taxonomic groups. Although the 

term “shrimp” is sometimes applied to smaller species, while “prawn” is more often used for larger forms, 

there is no clear distinction between both terms and their usage is often confused or even reverse in 

different countries or regions. Therefore, no attempt has been made here to restrict or define their meaning. 

Certain other crustaceans, such as the “mysid shrimps” (Mysidacea), “mantis shrimps” (Stomatopoda), 

and “mud shrimps” (Thalassinidea), are taxonomically not true shrimps. 

Altogether, there are about 3 047 species of shrimps and prawns known to date, subdivided into 4 major 

groups, namely Sergestoidea (about 94 species), Penaeoidea (about 376 species), Stenopodidea (at least 

60 species), and Caridea (at least 2 517 species). Although the Caridea comprise the majority of species, 

only some are abundant enough to be of interest to fisheries. Most of the commercial shrimps and prawns 

belong to the Penaeoidea. At present, only slightly less than 300 species of shrimps and prawns are of 

economic interest worldwide, and out of these, only about 100 comprise the principal share of the annual 

world catch. FAO’s Yearbook of Fishery Statistics reports in 1995 a worldwide production of all shrimps 

and prawns of around 3 200 000 t (both from capture fishery and aquaculture). Around 710 000 t of this 

production originated in the Western Central Pacific. 

The exact number of species of shrimps and prawns present in the Western Central Pacific is uncertain. 

This is especially true for the carideans and stenopodids, most species of which have no economic value 

and thus only very few studies exist on them. However, recent extensive studies on carideans from the 

Philippines and adjacent areas have shown that 528 species occur in that region alone. Although caridean 

shrimps are widely distributed in marine waters, brackish and fresh waters, and are found from high 

mountain regions to coral reefs and the deep sea, at present only the giant river prawn Macrobrachium 

rosenbergii is of high economic importance in the Western Central Pacific. This is a very large species, 

sometimes found in marine waters, and extensively fished and cultured in several countries. The other 

coastal or fresh-water caridean shrimps in the area are either too small or not abundant enough to be 

fished on a large scale, although a few of them may locally be used as food. It should be noted, however, 

that the present commercial fishing activities in the area are generally rather simple and mainly limited to 

shallow waters with depths less than 100 m. Several deep-sea caridean shrimps, mostly belonging to the 

family Pandalidae, can often be caught in large quantities during exploratory trawling operations and may 

eventually prove to be of commercial interest with the development of a deep-sea fishery. 

The Stenopodidea (with the single family Stenopodidae) generally have no economic importance, although 

a few of them, as well as some coral reef carideans, are sporadically seen in the aquarium trade and thus 

have some commercial value. 

Most of the commercial species of shrimps and prawns belong to the Penaeoidea. Studies on penaeoids 

are more comprehensive and at present 4 families including 191 species are known to occur in the Western 

Central Pacific, with the Penaeidae being the most important family. As species of the Penaeidae are 

generally of moderate to large size and often occur in large quantities in shallow waters along the 

continental shelf on trawlable bottoms, they are fished extensively by trawls, seines, set nets, traps, and 

artisanal gear. Large-scale pond culture of penaeid shrimps is practised in several countries. Species of 

the penaeoid families Aristeidae and Solenoceridae are mainly deep-water dwellers and largely 

unexploited. The fact that larger representatives of these 2 families are often caught on the basis of 

exploratory deep-sea trawling, indicates that they have a high commercial potential with the future 

development of a deep-sea fishery in the area. In contrast, species of the penaeoid family Sicyoniidae are 

generally small, nowhere abundant, and do not have any commercial potential. 

Sergestoid shrimps are usually small and of no interest to fisheries, except for the genus Acetes, 7 species 

of which are found in the Western Central Pacific. These epipelagic shrimps inhabit shallow coastal 

estuarine waters and often occur in great abundance. They are extensively fished by push nets, bag nets 

and seines, and are of considerable economic importance, particularly in the Southeast Asian countries of 

the area. 

Shrimps and prawns in the Western Central Pacific are generally marketed fresh or frozen, sometimes 

live, except for species of Acetes which are usually processed into shrimp paste. They are locally consumed 

or exported. In the Philippines, Indonesia and Thailand, altogether 94 200 t of shrimps and prawns were 

exported in 1987.


Guide to Major Groups GUIDE TO THE MAJOR GROUPS OF SHRIMPS AND PRAWNS 

OCCURRING IN THE AREA 

SERGESTOIDEA Page 858 

Sergestoid shrimps 

Usually small to microscopic, body strongly 

compressed laterally, shell rather soft; rostrum 

and last 2 pairs of legs (pereiopods) reduced 

(absent in Luciferidae); abdomen with posterior 

part of pleura (lateral plates) covering anterior 

part of succeeding pleura; males with large 

copulatory organ (petasma) on first pair of 

pleopods (abdominal appendages); generally 

pelagic, with eggs released directly into the water 

(eggs carried on second pair of legs in 

Luciferidae). 

PENAEOIDEA Page 866 

Penaeoid shrimps 

Small to large; all 5 pairs of legs (pereiopods) well 

developed, with first 3 pairs forming a pincer, 

none of the pincers particularly large; abdomen 

with posterior part of pleura (or lateral plates) 

covering anterior part of succeeding pleura; with 

large specific copulatory organ on first pair of 

pleopods (abdominal appendages) in males 

(petasma), and on posterior thoracic sternites in 

females (thelycum); eggs released directly into 

the water, not retained by the females. 

STENOPODIDEA Page 955 

Stenopodid shrimps 

Usually small; all 5 pairs of legs (pereiopods) well 

developed, with first 3 pairs forming a pincer, third 

pair huge and massive; abdomen with posterior 

part of pleura (lateral plates) covering anterior 

part of succeeding pleura; males and females 

without large specific copulatory organ on first 

pair of pleopods (abdominal appendages) or 

posterior thoracic sternites, respectively; females 

carry the eggs on the abdomen until hatching. 

CARIDEA Page 957 

Caridean shrimps 

Size very small to large; all 5 pairs of legs 

(pereiopods) well developed, the first 2 pairs with 

or without pincer, but third pair never bearing a 

pincer; second abdominal pleuron (lateral plate) 

greatly expanded, pear-shaped and overlapping 

posterior part of first pleuron and anterior part of 

third pleuron; males and females without large 

specific copulatory organ on first pair of pleopods 

(abdominal appendages) or posterior thoracic 

sternites, respectively; females carry the eggs on 

the abdomen until hatching. 

3 rd maxilliped 

2 

first 3 legs 

with 

pincers 

1 

2 

3 

3 

no pincer 

1 

2 

3 

1 

rostrum short or absent 

Sergestoidea 

4 

Penaeoidea 

1 

2 

Stenopodidea 

4 

5 

4 

5 

2 nd pleuron 

not expanded 

anteriorly 

5 

2 nd pleuron 

pear-shaped 

females carry eggs 

on abdomen 

Caridea 

3 rd leg enlarged 

2 nd pleuron not 

expanded 

anteriorly 

females 

carry eggs 

on 

abdomen 

3

List of Families 857 

List of Families LIST OF FAMILIES OCCURRING IN THE AREA 


Infraorder PENAEIDEA 

Superfamily SERGESTOIDEA 

LUCIFERIDAE 

SERGESTIDAE 

Superfamily PENAEOIDEA 

ARISTEIDAE 

SOLENOCERIDAE 

PENAEIDAE 

SICYONIIDAE 

Infraorder STENOPODIDEA 

STENOPODIDAE 

Infraorder Caridea 

Superfamily PASIPHAEOIDEA 

PASIPHAEIDAE 

Superfamily OPLOPHORIDEA 

OPLOPHORIDAE 

Superfamily ATYOIDEA 

ATYIDAE 

Superfamily BRESILIOIDEA 

BRESILIIDAE 

Superfamily NEMATOCARCINOIDEA 

EUGONATONOTIDAE 

NEMATOCARCINIDAE 

RHYNCHOCINETIDAE 

Superfamily PSALIDOPODOIDEA 

PSALIDOPODIDAE 

Superfamily STYLODACTYLOIDEA 

STYLODACTYLIDAE 

Superfamily CAMPYLONOTIDEA 

BATHYPALAEMONELLIDAE 

Superfamily PALAEMONOIDEA 

ANCHISTIOIDIDAE 

GNATHOPHYLLIDAE 

HYMENOCERIDAE 

PALAEMONIDAE 

Superfamily ALPHEOIDEA 

ALPHEIDAE 

HIPPOLYTIDAE 

OGYRIDIDAE 

Superfamily PROCESSOIDEA 

PROCESSIDAE 

Superfamily PANDALOIDEA 

PANDALIDAE 

THALASSOCARIDIDAE 

Superfamily CRANGONOIDEA 

CRANGONIDAE 

GLYPHOCRANGONIDAE


Infraorder Penaeidae, Superfamily Sergestoidea Infraorder PENAEIDEA 

Superfamily SERGESTOIDEA 

Sergestoid shrimps 

Diagnostic characters: Usually microscopic to small sized, with a body length from 1 to about 5 cm 

(exceptionally over 8.5 cm). Body strongly compressed laterally, shell soft. Carapace with crests and 

grooves poorly developed, often wanting. Rostrum very short and small, sometimes absent. In males, lower 

antennular flagella with a clasping organ. First leg with or without pincer, second and third legs bearing small 

pincers; fourth and fifth legs reduced or absent. Abdomen with posterior part of pleura (lateral plates) 

covering anterior part of succeeding pleura. Males with large copulatory organ (petasma) on first pair of 

pleopods (abdominal appendages). Generally pelagic; eggs released directly into the water (family Sergestidae), 

or carried on second pair of legs until hatching (family Luciferidae). 


3 rd 

maxilliped 

2 

3 

Habitat, biology, and fisheries: Members of this superfamily (including 2 families and 7 genera) mainly inhabit 

brackish and marine environments (a single species is found in pure fresh water). They can be found from 

shallow to deep waters (deeper than 2 100 m) and are generally pelagic, although a few (in the genus Sicyonella) 

have adapted to a benthic way of life. At present, 2 families and 4 genera of sergestoid shrimps are known from 

the Western Central Pacific, but all except the genus Acetes are without any economic importance as they are 

either too small, not abundant enough, or occur in very deep water. Members of the genus Acetes mainly occur 

in estuarine or shallow coastal waters and are seasonally very abundant. These are small shrimps with a body 

length of adults ranging between 1 and 4 cm. Their bodies are translucent or semi-translucent, with black eyes 

and several pairs of red pigment spots (chromatophores) on the bases of uropods. In the course of their fishing 

seasons, they are extensively caught by push nets, bag nets, and seines.They are mainly fished in the Southeast 

Asian countries of the area and are of significant commercial importance. From 1990 to 1995, the reported 

annual catch of sergestoid shrimps in the Western Central Pacific ranged from around 38 500 to 45 700 t (FAO 

Yearbook of Fishery Statistics).Only a small part of the catch is marketed fresh, and the greater fraction is dried, 

salted or fermented with salt and processed into shrimp paste. As only species of Acetes are of commercial 

interest, a key to species of this genus is given here. 

Other major groups of shrimps and prawns occurring in the area 

Penaeoidea: all 5 pairs of legs well developed, with first 

3 pairs forming a pincer, none of the pincers particularly 

large; rostrum usually well developed, extending 

beyond eyes; numerous branchiae (more than 8 on 

each side). 

1 

first 3 legs 

with pincer 

1 

2 

3 

rostrum well developed 

4 5 

Penaeoidea

Infraorder Penaeidae, Superfamily Sergestoidea 859 

Stenopodidea: all 5 pairs of legs well developed, first 3 pairs forming a pincer, third pair huge and massive; 

males without large copulatory organ on first pair of pleopods; females carry the eggs on the abdomen 

until hatching. 

Caridea: all 5 pairs of legs well developed, third pair without pincer; abdomen with pleuron of second 

segment greatly expanded, overlapping those of first and third segments; males without large copulatory 

organ on first pair of pleopods; females carry the eggs on the abdomen until hatching. 

1 

2 

Stenopodidea 


Key to families and genera 1/ of Sergestoidea occurring in the area 

1a. Head greatly elongated (Fig. 1); 

no branchiae; size very small, 

about 1 cm body length . . . . . . Luciferidae 

1b. Head not particularly elongate; 

branchiae present; size small, but 

generally adults with more than 

2cmbodylength. . . . . . (Sergestidae) → 2 

4 

5 

3 

females 

carry eggs 

on abdomen 

5 

4 

females carry 

eggs on abdomen 

2 nd pleuron 

expanded, 

pear-shaped 

2a. Fourth and fifth legs entirely lacking, 

reduced to a pair of protuberances 

(genital coxae) in males 

(Fig. 2); first maxillae and first 

maxillipeds without palp; second 

maxillae with a single undivided 

Fig. 1 Luciferidae 

lobe . . . . . . . . . . . . . . . . . . . Acetes 

2b. Fourth and fifth legs present, with fifth leg much shorter than fourth (Fig. 3); first maxillae 

and first maxillipeds with palp; second maxillae with 2 lobes . . . . . . . . . . . . . . other genera 


2 

1 

Fig. 2 Acetes 

3 rd leg 

genital coxa 

1 

3 rd 

maxilliped 

2 3 

maxillipeds 

2 

Caridea 

1 

3 

head elongated 

1 

2 

Fig. 3 Sergestes 

1/ Restricted to the identification of Acetes, the only genus of interest to fisheries in the area. 

3 

4 

5 th leg


Sergestidae SERGESTIDAE 

Key to sexes of Acetes 

1a. A pair of protuberances 

(genital coxae; Fig. 2) 

between third legs and 

first pleopods; lower 

antennular flagella with 

1 or 2 clasping spines, 

or modification of these 

(Fig. 4); petasma 

(Fig. 5) present on first 

pleopods . . . . . . . . . . . . male 

1b. No protuberance in 

genital area; lower antennular 

flagella without 

spine; petasma 

absent . . . . . . . . . . . . female 

Key to the species of Acetes occurring in the area 

Remark on key characters: see the respective species accounts for illustrations of the petasma and lower 

antennular flagellum of males, and the basis of third leg of females. 

Females 

1a. Apex of telson rounded or truncated (Fig. 6a) . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Apex of telson triangular (Fig. 6b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2a. Third thoracic sternite produced posteriorly (Fig. 7b) . . . . . . . . . . . . . . . Acetes japonicus 

2b. Third thoracic sternite not produced posteriorly (Fig. 7a) . . . . . . . . . . . . . Acetes serrulatus 

a) rounded or truncated b) triangular 

Fig. 6 apex of telson 

clasping 

spine 

Fig. 4 lower 

antennular flagellum 

of a male 

pars 

astringens 

capitulum 

processus 

ventralis 

capitulum 

Fig. 5 examples of the petasma 

a) b) 

Fig. 7 base of third leg 

3a. Procurved tooth present between bases of first pleopods . . . . . . . . . . . . . . . . . . . . . → 4 

3b. Procurved tooth absent between bases of first pleopods . . . . . . . . . . . . . . . . . . . . . → 6 

4a. Inner margin of basis of third leg with sharply pointed projection; third and fourth thoracic 

sternites deeply channeled longitudinally . . . . . . . . . . . . . . . . . . . . . . . .Acetes indicus 

4b. Inner margin of basis of third leg without sharply pointed projection; third and fourth 

thoracic sternites not channeled longitudinally. . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

5a. First segment of antennular peduncle at most as long as second and third segments 

together; distal inner margin of basis of third leg ending in blunt projection . . . Acetes intermedius 

5b. First segment of antennular peduncle longer than second and third segments together; 

distal inner margin of basis of third leg without projection . . . . . . . . . . . . . Acetes erythraeus

Sergestidae 861 

6a. Lower antennular flagellum with 20 segments or less; distal inner margin of basis of 

third leg ending in projection; pair of small protuberances on anterior part of third 

thoracic sternite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acetes sibogae 

6b. Lower antennular flagellum with 20 segments or more; distal inner margin of basis of 

third leg without projection; pair of large protuberances on anterior part of third thoracic 

sternite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Acetes vulgaris 

Males 

1a. Anterior margin of genital coxa rounded; petasma without pars astringens (Fig. 5) . . . . . . . → 2 

1b. Anterior margin of genital coxa pointed; petasma with pars astringens (Fig. 5) . . . . . . . . . → 4 

2a. Procurved tooth present between bases of first pleopods; lower antennular flagellum 

with 1 clasping spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acetesindicus 

2b. Procurved tooth absent between bases of first pleopods; lower antennular flagellum with 

2 clasping spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Lower antennular flagellum with triangular projection from upper end of first segment of 

main branch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acetes serrulatus 

3b. First segment of main branch of lower antennular flagellum without triangular projection 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acetes japonicus 

4a. Procurved tooth present between bases of first pleopods . . . . . . . . . . . . . . . . . . . . .→ 5 

4b. Procurved tooth absent between bases of first pleopods . . . . . . . . . . . . . . . . . . . . .→ 6 

5a. First segment of antennular peduncle shorter than second and third segments together; 

capitulum of petasma with 3 to 5 subequally large hooks along outer margin . . Acetes intermedius 

5b. First segment of antennular peduncle longer than second and third segments together; 

capitulum of petasma with 1 large hook at outer margin . . . . . . . . . . . . . . Acetes erythraeus 

6a. Lower antennular flagellum with 12 segments or less; capitulum of petasma with 1 large 

hook and often additionally 1 small hook along outer margin . . . . . . . . . . . . . Acetes sibogae 

6b. Lower antennular flagellum with 17 segments or more; capitulum of petasma with 3 large 

hooks along outer margin . . . . . . . . . . . . . . . . . . . . . . . . . . . .Acetes vulgaris 

List of genera and commercial species occurring in the area 


LUCIFERIDAE 

Genus Lucifer 

SERGESTIDAE 

Genus Acetes 

Acetes erythraeus Nobili, 1905 

Acetes indicus H. Milne Edwards, 1830 

Acetes intermedius Omori, 1975 

Acetes japonicus Kishinouye, 1905 

Acetes serrulatus (Krøyer, 1855) 

Acetes sibogae Hansen, 1919 

Acetes vulgaris Hansen, 1919 

Genus Sergestes 

Genus Sergia 

Genus Sicyonella 

References 

Miquel, J.C. 1984. Shrimps and Prawns. In FAO species identification sheets for fisheries purposes. Western Indian 

Ocean (Fishing Area 51), edited by W. Fischer and G. Bianchi. Rome, FAO. 

Omori, M. 1975. The systematics, biogeography, and fishery of epipelagic shrimps of the genus Acetes (Crustacea, 

Decapoda, Sergestidae). Bull. Ocean Res. Inst., Univ. Tokyo, 7:1-89. 

Pérez Farfante, I. and B.F. Kensley. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnosis 

for the families and genera. Mem. Mus. Natn. Hist. Nat., 175:1-233.


Acetes erythraeus Nobili, 1905 

En - Tsivakihini paste shrimp; Fr - Chevrette tsivakihini; Sp - Camaroncillo tsivakihini. 

Maximum body length 1.6 to 4.0 in females (rarely 4.8 cm) and 1.6 to 3.2 cm in males. Epipelagic, 

found over muddy or sandy bottoms, from the surface to a depth of 55 m. Marine or brackish, but 

usually brackish and fished in the intertidal zone and estuaries with mangroves. Probably the most 

common species of the genus in the area and of major commercial importance in its range, despite 

its very small size. Caught with triangular nets, lift nets, scoop nets, push nets, bag nets, set filter 

nets, and seines, occasionally encountered in penaeid shrimp culture ponds. Marketed dried, boiled, 

salted, fermented with salt, fresh, or processed in other ways; consumed locally, mainly in the form 

of seasoning such as shrimp paste or shrimp sauce, sometimes exported. Indo-West Pacific from 

the eastern coast of Africa to southern China and northeastern Australia. 

petasma 

Acetes indicus H. Milne Edwards, 1830 

En - Jawla paste shrimp; Fr - Chevrette jawla; Sp - Camaroncillo javlá. 

Maximum body length 2.3 to 4.0 cm (females) and 1.5 to 2.5 cm (males). Inhabits shallow, 

sometimes brackish coastal waters; epipelagic, usually swims in midwater or near the surface. One 

of the more common species of the genus in the area and of considerable economic importance. 

Caught with push nets, bag nets and seines, and sometimes by light fishing at night. Taken 

throughout its range. Marketed dried, boiled, salted, fermented with salt, fresh, or processed in other 

ways; consumed locally, mainly in the form of seasoning such as shrimp paste or shrimp sauce, 

sometimes exported. Indo-West Pacific from India to Viet Nam and Indonesia. 

sharp projection of basis 

1 large hook at 

outer margin 

bases of 3 rd legs (female) 

without pars 

astringens 

basis 


petasma 

(after Omori, 1975) 

lower antennular 

flagellum (male) 


flagellum (male)


Acetes intermedius Omori, 1975 

En - Taiwan mauxia shrimp; Fr - Chevrette mauxia de Formose; Sp - Camaroncillo mauxia dè Formosa. 

Maximum body length 2.0 to 2.6 cm (females) and 1.7 to 2.4 cm (males). Epipelagic, found mainly 

at sea. Within the area, reported from commercial catches made in the Philippines and Indonesia, 

and probably of moderately commercial importance. Caught with midwater trawls, triangular nets, 

lift nets, and scoop nets, and sometimes by light fishing at night. Marketed dried, boiled, salted, 

fermented with salt, fresh, or processed in other ways; consumed locally, mainly in the form of 

seasoning such as shrimp paste or shrimp sauce, sometimes exported. Found in the western Pacific, 

but so far only reported from Taiwan Province of China, Philippines, and the southern coast of Java. 

blunt projection of basis 


petasma 

3-5 large hooks 

along outer 

margin 

Acetes japonicus Kishinouye, 1905 




(after Yu, 1974) 

En - Akiami paste shrimp; Fr - Chevrette akiami; Sp - Camaroncillo akiami. 

Maximum body length 1.5 to 3.0 cm (females) and 1.1 to 2.4 cm (males). Epipelagic, inhabits shallow 

coastal waters over muddy bottoms. One of the more common species of the genus in the area and of 

considerable economic importance; reported from commercial catches made in Viet Nam, Thailand, and 

Malaysia. Caught with push nets, bag nets, and seines, and sometimes by light fishing at night.Marketed 

dried, boiled, salted, fermented with salt, fresh, or processed in other ways; consumed locally, mainly in 

the form of seasoning such as shrimp paste or shrimp sauce, sometimes exported.In 1995, the reported 

production from culture ponds of this species in Indonesia amounted to 3 500 t (FAO Aquaculture 

Production Statistics). Widely distributed in the Indo-West Pacific from the Persian Gulf to Japan and 

Indonesia. 


without 

pars 

astringens 

petasma 




2 clasping 

spines 

(after Hayashi, 1992)


Acetes serrulatus (Krøyer, 1855) 

En - Southern mauxia shrimp; Fr - Chevrette mauxia méridionale; Sp - Camaroncillo mauxia sureño. 

Maximum body length 1.5 to 2.1 cm (females) and 1.2 to 1.7 cm (males). Epipelagic, inhabits shallow 

coastal water. Probably less common than the other species of the genus. Within the area, reported 

from commercial catches made in Malaysia and Indonesia, but no further information on its 

economic status is presently available. Caught by push nets, bag nets, and seines. Marketed dried, 

boiled, salted, fermented with salt, fresh, or processed in other ways; consumed locally, mainly in 

the form of seasoning such as shrimp paste or shrimp sauce, sometimes exported. Western Pacific 

and so far known only from southern China, Singapore, Malaysia, and Indonesia. 


without pars 

astringens 

triangular 

projection 

petasma 


Acetes sibogae Hansen, 1919 



2 clasping 

spines 

En - Alamang shrimp; Fr - Chevrette alamang; Sp - Camaroncillo alamang. 

Maximum body length 1.4 to 3.4 cm (females) and 1.3 to 2.5 cm (males). Epipelagic, found over 

muddy bottoms in estuarine and marine waters to a depth of 55 m. Probably caught throughout its 

range in the area and of moderate commercial importance. Taken by triangular nets, lift nets and 

scoop nets. Marketed dried, boiled, salted, fermented with salt, fresh, or processed in other ways; 

consumed locally, mainly in the form of seasoning such as shrimp paste or shrimp sauce, sometimes 

exported. Indo-West Pacific from India to the Philippines and eastern Australia (the Australian 

population and certain specimens from India are sometimes considered to be 2 subspecies). 

projection of basis 


petasma 

1 large hook 

at outer 

margin 


12 

segments 

or less 


flagellum (male)


Acetes vulgaris Hansen, 1919 

En - Jembret shrimp; Fr - Chevrette jembre; Sp - Camaroncillo jembre. 

Maximum body length 2.0 to 3.4 cm (females) and 1.7 to 2.6 cm (males). Epipelagic, found over 

sandy and muddy bottoms in marine waters from depths of 9 to 55 m. One of the more common 

species of the genus in the area and of considerable economic importance; reported from 

commercial catches made in Thailand, Singapore, and Indonesia. Caught with push nets, bag nets, 

scoop nets, and seines, sometimes by light fishing at night. Marketed dried, boiled, salted, fermented 

with salt, fresh, or processed in other ways; consumed locally, mainly in the form of seasoning such 

as shrimp paste or shrimp sauce, sometimes exported. Western Pacific from southern China to the 

Strait of Malacca and Indonesia. 


petasma 

3 large hooks 

along outer 

margin 




17 segments 

or more 




Superfamily Penaeoidea Superfamily PENAEOIDEA 

Penaeoid shrimps 

Diagnostic characters: Small to large sized, with a body length from 2.5 to about 35 cm. All 5 pairs of 

legs well developed, with first 3 pairs of legs forming a pincer, none of the pincers particularly large. 

Abdomen with posterior part of pleura (lateral plates) covering anterior part of succeeding pleura. 

With large copulatory organ, on first pair of pleopods in males (petasma), and on posterior thoracic 

sternites in females (thelycum). Eggs are released directly into the water and not retained by the females 

on the abdomen. 

posterior part of pleura 

covering anterior part of 

succeeding pleura 

1 

first 3 legs 

with pincer 

2 

5 

4 

3 

Habitat, biology, and fisheries: Members of this superfamily are all marine, although the juveniles of some 

species of Penaeidae inhabit brackish water and occasionally are even found in almost fresh water. The 4 

families of penaeoids can all be found in the Western Central Pacific, with the family Penaeidae being of 

great economic importance in capture fisheries and aquaculture. Members of the 2 families Aristeidae and 

Solenoceridae mainly occur in deep water and are presently not exploited in the area. As some of them 

can reach a large size and are often taken on the basis of exploratory deep-water trawling, they may 

eventually prove to be of commercial interest. In contrast, species of the family Sicyoniidae are usually 

small and nowhere abundant. They are caught incidentally in prawn fisheries but do not have any economic 

importance. 


Sergestoidea: usually small sized to microscopic; body strongly compressed laterally; shell rather soft; 

rostrum as well as last 2 pairs of legs reduced or absent; branchiae few, not more than 8 on each side. 

Stenopodidea: third pincer very large and massive; males and females without large copulatory organ on 

first pair of pleopods or posterior thoracic sternites, respectively; females carry the eggs on the abdomen 


2 

3 

3 rd 

maxilliped 

1 


Sergestoidea 

1 

4 

2 

Stenopodidea 

5 

females 

carry 

eggs on 

abdomen 

3 

3 rd leg 

enlarged

Superfamily Penaeoidea 867 

Caridea: third leg always without pincer; pleuron 

of second abdominal segment greatly expanded 

and overlapping those of first and third segments; 

males and females without large copulatory 

organ on first pair of pleopods or posterior 

thoracic sternites, respectively; females carry the 

eggs on the abdomen until hatching. 

Key to the families of Penaeoidea occurring in the area 

1a. Either rostrum very short and armed with 1 or 2 upper teeth only, or upper antennular 

flagellum very short and attached to the base of distal antennular segment (Fig. 1) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aristeidae (p. 868) 

1b. Rostrum always armed with more than 3 upper teeth, and both upper and lower 

antennular flagella of similar length and attached to the tip of antennular peduncle . . . . . . . → 2 

2a. Pleopods (abdominal appendages) with 1 branch only; abdomen often with many 

distinct furrows and grooves (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . Sicyoniidae (p. 952) 

2b. Pleopods (abdominal appendages) with 2 branches; abdomen without or with very few 

distinct grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

upper 

antennular 

flagella 

very short 

(Aristeinae) 

Fig. 1 Aristeidae 

grooves 

3 rd and 4 th pleopods 

single-branched 

Fig. 2 Sicyoniidae 

3a. Cervical groove prominent and extending to about dorsal carapace; either postorbital 

or postantennal spine present (Fig. 3) . . . . . . . . . . . . . . . . . . . . Solenoceridae (p. 875) 

3b. Distinct part of cervical groove far from dorsal carapace; postorbital and postantennal 

spine absent (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Penaeidae (p. 889) 

postorbital 

spine 

(after Hayashi, 1992) 

cervical groove long 

Fig. 3 Solenoceridae 

1 

2 3 

no 

pincer 

4 

5 

females carry 


Caridea 

cervical groove short 

Fig. 4 Penaeidae 

2 nd pleuron 

expanded, 

pear-shaped


Aristeidae ARISTEIDAE 

Aristeid shrimps 

Diagnostic characters: Animals either a) with rostrum very long in females and young males, but 

becoming rather short in adult males, and always bearing more than 2 upper teeth (subfamily 

Aristeinae); orb)rostrum short, not extending beyond eyes and armed with 1 or 2 upper teeth 

(subfamily Benthesicyminae). No styliform projection at base of eyestalk, but a tubercle present on its 

inner border (very small in Aristaeomorpha). In the subfamily Aristeinae, upper antennular flagellum 

very short and attached to the base of distal antennular segment. Carapace lacks both postorbital 

and postantennal spines; cervical groove either long, extending almost to dorsal carapace, or very short. 

All 5 pairs of legs well developed, fourth leg bearing 2 well-developed arthrobranchs (hidden beneath 

carapace). In males, endopod of second pair of pleopods (abdominal appendages) with appendix 

masculina and appendix interna, but without lateral projection. Third and fourth pleopods divided into 

2 branches. Telson with 1 to 4 pairs of movable lateral spines. Colour: typical coloration of deep-sea 

crustaceans: body reddish or scarlet, sometimes pale white and with red cross bands on abdomen. 

rostrum short 

upper 

antennular 

flagellum 

short 

rostrum of male (Aristeinae) 

Subfamily Benthesicyminae 

rostrum long 

subfamily Aristeinae 

female 

Habitat, biology, and fisheries: All representatives of this family are marine and occur in very deep waters 

(generally deeper than 300 m), with the members of the subfamily Benthesicyminae being exclusively 

bathypelagic (to depths of at least 5 413 m), whereas those of the Aristeinae are benthic and prefer soft 

bottom. Aristeid shrimps are generally of large size and can reach a body length of 33 cm. The sexes are 

easily distinguished by the presence of a large copulatory organ (petasma) on the first pair of pleopods 

(abdominal appendages) of males, while the females have the posterior thoracic sternites modified into a 

large sperm receptacle process (thelycum) which holds the spermatophores or sperm sacs (usually whitish 

or yellowish in colour) after mating. The shape of the petasma and thelycum is often specific and very useful 

for species identification. The eggs are small and numerous, and are released directly into the water and 

not retained on the female abdomen. The larvae are planktonic and have the nauplius stage. At present, 

11 genera and 29 species of aristeid shrimps are known from the Western Central Pacific, but none of them 

are fished commercially because there is virtually no deep-sea fishery in the area. Nevertheless, the fact 

that some species reach a large size and are commonly taken on the basis of exploratory deep-water 

trawling, suggests they may have future commercial potential once that suitable deep-sea fishing gear is 

used in the area. In view of the present non-commercial status of the whole family in the area, no 

identification key to all species is provided here. Instead, a simplified key and species accounts are given 

for 3 species that have high potential interest.

Aristeidae 869 


Penaeidae: rostrum always 

armed with more than 3 upper 

teeth; both upper and lower 

antennular flagella of similar 

length, attached to tip of 

antennular peduncle; eyestalk 

without tubercle on inner 

border; in males, endopod of 

second pair of pleopods with 

appendix masculina only; a 

single well-developed arthrobranch 

on fourth leg (hidden 

beneath carapace). 

Penaeidae 

tubercle 

Aristeidae Penaeidae 

Sicyoniidae: shell generally hard and body “stony” in appearance; abdomen often with deep grooves and 

numerous tubercles; rostrum always armed with more than 3 upper teeth; both upper and lower antennular 

flagella of similar length, attached to tip of antennular peduncle; third and fourth pleopods single-branched. 

appendix 

appendix masculina interna 

Sicyoniidae 

grooves 



appendix 

interna 

Solenoceridae: either postorbital or postantennal spine present on carapace; rostrum always armed with 

more than 3 upper teeth; both upper and lower antennular flagella long, of similar length and attached to 

tip of antennular peduncle; telson usually armed with fixed lateral spines; in males, endopod of second pair 

of pleopods with appendix masculina, appendix interna, and lateral projection. 

Sergestidae: generally small sized; rostrum very short; body strongly compressed laterally, shell soft; last 

2 pairs of legs reduced or absent. 


upper antennular 

flagellum long 

Solenoceridae 


eyes 

lateral 

projection 

Aristeidae Penaeidae Solenoceridae 


various types of appendices masculinae 

on endopods of 2 nd pair of pleopods 

2 

3 

1 

Sergestidae


Stenopodidae: third pincer very large and massive; males and females without large copulatory organ on 



Shrimps of the infraorder Caridea: third leg without pincer; second abdominal pleuron (lateral plate) greatly 

expanded, overlapping posterior part of first pleuron as well as anterior part of third pleuron; males and 

females without large copulatory organ on first pair of pleopods or posterior thoracic sternites, respectively; 

females carry the eggs on the abdomen until hatching. 

2 nd pleuron 

expanded, 

pear-shaped 

1 

2 

Stenopodidae 

4 

5 


females 

carry 

eggs on 

abdomen 

3 

no 

pincer 

females carry 

eggs on 

abdomen 

Key to species with commercial potential in the area 

1a. Rostrum armed with more than 5 upper teeth; hepatic spine present (Fig. 1) (Aristaeomorpha) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Aristaeomorpha foliacea 

1b. Rostrum armed with 3 upper teeth only; hepatic spine absent (Fig. 2) . . . . . . . . . . . . . . → 2 

2a. Crests on carapace without sharp edges (Aristeus) . . . . . . . . . . . . . . . . . .Aristeus virilis 

2b. Crests on carapace very prominent and sharply edged (Plesiopenaeus) . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plesiopenaeus edwardsianus 

hepatic spine 


Fig. 1 Aristaeomorpha 

1 

2 

3 

4 

5 

Caridea 


Fig. 2 Aristeus




Aristaeomorpha foliacea (Risso, 1827) 

Aristeus mabahissae Ramadan, 1938 

Aristeus semidentatus Bate, 1881 

Aristeus virilis (Bate, 1881) 

Benthonectes filipes Smith, 1885 

Betheogennema pasithea (De Man, 1907) 

Bethesicymus altus Bate, 1881 

Bethesicymus bartletti Smith, 1882 

Bethesicymus investigatoris Alcock and Anderson, 1889 

Bethesicymus iridescens Bate, 1881 

Bethesicymus tirmiziae Crosnier, 1978 

Bethesicymus urinator Burkenroad, 1936 

Gennadas bouvieri Kemp, 1909 

Gennadas capensis Calman, 1925 

Gennadas gilchristi Calman, 1925 

Gennadas incertus (Balss, 1927) 

Gennadas kempi Stebbing, 1914 

Gennadas propinquus Rathbun, 1906 

Gennadas scutatus Bouvier, 1906 

Hemipenaeus carpenteri Wood-Mason, 1891 

Hemipenaeus spinidoralis Bate, 1881 

Hepomadus tener Smith, 1884 

Parahepomadus vaubani Crosnier, 1978 

Plesiopenaeus armatus (Bate, 1881) 

Plesiopenaeus edwardsianus (Johnson, 1867) 

Pseudaristeus crassipes (Wood-Mason, 1891) 

Pseudaristeus gracilis (Bate, 1888) 

Pseudaristeus kathleenae Pérez Farfante, 1987 

Pseudaristeus sibogae (De Man, 1911) 

References 

Crosnier, A. 1978. Crustacés Décapodes Péneides Aristaeidae (Benthesicyminae, Aristeinae, Solenocerinae). Faune 

de Madagascar, 46:1-197. 

Crosnier, A. 1989. Benthesicymidae, Aristeidae, Solenoceridae (Crustacea Penaeoidea). In Résultats des Campagnes 

MUSORSTOM, Vol. 5, edited by J. Forest. Mém. Mus. natn. Hist. nat., (A), 144:37-67.


Aristaeomorpha foliacea (Risso, 1827) 

Frequent synonyms / misidentifications: Aristaeomorpha rostridentata (Risso, 1827) / None. 

FAO names: En - Giant red shrimp; Fr - Gambon rouge; Sp - Gamba española. 

rostrum of male 

female 

Diagnostic characters: A large shrimp. Rostrum with 6 to 12 upper teeth (including 2 teeth on 

carapace); very long in females and extending far beyond antennal scale, but short in males and not 

exceeding tip of antennular peduncle. Carapace with antennal, hepatic, and branchiostegal spines. 

Upper antennal flagella very short. Third to sixth abdominal segments each bearing a strong 

posteromedian spine. Telson with 4 pairs of small movable lateral spines. Colour: body uniformly 

vermilion; eyes black. 

Size: Maximum body length 22.5 cm in females (carapace length 5.9 cm) and 17 cm in males (carapace 

length 4.5 cm); commonly between 12 and 16 cm. 

Habitat, biology, and fisheries: Found from depths of 61 to 1 300 m, but more often between 300 and 

750 m; prefers mud bottoms. Moves to midwater at night. One of the common larger shrimps caught during 

deep-water exploratory trawling operations, often encountered in large quantities. Not yet fished 

commercially in the area, but with high potential for deep-sea fisheries. 

Distribution: Cosmopolitan, 

reported to be widely 

distributed in the Western 

Atlantic, Mediterranean, and 

Indo-West Pacific.


Aristeus virilis (Bate, 1881) 


FAO names: En - Stout red shrimp; Fr - Gambon gaillard; Sp - Gambón colorado. 

rostrum of female 

(after Lee and Yu, 1977) 

Diagnostic characters: A large shrimp; body covered with pubescence. Rostrum armed with 3 upper 

teeth only (including 1 tooth on carapace); very long in females and extending far beyond antennal scale, 

but short in males and not exceeding tip of antennal scale. Carapace with antennal and branchiostegal 

spines but lacking hepatic spine. Crests on carapace without sharp edges. Upper antennal flagella 

very short. Legs with photophores; first to third legs armed with a movable spine on merus. Fourth 

to sixth abdominal segments each bearing a strong posteromedian spine. Telson with 4 pairs of small 

movable lateral spines. Colour: body pale white, with red bands on posterior margin of abdominal 

segments; eyes black; tip of rostrum, antennal scale, distal half of uropods, antennular and antennal flagella 

reddish; upper and lateral carapace, as well as legs and pleopods somewhat reddish; photophores on legs 

purple-red; some young individuals with body rather uniformly reddish. 

Size: Maximum body length about 22.2 cm in females (carapace length 6.1 cm) and 14.6 cm in males 

(carapace length 4.6 cm); commonly between 9 and 12 cm. 

Habitat, biology, and fisheries: Found on sand and mud bottom, at depths from 188 to 936 m, usually 

between 350 and 700 m, apparently not migrating into midwater at night. Not yet fished commercially in 

the area. However, the size of this species and the fact that it is commonly taken during experimental 

trawling operations in the Philippines, Indonesia, and New Caledonia suggest it has high potential with the 

development of a deep-sea fishery in these countries. 

Distribution: Indo-West Pacific 

from eastern coast of Africa to 

India, western Australia, the 

Philippines, Japan, Indonesia, 

New Caledonia, and Vanuatu. 

male


Plesiopenaeus edwardsianus (Johnson, 1867) 

Frequent synonyms / misidentifications: Aristaeopsis edwardsiana (Johnson, 1867) / None. 

FAO names: En - Scarlet shrimp; Fr - Gambon écarlat; Sp - Gamba carabinero. 

Diagnostic characters: Size very large. Rostrum armed with 3 upper teeth only (including 1 tooth on 

carapace); very long in females, reaching far beyond antennal scale, but short in males and not exceeding 

tip of antennal scale. Carapace with antennal and branchiostegal spines but lacking hepatic spine. Crests 

on carapace very sharp and prominent. Upper antennal flagella very short. Exopod of second 

maxilliped about 2 times as large as endopod. Legs without exopods. Third to sixth abdominal 

segments each bearing a sharp posteromedian spine. Telson with 4 pairs of small movable lateral 

spines. Colour: uniformly scarlet; eyes black. 

Size: Maximum body length 33.4 cm in females (carapace length 10.4 cm) and 19.3 cm in males (carapace 

length 6.3 cm); commonly between 15 and 19.5 cm. 

Habitat, biology, and fisheries: Found on sandy or muddy bottom, at depths of 200 to 1 850 m, usually 

between 400 and 900 m. A very large species, often trawled at depths of more than 500 m. With very high 

potential for deep-sea fisheries in the area, although so far only encountered during exploratory trawling 

operations off the Philippines, Willis and Fortuna islands. 

Distribution: Cosmopolitan, 

widely distributed in the 

Atlantic and the Indo-West 

Pacific.

Solenoceridae 875 

Solenoceridae SOLENOCERIDAE 

Solenocerid shrimps 

Diagnostic characters: Rostrum well developed (extending at least to middle of eye), always bearing 

more than 3 upper teeth (including those on carapace); no styliform projection at base of eyestalk, 

but a tubercle present on its inner border. Both upper and lower antennular flagella long (additionally 

strongly compressed laterally and tube-like in Solenocera), of the same length, and attached to tip 

of antennular peduncle. Carapace either with postorbital or postantennal spine; cervical groove 

long, extending to about dorsal carapace. All 5 pairs of legs well developed, fourth leg bearing 2 

well-developed arthrobranchs (hidden beneath carapace). In males, endopod of second pair of 

pleopods (abdominal appendages) with appendix masculina, appendix interna, and lateral projection. 

Third and fourth pleopods divided into 2 branches. Telson generally armed with at least 1 pair of fixed 

lateral spines. Colour: generally pink to red; sometimes with pale markings on antennular flagella and 

tips of uropods. 


cervical groove long 

antennular 

flagella long 

Habitat, biology, and fisheries: Found in deeper 

marine waters offshore from depths of 2 to over 5 

700 m (usually deeper than 20 m). Generally benthic 

animals with preference for soft bottoms. Species of 

the genus Solenocera often burrow in mud during the 

daytime, with only the tube-like antennular flagella 

sticking out for respiration. Their size range from 2.5 to 

21.5 cm body length but most species are of moderate 

size. The sexes are easily distinguished by the 

presence of a large copulatory organ (petasma) on the 

first pair of pleopods (abdominal appendages) of 

males, while the females have the posterior thoracic 

sternites modified into a large sperm receptacle 

process (thelycum) which holds the spermatophores 

or sperm sacs (usually whitish or yellowish in colour) 

cross-section 

respiratory 

tube 


upper 

antennular 

flagellum 

lower 

antennular 

flagellum 

(genus Solenocera) 

after mating. The shape of the petasma and thelycum is often specific and very useful for species 

identification. The eggs are small and numerous, and are released directly into the water and not retained 

on the female abdomen. The larvae are planktonic and have the nauplius stage. In the Western Central 

Pacific, about 8 genera and 36 species of solenocerid shrimps have been recorded.Since they are generally 

found in deeper waters, at present only a few of them are taken as bycatch in commercial trawl fisheries. 

Nevertheless, results of many exploratory deep-water trawling operations have shown that several species 

are abundant and have fishery potential. Nevertheless, no key to all species of Solenoceridae occurring in 

the area is given here as most of them are not yet commercially caught and the taxonomic status of some 

species is still unclear. Species accounts and a key are included below for 2 genera and 11 species which 

are either more commonly found in commercial catches or that can be easily confused with those 

commercial species.



Aristeidae: either rostrum very short and armed with 1 or 2 upper teeth only, or upper antennular flagellum 

very short and not attached to tip of antennular peduncle; no postorbital or postantennal spine present on 

carapace; telson armed only with movable lateral spines; in males, endopod of second pair of pleopods 

with appendix masculina, appendix interna, but without lateral projection. 

Penaeidae: no postorbital or postantennal spine present on carapace; cervical groove short, with distinct 

part always far from dorsal carapace; eyestalk without tubercle on inner border; in males, endopod of 

second pair of pleopods with appendix masculina only; a single well-developed arthrobranch on fourth leg 

(hidden beneath carapace). 

tubercle 

Solenoceridae Penaeidae 

upper 

antennular 

flagella 

very short 

(Aristeinae) 

eyes 

Aristeidae 

appendix 

interna 

appendix masculina 

appendix 

interna 

Aristeidae Penaeidae 

lateral 

projection 

Solenoceridae 

endopods of 2 nd pleopod in males 

Penaeidae 

cervical 

groove 

Sicyoniidae: shell generally hard and body “stony” in appearance; abdomen often with deep grooves and 

numerous tubercles; no postorbital or postantennal spine present on carapace, cervical groove generally 

indistinct or absent; third and fourth pleopods single-branched. 

Sergestidae: generally small sized; rostrum very short; body strongly compressed laterally, shell soft; last 

2 pairs of legs reduced or absent. 


Sicyoniidae 

grooves 



2 

3 


1 

2 

3 

1 

4 5 

Penaeidae 

Sergestidae






expanded, overlapping posterior part of first pleuron and anterior part of third pleuron; males and females 

without large copulatory organ on first pair of pleopods or posterior thoracic sternites, respectively; females 

carry the eggs on the abdomen until hatching. 

1 

2 

Stenopodidae 


females 

carry 

eggs on 

abdomen 


1a. Antennular flagella subcylindrical and 

thread-like (Fig. 1a); rostrum strongly convex 

(Fig. 2a); exopod of uropod armed with a 

distolateral spine (Haliporoides) . . Haliporoides sibogae 

1b. Antennular flagella flattened and tube-like 

(Fig. 1b); rostrum nearly horizontal (Fig. 2b); 

exopod of uropod without distolateral spine 

. . . . . . . . . . . . . . . . . . . . . (Solenocera ) → 2 

rostrum convex 

2a. Telson without lateral spine (Fig. 3a) . . . . . 

. . . . . . . . . . . . . . . . . . Solenocera crassicornis 

2b. Telson armed with lateral spines (Fig. 3b) . . . . . . → 3 

3a. Postrostral crest elevated . . . . . . . . . . . . . . → 4 

3b. Postrostral crest weak and low . . . . . . . . . . . → 10 

4 

a) Haliporoides 

5 

3 

1 

rostrum horizontal 

Fig. 2 carapace (lateral view) 

2 3 

no pincer 

5 

4 

females carry 

eggs on 

abdomen 

Caridea 

dorsal 

flagellum 

ventral 

flagellum 

2 nd pleuron 

expanded, 

pear-shaped 

a) Haliporoides b) Solenocera 

Fig. 1 cross-section of antennular flagella 

b) Solenocera 

a) Solenocera 

crassicornis 


b) Solenocera 

melantho


4a. Postrostral crest very high and plate-like (Figs 4 and 5) . . . . . . . . . . . . . . . . . . . . . → 5 

4b. Postrostral crest distinct but not plate-like (Figs 6 and 7) . . . . . . . . . . . . . . . . . . . . . → 6 

5a. Rostrum extending to 2/3 of eye; postrostral crest behind cervical notch with anterior 

part distinctly higher than posterior part (Fig. 4) . . . . . . . . . . . . . . . . . Solenocera choprai 

5b. Rostrum extending to 1/2 of eye; postrostral crest behind cervical notch with posterior 

part distinctly higher than anterior part (Fig. 5) . . . . . . . . . . . . . . . . Solenocera alticarinata 

postrostral ridge plate-like 

plate-like 

Fig. 4 Solenocera choprai 

Fig. 5 Solenocera alticarinata 

(from Crosnier, 1978) 


6a. Postrostral crest well separated from postrostral teeth by a distinct notch above cervical 

groove (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenocera koelbeli 

6b. No distinct notch present between postrostral teeth and postrostral crest (Fig. 7) . . . . . . . . 

notch 

→ 7 

Fig. 6 Solenocera koelbeli 


7a. Posterior part of hepatic groove and anterior part 

of brachiocardiac groove both very distinct and 

strongly curving downward; postrostral crest behind 

cervical groove sometimes with an upper 

tooth; median part of first abdominal segment 

very narrow and dorsal crest of second abdomi- 

dorsal crest 

distinct 

nal segment distinct (Fig. 8a) . . . . . . . Solenocera alfonso 

7b. Never both posterior part of hepatic groove and 

anterior part of brachiocardiac groove distinct 

and curving downward together; postrostral crest 

behind cervical groove without any teeth; median 

part of first abdominal segment moderately wide 

and dorsal crest of second abdominal segment 

indistinct (Fig. 8b) . . . . . . . . . . . . . . . . . . . . . → 8 

Fig. 7 Solenocera australiana 

(after Pérez Farfante and Grey, 1980) 

a) Solenocera alfonso b) Solenocera melantho 

Fig. 8 abdomen (dorsal view) 

8a. Rostrum with lower border nearly convex (Fig. 9a); male petasma with distal margin 

armed with many well-defined long spinules (Fig. 10a); female thelycum with posterior 

thoracic ridge almost straight (Fig. 11a) . . . . . . . . . . . . . . . . . . . Solenocera australiana 

8b. Rostrum with lower border razor-shaped, very straight or slightly concave (Figs 9b, c); 

male petasma with spinules on distal margin short and not very well defined; female 

thelycum with posterior thoracic ridge strongly bilobed (Fig. 11b, c) . . . . . . . . . . . . . . . → 9 

I 

II 

III 


a) Solenocera australiana b) Solenocera melantho (from Crosnier, 1989) c) Solenocera halli 

Fig. 9 rostrum (lateral view)


9a. Anterior end of hepatic crest very strongly 

convex (Fig. 12a); male petasma with dorsolateral 

lobule bearing 0 to 13 terminal 

spinules (Fig. 10a); female thelycum bearing 

2 or 3 pairs of protuberances in the middle, 

with submedian pair larger than lateral ones 

(Fig. 11b) . . . . . . . . . . . . . . Solenocera melantho 

9b. Anterior end of hepatic crest only slightly 

convex or nearly straight (Fig. 12b); male 

petasma with dorsolateral lobule bearing 18 

to 40 terminal spinules (Fig. 10c); female 

thelycum always bearing 2 pairs of protuberances 

in the middle, with submedian pair 

smaller than lateral pair (Fig. 11c) . . . Solenocera halli 

almost straight 

a) Solenocera 

australiana 

submedian 

protuberances 

larger 

bilobed 

b) Solenocera 

melantho 

Fig. 11 thelycum 


submedian 

protuberancess 

smaller 

dorsolateral 

lobe 

a) Solenocera 

australiana 

bilobed 

c) Solenocera 

halli 

dorsolateral 

lobe 

b) Solenocera 

melantho 

dorsolateral lobe 

c) Solenocera 

halli 

Fig. 10 petasma 


a) Solenocera melantho 

b) Solenocera halli 

Fig. 12 anterolateral corner of 

carapace 


10a. Rostrum with 6 or 7 large and well-separated upper teeth (Fig. 13); antennular flagella 

0.8 to 1.2 times as long as carapace and generally composed of less than 60 articles 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenocera pectinulata 

10b. Rostrum generally with 8 or 9 densely packed small upper teeth (Fig. 14); antennular 

flagella 1.3 to 1.9 times as long as carapace and composed of more than 60 articles 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Solenocera pectinata 

6-7 large teeth 

Fig. 13 Solenocera pectinulata 


8-9 small teeth 

Fig. 14 Solenocera pectinata 


hepatic 

crest 

hepatic 

crest




Cryptopenaeus clevai Crosnier, 1985 

Cryptopenaeus crosnieri Pérez Farfante and Kensley, 1985 

Gordonella kensleyi Crosnier, 1988 

Gordonella paravillosa Crosnier, 1988 

Hadropenaeus lucasii (Bate, 1881) 

Haliporoides cristatus Kensley, Tranter, and Griffin, 1987 

Haliporoides sibogae (De Man, 1907) 

Haliporus curvirostris Bate, 1881 

Haliporus taprobanensis Alcock and Anderson, 1899 

Hymenopenaeus equalis (Bate, 1881) 

Hymenopenaeus halli Bruce, 1966 

Hymenopenaeus laevis (Bate, 1881) 

Hymenopenaeus neptunus (Bate, 1881) 

Hymenopenaeus propinquus (De Man, 1907) 

Mesopenaeus brucei Crosnier, 1986 

Mesopenaeus mariae Pérez Farfante and Ivanov, 1982 

Solenocera alfonso Pérez Farfante, 1981 

Solenocera alticarinata Kubo, 1949 

Solenocera annectens (Wood-Mason, 1891) 

Solenocera australiana Pérez Farfante and Grey, 1980 

Solenocera barunajaya Crosnier, 1994 

Solenocera bedokensis Hall, 1962 

Solenocera choprai Nataraj, 1945 

Solenocera comata Stebbing, 1915 

Solenocera crassicornis (H. Milne Edwards, 1837) 

Solenocera faxoni De Man, 1907 

Solenocera halli Starobogatov, 1972 

Solenocera koelbeli De Man, 1911 

Solenocera melantho De Man, 1907 

Solenocera moosai Crosnier, 1985 

Solenocera pectinata (Bate, 1888) 

Solenocera pectinulata Kubo, 1949 

? Solenocera phuongi Starobogatov, 1972 

Solenocera rathbunae Ramadan, 1938 

Solenocera spinajugo Hall, 1961 

Solenocera waltairensis George and Muthu, 1970 

References 

Crosnier, A. 1978. Crustacés Décapodes Pénéides Aristaeidae (Benthesicyminae, Aristeinae, Solenocerinae). Faune 

de Madagascar, 46:1-197. 

Crosnier, A. 1985. Penaeoid shrimps (Benthesicymidae, Aristeidae, Solenoceridae, Sicyoniidae) collected in Indonesia 

during the Corindon II and IV expeditions. Mar. Res. Indonesia, 24:19-47. 

Crosnier, A. 1989. Benthesicymidae, Aristeidae, Solenoceridae (Crustacea Penaeoidea). In Résultats des Campagnes 

MUSORSTOM, Vol. 5, edited by J. Forest. Mém. Mus. natn. Hist. nat., (A), 144:37-67. 

Crosnier, A. 1994. Crustacea Decapoda: Penaeoidea récoltés lors de la campagne KARUBAR en Indonésie.In Résultats 

des Campagnes MUSORSTOM, Vol. 12, edited by A. Crosnier. Mém. Mus. natn. Hist. nat., (A), 161: 351-365.


Haliporoides sibogae (De Man, 1907) JAQ 

Frequent synonyms / misidentifications: Haliporoides sibogae australiensis Kensley, Tranter, and Griffin, 

1987; H. sibogae madagascariensis Crosnier, 1987; Hymenopenaeus sibogae (De Man, 1907) / None. 

FAO names: En - Jack-knife shrimp; Fr - Salicoque canif; Sp - Camarón cortapluma. 


Diagnostic characters: Body somewhat hairy. Both upper and lower antennular flagella very long, 

thread-like. Rostrum strongly convex and short, extending just beyond eyes; upper border armed 

with 4 to 7 teeth; lower border concave, armed with 1 to 3 teeth. Carapace with 1 postrostral and 1 

epigastric tooth, as well as antennal, postantennal, branchiostegal, hepatic, and suprahepatic 

spines. Cervical groove distinct, extending to dorsal carapace. Legs progressively longer posteriorly. 

Telson with a pair of fixed lateral spines. Colour: body orange to pink; antennal flagella whitish; eyes dark 

brown; uropods reddish with white tips. 

Size: Maximum body length 16.5 cm in males (carapace length 3.3 cm) and 20 cm in females (carapace 

length 4.9 cm), commonly between 7 and 10 cm. 

Habitat, biology, and fisheries: Deep sea from depths of 100 to 1 463 m, usually between 300 and 600 m, 

on soft bottom. Taken in large quantities on the basis of experimental deep-water trawling in the Philippines 

and Indonesia. Generally considered as a species with high potential for commercial deep-sea fisheries. 

In the area, so far only fished on a small scale off the northeastern coast of Australia (about 50 t in the 

annual period of 1989/1990). Marketed mainly frozen, consumed locally, sometimes exported. 

Distribution: Widely distributed 

in the Indo-West Pacific 

from Madagascar to Japan, 

Australia, and New Zealand 

(populations from Madagascar 

and eastern Australia are 

sometimes considered as 2 

different subspecies).


Solenocera choprai Nataraj, 1945 

Frequent synonyms / misidentifications: None / Solenocera alticarinata Kubo, 1949. 

FAO names: En - Ridgeback shrimp; Fr - Salicoque balafrée; Sp - Camarón costurón. 

(after Hall, 1962) 

Diagnostic characters: Rostrum short, extending to about 2/3 of eyes; upper border with 8 to 10 teeth 

(including 4 teeth on carapace); lower border moderately convex. Postrostral crest markedly elevated 

and plate-like, slightly interrupted by a small notch above cervical groove; height of posterior part 

progressively decreasing posteriorly. Carapace with orbital, postorbital, antennal and hepatic spines, 

but without pterygostomian spines. Antennular flagella moderately long and tube-like. Telson with a pair 

of lateral spines. Colour: body and legs reddish; antennae banded with dark red and white; uropods 

dark red, except for some white areas. 

Size: Maximum body length 13 cm (females) and 9.5 cm (males). 

Habitat, biology, and fisheries: Found on soft bottoms at depths between 50 and 175 m. Probably burrows 

in mud during the daytime, with only the tube-like antennular flagella sticking out for respiration. A bycatch 

of trawling operations in slightly deeper waters. Nowhere abundant, but of some commercial value because 

of its moderately large size. In the Philippines, this shrimp is occasionally marked together with other 

Solenocera species. Marketed mainly fresh for local consumption. Often confused with Solenocera 

alticarinata, which has a more northeastern distribution (from Japan to the Philippines), but they are both 

sometimes considered as a single species. 



from the eastern coast of Africa 

to the Philippines and Australia.


Solenocera crassicornis (H. Milne Edwards, 1837) 

Frequent synonyms / misidentifications: Solenocera indicus Nataraj, 1945; S. kuboi Hall, 1956; S. sinensis 

Yu, 1937; S. subnuda Kubo, 1949 / None. 

FAO names: En - Coastal mud shrimp; Fr - Salicoque des vases côtières; Sp - Camarón fanguero de orilla. 

Diagnostic characters: Rostrum short and nearly straight, about as long as eyes; upper border 

armed with 4 to 7 (mostly 5) teeth; lower border unarmed and somewhat convex. Carapace with 3 

postrostral teeth and 1 epigastric tooth, and orbital, postorbital, antennal and hepatic spines, but without 

pterygostomian spines; postrostral crest low and rounded. Antennular flagella moderately long and 

tube-like. Telson unarmed, without lateral spines. Colour: body pink to pinkish orange; posterior border 

of each abdominal segment covered with a red cross band; eyes dark brown; antennular flagella and 

distal part of tail fan reddish. 

Size: Maximum body length 9 cm (males) and 14 cm (females), commonly between 6 and 8 cm. 

Habitat, biology, and fisheries: Inhabits muddy bottoms close to shore, at depths from 20 to 85 m. 

Probably burrows in mud during the daytime, with only the tube-like antennular flagella sticking out for 

respiration. Mainly forms a bycatch of trawlers. In the area, this shrimp seems to be more common around 

Thailand and is of minor commercial importance. 



from the Persian Gulf to 

Japan and Indonesia.


Solenocera melantho De Man, 1907 

Frequent synonyms / misidentifications: Solenocera prominentis Kubo, 1949 / Solenocera alfonso Pérez 

Farfante, 1981; S. australiana Pérez Farfante and Grey, 1980; S. halli Starobogatov, 1972; S. koelbeli De Man, 

1911. 

FAO names: En - Razor mud shrimp. 


Diagnostic characters: Rostrum short, not extending beyond eyes; upper border with 6 to 10 teeth 

(including 3 teeth on carapace); lower border straight or slightly concave, razor-shaped. Postrostral 

crest distinct but not very high and plate-like, without a distinct notch above cervical groove. 

Carapace with orbital, postorbital, antennal and hepatic spines, but without pterygostomian spines. 

Anterior end of hepatic crest strongly concave and curving upward. Antennular flagella moderately 

long and tube-like. Telson with a pair of lateral spines. Colour: body pink to red and somewhat 

semi-transparent; some irregular red markings on abdomen; eyes black-brown; antennular flagella 

reddish with a white (or pale yellowish) band at midlength; distal part of uropods slightly yellowish. 

Size: Maximum body length 15 cm, commonly between 7 and 12 cm. 

Habitat, biology, and fisheries: Inhabits the upper slopes of continental shelves at depths from 78 to 

400 m, on sandy mud bottom. Mainly forms a bycatch of trawling operations in slightly deeper waters. 

Probably the most common species of the genus in the Philippines, but still not very abundant and only 

occasionally sold in local fish markets. Marketed mainly fresh for local consumption. This species, 

Solenocera alfonso, S. australiana, andS. halli are probably often confused with each other. 


and definitely known from 

Japan, Korea, Taiwan 

Province of China, coasts of 

China, the Philippines, and 

Indonesia.


Solenocera alfonso Pérez Farfante, 1981 

En - Deep-water mud shrimp. 

Maximum body length about 12 cm (at a maximum carapace length of 4 cm). Inhabits the upper 

slopes of island shelves at depths from 176 to 547 m, on bottoms of green mud or fine sandy mud. 

So far only taken on the basis of experimental deep-water trawling. However, the size of this species 

and the fact that it is sometimes found in large quantities suggest it may have commercial potential 

with the development of a deep-sea fishery in the area. Indo-West Pacific and so far only known 

with certainty from the Philippines, Indonesia, and Northwestern Australia. 

Solenocera alticarinata Kubo, 1949 

(after Pèrez Farfante, 1981) 

En - High ridge mud shrimp. 

Maximum body length 11 cm (females) and 9 cm (males), commonly between 7 and 9 cm. On sandy 

mud bottom, at depths from 50 to 180 m. Taken by trawls. Apparently restricted to the Western Pacific 

from Japan to Taiwan Province of China, the South China Sea, and the Philippines. Often confused 

with the closely related Solenocera choprai. In the area, so far only recorded from the Philippines 

where it is probably less common than S. choprai. 

(after Grey, Dall, and Baker, 1983)


Solenocera australiana Pérez Farfante and Grey, 1980 

En - Australia mud shrimp. 

Maximum body length 12 cm (females) and 9 cm (males). In shallow water at depths from 15 to 

40 m, over mud bottom with or without coral debris, rock, shell, or vegetation. Taken by commercial 

trawlers fishing for large penaeids in northern Australia, but so far without significant economic 

importance. Restricted to northern Australia. 

Solenocera halli Starobogatov, 1972 

(after Pèrez Farfante and Grey, 1980) 

En - Malayan mud shrimp. 

Maximum body length about 10 cm (at a maximum carapace length of 2.75 cm). Found in shallow 

waters at depths from 48 and 75 m. So far mainly caught by experimental trawlers but probably more 

common around Malaysia and Singapore. Indo-West Pacific; thus far known from Bay of Bengal, 

Strait of Malacca, Singapore, and the South China Sea. Often confused with Solenocera melantho. 

(after Hall, 1962)


Solenocera koelbeli De Man, 1911 

En - Chinese mud shrimp; Fr - Salicoque chinoise de vase; Sp - Camarón fanguero chino. 

Maximum body length 15 cm, commonly between 5 and 10 cm. On soft bottom at depths from 21 

to 241 m, usually between 60 and 90 m. A bycatch in trawl fisheries. Western Pacific from Japan to 

Taiwan Province of China, the South China Sea, Viet Nam, the Philippines, and Indonesia. 

Apparently not abundant in the area. Sometimes confused with Solenocera alticarinata, S. choprai, 

or S. melantho. 

Solenocera pectinata (Bate, 1888) 


En - Comb shrimp; Fr - Salicoque peigne; Sp - Camarón peine. 

Maximum body length 6 cm. On soft bottom at depths from 4 to 205 m. A common bycatch of 

commercial trawlers but of no economic importance, due to its small size. Widely distributed in the 

Indo-West Pacific, from the eastern coast of Africa to Japan and Wallis Island in the South Pacific. 

Often confused with Solenocera pectinulata but generally found at shallower depths. 

(after Hall, 1962)


Solenocera pectinulata Kubo, 1949 

En - False comb shrimp. 

Maximum body length 6.5 cm (carapace length to 2.2 cm), commonly between 3 and 5.5 cm. On 

bottoms of sand and/or mud, at depths from 75 to 350 m, usually deeper than 175 m. A common 

bycatch of commercial trawlers but without economic importance, due to its small size. Widely 

distributed in the Indo-West Pacific from the eastern coast of Africa to Japan and Indonesia. Often 

confused with Solenocera pectinata but generally has a deeper distribution. 

(after Motoh and Buri, 1984) 



Penaeidae 889 

Penaeidae PENAEIDAE 

Penaeid shrimps 

D 

cervical groove short 

iagnostic characters: Rostrum well developed 

and generally extending beyond 

eyes, always bearing more than 3 upper 

teeth. No styliform projection at base of eyestalk 

and no tubercle on its inner border. 

Both upper and lower antennular flagella of 

similar length, attached to tip of antennular 

peduncle. Carapace lacking both postorbital 

or postantennal spines. Cervical groove 

generally short, always with a distance from 

dorsal carapace. All 5 pairs of legs well developed, 

fourth leg bearing a single well-developed 

arthrobranch (hidden beneath 

carapace, occasionally accompanied by a 

second, rudimentary arthrobranch). In 

males, endopod of second pair of pleopods 

(abdominal appendages) with appendix masculina 

only. Third and fourth pleopods divided into 2 branches. Telson sharply pointed, with or without fixed 

and/or movable lateral spines. Colour: body colour varies from semi-translucent to dark greyish green or 

reddish, often with distinct spots, cross bands and/or other markings on the abdomen and uropods; 

live or fresh specimens, particularly those of the genus Penaeus, can often be easily distinguished 

by their coloration. 

Habitat, biology, and fisheries: Members of this family are usually marine, although juveniles and young are 

often found in brackish water or estuaries, sometimes with very low salinities (a few unconfirmed fresh-water 

records exist). Some penaeids, mainly those of the genera Parapenaeus and Penaeopsis, occur in deep water 

at depths of more than 750 m. Penaeids are mostly benthic and mainly found on soft bottom of sand and/or 

mud, but a few species (e.g. genus Funchalia) are pelagic and others are known to inhabit coral reefs (e.g. the 

genera Heteropenaeus, Trachypenaeopsis, also some Metapenaeopsis). Their size ranges from 2.5 to 35 cm 

body length. The sexes are easily distinguished by the presence of a very large copulatory organ (petasma) on 

the first pair of pleopods (abdominal appendages) of males, while the females have the posterior thoracic 

sternites modified into a large sperm receptacle process (thelycum) which holds the spermatophores or sperm 

sacs (usually whitish or yellowish in colour) after mating.The shape of the petasma and thelycum is often specific 

and very useful for species identification. The eggs are small and numerous, and are released directly into the 

water and not retained on the female abdomen. The larvae are planktonic and have the nauplius stage. 

mysis 

postlarvae 

juveniles 

eggs 

protozoa 

nauplius 

life-cycle of shrimps of the genus Penaeus 

adult


The life cycle of species of Penaeus and Metapenaeus, the 2 most important commercial shrimp genera, is complex 

(see figure on previous page). Adults generally move from shallow coastal waters to offshore and spawn at depths 

between 10 to 80 m. The eggs hatch within 14 to 24 hours and release very small, simple larvae, the nauplii. The 

nauplius larva passes through several substages before it metamorphoses into the mysis stage. These larvae are 

planktonic and are carried by currents toward shore where they arrive as postlarvae; this occurs about three weeks 

after hatching when the animals are 6 to 14 mm long and shrimp-like in appearance. The postlarvae invade inshore 

brackish waters, abandon their planktonic way of life, and become bottom dwellers living in shallow littoral areas. In 

these rich nursery grounds they grow rapidly, develop into juveniles and, as size increases, move gradually back 

toward the mouths of bays or estuaries, where they become subadults. Soon the shrimps migrate offshore, continue 

growing and mate, and when they finally reach the spawning grounds, the mature females spawn and the cycle is 

repeated; most shrimps in these grounds are about 1 year old, rarely older than 2 (or perhaps 3) years old. 

At present, 11 genera and 112 species of penaeids are known to occur in the Western Central Pacific. Among 

these, the genus Penaeus is of greatest economical importance. Species of Penaeus are caught extensively by 

trawls, seines, set nets, traps, and artisanal gear, with P. merguiensis and P. monodon probably being the 2 

most important species. Moreover, aquaculture of Penaeus is very popular in many countries, mostly using 

Penaeus monodon. The genus of secondary importance is Metapenaeus, often taken together with Penaeus 

and also extensively cultured in ponds. The third commercially important genus is probably Parapenaeopsis, 

which is quite common in the western part of the area and often forms a significant part of the bycatch in prawn 

fisheries.Therefore, species accounts are provided here for all species of Penaeus,Metapenaeus, and Parapenaeopsis 

that are known from the Western Central Pacific. The other genera seem to be less abundant, although species 

of Metapenaeopsis and Trachypenaeus are frequently found among catches of prawn fisheries and have some 

commercial value. The 2 deep-water genera Parapenaeus and Penaeopsis are not fished at present, although 

experimental deep-water trawling operations have shown that representatives of these 2 genera, particularly 

Parapenaeus, can be caught in large quantities and are potentially of interest to fisheries. The remaining 4 genera 

are all without economic interest: species of Atypopenaeus are generally too small and not abundant, 

Heteropenaeus and Trachypenaeopsis occur in coral reefs and are small, and Funchalia is a pelagic deep-sea genus. 

Since this family is of greatest commercial importance in the area, identification keys are given here for all penaeids, 

except for Metapenaeopsis, 40 species of which occur in the area, but most of these are either not common enough 

or found in deep waters, without any commercial potential. Therefore, an identification key is provided only for those 

species of Metapenaeopsis that may be found in commercial catches. For a full key to Metapenaeopsis, see the 

detailed revisions by A. Crosnier (1987-1994, published in “Bull. Mus. natn. Hist. nat., Paris”, and “Rèsultats des 

Campagnes MUSORSTOM”). 


The shapes of the male petasma and female thelycum are very important taxonomic characters in several 

genera of penaeid shrimps, such as Metapenaeopsis, Metapenaeus, Parapenaeopsis, Parapenaeus, and 

Trachypenaeus. However, as the petasma and thelycum are not fully developed in juveniles, a positive 

identification of juvenile specimens is often difficult. Therefore, it is highly recommended to use, if possible, 

adult specimens rather than juveniles when using the keys. 

For the differentiation between adult and juvenile males, it should be remembered that, in adult males, the 

left and right parts of the petasma are very rigid and strongly fused to each other (i.e.very difficult to separate), 

while in juvenile males the left and right parts of the petasma are either not fused or only weakly united (i.e. 

easy to separate) and somewhat soft with the scultpure not well defined. 

In adult females, the thelycum is clearly sculptured with the ridges and depressions very well marked, while 

in juvenile females the thelycum has only a shallow sculpture, not well defined. 


Aristeidae: either rostrum very short, armed with 1 or 2 upper teeth only, or upper 

antennular flagellum very short, not attached to tip of antennular peduncle; 

eyestalks generally with a tubercle on inner border; fourth leg bears 2 

well-developed arthrobranchs (hidden beneath carapace); in males, endopod of 

second pair of pleopods with appendix masculina, appendix interna, but without 

lateral projection. 

Sicyoniidae: shell generally hard and body “stony” in appearance; exopod present 

on first maxilliped only; abdomen often with deep grooves and numerous tubercles; 

third and fourth pleopods single-branched. 

Solenoceridae: carapace either with postorbital or postantennal spine; cervical 

groove distinct and extending to about dorsal carapace; eyestalks generally with 

a tubercle on inner border; fourth leg bears 2 well-developed arthrobranchs 

(hidden beneath carapace); in males, endopod of second pair of pleopods with 

appendix masculina, appendix interna, and lateral projection. 

postorbital 

spine 

Solenoceridae 

Penaeidae 

cervical 

groove

Penaeidae 891 

Sergestidae: size small; rostrum very short; body strongly compressed laterally; shell soft; last 2 pairs of legs 

reduced or absent. 





expanded, overlapping posterior part of first pleuron and anterior part of third pleuron; males and females without 

large copulatory organ on first pair of pleopods or posterior thoracic sternites, respectively; females carry the eggs 

on the abdomen until hatching. 

appendix masculina appendix 

interna 

appendix 

interna 

lateral 

projection 

Aristeidae Penaeidae Solenoceridae 

endopod of 2 nd tubercle 

tubercle 

pleopod in males 

Solenoceridae Aristeidae 

eyes 

Penaeidae 

Key to the genera of Penaeidae occurring in the area 

1a. Rostrum with lower teeth 

(Fig. 1) . . . . . . . . . . . . . . → 2 

1b. Rostrum without lower 

teeth (Fig. 2) . . . . . . . . . . . → 3 

2a. 

teeth on 

Abdomen with many lower border 

deep pubescent grooves 

(Fig. 3) . . . . . . . . Heteropenaeus 

(a single species, H. longimanus) 

2b. Abdomen glabrous and 

smooth (Fig. 4) . . . . . . . Penaeus 

3a. Telson with a pair of large subapical 

fixed lateral spines (Fig. 5) . . . . . . → 4 

3b. Telson without large subapical 

fixed lateral spines . . . . . . . . . . . . → 7 

rostrum (lateral view) 

Fig. 1 Penaeus 

Fig. 3 Heteropenaeus longimanus 



no lower teeth 

rostrum (lateral view) 

Fig. 2 Metapenaeus 

movable 

spines 

movable 

spines 

fixed 

spine 

fixed 

spine 

fixed 

spine 

a) Penaeopsis b) Metapenaeopsis c) Parapenaeus 


(after Lee and Yu, 1977)


4a. Body densely covered with short hairs, with grooves and crests on carapace obscure 

(Fig. 6a); petasma asymmetrical (Fig. 6b) . . . . . . . . . . . . . . . . . . . . . . Metapenaeopsis 

4b. Body almost naked, with crests and grooves on carapace distinct (Fig. 7a); petasma 

symmetrical (Figs 7b and 8b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

asymmetrical 

longitudinal suture 

a) carapace (lateral view) 

Fig. 6 Metapenaeopsis 

b) petasma 


vertical suture 


Fig. 7 Parapenaeus 

b) petasma 


5b. Carapace with longitudinal and vertical sutures (Fig. 7a); telson without movable lateral 

spines (Fig. 5c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapenaeus 

5a. Carapace lacking longitudinal and vertical sutures (Fig. 8a); telson with movable lateral 

spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

6a. Rostrum extending far beyond eye; pterygostomian spine present (Fig. 8a); deep water. .Penaeopsis 

6b. Rostrum short, not extending beyong eye; pterygostomian spine absent (Fig. 9); on 

reefs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachypenaeopsis 

(a single species, T. richtersii, in the area) 

pterygostomian spine 


Fig. 8 Penaeopsis 

b) petasma 

(dorsal view) 

Fig. 9 Trachypenaeopsis richtersii 


7a. Third maxilliped with epipod; male petasma asymmetrical (Fig. 10b); pelagic (Fig. 10a) . .Funchalia 

(a single species, F. taaningi, in the area) 

7b. Third maxilliped without epipod; male petasma symmetrical (Fig. 11b); benthic . . . . . . . . . → 8 

8a. Fifth leg without exopod (carapace without longitudinal or vertical sutures, Fig. 11a) . .Metapenaeus 

8b. Fifth leg with exopod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →9 

a) lateral view 

Fig. 10 Funchalia taaningi 


b) petasma 



Fig. 11 Metapenaeus 

b) petasma 

(ventral view)

Penaeidae 893 

9a. Carapace without longitudinal or vertical sutures; second leg with ischial spine; eyes 

small (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atypopenaeus 

(the genus Miyadiella is not included in this key, as its status is rather controversial and specimens are interpreted 

as juveniles of Atypopenaeus by some authors) 

9b. Carapace with both longitudinal and vertical sutures (Figs 13 and 14); second leg 

without ischial spine; eyes large . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →10 

10a. Body naked, with crests and grooves on carapace distinct; longitudinal suture usually 

long (Fig. 13); third leg without epipod . . . . . . . . . . . . . . . . . . . . . . . . Parapenaeopsis 

10b. Body usually hairy, with crests and grooves on carapace obscure (Fig. 14); longitudinal 

suture short; third leg generally with epipod . . . . . . . . . . . . . . . . . . . . . Trachypenaeus 




Fig. 12 Atypopenaeus 


Fig. 13 Parapenaeopsis 

vertical 

suture 


Fig. 14 Trachypenaeus 

vertical 

suture 

Key to the species of Atypopenaeus occurring in the area 

1a. Rostrum short, not extending beyond eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Rostrum extending far beyond eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2a. Hepatic spine present . . . . . . . . . . . . . . . . . . . . . . . . . . Atypopenaeus stenodactylus 

2b. Hepatic spine absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atypopenaeus dearmatus 

3a. Crests on fourth and fifth abdominal segments very high and blade-like; fifth leg very 

slender, extending beyond body; colour pale pink . . . . . . . . . . . . . . Atypopenaeus bicornis 

3b. Crests on fourth and fifth abdominal segments not blade-like; fifth leg about 2/3 the 

length of the body; colour reddish orange . . . . . . . . . . . . . . . . . Atypopenaeus formosus 

Key to species of Metapenaeopsis likely to be encountered in fishing activities in the area 

Note: in addition to the following key characters, the shape of the petasma and thelycum can be very helpful 

for quick identification of mature specimens (see figures shown in the respective species accounts). For a 

full key to Metapenaeopsis, see the revisions by A.Crosnier (1987-1994), in “Bull. Mus. natn. Hist. nat., 

Paris” and “Rèsultats des Campagnes MUSORSTOM”. 

1a. Posterolateral carapace with stridulating organs (Fig. 15). . . . . . . . . . . . . . . . . . . . . → 2 

1b. Posterolateral carapace without stridulating organs . . . . . . . . . . . . . . . . . . . . . . . . → 7 

2a. Stridulating ridges usually 4 to 6 (Fig. 15b) . . . . . . . . . . . . . . . . Metapenaeopsis stridulans 

2b. Stridulating ridges more than 7 (Fig. 15a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Rostrum far exceeding antennular peduncle, usually bearing 5 widely spaced upper 

teeth (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeopsis novaeguineae 

3b. Rostrum extending to about tip of antennular peduncle, bearing more than 6 regularly 

spaced upper teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

more than 

7 ridges 

6 ridges 

a) Metapenaeopsis toloensis b) Metapenaeopsis stridulans 

Fig. 15 stridulating ridges on posterolateral carapace 

Fig. 16 Metapenaeopsis novaeguineae 

(from Crosnier, 1994)


4a. Pterygostomian spine moderately to well developed (Fig. 17) . . . . . . . . . . . . . . . . . . → 5 

4b. Pterygostomian spine reduced (Fig. 18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 


Fig. 17 Metapenaeopsis barbata 


5a. Dorsal crest of third abdominal segment flat or 

very slightly concave (Fig. 19a) . . Metapenaeopsis barbata 

5b. Dorsal crest of third abdominal segment with 

deep median groove (Fig. 19b) . . Metapenaeopsis toloensis 

6a. Rostrum with 9 or 10 upper teeth . . Metapenaeopsis rosea 

6b. Rostrum with 7 or 8 upper teeth (Fig. 18) . . . 

. . . . . . . . . . . . . . . . . . Metapenaeopsis palmensis 

7a. Rostrum forming a crest (Fig. 20) . . . . . . . 

. . . . . . . . . . . . . . . . . . Metapenaeopsis lamellata 

7b. Rostrum not forming a crest . . . . . . . . . . . . . . . → 8 

8a. Pterygostomian spine very strong (Fig. 21) . . 

. . . . . . . . . . . . . . . . . . . . .Metapenaeopsis wellsi 

8b. Pterygostomain spine minute or absent 

(Fig. 22) . . . . . . . . . . . . . Metapenaeopsis mogiensis 

rostral 

crest 


Fig. 20 Metapenaeopsis lamellata 



Fig. 21 Metapenaeopsis wellsi 


Key to the species of Metapenaeus occurring in the area 

Remarks on key characters: the shape of the petasma and thelycum are the main taxonomic characters 

to separate species of this genus (see figures shown in the respective species accounts). Therefore, correct 

identification of juveniles and immature specimens is sometimes difficult. See Miquel (1982, Zool. Verh., 

195) for more information on the species of this genus. 

1a. Rostrum very short and high, not extending 

beyond eye (Fig. 23) . . . Metapenaeus lysianassa 

1b. Rostrum extending beyond eye . . . . . . . . . . → 2 


Fig. 18 Metapenaeopsis palmensis 


dorsal view 

dorsal 

ridge 

dorsal 

ridge 

dorsal view 

transverse view transverse view 

a) Metapenaeopsis b) Metapenaeopsis 

barbata 

toloensis 

Fig. 19 third abdominal segment 




Fig. 22 Metapenaeopsis mogiensis 


Fig. 23 Metapenaeus lysianassa

Penaeidae 895 

2a. Rostrum armed with teeth along entire upper border (Fig. 24) . . . . . . . . . . . . . . . . . . → 3 

2b. Rostrum unarmed in its distal 1/3 to 1/2 (Figs 25 and 26) . . . . . . . . . . . . . . . . . . . . →16 

epigastric tooth 

unarmed 

Fig. 24 Metapenaeus ensis 

Fig. 25 Metapenaeus eboracensis 

(after Miguel, 1982) 

3a. Telson with 3 pairs of large movable spines (Fig. 27b) . . . . . . . . . . . . . . . . . . . . . . → 4 

3b. Telson without large movable spines (although sometimes with many minute spinules; 

Fig. 27a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

distinct crest 

epigastric tooth 

Fig. 26 Metapenaeus tenuipes 


movable 

spines 

a) Metapenaeus ensis b) Metapenaeus anchistus 

Fig. 27 telson and left uropod (dorsal view) 


4a. In males, distolateral projections of petasma not reaching as far as distomedian ones, 

armed with a pair of spinules on each externodistal side; in females, thelycum with a 

posteromedian ogival boss, coxal projection of fourth leg very long, flat and truncate 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus intermedius 

4b. In males, distolateral projections of petasma exceeding distomedian ones, without 

external spinules; in females, thelycum without a posteromedian boss, coxal projection 

of fourth leg a conical process . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

5a. In males, a wide space between distomedian projections of petasma; in females, lateral 

plates of thelycum with raised posterior edge, a deep fissure between lateral plates and 

posterior transverse ridge . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus endeavouri 

5b. In males, a narrow space between distomedian projections of petasma; in females, 

lateral plates of thelycum without raised posterior edge and continuous to posterior 

transverse ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus anchistus 

6a. First leg with ischial spine subequal to basial spine . . . . . . . . . . . . . Metapenaeus suluensis 

6b. Ischial spine of first leg much smaller then basial spine, minute or absent . . . . . . . . . . . . → 7 

7a. In males, distomedian projection of petasma swollen; in females, lateral plates of 

thelycum with raised lateral or ventral ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

7b. In males, distomedian projection of petasma not swollen; in females, lateral plates of 

thelycum without raised ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 12


8a. In males, each distomedian projection of petasma forming a broad outward curved distal 

tooth; in females, anterior plate of thelycum with a pair of anterolateral rounded tubercles, 

each lateral plate with a short anteromedian ridge and a patch of setae . . . . . Metapenaeus insolitus 

8b. In males, each distomedian projection of petasma without distolateral teeth; in females, 

anterior plate of thelycum without tubercles, lateral plates with long ridges and without 

patches of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .→ 9 

9a. In males, distomedian projections of petasma concealing distolateral ones in ventral 

view, merus of fifth leg with a long inward curved spiniform process followed by a distinct 

row of tubercles; in females, ridges on lateral plates of thelycum subparallel or forming 

posteriorly distinctly inward curved processes . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

9b. In males, distomedian projections of petasma not concealing distolateral ones in ventral 

view, merus of fifth leg with a tubercle; in females, ridges on lateral plates of thelycum 

closer to each other posteriorly and without distinctly inward curved processes . . . . . . . . → 11 

10a. Rostrum armed with 7 to 9 upper teeth; in males, distomedian projections of petasma 

reaching about as far as distolateral ones; in females, lateral plates of thelycum with 

strongly raised crescent-shaped ventral ridges, coming close together posteromedially 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus conjunctus 

10b. Rostrum armed with 8 to 12 upper teeth; in males, distomedian projections of petasma 

reaching well beyond distolateral ones; in females, lateral plates of thelycum with 

strongly raised externolateral edges, a wide space separating them . . . . . . . Metapenaeus ensis 

11a. In males, distomedian projections of petasma directed forward, their inner margins 

almost parallel, tubercle on merus of fifth leg slightly bent inward; in females, ridges on 

lateral plates of thelycum curved outward posteriorly . . . . . . . . . . . . Metapenaeus papuensis 

11b. In males, distomedian projections of petasma directed anterolaterally, diverging, with 

distinct longitudinal grooves, tubercle on merus of fifth leg slightly bent outward; in 

females, ridges on lateral plates of thelycum curved inward posteriorly . . . . Metapenaeus elegans 

12a. Branchiocardiac crest reaching posterior extension of hepatic spine; in males, distomedian 

projections of petasma crescent-shaped; in females, posterior transverse ridge 

behind thelycum with 2 anterolateral rounded projections . . . . . . . . . . . .Metapenaeus affinis 

12b. Branchiocardiac crest ending near posterior 1/3 of carapace; in males, distomedian 

projections of petasma never crescent-shaped; in females, posterior transverse ridge 

behind thelycum without projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →13 

13a. In males, distolateral projections of petasma widening distally, distomedian projections 

not reaching as far as distolateral ones; in females, anterior plate of thelycum very broad 

in its distal half and very narrow posteriorly, lateral plates fused and rounded W-shape 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus demani 

13b. In males, distolateral projections of petasma tapering distally, distomedian projections 

exceeding distolateral ones; in females, anterior plate of thelycum about same width on 

anterior and posterior margins, lateral plates not rounded W-shape . . . . . . . . . . . . . . . →14 

14a. In males, distomedian projections of petasma tubular and diverging; in females, anterior 

margin of anterior plate of thelycum with 2 fang-like teeth and a median indistinct 

tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus dalli 

14b. In males, distomedian projections of petasma either tubular and subparallel or laminose 

and diverging; in females, anterior margin of anterior plate of thelycum with 3 tubercles . . . . . →15 

15a. In males, distomedian projections of petasma tubular and almost parallel, their distal 

part twisted dorsoventrally; in females, distal margin of anterior plate of thelycum 

triangular, median tubercle more prominent than lateral ones . . . . . . . . Metapenaeus bennettae 

15b. In males, distomedian projections of petasma laminose and diverging; in females, distal 

margin of anterior plate of thelycum convex, all tubercles of subequal size . . Metapenaeus moyebi 

16a. Telson armed with 4 pairs of large movable spines . . . . . . . . . . . . . . Metapenaeus macleayi 

16b. Telson without large movable lateral spines (though sometimes with many minute 

spinules) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →17

Penaeidae 897 

17a. In males, basial spine of third leg extremely long and barbed; in females, lateral plates 

of thelycum partially sheathing anterior plate . . . . . . . . . . . . . . . . . Metapenaeus dobsoni 

17b. In males, basial spine of third leg sample, not barbed; in females, lateral plates of 

thelycum not sheathing anterior plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →18 

18a. Rostrum without a distinct crest; adrostral groove extending beyond epigastric tooth 

(Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus eboracensis 

18b. Rostrum with a distinct crest; adrostral groove not reaching epigastric tooth (Fig. 26) . . . . . →19 

19a. In males, distolateral projections of petasma directed outward, distomedian projections 

with slender apical filaments; in females, thelycum with a large anterior and small lateral 

plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus brevicornis 

19b. In males, distolateral projections of petasma directed forward, distomedian projections 

with ribbon-like apical filaments; in females, thelycum with a small anterior and large 

lateral plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metapenaeus tenuipes 

Key to the species of Parapenaeopsis occurring in the area 1/ 

Note: in addition to the following key characters, the shape of the petasma and thelycum can also be very 

helpful for quick identification of mature specimens (see figures shown in the respective species accounts). 

1a. First and second legs with epipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. First and second legs without epipods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

2a. First leg without basial spine . . . . . . . . . . . . . . . . . . . . . . . Parapenaeopsis gracillima 

2b. First leg with basial spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Second leg with basial spine absent or minute; carapace with a 

large dorsoposterior dark brown patch . . . . . . . . Parapenaeopsis uncta 

3b. Second leg usually with a distinct basial spine; carapace without 

conspicuous dark patches on dorsal surface . . . . . . . . . . . . . . . → 4 

4a. Telson armed with 1 or 2 pairs of fixed lateral spines (Fig. 28) 

. . . . . . . . . . . . . . . . . . . . . . . . Parapenaeopsis coromandelica 

4b. Telson without fixed lateral spines . . . . . . . . . . . . . . . . . . . . → 5 

5a. Longitudinal suture short, reaching to about level of hepatic 

spine (Figs 29a, b); male petasma horn-like; in females, anterior 

and posterior plates of thelycum fused medially . . . . . . . . . . . . . → 6 

5b. Longitudinal suture long, extending near to posterior carapace 

(Figs 29c, d); male petasma not horn-like; in females, anterior 

and posterior plates of thelycum separated . . . . . . . . . . . . . . . → 7 

6a. Rostrum usually with 7 or 8 upper teeth (Fig. 29a); third leg without basial spine; in 

males, tip of distolateral projection of petasma with a small dorsal spiniform process; in 

females, posterior plate of thelycum without a median boss . . . . . . . . Parapenaeopsis cornuta 

6b. Rostrum usually with 9 or 10 upper teeth (Fig. 29b); third leg with a basial spine; in 

males, tip of distolateral projection of petasma without dorsal spiniform processes; in 

females, posterior plate of thelycum with a median boss . . . . . . . Parapenaeopsis maxillipedo 

suture 

short 

a) Parapenaeopsis cornuta 

telson 

Fig. 28 Parapenaeopsis 

coromandelica 

b) Parapenaeopsis maxillipedo 

suture 

long 

c) Parapenaeopsis hardwickii d) Parapenaeopsis sculptilis 


1/ The taxonomic status and the relationships of some species of this genus are still unclear. For example, 

Parapenaeopsis probata Hall, 1961 is here treated as a synonym of P. uncta Alcock, 1905, but its status probably 

should be re-examined.


7a. In males, distomedian projections wing-like, wider than long and with anterior margin 

often crenulate; in females, posterior plate of thelycum with a pair of anterolateral 

tooth-like projections and anteromedian margin bearing a transverse row of long hairs; 

body not strikingly cross-banded . . . . . . . . . . . . . . . . . . . . . Parapenaeopsis hardwickii 

7b. In males, distomedian projections rabbit ear-shaped, long and deeply concave ventrally; in 

females, posterior plate of thelycum without anterolateral tooth-like projections but with a 

median tubercle bearing a tuft of long hairs; body strikingly cross-banded . . Parapenaeopsis sculptilis 

8a. Epigastric tooth present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

8b. Epigastric tooth absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →10 

9a. Rostrum long and exceeding antennular peduncle; longitudinal suture extending almost 

to posterior carapace . . . . . . . . . . . . . . . . . . . . . . . . . Parapenaeopsis hungerfordii 

9b. Rostrum short and extending just beyond eyes; longitudinal suture only reaching as far 

as level of hepatic spine . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapenaeopsis venusta 

10a. Rostrum extending far beyond antennular peduncle; longitudinal suture reaching as far 

as level of hepatic spine; vertical suture absent . . . . . . . . . . . . . .Parapenaeopsis arafurica 

10b. Rostrum not exceeding second antennular segment; longitudinal suture extending far 

behind level of hepatic spine; vertical suture present . . . . . . . . . . . . Parapenaeopsis tenella 

Key to the species of Parapenaeus occurring in the area 

Note: the shape of the petasma and thelycum can be very helpful for quick identification of mature specimens, 

and these are therefore depicted at the end of the key (Figs 39, 40, and 41). For more information on the 

species of this genus, see the work of Crosnier (1986, Résultats des Campagnes MUSORSTOM, Vol 2). 

1a. Branchiostegal spine absent 

(Fig. 30) . . . . . . . . . Parapenaeus longipes 

1b. Branchiostegal spine present 

(Figs 31 to 38) . . . . . . . . . . . . . . . . → 2 

2a. Branchiostegal spine situated behind 

anterior edge of carapace 

(Figs 31and 32) . . . . . . . . . . . . . . . → 3 

2b. Branchiostegal spine situated on 

anterior edge of carapace 

(Figs 33 to 38) . . . . . . . . . . . . . . . → 4 

3a. Rostrum slightly curved downward; postrostral crest extending close to posterior border 

of carapace (Fig. 31) (see also Figs 39a and 41a) . . . . . . . . . . . . Parapenaeus investigatoris 

3b. Rostrum slightly curved upward; postrostral crest extending to slightly beyond midcarapace 

(Fig. 32; see also Figs 39b and 41b) . . . . . . . . . . . . . . . . . . . Parapenaeus murrayi 

branchiostegal spine 

Fig. 31 Parapenaeus investigatoris 


4a. Rostrum usually not exceeding 

basial segment of antennular 

peduncle (Figs 33; see also 

Figs 39c and 41c) . . . Parapenaeus fissurus 

4b. Rostrum extending far beyond 

basial segment of antennular 

peduncle . . . . . . . . . . . . . . . . . → 5 

Fig. 30 Parapenaeus longipes 



Fig. 32 Parapenaeus murrayi 



Fig. 33 Parapenaeus fissurus 

(from Crosnier, 1985)

Penaeidae 899 

5a. Epigastric spine approximately above level of hepatic spine (Fig. 34; see also Figs 39d and 

41d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapenaeus perezfarfante 

5b. Epigastric spine distinctly behind level of hepatic spine (Fig. 35) . . . . . . . . . . . . . . . . .→6 epigastric spine 

epigastric spine 

branchiostegal spine hepatic spine 

Fig. 34 Parapenaeus perezfarfante 


6a. Rostrum slender and almost straight 

(see also Fig. 40a) . . Parapenaeus ruberoculatus 

6b. Rostrum robust and sinuous . . . . . . . . . . → 7 

7a. Rostrum extending to about the middle 

of second segment of antennular 

peduncle (Fig. 35; see also 40b and 

41e) . . . . . . . . . Parapenaeus sextuberculatus 

7b. Rostrum extending beyond second 

segment of antennular peduncle 

(Figs 36 to 38) . . . . . . . . . . . . . . . . . → 8 

a) Parapenaeus 

investigatoris 

Fig. 37 Parapenaeus lanceolatus 


subdistolateral 

lobe 

b) Parapenaeus 

murrayi 

c) Parapenaeus 

fissurus 

Fig. 39 distal part of petasma 

(after Crosnier, 1985) 

hepatic spine 

Fig. 35 Parapenaeus sextuberculatus 


Fig. 36 Parapenaeus fissuroides 


Fig. 38 Parapenaeus australiensis 



lobe 

d) Parapenaeus 

perezfarfante 

ventral 

view 

8a. In males, petasma with subdistolateral lobes bifurcate (Fig. 40c); in females median part 

of thelycum bearing a pair of longitudinal swellings (Fig. 41f) . . . . . . . . Parapenaeus fissuroides 

8b. In males, petasma with subdistolateral lobes not bifurcate; in females, median part of 

thelycum without longitudinal swellings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

lateral 

view


9a. In males, petasma with subdistolateral lobes pointed (Fig. 40d); in females, thelycum 

with a distinct median pit (Fig. 41g) . . . . . . . . . . . . . . . . . . . . . Parapenaeus lanceolatus 

9b. In males, petasma with subdistolateral lobes rounded (Fig. 40e); in females, thelycum 

without a pit at middle (Fig. 41h, sometimes bearing a tubercle) . . . . . Parapenaeus australiensis 


ruberoculatus 


investigatoris 

e) Parapenaeus 

sextuberculatus 


lobe 


sextuberculatus 

anterior 

plate 

posterior 

plate 


fissurorides 


lobe 

Fig. 40 distal part of petasma (ventral view) 



murrayi 

f) Parapenaeus 

fissuroides 

longitudinal 

swellings 

Fig. 41 thelycum 



fissurus 

g) Parapenaeus 

lanceolatus 


lanceolatus 

median 

pit 


lobe 

e) Parapenaeus 

australiensis 


perezfarfante 

tubercle of some specimens 

h) Parapenaeus australiensis 

Key to the species of Penaeopsis occurring in the area 

Note: for more information on the species of this genus, see the work of Pérez Farfante (1980, Fish. Bull.: 77). 

1a. Telson generally with 3 pairs of movable spines (Fig. 42a); pterygostomian spine above 

anterolateral corner of carapace (Fig. 43a) . . . . . . . . . . . . . . . . . . . Penaeopsis rectacuta 

1b. Telson generally with 2 pairs of movable spines (Fig. 42b); pterygostomian spine at 

anterolateral corner of carapace (Fig. 43b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2

Penaeidae 901 

2a. In males, petasma with ventral costa produced distally into long spine considerably 

extending beyond level of row of cincinnuli; in females, thelycum with lateral plates 

turning abruptly mesially posterior to midlength, plate bearing short, pedunculate 

posteromedian protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . Penaeopsis eduardoi 

2b. In males, petasma with ventral costa ending distally in a blunt, short process or spine 

not extending beyond level of row of cincinnuli; in females, thelycum with lateral plates 

not turning abruptly mesially posterior to midlength, plate bearing short, subrectangular 

posteromedian protuberance . . . . . . . . . . . . . . . . . . . . . . . . . Penaeopsis challengeri 

3 pairs of 

movable 

spines 

a) Penaeopsis 

rectacuta 

b) Penaeopsis 

eduardoi 


2 pairs of 

movable 

spines 

Key to the species of Penaeus occurring in the area 2/ 

Note: live or fresh specimens of this genus can often be easily distinguished on the basis of their coloration. 

1a. Adrostral crest extending almost to posterior border of carapace; gastrofrontal crest 

present; generally 1 or 2 lower rostral teeth (Fig. 44a) . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Adrostral crest not extending beyond midcarapace; gastrofrontal crest absent; generally 

3 to 6 lower rostral teeth (Figs 44b, c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 




gastrofrontal 

crest 

hepatic crest 

a) Penaeus japonicus 

hepatic 

crest 

2a. Postrostral crest without median 

groove (Fig. 45a); usually 

2 lower rostral teeth (body uniform 

colour) ......Penaeusmarginatus 

2b. Postrostral crest with median 

groove (Figs 45b, c, d); usually 

1 lower rostral tooth . . . . . . . → 3 



spine 

spine 

b) Penaeopsis eduardoi, 

a) Penaeopsis rectacuta 

P. challengeri 

Fig. 43 anterolateral corner of carapace 

b) Penaeus semisulcatus 


a) Penaeus 

marginatus 

b) Penaeus 

japonicus 

postrostral 

crest 

c) Penaeus merguiensis 

c) Penaeus 

latisulcatus 

adrostral 

groove 

median groove 

Fig. 45 carapace (dorsal view) 

2/ This genus is sometimes subdivided into many subgenera or genera by some authors. 

d) Penaeus 

longistylus


3a. Telson unarmed (Fig. 46a; body covered with 

cross bands) . . . . . . . . . . . . Penaeus canaliculatus 

3b. Telson armed with movable lateral spines 

(Fig. 46b) . . . . . . . . . . . . . . . . . . . . . . . . → 4 

4a. First leg bearing a distinct ischial spine 

(Fig. 47a); median groove on postrostral crest 

less than half carapace length (Fig. 45d, body 

without cross bands). . . . . . . . . . Penaeus longistylus 

4b. First leg with ischial spine minute or absent 

(Fig. 47b); median groove on postrostral crest 

extending almost to posterior carapace 

(Figs 45b, c). . . . . . . . . . . . . . . . . . . . . . . → 5 

ischium 

with spine 

a) Penaeus longistylus 

5a. Rostrum with accessory 

ridges; gastrofrontal groove 

divided into 3 at posterior end 

(Fig. 48; body uniformly coloured) 

. . . . . . . . . . Penaeus plebejus 

5b. Rostrum without distinct accessory 

ridges; gastrofrontal 

groove divided into 2 at posterior 

end . . . . . . . . . . . . . . . . . → 6 

6a. Adrostral groove about as 

long as wide as postrostral 

Fig. 47 first leg 


accessory ridge 

gastrofrontal 

groove 

upper part of carapace 

(lateral view) 

Fig. 48 Penaeus plebejus 

a) Penaeus 

b) Penaeus 

canaliculatus 

japonicus 


b) Penaeus japonicus 

thelycum 


crest (Fig. 45b, body covered with cross bands; shape of thelycum unique and pouchlike, 

Fig. 49, those of all other species flap-like) . . . . . . . . . . . . . . . . . Penaeus japonicus 

6b. Adrostral groove distinctly wider than postrostral crest (Fig. 45c, body without cross 

bands) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Penaeus latisulcatus 

7a. Hepatic crest present; body covered with cross bands; generally 3 lower rostral teeth 

(Fig. 44b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

7b. Hepatic crest absent; body semi-translucent and densely covered with minute dark 

brown dots; generally 4 to 6 lower rostral teeth (Fig. 44c) . . . . . . . . . . . . . . . . . . . . →10 

8a. Fifth leg without exopod; hepatic crest nearly horizontal (Fig. 50); antennal flagella not 

banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Penaeus monodon 

8b. Fifth leg bearing a small exopod; hepatic crest sloping anteroventrally (Fig. 51); antennal 

flagella banded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9 



hepatic 

crest 

5 

(after Yu and Chan, 1986) 

th leg 

Fig. 50 Penaeus monodon 

exopod 

hepatic 

crest 

exopods 

5 

(after Yu and Chan, 1986) 

th leg 

Fig. 51 Penaeus semisulcatus

Penaeidae 903 

9a. Adrostral crest extending behind last postrostral 

tooth (Fig. 51); postrostral crest distinctly 

grooved (Fig. 53a); hepatic crest 

extending well behind antennal crest 

(Fig. 51) . . . . . . . . . . . . . . .Penaeus semisulcatus 

9b. Adrostral crest at most extending to last postrostral 

tooth (Fig. 52); postrostral crest without 

a distinct groove (Fig. 53b); hepatic crest 

not extending behind antennal crest (Fig. 52) 

. . . . . . . . . . . . . . . . . . . . . Penaeus esculentus 


hepatic crest 


Fig. 52 Penaeus esculentus 

10a. Third maxilliped of males with distal segment about half as long as second segment 

which bears a tuft of hairs at tip (Fig. 54a, rostral crest high and broadly triangular in 

both sexes; body yellowish to greenish, antennal flagella reddish brown) . . . Penaeus merguiensis 

10b. Third maxilliped of males with distal segment longer or as long as second segment . . . . . . → 11 

11a. Rostrum slightly curved at tip and 

sigmoidal shaped; antennal flagella 

yellowish (third maxilliped of 

males with second segment as 

long as distal segment and bearing 

a tuft of long hairs at tip, Fig. 54b; 

rostral crest slightly elevated in 

both sexes; body yellowish white to 

greyish green) . . . . . . . . . Penaeus indicus 

11b. Rostrum nearly horizontal straight; 

antennal flagella reddish brown . . . . . . → 12 

12a. Third maxilliped of males with distal 

segment 1.5 to 2.5 times longer 

than second segment which bearing 

a tuft of very long hairs at tip 

(Fig. 54c); rostral crest slightly elevated 

in males and moderately 

high in large females; body somewhat 

greenish . . . . . . . Penaeus penicillatus 

distinct 

groove 

a) Penaeus 

merguiensis 

a) Penaeus 

semisulcatus 

b) Penaeus 

esculentus 

Fig. 53 dorsal view of carapace 

b) Penaeus 

indicus 

c) Penaeus 

penicillatus 

d) Penaeus 

silasi 

Fig. 54 distal part of third maxilliped in adult males 

12b. Third maxilliped of males with distal segment as long as second segment which only bears 

a rudimentary tuft of hairs at tip (Fig. 54d); rostral crest slightly to moderately elevated in 

males but broadly triangular in large females; body yellowish white to pinkish . . . . . . Penaeus silasi 

Key to the species of Trachypenaeus occurring in the area 3/ 

Remarks on key characters: occasionally, spermatophores or sperm sacs (a white or yellowish cement-like 

mass) are attached to the thelycum of mature females, making observation difficult. Nevertheless, they can 

be easily removed by using a pin, forceps, or finger nail. 

1a. Second leg with epipod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Second leg without epipod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

3/ The taxonomic status and relationships of some species of this genus are still unclear, and this genus is sometimes 

subdivided into many genera by some authors.


2a. First leg without epipod . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachypenaeus villaluzi 

2b. First leg with epipod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Rostrum curved downward (female thelycum with sharply pointed apex) . Trachypenaeus gonospinifer 

3b. Rostrum straight or curved upward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

4a. Male petasma horn-like; female thelycum with sharply pointed apex . . . . . .Trachypenaeus sedili 

4b. Male petasma T-shaped, female thelycum 

with round apex . . . . . . . . . . . . . . . . . . → 5 

5a. Rostrum straight and armed with 8 to 11 

upper teeth; fourth and fifth abdominal segments 

without posteromedian incisions 

(Fig. 55a); fifth leg extending beyond antennal 

scale; posterior plate of female thelycum 

without distinct notch . . Trachypenaeus longipes 

5b. Rostrum upturned at tip and armed with 6 to 

8 upper teeth; fourth and fifth abdominal 

segments with posteromedian incisions 

(Fig. 55b); fifth leg not reaching tip of antennal 

scale; posterior plate of female thelycum 

having a distinct median notch ...... 

. . . . . . . . . . . . . . . . Trachypenaeus curvirostris 

6a. Male petasma T-shaped; posterior plate of female thelycum as a forwardly directed pocket 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Trachypenaeus malaiana 

6b. Male petasma horn-like; posterior plate of female thelycum not forming a pocket . . . . . . . . →7 

7a. In males, distolateral projections of petasma narrow; in females, anterior plate of 

thelycum without a backwardly directed projection . . . . . . . . . . . . Trachypenaeus anchoralis 

7b. In males, distolateral projections of petasma widely apart; in females, anterior plate of 

thelycum with a backwardly directed projection . . . . . . . . . . . . . Trachypenaeus granulosus 



Atypopenaeus bricornis Racek and Dall, 1965 

Atypopenaeus dearmatus De Man, 1907 

Atypopenaeus formosus Dall, 1957 

Atypopenaeus stenodactylus (Stimpson, 1860) 

Funchalia taaningi Burekenroad, 1940 

Heteropenaeus longimanus De Man, 1896 

Metapenaeopsis aegyptia Galil and Golani, 1990 

Metapenaeopsis angusta Crosnier, 1987 

Metapenaeopsis assimilis (De Man, 1920) 

Metapenaeopsis barbata (De Haan, 1844) 

Metapenaeopsis ceylonica Starobogatov, 1972 

Metapenaeopsis commensalis Borradaile, 1898 

Metapenaeopsis costata Crosnier, 1991 

Metapenaeopsis difficilis Crosnier, 1991 

Metapenaeopsis distincta (De Man, 1907) 

Metapenaeopsis evermanni (Rathbun, 1906) 

Metapenaeopsis fusca R.J.G. Manning, 1988 

Metapenaeopsis gaillardi Crosnier, 1991 

Metapenaeopsis gallensis (Pearson, 1905) 

Metapenaeopsis hilarula (De Man, 1911) 

Metapenaeopsis lamellata (De Haan, 1844) 

Metapenaeopsis laubieri Crosnier, 1991 

Metapenaeopsis liui Crosnier, 1987 

a) Trachypenaeus 

longipes 

III 

IV 

V 

VI 

b) Trachypenaeus 

curvirostris 

posteriormedian 

incisions 

Fig. 55 abdominal segments (dorsal view) 


Penaeidae 905 

Metapenaeopsis mannarensis De Bruin, 1965 

Metapenaeopsis marquesas Crosnier, 1991 

Metapenaeopsis menoui Crosnier, 1991 

Metapenaeopsis mogiensis (Rathbun, 1902) 

Metapenaeopsis novaeguineae (Haswell, 1879) 

Metapenaeopsis palmensis (Haswell, 1879) 

Metapenaeopsis parahilarula Crosnier, 1991 

Metapenaeopsis parapalmensis Crosnier, 1994 

Metapenaeopsis philippii (Bate, 1881) 

Metapenaeopsis propinqua Crosnier, 1991 

Metapenaeopsis provocatoria Racek and Dall, 1965 

Metapenaeopsis quinquedentata (De Man, 1907) 

Metapenaeopsis richeri Crosnier, 1991 

Metapenaeopsis rosea Racek and Dall, 1965 

Metapenaeopsis sibogae (De Man, 1907) 

Metapenaeopsis sinica Liu and Zhong, 1988 

Metapenaeopsis sinuosa Dall, 1957 

Metapenaeopsis spatulata Crosnier, 1991 

Metapenaeopsis stridulans (Alcock, 1905) 

Metapenaeopsis tarawensis Racek and Dall, 1965 

Metapenaeopsis toloensis Hall, 1962 

Metapenaeopsis velutina (Dana, 1852) 

Metapenaeopsis wellsi Racek, 1967 

Metapenaeus affinis (H. Milne Edwards, 1837) 

Metapenaeus anchistus (De Man, 1920) 

Metapenaeus benettae Racek and Dall, 1965 

Metapenaeus brevicornis (H. Milne Edwards, 1837) 

Metapenaeus conjunctus Racek and Dall, 1965 

Metapenaeus dalli Racek, 1957 

Metapenaeus demani (Roux, 1921) 

Metapenaeus dobsoni (Miers, 1878) 

Metapenaeus eboracensis Dall, 1957 

Metapenaeus elegans De Man, 1907 

Metapenaeus endeavouri (Schmitt, 1926) 

Metapenaeus ensis (De Haan, 1844) 

Metapenaeus insolitus Racek and Dall, 1965 

Metapenaeus intermedius (Kishinouye, 1900) 

Metapenaeus lysianassa (De Man, 1888) 

Metapenaeus macleayi (Haswell, 1879) 

Metapenaeus moyebi (Kishinouye, 1896) 

Metapenaeus papuensis Racek and Dall, 1965 

Metapenaeus suluensis Racek and Dall, 1965 

Metapenaeus tenuipes Kubo, 1949 

Parapenaeopsis arafurica Racek and Dall, 1965 

Parapenaeopsis cornuta (Kishinouye, 1900) 

Parapenaeopsis coromandelica Alcock, 1906 

Parapenaeopsis gracillima Nobili, 1903 

Parapenaeopsis hardwickii (Miers, 1878) 

Parapenaeopsis hungerfordi Alcock, 1905 

Parapenaeopsis maxillipedo Alcock, 1906 

Parapenaeopsis sculptilis (Heller, 1862) 

Parapenaeopsis tenella (Bate, 1888) 

Parapenaeopsis uncta Alcock, 1905 

Parapenaeopsis venusta De Man, 1907 

Parapenaeus australiensis Dall, 1957 

Parapenaeus fissuroides Crosnier, 1986 

Parapenaeus fissurus (Bate, 1881) 

Parapenaeus investigatoris Alcock and Anderson, 1899


Parapenaeus lanceolatus Kubo, 1949 

Parapenaeus longipes Alcock, 1905 

Parapenaeus murrayi Ramadan, 1938 

Parapenaeus perezfarfante Crosnier, 1986 

Parapenaeus ruberoculatus Hall, 1962 

Parapenaeus sextuberculatus Kubo, 1949 

Penaeopsis challengeri De Man, 1911 

Penaeopsis eduardoi Pérez Farfante, 1977 

Penaeopsis rectacuta (Bate, 1881) 

Penaeus canaliculatus (Olivier, 1811) 

Penaeus esculentus Haswell, 1879 

Penaeus indicus H. Milne Edwards, 1837 

Penaeus japonicus Bate, 1888 

Penaeus latisulcatus Kishinouye, 1896 

Penaeus longistylus Kubo, 1943 

Penaeus marginatus Randall, 1840 

Penaeus merguiensis De Man, 1888 

Penaeus monodon Fabricius, 1798 

Penaeus penicillatus Alcock, 1905 

Penaeus plebejus Hess, 1865 

Penaeus semisulcatus De Haan, 1844 

Penaeus silasi Muthu and Motoh, 1979 

Trachypenaeopsis richtersii (Miers, 1844) 

Trachypenaeus anchoralis (Bate, 1881) 

Trachypenaeus curvirostris (Stimpson, 1860) 4/ 

Trachypenaeus gonospinifer Racek and Dall, 1965 

Trachypenaeus granulosus (Haswell, 1879) 5/ 

Trachypenaeus longipes (Paulson, 1875) 

Trachypenaeus malaiana Balss, 1933 

Trachypenaeus sedili Hall, 1961 

Trachypenaeus villaluzi Muthu and Motoh, 1979 

References 

Crosnier, A. 1986. Crustacés Décapodes: Penaeidae les espéces indo-ouest-pacifiques du genre Parapenaeus. In 

Résultats des Campagnes MUSORSTOM, Vol. 2, edited by J. Forest. Mém. Mus. natn. Hist. nat., sér. A, Zool., 

133:303-353. 

Crosnier, A. 1991. Crustacea Decapoda: Les Metapenaeopsis indo-ouest-pacifiques sans appareil stridulant 

(Penaeidae). Deuxiéme partie. In Résultats des Campagnes MUSORSTOM, Vol. 9, edited by A. Crosnier. Mém. 

Mus. natn. Hist. nat., (A), 152:155-297. 

Crosnier, A. 1994. Crustacea Decapoda: Les Metapenaeopsis indo-ouest-pacifiques avec un appareil stridulant 

(Penaeidae). In Résultats des Campagnes MUSORSTOM, Vol. 12, edited by A. Crosnier. Mém. Mus. natn. Hist. 

nat., (A), 16:255-337. 

Crosnier, A. 1994. Crustacea Decapoda: Les Metapenaeopsis indo-ouest-pacifiques sans appareil stridulant 

(Penaeidae).Description de deux espéces nouvelles. In Résultats des Campagnes MUSORSTOM, Vol. 12, edited 

by A. Crosnier. Mém. Mus. natn. Hist. nat., (A), 16:339-349. 

Dall, W. and P.C. Rothlisberg. 1990. Taxonomy. In The biology of Penaeidae, edited by W. Dall, B.J. Hill, P.C. Rothlisberg 

and D.J. Staples. Adv. Mar. Biol., 27:55-126. 

Miguel, J.C. 1982. Le genera Metapenaeus (Crustacea, Penaeidae): taxonomie, biologie et peches mondiales. Zool. 

Verh., 195:1-137. 

Pérez Farfante, I. and B.F. Kensley. 1997. Penaeoid and Sergestoid shrimps and prawns of the world. Keys and diagnosis 

for the families and genera. Mém. Mus. natn. Hist. nat., 175:1-233. 

Yu, H.P. and T.Y. Chan. 1986. The illustrated penaeoid prawns of Taiwan. Taipei, Taiwan, Southern Material Center, Inc., 

183 p. 

4/ Includes Trachypenaeus asper Alcock, 1905 which is probably a synonym of T. curvirostris. 

5/ Includes Trachypenaeus pescadoreensis Schmitt, 1931 which is probably a synonym of T. granulosus. 



Penaeidae 907 

Metapenaeopsis barbata (De Haan, 1844) 


FAO names: En - Whiskered velvet shrimp; Fr - Crevette chamois barbulée; Sp - Camarón gamuza barbudo. 


Diagnostic characters: Body densely covered with short hairs; grooves and 

crests on carapace obscure.Rostrum directed slightly upward, almost straight 

and armed with 6 or 7 regularly spaced upper teeth (excluding epigastric tooth 

on carapace); rostrum extending to about tip of antennal scale. Pterygostomian 

spine well developed; generally 16 to 27 stridulating organs present 

on posterolateral carapace. Abdomen with dorsal crest on third segment 

narrow and more than 9 times as long as broad, median groove on crest 

indistinct or very shallow; sixth segment about 2 times as long as fifth segment. 

Petasma of males asymmetrical, left distoventral projection longer, bearing 

7 to 12 well-developed sharp projections at tip, while right distoventral 

projection bears 1 or 2 small spinule(s) at tip. Thelycum of females with 

thelycal plate broadly subquadrate, 0.5 to 0.6 times as long as broad; 

intermediate plate broadly trapezoidal and slightly concave, posterior ridge 

with a small but distinct median tubercle. Telson with 1 pair of fixed and 3 pairs 

of movable lateral spines. Colour: body whitish and mottled with irregular red 

blotches;eyes dark brown;antennal flagella indistinctly crossed with red and white 

bands; legs reddish; pleopods with white markings on sides; uropods reddish, 

with distal and basal parts pale yellowish. 

Size: Maximum body length 11.6 cm (females) and 7.8 mm (males), commonly 

between 7 and 9 cm. 

intermediate plate 

thelycum 


thelycal 

plate 

Habitat, biology, and fisheries: Found on sand, mud or sandy-mud bottoms, from depths of 2 to 219 m, 

usually less than 90 m. Taken mainly by trawls. Appears to be common but nowhere abundant throughout 

its range in the area. Of minor commercial importance, generally taken as bycatch. Marketed mainly fresh 

for local consumption. 

Distribution: Indo-West 

Pacific from the Gulf of Bengal 

to Japan and Indonesia. 

coxal 

plate 

right 

distolateral 

projection 


left 

distolateral 

projection 

petasma (ventral view)


Metapenaeopsis palmensis (Haswell, 1879) 

Frequent synonyms / misidentifications: Metapenaeopsis barbeensis Hall, 1962 / Metapenaeopsis 

novaeguineae (Haswell, 1879); M. stridulans (Alcock, 1905). 

FAO names: En - Southern velvet shrimp; Fr - Crevette chamois méridionale; Sp -Camaróngamuzasureño. 

thelycal 

plate 


Diagnostic characters: Body densely covered with hairs; grooves and 

crests on carapace obscure. Rostrum nearly straight or slightly curved 

upward, armed with 7 or 8 regularly spaced upper teeth (excluding 

epigastric tooth on carapace); rostrum extending just to distal antennular 

segment. Pterygostomian spine reduced and small; 8 to 13 (mostly 9 or 

10) stridulating organs present on posterolateral carapace. Abdomen 

with dorsal crest on third segment narrow and more than 9 times as long as 

broad, median groove on crest narrow but deep; sixth segment about 2 

times as long as fifth segment. Petasma of males asymmetrical, left 

distoventral projection longer, bearing about 10 rather blunt projections 

at tip; right distoventral projection usually without spinules at tip. 

Thelycum of females with thelycal plate slightly concave, inverted 

trapezoidal, 0.6 to 0.7 times as long as broad; intermediate plate concave, 

with lateral parts expanded into 2 small flaps, posterior edge very 

thick. Telson with 1 pair of fixed and 3 pairs of movable lateral spines. Colour: 

body whitish, mottled with irregular red stripes; eyes dark brown; antennal 

flagella pale red; legs whitish, with some red spots on the sides of 

pleopods; distal half of uropods reddish. 

right 

distolateral 

projection 

petasma (ventral view) 


left 

distolateral 

projection 

Size: Maximum body length 12 cm (females) and 8.5mm (males), commonly between 5 and 8 cm. 

Habitat, biology, and fisheries: Found on sandy-mud or mud bottoms, from depths of 5 to 100 m, usually 

less than 90 m. Taken mainly as bycatch in trawls. Probably the most common species of the genus in the 

area, but nowhere very abundant 

and only of limited commercial 

importance, due to its 

small size. Marketed mainly 

fresh for local consumption. 


Pacific from the western coast 

of Thailand to Japan and 

Australia. 

coxal 

plate 

thelycum 

(after Crosnier, 1994)

Penaeidae 909 

Metapenaeus anchistus (De Man, 1920) 

Frequent synonyms / misidentifications: None / Metapenaeus endeavouri Schmitt, 1926; 

M. intermedius (Kishinouye, 1900). 

FAO names: En - Spiny greasyback shrimp. 


covered with fine pubescence. 

Rostrum distinctly directed 

upward, bearing 10 to 12 teeth 

along entire upper margin, almost 

straight and slightly 

merus of fifth leg (male) 

curved downward at tip; rostrum extending to about distal segment of antennular 

peduncle. Postrostral crest low. Branchiocardiac crest distinct. First leg with 

distinct ischial spine. In adult males, merus of fifth leg with a basal notch followed 

by a prominent keel. Petasma of males with distolateral projections exceeding 

distomedian processes and without external spinules; distomedian 

processes triangular and close to each other. Thelycum of females with 

anterior plate expanded at anterior half; lateral plates generally leveled, 

posterior margin not raised and continuous with posterior transverse ridge; 

coxal projections of fourth leg conical. Telson with 3 pairs of large lateral 

distolateral 

projection 

anterior plate 


thelycum 

distomedian 

projection 


spines. Colour: body rather yellowish and covered with dense dark brown dots; eyes dark brown; rostrum 

and dorsal abdominal crest black-brown; antennal flagella reddish brown; legs somewhat yellowish or pink; 

pleopods more pinkish, with white markings on outer sides; distal part of uropods bluish, with margins 

reddish brown to somewhat purplish. 

Size: Maximum body length 16.5 cm (females) and 14.6 cm (males), commonly between 7 and 14 cm. 

Habitat, biology, and fisheries: Found mainly in depths to about 30 m. Caught by trawls and fish corrals. 

Commonly found in the Philippines and of moderate commercial importance because of its relatively large 

size; apparently less common 

or even rare in other parts of 

the area. Marketed mainly 



from the Strait of Malacca to the 

Philippines and Fiji.


Metapenaeus ensis (De Haan, 1844) 

Frequent synonyms / misidentifications: Metapenaeus ensis baramensis Hall, 1962; M. incisipes (Bato, 

1888); M. mastersii (Haswell, 1879); M. philippinensis Motoh and Muthu, 1979 / Metapenaeus monoceros 

(Fabricius, 1798). 

FAO names: En - Greasyback shrimp; Fr - Crevette glissante; Sp - Camarón resbaloso. 


Diagnostic characters: Body covered 

with fine pubescence. Rostrum 

armed with 8 to 11 teeth along 

entire upper margin, nearly 



straight and extending to about tip of antennular peduncle. Postrostral crest 

low. Branchiocardiac crest generally distinct and curved, almost reaching 

hepatic spine. First leg bearing a small ischial spine. In adult males, merus of 

fifth leg with a basal notch followed by a long, inwardly curved spine-like 

process and a row of tubercles. Petasma of males with distomedian process 

very large and triangular, covering almost entire distolateral projection in 

ventral view. Thelycum of females with long anterior plate; lateral plates with 

posterolateral edges strongly raised and curving inward, forming a pair of 

triangular projections. Telson without distinct lateral spines. Colour: body 


thelycum 

distomedian projection 


greyish green or dark green and covered with dense dark brown dots, large adults somewhat 

pinkish; eyes black-brown; antennal flagella reddish; legs generally whitish, in large adults basal segments 

covered with red bands; distal part of uropods somewhat bluish with reddish brown margins. 


Habitat, biology, and fisheries: Mainly in turbid waters down to a depth of 95 m over bottoms of mud, sandy-mud 

or silt. Juveniles are found in estuaries and backwaters, sometimes also in seagrass beds, mangrove banks, mud 

flats, and open channels. Caught by trawls, set nets, scoop nets, traps, and artisanal gear. Also a common 

byproduct or a secondary species in prawn culture. Probably the most abundant and the most commercially 

important species of the genus in the area, constituting a large part of the Metapenaeus catches and pond industry 

in the Philippines (609 t and 670 t, respectively, in 1987), Singapore, Thailand (11 400 t and 2 700 t, respectively, in 

1987), Indonesia (17 588 t and 13 784 t, respectively, in 1987), Viet Nam, and Malaysia. Fished commercially in 

northern Australia, with a catch of about 2 400 t (together with Metapenaeus endeavouri, catches from western 

Australia not included) during 

the annual period of 

1989/1990. Marketed mainly 

fresh or frozen, also cooked 

or salted and sometimes used 

as bait, consumed locally and 

exported. 


from the eastern coast of 

India and Sri Lanka to Japan 

and Australia.

Penaeidae 911 

Metapenaeus intermedius (Kishinouye, 1900) 


FAO names: En - Middle shrimp; Fr - Crevette ceinture; Sp - Camarón cintura. 

thelycum 


distolateral 

projection 



Rostrum distinctly directed 

upward and very straight, 


bearing 8 to 12 teeth along 

entire upper margin and ex- 


tending to about distal segment 

of antennular peduncle. Postrostral 

crest low. Branchiocardiac crest distinct. First leg bearing a distinct ischial spine. In adult males, 

merus of fifth leg with a basal notch followed by a distinct keel. Petasma of males with distomedian 

processes triangular and exceeding distolateral projections; outer margin of distomedian processes 

bearing an external spinule. Thelycum of females bearing a posteromedian ogival boss; 

posterior edges of lateral plates slightly raised; coxal projections of fourth leg long, flat and 

truncate. Telson with 3 pairs of large lateral spines. Colour: body somewhat whitish, becoming 

slightly pinkish in large adults, covered with dense dark brown dots; eyes dark brown; rostrum, outer 

margin of antennal scale, and dorsal abdominal crest black-brown; antennal flagella reddish brown; legs 

whitish; pleopods slightly reddish with white markings on outer sides; distal part of uropods bluish, with 

margins reddish brown to somewhat purplish. 


Habitat, biology, and fisheries: Found mainly in depths to 130 m, usually between 20 and 60 m. Taken 

mainly by trawls. Very common 

in the Strait of Malacca 

and Gulf of Thailand, probably 

also abundant in Viet Nam. Of 

moderate commercial importance 

because of its relatively 

large size. Marketed mostly 



Pacific from the Andaman 

Sea to Malaysia, southern 

coast of China, and Japan.


Metapenaeus moyebi (Kishinouye, 1896) 

Frequent synonyms / misidentifications: Metapenaeus burkenroadi Kubo, 1954 / Metapenaeus dalli Racek, 

1957; M. mastersii (Haswell, 1879) (= M. ensis (De Haan, 1844)). 

FAO names: En - Moyebi shrimp; Fr - Crevette moyebi; Sp - Camarón moyebi. 




Rostrum armed with 7 to 10 

teeth along entire upper margin, 

nearly straight or slightly 

uptilted at tip, extending to 



distolateral 

projection 

about middle of distal antennular article. Postrostral crest low. Branchiocardiac 

crest weak and indistinct. First leg with ischial spine small or nearly 

absent. In adult males, merus of fifth leg with a basal notch followed by a twisted 

keeled tubercle. Petasma of males with distomedian process enlarged and 

laminose or flap-like, strongly projected forward and diverging; distolateral 

projection directed anterolaterally. Thelycum of females with anterior plate 

distomedian 

projection 


thelycum 


flask-shaped, its anterior margin slightly convex and bearing 3 tubercles of subequal size; lateral 

plates kidney-shaped and often with angular contours. Telson without distinct lateral spines. 

Colour: body semi-translucent, somewhat pale green and covered with with dense dark brown dots; 

eyes black-brown; antennal flagella reddish; pleopods slightly pinkish; distal part of uropods somewhat 

yellowish green and with reddish brown margins. 

Size: Maximum body length 12.6 cm (females) and 8.3 cm (males), commonly between 5.5 and 9.5 cm. 

Habitat, biology, and fisheries: Found on mud or sandy-mud bottom in estuaries, backwaters, and 

nearshore waters to depths of about 45 m. Caught by trawls, set nets, seines, traps, and artisanal gear. 

Appears to be quite abundant throughout its range in the area and is frequently seen in local markets, but 

only of secondary economic 

importance due to its relatively 

small size. Also reported 

to be used for 

aquaculture in Malaysia and 

Singapore but probably not 

the main cultured species 

there. Marketed mainly fresh 

for local consumption. 



India and Sri Lanka to Japan 

and Indonesia.

Penaeidae 913 

Parapenaeopsis hardwickii (Miers, 1878) 

Frequent synonyms / misidentifications: None / Parapenaeopsis cultrirostris Alcock, 1906 (undetermined 

taxonomic status; generally considered as a synonym of Parapenaeopsis sculptilis (Heller, 1862). 

FAO names: En - Spear shrimp; Fr - Crevette javelot; Sp - Camarón lanzón. 


Diagnostic characters: Body naked and smooth. 

Rostrum armed with 9 to 11 upper teeth; in 

females, rostrum very long and of sigmoidal 

shape,with distal 1/3 to 1/2 toothless,extending 

far beyond antennular peduncle;in adult males, 

unarmed portion absent and slightly curving 

downward, only reaching to middle of second 

antennular segment. Longitudinal suture extending 

to about 3/4 or more carapace length. 

First and second legs bearing epipod and basial 

spine, basis of third leg unarmed. Petasma of 

rostrum of large males 

posterior plate 

thelycum 


distolateral 

projection 


lobe 


males with distomedian projection bluntly protruded and short, somewhat wing-like, anterior margin 

often crenulate; distolateral projection short and directed laterally. Thelycum of females with anterior plate 

concave and semi-circular in shape; posterior plate flat, its anterior margin slightly convex and bearing 

a transverse row of long hairs, with anterolateral angles strongly protruded forward. Telson bearing only 3 

to 5 pairs of minute movable lateral spinules. Colour: body greyish to greenish grey, sometimes pink, and 

densely covered with dark-coloured dots; eyes dark brown; rostrum black-brown; basal 1/3 of antennal flagella 

crossed with brown narrow bands;legs generally whitish to pinkish;pleopods reddish and with white and yellowish 

green markings on lateral surfaces; uropods dark reddish brown with yellowish margins. 


Habitat, biology, and fisheries: Found from the coastline to depths of about 90 m, usually less than 20 m, on 

bottom of mud, sandy-mud or sand. Juveniles mainly inhabit estuaries and backwaters. Caught by trawls, 

sometimes also by boat seines and stake nets, with females usually outnumbering males in the catches. 

Probably the most common species of the genus in the area and of moderate commercial importance. Appears 

to be rather abundant along the 

Malay Peninsula and in adjacent 

waters, where it often 

constitutes a significant part 

of the bycatch of prawn fishery. 

Marketed mainly fresh for 

local consumption. 


from Pakistan to Taiwan 

Province of China and Indonesia.


Penaeus canaliculatus (Olivier, 1811) 

Frequent synonyms / misidentifications: None / Penaeus japonicus Bate, 1888. 

FAO names: En - Witch prawn; Fr -Crevettesoricère;Sp - Camarón brujo. 

anterior 

process 

distomedian 

projection 


thelycum 

posterior 

process 

ventral 

costa 

Diagnostic characters: Carapace with grooves and crests very distinct, bearing 

both gastrofrontal and hepatic crests; rostrum armed with 10 or 11 upper teeth 

(including those on carapace) and 1 lower tooth; postrostral crest well developed 

and with a deep median groove throughout its length; adrostral groove extending 

almost to posterior margin of carapace and slightly wider than postrostral 

crest; posterior end of gastrofrontal groove divided into 2. First leg without 

ischial spine. Petasma of males with very short distomedian projections overhanging 

distal margin of costae. Thelycum of females formed by 2 subrectan- petasma (ventral view) 

gular lateral plates, with their anterolateral angles diverging; anterior process 

suboval; posterior process somewhat triangular. Telson without lateral spines. Colour: body pale 

yellowish and crossed with dark brown transverse bands; those on carapace not extending over 

lower half of carapace while those on last abdominal segment usually continuous to the ventral 

margin; eyes dark brown; antennal scale somewhat greenish and with white tips, flagella yellowish; legs 

yellowish to whitish; pleopods yellowish to reddish and with brown and white spots at bases; distal part 

of uropods with a patch of bright yellow, followed by another patch of bright blue, and with reddish 

margins. 


Habitat, biology, and fisheries: Found on sandy bottoms, from shallow water to depths of about 50 m. 

Taken by trawls and artisanal gear. An occasional bycatch in fisheries for other Penaeus species throughout 

its range in the area, but reported to be rather common in eastern New Guinea. Marketed fresh and frozen, 

often mixed with other species of Penaeus, and consumed locally. Can be easily confused with Penaeus 

japonicus in colour, and 

Penaeus latisulcatus when 

without colour. 


in the Indo-West Pacific, 


to the Red Sea, Taiwan 

Province of China, Okinawa, 

and Polynesia. 

lateral 

lobe

Penaeidae 915 

Penaeus esculentus Haswell, 1879 


FAO names: En - Brown tiger prawn; Fr - Crevette tigrée brune; Sp - Camarón tigre marrón. 


distolateral 

projection 


thelycum 

(after Dall, 1957) 

ventral costa 

Diagnostic characters: Carapace with grooves and crests distinct, rostrum 

generally armed with 5 to 7 upper teeth (including those on carapace) and 3or 

4 lower teeth; postrostral crest well developed and reaching nearly to posterior 

margin of carapace, without a distinct median groove; adrostral crest at most 

extending to last postrostral tooth; gastrofrontal crest absent; hepatic crest 

short, not extending behind antennal crest and slightly sloping anteroven- petasma (ventral view) 

trally. Fifth leg bearing an exopod (somewhat hidden beneath carapace). 

Petasma of males with distomedian projections overhanging distal margin of (after Dall, 1957) 

costae. Thelycum of females formed by 2 suboval lateral plates with tumid lips; anterior process rounded 

and with lateral edges somewhat raised, posterior process convex and partly inserted between lateral 

plates. Telson without lateral spines.Colour: body brownish and covered with mud-yellow cross bands; 

eyes light brown with many black dots; rostral teeth reddish brown; antennal flagella alternated with white 

and brown bands; both legs and pleopods reddish and with some white markings on basal segments; 

distal half of uropods brown with red margins. 

Size: Maximum body length 23.5 cm (females) and 19 cm (males), commonly between 15 and 20 cm. 

Habitat, biology, and fisheries: On the continental shelf from the coastline to a depth of 200 m, but usually 

between 10 and 20 m, on mud, sandy-mud or coarse bottoms. Juveniles inhabit shallow waters in estuaries, 

or are associated with seagrass beds, and sometimes found on the top of coral reef platforms. Feed 

primarily at night and are caught then by demersal otter trawls or beam trawls. Fished commercially in 

Australia, with a catch of 

about 3 300 t (together with 

Penaeus semisulcatus) in the 

Northern Prawn Fishery during 


1989/1990. Most of the catch 

is exported (mainly to Japan) 

and packed as frozen whole 

“green” (uncooked) prawns. 

Distribution: Endemic to 

Australia from Sharks Bay 

(Western Australia) to Wallis 

Lake (New South Wales).


Penaeus indicus H. Milne Edwards, 1837 PNI 

Frequent synonyms / misidentifications: None / Penaeus merguiensis De Man, 1888; P. penicillatus 

Alcock, 1905; P. silasi Muthu and Motoh, 1979. 

FAO names: En - Indian white prawn; Fr - Crevette royale blanche (des Indes); Sp - Camarón blanco de la India. 

posterior 


process 

Diagnostic characters: Carapace rather smooth, lacking gastrofrontal 

and hepatic crests; adrostral crest extending as far as or just before 

epigastric tooth; rostrum slightly curved at tip and sigmoidal-shaped, 

usually bearing 7 to 9 upper teeth (including those on carapace) and 3to 

6 lower teeth; rostral crest generally slightly elevated in large specimens 

including adult females (but still with crest in females slightly 

higher than in males); postrostral crest extending near to posterior margin 

of carapace; gastro-orbital crest distinct, extending over posterior 3/5 

to 2/3 of distance between hepatic spine and orbital margin. In adult 

males, third maxilliped with distal segment about as long as second 

segment which bears a tuft of dense long hairs (same length as distal 

distal 2 

segments of 

third maxilliped 

(male) 


thelycum 

distomedian 

projection 

segment)attip.Petasma of males with distomedian projections strongly curved and overhanging distal 

margin of costae. Thelycum of females formed by 2 semi-circular lateral plates, with their median margins 

forming tumid lips; anterior process slightly rounded and slightly convex; posterior process elongated and 

inserted between anterior part of lateral plates; both anterior and posterior processes rather distinct. 

Telson lacking lateral spines. Colour: body semi-translucent, somewhat yellowish white (small 

specimens) or greyish green and covered with numerous minute dark brown dots; eyes light brown 

and covered with some dark brown mesh-like stripes; rostral and abdominal dorsal crests reddish brown 

to dark brown; antennal flagella yellowish; antennular flagella of same colour as body and covered with 

many dark spots; legs translucent and somewhat whitish, pleopods yellowish to pinkish; distal part of 

uropods yellowish with red margins. 

Size: Maximum body length 23 cm (females) and 18.4 cm (males), usually less than 17 cm. 

Habitat, biology, and fisheries: On sandy and muddy bottoms, from the coastline to depths of about 90 m. 

Caught by trawls, fish corrals, gill nets, beach seines, and artisanal gear. Also a suitable candidate for the prawn 

pond industry. An abundant species and of commercial importance in the Philippines, Singapore, and Australia. 

Also reported to be very abundant in Thailand (where it is one of the main pond cultured prawn species), 

Malaysia, and Indonesia. However, as this shrimp is often confused with Penaeus silasi, its reported abundance 

from Thailand to Indonesia remains uncertain. In the Philippines, it is often mixed and sold together with Penaeus 

merguiensis. In northern Australia, it occurs in deeper waters (deeper than 35 m) and is sold at slightly higher 

prices than P. merguiensis (together their 

catches were about 3 000 t in the annual 

period of 1989/1990). Marketed fresh and 

frozen, consumed locally and exported. 

Distribution: Widely distributed in the Indo- 

West Pacific from the eastern coast of Africa 

to the Red Sea, Japan, and Australia. 

ventral 

costa 

lateral 

lobe 

petasma 

(ventral view)

Penaeidae 917 

Penaeus japonicus Bate, 1888 KUP 


FAO names: En - Kuruma prawn; Fr - Crevette kuruma; Sp - Camarón kuruma. 


both gastrofrontal and hepatic crests; rostrum generally armed with 9 or 10 upper 

teeth (including those on carapace) and 1 lower tooth, lacking distinct accessory 

crest on the blade; postrostral crest well developed and with a deep median groove 

throughout its length; adrostral groove extending near to posterior margin of 

carapace and almost as wide as postrostral crest; posterior end of gastrofrontal 

groove divided into 2. First leg without ischial spine. Petasma of males with very 

long distomedian projections overhanging distal margin of costae. Thelycum of 

females a well-developed pouch with double tubes, opened anteriorly; anterior 

and posterior processes fused, forming a subtriangular concave plate. Telson with 

3 pairs of movable lateral spines. Colour: body pale yellowish and crossed with 

dark brown transverse bands; those on carapace generally extending to lower 

half of carapace, last abdominal band interrupted; eyes dark brown; antennal 

scale somewhat greenish with white tips, flagella yellowish; legs whitish to yellowish 

(large specimens); pleopods yellowish to reddish (large specimens) and with 

brown and/or white spots at bases; distal part of uropods with a patch of bright 

yellow, followed by another patch of bright blue, and with red margins. 

distomedian 

projection 


thelycum 

petasma 


ventral 

costa 

Size: Maximum body length 30 cm (females) and 20 cm (males), commonly between 11 and 20 cm. 

Habitat, biology, and fisheries: Inhabits shelf areas from the coastline to depths of about 90 m, but usually 

less than 50 m, on sandy or sandy-mud bottoms. Adults usually are buried in the substrate during the daytime 

and are active at night. Caught by trawls, drift nets, set nets, traps, and artisanal gear. Common, but apparently 

not particularly abundant in the area; usually taken as bycatch in fisheries for other Penaeus species. Marketed 

fresh and frozen, often sold together 

with other species of 

Penaeus. Consumed locally 

and probably also exported to 

some extent. 




to the Red Sea, Japan, 

Australia, and Fiji; also entered 

the eastern Mediterranean 

through the Suez Canal. 

lateral 

lobe


Penaeus latisulcatus Kishinouye, 1896 WKP 

Frequent synonyms / misidentifications: Penaeus latisulcatus hathor Burkenroad, 1959 / None. 

FAO names: En -Westernkingprawn;Fr - Crevette royale occidentale; Sp - Camarón real. posterior 

process 

anterior 

process 


distomedian 

projection 


thelycum 

ventral 

costa 


both gastrofrontal and hepatic crests; rostrum generally armed with 10 or 11 

upper teeth (including those on carapace) and 1 lower tooth, lacking distinct 

accessory crest on the blade; postrostral crest well developed and with a 

deep median groove throughout its length; adrostral groove extending 

almost to posterior margin of carapace and distinctly wider than postrostral 

crest; posterior end of gastrofrontal groove divided into 2. Ischial spine on 

first leg indistinct or nearly absent. Petasma of males with short distomedian petasma (ventral view) 

projections slightly overhanging distal margin of costae. Thelycum of females 

formed by 2 subrectangular lateral plates and with anterolateral angles diverging; anterior process 

strongly bifurcate and horn-like; posterior process triangular. Telson with 3 pairs of movable lateral 

spines. Colour: body generally yellowish green, becoming slightly reddish in large adults; abdominal 

segments each with a short vertical black bar on pleuron; hinges on abdomen often bearing dark 

brown spots and posterolateral part of carapace also sometimes with 2 black stripes positioned at a right 

angle (these 2 markings usually absent in specimens from Australia); eyes brownish and with many dark 

dots; rostrum, postrostral crest, antennal crest, and dorsal crests of abdomen dark brown to reddish brown; 

antennal scale with tip whitish and outer margin dark brown, flagella whitish, becoming yellowish 

distally; legs whitish to bluish or reddish; pleopods yellowish, with bases somewhat reddish; uropods 

bright yellow, with distal half and outer margins of exopods bright blue, other margins reddish. 


Habitat, biology, and fisheries: From the coastline to depths of about 90 m, on bottoms of sand, mud, or 

gravel, with a clear preference for sandy substrates. Adults are buried in the substrate during the daytime and 

come out to feed at night. Mainly taken offshore by trawls, sometimes also caught in shallow waters by traps, 

fish corrals, and other artisanal gear. Common in the area. From 1990 to 1995, the reported annual catch of 

Penaeus latisulcatus in the Western Central Pacific (Thailand) ranged from 1 271 to 1 624 t (FAO Yearbook of 

Fishery Statistics). Its annual catch (together with Penaeus longistylus) in northern Australia was about 100 t in 

the period of 1989/1990. Marketed 

fresh or frozen, consumed 

locally and exported. 



and reported from the eastern 

coast of Africa to the Red Sea, 

Japan, Australia, and Fiji 

(specimens from the Indian 

Ocean are sometimes treated 

as a different subspecies). 

lateral 

lobe

Penaeidae 919 

Penaeus longistylus Kubo, 1943 

Frequent synonyms / misidentifications: Penaeus caesius Dall, 1957; P. jejunus Hall, 1956 / None. 

FAO names: En - Red-spot king prawn; Fr - Crevette royale à taches rouges; Sp - Camarón real manchado. 


posterior 

process 

lateral 

process 

distomedian 

projection 


thelycum 

ventral 

costa 


both gastrofrontal and hepatic crests; rostrum with tip distinctly upturned, armed 

with 10 to 12 upper teeth (including those on carapace) and 1 lower tooth, lacking 

distinct accessory crest on the blade; postrostral crest well developed, with a 

deep median groove extending to about half of its length; adrostral groove 

extending almost to posterior margin of carapace and about twice as wide as 

postrostral crest; posterior end of gastrofrontal groove divided into 2. First leg 

bearing a strong ischial spine. Petasma of males with short distomedian projec- 


tions not overhanging distal margin of costae. Thelycum of females formed by 2 

subrectangular lateral plates with tumid lips; anterior and posterior processes somewhat fused, with 

anterior end rounded. Telson with 3 pairs of movable lateral spines. Colour: body generally yellowish green, 

becoming slightly pinkish in large adults; lower border of carapace reddish; posteroventral carapace 

and abdominal pleura each with a short vertical black bar except fifth pleuron which bears 2 vertical 

bars; lateral side of third abdominal segment often with a large red-brown circular spot; eyes brownish 

and with many dark dots. Rostrum, postrostral crest, distal part of antennal scale, antennal carina, and dorsal 

crests of abdomen dark brown to red-brown; antennal flagella yellowish white; legs yellowish to reddish; 

pleopods yellowish; uropods yellowish at basal half and purplish at distal half, with margins red and black 

except outer margins of exopods bright blue. 


Habitat, biology, and fisheries: Inhabits reefs, sand or sandy-mud bottoms in depths from 18 to 60 m. Taken 

by trawlers offshore at night and in the vicinity of coral reefs, often together with Penaeus latisulcatus. Less 

common than P. latisulcatus in the area, except off northeastern Australia, where a commercial fishery has 

been developed for this species, with an annual catch ranging from 600 t to 1 800 t between 1983 and 1990. 

Marketed fresh or frozen, often 

sold together with P. latisulcatus, 

both locally consumed and 

exported. 

Distribution: Western Pacific; 

reported from Thailand, 

Malaysia, Singapore, South 

China Sea, Taiwan Province 

of China, the Philippines, 

Australia, and Lord Howe Island.


Penaeus marginatus Randall, 1840 

Frequent synonyms / misidentifications: Penaeus teraoi Kubo, 1949 / None. 

FAO names: En - Aloha prawn; Fr - Crevette aloha; Sp - Camarón aloha. 

(after Kubo, 1949) 

posterior 

process 

Diagnostic characters: Carapace with grooves and crests very distinct, 

bearing both gastrofrontal and hepatic crests; rostrum armed with 9 or 

10 upper teeth (including those on carapace) and 1 to 3 (mostly 2) lower 

teeth; postrostral crest well developed but lacking median groove; 

adrostral groove extending almost to posterior margin of carapace and 

about as long as wide as postrostral crest; posterior end of gastrofrontal 

groove divided into 2. First leg armed with a strong ischial spine. Petasma 

of males with very short distomedian projections. Thelycum of females 

formed by 2 subrectangular lateral plates and with anterolateral angles 

diverging; anterior and posterior processes fused with anterior 

end sharply pointed. Telson with 3 pairs of movable lateral spines. Colour: 

body generally yellowish brown and becoming pinkish brown in large 

adults; eyes brownish and with many dark dots; rostrum, and dorsal crests 

of abdomen dark brown; antennal scale reddish brown with white tips, 

flagella whitish but distally becoming somewhat yellowish; legs whitish 

at distal parts and orange-red at basal parts; pleopods pale red; uropods 

bright yellow at basal part and pinkish purple at distal part, submarginal 

regions bright blue, margins reddish. 

Size: Maximum body length 20.5 cm, commonly between 13 and 17 cm. 

distomedian 

projection 



thelycum 


ventral 

costa 


Habitat, biology, and fisheries: Found on bottoms of sand and sandy mud, from the surface to a depth 

of 300 m, with adults preferring deeper waters. Mainly caught offshore by trawlers and apparently not very 

common in the area. 




to Japan, Australia, Hawaii, 

and Cocos Islands.

Penaeidae 921 

Penaeus merguiensis De Man, 1888 PBA 

Frequent synonyms / misidentifications: None / Penaeus indicus H. Milne Edwards, 1837; P. penicillatus 

Alcock, 1905; P. silasi Muthu and Motoh, 1979. 

FAO names: En - Banana prawn; Fr - Crevette banane; Sp - Camarón banana. 

distomedian 

projection 

ventral 

costa 

distal 2 segments of 


(male) 

Diagnostic characters: Carapace rather smooth, lacking gastrofrontal and 

hepatic crests; adrostral crest extending to, or just before, epigastric tooth; tip 

of rostrum horizontally straight, and rostral crest becoming very high and 

broadly triangular in large specimens (even stronger in females), generally 

bearing 6 to 9 upper teeth (including those on carapace) and mostly3to5lower 

teeth; postrostral crest extending near to posterior margin of carapace; gastroorbital 

crest varying from distinct to nearly absent (in some specimens from 

the Philippines to Australia), extending over middle third to posterior 2/3 of 

the distance between hepatic spine and orbital margin. In adult males, third 

maxilliped with distal segment only about half as long as second segment 

which bears a tuft of dense short hairs (slightly shorter than distal segment) 

at tip. Petasma of males with distomedian projections short, not reaching distal 

posterior 

process 

margin of costae. Thelycum of females formed by 2 semi-circular lateral plates, with their median margins 

forming tumid lips; anterior process slightly rounded and concave, obscured by hairs; posterior process 

elongated and inserted between anterior part of lateral plates. Telson lacking lateral spines. Colour: body 

semi-translucent, somewhat yellowish (in youngs and moderate-sized specimens) to greenish (in very 

large specimens) and covered with numerous minute dark brown dots; eyes light brown and covered with 

some dark brown mesh-like stripes; rostral and abdominal dorsal crests reddish brown to dark brown; antennal 

flagella reddish brown; antennular flagella of same colour as body and covered with many dark spots; legs 

translucent and somewhat whitish; pleopods pinkish to reddish; distal part of uropods yellowish green with 

red margins; young specimens often with many longitudinal black broken lines on abdomen. 

Size: Maximum body length 24 cm (carapace length about 6 cm) in females and 20 cm (carapace length 

about 5 cm) in males, commonly between 13 and 15 cm. 

Habitat, biology, and fisheries: On bottoms of sand and mud, from the coastline and river mouths to depths of 

about 55 m, usually less than 20 m; prefers turbid waters. Sometimes forms very dense shoals and good catches 

are often linked with heavy rainfall. One of the economically most important shrimp species in the area. From 

1990 to 1995, the reported annual catch of P. merguiensis in the Western Central Pacific ranged from 44 303 to 

52 087 t. About 4/5 of this production originated in Indonesia. Caught by trawls, fish corrals, pocket netting, beach 

seining, cast nets, and artisanal gear. Also plays an important role in pond culture in Thailand (1 814 t in 1995), 

Malaysia (66 t in 1995), Indonesia (24 610 t in 

1995), and the Philippines (all these data 

from FAO Aquaculture Statistics). Marketed 

mostly fresh and frozen, consumed locally 

and exported. In the Philippines, this prawn 

has only about half the market value of P. 

monodon and in Australia it is sold at slightly 

lower prices than P. indicus. 

Distribution: Indo-West Pacific from the 

Arabian Sea to the South China Sea and Fiji. 

lateral 

lobe 

petasma 




thelycum


Penaeus monodon Fabricius, 1798 GIT 

Frequent synonyms / misidentifications: Penaeus bubulus Kubo, 1949; P. carinatus Dana, 1852; 

P. semisulcatus exsulcatus Hilgendorf, 1879 / None. 

FAO names: En - Giant tiger prawn; Fr - Crevette géante tigrée; Sp - Camarón tigre gigante. 

posterior 

process 

Diagnostic characters: Carapace with grooves and crests distinct, rostrum generally 

armed with 6 to 8 upper teeth (including those on carapace) and 3 lower teeth; 

postrostral crest well developed and reaching nearly to posterior margin of carapace, 

with or without a feeble median groove; adrostral crest extending to just before 

last postrostral tooth; gastrofrontal crest absent; hepatic crest almost horizontal 

and extending far behind antennal crest. Fifth leg without exopod. Petasma 

of males with distomedian projections slightly overhanging distal margin of costae. 

Thelycum of females formed by 2 suboval lateral plates with tumid lips; anterior 

process concave and rounded distally; posterior process subtriangular and partly 

inserted between lateral plates. Telson without lateral spines. Colour: body 

distomedian 

projection 



thelycum 

ventral 

costa 

greyish greenish or dark greenish blue; becoming reddish brown in large adults; carapace covered 

with mud-yellow transverse bands, while abdomen bears dark brown and mud-yellow cross bands; 

eyes light brown with many black dots; antennal flagella uniformly greenish brown; legs of same colour as 

body but sometimes reddish or provided with bright yellow and blue bands; pleopods somewhat reddish or pale 

red, with bases bright yellow and blue; distal half of uropods dark blue or dark brown with a red or 

mud-yellow median transverse band, and margins reddish. 

Size: Probably the largest known penaeid, with a maximum body length of 35 cm (females) and 26.8 cm 

(males), commonly between 12 and 20 cm. 

Habitat, biology, and fisheries: From the coastline to depths of about 150 m, usually less than 30 m, on bottoms 

of sand, mud, or slits. Juveniles usually inhabit seagrass beds, mangrove swamps, and estuaries. Taken by trawls, 

gill nets, seine nets, stake nets, traps, and artisanal gear. More abundant in the western part of the area and of 

major economic importance. From 1990 to 1995, the reported yearly catch of Penaeus monodon in the Western 

Central Pacific ranged from 8 513 to 17 194 t (FAO Yearbook of Fishery Statistics). Also commercially very 

important for aquaculture. Large-scale pond culture of this prawn is practised in many Southeast Asian countries 

such as Thailand (276 982 t in 1995), Malaysia (6 713 t in 1995), Indonesia (84 100 t in 1995), Philippines 

(88 815 t in 1995), and Australia 

(1 613 t in 1995). Marketed 

mostly fresh and frozen, consumed 

locally and exported. In 

the Philippines, it is an expensive 

food item and ranks above 

other Penaeus species. 




to the Red Sea, Japan, Australia, 

and Fiji. 

lateral 

lobe 


Penaeidae 923 

Penaeus penicillatus Alcock, 1905 REP 

Frequent synonyms / misidentifications: None / Penaeus indicus H. Milne Edwards, 1837; P. merguiensis 

De Man, 1988; P. silasi Muthu and Motoh, 1979. 

FAO names: En - Redtail prawn; Fr - Crevette queue rouge; Sp - Camarón colorado. anterior process 


and hepatic crests; adrostral crest extending just 

beyond epigastric tooth; tip of rostrum horizontally straight, 

and rostral crest generally slightly elevated in youngs and 

adult males, to moderately high in large females; rostrum 

usually armed with 7 to 9 upper teeth (including those on carapace) 

and 3 to 5 lower teeth; postrostral crest extending near 

to posterior margin of carapace; gastro-orbital crest distinct, 

occupying 1/2 to 1/3 the distance between hepatic spine and 

orbital margin. In adult males, third maxilliped with distal 

segment much longer than second segment which bears 

a tuft of dense long hairs (as long as distal segment) at 

tip. Petasma of males with distomedian projections slightly bent 

posterior 

process 

distal 2 segments 

of third maxilliped 

(male) 

distomedian 

projection 


thelycum 

and not reaching distal margin of costae. Thelycum of females formed by 2 semi-circular lateral plates, with 

theirmedianmarginsastumidlips;anterior process slightly rounded and obscured by hairs; posterior 

process elongated and inserted between anterior part of lateral plates. Telson lacking lateral spines. Colour: 

body semi-translucent, slightly greenish and covered with numerous minute dark brown dots; eyes light 

brown and covered with some dark brown mesh-like stripes; rostral and abdominal dorsal crests reddish brown 

to dark brown; antennal flagella reddish brown; antennular flagella of same colour as body and covered with 

many dark spots; legs translucent and somewhat whitish; pleopods rather reddish; distal half of uropods 

yellowish to greenish but always with reddish tips. 

Size: Maximum body length 21.2 cm (carapace length 3.3 cm) in females and 16.3 cm (carapace length 

3.1 cm) in males, commonly between 10 and 16 cm. 

Habitat, biology, and fisheries: On soft bottoms, from the coastline to a depth of about 90 m. Caught by 

trawls, seines, scoop nets, and artisanal gear. Marketed fresh and frozen. Reported to be rather common 

in Malaysia, but can be easily 

confused with Penaeus indicus, 

P. merguiensis, and P. 

silasi, and is probably not so 

common in the area. 


from Pakistan to Taiwan 

Province of China and Indonesia. 

costa 



Penaeus plebejus Hess, 1865 PNP 


FAO names: En - Eastern king prawn; Fr - Crevette royale orientale; Sp - Camarón real oriental. 


distomedian 

projection 


thelycum 

posterior 

process 

ventral 

costa 


both gastrofrontal and hepatic crests; rostrum armed with 10 or 11 upper teeth 

(including those on carapace) and 1 lower tooth, bearing distinct accessory 

crest on blade also in adults; postrostral crest well developed, with a deep 

median groove all along its length; adrostral groove extending almost to 

posterior margin of carapace and distinctly wider than postrostral crest; 

posterior end of gastrofrontal groove divided into 3. First leg without ischial 

spine. Petasma of males with short distomedian projections not overhanging 

distal margin of costae. Thelycum of females formed by 2 subrectangular lateral petasma (ventral view) 

plates and with anterolateral angles diverging; anterior process bearing 2 

minute, almost contiguous horns; posterior process somewhat triangular. Telson with 3 pairs of movable 

lateral spines. Colour: body generally yellowish cream, abdominal segments with faint dark spots at 

hinges and a faint dark vertical bar on each pleuron; rostrum dark brown with white tip; eyes deep 

brown; antennal scale with dark brown outer margins and white tip, flagella whitish; postrostral crest 

and dorsal crests of abdomen dark brown to red-brown; legs slightly pinkish; pleopods pinkish to slightly 

yellowish; uropods of same colour as body but with pale blue margins and red-brown fringes. 

Size: Maximum body length 30 cm (females) and 19 cm (males), commonly between 14 and 20.5 cm. 

Habitat, biology, and fisheries: Found on sandy bottoms of bare and vegetated areas from depths of 2 

to 220 m, with a migration pattern from estuaries to deeper marine waters northward along the coasts. 

This prawn forms the basis of an important fishery in eastern Australia, with a catch of about 3 250 t in the 

annual period of 1989/1990 (60% of the total commercial catch originating in southern Queensland). 

Subadults and adults are mainly caught by trawlers offshore at depths below 60 m. Juveniles are caught 

mainly within estuaries by trawling, set pocket, running netting, hauling, seining, and hand netting. Marketed 

locally fresh, frozen, or cooked. Larger specimens are exported mainly to Spain and Japan as “green” 

(uncooked), frozen, or headed prawns. 

Distribution: Restricted to 

eastern Australia, from southern 

Queensland to Victoria 

and Lord Howe Island.

Penaeidae 925 

Penaeus semisulcatus De Haan, 1844 TIP 

Frequent synonyms / misidentifications: Penaeus ashiaka Kishinouye, 1900; P. monodon manillensis 

Villaluz and Arriola, 1938; P. semisulcatus paucidentatus Parisi, 1919 / None. 

FAO names: En - Green tiger prawn; Fr - Crevette tigrée verte; Sp - Camarón tigre verde. 


posterior 

process 

distomedian 

projection 

Diagnostic characters: Carapace with grooves and crests distinct, rostrum 

generally armed with 6 to 8 upper teeth (including those on carapace) and 3 lower 

teeth; postrostral crest well developed and reaching nearly to posterior margin 

of carapace, with a distinct median groove; adrostral crest extending beyond 

last postrostral tooth; gastrofrontal crest absent; hepatic crest long and 

extending behind antennal crest, straight but distinctly sloping anteroventrally. 

Fifth leg with exopod (somewhat hidden beneath carapace). Petasma 

of males with distomedian projections short and not overhanging distal margin of 

costae. Thelycum of females formed by 2 suboval lateral plates with tumid lips; 

anterior process subtriangular and with raised edges, posterior process convex 

and partly inserted between lateral plates. Telson without lateral spines. Colour: 

body reddish brown to pale brown or dark green, carapace covered with 


thelycum 

ventral 

costa 

mud-yellow transverse bands while abdomen including tail fan bears greyish brown and mud-yellow 

cross bands; eyes light brown with many black dots; rostral teeth dark brown; antennal flagella 

alternated with white and brown bands; both legs and pleopods reddish and covered with some white 

markings, with tips of legs whitish, and bases of legs and pleopods also whitish; distal half of uropods 

dark reddish brown and with red margins. 


Habitat, biology, and fisheries: On the continental shelf from the coastline to depths of about 130 m, usually 

less than 60 m, over bottoms of sand, mud, or sandy-mud. Seems to prefer high salinity waters, with juveniles 

often associated with seagrass beds and sometimes found on the top of coral reef platforms. Reported to form 

small shoals and to be predominantly nocturnal, buried in the substrate during the daytime.Mainly taken offshore 

by trawls, sometimes also caught by fish corrals in coastal areas. Commonly seen in the markets of Thailand 

and Indonesia, and the most dominant prawn species in offshore fisheries in the Philippines. FAO’s Yearbook 

of Fishery Statistics records 650 t of this species taken in 1995 from the Western Central Pacific, this figure 

comprising exclusively catches from Thailand. Also caught commercially in Australia, with an annual catch of 

about 3 300 t (together with 

Penaeus esculentus) from the 

Northern Prawn Fishery during 


1989/1990. Marketed mostly 

fresh and frozen, consumed locally 

and exported. 




to Japan, Australia, and 

Fiji; also entered the eastern 

Mediterranean through the 

Suez Canal. 

lateral 

lobe 



Penaeus silasi Muthu and Motoh, 1979 

Frequent synonyms / misidentifications: None / Penaeus indicus H. Milne Edwards, 1837; 

P. merguiensis De Man, 1888; P. penicillatus Alcock, 1905. 

FAO names: En - False white prawn. 

(after Muthu and Motoh, 1979) 


and hepatic crests; adrostral crest extending to 

about epigastric tooth; tip of rostrum horizontally straight; 

rostral crest slightly to moderately elevated in youngs and 

adult males but high and broadly triangular in large females, 

with 7 to 9 upper teeth (including those on carapace) and 4or5 

lower teeth; postrostral crest extending near to posterior margin 

of carapace; gastro-orbital crest distinct, extending over 

posterior 3/5 to 2/3 of distance between hepatic spine and 

orbital margin. In adult males, third maxilliped with distal 

segment about as long as second segment which bears 

only a rudimentary tuft of hairs at tip. Petasma of males 

with distomedian projections slightly bent and not overhanging 

distal margin of costae. Thelycum of females formed by 2 semi- 

distal 2 segments of 


(male) 

posterior 

process 


thelycum 

anterior 

process 


ventral 

costa 

petasma 


(after Muthu and Motoh, 1979) 

circular lateral plates, with their median margins forming tumid lips; anterior process rounded and slightly 

concave, generally distinct and not obscured by hairs (more clear in adults); posterior process elongated 

and inserted between anterior part of lateral plates.Telson without lateral spines.Colour: bodysemi-translucent, 

somewhat yellowish white (small specimens) to pinkish (large specimens) and covered with numerous 

minute dark brown dots (dots becoming obscure in large specimens); eyes light brown and covered with 

some dark brown mesh-like stripes; rostral and abdominal dorsal crests reddish brown to dark brown; antennal 

flagella reddish brown; antennular flagella of same colour as body and covered with many dark spots; legs 

translucent and somewhat whitish, while pleopods yellowish to pinkish; distal part of uropods yellowish with 

red margins; young specimens often with short longitudinal black broken lines on abdomen. 

Size: Maximum body length 20 cm in females (carapace length 4.8 cm) and 15.3 cm in males (carapace 

length 4.1 cm), commonly between 12 and 16 cm. 

Habitat, biology, and fisheries: Found on muddy bottom in shallow waters to a depth of about 36 m. Caught 

by trawlers and probably artisanal gear. An abundant species in the markets of Singapore and of commercial 

importance. Probably also common in Thailand, Malaysia, and Indonesia, where it is often confused with 

Penaeus indicus. Marketed fresh and frozen, 

mainly for local consumption. 

Distribution: Indo-West Pacific; so far only 

reported from Thailand (Andaman Sea and 

Gulf of Thailand), Indonesia, Malaysia, and 

Singapore. Since this shrimp can be easily 

confused with P. indicus, P. merguiensis, 

and P. penicillatus, its actual distribution is 

likely to be wider in the Indo-Malay region.

Penaeidae 927 

Trachypenaeus curvirostris (Stimpson, 1860) TRV 

Frequent synonyms / misidentifications: ? Trachypenaeus asper Alcock, 1905 / Trachypenaeus 

longipes (Paulson, 1875); T. malaiana Balss, 1933. 

FAO names: En - Southern rough shrimp; Fr - Crevette gambri archée; Sp - Camarón fijador arquero. 

Diagnostic characters: Entire body hairy, with grooves and crests on carapace 

obscure. Rostrum slightly upcurved at tip, armed with 6 to 8 upper teeth 

(including those on carapace) and extending to about distal antennular article. 

Postrostral crest low and extending almost to posterior margin of carapace. 

Longitudinal suture on carapace short. First 3 pairs of legs bearing epipods, 

first leg additionally armed with a distinct ischial spine, fifth leg not exceeding 

tip of antennal scale. Abdomen with a small median tubercle on second 

segment, last 4 segments with a low dorsal crest, distinctly incised posteriorly. 

Petasma of males T-shaped, distolateral projections broadly wing-like 



thelycum 


distolateral 

projection 


and directed laterally, distomedian projections small. Thelycum of females with anterior plate concave 

and bluntly pointed anteriorly, bearing a median groove; posterior plate with a distinct median notch 

on anterior margin. Telson generally armed with 3 or 4 pairs of small movable lateral spines. Colour: 

body greyish pink to greyish blue, sometimes whitish on sides; eyes dark brown; antennal flagella 

reddish; legs pinkish and with some white patches; pleopods reddish with white markings on sides; 

uropods almost entirely reddish, with margins whitish to yellowish. 


Habitat, biology, and fisheries: Found on sand, mud, or sandy-mud bottom, from depths of 10 to 300 m, 

but usually between 30 and 60 m. Reported from many places in the area, but its actual distribution is 

unclear as it is often confused with other species of the genus. Probably of minor commercial importance. 

Caught mainly at night by 

trawls and bottom gill nets, 

also by artisanal gear. Marketed 

mainly fresh for local 

consumption. 


from eastern coast of Africa 

to Japan and northern 

Australia, also entered the 

eastern Mediterranean 

through the Suez Canal.


Trachypenaeus malaiana Balss, 1933 

Frequent synonyms / misidentifications: Trachypenaeus fulvus Dall, 1957; T. 

Trachypenaeus curvirostris (Stimpson, 1860). 

unicus Hall, 1961 / 

FAO names: En - Malayan rough shrimp; Fr - Crevette gambri malaise; Sp - Camarón fijador malayo. 

(these FAO names were previously used for Trachypenaeus sedili). 



distolateral 

projection 


thelycum 

distomedian 

projection 

Diagnostic characters: Entire body hairy, with grooves and crests on carapace 

obscure. Rostrum armed with 9 or 10 upper teeth (including those on 

carapace), nearly straight or slightly curved upward at tip, extending to about 

second antennular article. Postrostral crest low and extending almost to 

posterior margin of carapace. Longitudinal suture on carapace short. Epipod 

present only on third leg; ischial spine on first leg small to entirely absent; 

fifth leg extending beyond antennal scale. Abdomen bearing a small median 

tubercle on second segment, last 4 segments with a low dorsal crest. Petasma petasma (ventral view) 

of males T-shaped, with distolateral projections broadly wing-like and directed 

laterally, distomedian projections small. Thelycum of females with 


anterior plate semi-circular and edges slightly raised, bearing a low median groove; posterior plate 

with anterior margin broadly V-shaped and bearing a narrow but deep median notch. Telson generally 

armed with 1 pair of small movable lateral spines. Colour: body generally greyish to greyish blue, 

posterior margin of each abdominal segment covered with an indistinct narrow dark grey band; 

eyes dark brown; antennal flagella greyish brown; legs yellowish to pinkish; pleopods pinkish, with whitish 

to yellowish markings on sides; uropods almost entirely dark grey to brownish and with yellowish 

margins. 

Size: Maximum body length 10.5 cm (females) and 8 cm (males), commonly between 5.5 and 8 cm. 

Habitat, biology, and fisheries: Found in offshore waters at depths from 5 to 60 m, over muddy bottoms, 

juveniles in brackish water. Taken mainly by trawls. Probably the most common species of the genus in the 

area, but of limited commercial 

importance due to its 

small size and nowhere very 

abundant. Forms a bycatch in 

prawn fisheries. Marketed 

mainly fresh or frozen for local 

consumption. 


from the Strait of Malacca to 

the Philippines, South China 

Sea, and northern Australia.

Penaeidae 929 

Atypopenaeus formosus Dall, 1957 

En - Orange shrimp; Fr - Crevette orange; Sp - Camarón naranji. 

Maximum body length 10 cm (females) and 8 cm (males). Inhabits shallow inshore waters to depths 

of about 30 m, usually less than 10 m, over soft mud bottoms. Taken as bycatch in trawls. Of minor 

commercial importance. Its local name in Australia, “go home prawn”, refers to the common reaction 

of fishermen upon finding large numbers of this shrimp in their catches. Marketed fresh or frozen 

for local consumption. Restricted to the waters between New Guinea and northern Australia. 


Atypopenaeus stenodactylus (Stimpson, 1860) 



thelycum 

En - Periscope shrimp; Fr - Crevette périscope; Sp - Camarón periscopio. 

Maximum body length 5 cm (females) and 4 cm (males). On muddy bottoms close to the shore, from 

depths of 10 to 30 m. Taken by trawls, bag nets, and artisanal gear. Generally of no commercial 

importance in the area, due to its very small size, and not particularly abundant in the catches. Marketed 

mainly fresh for local consumption. Indo-West Pacific from India to Japan and northern Australia. 



thelycum 

distolateral projection 







Metapenaeopsis lamellata (De Haan, 1844) 

En - Humpback shrimp; Fr - Crevette bossue; Sp - Camarón jorobado. 

Maximum body length 10 cm (females) and 6 cm (males). At depths between 4 and 200 m, 

associated with hard bottoms of reef and coral debris. Sometimes taken by trawls. Not very common 

throughout its range in the area and of very limited commercial interest. Marketed fresh for local 

consumption. Western Pacific from the Gulf of Thailand to Japan, Australia, and New Caledonia. 


Metapenaeopsis mogiensis (Rathbun, 1902) 

thelycal 

plate 


thelycum petasma (ventral view) 


En - Mogi velvet shrimp; Fr - Crevette chamois mogi; Sp - Camarón gamuza mogi. 

Maximum body length 10.1 cm (females) and 8.2 cm (males); commonly between 5 and 7 cm. Over 

hard bottoms adjacent to coral reefs, from depths of 10 to 156 m, usually less than 50 m. Not particularly 

common in the area and of very limited commercial importance; sometimes taken as bycatch in trawls. 

Marketed fresh for local consumption. Widely distributed in the Indo-West Pacific from the eastern coast 

of Africa to Japan, Australia, and New Caledonia (sometimes divided into 4 subspecies). 


coxal 

plate 

thelycal plate 

coxal 

plate 

right 

distolateral 

projection 

right 

distoventral 

projection 

left 

distolateral 

projection 

left 

distoventral 

projection 


thelycum petasma (ventral view) 

(after Crosnier, 1991)

Penaeidae 931 

Metapenaeopsis novaeguineae (Haswell, 1879) 

En - Northern velvet shrimp; Fr - Crevette chamois nordique; Sp - Camarón gamuza norteño. 

Maximum body length 11.5 cm (females) and 7 cm (males). Over muddy to sandy bottoms, from depths 

of 5 to 30 m. Taken by trawls. A common bycatch in the prawn fishery operating in its range, but of minor 

commercial importance, due to its small size. Marketed fresh or frozen together with other small species 

and used for local consumption. Restricted to the waters between New Guinea and Australia. 


Metapenaeopsis rosea Racek and Dall, 1965 

coxal 

plate 

thelycum 

thelycal 

plate 

right 

distolateral 

projection 

intermediate 

plate 



left 

distolateral 

projection 

En - Pink velvet shrimp; Fr - Crevette chamois rosée; Sp - Camarón gamuza rosado. 

Maximum body length 12.2 cm (females) and 11 cm (males). On muddy bottoms, from depths of 7 

to 52 m. Restricted to the waters between New Guinea and Australia. Not abundant and of very 

limited commercial importance. 


coxal 

plate 

thelycum 

right 

distoventral 

projection 

thelycal 

plate 


left 

distoventral 

projection 



Metapenaeopsis stridulans (Alcock, 1905) 

En - Fiddler shrimp; Fr - Crevette violoneux; Sp - Camarón violinista. 

Maximum body length 10.6 cm (females) and 8.9 cm (males); commonly between 6 and 9 cm. On 

sandy or muddy bottoms, from depths of 9 to 90 m. Taken by trawls, gill nets, seines, and artisanal 

gear. Of limited commercial importance and apparently nowhere abundant in the area. Marketed 

mainly fresh for local consumption. Indo-West Pacific from the Persian Gulf to the South China Sea 

and New Caledonia. 

Metapenaeopsis toloensis Hall, 1962 

En - Tolo velvet shrimp; Fr - Crevette chamois tolo; Sp - Camarón gamuza tolo. 

Maximum body length 10 cm (females) and 8 cm (males); commonly between 6 and 9 cm. On sandy 

or muddy bottoms, from depths of 8 to 73 m. Taken by trawls and artisanal gear. Not particularly 

common in the area and of minor or no commercial importance. Marketed mainly fresh for local 

consumption. Indo-West Pacific from the Maldives to Japan and New Caledonia. 


intermediate 

plate 

coxal 

plate 

thelycum 

thelycum 

thelycal 

plate 

coxal 

plate 

thelycal 

plate 

intermediate 

plate 


right 

distoventral 

projection 

right 

distolateral 

projection 




left 

distolateral 

projection 

left 

distoventral 

projection

Penaeidae 933 

Metapenaeopsis wellsi Racek, 1967 

En - Velvet shrimp. 

Maximum body length 11 cm (females) and 11.8 cm (males). At depths between 13 and 78 m. 

Restricted to northern Australia from Shark Bay (Western Australia) to Gulf of Carpentaria (Queensland). 

Of minor commercial importance, taken as bycatch in trawls. Marketed fresh or frozen for 

local consumption. 


Metapenaeus affinis (H. Milne Edwards, 1837) 



En - Jinga shrimp; Fr - Crevette jinga; Sp - Camarón jinga. 

Maximum body length 18.6 cm for females (perhaps to 22.2 cm) and 14.6 cm for males; commonly 

between 10 and 14 cm. On mud or sandy-mud bottoms, from the coastline to depths of about 90 m, 

usually less than 55 m. Juveniles generally are found in estuaries and backwaters. Caught by 

trawlers, traps, seine nets, and artisanal gear. Marketed fresh or frozen, probably mainly for local 

consumption. Widespread in the Indo-West Pacific from the Persian Gulf to Taiwan Province of 

China, the Philippines, and Papua New Guinea. Can be easily confused with Metapenaeus ensis 

and seems to occur mainly in the western part of the area, from Viet Nam to Thailand, Malaysia, 

and Indonesia. 


thelycum 

thelycal 

plate 



thelycum 

right 

distolateral 

projection 

coxal 

plate 

intermediate 

plate 

left 

distolateral 

projection 




Metapenaeus benettae Racek and Dall, 1965 

En - Greentail shrimp; Fr - Crevette queue verte; Sp - Camarón rabo verde. 

Maximum body length 13 cm (females) and 10.5 cm (males). Found mainly on soft mud bottoms in 

estuaries, coastal lakes, and rivers, to a depth of 22 m. Juveniles generally inhabit upstream waters, 

mangrove canals or intertidal seagrass areas of low salinity and abundant algal cover. Caught by 

beam trawls in rivers and otter trawl in coastal waters, sometimes also by cast nets. Of commercial 

importance in eastern Australia, above all in southeastern Queensland (about 650 t in the annual 

period of 1989/1990). Marketed mainly fresh and consumed locally, also used as bait. Restricted to 

eastern Australia, from Rockhampton (Queensland) to eastern Victoria. 


Metapenaeus brevicornis (H. Milne Edwards, 1837) 

En - Yellow shrimp; Fr - Crevette jaune; Sp - Camarón amarillo. 

Maximum body length 13.2 cm (perhaps to 15.2 cm) for females and 9.8 cm for males. A marine to 

almost fresh-water species, found on sand or mud to depths of about 90 m, usually less than 30 m. 

Juveniles generally found in estuaries, backwaters, and deltas. A common species in the western part 

of the area. Mainly fished by set nets, traps, cast nets, scoop nets, drag nets, and artisanal gear, 

sometimes also by trawls. Often enters Penaeus culture ponds in Thailand, Singapore, Indonesia, and 

Viet Nam, and is harvested together with the cultured species.Marketed usually fresh or frozen, probably 

mainly for localconsumtion. Indo-West Pacific from Pakistan to Viet Nam, and Indonesia. 




thelycum 



thelycum 

distolateral 

projection 



distomedial 

projection 

distolateral 

projection 


Penaeidae 935 

Metapenaeus conjunctus Racek and Dall, 1965 

En - Wood shrimp; Fr - Crevette bois; Sp - Camarón leña. 

Maximum body length 14.3 cm (females) and 11.5 cm (males). Inhabits estuarine and brackish 

waters to depths less than 15 m. Taken by traps and seines. Generally not very common and of 

minor commercial importance. Found mainly mixed in the catches of Metapenaeus ensis or other 

species of the genus. Western Pacific from Thailand to Malaysia, Singapore, Indonesia, the 

Philippines, and northern Australia. 


Metapenaeus dalli Racek, 1957 

anterior 

plate 


thelycum 

distomedian 

projection 

petasma 


En - Western school shrimp; Fr -Crevettedali;Sp - Camarón dalí. 

Maximum body length 9.8 cm (females) and 7.8 cm (males). On bottoms of mud and sand in 

estuarine and brackish waters, to a depth of about 33 m. Caught mainly with hand nets, also by 

seines and traps. Not common in the area and of very limited commercial importance, due to its 

small size. Mainly known from western Australia but also found in the Philippines and Indonesia 

(southeastern coast of Java). 


anterior 

plate 


thelycum 

distolateral 

projection 

distomedian 

projection 



Metapenaeus demani (Roux, 1921) 

En - Demon shrimp; Fr - Crevette diable; Sp - Camarón diablo. 

Maximum body length 12.2 cm (females) and 10.4 cm (males). Over muddy bottoms in estuarine 

and coastal waters, to a depth of 50 m, usually less than 30 m. Caught mainly by trawl nets and 

artisanal gear. Of some commercial importance in the Gulf of Papua prawn fishery where it 

constitutes about 50% of the shrimp catches. Probably mainly consumed locally. Restricted to the 

waters between New Guinea and Australia (sometimes divided into 2, eastern and western, 

subspecies). 


Metapenaeus dobsoni (Miers, 1878) 

En - Kadal shrimp; Fr - Crevette kadal; Sp - Camarón kadal. 

Maximum body length 13 cm (females) and 11.8 cm (males). Occurs mainly in low salinity lagoons 

and adjacent marine areas on muds to a depth of 37 m. Juveniles inhabit estuarine and backwaters. 

Caught mainly by trawls, seines, stake nets, and artisanal gear. Generally not very common in the 

area although reported to be quite abundant in New Guinea and appears in culture ponds in 

Thailand. Indo-West Pacific from India to the Philippines and New Guinea. 



anterior 

plate 


thelycum 



thelycum 


distolateral 

projection 


distolateral 

projection 


petasma (ventral view) basial spine of 

third leg (male)

Penaeidae 937 

Metapenaeus eboracensis Dall, 1957 

En - York shrimp; Fr - Crevette york; Sp - Camarón york. 

Maximum body length 11.6 cm (females) and 9.8 cm (males). On sandy or muddy bottoms in inshore 

waters, rivers and estuaries, to a depth of 45 m, usually between 10 and 20 m. Caught by trawls, seines, 

hand nets, and artisanal gear. Of minor commercial importance, due to its small size. Constitutes only 

about 5% of the prawn fishery catches in the Gulf of Papua and supports a small amateur fishery in 

northern Australia. Mainly consumed locally. Restricted to the waters between southern New Guinea 

and northern Australia. 


Metapenaeus elegans De Man, 1907 

En - Fine shrimp; Fr - Crevette élégante; Sp - Camarón fino. 

Maximum body length 11.8 cm (females) and 8.4 cm (males). Usually in estuaries, ponds, and inland 

lagoons with low salinity, but also found at sea to a depth of 55 m, on mud or sandy-mud bottoms. Caught 

mainly by traps, also by trawls, push nets, set nets, and artisanal gear. Reported to be of limited 

commercial importance in the area, but can be easily confused with the commercially important 

Metapenaeus ensis; seems to be rather common in markets of the Philippines and probably is more 

commonly marketed in other countries as well. Marketed mainly fresh for local consumption. Indo-West 

Pacific from Sri Lanka to the Philippines and Fiji. 


(from Motoh and Buri, 1984) 


thelycum 



thelycum 


distolateral 

projection 





Metapenaeus endeavouri (Schmitt, 1926) 

En - Endeavour shrimp; Fr - Crevette devo; Sp - Camarón devo. 

Maximum body length 17.5 cm (females) and 14 cm (males). On sandy or sandy-mud bottoms, 

found from the coastline to depths of 50 or 60 m. Juveniles generally associated with seagrass areas 

in shallow estuaries. Caught mainly at night by demersal otter trawls, sometimes also by beam 

trawls. Caught commercially in northern Australia, with a catch of about 2 400 t (together with 

Metapenaeus ensis), catches from western Australia not included, in the annual period of 1989/1990. 

Marketed mainly frozen, cooked or salted, sometimes used as bait; consumed locally and also 

exported. In 1995, the reported aquaculture production of this species (probably a misidentification 

of Metapenaeus anchistus) in the Philippines amounted to 1 295 t (FAO Aquaculture Production 

Statistics). Restricted to northern Australia and the Gulf of Papua. 


Metapenaeus insolitus Racek and Dall, 1965 

En - Emerald shrimp; Fr - Crevette émeraude; Sp - Camarón esmeralda. 

Maximum body length 12 cm (females) and 8 cm (males). Over muddy or sandy bottoms in inshore 

waters, including creeks and estuaries, to a depth of 35 m, usually less than 8 m. Mainly caught by 

hand nets in amateur fisheries and consumed locally. Moderately abundant in inshore habitats, but 

of limited commercial importance, due to its small size. Restricted to northern Australia. 

(after Racek and Dall, 1965) 



thelycum 



thelycum 

distolateral 

projection 



distolateral 

projection 



Penaeidae 939 

Metapenaeus lysianassa (De Man, 1888) 

En - Bird shrimp; Fr - Crevette oiseau; Sp - Camarón parancero. 

Maximum body length 9 cm (females) and 6.1 cm (males). On muddy bottom in inshore waters, to 

depths of about 28 m. Caught by stake nets, traps, set nets, push nets, seines, and trawls. Abundant 

in the western part of the area, but of secondary commercial importance, due to its small size. 

Marketed fresh, frozen, or dried and mainly for local consumption. Indo-West Pacific from India to 

Viet Nam, Malaysia, and Indonesia. 


Metapenaeus macleayi (Haswell, 1879) 

En - Eastern school shrimp; Fr - Crevette de maclay; Sp - Camarón maclayo. 

Maximum body length 17.5 cm (females) and 14.6 cm (males). In estuaries and inshore waters, to a 

depth of 55 m. Juveniles inhabit seagrass areas within estuaries. Caught by means of trawling (mainly), 

hauling, and seining; also obtained on the basis of small-scale aquaculture. Of commercial importance 

in eastern Australia, but its catch in Queensland was only about 100 t in the annual period of 1989/1990. 

Marketed cooked or uncooked (“green”), mainly for local consumption. Restricted to eastern Australia, 

from Tin Can Bay (Queensland) to Corner Inlet (Victoria). 





thelycum 



thelycum 

distolateral 

projection 






Metapenaeus papuensis Racek and Dall, 1965 

En - Papua shrimp; Fr - Crevette papou; Sp - Camarón papuense. 

Maximum body length 11.8 cm (females) and 8.6 cm (males). Found in estuaries and inshore waters 

to depths of about 60 m. Probably not very common and without commercial importance.Very similar 

to Metapenaeus elegans. Indo-West Pacific, reported from the Philippines, Thailand, New Guinea, 

and probably also found in the Bay of Bangal. 


Metapenaeus suluensis Racek and Dall, 1965 



distolateral 

projection 


thelycum 



En - Sulu shrimp. 

Maximum body length 12 cm (females) and 9.9 cm (males). Found from the coastline to depths of 

about 40 m. Probably not very common and without commercial importance. Very similar to 

Metapenaeus ensis. Indo-West Pacific, reported from the Philippines, Gulf of Thailand, and New 

Guinea. 



distolateral 

projection 



thelycum 





Penaeidae 941 

Metapenaeus tenuipes Kubo, 1949 

En - Stork shrimp; Fr - Crevette cigogne; Sp - Camarón cigueña. 

Maximum body length 9.5 cm (females) and 7.5 cm (males). Occurs from the coastline and brackish 

waters to a depth of 30 m. Caught by trawls, set nets, traps, seines, and artisanal gear. Commonly 

found in the western part of the area, but nowhere very abundant and mainly forms a bycatch in 

prawn fisheries. Marketed fresh or frozen, mainly consumed locally. Western Pacific from Thailand 

to Malaysia, Singapore, and Indonesia. 


Parapenaeopsis arafurica Racek and Dall, 1965 

anterior 

plate 


thelycum 

distolateral 

projection 

distomedian 

projection 

ventral view 

petasma 

distal part 

in lateral 

view 

En - Arafura shrimp; Fr - Crevette arafura; Sp - Camarón arafura. 

Maximum body length 14 cm (females) and 8 cm (males). Found over soft mud bottoms at depths 

from 5 to 30 m. Caught mainly as bycatch in trawls. Apparently not very common and of no 

commercial importance. Restricted to the area between New Guinea and northern Australia. 

female 


male


Parapenaeopsis cornuta (Kishinouye, 1900) 

En - Coral shrimp; Fr - Crevette corail; Sp - Camarón coral. 

Maximum body length 10 cm (females) and 8.5 cm (males), commonly between 5 and 8 cm. 

Generally inhabits river mouths and estuaries, but sometimes found at sea to depths of about 40 m, 

on bottom of sandy-mud or mud. Caught by trawls, seines, stake nets, and artisanal gear. Reported 

to be found occasionally in large quantities in Thailand and the Philippines. However, as this species 

can be easily confused with Parapenaeopsis maxillipedo, its abundance in the area remains 

uncertain. Indo-West Pacific from the western coast of India to Japan and northern Australia. 

anterior 

plate 

posterior 

plate 

hairs 

Parapenaeopsis coromandelica Alcock, 1906 

distolateral 

projection of 

petasma 


thelycum 



En - Coromandel shrimp; Fr - Crevette coromandel; Sp - Camarón coromandel. 

Maximum body length about 12 cm (males and females). Found in shallow waters to a depth of 

about 11 m, mainly on mud. Caught mainly by seine nets and shrimp gill nets, also by artisanal gear. 

Probably not very common in the Western Central Pacific, although reported to be moderately 

abundant at the northwestern coast of Malaysia and along the western coast of Thailand. Indo-West 

Pacific from southern India to the Gulf of Thailand, Indonesian Archipelago, and Borneo. 


posterior plates 

thelycum 

distolateral projections 


Penaeidae 943 

Parapenaeopsis gracillima Nobili, 1903 

En -Thinshrimp. 

Maximum body length about 7 cm for females, males smaller. Prefers sandy bottoms, occasionally 

also on mud, at depths from 30 to 60 m. Caught by trawlers. Probably not common and of no 

commercial importance. Only known from the Strait of Malacca and northern Borneo of Malaysia. 


Parapenaeopsis hungerfordi Alcock, 1905 



thelycum 



En - Dog shrimp; Fr - Crevette chien; Sp - Camarón perro. 

Maximum body length 10.4 cm (females) and 7.8 cm (males), commonly between 4 and 9.5 cm. Found 

on mud or sandy mud bottoms, from depths of 5 to 45 m, usually less than 25 m. Taken mainly by trawls. 

Reported to be one of the dominant species in shrimp catches off northwestern Malaysia, the western 

coast of Thailand, and off the southern coast of China. Its abundance in the area is uncertain. Marketed 

fresh or frozen for local consumption. Indo-West Pacific from the western coast of Thailand to Malaysia, 

Indonesia, and along the southern coasts of China, including Hong Kong. 



thelycum 


anterior 

plate 



Parapenaeopsis maxillipedo Alcock, 1906 

En - Torpedo shrimp; Fr - Crevette torpille; Sp - Camarón torpedo. 

Maximum body length 12.5 cm (females; perhaps to 15 cm) and 10 cm (males). Found at sea in 

shallow depths of less than 30 m on mud-banks, sometimes also on sandy-mud bottom. Caught by 

trawls, bottom gill nets, push nets, and shore seines. Reported to be commonly found in commercial 

catches in Malayan waters. However, as this species is often confused with Parapenaeopsis cornuta, 

its abundance in the area remains uncertain. Indo-West Pacific from the western coast of India to 

the Philippines and northern Australia. 

Parapenaeopsis sculptilis (Heller, 1862) 


posterior 

plate 

En - Rainbow shrimp; Fr - Crevette arc-en-ciel; Sp - Camarón arco iris. 

Maximum body length 17 cm (females) and 13 cm (males). Usually inhabits shallow waters from 

the coastline to depths of about 90 m, but mainly less than 40 m, on sand, mud, or mixed bottoms. 

Caught mainly by stake nets, seines, and trawls. Reported to be of some commercial importance 

in Malaysia and Singapore, but its abundance in these areas is uncertain. Relatively common in 

inshore commercial catches in northern Australia, but only of minor importance and mostly used 

as bait. Marketed mainly fresh or frozen for local consumption. Indo-West Pacific from Pakistan to 

the Philippines and northern Australia. 


hairs 

thelycum 



thelycum 




distolateral 

projection 


lobe 


Penaeidae 945 

Parapenaeopsis tenella (Bate, 1888) 

En - Smoothshell shrimp; Fr - Crevette glabre; Sp - Camarón liso. 

Maximum body length 7 cm (females) and 5 cm (males), commonly between 4 and 6 cm. Found 

from depths of 5.5 to 50 m, but mostly around 10 m, on muddy or sandy mud bottoms. Taken mainly 

by trawls. Apparently common in the western part of the area, but of very limited commercial 

importance due to its small size. Marketed fresh for local consumption. Indo-West Pacific from 

Pakistan to Japan and northern Australia. 

Parapenaeopsis uncta Alcock, 1905 



thelycum 


En - Uncta shrimp; Fr - Crevette uncta; Sp - Camarón uncta. 

Maximum body length 13 cm (females) and 8.3 cm (males). Found from depths of 40 to 90 m on clean 

sand, sometimes mixed with shell fragments.Taken by trawls.One of the relatively larger representatives 

of the genus, but nowhere abundant and only sporadically found in shrimp catches. Marketed mainly 

fresh or frozen for local consumption. Indo-West Pacific from Kuwait to India, Malaysia, and Indonesia. 





thelycum 

distolateral 

projection 




Parapenaeopsis venusta De Man, 1907 

En - Adonis shrimp; Fr - Crevette adonis; Sp - Camarón adonis. 

Maximum body length 4.5 cm. Found from depths of 11 to 44 m on bottoms of sand, shells, stones, and 

mud. Taken mainly by trawls. Probably not common and of no commercial importance, due to its small 

size. Western Pacific from the Gulf of Thailand to Malaysia, Indonesia, and Queensland (Australia). 


Parapenaeus fissuroides Crosnier, 1986 



thelycum 

distolateral 

projection 



En - False rose shrimp. 

Maximum body length 14 cm (females) and 11.7 cm (males), commonly between 7 and 11 cm. Found 

from depths of 65 to 908 m, mainly between 110 and 400 m, on bottoms of sand, mud, sandy mud, and 

soft mud. Taken by trawls. Of limited commercial importance and only sporadically taken as bycatch in 

deeper waters. It may have some economic potential with the development of a deep-sea fishery. 

Marketed mainly fresh for local consumption.Widely distributed from the eastern coast of Africa to Japan 

and Indonesia (populations in the Indian Ocean are sometimes considered to be 2 subspecies). Often 

confused with Parapenaeus fissurus and in the past mostly reported under this name. 


anterior 

plate 

thelycum 


lobe 


Penaeidae 947 

Parapenaeus longipes Alcock, 1905 

En -Flamingoshrimp;Fr -Crevetteflamand;Sp - Camarón flamenco. 

Maximum body length 11.5 cm (females) and 8.5 cm (males), commonly between 5 and 8 cm. 

Found usually between depths of 30 and 90 m, sometimes to a depth of 165 m. Taken mainly as 

bycatch in trawls. This species has the shallowest vertical distribution in the genus, but is nowhere 

abundant and of very limited commercial importance. Marketed mainly fresh for local consumption. 

Indo-West Pacific from eastern coast of Africa to Japan and New Guinea. 


Penaeopsis eduardoi Pérez Farfante, 1977 



thelycum 

distomedian lobe 

distolateral 

lobe 


En - Four-spined needle shrimp. 

Maximum body length 13 cm (females) and 12 cm (males), commonly between 8 and 11 cm. Found 

on sandy mud bottoms from depths of 289 to 570 m, usually deeper than 300 m. So far taken mainly 

during exploratory trawling operations, but sometimes found in large quantities and may therefore 

have some economic potential with the development of a deep-sea fishery. Western Pacific from 

Japan to the Timor Sea and Fiji. Can easily be confused with Penaeopsis rectacuta. 

(after Pérez Farfante, 1977) 


thelycum 

anterior 

plate 

ventral 

costa 



Penaeopsis rectacuta (Bate, 1881) 

En - Needle shrimp; Fr - Crevette aiguille; Sp - Camarón aguji. 

Maximum body length 13.5 cm (females) and 11 cm (males), commonly between 8 and 11 cm. Found 

on sandy mud bottom from depths of 174 to 410 m, usually deeper than 300 m. Mainly taken by 

experimental trawlers, but sometimes found in fair quantities and may therefore have some commercial 

potential once that a deep-sea fishery is developed in the area. Western Pacific from Taiwan Province 

of China to the Philippines, South China Sea, and Timor Sea. Often confused with Penaeopsis eduardoi. 


Trachypenaeus anchoralis (Bate, 1881) 

anterior 

plate 


thelycum 

ventral 

costa 


(after Pérez Farfante, 1979) 

En - Hardback shrimp; Fr -Crevetteos;Sp - Camarón huesudo. 

Maximum body length 10.4 cm (females) and 7 cm (males). Found on bottoms of mud to coral debris, 

from depths of 12.5 to 60 m. Taken mainly by trawls. Caught incidentally in the northern prawn fishery 

of Australia, but without much economic importance, due to its relatively small size. Generally 

believed to be restricted to northern Australia from Shark Bay (western Australia) and Keppel Bay 

(Queensland), but probably also occurs in southern Taiwan Province of China. 




thelycum 

distolateral 

projection 

distomedian 

projection 


Penaeidae 949 

Trachypenaeus gonospinifer Racek and Dall, 1965 

En - Northern rough shrimp; Fr - Crevette gambri nordique; Sp - Camarón fijador norteño. 

Maximum body length 8 cm (females) and 5 cm (males). Found over muddy bottom from depths of 

13 to 52 m. Taken as incidental catch by trawlers. Not particularly abundant and without commercial 

importance, due to its small size. Restricted to the waters between New Guinea and northern 

Australia. 

(after Grey, Dall, and Baker, 1983) 

Trachypenaeus granulosus (Haswell, 1879) 

En - Coarse shrimp; Fr - Crevette gambri grenue; Sp - Camarón fijador de granos. 

Maximum body length 9.5 cm (females) and 7.2 cm (males), commonly between 6 to 8 cm. Found 

over bottoms of mud, hard sand or rocks, from depths of 5 to 81 m. Taken as incidental catch, mainly 

at night by trawlers, also with artisanal gear. Of minor economic importance because of its small 

size and hard shell. Marketed mainly fresh for local consumption. Indo-West Pacific from Persian 

Gulf to Taiwan Province of China and northern Australia. 



thelycum 


distolateral 

projection 



Trachypenaeus longipes (Paulson, 1875) 

En - Longlegged rough shrimp. 

Maximum body length 10.5 cm (females) and 8 cm (males). Found from nearshore waters to depths 

of about 220 m, usually between 40 and 60 m. Taken mainly by trawls. This species is often confused 

with Trachypenaeus curvirostris (sometimes under the name T. asper), and therefore its actual 

distribution and occurrence in the area is unclear. Probably not very common and of limited or no 

commercial importance. Indo-West Pacific from the Red Sea to Japan and the Philippines. 

Trachypenaeus sedili Hall, 1961 


anterior 

plate 


thelycum 

distolateral 

projection 


distomedian 

projection 


En - Singapore rough shrimp. (the FAO names previously used for this species are now used for 

Trachypenaeus malaiana) 

Maximum body length 8.8 cm (females) and 5.1 cm (males), commonly between 6 and 8 cm. Found 

on mud or sand bottom, from nearshore waters to depths of about 45 m.Taken by trawls and artisanal 

gear. Probably not a common species in the area and without commercial importance. Indo-West 

Pacific from India (perhaps Somalia) to the Malay Peninsula and South China Sea. 



thelycum 


distolateral 

projection 


Penaeidae 951 

Trachypenaeus villaluzi Muthu and Motoh, 1979 

En - Philippines rough shrimp. 

Maximum body length 7.3 cm (females) and 5.3 cm (males). Caught by otter trawls at a depth of 

about 7 m, on mud bottom. Probably not common and without commercial importance. So far only 

known from the Philippines. 




thelycum 


distolateral 

projection 





Sicyoniidae SICYONIIDAE 

Rock shrimps 

Diagnostic characters: Body generally 

robust, with shell very hard, of “stony” 

appearance; abdomen often with deep 

grooves and numerous tubercles. Rostrum 

well developed and extending beyond eyes, 

always bearing more than 3 upper teeth (including 

those on carapace); base of eyestalk 

with styliform projection on inner surface, but 

without tubercle on inner border. Both upper 

and lower antennular flagella of similar 

grooves 

length, attached to tip of antennular peduncle. 

Carapace lacks both postorbital and 

postantennal spines, cervical groove indistinct 

or absent. Exopod present only on 

first maxilliped. All 5 pairs of legs well developed, 

fourth leg bearing a single well-developed 

arthrobranch (hidden beneath 

carapace). In males, endopod of second pair 

of pleopods (abdominal appendages) with 

3 

appendix masculina only. Third and fourth pleopods single-branched. Telson generally armed with a 

pair of fixed lateral spines. Colour: body colour varies from dark brown to reddish; often with distinct 

spots or colour markings on carapace and/or abdomen - such colour markings are specific and 

very useful in distinguishing the species. 

Habitat, biology, and fisheries: All members of this family are marine and can be found from shallow to 

deep waters (to depths of more than 400 m). They are all benthic and occur on both soft and hard bottoms. 

Their sizes are generally small, about 2 to 8 cm, but some species can reach a body length over 15 cm. 

The sexes are easily distinguished by the presence of a large copulatory organ (petasma) on the first pair 

of pleopods of males, while the females have the posterior thoracic sternites modified into a large sperm 

receptacle process (thelycum) which holds the spermatophores or sperm sacs (usually whitish or yellowish 

in colour) after mating. The shape of the petasma and thelycum is often specific and very useful for species 

identification. The eggs are small and numerous, and are released directly into the water and not retained 

on the female abdomen. The larvae are planktonic and have the nauplius stage. This family contains a 

single genus only and at present about 14 species are recorded from the Western Central Pacific. However, 

the taxonomic status and relationships of these species are generally unclear. Moreover, none of them are 

large enough or abundant enough to be of commercial importance in the area. Therefore, no identification 

key is provided here, and a species account is given for Sicyonia lancifera only, the most common 

representative of this family in the Western Central Pacific. 

rd and 4th 1 

2 

3 

4 

5 

pleopods 



Aristeidae: body not “stony” in appearance, abdomen without deep grooves or tubercles; either rostrum very 

short, armed with 1 or 2 upper teeth only, or upper antennular flagellum very short, not attached to tip of 

antennular peduncle; third and fourth pleopods divided into 2 branches; telson without fixed lateral spine. 

Penaeidae: body not “stony” in appearance, abdomen without deep grooves or tubercles; exopod present 

posterior to first maxilliped; third and fourth pleopods divided into 2 branches. 

upper 

antennular 

flagella 

very short 

1 

2 

3 4 5 

Aristeidae 

1 

2 

3 

4 

Penaeidae 

5

Sicyoniidae 953 

Solenoceridae: body not “stony” in appearance, abdomen without deep grooves or tubercles; carapace 

either with postorbital or postantennal spine; cervical groove distinct, extending to about dorsal carapace; 

third and fourth pleopods divided into 2 branches. 

Sergestidae: size small; rostrum very short; body strongly compressed laterally; shell soft; last 2 pairs of 

legs reduced or absent. 



4 

3 

5 

Solenoceridae 

Stenopodidae: third pincer extraordinary large and massive; males and females without large copulatory 

organ on first pair of pleopods or posterior thoracic sternites, respectively; females carry the eggs on the 

abdomen until hatching. 


expanded, overlapping posterior part of first pleuron and anterior part of third pleuron;males and females without 

large copulatory organ on first pair of pleopods or posterior thoracic sternites, respectively; females carry the 

eggs on the abdomen until hatching. 

2 

1 

1 

2 

Stenopodidae 

females 

carry 

eggs on 

abdomen 

3rd 3 

leg 

enlarged 



Sicyonia benthophila De Man, 1907 

Sicyonia bispinosa (De Haan, 1844) 

Sicyonia curvirostris Balss, 1914 

Sicyonia fallax De Man, 1907 

Sicyonia furcata Miers, 1878 

Sicyonia inflexa (Kubo, 1949) 

Sicyonia laevis Bate, 1881 

Sicyonia lancifera (Olivier, 1811) 

Sicyonia nebulosa Kubo, 1949 

Sicyonia ocellata Stimpson, 1860 

Sicyonia ommanneyi Hall, 1961 

Sicyonia parvula (De Haan, 1850) 

Sicyonia rectirostris De Man, 1907 

Sicyonia trispinosa De Man, 1907 

4 

5 

3rd 3 

maxilliped 

References 

Chan, T.Y. and H.P. Yu. 1985. On the rock shrimps of the family Sicyoniidae (Crustacea: Decapoda) from Taiwan, with 

description of one new species. Asian Mar. Biol., 2:93-106. 

1 

2 

3 

no 

pincer 

2 

1 

Sergestoidae 

5 

4 

females carry 


Caridea 

2 nd pleuron 

pear-shaped


Sicyonia lancifera (Olivier, 1811) 

Frequent synonyms / misidentifications: Sicyonia cristata (De Haan, 1844) / None. 

FAO names: En - Knight rock shrimp; Fr - Boucot chevalier; Sp - Camarón de piedra lanzón. 


Diagnostic characters: Body robust, with shell very hard, of “stony”appearance. 

Rostrum nearly straight, with 3 to 6 upper teeth and 1 to 3 apical teeth, lower border 

usually bearing 1 tooth only. Carapace armed with 3 to 5 large, crest-like postrostral 

teeth as well as a very strong hepatic spine. Abdomen heavily sculptured 

and with each pleura ending in 2 or 3 sharp spines. Pleopods with a single 

branch only. Colour: body brownish, with a complicated pattern of white stripes and 

black dots; ventral surface somewhat reddish brown; dorsal surface of first abdominal 

segment whitish, with a pair of large black spots; eyes light brown; 

antennal flagella and thoracic appendages covered with white and brown bands; tail 

fan with a thick white band near base. 

Size: Maximum body length 8 cm (females larger), commonly between 3 and 5 cm. 

Habitat, biology, and fisheries: Found on sandy-mud bottoms, at depths from 25 

to 350 m, usually less than 100 m. Probably burrows in sand during the daytime. 

When it comes out, often walks on the bottom with the abdomen strongly curved 

upward. Probably the most common species of the family in the area, but still few in 

numbers and only caught incidentally during prawn trawling operations. Without 

commercial value throughout its range because of its small size and low quantities. 



from Mozambique to Japan 


abdomen 

(dorsal view) 

(after Motoh and Buri, 

1984)

Stenopodidae 955 

Stenopodidae Infraorder STENOPODIDEA 

Family STENOPODIDAE 

Stenopodid shrimps 

Diagnostic characters: Usually 

small-sized, with a body length from 

1 to 6 cm. All 5 pairs of legs well developed, 

with first 3 pairs of legs forming 

a pincer, third pair huge and massive. 

Abdomen with posterior part of pleura 

covering anterior part of succeeding 

pleura. Males and females without 

large copulatory organ on first pair of 

pleopods (abdominal appendages) or 

posterior thoracic sternites, respectively. 

Females carry the eggs on the abdomen 


Habitat, biology, and fisheries: This infraorder 

contains a single family and 

about 60 species (divided into 2 families 

by some authors). Altogether, 7 genera 

and 19 species have been reported from 

the Western Central Pacific. All species 

are marine and benthic, and can be found 

Spongicola venusta 

from shallow coral reef areas to deep sea 

(after Holthuis, 1993) 

at depths of more than 800 m, with some 

species living in symbiosis with other invertebrates or fishes. Amongst 

these, the cleaner shrimps of the genus Stenopus and the venus shrimp 

Spongicola venusta are best known. Stenopus shrimps inhabit coral reefs 

and set up “cleaning stations” which are regularly visited by fishes which 

allow the shrimps to clean their wounds, skins, and mouths. Spongicola 

venusta lives in pairs inside the body of the deep-sea hextactinellid 

sponges (i.e. the Venus’ flower basket). They enter the body of the 

sponges when they are in the postlarvae stage. As they grow, their size 

becomes too large and thus they cannot escape from the sponges where 

both the male and female spend the rest of their life. 

Members of this infraorder are generally without any economic importance. 

Therefore, no species accounts are included here and no key is provided.For 

keys to the genera of this infraorder users can refer to L.B. Holthuis (1993), 

Hextactinellid sponges 

“The recent genera of the Caridean and Stenopodidean shrimps (Crus- 

(Venus' flower basket) 

tacea, Decapoda): with an appendix on the order Amphionidacea,C.H.J.M. (after Tan et al., 1995) 

Fransen and C. van Achterberg eds, Nationaal Natuurhistorisch Museum, 

Leiden.” Only the cleaner shrimps of 

the genus Stenopus may sporadically 

enter the aquarium trade and have 

some commercial value. In the Western 

Central Pacific, the most commonly 

found species is Stenopus 

hispidus (Olivier, 1811). A key to species 

of Stenopus isgiveninJ.W.Goy 

(1992, J. Nat. Hist., 26:79-102), and 

good colour photos of this genus are 

included in H. Debelius and H.A. 

Baensch (1994, Marine Atlas: The 

joint aquarium care of invertebrates 

and tropical marine fishes, published 

by Mergus). 

Stenopus hispidus



Sergestoidea: body strongly compressed laterally; shell 

soft; rostrum and last 2 pairs of legs reduced or absent; 

males with large copulatory organ on first abdominal 

appendage; eggs usually released directly into water, 

not retained by the female. 

Penaeoidea: first 3 pairs of legs forming a pincer, none 

of them particularly large; with large copulatory organ, 

on first pair of pleopods in males, and on posterior 

thoracic sternites in females; eggs released directly into 

water, not retained by the female. 

3 

Caridea: third leg without pincer; pleuron of second 

abdominal segment greatly expanded and overlapping 

those of first and third segments. 

rd maxilliped 

first 3 

legs with 

pincer 


1 

2 

3 

4 5 

Penaeoidea 

Engystenopus palmipes Alcock and Anderson, 1894 

Engystenopus spinulatus Holthuis, 1946 

Microprosthema scabricaudatum Richters, 1880 

Microprosthema validum Stimpson, 1860 

Odontozona ensifera (Danna, 1852) 

Odontozona sculpticaudata Holthuis, 1946 

Paraspongicola pusilla De Saint Laurent and Cleva, 1981 

Spongicola henshawi Rathbun, 1906 

Spongicola holthuisi De Saint Laurent and Cleva, 1981 

Spongicola inflata De Saint Laurent and Cleva, 1981 

Spongicola venusta De Haan, 1841 

? Spongicoloides japonica (Kubo, 1942) 

(after Liu, 1955) 

Stenopus chrysexanthus Goy, 1992 

Stenopus cyanoscelis Goy, 1984 

Stenopus devaneyi Goy and Randall, 1984 

Stenopus hispidus (Olivier, 1811) 

Stenopus pyronotus Goy and Davaney, 1980 

Stenopus tenuirostris De Man, 1888 

Stenopus zanzibaricus Bruce, 1976 

1 

2 

3 

no 

pincer 


Sergestoidea 

4 

5 

females carry 


Caridea 

2 nd pleuron 

expanded, 

pear-shaped

Infraorder Caridea 957 

Infraorder Caridea Infraorder CARIDEA 

Caridean shrimps 

Diagnostic characters: Very small to 

large-sized, with a body length from 0.5 to 

32 cm. All 5 pairs of legs well developed, the 

first 2 pairs with or without a pincer, but third 

leg never bearing a pincer. Second abdominal 

pleuron (lateral plate) greatly expanded, 

pear-shaped and overlapping 

posterior part of first pleuron as well as 

anterior part of third pleuron. Males and 

females without large copulatory organ on 

first pair of pleopods (abdominal appendages) 

or posterior thoracic sternites, respectively. 

Females carry the eggs on the abdomen 


3 

Habitat, biology, and fisheries: This large 

infraorder contains at least 2 517 species in 28 

families (the number of families in this infraorder 

is controversial among crustacean taxonomists). 

They can occur in all kinds of aquatic 

habitats such as high mountain streams (at 

altitudes of more than 2 500 m), lakes, caves, exopod 

underground waters, rivers, estuaries, littoral 

exopod 

appendix 

zones, beaches, bays, coral reefs, continental 

interna 

shelves, and the deep sea (at depths of at least 

appendix 

6 364 m, perhaps even 10 912 m for a red 

interna 

“shrimp” sighted by a bathyscaphe at Chal- 

endopod 

lenger Deep, near Guam). A few species inhabit 

the upper littoral zone and are able to endopod 

endure short periods of desiccation. Both ben- 

appendix 

male masculina 

female 

thic and/or pelagic (including epi- and bathypelagic) 

ways of life are found in members of 

second abdominal leg (pleopod) 

this infraorder, and a large number of marine 

species live in symbiosis with other invertebrates or fishes. The sexes are generally separated, but certain 

species, such as some Pandalus, commonly first undergo a male phase and later transform into females. 

The gonopores are situated at the bases of fifth or third leg in males and females, respectively. However, 

as many caridean shrimps are of very small size, with the thoracic sternum being narrow, sexing caridean 

shrimps by observing the position of gonopores is often difficult. A more simple way to distinguish the sexes 

of caridean shrimps is to determine the presence or absence of the so-called appendix masculina on the 

second pleopods (or abdominal appendages; see figure above). In females, the endopod of the second 

pair of pleopods bears an appendix interna only. In males, the endopod of the second pleopods usually 

has an appendix interna as well, but additionally bears an appendix masculina, which, when observed 

under magnification, is distinct, even in juveniles. Besides, a remarkable sexual dimorphism (e.g. the size 

of the second pincer, the curvature of the rostrum, the shape of the abdominal pleura, etc.) is present in 

many species, but such a dimorphism is often specific and not necessarily the same in different species. 

The females of caridean shrimps carry the eggs on the abdomen. Their larvae leave the eggs in relatively 

advanced stages (i.e. lacking the nauplius stage) and some even directly as juveniles. 

At present, 22 families of caridean shrimps are known to occur in the Western Central Pacific, but the 

exact number of species in the area is unclear. This is mainly due to fact that most of them are without 

any economic importance and therefore, have rarely been studied. However, recent extensive studies 

on carideans from the Philippines and adjacent areas have shown that 528 species are found in that 

region alone. Despite the large number of species, most caridean shrimps are small and do not occur in 

sufficient quantities to be actively fished, and/or live in very deep sea. Therefore, they are generally of 

no commercial importance in the Western Central Pacific. So far, only the giant river prawn Macrobrachium 

rosenbergii is both actively fished and extensively cultured in the area. It is mainly marketed 

live or fresh for local consumption, sometimes also exported. Several other fresh-water and coastal 

caridean shrimps in the area also have a relatively larger size or are easy to catch, and are likely to be 

used as food by natives in several countries. However, information on local use of caridean shrimps is 

mostly lacking and the exact identities of the species in question are often uncertain and/or confusing in 

literature. On the other hand, some deep-sea caridean shrimps, mostly belonging to the family Pandalidae 

rd leg without 

2 

pincer 

females carry eggs 

on abdomen 

nd pleuron 

pear-shaped


(particularly those of the genus Heterocarpus), are large in size and are often caught in great numbers 

during exploratory trawling operations. They are generally considered to have commercial potential with the 

development of a future deep-sea fishery in the area. Finally, it should be mentioned that some coral reef 

carideans (mainly members of the families Alpheidae, Gnathophyllidae, Hippolytidae, Hymenoceridae, Palaemonidae, 

and Rhynchocinetidae) have an attractive coloration and can be sporadically found in the aquarium 

trade, where those uncommon species often command a high price. Nevertheless, their supply are usually 

unstable and they are commercially much less important as marine coral fishes. Therefore, individual 

identification sheets are provided here for several selected species only, as representatives of the infraorder in 

the area. For keys to the families and genera of carideans, users may consult L.B. Holthuis (1993), “The recent 

genera of the Caridean and Stenopodidean shrimps (Crustacea, Decapoda): with an appendix on the order 

Amphionidacea, C.H.J.M. Fransen and C. van Achterberg (eds), Nationaal Natuurhistorisch Museum, Leiden.” 

For the identification of species, users may refer to the keys published by F.A. Chace, Jr (1976-1997: Smithson. 

Contrib. Zool. 222, 277, 381, 384, 397, 411, 432, 466, 543, 587), or are encouraged to send the sample(s) to 

the author of the present contribution. 


Sergestoidea: last 2 pairs of legs reduced or absent; 

anterior part of second abdominal pleuron not overlapping 

first abdominal pleuron; males with large copulatory 

organ on first pair of pleopods; eggs usually released 

directly into water, not retained by the female. 

Penaeoidea: third leg with pincer; anterior part of second 

abdominal pleuron not overlapping first abdominal 

pleuron; with large copulatory organ, on first pair of 

pleopods in males, and on posterior thoracic sternites in 

females; eggs released directly into water, not retained 

by the female. 

3 

Stenopodidea: third leg bearing a very large pincer; anterior 

part of second abdominal pleuron not overlapping 

first abdominal pleuron. 

rd 2 

3 

maxilliped 

1 

2 

3 

4 5 

2 

(after Liu, 1955) 

nd pleuron 

pincer 

not expanded 

anteriorly 

Penaeoidea 

List of families and species treated in this contribution 


Infraorder CARIDEA 

Superfamily PASIPHAEOIDEA 

PASIPHAEIDAE 

Superfamily OPLOPHORIDEA 

OPLOPHORIDAE 

Superfamily ATYOIDEA 

ATYIDAE 

Atyopsis moluccensis (De Haan, 1849) 

Atyopsis spinipes (Newport, 1847) 

Caridina weberi De Man, 1892 

1 

1 


Sergestoidea 

3 4 

2 

Stenopodidea 

5 

3 rd leg 

enlarged

Infraorder Caridea 959 

Superfamily BRESILIOIDEA 

BRESILIIDAE 

Superfamily NEMATOCARCINOIDEA 

EUGONATONOTIDAE 

NEMATOCARCINIDAE 

RHYNCHOCINETIDAE 

Rhynchocinetes durbanensis Gordon, 1936 

Superfamily PSALIDOPODOIDEA 

PSALIDOPODIDAE 

Superfamily STYLODACTYLOIDEA 

STYLODACTYLIDAE 

Superfamily CAMPYLONOTIDEA 

BATHYPALAEMONELLIDAE 

Superfamily PALAEMONOIDEA 

ANCHISTIOIDIDAE 

GNATHOPHYLLIDAE 

HYMENOCERIDAE 

Hymenocera picta Dana, 1852 

PALAEMONIDAE 

Exopalaemon styliferus (H. Milne Edwards, 1840) 

Exopalaemon vietnamicus Nguyên, 1992 

Leandrites indicus Holthuis, 1950 

Leptocarpus potamiscus (Kemp, 1917) 

Macrobrachium equidens (Dana, 1852) 

Macrobrachium lar (Fabricius, 1798) 

Macrobrachium mirabile (Kemp, 1917) 

Macrobrachium rosenbergii (De Man, 1879) 

Nematopalaemon tenuipes (Henderson, 1893) 

Palaemon concinnus Dana, 1852 

Superfamily ALPHEOIDEA 

ALPHEIDAE 

HIPPOLYTIDAE 

Lysmata amboinensis (De Man, 1888) 

Lysmata debelius Bruce, 1983 

Saron neglectus De Man, 1902 

OGYRIDIDAE 

Superfamily PROCESSOIDEA 

PROCESSIDAE 

Superfamily PANDALOIDEA 

PANDALIDAE 

Heterocarpus hayashii Crosnier, 1988 

Heterocarpus parvispina De Man, 1917 

Heterocarpus sibogae De Man, 1917 

THALASSOCARIDIDAE 

Superfamily CRANGONOIDEA 

CRANGONIDAE 

GLYPHOCRANGONIDAE


Atyidae ATYIDAE 

Atyopsis moluccensis (De Haan, 1849) 

En - Moluccas brush shrimp. 

Maximum total length 7.7 cm (occasionally 8.6 cm); males larger. Inhabits upper or middle parts of 

fast-flowing streams. Probably reproduce in brackish water and juveniles are tolerant of salt water. 

Abundance of this shrimp in the area is uncertain. Atyid shrimps are reported to be used as food or 

fertilizer in many parts of the area. Although the size of this species is large for atyids, its economic 

importance is probably still minor. Atyids are mainly marketed fresh or dry for local consumption. 

Recently, live specimens, probably originating from Indonesia, have been seen in the aquarium 

trade, for which this shrimp is a suitable candidate, due to the attractive bandings on the body and 

since it can easily be kept in captivity. Indo-West Pacific, and known with certainty from Sri Lanka 

to Thailand, Malaysia, Indonesia, and perhaps the Philippines. Often confused with Atyopsis 

spinipes. 

2 

with tufts 

of hairs 

1 

3 

Atyopsis spinipes (Newport, 1847) 

4 

5 

(adapted from Chace, 1983) 

En - Soldier brush shrimp; Fr - Saltarelle soldat; Sp - Camarón soldado. 

Maximum total length 7.1 cm, commonly between 4 and 5 cm (females larger). Adults mainly occcur 

in fresh water, in upper or middle parts of fast-flowing streams. Reproduce in brackish water. 

Abundance of this shrimp in the area is uncertain. Although atyid shrimps are reported to be used 

as food or fertilizer in many parts of the area and this species is relatively large amongst the atyids, 

its economic importance is probably only minor. Atyids are mainly marketed fresh or dry for local 

consumption. This shrimp has recently been introduced to the aquarium trade (place of origin 

probably Indonesia), because of the attractive bandings on the body and as it can easily be kept in 

captivity. Western Pacific, and known with certainty from Ryukyus, Taiwan Province of China, the 

Philippines, Lesser Sunda Islands, Palau, Fiji, and Samoa. 

1 

2 

3 

4 

with tufts 

of hairs (adapted from Chace, 1983) 

5

Atyidae/Hippolytidae 961 

Caridina weberi De Man, 1892 

En - Pugnose caridina; Fr - Saltarelle nez-camus; Sp - Camarón ñata. 

Maximum total length about 3 cm, commonly between 1.5 and 2 cm (females usually larger).Inhabits 

mainly in middle or lower parts of rivers, often around heavy vegetation. Reproduce in brackish 

water and juveniles are tolerant of salt water. Abundance of this shrimp in the area is uncertain 

although it has been reported to be of some economic value in Indonesia. Considering their very 

small size, all species of Caridina probably have very limited, if any, commercial importance. 

Indo-West Pacific from India to Japan and Polynesia. 

with tufts 

of hairs 

1 

2 

3 4 5 

(adapted from Holthuis, 1993) 

Atyidae/Hippolytidae 

Hippolytidae HIPPOLYTIDAE 

Lysmata amboinensis (De Man, 1888) 

En - Common cleaner shrimp. 

Body length around 5 cm. Shallow marine reef areas. Well known for its fish cleaning behaviour and 

sometimes encountered in large groups. Popular and often seen in the marine aquarium trade where 

it is sold at moderate prices. Nevertheless, its unstable supply suggests that this shrimp is not 

abundant in its natural habitats. Widely distributed in the Indo-West Pacific from Kenya to Japan, 

French Polynesia, and Hawaii. 

2 

1 

4 

3 

5 

(adapted from Holthuis, 1947)


Lysmata debelius Bruce, 1983 

En -Cardinalshrimp. 

Body length around 4 to 5 cm. Marine reef areas in depths from 10 to 28 m. Normally found in pairs 

and behave as fish cleaners. A popular shrimp in the marine aquarium trade and sold at somewhat 

higher prices than most of the other marine aquarium shrimps. However, its supply is not large and 

unstable, indicating that this shrimp is not abundant in its natural habitats. Known with certainty from 

the Philippines, Indonesia, and Sri Lanka. 

2 

3 

subdivided into 

many segments 

Saron neglectus De Man, 1902 

1 

4 

5 

(adapted from Bruce, 1983) 

En - Spotted marbled shrimp. 

Body length around 2 to 4 cm. Shallow marine reef areas. Nocturnal and usually hiding under rocks 

and caves, no fish cleaning behaviour reported. Males with first leg greatly enlarged. Probably the 

most common species of the genus seen in the marine aquarium trade (live specimens originated 

probably mainly from Indonesia). Sold at moderate prices, due to its unstable supply. Widely 

distributed in the Indo-West Pacific from Madagascar to the Red Sea, Japan, and New Caledonia. 

Can be easy confused with Saron marmoratus (Olivier, 1811). 

1 

2 

3 4 

5 


Hymenoceridae 963 

Hymenoceridae HYMENOCERIDAE 

Hymenocera picta Dana, 1852 

En - Painted harlequin shrimp. 

Body length around 2 to 5 cm. Shallow marine reef areas in depths from 1 to 20 m. Usually live in 

pairs and reported to be strongly territorial. No fish cleaning behavior observed but reported to kill 

and feed on starfishes, at least under aquarium conditions. A popular shrimp in the aquarium trade 

because of its bizarre looking and amazing coloration. Sold at moderately high prices due to its rare 

supply, indicating that this shrimp is not commonly found in its natural habitats. Widely distributed 

in the Indo-West Pacific from eastern Africa to Japan, Hawaii, and French Polynesia. 

2 

1 

3 4 

5


Palaemonidae PALAEMONIDAE 

Macrobrachium rosenbergii (De Man, 1879) PRF 

Frequent synonyms / misidentifications: Macrobrachium rosenbergii dacqueti (Sunier, 1925) / 

Macrobrachium carcinus (Linnaeus, 1758). 

FAO names: En -Giantriverprawn;Fr - Bouqet géant; Sp - Camarón gigante. 

1 

3 

2 

Diagnostic characters: Rostrum long, well extending beyond antennal scale; forming a high basal 

crest above the eye; armed with 11 to 14 upper teeth (including those on carapace and with distal teeth 

more widely spaced) and 8 to 14 lower teeth. Hepatic spine situated distinctly below antennal spine. 

Second legs very large, robust and of same size, with carpus longer than merus; in adult males, 

entire second leg densely covered with spines and sharp tubercles, cutting edges of fingers bearing 

only 1 or 2 large basal teeth and without rows of tubercles on either side, movable finger very hairy 

except at tip, carpus shorter than pincer. Telson tapering posteriorly, with tip exceeding posterolateral 

spines. Colour: body generally dark green to greyish blue, with longitudinal or irregular streaks of darker 

and lighter colour, hinges of abdominal segments often orange; eyes dark brown; antennal flagella dark 

blue to greyish; large pincer bluish to dark blue; eggs yellowish; youngs and berried females often with 

some longitudinal golden strips on the sides of body. 

Size: The largest known caridean shrimp, maximum body length 34 cm (females) and 26 cm (males), 

commonly between 10 and 20 cm. 

Habitat, biology, and fisheries: Inhabits mainly estuarine areas and rivers but sometimes also found at 

sea; requires brackish water for spawning and nursing up to postlarval stage, while juveniles are mainly 

found in fresh-water zones. An omnivorous, very large and common fresh-water shrimp that is extensively 

caught in the area. Taken by bamboo barriers, fish corrals, traps, set nets, cast nets, hook-and-line, and 

artisanal gear. Big catches are often linked to heavy rains. Also an important candidate for fresh-water 

aquaculture in many countries. In 1995, the harvest of this species reported from the area amounted to 

9 732 t from capture fishery in Indonesia and Thailand, and 5 040 t from aquaculture in Thailand. Marketed 

live, fresh or frozen, mainly for 

local consumption but sometimes 

also for export. 


from Pakistan to Viet Nam, 

the Philippines, New Guinea, 

and northern Australia (the 

western form from India to Malaysia 

is sometimes treated as 

a different subspecies). This 

shrimp has been introduced to 

many parts of the world for use 

in aquaculture. 

4 

5

Palaemonidae 965 

Exopalaemon styliferus (H. Milne Edwards, 1840) 

En - Roshna prawn; Fr - Bouqet rosna; Sp - Camarón rosna. 

Maximum total length for males 9 cm; egg-bearing females 6.8 to 8.6 cm. Inhabits shallow coastal 

waters, brackish or marine, occasionally also in fresh water. A small species, abundance in the area 

uncertain, probably without commercial importance and only caught incidentally in fisheries for other 

shrimps. Indo-West Pacific from Pakistan to Thailand and Borneo. 

6-10 

lower 

teeth 

2 

1 

(after Holthuis and Miguel, 1984) 

Exopalaemon vietnamicus Nguyên, 1992 

3 

4 

5 

En - Vietnamese crest prawn. 

Maximum body length 7.7 cm; egg-bearing females more than 4 cm body length. Inhabits estuaries 

and shallow coastal mud flat areas near river mouths. Caught by conical set nets and push nets. 

Often occurs in large quantities in coastal rice shrimp farming areas and semi-extensive shrimp 

culture ponds. Sometimes too abundant, becoming a food competitor of cultured penaeid prawns. 

In certain areas it constitutes 40 to 50% of the total shrimp harvest at the end of the rainy season. 

Marketed dried or fresh and usually mixed with other small penaeids; an important food source for 

local consumption. Restricted to southeastern Viet Nam near Ho Chi Min City. 

12-15 

lower 

teeth 

5-8 upper teeth 

1 

rostral crest 

3 

2 

4 

5 

(after Nguyên, 1992)


Leandrites indicus Holthuis, 1950 

En - Indian small prawn. 

Maximum total length about 3 cm. Inhabits brackish water in mangrove areas. A bycatch of fisheries 

for Acetes species, caught by conical set nets and push nets. In certain parts of Viet Nam, this small 

species is very abundant and constitutes a good food source for local consumption. Sometimes also 

enters penaeid culture ponds. Marketed fresh and mixed with Acetes species, also used in the 

processing of native shrimp paste. Western Pacific, and so far only recorded from Makasar (Celebes) 

and Viet Nam. 



spine 

1 

2 

3 

4 

5 

(after Nguyên, 1992) 

Leptocarpus potamiscus (Kemp, 1917) 

En - Bombay prawn; Fr - Bouqet bombay; Sp - Camarón de Bombay. 

Maximum total length about 6 cm (females) and 4.5 cm (males). Inhabits fresh to brackish water. In 

Viet Nam, this shrimp is reported to occur in great quantities in the irrigation ditches designed for 

culture of Macrobrachium rosenbergii and is a valuable species for local fisheries. Indo-West Pacific 

from India to southern China and Indonesia. 


1 

2 

3 

5 

4 

(after Nguyên, 1992) 



Palaemonidae 967 

Macrobrachium equidens (Dana, 1852) 

En -Roughriverprawn;Fr - Bouqet chagrin; Sp - Camarón lija. 

Maximum total length 9.8 cm (males usually larger). Inhabits lower parts of streams, river mouths, 

estuaries, and brackish waters of high salinity; rarely found in pure fresh water but often in sea water 

(near river mouths) to a depth of at least 30 m. Reproduce in brackish and sea water, larvae have 

about 11 stages and transform into postlarvae in 43 days. A common species in the area but nowhere 

abundant. Similarly to other small to medium-sized species of the genus, it is frequently found in 

mixed catches of fresh-water shrimps. Since this shrimp also occurs in pure sea water, it is often 

caught incidentally in penaeid fisheries. Marketed mainly live or fresh, generally sold at low prices 

and consumed locally. Widely distributed in the Indo-West Pacific from eastern Africa to the Ryukyu 

Islands, New Caledonia, and Solomon Islands, also introduced to Nigeria. 

2 

1 

3 


Macrobrachium lar (Fabricius, 1798) 

4 

hepatic spine 

5 

(after Holthuis and Miguel, 1984) 

En - Monkey river prawn; Fr - Bouqet singe; Sp - Camarón mono. 

Maximum total length 18.1 cm (males usually much larger). Adults occur in fresh water, mainly in 

upper and middle parts of rivers; able to endure short time of desiccation and occasionally observed 

to crawl on land during heavy rainfall. Migrate to the river mouth or estuary for reproduction, with 

juvenile stages in brackish and sometimes also in salt water. Fished throughout its range. Caught 

by traps and artisanal gear. A large species, but nowhere very abundant and marketed mainly locally, 

live or fresh. In the Philippines, this shrimp is not always available in the fish markets, sold at about 

half the price of Macrobrachium rosenbergii. Aquaculture experiments have been carried out by 

many countries but were so far unsuccessful, due to the very long larval stage in this species (at 

least 100 days). Widely distributed in the Indo-West Pacific, from eastern Africa to Ryukyu Islands 

and the Marquesas Islands; probably not indigenous (i.e. introduced) in Hawaii. 


3 1 4 

5 

hepatic spine 

2 

(after Holthuis, 1993)


Macrobrachium mirabile (Kemp, 1917) 

En - Shortleg river prawn; Fr - Bouqet tipattes; Sp - Camarón patojo. 

Maximum total length 6 cm (females) and 4 cm (males). Inhabits fresh and brackish water. Similarly 

to the other small species of the genus, its fishery in the area is unclear, probably caught incidentally 

with other fresh-water shrimps and used as food whenever available. Indo-West Pacific, and 

recorded from eastern India, Bangladesh, Myanmar, Thailand, Malaysia, and Borneo. 

rostral 

crest 

2 


1 

hepatic spine 

(after Kemp, 1917) 

Nematopalaemon tenuipes (Henderson, 1893) 

3 

4 

5 

En - Spider prawn; Fr - Bouqet araignée; Sp - Camarón araña. 

Maximum total length 8 cm. Inhabits shallow coastal waters to depths of about 20 m, and also found 

in estuarine and brackish waters. Abundance of this species in the area is unclear, but it has been 

reported to be used as food in the Philippines. Marketed dried or salted for local consumption. Widely 

distributed in the Indo-West Pacific from eastern Africa to Taiwan Province of China and the 

Philippines, probably also from New Zealand. 

3 

2 

1 

4 

5

Palaemonidae/Pandalidae 969 

Palaemon concinnus Dana, 1852 

En - Mangrove prawn; Fr - Bouqet mangrove; Sp - Camarón de manglar. 

Maximum total length 7 cm (females), smallest egg-bearing females 4.9 cm. Restricted to brackish 

water in the lower part of rivers, rarely found in pure fresh or sea water. In Viet Nam, this shrimp 

forms a bycatch of fisheries for Macrobrachium species and is offered for sale in local markets. 

Probably also used as food in the Philippines. Widely distributed in the Indo-West Pacific, from 

eastern Africa to Taiwan Province of China and Polynesia. 

unarmed 



spine 

2 

(after Ngugên, 1992) 

Palaemonidae/Pandalidae PANDALIDAE 

Heterocarpus hayashii Crosnier, 1988 

1 

3 

4 

5 

En - Japanese nylon shrimp. 

Maximum body length about 11 cm (females larger), commonly between 6 and 10 cm. Found on 

bottoms of sand and mud, at depths from 150 to 625 m, usually around 200 m, or deeper. So far 

not fished commercially but sometimes caught in large quantities on the basis of exploratory 

deep-water trawling. The moderate size of this shrimp and the fact it is found in not very deep water 

suggests that it may have commercial potential with the development of a deep-sea fishery in the 

area. Western Pacific, reported from Japan, Taiwan Province of China, the Philippines, Australia, 

New Caledonia, Chesterfield Islands, and Hawaii. Often confused with Heterocarpus ensifer A.Milne 

Edwards, 1881, H. parvispina, andH. sibogae. 

low ridge 

II 

III 

IV 

(adapted from Chace, 1985)) 

shorter than spine 

of 3 rd somite


Heterocarpus parvispina De Man, 1917 

En - Short-spined nylon shrimp. 


sandy-mud bottoms, at depths from 230 to 815 m, mostly less than 600 m. Not fished commercially 

at present. Occasionally caught in large quantities on the basis of exploratory deep-water trawling 

(more abundant from 350 m depth downward) and may have commercial potential with the 

development of deep-sea fisheries in the area. Western and southern Pacific from Taiwan Province 

of China to Indonesia, Australia, and French Polynesia. Often confused with Heterocarpus ensifer 

A. Milne Edwards, 1881, H. hayashii, andH. sibogae. 

not ridged 

Heterocarpus sibogae De Man, 1917 

II 

III 

IV 


En - Mino nylon shrimp; Fr - Crevette nylon mino; Sp - Camarón nailón mino. 


bottoms of sand and mud from depths of about 150 to 950 m. Although at present not commercially 

fished, it is the most common caridean shrimp caught during exploratory deep-water trawling 

operations in the area. Often caught in large quantities from moderate depths (about 200 m 

downward) in the area (e.g. the Philippines, Indonesia, Australia, New Caledonia, Fiji, Vanuatu, 

Samoa, Tonga, French Polynesia, and probably also Palau) and therefore has a very high commercial 

potential. Widely distributed in the Indo-West Pacific from Madagascar to Japan and French 

Polynesia. Often confused with Heterocarpus ensifer A. Milne Edwards, 1881, H. hayashii, andH. 

parvispina. 

II 

III 

IV 


very 

short 

as long as spine 

of 3 rd somite

Rhynchocinetidae 971 

Rhynchocinetidae RHYNCHOCINETIDAE 

Rhynchocinetes durbanensis Gordon, 1936 

En - Striped hinge-beak shrimp. 

Body length around 3 to 4 cm. Shallow marine reef areas. Gregarious and usually hiding under rocks 

and caves, also active during the daytime (afternoon); no fish cleaning behavior reported. Probably 

the most common shrimp in the area seen in the marine aquarium trade (with live specimens 

originating from Sri Lanka, the Philippines, and Indonesia), where it is regularly offered for sale. 

Very popular for its attractive coloration and as it easily adapts to captivity; sold at inexpensive prices. 

Widely distributed in the Indo-West Pacific from the eastern coast of South Africa to the Ryukyu 

Islands and Indonesia. Previously often confused with Rhynchocinetes uritai Kubo, 1942. 

rostrum movable 



LOBSTERS 

by T.Y. Chan

974 Lobsters 


body length 

antenna 

rostrum 

length of 

carapace 

length of abdomen 

(“tail”) 

antennule 

uropod 

rostrum 

abdominal 

segments 

general shape (dorsal view) of a true lobster (Metanephros spp.) 

supraorbital spine 

postrostral 

post-supraorbital spine 

spines 

gastric tubercle 

postrostral carina 

postorbital margin 

orbit 

subdorsal carina 


hepatic 

postcervical 

spine 


groove cervical 

groove 


V 

I 

II 

III 

IV 

VI 

leg 1 

median carina 

telson 

abdominal 

sculpture 

tail fan 

carapace of a true lobster (lateral view) 

first 3 legs 

with pincers 

(first pincer 

enlarged) 

leg 2 

leg 3 

leg 5 

leg 4 

submarginal posterior 

groove 

marginal posterior 

ridge 

longitudinal 

ridges


antennular 

flagella 

antennular 

peduncle 

body 

length 

length of 

carapace 

length of 

abdomen 

(“tail”) 

frontal 

horn 

eye 

antennular 

plate 

abdominal 

segments 

uropod 

telson 

general shape (dorsal view) of a spiny lobster 

(Panulirus spp.: no rostrum, no pincers) 

simple dactylus (Panulirus) 

false pincer (Justitia) 

I 

II 

III 

IV 

V 

VI 

leg 1 

transverse 

grooves 

leg 5 

antennal 

peduncle 

leg 2 

leg 3 

tail 

fan 

leg 4 

antennular 

peduncle 

principal 

spines 

frontal 

horn 

abdominal 

appendages 

(pleopods) 

types of terminal segments of legs 

antennular plate 

pleura of 2 nd and 

3 rd segments 

I 

II 

antennal 

peduncle 

antennular somite of a spiny lobster 

(left antenna and eye omitted) 

articulated 

parts non-articulated 

parts 

III 

uropod 

VI 

IV 

telson 

tail (abdomen) in lateral view 

true pincer (Metanephrops) 

false pincer (Enoplometopus) 

stridulating 

organ 

V 

eye

976 Lobsters 


Lobsters are generally large-sized crustaceans with a body length (measured dorsally from the orbital 

margin to the end of the tail, excluding the rostrum and any of the appendages) from a few to more than 

60 cm. Like shrimps and prawns, lobsters have a well-developed and extended abdomen or “tail”.In addition 

to their usually thicker shell, lobsters generally differ from shrimps and prawns by having the body more 

dorsoventrally depressed (particularly at posterior abdominal segments), the pleopods (i.e. legs of the 

abdomen) less developed, the thoracic sternum wide and distinct, the first abdominal pleuron considerably 

more reduced than the posterior pleura, and the posterior margin of the telson usually broadly convex or 

truncate. Certain other lobster-like crustaceans, such as the “squat lobsters” (Galatheoidea), “mud lobsters” 

(Thalassinidea: Thalassinidae), “mud shrimps” (Thalassinidea: Upogebiidae), and “ghost shrimps” 

(Thalassinidea: Callianassidae) are taxonomically not “true” lobsters. These groups are of no or only very 

minor importance to fisheries in the Western Central Pacific and therefore not included in this field guide. 

lobsters 

shrimps and prawns 


pleuron 

reduced 


pleuron 

well developed 


pleopods 

short 

pleopods 

long 

conspicuous morphological differences between lobsters and shrimps 

body dorsoventrally 

depressed 


broadly convex 

body laterally 

compressed 


pointed

General Remarks/Guide to Families 977 

In the Western Central Pacific, the lobsters are represented by 8 families, 22 genera and about 89 species. 

Although the production of most of the species is not very large, many of them are of moderate importance 

to fisheries because of their large size and high market value. 

Most spiny and slipper lobsters (Palinuridae and Scyllaridae, respectively), and the families Synaxidae and 

Enoplometopidae, inhabit shallow waters of rocky and reef bottoms or bottoms with coarse sediments. 

Among these, the genera Panulirus, Scyllarides and Parribacus are actively fished throughout the area 

because of their large size.On the other hand, juveniles of Panulirus species and the rare Enoplometopidae 

and Synaxidae are often highly valued in the aquarium trade. These shallow-water lobsters are mainly 

taken by hand while diving or by spears during night fishing, but sometimes also taken by tangle nets, 

lobster pots, or traps. 

Most Nephropidae, Thaumastochelidae, Glypheidae, Polychelidae, a few Palinuridae, and Scyllaridae are 

found in deeper waters on soft bottoms with sand and/or mud. They are usually caught by trawlers. At 

present, only the two shallow-water genera Ibacus and Thenus (both belonging to the family Scyllaridae) 

are landed in larger amounts and are of moderate commercial importance. However, exploratory fishing 

indicates that several deep-water species of the genera Metanephrops, Nephropsis, Acanthacaris, 

Linuparus and Puerulus (the first 3 genera belonging to the family Nephropidae, while the latter 2 belong 

to the family Palinuridae) are large and occasionally abundant and may eventually be of commercial 

interest. Species of the other 3 families (Thaumastochelidae, Glypheidae, and Polychelidae) are generally 

rare and of no interest to fisheries in the area, and are not treated here in separate family or species 

accounts. 

Lobsters in the Western Central Pacific are generally locally consumed and marketed fresh or live.In certain 

countries, such as the Philippines and Indonesia, a fair amount of lobsters are exported (live, fresh, cooked 

whole, or tailed). 

General Remarks/Guide GUIDE to Families TO FAMILIES OCCURRING IN THE AREA 

NEPHROPIDAE Page 982 

True lobsters and lobsterettes 

Body tubular, surfaces almost naked or 

covered with thick fur; rostrum well developed; 

antennae long and thread-like; antennal 

scale, if present, with inner margin unarmed 

and curved; first 3 pairs of legs with true 

pincers, first pair much larger than others; 

abdominal pleura ending in acute ventral 

tooth; tail fan entirely hardened, telson armed 

with fixed spines and with posterior margin 

broadly convex. 

THAUMASTOCHELIDAE 

Pincer lobsters 

Body slightly depressed dorsoventrally; eyes 

strongly reduced, cornea lacking pigmentation; 

rostrum well developed; antennae long and 

thread-like, antennal scale bearing several 

large teeth along inner margin; first 3 pairs of 

legs (occasionally also fifth legs) with true 

pincers, first pair large but very unequal; 

abdominal pleura short, quadrangular and 

without large ventral tooth; tail fan entirely 

hardened, telson quadrangular and unarmed. 

Only 2 deep-water species known from the 

area, very rare and of no interest to fisheries. 

1 st pincer 

enlarged 

strong rostrum 

1 st 

pincers 

very 

unequal 

1 

2 

first 3 legs with 

true pincers 

1 

2 

first 3 legs with 

true pincers 

3 

3 

4 

4 

5 

5

978 Lobsters 

ENOPLOMETOPIDAE Page 995 

Reef lobsters 

Body tubular and distributed with tufts of 

long stiff hairs; carapace with a 

well-developed rostrum; antennae long and 

thread-like, antennal scale with inner 

margin unarmed and curved; first pair of 

legs as large pincer, second and third legs 

slender and forming false pincers; 

abdominal pleura more or less rounded and 

sometimes ending in a strong ventral tooth; 

tail fan entirely hardened, telson bearing 

movable spines and with posterior margin 

broadly convex. 

POLYCHELIDAE 

Blind lobsters 

Eyes small, cornea lacking pigmentation; 

carapace box-like, rostrum absent or 

rudimentary; antennae thread-like, shorter 

than body; first 4 or all legs with true pincers, 

first pair long and slender; tail fan entirely 

hardened, telson pointed. All species found 

in very deep waters and of no interest to 

fisheries in the area. 

GLYPHEIDAE 

Fenix lobsters 

Body somewhat flattened dorsoventrally; 

eyes large and black, inserted on a median 

elevation of cephalon; carapace with a 

well-developed rostrum; antennae long and 

thread-like; first 2 legs forming false pincers, 

first pair very strong; uropods of tail fan 

entirely hardened. A single deep-water 

species, rare and of no commercial 

importance. 

1 

1 st pincer 

enlarged 

1 st leg with 

large pincer 

1 

antenna shorter 

than body 

1 

2 

3 

2 nd and 3 rd legs 

with false pincers 

2 

4 

small pincers 

false pincers 

2 3 4 

simple dactylus 

3 

4 

5 

5 

tufts of long 

stiff hairs 

5 

telson pointed

General Remarks/Guide to Families 979 

SYNAXIDAE Page 1001 

antenna 

Furry lobsters 

short 

Body somewhat flattened dorsoventrally and 

very hairy, without enlarged spines; carapace 

laterally angular, with a broad and flat triangular 

or rounded rostrum; antennae whip-like, shorter 

than carapace; legs without pincers, first pair 

much more robust than the others; posterior half 

of tail fan soft and flexible. 

2 

PALINURIDAE Page 1005 

Spiny lobsters, langoustes 

Body tubular or slightly flattened dorsoventrally; 

hairs, if present, few and scattered; rostrum 

absent or reduced to a small spine; carapace 

subcylindrical or prismatic, laterally rounded or 

straight, surface spiny and with a pair of large 

frontal horns above eyes; antennae very long 

and rather thick, whip-like or spear-like; legs 

without true pincers and first pair (except in 

Justitia) not or only slightly longer than the 

following legs, but often somewhat more robust; 

posterior half of tail fan soft and flexible. 

SCYLLARIDAE Page 1028 

Slipper lobsters 

Body strongly flattened dorsoventrally; 

carapace depressed and laterally angular; 

rostrum absent or minute; eyes enclosed by 

distinct orbits and without large frontal horns; 

antennae plate-like; legs without pincers, none 

of them enlarged; posterior half of tail fan soft 

and flexible. 

1 

3 

1 

no rostrum 

antenna 

plate-like 

distinct orbits 

2 

4 

all legs without 

pincers 

all legs without true pincers 

frontal 

horns 

1 

thick, long 

antenna 

2 

5 

3 4 5 

3 

4 

all legs without pincers 

body flat 

5

980 Lobsters 

List of FamiliesLIST and Species OF FAMILIES AND SPECIES OCCURRING IN THE AREA 


NEPHROPIDAE 

Acanthacaris tenuimana Bate, 1888 

Metanephrops andamanicus (Wood-Mason, 1891) 

Metanephrops arafurensis ((De Man, 1905) 

Metanephrops australiensis (Bruce, 1966) 

Metanephrops neptunus (Bruce, 1965) 

Metanephrops sibogae (De Man, 1916) 

Metanephrops sinensis (Bruce, 1966) 

Metanephrops thomsoni (Bate, 1888) 

Metanephrops velutinus Chan and Yu, 1991 

Nephropsis acanthura Macpherson, 1990 

Nephropsis ensirostris Alcock, 1901 

Nephropsis holthuisi Macpherson, 1993 

Nephropsis serrata Macpherson, 1993 

Nephropsis stewarti Wood-Mason, 1873 

Nephropsis suhmi Bate, 1888 

Nephropsis sulcata Macpherson, 1990 

THAUMASTOCHELIDAE 

Thaumastocheles japonicus Calman, 1913 

Thaumastochelopsis wardi Bruce, 1988 

ENOPLOMETOPIDAE 

Enoplometopus chacei Kensley and Child, 1986 

Enoplometopus daumi Holthuis, 1983 

Enoplometopus debelius Holthuis, 1983 

Enoplometopus gracilipes (De Saint Laurent, 1988) 

Enoplometopus holthuisi Gordon, 1968 

Enoplometopus occidentalis (Randall, 1840) 

POLYCHELIDAE 

Polycheles baccatus Bate, 1878 

Polycheles carpenteri (Alcock, 1894) 

Polycheles enthrix (Bate, 1878) 

Polycheles gracilis (Bate, 1888) 

Polycheles laevis (Bate, 1878) 

Polycheles obscurus (Bate, 1878) 

Polycheles typhlops Heller, 1862 

Stereomastis andamanensis (Alcock, 1894) 

Stereomastis auriculata (Bate, 1878) 

Stereomastis helleri (Bate, 1878) 

Stereomastis phosphoreus (Alcock, 1894) 

Stereomastis sculpta (S.I. Smith, 1880) 

Stereomastis trispinosa (De Man, 1905) 

GLYPHEIDAE 

Neoglyphea inopinata Forest and De Saint Laurent, 1975 

SYNAXIDAE 

Palibythus magnificus Davie, 1990 

Palinurellus wieneckii (De Man, 1881)


PALINURIDAE 

Justitia chani Poupin, 1994 

Justitia japonica (Kubo, 1955) 

Justitia longimanus (H. Milne Edwards, 1837) 

Justitia vericeli Poupin, 1994 

Linuparus sordidus Bruce, 1965 

Linuparus trigonus (Von Siebold, 1824) 

Palinustus unicornutus Berry, 1963 

Palinustus waguensis Kubo, 1963 

Panulirus albiflagellum Chan and Chu, 1996 

Panulirus homarus (Linnaeus, 1758) 

Panulirus longipes (A. Milne Edwards, 1868) 

Panulirus ornatus (Fabricius, 1798) 

Panulirus pascuensis Reed, 1954 

Panulirus penicillatus (Olivier, 1791) 

Panulirus polyphagus ((Herbst, 1793) 

Panulirus stimpsoni Holthuis, 1963 

Panulirus versicolor (Latreille, 1804) 

Puerulus angulatus (Bate, 1888) 

Puerulus velutinus Holthuis, 1963 

SCYLLARIDAE 

Arctides regalis Holthuis, 1963 

Ibacus brevipes Bate, 1888 

Ibacus brucei Holthuis, 1977 

Ibacus ciliatus (Von Siebold, 1824) 

Ibacus novemdentatus Gibbes, 1850 

Ibacus peronii Leach, 1815 

Ibacus pubescens Holthuis, 1960 

Parribacus antarcticus (Lund, 1793) 

Parribacus caledonicus Holthuis, 1960 

Parribacus holthuisi Forest, 1954 

Parribacus scarlatinus Holthuis, 1960 

Scyllarides haanii (De Haan, 1841) 

Scyllarides squammosus (H. Milne Edwards, 1837) 

Scyllarus aesopius Holthuis, 1960 

Scyllarus aureus Holthuis, 1963 

Scyllarus aurora Holthuis, 1982 

Scyllarus batei Holthuis, 1946 

Scyllarus bertholdii Paulson, 1875 

Scyllarus bicuspidatus (De Man, 1905) 

Scyllarus cultrifer (Ortmann, 1897) 

Scyllarus demani Holthuis, 1946 

Scyllarus gibberosus (De Man, 1905) 

Scyllarus martensii Pfeffer, 1881 

Scyllarus rapanus Holthuis, 1993 

Scyllarus rugosus H. Milne Edwards, 1837 

Scyllarus sordidus (Stimpson, 1860) 

Scyllarus timidus Holthuis, 1960 

Scyllarus umbilicatus Holthuis, 1963 

Scyllarus vitiensis (Dana, 1852) 

Thenus orientalis (Lund, 1793) 



982 Lobsters 

Nephropidae NEPHROPIDAE 

True lobsters and lobsterettes 

Diagnostic characters: Moderate to 

large-sized crustaceans. Body tubular, 

surface almost naked or covered with thick 

fur. Carapace (or “head”) with a well-developed 

rostrum, ornamented with spines or 

first 3 legs 

nodules, occasionally smooth. Eyes usually with 

well developed and black, or small and lack- pincers 

ing pigmentation, or even absent. Antennae 

very long and thread-like; antennal scale, if 

present, with inner margin unarmed and 

1 

curved. First 3 pairs of legs forming true 

pincers, with the first pair greatly enlarged 

and long. Both abdomen and tail fan 

well developed and powerful. Abdomen 

smooth or variously sculptured, pleura ending 

in acute ventral tooth. Tail fan entirely 

hardened, telson with fixed spines and 

posterior margin broadly convex. Colour: 

with the typical coloration of deep-sea crustaceans 

- from white to pink or red, some 

species with special markings (usually red 

and/or white) on body; setae or hairs, if present, 

are light brown. 

Habitat, biology, and fisheries: All species 

are deep-sea forms and found at depths from 

150 to more than 1 893 m. Their adult sizes 

are ranging from 5 cm to over 40 cm. They 

are bottom-dwellers with a preference for 

soft substrate. Some species are known to 

live in self-made burrows. The sexes are easily 

distinguished by the position of the 

gonopores, which are situated at the bases 

of the third and fifth legs in the females and 

males, respectively. In addition, the first 

pleopods (“legs” of the abdomen) of males 

are transformed into a strong and rigid penlike 

copulatory organ while they are reduced and thread-like in females. The females bear large and 

colourful eggs (white, blue or red, and other colours) on the ventral side of the abdomen. They seem to 

have a long incubation period, but a short larval development. At present, none of the species of this family 

is of significant commercial importance in the Western Central Pacific. However, the large size of several 

species and the fact that some of them are commonly caught on the basis of exploratory trawling suggest 

that they may have some commercial potential with the development of deep-sea fisheries. This is 

especially the case with members of the genus Metanephrops, which are treated here in separate species 

accounts. 

st 1 

rostrum 

well 

developed 

pincer 

enlarged 

2 

3 

4 

5 

median 

carina 

pleura with 

ventral tooth 

tail fan hardened 

gonopore 

leg 3 

leg 4 

leg 5 

male 

sperm 

receptacle 

female 

bases of last 3 legs in nephropids 

gonopore 

abdominal 

sternite 

copulatory 

stylets 

male female 

first pleopods of nephropids 

reduced, 

thread-like

Nephropidae 983 


Thaumastochelidae: body somewhat flattened dorsoventrally; 

antennal scale with several large teeth along inner margin; first 

pair of pincers very unequal; abdominal pleura short, quadrangular 

and not ending in a point; telson unarmed. 

Enoplometopidae: body distributed with tufts of long stiff hairs; 

second and third legs forming false pincers; telson bearing 

movable spines. 

Polychelidae: carapace box-like, rostrum absent or rudimentary; 

first 4 or all legs with pincers, first pair long but slender; 

telson pointed. 

Glypheidae: body somewhat flattened dorsoventrally; eyes 

inserted on a median elevation of cephalon; first 2 legs forming 

false pincers. 

Synaxidae: body somewhat flattened dorsoventrally and hairy;all 

legs simple and without pincers, the first pair heavier than the 

others; antennae whip-like and shorter than carapace; posterior 

half of tail fan soft and flexible. 

movable 

spines 

2 nd and 3 rd legs 

with false 

pincers 

telson 

rostrum 

2 

Enoplometopidae 

1 false pincers 

3 

4 


Glypheidae 

5 

5 

3 

4 

1 

2 

tufts of 

long stiff 

hairs 

1 st pincers very unequeal 

antennal 

scale 

1 st leg 

long and 

slender 

pleura 

quadrangular 

telson 

pointed 

1 

2 

3 

4 

no pincers Polychelidae 

1 2 

antenna 

short 

3 

Synaxidae 

5 

1 

2 

Thaumastochelidae 

5 

3 

4 

first 4 or all 

legs with 

pincers 

4 

5 

posterior half of 

tail fan soft

984 Lobsters 

Palinuridae: carapace without rostrum or with rostrum reduced to a small spine, but with a pair of large 

frontal horns over the eyes; antennae very long and thick, whip-like or spear-like; legs without true pincers 

and first pair generally not enlarged (except in Justitia); posterior half of tail fan soft and flexible. 

Scyllaridae: body strongly flattened dorsoventrally; rostrum absent or rudimentary; eyes enclosed by 

distinct orbits; antennae plate-like; legs without true pincers and none of them enlarged; posterior half of 

tail fan soft and flexible. 

no pincers 

Key to the genera of Nephropidae occurring in the area 

1a. Eyes large and black; antennal scale present; body provided with some spines but never 

uniformly spinulose (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metanephrops 

1b. Eyes minute, cornea lacking pigmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

2a. Antennal scale present; body more or less uniformly spinulose and not covered with soft 

pubescence (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Acanthacaris 

(a single species, A. tenuimana, in the area) 

2b. Antennal scale absent; body not uniformly spinulose but covered with thick pubescence 

(Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nephropsis 

Fig. 1 Metanephrops 

1 

Palinuridae 

antennal 

scale 

2 

5 

3 

4 

frontal 

horns 

posterior half 

of tail fan soft 

no 

pincers 

antennal 

scale 

Scyllaridae 

antenna 

plate-like 

5 

1 

4 

2 

3 

body flat 



Fig. 2 Acanthacaris Fig. 3 Nephropsis


Key to the species of Metanephrops occurring in the area 

1a. Carapace rather uniformly spinulose 

(Fig. 4a); dorsal surface of 

uropods covered with spinules . . . . . → 2 

1b. Carapace smooth between the 

ridges and large spines (Fig. 4b); 

uropods unarmed dorsally . . . . . . . . → 4 

2a. Abdominal segments each with 

2 transverse grooves (Fig. 5a); 

large pincer with finger distinctly 

longer than palm 

(Fig. 6a) . . . . . . Metanephrops neptunus 

2b. Abdominal segments each with 

surface 

uniformly 

spinulose 

no 

spinules 

between 

large 

spines 

and 

ridges 

a) Metanephrops neptunus b) Metanephrops sibogae 


1 transverse groove only (Fig. 5b, c); large pincer with finger shorter than palm (Fig. 6b) . . . . . → 3 

3a. Abdomen with deep longitudinal furrows (Fig. 5b); large pincer covered with sharp 

tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metanephrops arafurensis 

3b. Abdomen without distinct longitudinal furrows (Fig. 5c); large pincer finely granular to 

nearly smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metanephrops australiensis 

a) Metanephrops 

neptunus 

longitudinal 

furrows 

2 

transverse 

grooves 

b) Metanephrops 

arafurensis 

1 

transverse 

groove 

Fig. 5 first 3 abdominal segments (dorsal view) 

c) Metanephrops 

australiensis 

4a. Abdomen with distinct transverse and longitudinal grooves (Fig. 7); large pincers 

distinctly ridged (Fig. 8a), with outer border flat . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

4b. Abdomen smooth or with only narrow transverse grooves; large pincer smooth or weakly 

ridged (Fig. 8b), with outer border always angular . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

5a. Elevated parts of abdomen naked and smooth (Fig. 7a) . . . . . . . . Metanephrops andamanicus 

5b. Elevated parts of abdomen coarse and pubescent (Fig. 7b) . . . . . . . . Metanephrops velutinus 

elevated 

parts 

naked 

and 

smooth 

longitudinal 

grooves pubescent 

a) Metanephrops andamanicus b) Metanephrops velutinus 


finger 


neptunus 

palm 

Fig. 6 large pincer 

distinctly 

ridged 

finger 


australiensis 

weakly 

ridged 



andamanicus 

sibogae 


palm

986 Lobsters 

6a. Abdomen smooth, without grooves (Fig. 9a) . . . . . . . . . . . . . . . . . Metanephrops sibogae 

6b. Abdomen with narrow transverse grooves (Fig. 9b, c) . . . . . . . . . . . . . . . . . . . . . . → 7 

surface 

smooth 

no transverse 

groove on 1 st 

segment 

a) Metanephrops sibogae b) Metanephrops thomsoni 


transverse 

groove on 1 st 

segment 

c) Metanephrops sinensis 

7a. Two postorbital spines present (Fig. 10a); large pincers with inner margin naked but 

often bearing some large spines (Fig. 11a); first abdominal segment generally without 

distinct transverse grooves (Fig. 9b) . . . . . . . . . . . . . . . . . . . . . Metanephrops thomsoni 

7b. Three postorbital spines present (Fig. 10b); large pincers without large spines along 

inner margin, lateral margin of movable fingers bearing a brush of setae (Fig. 11b); first 

abdominal segment usually with short, lateral, transverse grooves (Fig. 9c) . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metanephrops sinensis 

2 postorbital 

spines 

3 postorbital 

spines 

a) Metanephrops thomsoni b) Metanephrops sinensis 


large 

spines 

brush of 

setae 

a) Metanephrops thomsoni b) Metanephrops sinensis 


Key to the species of Nephropsis occurring in the area 

1a. Rostrum without lateral teeth (Fig. 12a) . . . . . . . . . . . . . . . . . . . . Nephropsis ensirostris 

1b. Rostrum with lateral teeth (Fig. 12b-d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

2a. Rostrum with 2 pairs of lateral teeth (Fig. 12b) . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2b. Rostrum with 1 pair of lateral teeth (Fig. 12c, d) . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

no lateral 

teeth on 

rostrum 

2 pairs of 

lateral teeth 

subdorsal 

ridges 

spinous 

1 pair of 

lateral teeth 

subdorsal 

ridges 

lacking 

spines 

a) Nephropsis ensirostris b) Nephropsis sulcata c) Nephropsis serrata d) Nephropsis stewarti 

Fig. 12 carapace (dorsal view)


3a. Abdomen with a median longitudinal carina (Fig. 13) . . . . . . . . . . . . . . .Nephropsis sulcata 

3b. Abdomen without a median longitudinal carina . . . . . . . . . . . . . . . . . . Nephropsis suhmi 

4a. Basal part of telson with an erect dorsal spine (Fig. 14a) . . . . . . . . . . . Nephropsis acanthura 

4b. Basal part of telson without an erect dorsal spine (Fig. 14b) . . . . . . . . . . . . . . . . . . . → 5 

5a. Abdomen with a median longitudinal carina . . . . . . . . . . . . . . . . . . . Nephropsis holthuisi 

5b. Abdomen without a median longitudinal carina . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

6a. Subdorsal ridges of carapace spinose (Fig. 12c) . . . . . . . . . . . . . . . . . Nephropsis serrata 

6b. Subdorsal ridges of carapace lacking spines (Fig. 12d) . . . . . . . . . . . . . Nephropsis stewarti 

abdomen (dorsal view) 

Fig. 13 Nephropsis sulcata 

carinae 




Metanephrops andamanicus (Wood-Mason, 1891) 

Metanephrops arafurensis (De Man, 1905) 







Nephropsis acanthura Macpherson, 1990 

Nephropsis ensirostris Alcock, 1901 

Nephropsis holthuisi Macpherson, 1993 

Nephropsis serrata Macpherson, 1993 


Nephropsis suhmi Bate, 1888 

Nephropsis sulcata Macpherson, 1990 

no spine 

b) Nephropsis holthuisi, 

a) Nephropsis acanthura 

N. serrata, N. stewarti 

Fig. 14 tail fan (lateral view) 

References 

Chan, T.Y. and H.P. Yu. 1993. The Illustrated lobsters of Taiwan. Taipei, SMC Publishing Inc., 248 p. 

Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue 

of species of interest to fisheries known to date. FAO Fish. Synop., 125(13):1-292. 

spine

988 Lobsters 



Phoberus tenuimanus Bate, 1888; P. caecus 

sublevis Wood-Mason and Alcock, 1891; 

Acanthacaris opipara Burukovsky and Musy, 

1976; Phoberus brevirostris Thung and 

Wang, 1985 /None. 

FAO names: En - Prickly deep-sea lobster; 

Fr - Langoustine spinuleuse; Sp - Cigala 

raspa. 

Diagnostic characters: A large lobster. 

Body tubular, entirely covered with small 

spines and sharp tubercles, but not covered 

with soft pubescence. Carapace with 

a well-developed rostrum. Eyes minute, 

cornea lacking pigmentation. Antennae 

long and thread-like, antennal scale well 

developed. First 3 pairs of legs ending in 

pincers; first pair equal-sized, very slender 

and longer than body, with pincers 

bearing long teeth along cutting edges; 

second pair of legs much longer but less 

spiny than third pair. Tail fan entirely hardened; 

posterior margin of telson truncate. 

Colour: uniform delicate pink. 

Size: Maximum carapace length 21 cm 

(total length to 40 cm), ovigerous females 

between 11 and 19 cm carapace length. 

Habitat, biology, and fisheries: Found in 

deep water at depths from 600 to 1 670 m 

on muddy bottoms where it lives in burrows. 

This species is occasionally taken during 

exploratory trawling operations in deep 

water, but only in very small quantities. Due 

to its large size it may have some fishery 

potential once suitable fishing grounds are 

found. 

Distribution: Indo-West Pacific from the 

eastern coast of Africa to Japan, the East 

and South China Sea, Indonesia, and 

New Caledonia.




Nephrops sibogae De Man, 1916 / None. 

FAO names: En - Siboga lobster. 

Diagnostic characters: A small to mediumsized 

lobster. Body cylindrical and naked. 

Carapace spiny, but not uniformly so; 

rostrum well developed, armed with lateral 

and ventral teeth only; 4 pairs of 

postrostral teeth and 3 postorbital spines 

present. Eyes large and pigmented. 

Antennae long and thread-like, antennal 

scale present. First 3 pairs of legs ending in 

pincers; first pair enlarged and long; 

pincers feebly ridged and finely granular, 

their inner margin without pubescence 

but occasionally bearing a few large 

spines. Abdomen naked and smooth, 

without distinct sculptures. Tail fan 

entirely hardened; posterior margin of telson 

broadly convex. Colour: body uniformly 

orange-pink. 

Size: Maximum body length 18 cm, commonly 

to about 13 cm. 

Habitat, biology, and fisheries: At depths 

from 246 to 320 m on soft sandy sediment; 

probably lives in burrows. Of minor importance 

to fisheries in northern Australia, 

where it is trawled commercially during the 

northern prawn fishery closed seasons. 

Also taken in a fair amount during experimental 

trawling operations in southern Indonesia. 

Distribution: Only known from southern 

Indonesia and northern Australia. 

(after De Man, 1916)

990 Lobsters 



Nephrops thomsoni Bate, 1888 / None. 

FAO names: En - Red-banded lobster. 


lobster. Body cylindrical and naked. 

Carapace spiny, but not uniformly so; rostrum 

well developed, armed with lateral and 

ventral teeth only;3 pairs of postrostral teeth 

and only 2 postorbital spines present. Eyes 

large and pigmented. Antennae long and 

thread-like, antennal scale present. First 3 

pairs of legs ending in pincers; first pair enlarged 

and long; pincers feebly ridged and 

finely granular, inner margin without pubescence 

but often bearing some large 

spines. Abdomen without median longitudinal 

carina and weakly sculptured; transverse 

grooves very shallow (nearly absent 

on first segment) and broadly interrupted 

medially, longitudinal grooves absent.Tail 

fan entirely hardened; posterior margin of telson 

broadly convex. Colour: body almost 

uniformly orange-pink; post-orbital margin, 

tips of fingers of large pincers, margins of 

abdominal pleura, and posterior margin of tail 

fanwhitish. Eyesdarkbrown.Eggs blue, 

becoming dirty white when eye spots appear. 

Size: Maximum body length 15 cm, commonly 

between 9 and 12 cm. 

Habitat, biology, and fisheries: On sandy 

mud bottoms at depths from 50 to 509 m. 

Caught in the Philippines by lobster cages 

called “Panak”, but not in large quantities 

and only occasionally sold in local fish markets. 

Often taken on the basis of exploratory 

deep-water trawling around the northwest 

coasts of the Philippines and may therefore 

have more fishery potential in the area. 

Distribution: Western Pacific from Japan, the 

Ryukyu Islands, the East and 

South China Sea, Taiwan 

Province of China, and the 

Philippines. 

Remarks: Thecommonname 

“Red-banded lobster” refers 

to the presence of red bands 

on the first pair of legs. However, 

specimens known from 

the area (i.e. the Philippines) 

differ from those found elsewhere 

in its range by the lack 

of these red bands. 

(after Chan and Yu, 1988)




None / None. 

FAO names: En - Indian Ocean lobsterette; 

Fr - Langoustine indienne; Sp - Cigala del 

Oceano Indico. 


lobster. Body cylindrical, covered 

with thick fur. Carapace with a well-developed 

rostrum armed with 1 pair of lateral 

spines; anterior carapace bearing only 

supraorbital and antennal spines; subdorsal 

ridges without spines. Eyes minute, 

cornea lacking pigmentation. Antennae 

long and thread-like; antennal scale 

absent. First 3 pairs of legs ending in pincers; 

first pair rather stout and very hairy. 

Abdomen without median longitudinal 

carina, all pleura sharply pointed ventrally 

but lacking spines on front edges. Tail fan 

entirely hardened; outer blade showing a 

transverse fissure; telson with a pair of 

fixed posterolateral spines but unarmed 

dorsally. Colour: body whitish and covered 

with thick grey fur. Anterior carapace 

including rostrum, ventral surface, mouth 

parts and tail fan pink-red. Antennal and 

antennular flagella orange. Legs orangepink, 

with distal segments reddish; large 

pincers sometimes slightly orange. Eggs 

white. 

Size: Maximum known body length about 

20 cm, commonly between 10 and 15 cm. 

Habitat, biology, and fisheries: Deep sea 

at depths from 170 to 1 060 m, mostly between 

500 and 750 m on soft muddy substrate. 

A common bycatch of deep-water 

trawling operations throughout its range. Although 

it is probably the largest and most 

common species of the genus, its quantities 

are at present too small for significant interest 

to fisheries. 


in the Indo-West Pacific, 

from eastern Africa to Japan, the 

Philippines, Indonesia, and 

northwestern Australia. 

(after Alcock and Anderson, 1896)

992 Lobsters 

Metanephrops andamanicus (Wood-Mason, 1891) NEA 

En - Andaman lobster; Fr - Langoustine andamane; 

Sp - Cigala de Andamán. 

Maximum total body length 20 cm, commonly between 

15 and 18 cm. On hard mud in depths from 

250 to 750 m, but mostly from 300 to 450 m; probably 

lives in burrows. In the Western Central Pacific, 

so far only taken in very small numbers during 

experimental deep-water trawling operations, but 

its high abundance just north of the Philippines in 

the South China Sea may indicate that it has some 

fishery potential in the area. Indo-West Pacific from 

eastern Africa to the Andaman Sea, the South China 

Sea, Indonesia, and perhaps also Papua New 

Guinea. 


En - Northwest lobster. 

Maximum total body length 18 cm. At depths from 

418 to 500 m, on firm sediments such as Globigerina 

ooze; probably lives in burrows. In the area, 

so far only taken in few numbers during experimental 

deep-water trawling operations. However, as 

this lobster is the main component of the commercial 

Metanephrops fishery in northwestern Australia, 

more knowledge of its fishing grounds may 

reveal that it has also some fishery potential elsewhere 

in the Western Central Pacific. Known from 

the Philippines, Indonesia, and northwestern Australia. 

(after Alcock, 1894) 

(after Bruce, 1966)



En - Neptune lobster. 

Maximum total body length 25 cm; probably the largest 

species of the genus. On soft bottoms at depths 

to 940 m, mostly more than 500 m; probably lives in 

burrows. Occasionally caught in rather small quantities 

during deep-water trawling operations. With the 

development of deep-sea fishing gear this lobster is 

potentially very attractive for fisheries, due to its large 

size.Western Pacific from the South China Sea to the 

Philippines, Indonesia, and northwestern Australia. 


En - China lobster. 

Maximum total body length 15 cm. Found on 

muddy bottoms in deep water at depths from 203 

to 407 m, sometimes with shells. Not fished commercially 

at present, but often caught in large numbers 

during experimental deep-water trawling 

operations around the Philippines and may therefore 

have some fishery potential in the area. So far 

only known from the South China Sea and the 

northwestern coast of the Philippines. 

(after Bruce, 1965) 

(after Bruce, 1966)

994 Lobsters 


En - Velvet lobster. 

Maximum carapace length 8.6 cm. In deep water 

at depths of 238 to 702 m, mostly between 350 and 

450 m on hard muddy substrate. A commercial 

species in northwestern Australia, but not yet 

fished in the area. Often caught on the basis of 

exploratory deep-water trawling operations around 

the Philippines and Indonesia and may therefore 

also have some fishery potential in area. So far 

only found in the Philippines, Indonesia, and Western 

Australia. 




Enoplometopidae 995 

Enoplometopidae ENOPLOMETOPIDAE 

Diagnostic characters: Moderate- to smallsized 

crustaceans. Body cylindrical, distributed 

with tufts of long stiff hairs. 

Carapace (or “head”) with a well-developed rostrum. 

Eyes well developed and black. Antennae 

long and thread-like, antennal scale with 

inner margin unarmed and curved. First pair 

of legs greatly enlarged and forming true 

pincers; second to fifth legs slender and 

ending in false pincers (less distanct in posterior 

legs). Abdomen well developed and powerful, 

pleura more or less rounded and 

sometimes bearing strong spines. Tail fan 

entirely hardened; telson bearing movable 

spines, its posterior margin broadly convex. 

Colour: brilliant and attractive - orange-red, 

red, purple and/or white, with conspicuous 

spots on body, sometimes also with stripes. 

Habitat, biology, and fisheries: Reef lobsters 

usually live in coral and rocky reefs or in deeper 

parts of reef slopes at depths of 30 cm to 300 m. 

As in nephropids, the sexes can be determined 

by the position of the gonopores at the bases of 

legs. In addition, the first pleopods (“legs” of the 

abdomen) of males are large and leaf-like (thin 

but rigid), while they are small and thread-like 

in females. A large sperm receptacle process is 

present on the thoracic sternum between the 

last 3 legs in females. The eggs are small (about 

0.5 mm in diameter) and numerous. They hatch 

within a short time (about 6 days for 

Enoplometopus debelius) but the larvae are 

very difficult to rear. Since reef lobsters are 

nocturnal and shy, they are very difficult to 

catch. However, they are often highly valued in 

the aquarium trade for their attractive 

coloration. Reef lobsters presumably 

originating from Indonesia or the Philippines 

can be found in aquarium shops of other Asian 

countries, Europe, and the USA. Therefore, 

species accounts are provided here for the 

more common species. 

gonopore 

leg 1 

leg 2 

leg 3 

leg 4 

sperm 

receptacle 

leg 5 

female male 

bases of legs in reef lobsters 

Reef lobsters 

1 st leg with 

large pincer 

2 nd to 5 th 

legs with 

false 

pincers 

tufts of stiff 

hairs 

gonopore 

movable 

spines 

5 

telson 

first pleopod of male (Enoplometopus) 

4 

3 

1 

2

996 Lobsters 


Nephropidae: body almost naked or covered with thick fur; first 

3 pairs of legs ending in true pincers, with the first pair much 

larger than the others; telson only bearing immovable spines. 

Thaumastochelidae: body slightly depressed dorsoventrally; 

eyes strongly reduced, cornea lacking pigmentation; antennal 

scale bearing several large teeth along inner margin; first 3 pairs 

of legs (occasionally also fifth legs) ending in true pincers, first 

pair large but very unequal; abdominal pleura short, 

quadrangular and without large ventral tooth; telson unarmed. 

Polychelidae: eyes small, cornea lacking pigmentation; carapace 

box-like, rostrum absent or rudimentary; first 4 or all legs ending 

in true pincers, first pair long and slender; telson pointed. 

Glypheidae: body somewhat flattened dorsoventrally; eyes 

inserted on a median elevation of cephalon; first 2 legs forming 

false pincers with first pair very strong, third leg simple. 

Synaxidae: body somewhat flattened dorsoventrally and 

uniformly hairy; legs simple and without pincers, first pair 

much more robust than others; antennae rather thick and 

whip-like, shorter than carapace; posterior half of tail fan soft 

and flexible. 

1 st pincers very 

unequal 

true pincers 


pleura 

quadrangular 

1 

2 

rostrum 

3 

4 


false pincers 

Glypheidae 

5 


2 

1 

3 

4 

telson 

unarmed 

eye reduced 

5 

antenna 

short 

true 

pincers 

1 st leg long 

and slender 

eyes 

reduced 

3 

4 

5 

2 

2 

3 

4 

5 

1 

Nephropidae 

Polychelidae 

1 

Synaxidae 

1 

telson with 

immovable spine 

no pincers 

2 

3 

telson 

pointed 

5 

4 

true 

pincers 


of tail fan soft


Palinuridae: body tubular or slightly flattened dorsoventrally; rostrum absent or reduced to a small spine; 

carapace spiny and with a pair of large frontal horns above eyes; antennal flagella rather thick and very 

long, whip-like or spear-like; legs without true pincers and first pair generally not enlarged (except in 

Justitia); posterior half of tail fan soft and flexible. 

Scyllaridae: body strongly flattened dorsoventrally; rostrum absent or minute; eyes enclosed by distinct 

orbits; legs without true pincers and none of them enlarged; antennae plate-like; posterior half of tail fan 

soft and flexible. 

legs 

without 

true 

pincers 

1 

Palinuridae 

2 

3 

4 

5 

frontal 

horns 

posterior 

half of 

tail fan 

soft 

Key to the species of Enoplometopidae occurring in the area 

Note: since reef lobsters generally have particular colour patterns, live specimens of the various species 

are easily distinguished by their coloration. 

1a. Carapace with or without 1 postcervical spine (Fig. 1a, b); abdominal pleura broadly 

convex, with a blunt posterolateral angle (Fig. 2a); lateral margins of telson armed with 

1 pair of median spines (Fig. 3a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Carapace with 2 postcervical spines (Fig. 1c); abdominal pleura bearing a strong tooth 

(Fig. 2b); lateral margins of telson armed with 2 pairs of median spines (Fig. 3b) . . . . . . . . → 5 

2a. Rostrum bearing 2 pairs of lateral teeth; body orange-red and with colour markings 

mainly limited to lower carapace and posterior margins of abdominal segments . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enoplometopus chacei 

2b. Rostrum bearing 3 or more pairs of lateral teeth; colour spots and/or stripes present on 

whole body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

rostrum 

antennular 

peduncle 

a) Enoplometopus 

daumi 

4 median 

spines 

1 postcervical 

spine 

b) Enoplometopus 

occidentalis 

5 median 

spines 

2 postcervical 

spines 


c) Enoplometopus 

holthuisi 

no pincers 

body flat 



Scyllaridae 

antenna 

plate-like 

blunt 

angles strong teeth 

a) Enoplometopus 

occidentalis 

5 

1 

3 

4 

2 

b) Enoplometopus 

holthuisi 

Fig. 2 abdomen (lateral view)

998 Lobsters 

3a. Carapace with 5 median spines; rostrum reaching to or beyond antennular peduncle 

(Fig. 1b); colour orange-red with some conspicuous white spots on abdomen, fewer on 

carapace . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enoplometopus occidentalis 

3b. Carapace with 4 median spines; rostrum reaching to about middle of distal segment of 

antennular peduncle (Fig. 1a); colour mainly purplish . . . . . . . . . . . . . . . . . . . . . . . → 4 

1 spine 2 spines 

a) Enoplometopus occidentalis b) Enoplometopus holthuisi 

Fig. 3 tail fan (dorsal view) 

4a. Postcervical tooth distinct; body whitish, almost uniformly covered with small purple dots 

(Fig. 4a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enoplometopus debelius 

4b. Postcervical tooth rather indistinct; carapace with vertical reddish brown stripes; abdomen 

provided with many white spots (Fig. 4b). . . . . . . . . . . . . . . . . Enoplometopus daumi 

5a. Body pale pink and almost uniformly covered with small non-circular red spots (Fig. 4c) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enoplometopus gracilipes 

5b. Body reddish; lateral carapace with a large ocellated spot and some vertical white 

stripes (Fig. 4d); abdomen provided with many white spots . . . . . . . . Enoplometopus holthuisi 

purple spots 

a) Enoplometopus debelius b) Enoplometopus daumi 

non-circular 

red spots ocellated 

spot 

c) Enoplometopus gracilipes d) Enoplometopus holthuisi 




Enoplometopus chacei Kensley and Child, 1986 



Enoplometopus gracilipes (De Saint Laurent, 1988) 



reddish 

brown 

stripes 

Reference 

Holthuis, L.B. 1983. Notes on the genus Enoplometopus, with descriptions of a new subgenus and two new species 

(Crustacea Decapoda, Axiidae). Zool. Med. Leiden, 56(22):281-298.



En - Striped reef lobster. 

Carapace length of adults between 1.3 and 

2.6 cm, body length between 4 and 6 cm. 

Shallow coral reef areas. Shy, generally hide in 

rock cavities, often with only the pincers visible. 

Territorial and extremely aggressive against 

members of the same species except during 

mating. Collected by rotenone and probably 

also by divers. Not common, but a favourite for 

the aquarium trade because of its small size 

and special coloration. Live specimens are 

probably regularly exported from the 

Philippines and Indonesia. With certainty only 

known from Indonesia and the Philippines. 


En - Violet-spotted reef lobster. 

Carapace length of adults between 2.2 and 

2.4 cm, body length reaching 5 cm or more. On 

reef slopes in depths from 15 to 25 m. Collected 

by hand net during night diving. Appears to be 

slightly less aggressive than other species of 

the genus. An uncommon species that is sold 

at high prices in the aquarium trade for its 

attractive coloration. Specimens are mostly 

exported from Indonesia. Western Pacific from 

Indonesia to Hawaii and possibly Japan.

1000 Lobsters 


En - Bullseye reef lobster. 

Maximum body length about 12 cm. Lives on rocky 

reef slopes and deeper parts of reefs at depths of 

about 20 m to 80 m. Less shy and more aggressive 

than other species of the genus. Occasionally found 

in the aquarium trade and sold at a high price. 

Probably caught by night diving. Western Pacific 

from the Philippines, Indonesia, Eniwetok Atoll 

(Marshall Islands), Austral Islands, and Hawaii. 


En - Red reef lobster. 

Body length between 4 and 14 cm. Inhabits coral 

or rocky reefs and often found in deeper areas at 

the fringe of reefs, at depths of a few meters to 

about 100 m. Nocturnal and shy, usually found 

hiding in crevices and rocks. Very aggressive 

against members of the same species except 

during mating. Probably the most common species 

of the genus but still rather rare and only 

occasionally caught by divers collecting spiny 

lobsters or aquarium fishes. Small specimens are 

occasionally found in the aquarium trade and sold 

at a high price. Indo-West Pacific from eastern 

Africa to Japan, eastern Australia, and Hawaii.

Synaxidae 1001 

Synaxidae SYNAXIDAE 

Furry lobsters 

Diagnostic characters: Moderate- to small-sized crustaceans. 

Body somewhat flattened dorsoventrally and covered with a 

dense fur of short hair. Carapace long and laterally angular, 

covered with small rounded granules only, without enlarged spines; 

rostrum broad and flat triangular or rounded. Eyes small but 

distinct. Antennae rather thick and whip-like, but shorter than 

carapace; antennulae with flagella shorter than peduncle;stridulating 

organ sometimes present between bases of antennae and 

antennular plate.Legs without pincers but first pair much heavier 

than others. Both abdomen and tail fan well developed and powerful, 

posterior half of tail fan soft and flexible; abdomen hairy, with a 

low smooth keel along dorsal midline, but without transverse 

grooves. Colour: uniformly orange to bright orange or orange-red. 

Eyes black. 

Habitat, biology, and fisheries: This family contains only 2 genera 

and a total of 2 species in the Western Central Pacific. The shallow 

water genus Palinurellus is smaller (maximum total length about 

20 cm) than the slightly larger deep-water genus Palibythus 

(maximum total length 27 cm). Both genera live on hard bottom and 

are difficult to catch. Furry lobsters generally occur in few numbers 

and have therefore a very limited commercial potential. However, 

the “exotic” appearence and bright coloration of Palinurellus 

species has caught the attention of the tropical marine aquarium 

trade. Specimens presumably originating from the Philippines and 

Indonesia are occasionally sold for high prices in aquarium shops 

of other Asian countries, Europe, and the USA. The sexes of furry 

lobsters can be determined by the position of the gonopores as in 

the other families of lobsters. Furthermore, the first pair of pleopods 

(“legs” of the abdomen) is absent in males but present in females. 

antennae 

short, 

antennules 

whip-like rostrum 

no 

enlarged 

spines 

1 st leg very 

robust 

body 

covered 

with short 

hairs 

all legs 

without 

pincers 


Nephropidae: body tubular, almost naked or covered with thick 

fur; antennae very long and thread-like; first 3 pairs of legs ending in pincers, first pair greatly enlarged; 

tail fan entirely hardened. 

Thaumastochelidae: eyes strongly reduced, cornea lacking pigmentation; antennae very long and 

thread-like, scale with several large teeth along inner margin; first 3 pairs of legs (occasionally also fifth 

legs) ending in pincers, first pair large but very unequal; tail fan entirely hardened. 

pincers 

tail fan 

hardened 

Nephropidae 

1 

2 

3 

antennae long, 

thread-like 

5 

4 

antennae long, 

thread-like 


eye 

reduced 

tail fan 

hardened 

2 

5 

1 


3 

4 

1 

pincers 

2 

3 

4 

5

1002 Lobsters 

Enoplometopidae: body tubular, distributed with tufts of long 

stiff hairs; antennae very long and thread-like; first pair of legs 

enlarged and forming true pincer, second and third legs 

slender and ending in false pincers; tail fan entirely hardened. 

Polychelidae: eyes small, cornea lacking pigmentation; 

rostrum absent or rudimentary; antennae thread-like and 

shorter than body; first 4 or all legs with pincers, first pair long 

and slender; tail fan entirely hardened, with telson pointed. 

Glypheidae: eyes inserted on a median elevation of 

cephalon; antennae very long and thread-like; first 2 legs 

forming false pincers, with first pair very strong, third legs 

simple; uropods of tail fan entirely hardened. 

Palinuridae: body naked or with few and scattered hairs; 

carapace spiny and with a pair of large frontal horns over 

eyes, but lacking a median rostrum or with rostrum greatly 

reduced; antennae very long and whip-like or spear-like; 

legs without pincers and first pair generally not enlarged 

(except in one species of Justitia). 

Scyllaridae: body strongly flattened dorsoventrally; 

rostrum absent or minute; antennae plate-like; legs without 

pincers and none of them enlarged. 

1 st leg 

long and 

slender 

antennae 

long 

eyes 

reduced 

2 

4 

3 

5 

1 

Polychelidae 

1 

2 

Palinuridae 

telson 

pointed 

pincers 

frontal horns 

5 

3 

4 

carapace 

spiny 

antennae 

thread-like 

and long 

false 

pincers 

tufts of long 

stiff hairs 

true 

pincer 

tail fan 

hardened 


antennae thread-like 

and long 

1 

2 

3 

false pincers 

4 

Glypheidae 

Scyllaridae 

5 

antennae 

plate-like 

5 

1 

2 

4 

3 

5 

3 

4 

1 

2 

body flat

Synaxidae 1003 

Key to the species of Synaxidae occurring in the area 

1a. Antennular plate with stridulating organ (Fig. 1a); rostrum transversely oval, wider than 

long; lateral margin of carapace with distinct teeth behind anterolateral tooth (Fig. 2a) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palibythus magnificus 

1b. Antennular plate without stridulating organ (Fig. 1b); rostrum triangular, longer than 

wide; lateral margin of carapace without teeth behind anterolateral tooth (Fig. 2b) . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palinurellus wieneckii 

stridulating 

organ 

(from Davie, 1990) (from Davie, 1990) 

a) Palibythus magnificus b) Palinurellus wieneckii 

Fig. 1 lateral part of orbital region and antennular plate 




Palinurellus wieneckii (De Man, 1881) 

rostrum 

anterolateral 

tooth 

Reference 

Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue 

of species of interest to fisheries known to date. FAO Fish. Synop., 125(13):1-292. 

teeth 

(after Davie, 1990) 

a) Palibythus magnificus b) Palinurellus wieneckii 

Fig. 2 anterior part of carapace

1004 Lobsters 


En - Musical furry lobster. 

Maximum body length about 27 cm. On rocky 

bottoms at depths between 90 and 300 m. 

Although this furry lobster is of a fair size, it is 

rare and so far only caught by experimental 

trapping. Only known from Western Samoa and 

Tuamotu Archipelago. 

Palinurellus wieneckii (De Man, 1881) 

En - Indo-Pacific furry lobster; Fr - Cacahouète 

indopacifique; Sp - Langosta del Indo-Pacifico. 

Maximum body length about 20 cm, commonly 

between 7 and 14 cm. In shallow waters on 

coral reefs at depths from 9 to 27 m. Probably 

nocturnal and often found in deep caves. 

Occasionally caught by divers (using hand) or 

gill nets. Too rare and small to be of significant 

importance as food, but highly valued in the 

aquarium trade for its bright colour and rarity. 

Specimens in the aquarium trade are 

presumably originated from the Philippines and 

Indonesia. Widely distributed in the Indo-West 

Pacific from the eastern coast of Africa to the 

Red Sea, southern Japan, New Caledonia, 


(after Davie, 1990) 



Palinuridae 1005 

Diagnostic characters: Moderate- to 

large-sized crustaceans. Body tubular or 

slightly flattened dorsoventrally; hairs, if present, 

few and scattered. Carapace (or “head”) 

subcylindrical or box-like, laterally rounded or 

straight, without a well-developed rostrum, 

ornamented with spines and granules of 

various sizes, sometimes with scale-like 

sculpture (Justitia). Eyes well developed, 

each protected by a strong, spiny frontal 

projection of the carapace (frontal horns). 

Antennae rather thick and very long, whiplike 

or spear-like; antennal scale absent; antennulae 

slender and each with 2 long or short 

flagella. Bases of antennae often separated 

by a broad antennular plate usually bearing 

some spines. In some genera a projection 

from the base of antenna is developed 

and forms with the rim of the antennal plate a 

stridulating organ which can produce a grating 

sound by movement of the antenna. Legs 

generally simple, without true pincers; the 

first pair not or only slightly longer than the 

following legs (except in male of Justitia 

longimanus), but often somewhat more robust. 

Both abdomen and tail fan well developed 

and powerful, posterior half of tail fan 

soft and flexible. Abdominal segments either 

smooth or each provided with 1 or more transverse 

grooves. Colour: mostly brightly coloured 

and provided with special markings, 

bands or spots, or uniformly coloured. 

antennular flagella 

antennal 

peduncle 

PALINURIDAE 

Spiny lobsters, langoustes 

all legs without 

true pincers 

frontal 

horn 

cervical 

groove 

transverse 

grooves 

(in some 

species) 

antennular 

plate 

1 

dorsal view (Panulirus) 

2 

5 

antennal flagellum 

abdominal pleura 

lateral view (Panulirus) 

antennule 

3 

4 

thick, 

long 

antenna

1006 Lobsters 

gonopore 

female 

leg 1 

leg 2 

leg 3 

leg 4 

leg 5 

gonopore 

bases of legs in spiny lobsters 

male 

simple 

dactylus 

terminal segments of 5 th leg in spiny lobsters 

Habitat, biology, and fisheries: The maximum body length of species in the area ranges from about 14 cm 

to over 60 cm. The sexes are easily distinguished by the position of the gonopores which are situated at 

the bases of the third and fifth legs in females and males, respectively. Furthermore, the last leg forms a 

false (mostly) or true pincer in mature females, but is simple in mature males. Members of this family are 

characterized by the eggs (usually orange in colour) being very small and numerous, and by having a 

relatively shorter carrying time and very long planktonic larval stages (i.e. the phyllosoma). Spiny lobsters 

are all bottom-dwelling and can be found from very shallow water to a depth of 683 m. The shallow-water 

genus Panulirus comprises most of the species which are mainly nocturnal and live in coral or rocky reefs 

in depths less than 40 m. These are traditionally considered as excellent seafood and have a high economic 

value in the area. Juveniles are sometimes also seen in the aquarium trade. All species of Panulirus are 

actively fished throughout the area by divers (taken both by hand and spears), tangle nets, traps, or 

sometimes even by trawls, although they are not landed anywhere in large quantities. From 1990 to 1995, 

FAO’s Yearbook of Fishery Statistics reports a range of yearly catch of Panulirus of 2 450 to 4126 t from 

the Western Central Pacific. The other 4 genera occurring in the area are more commonly found in deeper 

water, some of them living in rocky areas (e.g. Justitia and Palinustus) and others on soft substrate (e.g. 

Linuparus and Puerulus). All these deep-water species are not very abundant, some even rare, and so far 

only taken as bycatch. Nevertheless, the rarer species are also used for human consumption. Due to their 

rather large size, some species may have more commercial potential with the development of deep-sea 

fisheries. Therefore, all spiny lobsters occurring in the area are treated here in separate species accounts. 

hatching 

OCEANIC PHASE 

mid 

larval 

stage 

early 

larval 

stage 

berried 

female 

late 

larval 

stage 

young 

adult 

female 

Puerulus 

settlement 

stage 

male 

life cycle of spiny lobsters of the genus Panulirus 

false pincer 

NURSERY AREAS INSHORE 

juvenile 

juvenile 

(after Phillips, 1985)



Nephropidae: body almost naked or covered with thick fur; rostrum well 

developed; antennae thin and thread-like; first 3 pairs of legs ending in 

pincers, first pair greatly enlarged; tail fan entirely hardened. 

Thaumastochelidae: eyes strongly reduced, cornea lacking 

pigmentation; rostrum well developed; antennae thin and 

thread-like, scale with several large teeth along inner margin; first 3 

pairs of legs (occasionally also fifth legs) forming pincers, first pair 

large but very unequal; tail fan entirely hardened. 

Enoplometopidae: body distributed with tufts of long stiff hairs; 

carapace with a well-developed rostrum; antennae thin and 

thread-like, antennal scale present; first pair of legs enlarged and 

forming true pincers, second and third legs ending in false pincers; 

tail fan entirely hardened. 

Polychelidae: eyes small, cornea lacking pigmentation; antennae 

thin and thread-like, shorter than body; first 4 or all legs ending in 

pincers, with first pair long and slender; telson entirely hardened 

and with telson pointed. 

Glypheidae: eyes inserted on a median elevation of cephalon; 

carapace with a well-developed rostrum; antennae thin and 

thread-like; first 2 legs forming false pincers, the first pair very 

strong, third legs simple; uropods of tail fan entirely hardened. 

true pincers 

rostrum 

eye 

reduced 

tail fan 

hardened 

eyes 

reduced 

telson 

pointed 

1 

2 

3 

4 

5 

5 

1 

2 

Polychelidae 

3 


4 

true pincers 

large 

pincer 

false 

pincers 

tufts of long 

stiff hairs 

tail fan 

hardened 

1 

true 

pincers 


rostrum 

2 

tail fan 

hardened 

3 

4 

false pincers 

Glypheidae 

Nephropidae 

5 

4 

5 

3 

1 


rostrum 

rostrum 

2 

1 

uropods 

hardened 

2 

3 

5 

4

1008 Lobsters 

Synaxidae: body very hairy; carapace laterally angular, with a broad and flat triangular or rounded rostrum, 

covered with small rounded granules but without enlarged spines; antennae whip-like but shorter than 

carapace; legs without pincers but first pair much heavier than others. 

Scyllaridae: body strongly flattened dorsoventrally; carapace depressed and laterally angular; eyes 

enclosed by distinct orbits and lacking large frontal horns; antennae plate-like; legs without pincers and 

none of them enlarged. 

no enlarged 

spines 

Synaxidae 

1 2 

antenna 

short 

3 

4 

5 

rostrum 

antenna 

plate-like 

eyes 

enclosed 

within 

orbits 

body flat 

Key to the genera of Palinuridae occurring in the area 

Note: members of this family, particularly those of the genus Panulirus, generally have distinctive colour 

markings on the body and live and fresh specimens of the various species are often easily distinguished 

by their coloration. 

1a. Carapace covered with scale-like sculpture; each abdominal segments with 4 or more 

transverse grooves; first pair of legs sometimes enlarged and forming false pincers in 

males (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Justitia 

1b. Carapace without scale-like sculpture; abdomen smooth or each segment at most with 

2 transverse grooves; first pair of legs simple and never enlarged . . . . . . . . . . . . . . . . → 2 

2a. Frontal horns truncated; proximal segment of antennular peduncle longer than antennal 

peduncle; legs very spinous (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palinustus 

2b. Frontal horns pointed; proximal segment of antennular peduncle distinctly shorter than 

antennal peduncle; legs sparsely covered with spines . . . . . . . . . . . . . . . . . . . . . . → 3 

scale-like 

sculpture 

Fig. 1 Justitia 

false 

pincer 

transverse 

grooves 

antennal 

peduncle 

Scyllaridae 

Fig. 2 Palinustus 

5 

1 

2 

4 

3 

proximal 

segment of 

antennular 

peduncle 

legs 

spinous


3a. Carapace subcylindrical, without median keel (Fig. 3); antennule with flagellum longer 

than peduncle (Fig. 4a); abdominal pleura ending in 1 strong tooth only (Fig. 5a) . . . . . Panulirus 

3b. Carapace box-like, with a median keel; antennule with flagellum much shorter than 

peduncle (Fig. 4b); abdominal pleura ending in 2 or more strong teeth . . . . . . . . . . . . . .→ 4 

Fig. 3 Panulirus 

no median 

keel 

peduncle 

flagellum 

flagellum 

peduncle 

a) Panulirus b) other genera 

Fig. 4 antennule of spiny lobsters 

4a. Frontal horns small and fused at the middle of anterior carapace (Fig. 6a); abdominal 

pleura ending in short teeth (Fig. 5b); antennae thick, inflexible and shorter than body 

length (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Linuparus 

4b. Frontal horns large and widely separated (Fig. 6b); abdominal pleura ending in 2 long 

teeth (Fig. 5c); antennae slender and much longer than body length (Fig. 8) . . . . . . . Puerulus 

pleura ending in 1 

strong tooth 

pleura ending in 2 or 

more short teeth 

a) Panulirus b) Linuparus 

pleura ending in 

c) Puerulus 2 long teeth 

Fig. 5 abdomen (lateral view) 

antennal 

flagellum 

frontal horns 

small, fused 

frontal horns large, 

widely separated 

a) Linuparus b) Puerulus 

Fig. 6 anterior part of carapace (dorsal view) 

Fig. 7 Linuparus Fig. 8 Puerulus

1010 Lobsters 

Key to the species of Justitia occurring in the area 

1a. Median spine on anterior margin of carapace flanked by 2 to 3 pairs of spines (Fig. 9a); 

antennular peduncle distinctly exceeding antennal peduncle; first pair of legs of males 

extremely long and forming false pincers (Fig. 10); abdomen not banded . . . . Justitia longimanus 

1b. A single median spine present on anterior margin of carapace (Fig. 9b); antennular 

peduncle reaching as far as antennal peduncle; first pair of legs short and simple in both 

sexes; abdomen covered with red bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

2 or 3 pairs 

of spines 

frontal 

horns 

1 median 

spine 

median 

spine 

smooth 

false pincer 

of males 

a) Justitia longimanus 

b) Justitia japonica 


Fig. 10 Justitia longimanus 

2a. Distal antennal segment with dorsal spine (Fig. 11a); frontal horns usually with 2 dorsal 

teeth; abdomen with intermediate grooves, usually interrupted and poorly defined, 

present between main grooves (Fig. 12a); red bands on abdomen located at posterior 

margin of each segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Justitia chani 

2b. Distal antennal segment without dorsal spine; frontal horns usually with 3 dorsal teeth; 

abdominal grooves simple, without intermediate grooves; red bands on abdomen 

located at anterior margin of each segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Antennal peduncle with disto-outer spine much larger than disto-inner spine (Fig. 11b); 

third to fifth abdominal segments each with 7 transverse grooves (Fig. 12b); lateral 

carapace without special coloured patches . . . . . . . . . . . . . . . . . . . . . Justitia japonica 

3b. Disto-outer and disto-inner spines of antennal peduncle similar in length (Fig. 11c); third 

to fifth abdominal segments each with 5 transverse grooves (Fig. 12c); lateral carapace 

with 2 red patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Justitia vericeli 

a) Justitia 

chani 

disto-inner 

spine 

disto-outer 

spine 

distal 

segment 

dorsal 

spine 

b) Justitia 

japonica 

disto-outer 

spine 

c) Justitia 

vericeli 

disto-inner 

spine 

Fig. 11 right antennal peduncle (dorsal view) 

intermediate grooves 

main 

grooves 

1 

1 

2 

3 

4 

5 

6 

2 3 

45 

6 

a) Justitia chani b) Justitia japonica 

1 

2 

3 

4 

c) Justitia vericeli 5 


(from Poupin, 1994) 

7 transverse grooves 

5 transverse 

grooves


Key to the species of Palinustus occurring in the area 

1a. Elevated parts of abdomen almost naked; 

anterior margin of carapace between 

frontal horns generally with a 

single median spine (Fig. 13a); postorbital, 

antennal and branchiostegal 

spines moderately long and similar in 

median 

spine 

frontal horns 

irregular 

spines 

size (Fig. 14a); antennules uniformly 

a) Palinustus unicornutus b) Palinustus waguensis 

orange-red and legs covered with 

broad pale bands . . Palinustus unicornutus Fig. 13 anterior part of carapace (dorsal view) 

1b. Elevated parts of abdomen distinctly pubescent; anterior margin of carapace between 

frontal horns provided with 1 to 8 irregularly arranged spines, or spines absent 

(Fig. 13b); postorbital spine distinctly shorter than antennal spine and branchiostegal 

spines (Fig. 14b); antennules banded and legs covered with dense narrow red rings 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Palinustus waguensis 

postorbital 

spine 

antennal 

spine 


spine 

a) Palinustus unicornutus b) Palinustus waguensis 

Fig. 14 anterior part of carapace (lateral view) 

postorbital 

spine 

antennal 

spine 


spine 

Key to the species of Linuparus occurring in the area 

1a. Strong median teeth present between 

frontal horns (Fig. 15a); epistome 

ridges with strong anterior 

teeth (Fig. 16a); second and third 

abdominal pleura bearing basal 

median teeth 

frontal horns 

teeth (Fig. 17) . . . . . Linuparus trigonus 

1b. No strong median teeth between 

frontal horns (Fig. 15b); epistome 

a) Linuparus trigonus b) Linuparus sordidus 

ridges without strong anterior teeth 

(Fig. 16b); second and third abdominal 

pleura without basal teeth . . . . 

. . . . . . . . . . . . . Linuparus sordidus 

Fig. 15 anterior part of carapace (dorsal view) 

a) Linuparus trigonus: 

epistome 

anterior tooth 

epistomal 

ridge 

Fig. 16 

epistome 

epistomal 

ridge 

b) Linuparus sordidus: 



basal teeth of pleura 

abdomen (lateral view) 

Fig. 17 Linuparus trigonus

1012 Lobsters 

Key to the species of Puerulus occurring in the area 

1a. Body heavily pubescent; postorbital 

spine present; median keel of carapace 

with 6 postcervical and 6 intestinal 

tubercles; eyes large, much 

broader than long (Fig. 18a) . . . . 

1b. 

. . . . . . . . . . . . . Puerulus velutinus 

Body only slightly pubescent; postorbital 

spine absent; median keel of 

carapace with 3 postcervical and 2 

intestinal teeth; eyes smaller, 

longer than broad (Fig. 18b) . . . . 

. . . . . . . . . . . . . Puerulus angulatus 

6 post- 

3 postcervicalcervical 

tubercles 

teeth 

6 intestinal 

2 intestinal 

tubercles 

teeth 

a) Puerulus velutinus b) Puerulus angulatus 


Key to the species of Panulirus occurring in the area 

1a. Abdomen provided with transverse grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Abdomen without transverse grooves or only with broad sunken pubescent areas . . . . . . . → 6 

2a. Anterior margins of transverse grooves on abdomen crenulated, grooves incomplete or 

interrupted in the middle (Fig. 19a); antennular plate bearing 4 well-separated principal 

spines and some small spinules (Fig. 20a); body greenish to brown, regions between 

eyestalks with bright orange and blue markings; legs blotched . . . . . . . . . Panulirus homarus 

2b. Transverse grooves on abdomen with straight anterior margins, not crenulated . . . . . . . . .→ 3 

3a. Antennular plate armed with 4 close-set principal spines (Fig. 20b); anterior margin of 

third abdominal pleuron spinous (Fig. 19b); body dark blue and brown, membranes at 

outer bases of antennae light blue; legs striped . . . . . . . . . . . . . . . . Panulirus penicillatus 

3b. Antennular plate with 2 principal spines (Fig. 20c); anterior margin of third abdominal 

pleuron not spinous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

1 

crenulated 

interrupted 

a) Panulirus homarus 

1 

2 

2 

e) Panulirus polyphagus 

3 

transverse 

grooves 

yellowish 

white bands 

3 

4 

4 

spinous 

1 2 

transverse 

grooves 

3 

4 

b) Panulirus penicillatus 

streak 

eye 

spot 

1 

2 

broad dark 

bands 

3 

f) Panulirus ornatus 

4 

1 

pleural groove 

2 

transverse 

groove 

c) Panulirus pascuensis 

Fig. 19 abdominal segments (lateral view) 

1 

2 

3 

4 

sunken 

pubescent areas 

3 

4 

1 2 

pleural groove 

transverse 

groove 

d) Panulirus longipes 

sunken pubescent 

1 areas 

2 

g) Panulirus versicolor h) Panulirus stimpsoni 

3 

3 

4 

4


4a. Transverse groove of second abdominal segment not joining corresponding pleural groove 

(Fig. 19c); body dark purple with some greenish, not speckled; irregular pale bands along 

posterior margin of abdominal segments, sometimes with whitish spots mixed in with them; 

spots on basis of tail fan; antennules not banded; legs striped . . . . . . . . . . Panulirus pascuensis 

4b. Transverse groove of second abdominal segment joining corresponding pleural groove (Fig. 19d) . . . . → 5 

4 principal 

spines 

2 principal 

spines 

eye 

additional 

spinules 

a) Panulirus homarus 

d) Panulirus polyphagus 

4 principal 

spines 

4 close-set 

principal spines 

frontal 

horns 

b) Panulirus penicillatus 

4 principal 

spines 

2 principal 

spines 

e) Panulirus ornatus 

f) Panulirus versicolor 

Fig. 20 antennular plate 

additional 

spinules 

c) Panulirus longipes 

4 principal 

spines 

additional spinules 

g) Panulirus stimpsoni 

5a. Median area at anterior carapace behind frontal horns usually bearing a longitudinal row of 

3 spines only (Fig. 21a); ventral surfaces of distal 2 antennal segments each with 2 large 

spines only (Fig. 22a); thoracic sternum with 2 strong submedian protrusions (Fig. 23a); 

antennules with outer flagella dark brown and inner flagella entirely whitish; antennal 

peduncle including stridulating pad pinkish; lateral carapace with 2 complete longitudinal 

white strips extending along the entire carapace; legs striped . . . . . . . . Panulirus albiflagellum 

5b. Median area at anterior carapace behind frontal horns always bearing some smaller, 

irregular spines in addition to the regular row of 3 spines (Fig. 21b); ventral surfaces of distal 

2 antennal segments each with 1 large spine (sometimes also with several other scattered 

spinules, Fig. 22b); thoracic sternum without strong submedian protrusions (Fig. 23b); 

antennules alternated with dark brown and white bands; antennular peduncle brown to 

purple and with stridulating pad bright blue; lateral carapace with 1 short (upper) and 1 long 

(lower) longitudinal white stripes; legs striped or spotted 

frontal horns 

. . . . . . . . . . . . . . Panulirus longipes 

row of 3 

spines only 

additional 

spines 

a) Panulirus albiflagellum b) Panulirus longipes 

Fig. 21 anterior part of carapace 

2 large 

spines 

on each 

segment 

1 large 

spine on 

each 

segment 


Fig. 22 distal 2 segments of antenna (ventral view) 

6a. Abdomen naked and smooth (Fig. 19e, f); legs blotched . . . . . . . . . . . . . . . . . . . . . → 7 

6b. At least second and third abdominal segments with broad sunken pubescent areas 

(Fig. 19g, h); legs striped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

7a. Antennular plate armed with 1 pair of principal spines (Fig. 20d); body pale green and 

abdomen with narrow transverse yellowish white bands (Fig. 19e) . . . . . . Panulirus polyphagus 

7b. Antennular plate armed with 2 pairs of principal spines (Fig. 20e); body greenish and 

abdomen with broad transverse dark bands (Fig. 19f), legs and antennules conspicuously 

ringed with light yellow and black . . . . . . . . . . . . . . . . . . . . . . Panulirus ornatus

1014 Lobsters 

8a. Antennular plate armed with 2 pairs of principal spines only (Fig. 20f); fourth to sixth 

abdominal segments smooth (Fig. 24a); body deep blue and green, abdomen with 

narrow transverse white bands, antennal and antennular flagella whitish . . . . Panulirus versicolor 

8b. Antennular plate with many small spinules in additional to the 2 pairs of principal spines 

(Fig. 20g); sunken pubescent areas present on all abdominal segments (Fig. 24b); body 

greenish, region between eyestalks not brightly marked and abdomen not banded 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panulirus stimpsoni 

leg 1 

leg 2 

strong 

protrusions 

leg 3 

leg 4 

leg 5 

leg 1 

leg 2 


Fig. 23 thoracic sternum (ventral view) 

leg 3 

leg 4 

leg 5 

sunken 

pubescent 

areas 













Panulirus longipes (A. Milne Edwards, 1868) 




Panulirus polyphagus (Herbst, 1793) 





References 

Chan, T.Y. and K.H. Chu. 1996. On the different forms of Panulirus longipes femoristriga (von Martens, 1872) 

(Crustacea: Decapoda: Palinuridae), with description of a new species. J. Nat. Hist., 30:367-387. 

Chan, T.Y. and H.P. Yu. 1993. The illustrated lobsters of Taiwan. Taipei, SMC Publishing Inc., 248 p. 

Chan, T.Y. and H.P. Yu. 1995. The rare lobster genus Palinustus A. Milne Edwards, 1880 (Decapoda: Palinuridae), with 

description of a new species. J. Crust. Biol., 15(2):376-394. 

Holthuis, L.B. 1991. Marine lobsters of the world. FAO species catalogue. Vol. 13. An annotated and illustrated catalogue of 

species of interest to fisheries known to date. FAO Fish. Synop., 125(13):1-292. 

Poupin, J. 1994. The genus Justitia Holthuis, 1946, with the description of J. chani and J. vericeli spp. nov. (Crustacea, 

Decapoda, Palinuridae). J. Taiwan Mus., 47(1):37-56. 

1 

2 

3 

4 

5 

6 

sunken 

pubescent 

areas 

1 

2 

3 

4 

5 

6 

a) Panulirus versicolor b) PanulIrus stimpsoni 

Fig. 24 abdomen (dorsal view)




Panulirus longipes (A. Milne Edwards, 1868); 

P. longipes femoristriga (Von Martens, 1872). 

FAO names: En - White whisker spiny lobster. 

Diagnostic characters: Carapace rounded and 

spiny; rostrum absent; anterior margin armed 

with irregular-sized spines; height of frontal 

horn about 2.5 times the eye height; median 

area behind frontal horns generally with a 

longitudinal row of 3 spines only; cervical 

groove distinctly wider than posterior 

marginal groove. Antennules with flagella 

longer than peduncle; antennular plate at 

bases of antennae bearing 1 pair of 

well-separated principal spines and some 

scattered spinules; ventral surfaces of distal 

2 antennal segments each with a row of 2 

equal-sized large spines only. First 4 pairs of 

legs without pincers. Thoracic sternum with 2 

strong submedian protrusions. Abdominal 

segments with a complete transverse groove 

joining the pleural groove; abdominal pleura 

only with that of second segment sometimes 

bearing spinules. Posterior half of tail fan soft 

and flexible. Colour: body dark brown to indigo, 

covered with numerous white spots and 

markings. Eyes black-brown. Lateral carapace 

with 2 longitudinal white stripes running 

along the entire carapace. Inner surface of 

antennal peduncle (including stridulating 

pad) and antennular plate pink; antennal 

flagella dorsally brown, ventrally whitish to pink; 

antennules with peduncle dark brown but 

laterally white, outer flagella dark brown and 

inner flagella entirely whitish. Legs striped 

with conspicuous white lines. Abdomen 

covered with numerous medium-sized white 

spots. Soft part of tail fan orange-brown with 

distal margin whitish. Pleopods somewhat 

greenish with white margins. Eggs orange. 

(after George and Holthuis, 1965) 

Size: Maximum body length about 25 cm, commonly between 18 and 21 cm. 

Habitat, biology, and fisheries: In shallow reef areas to a depth of about 20 m. Caught throughout its 

range, but apparently nowhere abundant. Nevertheless, in Indonesia this lobster can periodically be found 

in markets (live and fresh) and is sometimes even exported (live). No separate statistics are reported for 

this species because of former confusion with Panulirus longipes. 


Pacific from the Maldive 

Islands, Viet Nam, Indonesia, 

Great Barrier Reef (Australia), 

Taiwan Province of China, and 

Japan; probably also occurs in 

the Solomon Islands, New 

Hebrides, 

Polynesia. 

and French

1016 Lobsters 



Panulirus dasypus (H. Milne Edwards, 1837); 

P. burgeri (De Haan, 1841) / None. 

FAO names: En - Scalloped spiny lobster; 

Fr - Langouste festonée; Sp - Langosta 

festoneada. 

Diagnostic characters: Carapace rounded 

and spiny, sometimes with branchiostegal 

areas slightly inflated; rostrum absent; 

anterior margin armed with 4 regularly 

spaced large spines other than frontal horns; 

height of frontal horns about 2 times the 

eye height, without spinules in between. 

Antennules with flagella longer than 

peduncle, antennular plate at bases of 

antennae bearing 2 pairs of well-separated 

principal spines (anterior pair slightly 

larger) and some spinules. First 4 pairs of 

legs without pincers. Abdominal segments 

with a slightly crenate transverse groove, 

sometimes interrupted at the middle. 

Posterior half of tail fan soft and flexible. 

Colour: body greenish to brownish. Eyes 

dark brown. Anterior carapace and region 

between eyestalks with bright orange and 

blue markings. Frontal horns banded with 

black and white. Antennular flagella 

alternated with brown and white bands. 

Legs blotched. Abdomen covered with tiny 

white spots. Pleopods red-brown. Eggs 

orange. 

Size: Maximum body length 31 cm, 


Habitat, biology, and fisheries: In reef areas 

with sand in the surf zone and sometimes also 

in turbid waters at depths from 1 to 5 m, but 

can be found down to a depth of 90 m. 

Gregarious and nocturnal. The females produce 100 000 to 900 000 eggs per brood and hatching occurs 

after 25 to 59 days. The phyllosoma larvae last 4 to 7 months and have 9 stages. The juveniles molt every 

few weeks and become sexually mature after 2 to 3 years (carapace length 5 to 6 cm) after larval settlement. 

In the fourth year the reproductive potential is highest (carapace length 7 to 7.9 cm). The adults molt about 

4 times a year and the life span of this lobster is estimated to be 8 to 10 years. Actively fished throughout 

its range by hand, with traps, gill nets, cast nets, and baited lines. Big catches are often possible after 

typhoons or heavy rains. The fishery of this lobster is mostly local though it is exported in some areas such 

as the Philippines and Indonesia (sometimes live) together with other species of the genus. 




Africa to Japan, Australia, and 

the Marquesas Archipelago.


Panulirus longipes (A. Milne Edwards, 1868) LOJ 


Panulirus longipes femoristriga (Von Martens, 

1872) / Panulirus japonicus (Von Siebold, 1824); 

P. albiflagellum Chan and Chu, 1996. 

FAO names: En - Longlegged spiny lobster; 

Fr - Langouste diablotin; Sp - Langosta duende. 


and spiny; rostrum absent; anterior margin 

armed with irregular-sized spines; height of 

frontal horns about 2.5 times the eye height; 

median area behind frontal horns always 

bearing some additional spinules other than 

the regular longitudinal row of 3 spines; 

cervical groove about as wide as posterior 

marginal groove. Antennules with flagella 


bases of antennae bearing 1 pair of wellseparated 

principal spines and some 

scattered spinules; ventral surfaces of 

distal 2 antennal segments each with 1 large 

spine, often flanked by some scattered 

spinules. First 4 pairs of legs without pincers. 

Thoracic sternum without strong submedian 

protrusions. Abdominal segments 

with a complete transverse groove joining 

the pleural groove; abdominal pleura only 

with that of second segment sometimes 

Panulirus longipes longipes 

white or pale 

lines on legs 

Panulirus longipes 

femoristriga 

bearing spinules. Posterior half of tail fan soft and flexible. Colour: body dark brown to indigo and covered 

with numerous white spots and markings. Eyes black-brown. Lateral carapace with 1 short (upper) and 

1 long (lower) longitudinal white stripe. Inner surfaces of antennae and antennular plate brown to 

purple and with stridulating pad bright blue; antennal flagella brownish with ventral surface lighter in 

colour; antennules dark brown and alternated with conspicuous white bands. Legs covered with 

prominent white spots connected by orange lines, or only striped with white or pale lines. Abdomen 

covered with numerous small to medium-sized white spots. Soft part of tail fan orange-brown with 

posterior margin whitish. Pleopods somewhat greenish with white margins. Eggs orange. 

Size: Maximum body length 35 cm, commonly between 18 and 25 cm. 

Habitat, biology, and fisheries: Found in shallow coral or rocky reefs (but can be found down to a depth 

of 130 m), usually in clear waters with moderate currents, sometimes in slightly turbid waters. Nocturnal 

and not gregarious; females produce an average of 132 000 eggs per brood. Actively fished throughout its 

range, but apparently nowhere very abundant. Taken by hand during night diving or with spears, also with 

traps, tangle nets and lobster pots. The fishery of this lobster is mostly of local interest. In some regions, 

such as the Philippines and Indonesia, occasionally exported live, together with other species of this genus. 

Distribution: Widely distributed in the Indo-West Pacific from the eastern coast of Africa to Japan and Fiji. 

Two subspecies are recognized: the western or the spotted-legged form Panulirus longipes longipes 

occurring from eastern Africa to Thailand, Taiwan Province of China, the Philippines, and Indonesia; the 

eastern or the striped-legged 

form P. longipes femoristriga 

is known from Japan to the 

Bonin Islands, Taiwan 

Province of China, the 

Philippines, Indonesia, 

Australia, New Caledonia, the 

Loyalty Islands, New 

Hebrides, and Fiji; probably 

also in Micronesia, Papua New 

Guinea, Tonga, and the Cook 

Islands.

1018 Lobsters 



FAO names: En - Ornate spiny lobster; 

Fr - Langouste ornée; Sp - Langosta 

ornamentada. 

Diagnostic characters: Carapace 

rounded and spiny; rostrum absent; 

anterior margin bearing irregular-sized 

spines other than frontal horns; height 

of frontal horns about 2 times the eye 

height, without spinules in between. 

Antennules with flagella longer than 

peduncle; antennular plate at bases of 

antennae bearing 2 pairs of wellseparated 

principal spines (anterior 

pair considerably larger), sometimes 

also with several spinules.First4pairs 

of legs without pincers. Abdomen 

naked and smooth, without transverse 

grooves or sunken pubescent 

areas. Posterior half of tail fan soft and 

flexible. Colour: body greenish with 

carapace slightly bluish. Eyes blackbrown. 

Frontal horns intricately 

banded with yellowish white and 

brown markings. Antennal peduncle 

bluish with stridulating pad somewhat 

pinkish. Antennules and legs 

conspicuously ringed with pale 

yellow and black. Abdomen covered 

with broad transverse dark bands 

over middle of each segment and 

bearing large pale yellowish spots 

near hinges. Pleopods yellowish. Eggs 

orange. 

Size: Maximum body length about 60 cm, commonly between 20 and 35 cm. Maximum weight over 6 kg. 

Probably the largest species of the genus. 

Habitat, biology, and fisheries: Usually occurs at depths from 1 to 10 m, but can be found to a depth of 

200 m. In calm areas of coral and rocky reefs or reef slopes, sometimes also found on muddy substrate in 

river mouths with fairly turbid water. Lives solitary or in pairs; seasonal mass migrations have been observed 

in Torres Strait populations. Actively fished throughout its range, mostly by divers (using hand and spear), 

sometimes by hand nets and trawls (formerly a main fishing method in Torres Strait, but now banned). Sold 

mostly fresh or frozen in local markets, sometimes exported (as from the Philippines and Australia, live or 

tailed). In most parts of the area the catches are not very large but a commercial fishery for this species 

has been developed in Papua New Guinea (off the Gulf of Papua) and Australia (off Torres Strait and N.E. 

Queensland) since 1966, with an annual catch of about 250 t “tail weight” in 1990. It is reported that traps 

are not effective to catch this 

species and mass mortality 

may occur in breeding 

lobsters. 



from East Africa to Japan, 

Australia, and Fiji. Recently 

also reported to enter the 

Mediterranean from the Red 

Sea. 






FAO names: En - Pronghorn spiny lobster; 

Fr - Langouste fourchette; Sp - Langosta 

horquilla. 


and spiny, with branchiostegal areas slightly 

inflated; rostrum absent; anterior margin armed 

with 4 large and regularly spaced large spines 

other than frontal horns; height of frontal 

horns about 2 times the eye height; median 

area behind frontal horns with a longitudinal 

row of spinules. Antennules with flagella 


bases of antennae armed with 4 close-set 

principal spines (posterior pair larger).First 

4 pairs of legs without pincers. Abdominal 

segments with a transverse groove, not 

continuous with pleural groove; anterior 

margins of pleura spinous. Posterior half of 

tail fan soft and flexible. Colour: body dark blue 

and brown; males usually darker than females. 

Eyes black. Tips of large spines on carapace 

yellowish. Antennular peduncle striped with 

white lines, flagella uniformly brownish; 

membranous areas at outer base of antenna 

light blue. Legs conspicuously striped with 

white lines. Abdomen with tiny pale dots. 

Pleopods and soft part of tail fan black. 


commonly between 20 and 30 cm (males 

usually larger). 


shallow waters, usually at depths from 1 to 

4 m (maximum depth 16 m) at seaward 

edges of reefs, in clear waters not influenced 

by rivers. Nocturnal and usually not gregarious, but sometimes occurs in a “harem” of mixed sexes; often 

found in deep caves during the daytime and strongly clinging to rocks at surf zones or areas with strong 

currents such as surge channels. Good catches are often possible during dark nights, particularly after the 

full moon. The phyllosoma larval stage of this species probably lasts 7 to 8 months and has 10 substages. 

The females seem to be reproductive all year around in the south western Pacific. Requires silt-free clear 

waters and is therefore found in optimal conditions around oceanic islands; it is the predominant spiny 

lobster in the South Pacific Islands. Fished throughout its range and mostly taken during day and night 

diving by hand and spear, sometimes also by trammel nets and traps, but less effectively so. However, the 

catches are generally not very abundant and it is mostly sold fresh, live, cooked whole or tailed for local 

consumption, but also exported in some regions, such as from the Philippines and Indonesia. Considered a 

commercially threatened 

species by the World 

Conservation Union (IUCN). 

Distribution: Probably the 

most widely distributed species 

of the genus and can be found 

in the Indo-Pacific from the 

eastern coast of Africa to the 

Red Sea, Japan, Australia, 

French Polynesia, Hawaii, and 

the offshore islands near the 

western coasts of America (e.g. 

the Galapagos Archipelago).

1020 Lobsters 

Panulirus polyphagus (Herbst, 1793) 

Frequent synonyms / misidentifications: Panulirus fasciatus (Fabricius, 1798) / None. 

FAO names: En - Mud spiny lobster; 

Fr - Langouste de vase; Sp - Langosta 

fanguera. 


rounded and spiny; rostrum absent; 

anterior margin with irregular-sized 

spines other than frontal horns; height 

of frontal horns less than 2 times the 

eye height, without spinules in 

between. Antennules with flagella 

longer than peduncle; antennular plate 

at bases of antennae armed with 1 

pair of well-separated principal 

spines only. First 4 pairs of legs without 

pincers. Abdomen naked and smooth; 

without transverse grooves or 

sunken pubescent areas. Posterior 

half of tail fan soft and flexible. Colour: 

body dull green. Eyes black-brown. 

Spines on carapace with yellowish 

brown tips, orbital margin and 

posterior marginal groove yellowish 

white. Antennular plate with a medial 

longitudinal yellowish white line; 

antennular peduncle alternated with 

yellowish white and pale green 

bands, flagella banded with yellowish 

white and dark brown. Legs light 

brown with yellowish white blotches. 

Abdomen with tiny pale dots; a 

yellowish white band with brown 

margins near posterior border of 

each segment. Pleopods and soft part 

of tail fan orange-brown with yellowish 

white margins. 

Size: Maximum body length about 

40 cm, commonly between 20 and 

25 cm. 

Habitat, biology, and fisheries: Mainly 

found on muddy bottoms (sometimes 

also on rocky bottoms) in turbid waters near river mouths at depths from 3 to 90 m, but usually less than 

40 m deep. Unlike other spiny lobsters, this species is mainly taken by trawling, sometimes also by set 

nets, and seines, but rarely enters traps. Uncommon or absent in most parts of the area because of its 

preference for muddy substrate. Commercially and economically important mostly from the Gulf of Thailand. 

Sold fresh or frozen in local 

markets, and mounted dry 

specimens are sold as 

souvenirs to tourists. 


Pacific from Pakistan to India, 

Thailand, Viet Nam, Taiwan 


Philippines, Indonesia, Papua 

New Guinea, and northern 

Australia.



Frequent synonyms / misidentifications: None / Panulirus fasciatus (Fabricius, 1798) (= P. polyphagus 

(Herbst, 1793)). 

FAO names: En - Painted spiny lobster; 

Fr - Langouste barriolée; Sp - Langosta colorete. 

Diagnostic characters: Carapace rounded and 

spiny; rostrum absent; anterior margin bearing 4 

regularly spaced large spines other than frontal 

horns; height of frontal horns more than 3 times 

the eye height, without spinules in between. 

Antennules with flagella longer than peduncle; 

antennular plate at bases of antennae armed with 

2 pairs of well-separated principal spines only 

(anterior pair larger). First 4 pairs of legs without 

pincers. Abdomen more or less smooth, with 

broad but shallow sunken pubescent areas only 

present at each half of second and third 

segments. Posterior half of tail fan soft and flexible. 

Colour: body of adults generally blue and green; 

more greenish in large individuals. Carapace, 

including frontal horns, with a mosaic pattern of 

green, white and blue. Eyes black-brown. Antenna 

with inner surface pink and outer surface blue; 

inner surface of antennular peduncle white,outer 

surface blue; flagella whitish. Legs blue, 

distinctly striped with white lines. Abdomen 

greenish, having white lines with blue margins 

along posterior margin of each segment. Soft part 

of tail fan green and blue or orange-brown (large 

individuals). Pleopods blue, with white margins and 

a conspicuous medial white line. Coloration of 

juveniles somewhat different: body bluish to dark 

blue with conspicuous longitudinal white lines on 

lateral carapace; antennae and antennular plate 

almost entirely whitish; antennules bluish and 

distally whitish; legs with dark blue stripes; 

pleopods without medial white line. 

Size: Maximum body length about 40 cm, commonly between 20 to 30 cm. 

Habitat, biology, and fisheries: Found in reef areas at depths of usually less than 16 m (mostly between 4 

and 12 m) in clear or sometimes turbid water with strong currents, often on seaward edges of the reef plateau. 

Nocturnal and not gregarious; hides in crevices during the daytime with only the white antennae visible. Actively 

fished throughout its range by divers (using hand or spear), but apparently rarely enters traps or pots. The 

catches of this lobster are nowhere very large and it is mainly locally consumed live, fresh, cooked whole, or 

tailed, but in some regions, such as from the Philippines, exported live or tailed. The attractive coloration of the 

juveniles has caught the attention of the tropical marine aquarium trade and specimens presumably originating 

from the Philippines and Indonesia are sometimes sold in the aquarium shops of other Asian countries as well 

as in Europe and the USA 

(juveniles of other Panulirus 

species are occasionally also 

found in the aquarium trade, 

but usually sold at lower 

prices). 




to the Red Sea, Japan, 

Micronesia, Melanesia, 

northern Australia, and French 

Polynesia.

1022 Lobsters 


En - Small furrow lobster. 

Body length between 11 and 15.5 cm; probably 

the smallest species of the genus. On rocky 

substrate in depths from 150 to 340 m. Caught 

by tangle nets and dredges. Rare and of very 

limited interest to fisheries. Known from the 

western Pacific and so far only recorded from 

Japan, Taiwan Province of China, and New 

Caledonia. 


En - Japanese furrow lobster. 


between 15 and 25 cm; probably the largest 

species of the genus. On rocky bottoms at depths 

from 90 to 340 m. A limited fishery potential 

because of its deep, untrawlable habitat; seldom 

caught by lobster pots and trap nets. Indo-West 

Pacific from Madagascar to Taiwan Province of 

China, Japan, New Caledonia, and the 

Chesterfields Islands. 

(after Poupin, 1994) 

(after Baba et al., 1986)



En - Longarm furrow lobster; Fr - Langouste 

gibbon; Sp - Langosta de muelas. (The old FAO 

name of this species is re-used because recent 

research showed that J. longimanus and J. 

mauritiana represent the same species). 

Maximum body length 18.5 cm, commonly 

between 11 and 16 cm. At depths from 23 to 

454 m, usually between 50 and 150 m. Inhabits 

the outer parts of coral or rocky reef slopes. 

Although perhaps the most common species of 

the genus, it is only occasionally caught by 

traps, tangle nets, or divers. Of limited interest 

to fisheries because of its rocky habitat and 

usual occurrence in deeper waters. Worldwide 

distribution in tropical and subtropical seas; 

recorded in the western Atlantic from Bermuda 

to Brazil and in the Indo-Pacific from 

Madagascar to Taiwan Province of China, 

Japan, Hawaii, and French Polynesia. 


En - Polynesian furrow lobster. 

Body length about 18 cm. On hard coral bottoms 

at depths from 160 to 320 m. Apparently very rare 

and only caught during experimental trapping 

operations. So far only known from the Tuamotu 

Archipelago (French Polynesia). 

(after Miers, 1882) 

(after Poupin, 1994)

1024 Lobsters 


En - Oriental spear lobster. 

Body length between 17 and 27 cm. On sandy 

mud and limestone rocks in depths from 200 to 

500 m. A rare species, occasionally caught by 

trawls or traps.However, because of its large size 

and occurence in only moderately deep water, 

this lobster is potentially attractive for fisheries. 

Reported in the Indo-West Pacific from Taiwan 

Province of China, the South China Sea, 

northwestern Australia, and New Caledonia. 


En - Japanese spear lobster. 

Maximum body length 47 cm, commonly between 

20 and 35 cm. On sand or mud bottoms in depths 

from 30 to 318 m. Occasionally taken by trawls, but 

apparently nowhere very abundant. Rarely sold in 

fish markets in the Philippines. Caught 

commercially on a small scale off northern 

Australia and Queensland during the northern 

prawn fishery closed seasons. The lobsters are 

tailed because their rigid antennae make them 

difficult to pack up. Indo-West Pacific from Japan to 

Taiwan Province of China, Viet Nam, the 

Philippines, Indonesia, and Australia. 

(after Bruce, 1965) 

(from Ho and Yu, 1979)



En - Unicorn blunthorn lobster. 

Maximum body length about 14 cm. On hard 

bottoms at depths from 205 to 670 m. Rare and 

mainly taken during experimental trawling and 

trapping operations. Indo-West Pacific from the 

eastern coast of South Africa to La Réuion, 

Comoro Islands, Japan, Indonesia, New 

Caledonia, and French Polynesia. 


En - Japanese blunthorn lobster. 


to 10 cm. Inhabits the outer parts of coral and 

rocky reef slopes at depths from 72 to 180 m. 

The most common species of the genus 

although still moderately rare and only 

occasionally caught by tangle nets and trawls. 

A small species of very limited interest to 

fisheries. In Thailand, dried specimens are 

occasionally sold as souvenirs to tourists. 

Indo-West Pacific from Japan to Taiwan 

Province of China, the Philippines, Indonesia, 

Thailand, India, and Madagascar. 

(after Berry, 1979) 

(after Kubo, 1963)

1026 Lobsters 


En - Easter Island spiny lobster. 

Total body length of adults between 15 and 

25 cm. Lives on rocky substrate in shallow 

waters to a depth of 50 m; breeding season 

around December. Caught by divers during 

both day and night by hand and spears, also 

by gill nets and lobster pots. Sold fresh and 

consumed locally. Only known from Easter 

Island, Pitcairn Islands and Austral Islands of 

French Polynesia. 


En - Chinese spiny lobster. 

Maximum body length 35 cm, commonly 

between 16 and 23 cm; maximum weight about 

3 kg. Found in reef areas at depths of less than 

40 m. Mainly distributed along the southern 

coast of China, including Hong Kong, and the 

western coast of Taiwan Province of China. In 

the Western Central Pacific, only recently found 

in the Gulf of Thailand where it is apparently 

rare. 

(after George and Holthuis, 1965) 

(from Ho and Yu, 1979)



En - Banded whip lobster. 


between 11 and 17 cm. On soft substrate at 

depths from 180 to 536 m. Often taken as 

bycatch during deep-sea trawling operations, 

but usually not in large numbers. Of potential 

interest to fisheries. Indo-West Pacific from the 

eastern coast of Africa to Japan, Taiwan 

Province of China, the Philippines, Indonesia, 

New Guinea, northwestern Australia, New 

Caledonia, and Marquesas Islands. 


En - Velvet whip lobster. 

Maximum body length about 19 cm. On sand or 

mud bottoms at depths from 485 to 683 m. So 

far only taken during experimental trawling 

operations, but sometimes collected in 

relatively great numbers and may therefore be 

of potential interest to fisheries. Only known 

from the Philippines, Indonesia, and 

northwestern Australia. 

(after Berry, 1969) 

(after De Man, 1916) 



1028 Lobsters 

Scyllaridae SCYLLARIDAE 

Diagnostic characters: Small to large crustaceans. 

Body strongly depressed. 

Carapace laterally angular, rostrum absent 

or minute; eyes small but distinct and pigmented, 

enclosed within distinct orbits; no 

large frontal horns. Antennae short, broad, 

extremely flattened and plate- or scale-like; 

antennules short and slender, with short flagella. 

All legs similar in size and generally 

without pincers (except in Scyllarus cultrifer, 

third and fourth legs ending in false pincers). 

Both abdomen and tail fan well developed and 

powerful, posterior half of tail fan soft and 

flexible. Colour: usually drab, somewhat 

brownish in various shadings; dorsal surface 

of first abdominal segment often with a 

characteristic pattern of brightly coloured 

large spot(s). 

Habitat, biology, and fisheries: Six genera 

and about 30 species of slipper lobsters are 

currently known from the Western Central 

Pacific. Their adult sizes are ranging from 2 to 

50.5 cm body length. All species are bottom 

dwelling and are found from very shallow water 

to a depth of more than 484 m. As in spiny 

lobsters, the sexes can easily be distinguished 

by the position of the gonopores at the bases 

of the third or fifth legs. Mature female slipper 

lobsters also have the tip of the fifth leg 

transformed into a small false or true pincer 

(except in Thenus). The eggs are generally very 

small and numerous and orange or yellowish in 

colour. Slipper lobsters have a long planktonic 

phyllosoma larval stage similar to that of spiny 

lobsters. Members of the genera Arctides, 

Parribacus, and Scyllarides live in shallow 

coral and rocky reefs. Arctides species are rare 

but species of the latter 2 genera often form the 

Slipper lobsters 

cervical 

incision 

branchial 

carina 

body 

flattened 

antenna 

plate-like 

bycatches of fisheries for spiny lobsters and are of moderate commercial value. The genera Ibacus and 

Thenus mainly occur on soft bottoms (the former one in deeper waters) and can be taken by commercial 

trawlers. They reach a high production in some regions and may be used for export as well as for local 

consumption. Yearly production of all slipper lobsters in the Western Central Pacific from 1990 to 1995 

ranged from 1 641 to 3115 t (FAO Yearbook of Fishery Statistics). About 1/3 to 1/2 of this production 

originated in the Gulf of Thailand. The genus Scyllarus has the most numerous species in the family which 

occur in both soft and hard bottoms from shallow to deep waters. However, they are usually small in size 

and few in numbers and are therefore without economic value. Since the taxonomic status of many species 

of the genus Scyllarus is still unclear, a key to the species of this genus is not presented here, and only 

abbreviated species accounts are provided for 2 of the common species in the Western Central Pacific to 

give an idea of their general appearances. 


None. No other family of lobsters has such a strongly flattened body or plate-like antennae. 

distinct orbit

Scyllaridae 1029 

Key to the genera of Scyllaridae occurring in the area 

1a. Carapace with deep cervical incisions and lateral margins cut into large teeth; abdominal 

pleura directed laterally (Figs 1, 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .→ 2 

1b. Carapace with very shallow cervical incisions and lateral margins not cut into large teeth; 

abdominal pleura directed downwards (Figs 3 to 6) . . . . . . . . . . . . . . . . . . . . . . . .→ 3 

2a. Dorsal surface of body rather smooth and provided with distinct branchial carinae; fifth 

abdominal segment bearing a posteromedian spine; body colour plain (Fig. 1) . . . . . . . . Ibacus 

2b. Dorsal surface of body entirely covered with scale-like tubercles; branchial carina 

absent; fifth abdominal segment without posteromedian spine; body coloration mottled 

(Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parribacus 

surface 

smooth 

cervical 

incision 

deep 

branchial 

carina 

posteromedian spine 

on 5 th segment 

no spine 

cervical 

incision 

deep 

surface with 

scale-like 

tubercles 

Fig. 1 Ibacus 

Fig. 2 Parribacus 

3a. Body strongly depressed; carapace trapezoid and narrowing posteriorly; orbits located 

at anterolateral angles of carapace (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . Thenus 

(a single species, T. orientalis, in this genus) 

3b. Body not strongly depressed, sometimes slightly vaulted; carapace more or less rectangular; 

orbits situated on anterior margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

4a. Size large (up to 50.5 cm body length); distal margin of antenna finely crenate; abdomen 

uniformly granulate and not particularly sculptured (Fig. 4) . . . . . . . . . . . . . . . . Scyllarides 

4b. Size medium or small (body length less than 17 cm); distal margin of antenna cut into 

distinct teeth; abdomen with transverse grooves or arborescent sculpture . . . . . . . . . . . . → 5 

shallow 

cervical 

incision 

Fig. 3 Thenus orientalis 

orbit 

carapace 

trapezoid 

shallow 

cervical 

incision 

Fig. 4 Scyllarides 

finely 

crenate 

orbit 

carapace 

rectangular 

abdomen not 

sculptured

1030 Lobsters 

5a. Medium size (adults more than 12 cm body length); distal margin of antenna with more 

than 20 small teeth; exopods of maxillipeds with multiarticulate flagella (Fig. 5) . . . . . . Arctides 

(a single species, Arctides regalis, in the area) 

5b. Small size (usually less than 10 cm body length); distal margin of antenna with less than 

10 large teeth; exopods of maxillipeds without flagellum or with flagellum transformed 

to a single laminate segment (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scyllarus 

(generally, the species of Scyllarus can be separated into 2 groups: 1 has arborescent sculpture on the abdomen 

while the other group has broad transverse grooves on the abdomen and lacks an arborescent sculpture) 

orbit 

shallow 

cervical 

incision 

Fig. 5 Arctides regalis 

many small 

teeth 

carapace 

rectangular 

arborescent 

sculpture 

Key to the species of Ibacus occurring in the area 

shallow 

cervical 

incision 

carapace 

rectangular 

Fig. 6 Scyllarus 

few large 

teeth 

orbit 

margin 

with 

many 

teeth 

arborescent 

sculpture 

transverse 

grooves 

abdomen 

(Scyllarus rugosus) 

1a. Carapace with 6 to 9 posterolateral teeth (Fig. 7a-c) . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Carapace with more than 9 posterolateral teeth (Fig. 7d-f) . . . . . . . . . . . . . . . . . . . . → 4 

2a. Merus of third maxilliped concave on ventral surface and without deep incisions on inner 

margin (Fig. 8a); cervical incision very wide (Fig. 7a) . . . . . . . . . . . . . . . . . . Ibacus brucei 

2b. Merus of third maxilliped convex on ventral surface and provided with deep incisions on 

inner margin (Fig. 8b, c); cervical incision narrow (Fig. 7b, c) . . . . . . . . . . . . . . . . . . .→ 3 

3a. Branchial carina nearly straight (Fig. 7b); posterior incision of orbit without tubercle. 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ibacus novemdentatus 

3b. Branchial carina strongly convex (Fig. 7c); posterior incision of orbit with a distinct 

tubercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ibacus peronii 

cervical 

incision 

wide 

branchial carina 

straight 

7-8 


teeth 

a) Ibacus brucei b) Ibacus novemdentatus 

body heavily 

pubescent 

12-17 


teeth 

branchial carina 

strongly convex 

d) Ibacus brevipes e) Ibacus pubescens f) Ibacus ciliatus 


7-8 


teeth 

11-15 


teeth 

c) Ibacus peronii 

6 or 7 


teeth 

body naked 

10-12 


teeth


4a. Merus of third maxilliped convex on ventral surface and provided with deep incisions on 

inner margins (Fig. 8d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ibacus brevipes 

4b. Merus of third maxilliped concave on ventral surface and without deep incisions on inner 

margin (Fig. 8e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

anterior 

teeth of 

a) Ibacus brucei 

epistome incision 

c) Ibacus peronii 

merus of 3 rd 

maxilliped 

merus of 3rd maxilliped 

incision 

d) Ibacus brevipes 

Fig. 8 mouth field (ventral view) 

5a. Body heavily pubescent (Fig. 7e); 

posterior margin of fifth abdominal 

segment evenly serrated 

(Fig. 9a) . . . . . . . . . . . Ibacus pubescens 

5b. Body except distal segment of antenna 

naked (Fig. 7f); posterior 

margin of fifth abdominal segment 

only with a median spine and 3 to 

4 lateral tubercles (Fig. 9b) . . . Ibacus ciliatus 

anterior teeth 

of epistome 

b) Ibacus novemdentatus 

merus of 

3 rd 

maxilliped 

evenly 

serrated 

median 

spine 

incision 

e) Ibacus ciliatus 

a) Ibacus pubescens 

merus of 3 rd 

maxilliped 

4 

5 

6 

merus of 3 rd 

maxilliped 

b) Ibacus ciliatus 

Fig. 9 posterior part of abdomen (dorsal view) 

Key to the species of Parribacus occurring in the area 

median carina elevated transverse groove 

wide and naked 

1a Median carina on second and third 

abdominal segments markedly elevated; 

transverse grooves separating 

articulated and non-articulated parts 

articulated part 

of abdominal segments wide and almost 

naked (Fig. 10a) . . . Parribacus antarcticus 

a) Parribacus antarcticus 

1b Median carina on second and third 

non-articulated part 

abdominal segments low; transverse 

articulated 

grooves separating articulated and 

part smooth 

non-articulated parts of abdominal 

segments narrow and hairy (Fig. 10b) . . . . . → 2 

transverse groove 

narrow and hairy 

median 

carina 

low 

4 

5 

6 

b) Parribacus holthuisi 

Fig. 10 third abdominal segment (dorsal view)

1032 Lobsters 

2a. Articulated parts of abdominal segments bearing distinct tubercles (Fig. 11a); fourth 

segment of antenna with 7 outer teeth (apical tooth not included) . . . . . . Parribacus caledonicus 

2b. Articulated parts of abdominal segments more or less smooth (Fig. 10b); fourth segment 

of antenna with 5 to 6 outer teeth (apical tooth not included) . . . . . . . . . . . . . . . . . . .→ 3 

3a. Fourth segment of antenna bearing 5 outer teeth (apical tooth not included); the 

posterior of the 2 lateral teeth of carapace before cervical incision much smaller than 

the first (Fig. 11b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parribacus holthuisi 

3b. Fourth segment of antenna bearing 6 outer teeth (apical tooth not included); the 2 lateral 

teeth of the carapace before cervical incision only slightly unequal in size (Fig. 11c) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parribacus scarlatinus 

7 teeth on 

outer margin 

of 4th antennal 

segment 

teeth 

unequal 

a) Parribacus caledonicus b) Parribacus holthuisi c) Parribacus scarlatinus 

Key to the species of Scyllarides occurring in the area 

5 teeth on 

outer margin 

of 4 th 

antennal 

segment 

teeth 

strongly 

unequal 

Fig. 11 carapace and anterior abdominal segments (dorsal view) 

6 teeth on 

outer 

margin of 

4 th antennal 

segment 

teeth 

slightly 

unequal 

1a. Carapace with a deep cervical groove and with posterior part distinctly wider than 

anterior half (Fig. 12a); fourth abdominal segment always produced into a remarkable 

hump medially; posterior margin of second abdominal pleuron somewhat concave 

(Fig. 13a); only diffuse spots present on first abdominal segment . . . . . . . . . Scyllarides haanii 

1b. Carapace with a shallow cervical groove and with anterior part more or less as wide as 

posterior half (Fig. 12b); middle of fourth abdominal segment never with a hump, only 

moderately ridged; posterior margin of second abdominal pleuron somewhat convex 

(Fig. 13b); first abdominal segment with large distinct spots . . . . . . . . Scyllarides squammosus 

dorsal protuberances 

prominent 

1 2 

3 4 

5 

hump 

6 

cervical 

groove 

deep 

a) Scyllarides haanii 

dorsal protuberances 

weak 

cervical 

groove 

shallow 

Fig. 12 carapax (lateral view) 

b) Scyllarides squammosus 

1 2 3 

convex 

concave 

4 

5 

6 


a) Scyllarides 

haanii 

b) Scyllarides 

squammosus





Ibacus brevipes Bate, 1888 

Ibacus brucei Holthuis, 1977 



Ibacus peronii Leach, 1815 








Scyllarus aesopius Holthuis, 1960 

Scyllarus aureus Holthuis, 1963 

Scyllarus aurora Holthuis, 1982 

Scyllarus batei Holthuis, 1946 


Scyllarus bicuspidatus (De Man, 1905) 

Scyllarus cultrifer (Ortmann, 1897) 

Scyllarus demani Holthuis, 1946 

Scyllarus gibberosus (De Man, 1905) 

Scyllarus martensii Pfeffer, 1881 

Scyllarus rapanus Holthuis, 1993 


Scyllarus sordidus (Stimpson, 1860) 

Scyllarus timidus Holthuis, 1960 

Scyllarus umblicatus Holthuis, 1963 

Scyllarus vitiensis (Dana, 1852) 


References 

Chan, T.Y. and H.P. Yu. 1993. The illustrated lobsters of Taiwan. Taipei, SMC Publishing Inc., 248 p. 

Holthuis, L.B. 1991. FAO species catalogue. Marine lobsters of the world. Vol. 13. An annotated and illustrated catalogue 

of species of interest to fisheries known to date. FAO Fish. Synop., 125(13):1-292.

1034 Lobsters 



FAO names: En - Japanese fan lobster. 

Diagnostic characters: Body extremely 

flattened with dorsal surface rather smooth or 

slightly pitted; only distal segment of antenna 

heavily pubescent, particularly in adult males. 

Carapace with well-developed branchial carinae 

and deep cervical incisions, posterolateral 

margin cut into 10 to 12 large teeth. Eyes small 

and subspherical; orbits not closed and situated 

on anterior margin of carapace. Antennae broad, 

flattened and plate-like. Merus of third 

maxilliped ventrally concave, with only 

shallow incisions on inner margin. All legs 

without pincers and similar in size. Posterior 

margin of fifth abdominal segment armed with 

a median spine and 3 or 4 lateral tubercles; 

abdominal pleura directed laterally. Posterior 

half of tail fan soft and flexible. Colour: body 

purplish brown all over. Eyes dark brown. Legs 

orange-brown. Soft part of tail fan transparent and 

somewhat reddish brown. Setae light brown. 

Eggs orange. 

Size: Maximum body length 23 cm, commonly to 

about 15 cm. 

Habitat, biology, and fisheries: Foundonsoft 

substrate of sand, mud, or clay at depths from 48 

to 314 m, mostly between 100 and 250 m. Taken 

by commercial trawlers and common in fish 

markets of the Philippines, but sold at lower prices 

than spiny lobsters. The females carry the eggs 

for about 30 days until the larvae hatch out; the 

phyllosoma larvae passes through 9 stages in 76 

days and metamorphose to the benthic reptant 

larvae. 


from Japan, Korea, coast of 

China, Taiwan Province of 

China, the east coast of the 

Philippines, and Thailand. 

(from Holthuis, 1985)



Frequent synonyms / misidentifications: None / Ibacus ciliatus (Von Siebold, 1824); I. pubescens 

Holthuis, 1960; I. peronii Leach, 1815. 

FAO names: En - Smooth fan lobster; 

Fr - Cigale glabre; Sp - Cigarra liso. 


flattened and with dorsal surface rather 

smooth or slightly pitted. Carapace with 

nearly straight branchial carinae; 

cervical incisions deep but narrow; 

posterolateral margin cut into 7 or 8 

large teeth. Eyes small and subspherical; 

orbits not closed and without tubercle at 

posterior incision, situated on anterior 

margin of carapace. Antennae broad, 

flattened and plate-like. Merus of third 

maxilliped ventrally convex, with deep 

incisions on inner margin. All legs 

without pincers and similar in size. 

Posterior margin of fifth abdominal 

segment evenly serrated, bearing a 

median spine; abdominal pleura 

directed laterally. Posterior half of tail fan 

soft and flexible. Colour: body pale 

yellowish brown and marbled with reddish 

brown patches on dorsal surface. Ventral 

surfaces of carapace and legs brown and 

white. Eyes dark brown. Soft part of tail fan 

transparent and somewhat pale yellowish. 

Eggs orange. 



Habitat, biology, and fisheries: Found at 

depths from 37 to 400 m on levelled bottom 

of sand and mud. The phyllosoma larva of 

this species has 7 stages and lasts for 65 

days. Taken by commercial trawlers in the 

Philippines, but in smaller catches than 

Ibacus ciliatus. Also used as food and sold 

in local fish markets (with same local 

(from Holthuis, 1985) 

names as I. ciliatus). 


from the east coast of Africa 

to Japan, the Philippines, 

Indonesia, and northwestern 

Australia.

1036 Lobsters 


Frequent synonyms / misidentifications: Ibacus ciliatus pubescens Holthuis, 1960 / Ibacus ciliatus (Von 

Siebold, 1824). 

FAO names: En - Hairy fan lobster. 


flattened and with dorsal surface 

uniformly covered with dense short 

pubescence. Carapace with welldeveloped 

branchial carinae and deep 

cervical incisions, posterolateral 

margin cut into 11 to 15 distinct teeth. 

Eyes small and subspherical; orbits not 

closed and situated on anterior margin of 

carapace. Antennae broad, flattened and 

plate-like. Merus of third maxilliped 

ventrally concave, with only shallow 

incisions on inner margin. All legs 

without pincers and similar in size. 

Posterior margin of fifth abdominal 

segment evenly serrated, bearing a 

median spine; abdominal pleura 

directed laterally. Posterior half of tail 

fan soft and flexible. Colour: body pale 

brown with lateral teeth on carapace 

somewhat whitish. Eyes dark brown. Soft 

part of tail fan transparent. Setae light 

brown. 

Size: Maximum carapace length 7.8 cm, 

with a maximum body length of about 

20 cm. 

Habitat, biology, and fisheries: Over 

sand or mud bottoms at depths from 150 

to 391 m. Taken by trawls. The hairy fan 

lobster was formerly often confused with 

Ibacus ciliatus, and it is still not known 

which of these 2 species is more 

common in the Western Central Pacific. 

Very likely mixed with I. ciliatus in the 

markets of the Philippines. 


Distribution: So far only 

known amongst the islands 

and along the west coast of 

the Philippines, Indonesia, 

and northwestern Australia.



Frequent synonyms / misidentifications: Parribacus ursus major (Herbst, 1793) / None. 

FAO names: En - Sculptured mitten lobster; 

Fr - Cigale savate; Sp - Cigarra chinesa. 


flattened, with dorsal surface uniformly 

covered with scale-like tubercles and short 

hairs. Carapace with distinct rostral tooth; 

without branchial ridge but with deep cervical 

incisions; lateral margin cut into large teeth. 

Eyes small and subspherical; orbits not closed 

and situated on anterior margin of carapace. 

Antennae broad, flattened and plate-like; 

fourth segment armed with 6 large outer 

teeth (sometimes bifurcated and excluding 

apical tooth). All legs without pincers and 

similar in size. Abdomen with median carina 

on second and third segments markedly 

elevated; transverse grooves separating 

articulated and non-articulated parts of 

each segment are wide and almost naked; 

fifth segment without posteromedian 

spine; pleura directed laterally. Posterior 

half of tail fan soft and flexible. Colour: body 

yellowish and mottled with brown and black 

patches. Rostrum and orbital margin 

purplish. Eyes black. Large teeth on lateral 

carapace and antennae intricately banded 

with yellow, orange, light purple, deep 

brown and black distally. Abdomen with 2 

black lateral lines; central region of first 

segment yellowish with some brown 

patches. Ventral surface of body greenish 

yellow, with pale green spots. Legs 

greenish yellow and covered with green 

bands, becoming rather inconspicuous on 


ventral surfaces. Tail fan light brown with 

deep blue dots. 

Size: Maximum body length about 20 cm, commonly between 12 and 15 cm. 

Habitat, biology, and fisheries: Lives in coral and stone reefs from shallow waters to a depth of 20 m. 

Nocturnal and sometimes found in small groups, hiding inside crevices and undercuts of reefs during 

daytime. Fished throughout its range but nowhere abundant. A common incidental catch for fisheries of 

spiny lobsters and caught by divers (using hand and spear), or by dipnets or tangle nets. Often considered 

as a delicacy, comparable to spiny lobsters, and mainly sold live or fresh in local markets. 

Distribution: Worldwide in 

tropical seas and has been 

recorded in the Western 

Atlantic from the Caribbeans 

to Brazil, and in the Indo-West 

Pacific from the east coast of 

Africa to Taiwan Province of 

China, Hawaii, and French 

Polynesia. 



1038 Lobsters 



FAO names: En - Aesop slipper lobster. 

Diagnostic characters: Body vaulted, 

pubescent and covered with rounded 

tubercles. Eyes small and subspherical. 

Carapace somewhat rectangular, but 

posterior half distinctly wider than anterior 

half; cervical groove strong; pregastric, 

gastric and cardiac teeth all strongly 

protruding. Antennae broad, flattened and 

plate-like, with distal margin finely crenate. All 

legs without pincers and similar in size. 

Abdomen uniformly granulate and not 

particularly sculptured; dorsal midline of 

second and third segments strongly 

ridged, and that of fourth segment 

produced into a remarkable hump; pleura 

directed downwards, with posterior margin 

of second pleuron somewhat concave. 

Posterior half of tail fan soft and flexible. 

Colour: body and legs yellowish white and 

covered with purplish red patches; dorsal 

surfaces more purplish, ventral surfaces more 

yellowish. Eyes dark brown. Antennae with 

purplish margins; antennules somewhat 

orange except distal segment of peduncle and 

flagella purplish. First abdominal segment 

yellowish with 3 diffused purplish red 

spots. Soft part of tail fan light brown with 

numerous purple dots. 

Size: Maximum body length 50.5 cm, 

commonly between 16 and 30 cm; probably 

the largest species of the genus. 


coral or rocky reefs at depths from 10 to 

135 m, usually less than 50 m. Apparently 

nowhere abundant and in some places even 

uncommon. Despite of its large size therefore 

mostly incidentally taken during fishing 

activities for spiny lobsters (e.g. by hand or by lobster pots). When caught, this slipper lobster is highly 

esteemed as food and sold (usually live or fresh) in local markets at prices comparable to that of spiny 

lobsters. 


Pacific from Mauritius to the 

Red Sea, Japan, Hawaii, and 

Australia.




FAO names: En - Blunt slipper lobster; 

Fr - Cigale grenue; Sp - Cigarra ñato. 

Diagnostic characters: Body vaulted, 

pubescent and densely covered with 

rounded tubercles. Eyes small and 

subspherical. Carapace rectangular, with 

anterior half about as wide as posterior 

half; cervical groove shallow and 

constricted at middle; pregastric tooth 

slightly elevated, gastric tooth low and 

cardiac tooth almost levelled. Antennae 

broad, flattened and plate-like, with distal 

margin finely crenate. All legs without 

pincers and similar in size. Abdomen 

uniformly granulate and not particularly 

sculptured; dorsal midline of second to 

fourth segments weakly to moderately 

ridged and progressively higher 

posteriorly; pleura directed downwards, 

with posterior margin of second pleuron 

somewhat convex. Posterior half of tail fan 

soft and flexible. Colour: body red-brown 

with depressed areas and ventral surfaces 

rather pale yellowish. Eyes black-brown. 

Margins of carapace orange-red; antennae 

yellowish with orange-red margins; 

antennules bright yellow with purplish 

flagella. Legs bright yellow with regions near 

joints distinctly purplish. First abdominal 

segment with 3 distinct red spots, the 

central one much larger and anteriorly 

fused with lateral spots. Soft part of tail fan 

light brown with red-brown dots. 


commonly to about 20 cm. 

Habitat, biology, and fisheries: Nocturnal; inhabits coral and rocky reefs to a depth of about 80 m, mostly 

between 20 and 50 m. A common bycatch of fisheries for spiny lobsters and taken by hand during day and 

night diving, wire traps and tangle nets, but apparently nowhere abundant in the Western Central Pacific. 

Like other species of the genus, it is often considered as a delicacy and sold in local markets (fresh or live) 

at slightly lower prices than spiny lobsters. 


Pacific from the eastern coast 

of Africa to Japan, Australia, 

New Caledonia, and Hawaii.

1040 Lobsters 



FAO names: En - Flathead lobster; Fr - Cigale raquette; 

Sp - Cigarra chata. 

Diagnostic characters: Body markedly depressed, 

with surface pubescent and granulate. Carapace 

trapezoid, narrowing posteriorly, with weak 

branchial carinae and shallow cervical incisions; 

anterior part of lateral margin with only 2 teeth, 

posterior 3/4 without teeth. Eyes small and 

subspherical; orbits situated at anterolateral angles 

of carapace. Antennae broad, flattened and plate-like. 

All legs without pincers and similar in size. Abdomen 

with median carina and transverse grooves, fifth 

segment armed with a strong posteromedian spine; 

pleura directed downwards. Posterior half of tail fan 

soft and flexible. Colour: body brownish with reddish 

brown granules, ventral surface somewhat yellowish 

white. Eyes deep brown. Antennules yellowish white 

with red-brown bands. Legs yellowish white and distally 

brownish. Soft part of tail fan and pleopods somewhat 

orange-red. Setae light brown. 

Size: Maximum body length 25 cm, commonly to about 

16 cm; maximum weight over 0.5 kg. 

Habitat, biology, and fisheries: Inhabits bottoms of 

soft substrate, sand and/or mud, sometimes with shells 

or gravel; at depths from 8 to more than 200 m, but 

usually between 10 and 60 m. It buries into the soft 

substrate with only the eyes and antennules visible 

during daytime; actively swims during nocturnal 

foraging, which covers long distances. The planktonic 

larval stage lasts for about 3 months. Common 

throughout its range but nowhere very abundant. Forms 

a bycatch of trawling operations and is sometimes also 

taken by diving (with a catch of over 500 t in Australia 

between 1989 and 1990). Marketed fresh or frozen in 

fish markets and used for food. Similar to the species of 

Ibacus, it is generally considered to have a good taste 

(in Queensland this species is even ranked above 

Ibacus) and sold at slightly lower prices than spiny 

lobsters. 



from the east coast of Africa to 

the Red Sea, Japan, and the 

northern coast of Australia.



En - Royal Spanish lobster. 

Maximum body length up to 17 cm. In shallow 

waters at depths from 5 to 50 m on the outer 

edges of coral reefs. Nocturnal and hides in 

cavities during daytime. Too rare to be of 

significant importance to fisheries, but highly 

valued in the aquarium trade for its bright 

coloration. Indo-West Pacific from Mauritius, La 

Réunion, New Caledonia, Hawaii, and Easter 

Islands. 


En - Caledonian mitten lobster. 

Maximum body length about 18 cm. In shallow 

water to a depth of 6 m on reefs, usually on the 

exposed side and in surge channels. Hides in 

crevices during the daytime, often attached to 

the ceiling of caves. In Fiji, this species is often 

found in the same habitat with Panulirus 

penicillatus. Caught by divers with gloved 

hands and mainly for local consumption. 

Southern Pacific and known from Queensland 

(Australia), New Caledonia, Loyalty Islands, 

New Hebrides, Fiji, and Samoa. 


1042 Lobsters 


En - Red-spotted mitten lobster. 


water at depths from 1 to 5 m on the sandy 

bottoms of coral reefs. Used as food locally and 

collected at night with torches. Only known 

from the Society, Tuamotu, and Gambier 

Islands (French Polynesia). 


En - Marbled mitten lobster. 


reefs areas. Like other species of mitten 

lobsters, this species is used locally for human 

consumption. Central and South Pacific from 

Kapingamarangi through the Marshall, Gilbert, 

and Phoenix islands to the Marquesas. 

(from Forest, 1954) 




En - Two-spot locust lobster. 


between 3 and 5 cm. On soft bottoms at depths 

from 15 to 150 m, commonly between 40 and 

75 m. Caught incidentally during trawling 

operations. Like the other Indo-West Pacific 

species of the genus, it is generally considered 

to be of no economic importance due to its small 

size and limited abundance. Western Pacific 

and western Australia; from the East and South 

China Sea, Taiwan Province of China to Viet 

Nam, Thailand, the Philippines, Indonesia, 

northern and western Australia. 


En - Hunchback locust lobster. 


between 4 and 5 cm. On soft bottoms at depths 

from 20 to 60 m, rarely to 200 m. Commonly 

and incidentally taken by trawlers fishing for 

other species. Nowhere abundant and of limited 

commercial importance due to its small size. 

Indo-West Pacific from East Africa to the Red 

Sea, Japan, and northeastern Australia. 




CRABS 

by P.K.L. Ng

1046 Crabs 


anterolateral 

margin and teeth 

orbit 


region 


margin 

eye 

frontal view of body 

dactylus 

rostrum antennule 

carapace 

abdomen 

(folded under carapace) 

suborbital 

region 

pterygostomial 

region 

antenna 

palm 

general shape (dorsal view) of a brachyuran crab (family Portunidae) 

basal segment 

of antennule 

antennule 

epistome endostome 

orbit 

antennular 

fossa 

frontal 

margin 

posterior 

border 

flagellum of 

antenna 

eyestalk 

basal antennal 

segment 

eyes, antennae, and antennules (ventral view) 

cornea 

hand 

epistome 

palp 

(carpus, 

propodus, 

dactylus) 

cheliped 

4 

lateral tooth 

or spine 

3 

1 

2 

walking 

legs 1- 4 

front antennule 

buccal 

cavern 

mouth field 

ischium 

merus 

antenna 

eye 

exopod 

orbit 

3 rd 

maxilliped


outer 

margin 

coxa 

gut 

telson 

merus 

carpus 

fused 

basis-ischium 

cheliped 

3 

inner 

margin 

6 

5 

4 

inner carpal spine 

(or tooth) 

palm 

4 

abdomen of female 

pollex 

claw 

(or chela, pincer) 

teeth of cutting 

edge 

5 

6 

7 

dactylus 

finger 

tip 

thoracic 

sternites 

coxa 

merus 

basis ischium 

abdomen keel 

thoracic sternum and abdomen (ventral view) 

sternum 

female abdominal cavity and vulvae 

groove 

for 2 nd 

gonopod 

first gonopod 

distal part 

basal part 

abdominal cavity 

vulva 

abdomen 

male gonopods 

sternum 

gut 

carpus 

walking leg 

6 

5 

4 

3 

abdomen of male 

4 

5 

propodus 

telson 

6 

7 

8 

abdomen 

abdominal 

cavity 

abdomen 

dactylus 

locking mechanism 

for abdomen 

male abdominal cavity and gonopods 

second gonopod 

distal 

segment 

basal 

segment 

1 st gonopod 

penis (male 

genital opening) 

2 nd gonopod

1048 Crabs 


Introduction 

Like the shrimps and lobsters, crabs belong to the order Decapoda (= “ten-legged”, referring to the 10 

thoracic appendages normally present in these crustaceans). Crabs can be classified into 2 main 

groups, brachyuran crabs (infraorder Brachyura) and anomuran crabs (infraorder Anomura). Most species 

of Brachyura, or true crabs, can easily be separated from the so-called “false crabs” belonging to the 

infraorder Anomura by having 4 pairs of well-developed walking legs. Anomuran crabs always have only 3 

pairs of walking legs clearly visible, while the fourth (last) pair is very small, normally tucked under the body 

and hardly noticeable. However, this is just a general rule rather than a distinct separating character as 

there are a number of true crabs which have their fourth pair of legs greatly reduced as well (e.g. 

Dynomenidae and Retroplumidae) or even completely reduced (Hexapodidae). 

A more recent compilation of the actual number of all species of crabs known to date is pending. The last 

census was done by Fenner Chace Jr. (1951), who recorded worldwide 4 428 and 1 270 species of 

brachyuran and anomuran crabs, respectively. The late Raoul Serène (1968) estimated that perhaps some 

1 000 species of brachyuran crabs occur in the Indo-Malayan area. However, these numbers have 

substantially increased over the last 40 years, due to the rapid pace of crab discoveries. It is not 

unreasonable to believe that the current number of brachyuran and anomuran crabs in the world ranges 

from 5 000 to 6 000 and 1 500 to 2 000 species, respectively. Of these, the largest proportion is found in 

the Western Central Pacific, where around 1 500 to 2 000 brachyuran crab species (marine and fresh-water 

taxa) are probably present. 

The present contribution focuses on 15 families of brachyuran crabs and a single family of anomuran crabs 

which include commercially important species in the Western Central Pacific. The majority of edible crab 

species belong to the Brachyura, and accordingly, a large number of brachyuran crabs are caught for human 

consumption in the Western Central Pacific. It is important to note, however, that a much greater number 

of brachyuran crab species than listed here are collected for food by many poorer communities and 

indigenous people in the area. Any edible species which are common enough to be collected in great 

numbers can be eaten, even if they are small in size. To these belong many ocypodids such as soldier 

crabs (Dotilla spp.), fiddler crabs (Uca spp.), and periscope crabs (Macrophtalmus spp.), but also several 

medium-sized species of vinegar crabs (Sesarminae, Grapsidae). In addition, many medium-sized species 

of reef crabs of the families Xanthidae and Eriphiidae are locally consumed among natives of several Pacific 

islands. However, it is unrealistic to list and discuss every single species that is eaten once in a while or 

might be collected for food. Therefore, a selection has been made here of those species which at present 

have a distinct fishery value, are larger and more common, or have a good potential in the future as their 

fisheries develop. It is also worth noting that several species of fresh-water crabs of the families Potamidae 

and Parathelphusidae are consumed in many parts of Southeast Asia and Indo-China. 

In contrast to the brachyurans, few anomuran crabs have a major fishery value in the Western Central 

Pacific, with a single species (the “coconut crab”, Birgus latro) being of distinct commercial importance. 

The stone crabs (Lithodidae) are represented by several species within the area, but none of them are 

harvested so far, although some species are utilized for food in other regions of the world (the best known 

example is the large fishery for the “Alaskan king crab”, Paralithodes camtschaticus, in the northern Pacific). 

The lithodid species occurring in the Western Central Pacific, however, are generally too rare to show any 

significant commercial importance, although it may be possible that a number of species can be utilized in 

the future. Reports that some large hermit crabs (Paguridae and Diogenidae) are sometimes caught for 

food are actually not very reliable, and almost certainly none of these show any commercial importance. 

Many species of land hermit crabs (genus Coenobita), however, are regularly collected for the pet trade. 

The so-called squat lobsters (Anomura: Galatheidae), which actually have a more crab-like than lobster-like 

appearance, are represented by a few edible species in the Atlantic, but none of the species in the Western 

Central Pacific are large or common enough to have any food value. The same is true for the deep-water 

chirostylids (deep-water squat lobsters). However, because of their crab-like shape, galatheids and 

chirostylids have been included in the present key to families of marine crab-like Anomurans. On the other 

hand, several anomurans of clearly lobster- or shrimp-like appearance, such as the mud lobsters (Thalassina 

spp., Thalassinidae, notably T. anomala) and mud shrimps (Upogebia spp., Upogebiidae), are 

occasionally caught for food in the Western Central Pacific.


Poisonous Crabs 

Although poisonous crabs have been known for a very long time, only in recent years have the necessary 

biochemical studies been done to quantify and qualify the toxins involved. Some people become violently 

sick after consumption of crabs because of allergic responses, a response not related to poison. In general, 

there are 2 categories of poisonous crabs: 

The first category includes the permanent highly toxic species. These crabs are always poisonous, and 

include taxa such as the “mosaic crab” (Lophozozymus pictor), “demon crabs” (Demania spp.), “jewel crab” 

(Zosimus aeneus), “crested reef crab” (Platypodia granulosa), and “green egg crab” (Atergatis floridus). 

The consumption of any of these crabs, even if well cooked, is extremely dangerous and has proved fatal 

in several instances. It is important to note here that all these species belong to the family Xanthidae and 

they all have distinctive colour markings or striking colours, presumably to warn potential predators. All 

species are of moderately large size, reaching carapace widths from 7 to 10 cm, and as such, may be 

picked up by fishermen or collectors. The toxins that have been identified include palytoxins, saxitoxins, 

and tetrotoxins, and occur throughout the tissues and exoskeleton, being most concentrated in the liver 

and gonads. All these toxins act on the nervous system. As they loose their toxins when kept in captivity 

and fed on normal food, it is believed that the crabs obtain these toxins directly or indirectly from the food. 

The 2 most notorious genera are Lophozozymus and Demania, and a number of human deaths have been 

attributed to them. Tests on Lophozozymus pictor have shown that, although the degree of toxicity varies 

from individual to individual, they all contain enough toxins to kill an adult human. In a single analysis, 1 g 

of the crab’s flesh contained enough toxins to kill 42 000 mice. A large specimen of L. pictor, however, can 

easily reach a weight of 100 g. This makes it one of the (if not THE) most poisonous crabs known. Not all 

species of Lophozozymus and Demania have been analyzed biochemically, but the general consensus is 

that most, if not all their members are highly toxic. 

The second category of toxic crabs are those which are mildly poisonous and/or occasionally poisonous. 

The consumption of such crabs may cause illness but rarely death. The species involved here include “reef 

crabs” (Carpilius spp., Carpiliidae), “red-eyed crabs” (Eriphia spp., Eriphiidae), coral reef crabs like Etisus 

spp. and Atergatis spp. (Xanthidae) and “land crabs” (Cardisoma spp., Gecarcinidae). In most instances, 

these crabs are also not always poisonous, with their toxicity varying with place and time of year. This is 

very likely to be associated with the food habits of the crabs. In some cases, this is because the crabs have 

consumed poisonous fruits or leaves (e.g. for land crabs like Cardisoma). Poisonous crabs have also been 

associated with red-tide algal or dinoflagellate blooms. Species like the “red egg crab” (Atergatis integerrimus) 

are probably poisonous because they only occasionally feed on organisms which are toxic and only 

in small quantities. This second category of poisonous crabs poses problems for fishery officers as a 

species which is poisonous in one area may be totally harmless in another. Obviously, great care has to 

be taken in harvesting and consumption of those species. 

Zosimus aeneus Lophozozymus pictor 

Atergatis floridus Demania cultripes 

(after Garth and Alcala, 1977)

1050 Crabs 

Platypodia granulosa 

(after Garth and Alcala, 1977) 

Demania toxica 

Notes on the classification of brachyurans 

The total number of families of Brachyura is still undetermined. Although many authors still follow the 

classification presented by Balss (1957), more recent studies by Guinot (1978, 1979) have shown that this 

system is artificial. Unfortunately, not all the brachyuran families currently recognized were dealt with by 

Guinot in detail and the status of a number of them remains unresolved. For the present report, the writer 

has essentially adopted Guinot’s (1978) system of higher classification. Accordingly, 53 families are 

recognized, following mainly Guinot (1978) and Manning and Holthuis (1981). Out of these, 8 families are 

found in fresh water only and thus are outside the scope of the present contribution. Nevertheless, it is 

relevant to note that out of these 8 fresh-water families, 3 occur in the Western Central Pacific, namely the 

Potamidae (= Isolapotamidae), Gecarcinucidae, and Parathelphusidae (= Sundathelphusidae) (Ng, 1988). 

Of the 45 marine families, 40 have been recorded in the Western Central Pacific thus far, with only the 

Orithyiidae, Thiidae, Cheiragonidae, Pirimelidae, and Platyxanthidae apparently being absent from the 

area. 

Some of the families recognized here have undergone nomenclatural changes. The Eriphiidae has 

previously been known as the Menippidae and Oziidae. However, Eriphiidae is the oldest name and thus 

has nomenclatural priority. The Mimilambridae Williams, 1979, is considered a junior synonym of Parthenopidae 

MacLeay, 1838 (see Ng and Rodriguez, 1986). The recognition of a separate family for the 

Eumedonidae, symbionts on echinoderms, follows Stevcic et al. (1988). Stevcic (1988) recognized a 

separate family, Cheiragonidae, for crabs previously classified in the Telmessinae (Atelecylidae). Finally, 

the Camptandriidae, previously considered to be a subfamily of the Ocypodidae, is regarded here as a 

separate family, following the suggestions of Harminto and Ng (1991). 

Characters useful for identification 

The teeth of the anterolateral margins of the carapace are also known as the epibranchial teeth. The first 

anterolateral tooth is often called the “external orbital” or “exo-orbital” angle (or tooth) and is counted 

separately from the following anterolateral teeth by many authors (but not here). The frontal margin (or 

front) becomes elongate and/or spiniform in many crabs such as the homolids (deep-water porter crabs) 

and majids (spider crabs), and is then frequently called a rostrum. 

The maximum carapace width is used as principal measurement indicating the size of a crab, measured 

as the greatest distance between the lateral margins of the carapace. 

The buccal cavern (location of the mouthparts), is bordered on both sides by the pterygostomial regions, 

and above by the epistome. The calcareous plate inside the buccal cavern is called the endostome.Usually, 

only the anterior part of the endostome is visible, even when the mouthparts are moved aside. The outer mouth 

parts or third maxillipeds are often just referred to as “the mouthparts”, even though there are actually 6 

pairs of feeding appendages. Underneath the third maxillipeds, the second maxillipeds and first maxillipeds 

are located, normally covered by the third maxillipeds in life.Two smaller feeding appendages are situated below 

the 3 pairs of maxillipeds: the first maxilla (or maxilla) and second maxilla (or maxillules). Finally, the mouth is 

bordered by a pair of well-calcified, jaw-like, and highly modified appendages, the mandibles. 

The 5 pairs of locomotory appendages of a crab (the pereiopods) are made up of a pair of usually powerful 

chelipeds (legscarryingachela or pincer) and normally of 4 pairs of walking (or ambulatory) legs.For 

the present contribution, the first appendage is referred to as the cheliped and the last 4 appendages 

(walking legs) as legs. The claw (or chela) itself consists of a palm (or manus) and 2 fingers, one of which 

is movable (the dactylus or movable finger), whereas the other one (pollex) is fixed. The tips or edges 

of the fingers may be pectinated. In some families the last pair or all walking legs are modified for swimming 

or burrowing, as seen in the Portunidae and the Matutinae (the latter a subfamily of the Calappidae).


Adult male and female crabs are easily distinguished by the shape of their abdomen. In males, the abdomen 

is triangular to broadly T-shaped, whereas in females it is broad, usually semicircular, often covering most part 

of the ventral surface. Almost all crabs have 7 abdominal segments (although the seventh segment or telson 

is actually not a true segment), but in a number of families, several segments are partially or completely fused. 

This fusion may be complete (i.e. with the sutures between segments no longer visible) or incomplete (i.e. with 

parts of the sutures still present or obscure). In both cases, however, the segments are immovable. 

Many crab species show a sexual dimorphism, with the males usually being larger or possessing special 

or excessively developed structures. In some species, however, it is the female which grows larger. Males 

possess 2pairsof gonopods, that is, modified pleopods specifically adapted for copulation (most crabs 

practice internal fertilization). The pleopods (abdominal appendages) of females are branched, setose and 

serve to carry the eggs: fertilized eggs are exuded, attached to the setose pleopods of females, and kept 

there for several weeks until the planktonic larvae (zoeae) hatch out. The larvae pass several stages before 

they finally metamorphose to a young crab. 

Many species of crabs possess pubescence to varying degrees on their body and appendages. The hair 

(or more appropriately called setae) may be soft or stiff, simple or plumose (plume-like), or so short that 

it becomes pile-like, sometimes even short and dense, giving a velvet-like appearance. The setae may 

sometimes be hard and spine-like, especially on the propodus and dactylus of legs. Unlike real spines, 

however, those stiff setae are never calcareous. Majids often possess hook-like setae that attach to 

sponges, algae, and debris (similar in action to velcro), supporting the camouflage of the crab. In other 

species, the longer and/or plumose setae gather dirt and mud in order to obscure the animal’s outline. 

Most of the softer setae on the legs and chelae have a sensory function. 

Carapace types 

The shape of the carapace is often used as a descriptive character in many guides and keys. Unfortunately, 

a large variety of terms have been introduced in the past, not always applied with exactly the same meaning. 

Therefore, an approximate categorization has been attempted here and those carapace types which belong 

to a respective category are illustrated below. It should be remembered, however, that there are sometimes 

no clear lines separating the different carapace types, and so the designation of a particular type may be 

somewhat subjective in certain cases. Nevertheless, the use of carapace shapes is still a useful character 

in many instances. 

The carapace types utilized here are shown in Figures A to N. 

longitudinally rectangular 

Figure A 

transversely rectangular 

Figure B 

trapezoidal 

Figure D 

squarish 

Figure C

1052 Crabs 

pentagonal 

Figure E 

transversely hexagonal 

Figure G 

transversely ovate 

Figure H 

hexagonal 

Figure F


longitudinally ovate 

Figure I 

transversely subovate 

Figure J 

triangular 

Figure K

1054 Crabs 

circular 

Figure L 

subcircular 

Figure M 

pyriform 

Figure N

Imported Crabs of Commercial Importance 1055 

Imported Crabs of Commercial IMPORTED Importance CRABS OF COMMERCIAL IMPORTANCE 

Several species of non-Western Central Pacific crabs are regularly imported into the area, notably to 

Singapore, Malaysia, and Thailand. They command very high market values and are popular not only 

among locals but also the relatively large expatriate community (especially Japanese) in these countries. 

In most cases, they are brought in alive for better value and for the live-seafood restaurant trade.Therefore, 

one would probably frequently encounter these species in markets or retailers. The main species imported 

are “Chinese mitten or hairy crabs” (Eriocheir sinensis, E. hepuensis), “Japanese mitten crab” (E. 

japonicus), “giant Tasmanian crab” (Pseudocarcinus gigas), “snow crab” (Chionoecetes opilio and C. 

japonicus), “queen crab” (Erimacrus isenbeckii), and “Alaskan king crab” (Paralithodes camtschaticus). 

Eriocheir sinensis Eriocheir hepuensis 

(from Shen, 1932) 

Eriocheir japonicus 

(from De Haan, 1833) 

Chionoecetes opilio 

(after Kobyakowa, 1955) 

Pseudocarcinus gigas 

Erimacrus isenbeckii 


Paralithodes camtschaticus (male) Paralithodes camtschaticus (female) 




1056 Crabs 

Guide toGUIDE FamiliesTO 

FAMILIES OF INTEREST TO FISHERIES OCCURRING IN THE AREA 

HOMOLIDAE Page 1083 

Deep-water carrier crabs 

Carapace longitudinally rectangular; dorsal surface 

granulose to spinose; front narrow, usually with 3 

long horn-like projections (rostra). Male chelipeds 

long. Last pair of legs inserted obliquely on carapace 

and directed upwards, reduced, subchelate to 

chelate, modified to carry sponges. All male abdominal 

segments distinct, movable. 

DROMIIDAE Page 1085 

Sponge crabs 

Carapace circular to hexagonal; dorsal surface gently 

to strongly convex longitudinally and transversely, 

smooth or granular, usually setose; front narrow, 

usually entire; anterolateral margins of carapace 

strongly convex, unarmed, or with small spines. Last 

2 pairs of legs inserted obliquely 

on carapace and directed upwards, 

strongly reduced, subchelate, 

modified to carry 

sponges or tunicates on back of 

carapace. All male abdominal 

segments distinct, movable; a 

small platelet-like structure usually 

intercalated between edges 

of sixth abdominal segment and 

telson. Male first gonopod stout, 

simple; male second gonopod 

long, usually subequal or longer 

than length of male first gonopod. 

Male and female genital openings 

sternal. 

male 

abdomen 

RANINIDAE Page 1089 

Spanner crabs 

Carapace longitudinally ovate; dorsal surface 

strongly granulose or squamose to smooth; 

front triangular, narrow. Third maxillipeds very 

narrow, merus distinctly triangular. Eyestalks 

long, longer than front. Fingers of chela 

strongly bent; at least 1 pair of legs with last 2 

or 3 segments paddle-like. Thoracic sternum 

very narrow, especially sternites 5 to 7. All 

male abdominal segments distinct, movable. 

6 

5 

4 

3 

plate-like 

structure 

triangular 

merus 

right 3 rd 

maxilliped 

last 2 pairs of legs 

subchelate 

narrow front 

(or rostrum) 

carapace 

longitudinally 

rectangular 

last legs 

reduced, 

subchelate 

to chelate 

carapace 

circular to 

hexagonal 

fingers 

strongly 

bent 

carapace 


ovate 

at least 1 pair of legs paddle-like

Guide to Families 1057 

CALAPPIDAE Page 1091 

Box and moon crabs 

Carapace circular, ovate to transversely ovate and subovate, sides of carapace may be expanded to 

form a clypeiform process (= expanded posterior edge). Merus of third maxillipeds distinctly triangular; 

opening for afferent respiratory current at base of chela, no canal present along sides of buccal cavern 

even when third maxillipeds pushed aside. Larger chela may have specialized cutting tooth for cutting 

gastropod shells. Propodus and dactylus of legs may be paddle-like. Male abdominal segments 3 to 5 

completely fused; male genital openings always coxal. 

subfamily 

Calappinae 

XANTHIDAE Page 1098 

Stone and mud crabs 

Carapace hexagonal, transversely hexagonal to transversely 

ovate, sometimes circular; dorsal surface usually 

ridged or granulose;frontal margin usually notched medially; 

usually 2 to 6 spines, teeth and/or lobes on each anterolateral 

margin. Longitudinal ridges which define efferent respi- 

ratory current usually absent or strong only on posterior part 

of endostome, ridges not visible on anterior 

part of endostome when mouthparts 

pushed aside. Fingers of chela may be 

spoon-tipped. Legs varying in structure; 

propodus and dactylus may show a special 

dactylo-propodal articulation. Male abdominal 

segments 3 to 5 immovable, fused 

completely or incompletely. Male first 

gonopod slender, slightly sinuous; male 

second gonopod very short, less than 1/4 

the length of male first gonopod. 

right chela with 

specialized tooth 

expanded 

“clypeiform 

process” 

segments 3-5 

fused (completely 

or incompletely) 

6 

5 

4 

3 

2 

1 

male abdomen 

ERIPHIIDAE Page 1103 

Stone and mud crabs 

Carapace hexagonal, transversely rectangular to transversely 

ovate; dorsal surface ridged or granulose; frontal 

margin notched medially; 4 teeth and/or lobes on each 

anterolateral margin.Legs normal.Longitudinal ridges which 

define efferent respiratory current well developed along entire 

endostome, ridges visible on anterior part of endostome 

when mouthparts pushed aside. All male abdominal segments 

distinct, movable. Male first gonopod stout, almost 

straight or gently curved; male second gonopod elongate, 

longer or subequal in length to male first gonopod. 

subfamily 

Matutinae 

all male abdominal 

segments freely 

movable 

often a 

long 

lateral 

spine 

paddle-like legs

1058 Crabs 

CARPILIIDAE Page 1110 

Reef crabs 

Carapace transversely ovate; dorsal surface 

smooth, distinctly convex longitudinally and transversely; 

front entire; a single low, small tooth on 

each anterolateral margin. Legs simple. Longitudinal 

ridges which define efferent respiratory 

current usually absent or 

strongly developed on posterior part 

of endostome only; ridges not clearly 

visible on anterior part of endostome 

when mouthparts pushed aside. 

Male abdominal segments 3 to 5 immovable, 

completely fused. Male 

first gonopod stout, almost straight 

or gently curved; male second 

gonopod elongate, longer or subequal 

in length to male first gonopod. 

male 

abdomen 

PILUMNIDAE Page 1112 

Hairy crabs 

Carapace hexagonal, transversely rectangular or 

transversely ovate; dorsal surface convex, smooth 

to granulated; frontal margin entire to multilobate; 

usually 1 to 4 teeth or lobes on each anterolateral 

margin. Longitudinal ridges defining efferent respiratory 

current usually well developed along entire 


when mouthparts pushed aside. Legs normal. 

Male abdominal segments 3 to 5 freely movable. 

Male first gonopod slender, S-shaped, distal part 

simple; male second gonopod very short, sigmoid. 

GONEPLACIDAE Page 1114 

Rhomboid crabs 

Carapace hexagonal, transversely rectangular, 

trapezoidal, or transversely ovate; dorsal surface 

convex, usually smooth; frontal margin usually entire, 

sometimes multilobate; anterolateral margin 

usually armed with 1 to 4 teeth or lobes, or entire. 

Male abdominal segments 3 to 5 distinct, movable 

or fused and immovable. Male first gonopod moderately 

stout, gently curved or sinuous; male second 

gonopod relatively short to elongate, but usually 

shorter than male first gonopod. 

6 

5 

4 

3 

2 

1 

segments 3-5 

completely 

fused 

male chelipeds elongate 

anterolateral margin 

with a single tooth

Guide to Families 1059 

PORTUNIDAE Page 1115 

Swimming crabs 

Carapace hexagonal, transversely 

ovate to transversely hexagonal, 

sometimes circular; dorsal surface 

relatively flat to gently convex, 

usually ridged or granulose; front 

broad, its margin usually multidentate; 

usually 5 to 9 teeth on each 

anterolateral margin of carapace; 

posterolateral margins usually distinctly 

converging. Legs laterally 

flattened to varying degrees, last 2 

segments of last pair paddle-like. 

Male abdominal segments 3 to 5 

completely fused, immovable. 

male 

abdomen 

GERYONIDAE Page 1132 

Golden crabs 

Carapace hexagonal; dorsal surface relatively smooth 

to granular; frontal margin with 4 teeth; anterolateral 

margins distinctly convex, each with 3 to 5 low, sometimes 

indistinct teeth. Dactylus of walking legs Tshaped 

in cross-section. Male abdominal segments 3 

to 5 fused, functionally immovable, but sutures still 

visible. 

MAJIDAE Page 1136 

Spider crabs 

Carapace pyriform, circular to subovate, anterior 1/2 

to 1/3 usually distinctly narrower than posterior part; 

dorsal surface gently convex, spinulose, granulose 

and/or ridged; front narrow, often with 2 long horn-like 

projections (rostra); orbits poorly developed to absent; 

anterolateral margins of carapace often armed with 

well-developed spines. Legs spinulose and/or granulose, 

often with stiff setae. All male abdominal segments 

usually freely movable in most species. 

5 

4 

3 

segments 

3-5 fused 

male abdominal segments 

3-5 immovable, but 

sutures visible 

orbits 

incomplete 

last legs 

paddle-like 

often 2 horn-like 

projections 


with 5-9 teeth 

carapace 

hexagonal 

dactylus 

T-shaped in 

cross-section 


carapace

1060 Crabs 

GRAPSIDAE Page 1138 

Sally-light-foots, vinegar crabs, and paddler crabs 

Carapace squarish, transversely rectangular, trapezoidal or circular; dorsal surface flat to gently 

convex, with low oblique or transverse ridges; front much broader than orbits; orbits occupying almost 

entire anterior border (excluding front); antero- and posterolateral margins of carapace usually not 

clearly demarcated, lateral margins appearing almost straight or gently convex, usually armed with 1 

or 2 teeth anteriorly. Rhomboidal gap usually present between third maxillipeds; mandibles often 

exposed. Dactylus of legs with distinct spines. Male abdominal segments 3 to 5 freely movable in most 

species. 

subfamily 

Grapsinae 

carapace 

circular 

GECARCINIDAE Page 1147 

Land crabs 

Carapace circular to transversely ovate; dorsal surface 

smooth, strongly convex longitudinally and 

transversely; frontal margin entire; anterolateral 

margins unarmed or each with a single tooth; rhomboidal 

gap present between the third maxillipeds. 

Legs stout, dactylus longitudinally ridged, often 

with dense, stiff setae, margins with spines. All male 

abdominal segments distinct, movable. 

OCYPODIDAE Page 1152 

Ghost crabs 

Carapace squarish, transversely rectangular, 

trapezoidal or transversely ovate; dorsal surface 

gently convex, usually smooth or with grooves; frontal 

margin entire, relatively narrow; orbits broad, 

occupying almost entire anterior border (excluding 

the front), antero- and posterolateral margins of 

carapace usually not clearly demarcated, lateral 

margins appearing almost straight or gently convex, 

lateral margins unarmed. Eyestalks long, 

longer than width of orbit. No rhomboidal gap between 

third maxillipeds. Dactylus of legs with numerous 

stiff setae. Ventral surface of abdomen or 

base of legs may have tufts of fine setae. All male 


subfamily 

Sesarminae 

carapace 

squarish 

very narrow 

front long orbits

Guide to Families/Key to Brachyura 1061 

COENOBITIDAE Page 1154 

Land hermit crabs and coconut crabs 

Carapace relatively well calcified; eyestalk laterally 

flattened; eyes usually held subparallel to each 

other. Antennae laterally flattened. Coxae of third 

maxillipeds close to each other, without distinct 

gap between them. Chelipeds short, stocky, equal 

or unequal; when unequal, left chela larger. First 2 

pairs of walking legs well developed, last 2 pairs 

reduced, third legs chelate. Abdomen bilaterally 

asymmetrical, not clearly divided into segments. 

Either hermit crabs or distinctly crab-like animals 

with abdomen tucked under carapace; uropods 

modified into a “rasp” used for clinging interior of 

gastropod shells (except in Birgus latro). 

Guide Key to to Brachyura Families/Key to KEY Brachyura TO THE FAMILIES OF BRACHYURAN CRABS 

Note: the following key covers all marine families of brachyuran crabs recognized in this work, most of which 

have been reported from the Western Central Pacific. The 5 families which so far are known only from other 

regions of the world have been included in the anticipation that some of them may be recorded in the future 

from the area. Wherever possible, external and easily viewed characters are utilized, and in most cases, 

no smaller structures have been chosen, such as male gonopods and structures of male and female 

genitalia (which are very important in crab classification). Due to the diversity in some families (e.g. 

Xanthidae and Goneplacidae), not all their members can be identified to the family level with this key, 

although it should work for the majority of species encountered. There are also a number of unusual species, 

the familial classification of which is still contentious. For a more comprehensive key, see Sakai (1976) and 

Dai and Yang (1991). For the identification of some more difficult species, the only safe way is to send the 

sample(s) to an expert of decapod taxonomy. 

1a. Only 3 pairs of legs visible, fourth 

pair absent (Fig. 1) . . . . . . . . Hexapodidae 

1b. Four pairs of legs visible . . . . . . . . . . → 2 

2a. Basal segment of eyestalk much 

longer than terminal article, from 

dorsal view, eyestalk appears to be 

2-segmented (Figs 2a and 3) . . . Latreillidae 

2b. Basal segment of eyestalk much 

shorter than terminal article, from 

dorsal view, eyestalk appears to be 

unsegmented (Fig. 2b) . . . . . . . . . . . → 3 

basal 

article 

very long 

basal article 

very short 

a) Latreillidae b) other crabs 

Fig. 2 eyestalk 

last leg strongly 

reduced 

Fig. 1 Hexapodidae 

Fig. 3 Latreillidae 

antenna laterally 

flattened 

eye laterally 

flattened 

3 rd leg 

chelate

1062 Crabs 

3a. Fourth (last) last pair of legs distinctly subchelate to chelate (Fig. 4a-f) or strongly 

reduced to just 3 articles, inserted obliquely on carapace and directed upwards . . . . . . . . → 4 

3b. Fourth (last) last pair of legs normal in structure or reduced in size but not subchelate 

or chelate (Fig. 4g) and never reduced to just 3 articles, inserted laterally on carapace 

and directed laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

subchela 

a) b) c) d) e) f) g) 

Fig. 4 last leg: (a-f) various types of subchelate to chelate conditions of propodus and dactylus; (g) normal leg 

4a. Merus of third maxilliped distinctly triangular 

in shape (Fig. 5a) . . . . . . . . . . . → 5 

4b. Merus of third maxilliped quadrate to 

subquadrate, never clearly triangular 

in shape (Fig. 5b, c). . . . . . . . . . . . . . . → 7 

5a. Third maxillipeds not covering a large 

anterior part of buccal cavern which is 

normally filled by the exposed second 

maxillipeds (Fig. 6a); crab carries inanimate 

objects (e.g. shells and 

leaves) or sea anemones (Fig. 7) . . . Dorippidae 

a) b) 

Fig. 5 third maxilliped 

c) 

5b. Third maxillipeds covering or almost completely covering anterior part of buccal cavern, 

second maxillipeds always covered and not exposed (Fig. 6b); crab usually carries 

pieces of inanimate objects (e.g. dead shells and sticks) when alive . . . . . . . . . . . . . . . → 6 

buccal cavern not covered 

by 3 rd maxillipeds anteriorly 

a) Dorippidae b) Cyclodorippidae 

Fig. 6 mouth field 

buccal cavern covered by 

3 rd maxillipeds anteriorly 

6a. Carapace hexagonal to subovate 

(Fig. 9); orbits distinct; exopod of 

third maxilliped usually without 

flagellum (Fig. 8a) . . . . . . Cyclodorippidae 

6b. Carapace rectangular to squarish 

(Fig. 10); orbits absent; exopod of 

third maxilliped with distinct flagellum 

(Fig. 8b) . . . . . . . . Cymonomidae 

triangular 

merus 

a) Cyclodorippidae 

quadrate 

merus 

subcircular (or 

subovate) merus 

Fig. 7 Dorippidae 

(after Shen, 1932) 

exopod with flagellum 

exopod 

without 

flagellum 

b) Cymonomidae 

Fig. 8 third maxilliped

Key to Brachyura 1063 

Fig. 9 Cyclodorippidae Fig. 10 Cymonomidae 

7a. Carapace pyriform (pear-shaped) (Fig. 12); orbits incomplete; carapace, chelipeds, and 

legs often with hooked setae; vulvae of adult female on thoracic sternum (Fig. 11a) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Majidae (in part) (p. 1136) 

7b. Carapace shape not as above; orbits usually complete; carapace, chelipeds, and legs 

without hooked setae; vulvae of adult female on coxae of third legs (Fig. 11b) . . . . . . . . . . → 8 

vulva on 

sternum 

vulva on coxa 

of 3 rd leg 

a) Majidae b) others 

Fig. 11 position of female vulvae on ventral side of body Fig. 12 Majidae 

(abdomen omitted) 

8a. Carapace longitudinally rectangular, dorsal surface glabrous or with scattered stiff 

setae; only fourth pair of legs with dactylus and propodus subchelate to chelate (Fig. 13) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homolidae (p. 1083) 

8b. Carapace longitudinally ovate, circular or hexagonal, dorsal surface usually with dense, 

soft setae; both third and fourth legs with dactylus and propodus subchelate to chelate; 

carries sponges and sea anemones when alive . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

9a. Carapace circular to hexagonal; a small platelet-like structure usually intercalated 

between edges of sixth abdominal segment and telson (Fig. 14a); crab carries sponges, 

tunicates, and bivalve shells (Fig. 15) . . . . . . . . . . . . . . . . . . . . . Dromiidae (p. 1085) 

9b. Carapace longitudinally ovate; no platelet-like structure intercalated between edges of 

sixth abdominal segment and telson (Fig. 14b); crab believed to carry sponges or related 

objects (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Homolodromiidae 

Fig. 13 Homolidae 

telson 

6 

5 

4 

3 

platelet-like 

structure 

a) b) 

Fig. 14 male abdomen 

6 

5 

4 

3

1064 Crabs 

Fig. 15 Dromiidae 

10a. Merus of third maxilliped distinctly triangular 

in shape (Fig. 17a) . . . . . . . . . . . . . . . → 11 

10b. Merus of third maxilliped quadrate to ovoid, 

never distinctly triangular in shape 

(Fig. 17b) . . . . . . . . . . . . . . . . . . . . . → 14 

11a. Carapace longitudinally ovate; sternum 

very narrow, thoracic sternites 5 to 7 very 

narrow (Fig. 18a); fingers of chela strongly 

bent (Fig. 19) . . . . . . . . . . . Raninidae (p. 1089) 

11b. Carapace shape not as above; sternum 

normal, thoracic sternites 5 to 7 not 

strongly narrowed (Fig. 18b); fingers of 

chela not strongly bent . . . . . . . . . . . . . . → 12 

very 

narrow 

sternites 

5-7 

coxa of 

cheliped 

sternites 

5-7 

coxae of 

legs 

coxa of 

cheliped 

coxae 

of 

legs 

a) Raninidae b) other crabs 

Fig. 16 Homolodromiidae 

(after Alcock and Anderson, 1899) 

narrow, 

triangular 

merus 

a) Raninidae b) others 

sternites 

5-7 


quadrate 

merus 

Fig. 18 thoracic sternum 

Fig. 19 Raninidae 

(ventral side of body, between legs) 

12a. Carapace subcircular with strong teeth on lateral margins; only dactylus of fourth leg 

paddle-like (Fig. 21); vulvae of adult female exposed, not covered by abdomen 

(Fig. 20a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orithyiidae 

(family not recorded from the area so far) 

12b. Carapace either of differing shape, or, if subcircular, never with strong teeth on lateral 

margins; dactylus of fourth leg normal and styliform, or, if paddle-like, all other dactyli 

of legs are also similarly structured; vulvae of adult female always covered by abdomen 

(Fig. 20b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 13 

abdomen 

vulva 

(covered by 

abdomen) 

a) 

vulva (not 

covered by 

abdomen) 

b) 

Fig. 20 position of female vulvae 

paddle-like 

Fig. 21 Orithyiidae 

(after Shen, 1932)


13a. Opening for afferent respiratory current below orbits, adjacent to endostome, with 

distinct canal present along sides of buccal cavern (Fig. 22a) when third maxillipeds 

pushed aside; sides of carapace never expanded to form a clypeiform process (= expanded 

posterior edge); larger chela never with specialized cutting tooth; legs never 

paddle-like (Fig. 23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leucosiidae 

13b. Opening for afferent respiratory current at base of chela, no canal present along sides 

of buccal cavern even when third maxillipeds pushed aside (Fig. 22b); sides of carapace 

may be expanded to form a clypeiform process; larger chela may have a specialized 

cutting tooth (Fig. 24); propodus and dactylus of legs may be paddle-like (Fig. 25) . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calappidae (p. 1091) 

3 rd 

maxilliped 3rd respiratory maxilliped 

canal 

a) Leucosiidae b) others 

Fig. 22 presence or absence of respiratory canal 


Fig. 24 right chela (Calappidae) 

no respiratory 

canal 

Fig. 23 Leucosiidae 


a) subfamily Calappinae b) subfamily Matutinae 

Fig. 25 Calappidae 

14a. A small platelet-like structure always intercalated present between edges of sixth 

abdominal segment and telson (Fig. 26a); fourth leg strongly reduced, present only as 

a short appendage (only partially subchelate in 1 Atlantic species); no known carrying 

behaviour (Fig. 27). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dynomenidae 

14b. No platelet-like structure intercalated between edges of sixth abdominal segment and 

telson (Fig. 26b); fourth leg reduced but always distinct, with most segments slender, 

relatively long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 15 

6 

5 

4 

3 

platelet-like 

structure 

a) b) 

6 

5 

4 

3 

Fig. 26 male abdomen Fig. 27 Dynomenidae

1066 Crabs 

15a. Fourth leg strongly reduced compared to other legs (Fig. 28a) . . . . . . . . . . . . . . . . . → 16 

15b. Fourth leg subequal to other legs, or if smaller, not greatly reduced in size compared to 

third leg (Fig. 28b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 17 

last leg 

strongly 

reduced 

a) b) 

16a. Carapace circular; dorsal surface 

granulated; fourth leg reduced but 

normal in structure, dactylus 

curved, tapering to a sharp point 

(Fig. 29) . . . . . . . . . . . . . . . . Palicidae 

16b. Carapace hexagonal, dorsal surface 

usually with 1 or 2 distinct 

transverse ridges; fourth pair of 

legs very slender, almost filamentlike, 

often appearing feather-like, 

dactylus straight, with rounded tip 

(Figs 30 and 31) . . . . . . . . Retroplumidae 

Fig. 30 Retroplumidae (ventral view) 

(from Alcock and Anderson, 1899) 

Fig. 28 last 2 pairs of legs 

last leg 

feather-like 

dactylus of 

last leg 

Fig. 29 Palicidae 


Fig. 31 Retroplumidae (dorsal view) 


last leg 

normal 

17a. Carapace longitudinally rectangular (Fig. 33); female genital opening on coxa of third 

walking leg (Fig. 32a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poupiniidae 

17b. Carapace not longitudinally rectangular; female genital opening on sternum (Fig. 32b) . . . . → 18 

sternum 

vulva on coxa 

of 3 rd leg 

vulva on 

sternum 

a) b) 

sternum 

Fig. 32 position of female vulvae Fig. 33 Poupiniidae


18a. Antennae (excluding basal segment) strongly reduced (Figs 34a and 35) . . . . . . . . . Belliidae 

18b. Antennae (excluding basal segment) normal, with distinct flagellum (Fig. 34b, c). . . . . . . . → 19 

small antenna 

well-developed antenna 

a) Belliidae 

b) Majidae 

Fig. 34 front and antenna 

19a. Fossae (sockets) for antennulae squarish 

to longer than broad, antennulae fold 

longitudinally or almost so (Fig. 36a, b) . . . . → 20 

19b. Fossae for antennulae broader than 

long, antennulae fold transversely or 

obliquely (Fig. 36c, d) . . . . . . . . . . . . . . → 27 

a) 

antennules folding 

vertically 

20a. Carapace pyriform, subpyriform, triangular, 

circular, or subcircular; orbits incomplete 

to absent (Fig. 37a) . . . . . . . . . → 21 

20b. Carapace longitudinally and transversely 

ovate, hexagonal, circular, or 

subcircular; orbits complete (Fig. 37b) . . . . → 22 

b) 

Fig. 36 anteriormost part of body (ventral view) 

c) other crabs 

Fig. 35 Belliidae 

antennules folding transversely 

or obliquely 

21a. Carapace well calcified, dorsal surface gently to strongly convex, almost always covered 

with spines or granules (Fig. 40); hooked setae often present; abdomen usually with 6 

segments and a telson (Fig. 38a) (rarely 5 segments and a telson); male genital openings 

coxal (i.e. situated at basis of coxae of fourth legs, Fig. 39a) . . . . . . . Majidae (in part) (p. 1136) 

21b. Carapace poorly calcified, soft, dorsal surface flat to almost flat, never covered with 

spines or granules (Fig. 41); hooked setae absent; abdomen always with only 5 

segments and a telson (Fig. 38b) (or fewer segments, some completely fused); male 

genital openings sternal (Fig. 39b) . . . . . . . . . . . . . . . . . . . . . . . . Hymenosomatidae 

6 

5 

4 

3 

2 

1 

a) 6 segments 

and telson 

Fig. 38 abdomen 

5 

4 

3 21 

b) 5 segments 

and telson 

abdominal 

cavity 

penis 

c) 

incomplete 

orbit 

d) 

a) b) 

coxa of 4 th 

(last) leg 

Fig. 37 orbits 

abdominal 

cavity 

penis 

abdomen 

abdomen 

a) coxal position b) sternal position 

Fig. 39 position of male genital opening 

complete 

orbit 

coxa of 4 th 

(last) leg

1068 Crabs 

Fig. 40 Majidae 

22a. Antennal flagellum distinctly setose 

(Fig. 42a) . . . . . . . . . . . . . . . . . . . → 23 

22b. Antennal flagellum slightly setose to 

glabrous (Fig. 42b) . . . . . . . . . . . . . . → 26 

23a. Front entire, without teeth or lobes; anterolateral 

and posterolateral margins 

of carapace lined with dense, long setae 

forming distinct fringe (Fig. 43) . . . . . Thiidae 


23b. Front with teeth or lobes; anterolateral 

a) highly setose b) slightly setose 

and posterolateral margins of carapace 

with relatively dense setae, but 

Fig. 42 antennal flagellum 

not forming distinct fringe . . . . . . . . . . . → 24 

24a. Antennae very long, longer than or as long as carapace length, strongly setose (Fig. 44). . Corystidae 

24b. Antennae short, much shorter than carapace length, not strongly setose. . . . . . . . . . . . → 25 

Fig. 43 Thiidae 

(after Christiansen, 1969) 

25a. Carapace usually rounded to longitudinally 

ovate; vulvae of adult 

female completely covered by abdomen 

(Figs 45a and 46) . . . . Atelecylidae 

25b. Carapace squarish to hexagonal; 

vulvae of adult female exposed, 

not covered by abdomen 

(Figs 45b and 47) . . . . . . . Cheiragonidae 


Fig. 41 Hymenosomatidae 


long 

antenna 

Fig. 44 Corystidae 

abdomen vulva 

abdomen 

(covered by 

abdomen) 

a) b) 

Fig. 45 position of female vulvae 

vulva (not 

covered by 

abdomen)


Fig. 46 Atelecylidae 

(after Alcock and Anderson, 1899) 

Fig. 47 Cheiragonidae 

(after Sakai, 1976) 

26a. Carapace hexagonal, its width and length subequal; front broad compared to maximum 

carapace width; anterolateral margins of carapace weakly convex, each with 4 or 5 

well-developed teeth (Fig. 48) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pirimelidae 


26b. Carapace transversely ovate; front narrow compared to maximum carapace width; 

anterolateral margins of carapace strongly convex, each usually with well-defined teeth 

or lobes (Fig. 49) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cancridae 

broad front 

narrow front 

Fig. 48 Pirimelidae 

Fig. 49 Cancridae 



27a. Carapace triangular or hexagonal; front triangular, forked or spiniform; chelipeds usually 

stout, very elongate (Fig. 50) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parthenopidae 

27b. Carapace shape not as above; front usually truncate or multidentate; chelipeds usually 

stout to slender, relatively short; if chelipeds long, usually slender . . . . . . . . . . . . . . . → 28 

a) 


Fig. 50 Parthenopidae 

b)

1070 Crabs 

Key 28a. toMerus Brachyura of third maxilliped strongly reduced compared to ischium and other segments 

(Fig. 51a), sometimes completely fused with ischium (Fig. 51b); male genital openings 

always sternal (Fig. 52a); males always much smaller than females; parasites on hard 

corals, or symbionts on molluscs, echinoderms or worms . . . . . . . . . . . . . . . . . . . → 29 

28b. Merus of third maxilliped well developed, usually quadrate in shape (Fig. 51c); male 

genital openings usually coxal (i.e. situated at basis of coxae of fourth legs, Fig. 52b), 

occasionally sternal; males not much smaller, subequal or larger in size to females; 

free-living crabs, not parasites of hard corals, not symbionts on molluscs or worms . . . . . . → 30 

merus strongly 

reduced 

normal 

merus 

ischium 

merus 

and 

ischium 

fused 

a) Cryptochiridae b) Pinnotheridae c) others 


29a. Carapace longitudinally ovate to 

longitudinally rectangular; dorsal 

surface of carapace usually with 

small spines or tubercles; merus of 

third maxilliped always separated 

from ischium, dactylus terminally 

attached to propodus; dactylus of 

legs strongly hooked; parasites on 

hard corals (Fig. 53) . . . . . . Cryptochiridae 

29b. Carapace circular to transversely 

ovate; dorsal surface of carapace 

smooth; merus of third maxilliped 

may be fused with ischium; dactylus 

usually subterminally to basally 

attached to propodus with palp 

often appearing bifurcated; dactylus 

of legs weak, not strongly 

hooked; parasitic or commensal on 

molluscs or worms (Fig. 54). . . Pinnotheridae 

sternum 

a) 

sternum 

male 

genital 

openings 

sternal 

b) coxa of 

4th male 

genital 

1 

2 

openings 

coxal 

3 

4 

leg 

Fig. 52 position of male genital opening 

a) male 

Fig. 53 Cryptochiridae 

(after Takeda and Tamura, 1980) 

a) male b) female 

Fig. 54 Pinnotheridae 


b) female


30a. Last pair of legs with dactylus flattened, paddle-like (Fig. 55a) (with the exception of a 

few mud-dwelling and obligate coral-symbionts) (Fig. 56) . . . . . . . . . . . Portunidae (p. 1115) 

30b. Last pair of legs with normal dactylus (Fig. 55b) . . . . . . . . . . . . . . . . . . . . . . . . → 31 

31a. Distinct rhomboidal gap between third maxillipeds; mandibles usually visible when 

mouthparts closed (Fig. 57a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 32 

31b. No distinct rhomboidal gap between third maxillipeds (Fig. 57b); mandibles never visible 

when mouthparts closed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 33 

dactylus paddle-like 

a) 

b) 

dactylus 

normal 

Fig. 55 fourth leg Fig. 56 Portunidae 

32a. Carapace transversely ovate to 

circular, dorsal surface usually 

very smooth, rounded; land crabs 

(Fig. 58) . . . . . . . Gecarcinidae (p. 1147) 

32b. Carapace squarish, transversely 

rectangular, trapezoidal or circular, 

dorsal surface rough, rugose 

or setose (Fig. 59) . . . . . . . . 

. . . . . . . . . Grapsidae (in part) (p. 1138) 

a) subfamily Grapsinae 

rhomboidal gap distinct, 

mandibles visible 

no rhomboidal gap, 

mandibles hidden 

third 

maxillipeds 

a) b) 

Fig. 57 third maxillipeds 

Fig. 58 Gecarcinidae 

b) subfamily Sesarminae 

Fig. 59 Grapsidae 

33a. Front simple, triangular, very narrow compared to broader carapace (Fig. 60a) . . . . . . . . → 34 

33b. Front truncate, multilobate or multidentate, relatively broad compared to transverse 

carapace (Fig. 60b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 35 

very narrow front 

broad front 

a) Ocypodidae b) other crabs 

Fig. 60 anterior part of carapace (dorsal view)

1072 Crabs 

34a. Orbits long, eyes elongate; terrestrial to semiterrestrial crabs (Fig. 61) . . . . Ocypodidae (p. 1152) 

34b. Orbits absent, eyes relatively short; semiterrestrial crabs (Fig. 62) . . . . . . . . . . . Mictyridae 

a) 

Fig. 61 Ocypodidae 


35a. Cross-section of dactylus of walking leg T-shaped (Figs 63 and 64). . . . . . Geryonidae (p. 1132) 

35b. Cross-section of dactylus of walking leg not T-shaped, usually quadrate to ovate 

(Fig. 63b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 36 

Fig. 62 Mictyridae 

36a. Male abdominal segments 2 and 3 always immovable, completely to incompletely fused 

(Fig. 65a); male first gonopods strongly bent (Fig. 66a); small semiterrestrial crabs 

usually associated with estuarine habitats and mangroves (Fig. 67) . . . . . . . . Camptandriidae 

36b. Male abdominal segments 2 and 3 always movable, never fused (Fig. 65b); male first 

gonopods normal (Fig. 66b); aquatic to semiterrestrial crabs . . . . . . . . . . . . . . . . . . → 37 

Fig. 64 Geryonidae 

b) 

a) T-shaped b) ovate 

Fig. 63 cross-section of dactylus of walking leg 

6 

5 

4 

3 

2 

1 

segments 2 

and 3 

immovable 

a) b) 


6 

5 

4 

3 

2 

1 

segments 

2 and 3 

freely 

movable


strongly 

recurved 

a) Camptandriidae b) others 

Fig. 66 male first gonopod 

Fig. 67 Camptandriidae 


37a. Carapace squarish to longitudinally rectangular, posterolateral margins subparallel, 

dorsal surface flat or gently convex; free-living intertidal, estuarine, or fresh-water crabs 

(Fig. 68) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grapsidae (in part) (p. 1138) 

37b. Carapace shape not as above; dorsal surface usually convex; posterolateral margins 

gently to strongly converging; sublittoral to deep-water crabs . . . . . . . . . . . . . . . . . . → 38 

38a. Legs with special dactylo-propodal articulation (formed by a rounded, submedian 

extension of the lateral margin of the propodus, sliding underneath a projecting button 

on the subproximal edge of the dactylus; Fig. 69a) . . . . . . . . . . . . . . . . . . . . . . . → 39 

38b. Legs without special articulation between propodus and dactylus (Fig. 69b) . . . . . . . . . . → 41 

projecting button 

of dactylus 

a) special dactylopropodal 

articulation 

propodus 

b) normal articulation 

Fig. 68 Grapsidae (subfamily Varuninae) 

Fig. 69 distal end of walking leg 


39a. Fingers of chelipeds distinctly spoon-tipped (Fig. 70a); male abdominal segments 3 to 

5 always immovable, completely fused although sutures may be partially visible 

(Figs 71a and 72) . . . . . . . . . . . . . . . . . . . . . . . . . . . Xanthidae (in part) (p. 1098) 

39b. Fingers of chelipeds with sharp tip, not distinctly spoon-tipped (Fig. 70b); male abdominal 

segments 3 to 5 freely movable (Fig. 71b) or fused . . . . . . . . . . . . . . . . . . . . . → 40 

fingers 

spoon-tipped 

a) 

tip sharp 

Fig. 70 pincer 

b) 

segments 3-5 fused 

(sutures completely absent) 

6 

5 

4 

3 

2 

1 

6 

5 

4 

3 

2 

1 


(sutures visible 

laterally) 

segments 

3-5 freely 

movable 

a) Xanthidae b) others 


6 

5 

4 

3 

2 

1

1074 Crabs 

40a. Male first gonopod very slender, S-shaped (Fig. 73a); male second gonopod very short, 

sigmoid (very short, stout, comma-shaped) less than 1/4 the length of male first 

gonopod (Fig. 74b); internal and external commensals of echinoderms (Fig. 75) . . . Eumedonidae 

40b. Male first gonopod moderately stout, slightly sinuous to almost straight (Fig. 73b); male 

second gonopod short but not sigmoid, always longer than 1/4 the length of male first 

gonopod (Fig. 74b); commensals on corals (Fig. 76) . . . . . . . . . . . . . . . . . . .Trapeziidae 

a) slender, 

S-shaped 

b) stout, 

slightly sinous 


41a. Male abdominal segments 3 

to 5 distinct, movable 

(Fig. 77a); regions on carapace 

usually poorly defined . . . . → 42 

41b. Male abdominal segments 3 

to 5 fused, immovable, although 

sutures may be partially 

visible (Fig. 77b, c); 

regions on carapace usually 

well defined . . . . . . . . . . . . . → 46 

a) b) 

a) sigmoid 

(short, stout) 

Fig. 72 Xanthidae 

b) normal, 

not sigmoid 

Fig. 74 male second gonopod Fig. 75 Eumedonidae 

6 

5 

5 

4 

3 

2 

1 

b) segments 3-5 fused 

(sutures visible laterally) 

4 

3 

2 

1 

a) segments 3-5 

freely movable 


6 

a) b) 

Fig. 76 Trapeziidae 

6 

5 

3 

4 

2 

1 

c) segments 3-5 

completely fused


42a. Longitudinal ridges which define efferent respiratory current absent, or strong only on 

posterior part of endostome, ridges not clearly visible on anterior part of endostome 

when mouthparts pushed aside (Fig. 78a, b); carapace and legs never spinose . . . . . . . . → 43 

42b. Longitudinal ridges which define efferent respiratory current well developed along entire 

endostome, ridges visible on anterior part of endostome when mouthparts pushed aside 

(Fig. 78c); or carapace and legs strongly spinose . . . . . . . . . . . . . . . . . . . . . . . . → 44 

endostomial ridges 

almost absent 

a) b) 

43a. Carapace transversely ovate; endopodite of 

first maxilliped usually with distinct lobe on 

inner angle (Fig. 79a); deep-water crabs, 

associated with hydrothermal vents 

(Fig. 80) . . . . . . . . . . . . . . . . Bythograeidae 

43b. Carapace hexagonal; endopodite of first 

maxilliped with small lobe or without small 

lobe on inner angle (Fig. 79b); mainly shallow-water 

crabs, if in deep waters, not 

known to be associated with hydrothermal 

vents (Fig. 81) . . . . . . . . . . . . . Platyxanthidae 


ridges strong 

only posteriorly 

Fig. 78 endostome (= part of the "mouth field") 

Fig. 80 Bythograeidae Fig. 81 Platyxanthidae 

44a. Male first gonopod stout, straight (Fig. 82a); male second gonopod very elongate, subequal 

in length to, or longer than, male first gonopod (Figs 83a, b and 84) . . . . . . . Eriphiidae (p. 1103) 

44b. Male first gonopod slender to stout, slightly sinuous to almost straight (Fig. 82b); male 

second gonopod distinctly shorter than length of male first gonopod (Fig. 83c). . . . . . . . . → 45 

a) stout, 

straight 

b) slender, 

sinuous 

c) slightly 

sinuous 


a) very long, 

whip-like 

b) long c) very 

short 

distinct 

lobe 

small 

lobe 

exopod 

c) 

ridges well 

developed 

exopod 

a) Bythograeidae b) Platyxanthidae 

Fig. 79 first maxilliped 

Fig. 83 male second gonopod Fig. 84 Eriphiidae

1076 Crabs 

45a. Male first gonopod slender, distinctly S-shaped to almost straight (Fig. 85a); male second 

gonopod sigmoid (very short, stout, comma-shaped) (Figs 86a and 87) . . . . . Pilumnidae (p. 1112) 

45b. Male first gonopod moderately stout, slightly sinuous (Fig. 85b); male second gonopod 

about 1/3 to 1/2 the length of male first gonopod (Figs 86b and 88) . .Goneplacidae (in part) (p. 1114) 

a) slender, 

sinuous 

b) slightly 

sinuous 


a) very 

short 

b) short 

Fig. 86 male second gonopod 

Fig. 87 Pilumnidae 


46a. Anterolateral margins entire, only 1 rounded lateral tooth present on each margin 

(Fig. 89) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carpiliidae (p. 1110) 

46b. Anterolateral margins multidentate or multilobate . . . . . . . . . . . . . . . . . . . . . . . . → 47 

Fig. 88 Goneplacidae Fig. 89 Carpiliidae 

47a. Carapace usually transversely rectangular, subcircular, sometimes transversely ovate; 

male second gonopod usually 0.3 times the length, to subequal length, of male first 

gonopod (Figs 90a and 91) . . . . . . . . . . . . . . . . . . . . . . . . . Goneplacidae (in part) 

47b. Carapace usually hexagonal or transversely ovate; male second first gonopod short, 

subequal to, or less than, 0.3 times the length of male first gonopod but not distinctly 

sigmoid (stout, comma-shaped) (Fig. 90b) . . . . . . . . . . . . . . . . . . . . . . . . . . . → 48 

male 1 st 

gonopod 

male 2 nd 

gonopod 

male 1 st 

gonopod 

a) b) 

male 2 nd 

gonopod 

Fig. 90 relative length of male first and second gonopods 

Fig. 91 Goneplacidae 

(after Shen, 1932)

Key to Anomura 1077 

48a. Male first gonopods very slender, usually S-shaped, 

tip relatively simple, often with long setae subdistally 

and distally (Figs 92 and 93) . . . . Xanthidae (in part) (p. 1097) 

48b. Male first gonopods moderately stout, sometimes 

slightly sinuous, tip usually with numerous complex 

folds (Figs 92b and 94) . . . . . . . . . . . . . . . . Panopeidae 

Fig. 93 Xanthidae Fig. 94 Panopeidae 


Key to Anomura KEY TO THE FAMILIES OF CRAB-LIKE ANOMURA 

Notes: the following key covers all marine families of crab-like anomuran crabs recognized here, most of 

which (except the Lomidae) have been reported from the Western Central Pacific. The key uses, wherever 

possible, external and easily viewed characters. Due to the diversity in some families, however, not all their 

members can be identified to the family level with this key, although it should work for the majority of species 

encountered. A specialist should be consulted for the more difficult species. For further useful information 

on crab-like anomurans and allies see De Man (1928), Miyake (1982), Macpherson (1988), McLaughlin 

(1997), and Baba (1988). 

1a. Carapace usually longitudinally ovate; 

dactyli of second to fourth legs curved, 

laterally flattened (Fig. 95a-c); burrowing 

crabs . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Carapace usually not longitudinally ovate; 

dactyli of second to fourth legs normal, 

not laterally flattened, usually non-burrowing 

crabs . . . . . . . . . . . . . . . . . . . . → 3 

2a. Carapace dorsoventrally flattened; first 

leg subchelate, with pincer-like structure 

(Fig. 96) . . . . . . . . . . . . . . . . . . Albuneidae 

2b. Carapace subcylindrical in cross-section; 

first leg simple, not subchelate (Fig. 97) . . . Hippidae 

Fig. 96 Albuneidae 

1 st leg with 

“pincer” 

1 st leg simple 

complex 

lobes 

a) Xanthidae b) Panopeidae 

Fig. 92 male first gonopods 

a) b) c) 

Fig. 95 dactylus of legs 2-4 

Fig. 97 Hippidae

1078 Crabs 

3a. Uropods usually structurally 

modified into a “rasp” 

(Fig. 98a) and used for clinging 

onto interior of gastropod 

shells or hollow wood or underside 

of bivalve shells, at 

least in early crab stages . . . . . . → 4 

3b. Uropods (may be absent) 

not modified for clinging purposes, 

no “rasp” present 

(Fig. 98b); crabs not associated 

with hollow or concave 

objects even in early crab 

stages . . . . . . . . . . . . . . . . → 9 

4a. Abdomen bilaterally symmetrical, clearly divided 

into segments, usually well calcified even at early 

crab stages, pleurites (side plates) distinct; 

shrimp- to lobster-like appearance, abdomen not 

tucked under carapace (Fig. 99) . . . . . . . . Pylochelidae 

4b. Abdomen usually bilaterally asymmetrical, not 

clearly divided into segments, usually weakly 

calcified or membranous, at least in early crab 

stages, pleurites (side plates) absent; typical 

hermit crabs or distinctly crab-like with abdomen 

tucked under the carapace . . . . . . . . . . . . → 5 

5a. Coxae of third maxillipeds close to each other, no 

distinct gap between them (Fig. 100a); chelipeds 

equal or unequal, when unequal, left chela larger ..... → 6 

5b. Coxae of third maxillipeds distinctly separated 

from each other, wide gap between them 

(Fig. 100b); chelipeds unequal, with right chela 

larger . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

6a. Eyestalk laterally flattened (Fig. 101a); eyes usually 

held subparallel to each other (Fig. 102a); 

antennae laterally flattened, short; completely 

terrestrial crabs (Fig. 103). . . . . . Coenobitidae (p. 1154) 

6b. Eyestalk circular to subcircular in cross-section, 

not laterally flattened (Fig. 101b); eyes 

held obliquely to each other (Fig. 102b); antennae 

long, subcylindrical in cross-section; completely 

aquatic crabs (Fig. 104) . . . . . . . . . Diogenidae 

eyes 

laterally 

flattened 

eyes 

cylindrical 

a) Coenobitidae b) Diogenidae 

Fig. 101 eye in dorsal view 

rasp-like 

structures 

of uropod 

uropod without 

a) rasp-like structures b) 

Fig. 98 posterior portion of body (dorsal view) 

eyes held 

subparallel 

narrow gap 

a) Coenobitidae 

and Diogenidae 

Fig. 99 Pylochelidae 

coxa 

wide gap 

b) Lithodidae, Paguridae, 

and Parapaguridae 


eyes held 

obliquely 

a) Coenobitidae b) Diogenidae 

Fig. 102 relative position of eyes (dorsal view)

Key to Anomura 1079 

a) 

Fig. 103 Coenobitidae 

7a. Adults distinctly crab-like; carapace well calcified in 

adults, often with numerous strong granules or spines; 

rostrum spiniform, distinct; second to fourth legs distinct, 

fifth pair strongly reduced, inserted into branchial 

chamber (Fig. 105) . . . . . . . . . . . . . . . . . . . Lithodidae Fig. 104 Diogenidae 

7b. Adults typical hermit crabs; carapace weakly calcified in adults, rarely with granules or 

spines; rostrum weak to indiscernible; second to third legs distinct, fourth and fifth pair 

strongly reduced, but not inserted into branchial chamber . . . . . . . . . . . . . . . . . . . . → 8 

Fig. 105 Lithodidae 

Fig. 106 sixth segment and telson (dorsal view) 

8a. First maxilliped with distinct flagellum; telson usually with median constriction, lateral 

margins with median cleft (Fig. 106a); mostly shallow water with some deep-water 

species (Fig. 107) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paguridae 

8b. First maxilliped without flagellum; telson without median constriction, lateral margins 

entire (Fig. 106b); deep-water species (Fig. 108) . . . . . . . . . . . . . . . . . . Parapaguridae 

b) 

convex 

margin 

median 

constriction 

a) Paguridae b) Parapaguridae 

Fig. 107 Paguridae Fig. 108 Parapaguridae

1080 Crabs 

9a. No tail fan (telson and uropods) present; eye peduncles broad and flat, cornea inserted 

laterally on peduncle (Fig. 109) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lomidae 


9b. Distinct tail fan (telson and uropods) present; eye peduncles normal; cornea inserted 

terminally on peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

10a. Carapace circular, subcircular to longitudinally ovate; rostrum short to absent; most of 

abdomen tucked underneath carapace (Fig. 110) . . . . . . . . . . . . . . . . . . . Porcellanidae 

10b. Carapace longitudinally rectangular to longitudinally ovate, rarely circular; rostrum 

distinct; most of abdomen not tucked underneath carapace . . . . . . . . . . . . . . . . . . → 11 

Fig. 109 Lomidae Fig. 110 Porcellanidae 

11a. Antennular peduncle with 4 movable articles; 

telson divided into 2 or more smaller 

plates (Figs 111a and 112) . . . . . . . . Galatheidae 

11b. Antennular peduncle with 5 movable articles; 

telson entire, not divided into smaller 

plates (Figs 111b and 113) . . . . . . . Chirostylidae 

List of Families LIST OF FAMILIES OF MARINE BRACHYURA AND CRAB-LIKE ANOMURA 

PRESENTLY RECOGNIZED 

Note: the following list also includes 5 families of Brachyura and a single family of anomuran crab which 

have not been recorded yet from the Western Central Pacific. They are included in the list and above keys 

since it is likely that some of these families may be discovered in the future in the area. These families are 

marked in the list by an asterisk (*). 


Infraorder BRACHYURA Latreille, 1803 

ATELECYLIDAE Ortmann, 1893 

BELLIIDAE Dana, 1852 (= Acanthocyclidae Dana, 1852) 

BYTHOGRAEIDAE Williams, 1980 

CALAPPIDAE De Haan, 1833 

CAMPTANDRIIDAE Stimpson, 1858 

CANCRIDAE Latreille, 1803 (= Trichoceridae Dana, 1852) 

a) divided into 2 or 

more small plates 

b) not divided into 

small plates 


Fig. 112 Galatheidae Fig. 113 Chirostylidae

List of Families 1081 

* CHEIRAGONIDAE Ortmann, 1893 (= Telmessidae Guinot, 1977) 

CARPILIIDAE Ortmann, 1893 

CORYSTIDAE Samouelle, 1819 (= Nautilocorystidae Ortmann, 1893; Euryalidae Rathbun, 1930) 

CRYPTOCHIRIDAE Paulson, 1875 (= Hapalocarcinidae Calman, 1900) 

CYCLODORIPPIDAE Ortmann, 1892 (= Tymolidae Alcock, 1896) 

CYMONOMIDAE Bouvier, 1898 

DORIPPIDAE MacLeay, 1838 

DROMIIDAE De Haan, 1833 

DYNOMENIDAE Ortmann, 1892 

ERIPHIIDAE MacLeay, 1838 (= Menippidae Ortmann, 1893; Oziidae Dana, 1851) 

EUMEDONIDAE Dana, 1852 

HEXAPODIDAE Miers, 1886 

HOMOLIDAE De Haan, 1839 (= Thelxiopeidae Rathbun, 1937) 

HOMOLODROMIIDAE Alcock, 1899 

HYMENOSOMATIDAE MacLeay, 1838 

GECARCINIDAE MacLeay, 1838 

GERYONIDAE Colosi, 1923 

GONEPLACIDAE MacLeay, 1838 

GRAPSIDAE MacLeay, 1838 

LATREILLIDAE Stimpson, 1858 

LEUCOSIIDAE Samouelle, 1819 

MAJIDAE Samouelle, 1819 (= Inachidae MacLeay, 1838; Epialtidae MacLeay, 1838; Blastidae 

Stebbing, 1902; Mamaiidae Stebbing, 1902) 

MICTYRIDAE Dana, 1852 

OCYPODIDAE Rafinesque, 1815 

* ORTHIYIIDAE Dana, 1852 

PALICIDAE Bouvier, 1897 (= Cymopoliidae Faxon, 1895) 

PANOPEIDAE Ortmann, 1893 

PARTHENOPIDAE MacLeay, 1838 (= Mimilambridae Williams, 1979) 

PILUMNIDAE Samouelle, 1819 

PINNOTHERIDAE De Haan, 1833 

* PIRIMELIDAE Alcock, 1899 

* PLATYXANTHIDAE Guinot, 1977 

PORTUNIDAE Rafinesque, 1815 (= Megalopidae Haworth, 1825; Carcinidae MacLeay, 1838; Xaividae 

Berg, 1900) 

POUPINIIDAE Guinot, 1993 

RANINIDAE De Haan, 1839 

RETROPLUMIDAE Gill, 1894 

* THIIDAE Dana, 1852 

TRAPEZIIDAE Miers, 1886 

XANTHIDAE MacLeay, 1838 

Infraorder ANOMURA H. Milne Edwards, 1832 (crab-like families listed only) 

ALBUNEIDAE Stimpson, 1858 

CHIROSTYLIDAE Ortmann, 1892 

COENOBITIDAE Dana, 1851 

DIOGENIDAE Ortmann, 1892 

GALATHEIDAE Samouelle, 1819 

HIPPIDAE Latreille, 1825 

LITHODIDAE Samouelle, 1819 

PAGURIDAE Latreille, 1803 

PARAPAGURIDAE Smith, 1882 

PORCELLANIDAE Haworth, 1825 

PYLOCHELIDAE Bate, 1888 (= Pomatochelidae Stebbing, 1914) 

* LOMIDAE Bouvier, 1895

1082 Crabs 

References 

Abele, L.G. and E. Felgenhauer. 1982. Decapoda. In McGraw-Hill, Synopsis and classification of living organisms, 

edited by S.S. Parkes. McGraw-Hill Book Co., pp. 296-326. 

Baba, K. 1988. Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the “Albatross” Philippine Expedition, 

1907-1910. Researches on Crustacea, special number 2: 1-203. 

Balss, H. 1957. Decapoda, VIII: Systematik. In H.G. Bronn, Klassen und Ordnungen des Tierreichs. Band 5, Abteilung 

1, 7(12):1505-1672. 

Bowman, T.E. and L.G. Abele. 1982. Classification of the recent crustacea. In The biology of the Crustacea. Vol. 1, 

Systematics, the fossil record and biogeography, edited by D.E. Bliss. Academic Press. 

Burggren W.W. and B.R. McMahon (eds). 1988. Biology of of land crabs. Cambridge University Press, 479 p. 

Chace, F. A., Jr. 1951. The number of species of decapod and stomatopod Crustacea. J. Wash. Acad. Sci., 41(11):370-372. 

Chang, C.M. 1965. Edible Crustacea of Taiwan. Taipei, JCRR Publication, 60 p. 

Davidson, A. 1976. Seafoods of South-east Asia. Singapore, Federal Publications, 343 p. 

Dawson, E.W. and W.R. Webber (eds). 1991. The deep-sea red crab Chaceon (“Geryon”). A guide to information and 

a reference list of the family Geryonidae. National Museum of New Zealand, miscellaneous series, 24 p. 

De Man, J.G. 1928. The Decapoda of the Siboga-Expedition. Part VII. The Thalassinidae and Callianassidae collected 

by the Siboga-Expedition with some remarks on the Laomediidae. Leiden, Siboga-Expeditie, 39a, 90 p. 

Fielder, D. F. and M. P. Heasman. 1978. The mud crab. Queensland Museum Booklet, 15 p. 

Guinot, D. 1967. Les crabes comestibles de l’Indo-Pacifique. Editions de la Fondation Singer-Polignac, 1966, 145 p. 

Guinot, D. 1978. Principes d’une classification evolutive des Crustaces Decapodes Brachyoures. Bulletin du biologique 

France et Belgique, new series, 112(3):211-292. 

Guinot, D. 1979. Donnees nouvelles sur la morphologie, la phylogénèse et la Crustacés Décapodes Brachyoures. Mém. 

Mus. natn. Hist. nat., Paris, (A) Zoologie, 112:1-354. 

Guo, J.Y., N.K. Ng, A. Dai, and P.K.L. Ng. 1997. The taxonomy of three commercially important species of mitten crabs of 

the genus Eriocheir De Haan, 1835 (Crustacea: Decapoda: Brachyura: Grapsidae). Raffles Bull. Zool., 45:445-476. 

Harminto, S. and P.K.L. Ng. 1991. A revision of the Camptandriine genus Baruna Stebbing, 1904 (Crustacea: Brachyura: 

Decapoda: Ocypodidae), with descriptions of two new species. Raffles Bull. Zool., 39(1):187-207. 

Kailola, P.J., M.J. Williams, P.C. Stewart, R.E. Reichelt, A. McNee, and C. Grieve (compilers). 1993. Crustaceans. In 

Australian fisheries resources. Sydney, Bureau of Resource Sciences and Fisheries Research and Development 

Corporation, pp. 113-179. 

Manning, R. B. and L.B. Holthuis. 1981. West African brachyuran crabs (Crustacea: Decapoda). Smithson. Contrib. 

Zool., (306):1-379. 

McLaughlin, P.A. 1997. Crustacea Decapoda: Hermit crabs of the family Paguridae from the KARUBAR Cruise in 

Indonesia. Résultats des Campagnes MUSORSTOM, edited by A. Crosnier and P. Bouchet, Vol. 16. Mém. Mus. 

natn. Hist. nat., 172:433-572. 

Miyake, S. 1982. Japanese crustacean decapods and stomatopods in color. Vol. I. Macrura, Anomura and Stomatopoda. 

Osaka, Hoikusha Publishing Co., 261 p. 

Motoh, H. 1980. Field guide to the edible Crustacea of the Philippines. Iloilo, SEAFDEC Publication, 96 p. 

Ng, P.K.L. 1988. The freshwater crabs of Peninsular Malaysia and Singapore. Department of Zoology, National 

University of Singapore, Shinglee Press, Singapore, 156 p. 

Ng, P.K.L., D.G.B. Chia, E.G.L. Koh and L.W.H. Tan. 1992. Poisonous Malaysian crabs. Nature Malaysiana, 17(1):4-9. 

Ng, P.K.L. and G. Rodriguez. 1986. New records of Mimilambrus wileyi Williams, 1979 (Crustacea: Decapoda: 

Brachyura), with notes on the systematics of the Mimilambridae Williams, 1979 and Parthenopoidea MacLeay, 

1838 sensu Guinot, 1978. Proc. Biol. Soc. Wash., 99(1):88-99. 

Poupin, J. 1994. Quelques crustacés décapodes communes de Polynesia Francaise. Rapport Scientifique du Service 

Mixte de Surveillance Radiologique et Biologique, 86 p. 

Sakai, T. 1976. Crabs of Japan and the adjacent seas. In three volumes; English Text, p. xxix + 773p., Japanese Text, 

pp. 1-461, Plates Volume, pp. 1-16, Pls. 1-251. Tokyo, Kodansha Ltd. 

Sakai, T., T. Tomiyama and T. Hibiya. 1983. Crabs. Tokyo, Fisheries in Japan, 177 p. 

Schreiber, A. and E. Cases. 1984. Edible Crustacea of the central Philippines (Decapoda: Reptantia).Philipp. Sci., 21:11-50. 

Serène, R. 1968. The Brachyura of the Indo Pacific Region. In Prodromus for a check list of the non-planctonic marine 

fauna of South East Asia. Singapore National Academy of Science, Special Publication number 1:33-120. 

Serène, R. 1984. Crustacés Décapodes Brachyoures de l’Ocean Indien occidental et de la Mer Rouge. Xanthoidea: 

Xanthidae et Trapeziidae.Addendeum Carpiliidae et Menippidae - A.Crosnier.Faune Tropicale (ORSTOM), 24:1-400. 

Stevcic, Z. 1988. The status of the family Cheiragonidae Ortmann, 1893. Oebalia, new series, 14:1-14. 

Stevcic, Z., P. Castro, and R.H. Gore. 1988. Re-establishment of the Family Eumedonidae Dana, 1853 (Crustacea: 

Brachyura). J. Nat. Hist., 22:1301-1324. 

Titcomb, M. 1978. Native use of marine invertebrates in old Hawaii. Pac. Sci., 32:325-386. 

Wee, D.P.C. and P.K.L. Ng. 1995. Swimming crabs of the genera Charybdis De Haan, 1833, and Thalamita Latreille, 

1829 (Crustacea: Decapoda: Brachyura: Portunidae) from Peninsular Malaysia and Singapore. Raffles Bull. 

Zool., Supplement 1, 128 p. 



Infraorder Brachyura - Homolidae 1083 

Infraorder Brachyura - Homolidae Infraorder BRACHYURA 

HOMOLIDAE 

Deep-water carrier crabs 

Diagnostic characters: Carapace longitudinally rectangular; dorsal surface granulose to spinose; 

front narrow, usually with 3 long horn-like projections (rostra). Male chelipeds long. Last (fourth) 

pair of legs inserted obliquely on carapace and directed upwards, reduced, subchelate to chelate, 

modified to carry sponges. All male abdominal segments distinct, movable. 

carapace 


rectangular 

last leg 

reduced, 

subchelate 

to chelate 

Habitat, biology, and fisheries: Benthic deep-water crabs, usually occurring in depths below 200 m. Most 

species are of minor commercial value, being either too small or only occasionally caught. Only the large 

Paromola japonica and P. macrochira have some fishery value. P. macrochira, however, is known only 

from Japanese and Taiwanese waters, but not from the Western Central Pacific. 


The only other crab families which have the last (fourth) pair of legs modified to carry objects are the 

Dromiidae (sponge crabs), Homolodromiidae (deep-water sponge crabs), Latreillidae (spindle crabs), 

Cymonomidae and Cyclodorippidae (deep-water porter crabs) and Dorippidae (porter crabs); some 

Majidae (spider crabs) also have a similar structure. All these families, however, differ markedly in body 

shape (not longitudinally rectangular/subrectangular) from the Homolidae and none of them include 

species of commercial interest. 

Poupiniidae (deep-water hedgehog crabs, non-commercial): in body shape, these recently discovered 

deep-water crabs are most similar to the Homolidae, but poupiniids have the fourth (last) walking leg 

unmodified (not subchelate or chelate). 

References 

Guinot, D. and B. Richer de Forges. 1995. Crustacea Decapoda Brachyura: Révision de la famille des Homolidae de 

Haan, 1839. In Résultats des Campagnes MUSORSTOM, Vol. 13, edited by A. Crosnier. Mém. Mus. natn. Hist. 

nat., 163:283-517. 

Serène, R. and P. Lohavanijaya. 1973. The Brachyura (Crustacea: Decapoda) collected by the NAGA Expedition, 

including a review of the Homolidae. Naga Report, Scientific Results of Marine Investigation of the South China 

Sea and the Gulf of Thailand 1959-1961, 4(4), 186 p.

1084 Crabs 

A single species of interest to fisheries occurring in the area. 

Paromola japonica Parisi, 1915 

Frequent synonyms / misidentifications: Latreillopsis hawaiiensis Edmondson, 1932 / Paromola 

macrochira Sakai, 1961. 

FAO name: En - Japanese deepwater carrier crab. 

Diagnostic characters: Carapace rectangular, longer than broad, surface granular; lateral margins 

granular to spiniform; frontal margin (rostrum) with 3 spiniform projections. Last pair of legs short, with 

dactylus and propodus subchelate, modified for carrying objects. Colour: light to reddish brown overall. 

Size: Maximum carapace length 18 cm (males). 

Habitat, biology, and fisheries: A deep-water crab, found in depths from 150 to 250 m. No targeted 

fisheries are known for this species. Taken as incidental catch by benthic trawls and occasionally by traps. 

Commands low prices when sold in markets. 

Distribution: West Pacific, including Southeast Asia and Hawaii. 

Remarks: The only other species that Paromola japonica might be confused with is P. macrochira which 

is also taken incidentally by trawls and traps, but so far known only from Japanese and Taiwanese waters. 

P. japonica is easily distinguished from P. macrochira by the more spiniform lateral margins of carapace 

and the basal antennal segment bearing several sharp tubercles (absent in P. macrochira).

Dromiidae 1085 

Dromiidae DROMIIDAE 

Sponge crabs 

Diagnostic characters: Carapace circular to hexagonal; dorsal surface gently to strongly convex 

longitudinally and transversely, smooth or granular, usually setose; front narrow, usually entire; 

anterolateral margins of carapace strongly convex, unarmed, or with small spines. Anterior 2 pairs of legs 

normal; last 2 pairs of legs inserted obliquely on carapace and directed upwards, strongly reduced, 

subchelate, modified to carry sponges or tunicates on back of carapace. All male abdominal segments 

distinct, movable; a small platelet-like structure usually intercalated between edges of sixth abdominal 

segment and telson. Male first gonopod stout, simple; male second gonopod long, usually subequal 

or longer than length of male first gonopod. Male and female genital openings sternal. 

carapace 

circular to 

hexagonal 

last 2 pairs of 

legs subchelate 

male abdomen 

Habitat, biology, and fisheries: Benthic crabs, with most species occurring in or near reefs or on 

soft-substrate bottoms. Omnivorous and known to feed on sea stars (Asteroidea). Best known for their habit 

of carrying sponges and colonial tunicates on their backs for camouflage. Only of minor commercial 

importance, as most species are too small to have any food value. Large members of the genera Dromia 

and Lauridromia are occasionally collected by trawls or traps and are sold in local markets. 


The only other crab families which have their last 2 pairs of legs turned upwards and adapted for carrying 

objects are the Homolodromiidae, Cymonomidae, Cyclodorippidae, and Dorippidae. 

Homolodromiids (non-commercial): closest to the dromiids in general body shape, but dromiids are 

generally more rounded to quadrate in shape, and only dromiids possess a pair of intercalated platelet-like 

structures between the abdominal segments 6 and the telson. 

Cymonomidae, Cyclodorippidae, and Dorippidae (all non-commercial): carapace much more rounded and 

flatter; legs proportionately much longer; merus of third maxillipeds triangular in shape (distinctly rectangular 

in dromiids). 

telson 

6 

5 

4 

3 

plate-like 

structure

1086 Crabs 


1a. Dactylus of fourth leg without spine on outer margin (Fig. 1a) . . . . . . . . . . . . Dromia dormia 

1b. Dactylus of fourth leg with distinct spine on outer margin (Fig. 1b) . . . . . . . . . . . . . . . . → 2 

2a. Carapace wider than long, surfaces with soft pubescence; 6 teeth on each anterolateral 

margin; 4 to 8 spines on inner margins of dactyli of first 2 pairs of legs (Fig. 2a) . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lauridromia indica 

2b. Carapace as wide as long, surfaces with coarse pubescence; 4 teeth on each anterolateral 

margin; 16 to 20 spines on inner margins of dactyli of first 2 pairs of legs (Fig. 2b) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lauridromia dehaani 

dactylus 

unarmed 

dactylus with 

distinct spine 

a) Dromia dormia b) genus Lauridromia 

Fig. 1 fourth leg 

List of species of interest to fisheries occurring in the area 


Dromia dormia (Linnaeus, 1763) 

Lauridromia dehaani (Rathbun, 1923) 

Lauridromia indica (Gray, 1831) 

a) Lauridromia indica b) Lauridromia dehaani 

Fig. 2 dactylus of first 2 legs 

References 

Alcock, A.A. 1901. Catalogue of the Indian decapod Crustacea in the collection of the Indian museum. Part I. 

Brachyura. Fascicle I. Introduction and Dromides or Dromiacea (Brachyura Primigenia). Calcutta, Trustees 

of the India Museum, 80 p. 

McLay, C.L. 1993. Crustacea: Decapoda: The sponge crabs (Dromiidae) of New Caledonia and the Philippines with a 

review of the genera. In Résultats des Campagnes MUSORSTOM, 10, edited by A. Crosnier. Mém. Mus. natn. 

Hist. nat., 156:111-251.

Dromiidae 1087 

Dromia dormia (Linnaeus, 1763) 

Frequent synonyms / misidentifications: Cancer dormitator Herbst, 1790; Dromia rumphii Weber, 

1795; D. hirsutissima Dana, 1852 / None. 

FAO name: En - Common sponge crab. 


rounded, as wide as or slightly wider 

than long; surfaces convex, with dense 

pubescence; 5 anterolateral teeth, median 

ones largest. No spine present on 

outer margin of dactylus of last walking 

leg. Colour: light brown overall with pink 

fingers. 

Size: Maximum carapace width 20 cm 

(males) and 12 cm (females). 

Habitat, biology, and fisheries: Prefers 

moderately shallow waters with 

rocky-muddy substrates, at depths from 

5 to 50 m; sometimes found near reefs. 

Occasionally appears in markets in 

parts of eastern Indonesia and Philippines. 

Caught incidentally in nets, fish 

or crab traps, and sometimes by benthic 

(from Alcock, 1901) 

trawls. 

Distribution: Southern Philippines, Ambon, southern China, New Caledonia, and Hawaii. 

Remarks: Only 3 species of Dromia are known to occur in the Indo-West Pacific, and D. dormia is one of 

the largest representatives. It is the only Indo-West Pacific species of Dromia with all the 5 anterolateral 

teeth well developed and large.

1088 Crabs 

Lauridromia indica (Gray, 1831) 

Frequent synonyms / misidentifications: Dromia orientalis Miers, 1880; Dromidiopsis cranioides 

(De Man, 1888) / None. 

FAO name: En - Cannonball sponge crab. 

Diagnostic characters: Carapace rounded, 

much wider than long; surfaces convex, with 

dense pubescence; 6 anterolateral teeth. 

Spine present on outer margin of dactylus of 

last walking leg. Colour: light brown with 

bright pink fingers. 

Size: Maximum carapace width 8 cm (males) 

and 7 cm (females). 

Habitat, biology, and fisheries: In muddy 

substrates from depths of 10 to 60 m. A relatively 

common species, caught incidentally 

by trawlers and benthic nets in various parts 

of Southeast Asia, sometimes in very large 

numbers. No targeted fisheries are known for 

this species, which has a minor commercial 

value due to the poor quality of its flesh. 

Distribution: Thailand, Malaysia, 

Singapore, northern 

Borneo, and southern Philippines. 

Remarks: Only 3 species of 

this Indo-West Pacific genus 

are known. Lauridromia indica 

can easily be distinguished 

from other species of 

the genus by its proportionately 

wider carapace with 

relatively soft pubescence 

and by having 6 teeth on each 

anterolateral margin. 

Lauridromia dehaani (Rathbun, 1923) 

En - Japanese sponge crab. 

Maximum carapace width 10 cm (males) and 8 cm (females). On mud or sandy-muddy substrates 

from depths of 50 to 150 m. Locally consumed by some rural communities, rarely sold in markets. 

Japan, Taiwan Province of China, China, Hong Kong, Java, India, and Gulf of Aden.

Raninidae 1089 

Raninidae RANINIDAE 

Spanner crabs 

Diagnostic characters: Carapace longitudinally 

ovate; dorsal surface strongly granulose 

or squamose to smooth; front triangular, narrow. 

Third maxillipeds very narrow, merus 

distinctly triangular. Eyestalks long, longer than 

front. At least one pair of legs with last 2 or 3 

segments paddle-like. Thoracic sternum very 

narrow, especially sternites 5 to 7. Allmale 



Spanner crabs burrow in soft 

substrates and tend to occur in 

moderately shallower waters. 

They are absent or rare in continental 

shelf waters and prefer 

more saline waters. They feed on 

a variety of worms and softshelled 

molluscs. Most species 

are moderately small and are not 

often encountered. A single species, 

Ranina ranina, is large and 

triangular 

merus 

abundant enough to be fished 

commercially. 

right 3 

carapace 


ovate 

rd 

maxilliped 


The combination of a longitudinally ovate, elongate carapace 

with its very narrow thoracic sternum, and the narrow 

third maxillipeds with a triangular merus easily distinguish 

raninids from other crab families. The following 2 families 

look superficially similar to the Raninidae: 

Homolidae: carapace similarly longitudinally elongate, but 

subcylindrical in shape; last pair of legs reduced, subchelate, 

turned upwards and adapted for carrying objects. 

Corystidae (non-commercial): many species similar in 

body shape, but have broader and more rectangular 

mouthparts, a broader thoracic sternum, and never have 

any of their legs paddle-like and possess a pair of very 

long and highly setose antennae. 

fingers 

strongly bent 

at least 1 

pair of legs 

paddle-like 

Homolidae 

last leg reduced, 

subchelate 

References 

Brown, I. W. 1986. South Queensland’s spanner crabs - a growing fishery. Australian Fisheries, 45(10):3-7. 

Ihle, J.E.W. 1918. Die Decapoda Brachyura der Siboga-Expedition. III. Oxystomata: Calappidae, Leucosiidae, Raninidae. 

Siboga Exped. Monogr., 39b(2):159-322.

1090 Crabs 


Ranina ranina (Linnaeus, 1758) 

Frequent synonyms / misidentifications: Ranina dentata H. Milne Edwards, 1837 / None. 

FAO name: En - Spanner crab. 

 

Diagnostic characters: Carapace very elongate, much longer than broad; anterior part much broader 

than narrow, posterior part waist-like. Abdomen clearly visible from dorsal view. Chelae greatly outsized in 

males; chelae and legs laterally flattened, spade-like. Colour: orange to red overall. 

Size: Maximum carapace length 15 cm (males) and 12 cm (females); weight up to 900 g (males) and 400 g 

(females). 

Habitat, biology, and fisheries: Mainly in more oceanic waters, but also in intertidal waters, to depths of 

more than 100 m, with preference for open sandy areas. Harvested throughout its range, taken by trawls, 

dredges, baited tangle nets, and bottom nets. A widely exploited species in the Philippines, eastern 

Indonesia, East Asia, and eastern and northern Australia. The fishery for Ranina ranina in Australia has 

grown substantially in the 1980s and is probably one of the largest for this species, with almost 700 t landed 

in Queensland and New South Wales from 1989 to 1990. Large specimens command very high prices, 

especially in live-seafood markets. Prices in Australia amount to about US$2 to US$3 per kg, while live 

specimens in Hong Kong (China) and Taiwan Province of China are sold for US$5 to US$10 per kg. The 

fishery for this species is managed in Australia but not elsewhere. 

Distribution: Indo-West Pacific, including Australia, Guam, New Caledonia, and Hawaii.

Calappidae 1091 

Calappidae CALAPPIDAE 

Box and moon crabs 

Diagnostic characters (principal family characters): Carapace circular or ovate to transversely ovate 

or subovate; frontal margin triangular, narrow. Merus of third maxillipeds distinctly triangular. 

Opening for afferent respiratory current at base of chela, no canal present along sides of buccal 

cavern (even when third maxillipeds pushed aside). Male abdominal segments 3 to 5 completely fused; 

male genital openings always coxal. 

Subfamily Calappinae (box crabs): carapace circular or subcircular to transversely ovate or subovate; 

dorsal surface strongly convex longitudinally and transversely, smooth to granular, and ridged; anterolateral 

margins armed with numerous small teeth and lobes, posterolateral parts of carapace sometimes 

strongly expanded to form a clypeiform structure (= expanded posterior edge) which at least partially 

conceals the legs. Chelae laterally flat- 

right chela with 

tened, dorsal margin with high, multiden- 


tate crest, right (larger) chela with special 

tooth on base of pollex for peeling gastropods, 

left chela with forceps-like fingers. 

Legs smooth, laterally flattened to varying 

degrees but never paddle-like. 

right chela 

specialized 

cutting tooth 

Subfamily Matutinae (moon crabs): carapace 

circular to ovate; dorsal surface usually 

almost smooth to granulose; junction of 

antero- and posterolateral margins well developed, 

often with long spine. Legs distinctly 

flattened laterally, last 2 segments 

of all legs paddle-like. 

triangular front (or rostrum) 

triangular merus 

of 3rd maxilliped 


Habitat, biology, and fisheries: Burrowing crabs on soft and mud substrates. Most species of minor 

commercial importance, with only Calappa lophos, C. philargius, and Ashtoret lunaris being more 

commonly seen in markets. 


Portunidae: may be confused with members of the Matutinae 

(also with paddle-like legs and often long lateral spines), but 

can be readily distinguished by the following combination of 

characters: carapace hexagonal, transversely ovate to transversely 

hexagonal; only the last pair of legs paddle-like; meri 

of third maxillipeds quadrate. 

Dorippidae (non-commercial): also with triangular meri of the 

third maxillipeds, but differ by having the last 2 pairs of legs 

small, inserted obliquely on carapace and directed upwards 

for carrying objects; sides of the carapace never expanded 

into a clypeiform process; chelae relatively delicate, never 

with specialized teeth for opening mollusc shells. 

subfamily Calappinae 

subfamily Matutinae 

Portunidae 

expanded 

clypeiform 

process 

all legs 

paddle-like 

only last pair of 

legs paddle-like

1092 Crabs 

Leucosiidae (non-commercial): small, pea-like crabs, also possess triangular meri of the third maxillipeds, 

but differ by having the opening for afferent respiratory current located below the orbits, adjacent to the 

endostome; a distinct canal present along sides of buccal cavern when third maxillipeds are pushed aside; 

sides of carapace never expanded into a clypeiform process; chelae usually delicate, never with specialized 

teeth for opening mollusc shells; legs never paddle-like. 

Key to the subfamilies of Calappidae 

1a. Dactylus of legs normal, not paddle-like; 

right (larger) chela with 

specialized cutting tooth (Fig. 1a); 

posterolateral part of carapace 

often strongly expanded to form a 

clypeiform structure (= expanded 

posterior edge) which covers legs 

(Fig. 2) . . . . . . . . . . . . . . .Calappinae 

specialized 

cutting tooth 

a) Calappinae b) Matutinae 

Fig. 1 right chela 

1b. Dactylus of legs paddle-like; larger chela normal, without specialized cutting tooth 

(Fig. 1b); carapace round, lateral spine very strong; posterorlateral part of carapace 

never expanded; propodus and dactylus of all legs paddle-like (Fig. 3) . . . . . . . . . . Matutinae 

Fig. 2 Calappinae Fig. 3 Matutinae 

Key to food species of Calappinae occurring in the area 

1a. Clypeiform part (= expanded posterior edge) of carapace with margin smooth, entire, 

not armed with spines or teeth (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . Calappa calappa 

1b. Clypeiform part of carapace with margin armed with spines or teeth . . . . . . . . . . . . . . . → 2 

2a. Lateral part of clypeiform part (= expanded posterior edge) of carapace with strong, 

transverse, outwardly pointing projections; carapace with purple lines on lateral regions, 

purple spots on median and posterior regions in life (Fig. 5) . . . . . . . . . . . . . Calappa lophos 

2b. Lateral part of clypeiform part of carapace with dentate margin, colour pattern of life 

specimens not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

margin 

smooth, 

unarmed Fig. 4 Calappa calappa 

strong projections 

Fig. 5 Calappa lophos


3a. Entire posterior margin of carapace and clypeiform part (= expanded posterior edge) 

armed with strong teeth; large purple spot on palm and carpus of cheliped, and around 

each orbit in life (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calappa philargius 

3b. Only anterior edge of clypeiform part of carapace gently denticulate; life colour a uniform 

yellowish grey to grey (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calappa hepatica 

strong 

teeth 

Fig. 6 Calappa philargius 

only anterior 

edge with teeth 

Key to food species of Matutinae occurring in the area 

1a. Outer surface of palm with ridge subparallel to 

ventral margin (Fig. 8a) . . . . . . . . Ashtoret lunaris 

1b. Outer surface of palm with oblique ridge 

(Fig. 8b) . . . . . . . . . . . . . . . . . . . . . . → 2 

Fig. 7 Calappa hepatica 

ridge subparallel 

to ventral margin 

ridge oblique to ventral 

margin 

2a. Carapace surface with pattern of fine red lines 

forming mesh-like pattern; ventral margin of 

palm entire (Fig. 9) . . . . . . . . . . Matuta planipes 

a) Ashtoret lunaris 

Fig. 8 chela 

b) Matuta 

2b. Carapace surface with numerous small spots; ventral margin of palm serrated (Fig. 10) . . Matuta victor 

Fig. 9 Matuta planipes Fig. 10 Matuta victor 



Subfamily CALLAPINAE 

Calappa calappa (Linnaeus, 1758) 

Calappa hepatica (Linnaeus, 1758) 

Calappa lophos (Herbst, 1785) 

Calappa philargius (Linnaeus, 1758) 

Subfamily MATUTINAE 

Ashtoret lunaris (Forsskål, 1775) 

Matuta planipes Fabricius, 1798 

Matuta victor (Fabricius, 1781) 

References 

Chen, H.L. 1993. The Calappidae (Crustacea: Brachyura) of Chinese waters. In The marine biology of the South China 

Sea, by E.B. Morton, pp. 675-704. 

Galil, B.S. 1997. Crustacea Decapoda: A revision of the Indo-Pacific species of the genus Calappa Weber, 1795 

(Calappidae). In Résultats des Campagnes MUSORSTOM, Vol. 18, edited by A. Crosnier. Mém. Mus. natn. Hist. 

nat., 176:271-335. 

Ihle, J.E.W. 1918. Die Decapoda Brachyura der Siboga-Expedition. III. Oxystomata: Calappidae, Leucosiidae, Raninidae. 

Siboga Exped. Monogr., 39b(2):159-322.

1094 Crabs 

Calappa lophos (Herbst, 1785) 

Frequent synonyms / misidentifications: Calappa guerini Brito Capello, 1871 / None. 

FAO name: En - Common box crab. 

Diagnostic characters: Clypeiform posterolateral part 

of carapace with strong lateral projections. Colour: 

ground colour yellowish beige to yellow; posterior 1/3 

of carapace with distinct red spots, posterolateral part 

with transverse red stripes; outer surface of cheliped 

with red streaks and spots. 

Size: Maximum carapace width 10 cm. 

Habitat, biology, and fisheries: In sandy-muddy areas 

from depths of 10 to 100 m. Irregularly sold in 

markets, like most of the larger species of Calappa. 

They are more frequently marketed in the Philippines 

and East Asia, but do not command high prices. The 

crabs are caught mainly by trawlers, benthic nets, and 

sometimes in traps. 

Distribution: Japan, China, Southeast 

Asia, and Australia; westwards to Sri 

Lanka. 

Remarks: Aside from Calappa lophos, 

several larger species of Calappa are 

found in the area, of which only C. philargius, 

C. hepatica, and C. calappa are large 

and/or common enough to be sold in markets. 

Matuta planipes Fabricius, 1798 


FAO name: En - Flower moon crab. 

Diagnostic characters: Carapace rounded, with 2 long, 

well-developed lateral spines; anterolateral margins unevenly 

serrated. Outer surface of palm with strong oblique 

ridge. Colour: mosaic to reticulate network of maroon lines 

on a white background. 

Size: Maximum carapace width 

(excluding lateral spines). 


Mainly in soft substrates from 

depths of 10 to 40 m. Taken 

mainly as a bycatch of trawlers, 

but rarely caught in sufficient numbers to 

have significant market value. Sometimes 

caught in nets and consumed locally. 

Distribution: China, Japan, Southeast 

Asia, and Australia; westwards to India. 

Remarks: The coloration of this species is 

very distinctive and it cannot be confused 

with any other species in the area. 

chela (outer surface)


Matuta victor (Fabricius, 1781) 

Frequent synonyms / misidentifications: Matuta lunaris Forsskål, 1775 (in part); M. peronii Leach, 

1817; M. lesuerii Leach, 1817; M. crebripunctata Miers, 1877 / None. 

FAO name: En - Common moon crab. 

chela (outer surface) 

Diagnostic characters: Carapace rounded, with 2 long, well-developed lateral spines; anterolateral 

margins gently serrated. Outer surface of palm with low but distinct oblique ridge. Colour: yellowish ground 

colour with numerous fine black spots and several larger ones on carapace; legs and chelae bright yellow. 

Size: Maximum carapace width 5 cm (excluding lateral spines). 

Habitat, biology, and fisheries: Prefers sandy areas, from the intertidal zone to depths of about 20 m. 

Often caught by local communities in nets, by hand, or beach seines. 

Distribution: Southeast Asia to the Philippines, New Caledonia, Fiji, and New Hebrides. 

Remarks: The identity of Matuta victor has been badly confused with Ashtoret lunaris (Forsskål, 1775) 

and Matuta banksii Leach, 1817. The recent revision by Galil and Clark (1994) has shown that the type 

material of Cancer lunaris Forsskål, 1775, is mixed and Matuta banksii is in fact a junior synonym of 

Ashtoret lunaris. The common Indo-West Pacific species with the carapace pattern of fine black spots 

which has been identified at one time or another as Matuta lunaris, M. victor, or M. banksii, is actually 

either Matuta victor or Ashtoret lunaris.

1096 Crabs 

Ashtoret lunaris (Forsskål, 1775) 

En - Yellow moon crab. 

Maximum carapace width 5 cm. Found in sandy substrates, often near reefs or seagrass beds, from 

the intertidal zone to a depth of 50 m. Caught in nets for food in some parts of its range, often in 

good numbers. Indo-West Pacific, eastwards to Papua New Guinea and Australia. 

Calappa calappa (Linnaeus, 1758) 

(from Rüppell, 1830) chela (outer surface) 

En -Giantboxcrab. 

Maximum carapace width 13 cm. Two colour morphs are known: one uniform tan and the other 

speckled with numerous red to maroon spots. Found in rocky to shelly substrates, from depths of 

10 to 50 m. Usually caught in traps or nets. Of interest to fisheries due to its large size, but nowhere 

common enough to have major commercial importance. Also popular as a curiosity (e.g. in Hawaii). 

Indo-West Pacific, including Japan, Australia, Papua New Guinea, New Caledonia, and Hawaii.


Calappa hepatica (Linnaeus, 1758) 

En - Reef box crab. 

Maximum carapace width 8 cm. In sandy and shelly substrates, often in reefs and among seagrass 

beds, from the intertidal zone to a depth of about 100 m. Collected for food, occasionally by hand 

or in traps. Indo-West Pacific, reaching Australia, Hawaii, and French Polynesia. 

Calappa philargius (Linnaeus, 1758) 

En - Spectacled box crab. 

Maximum carapace width 12 cm. Prefers sandy to slightly muddy substrates at depths from 10 to 

100 m. Usually collected in nets or trawls. Caught for food in many parts of its range but nowhere 

very important and rarely sold in markets. Indo-West Pacific, including Korea and Japan.

1098 Crabs 

Xanthidae XANTHIDAE 

Xanthid stone and mud crabs 

Diagnostic characters: Carapace hexagonal, transversely hexagonal to transversely ovate,sometimes 

circular; dorsal surface usually ridged or granulose; frontal margin usually notched medially; 

usually 2 to 6 spines, teeth and/or lobes on each anterolateral margin. Longitudinal ridges which define 

the efferent respiratory current usually absent or strong only on posterior part of endostome; ridges not 

visible on anterior part of endostome when mouthparts pushed aside. Fingers of chela may be spoontipped. 

Legs varying in structure; propodus and dactylus with or without a special dactylo-propodal 

articulation, which is formed by a rounded submedian extension of the lateral margin, shaped to slide 

underneath a projecting button on the subproximal edge of the dactylus. Male abdominal segments 3 to 

5 immovable, fused completely or incompletely. Male first gonopod slender, slightly sinuous; distal part 

relatively simple, without complex folds, long setae usually present distally or subdistally; male second 

gonopod very short, less than 1/4 the length of male first gonopod. 

6 

5 

4 

3 

2 

1 

male abdomen 


(completely or 

incompletely) 

Habitat, biology, and fisheries: Benthic crabs with diverse habits. Most species of minor or no commercial 

importance. A single species of Atergatopsis (“egg crabs”) and 4 of the reef species of Etisus (“spooner 

crabs”) are of interest to fisheries in the area.It is important to note here that several species of xanthids 

are highly poisonous, particularly Zosimus aeneus, Lophozozymus pictor, andAtergatis floridus, and 

their consumption has caused a number of human deaths (see General Remarks). 


The Xanthidae is a very diverse group and can easily be confused with a number of families. Particularly 

difficult to distinguish are the Eriphiidae (likewise called “stone and mud crabs”) and Carpiliidae (reef crabs), 

both of which were previously included in the Xanthidae. 

Carpiliidae, Eriphiidae: adult males can be distinguished from xanthids by having the male first gonopods 

stout and cylindrical (rather than slender and sinuous), and the male second gonopods very slender, longer 

than the male first gonopod (rather than very short). Male eriphiids can also be distinguished from xanthids 

by having all the male abdominal segments freely movable, with the sutures clearly visible (versus male 

abdominal segments 3 to 5 completely fused, with sutures not dicernible). 

all male 

abdominal 

segments freely 

movable 

Eriphiidae 

Carpilidae 

anterolateral 

margin with 

only 1 tooth

Xanthidae 1099 


1a. Anterolateral margins almost entire except for a 

weak lateral tooth (Fig. 1) . . . . . . . . Atergatopsis signatus 

1b. Anterolateral margins multidentate to multispinate . . . . . → 2 

2a. Anterolateral margins each with first 2 teeth lobiform, 

not spine-tipped; adult chelipeds elongate; 

margins of legs unarmed (Fig. 2) . . . . . . Etisus laevimanus 

2b. Anterolateral margins with teeth distinct, sharp; 

adult chelipeds normal; margins of legs armed 

with sharp granules or spines . . . . . . . . . . . . . . . → 3 

3a. Anterolateral margins each with 8 strong, equal-sized teeth which curve distinctly 

forward (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etisus utilis 

3b. Anterolateral margins each with 6 strong, equal-sized triangular teeth which do not 

curve distinctly forward, often with 4 to 5 smaller teeth between them (Figs 4 and 5) . . . . . . → 4 

4a. Frontal lobes separated by narrow groove; carpus of cheliped with 1 spine on inner 

margin (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etisus dentatus 

4b. Frontal lobes separated by shallow, broad, V-shaped sulcus (cleft); carpus of cheliped 

with 2 spines on inner margin (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . Etisus splendidus 



Atergatopsis signatus (Adams and White, 1848) 

Etisus dentatus (Herbst, 1785) 

Etisus laevimanus Randall, 1840 

Etisus splendidus Rathbun, 1906 

Etisus utilis Jacquinot, 1852 

 

Fig. 2 Etisus laevimanus 

Fig. 4 Etisus dentatus 

first 2 teeth 

lobiform 

narrow 

groove 

a single weak 

lateral tooth 

Fig. 1 Atergatopsis signatus 

Fig. 3 Etisus utilis 

Fig. 5 Etisus splendidus 

8 strong, 

equal-sized 

teeth 

V-shaped 

sulcus 

References 

Alcock, A., 1898. Materials for a carcinological fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The family 

Xanthidae. J. Asiat. Soc. Bengal, 67(2), No. 1:67-233. 


Xanthidae et Trapeziidae. Addendeum Carpiliidae et Menippidae - A. Crosnier. Faune Tropicale (ORSTOM), 

24:1-400.

1100 Crabs 

Etisus laevimanus Randall, 1840 

Frequent synonyms / misidentifications: Etisus macrodactylus Bianconi, 1851; E. convexus Stimpson, 

1858; E. maculatus Heller, 1861 / None. 

FAO name: En - Smooth spooner. 

Diagnostic characters: Carapace distinctly broader than long, surfaces very smooth; anterolateral margin 

with 5 lobiform teeth (first 2 teeth never spine-tipped). Chelae very long in adults, reaching or almost 

reaching maximum width of carapace. Colour: quite variable, from dark grey to reddish brown, often with 

a patchwork of grey and dark brown. 


Habitat, biology, and fisheries: Inhabits reefs from the intertidal zone to a depth of about 20 m. Caught 

incidentally on reefs, using nets and fish traps; also collected by hand in some parts of its range. Of low 

market value, although it may be very common in certain regions, especially on disturbed reef flats. 

Distribution: Indo-West Pacific, reaching eastwards to Guam, Hawaii, and French Polynesia. 

Remarks: One of the most 

distinctive members of the 

genus because of its very 

broad and smooth carapace 

with lobiform anterolateral 

margins, and the elongate 

chelipeds.

Xanthidae 1101 

Etisus splendidus Rathbun, 1906 

Frequent synonyms / misidentifications: None / Etisus 

utilis Jacquinot, 1852; E. dentatus (Herbst, 1785). 

FAO name: En - Splendid spooner. 

Diagnostic characters: Carapace ovate, surfaces 

smooth; 8 large teeth on each anterolateral margin (often 

with smaller denticles between them); front divided into 2 

distinct lobes, separated by distinct V-shaped cleft. 

Carpus of cheliped with 2 spines on inner margin. Colour: 

red to reddish brown overall. 


Habitat, biology, and fisheries: A coral reef crab. Never 

collected in large numbers, but prized when caught, 

because of its large size. Often caught in benthic nets and 

fish traps, or by hand in intertidal reef areas. 

Distribution: Indo-West Pacific, eastwards to Hawaii and 

French Polynesia, but not yet known from Southeast Asia. 

Remarks: There are 2 species in the area 

similar to Etisus splendidus in size and 

general morphology, namely E. utilis and E. 

dentatus (see species accounts below). E. 

utilis is easily recognized by its 8 anterolateral 

teeth which are curved forwards and 

dorsoventrally flattened, whereas E. dentatus 

can be distinguished by the distinct sinus 

between the 2 frontal lobes being very narrow 

and the carpus of the cheliped possessing 

only 1 spine on the inner margin. 

Etisus utilis Jacquinot, 1852 

Frequent synonyms / misidentifications: None / Etisus 

splendidus Rathbun, 1906; E. dentatus (Herbst, 1785). 

FAO name: En - Sawedged spooner. 

Diagnostic characters: Carapace ovate, surfaces 

smooth; 8 large teeth on each anterolateral margin 

(often with smaller denticles between them); front 

divided into 2 distinctly truncate lobes, separated by 

narrow fissure. Carpus of cheliped with 2 large spines 

on inner margin. Colour: reddish brown overall; tips of 

dactylus of legs red. 


Habitat, biology, and fisheries: A coral reef 

crab. Caught by hand, in benthic nets, and fish 

traps. Often collected for food, although rarely 

in large numbers. There is some evidence that 

this species may be temporarily mildly 

poisonous in some parts of its range. 

Distribution: Indo-West Pacific, reaching 

eastwards to New Caledonia. 

Remarks: See Etisus splendidus.

1102 Crabs 

Atergatopsis signatus (Adams and White, 1848) 

En - Giant egg crab. 

Maximum carapace width 12 cm. Inhabits reefs from the intertidal zone to a depth of 25 m. 

Occasionally collected by hand or in traps because of its large size, but not a common species and 

therefore only locally of importance. Indo-West Pacific to Japan. 

Etisus dentatus (Herbst, 1785) 

En - Spiny spooner. 

Maximum carapace width 12 cm. In reefs or among rocky substrates, from the intertidal zone to a 

depth of 20 m. Occasionally collected by hand or in traps, but nowhere of significant importance. 

Indo-West Pacific, eastwards to Tahiti and Hawaii, but not recorded from most of Southeast Asia. 



Eriphiidae 1103 

Eriphiidae ERIPHIIDAE 

(= Menippidae, Oziidae) 

Eriphiid stone and mud crabs 

Diagnostic characters: Carapace hexagonal, 

transversely rectangular to transversely 

ovate; dorsal surfaces ridged or granulose; 

frontal margin notched medially; 4 teeth 

and/or lobes on each anterolateral margin. Legs 

normal. Longitudinal ridges which define efferent 

respiratory current well developed along entire 


when mouthparts pushed aside. All male 

abdominal segments distinct, movable. Male 

first gonopod stout, almost straight or gently 

curved; male second gonopod elongate, longer 

or subequal in length to male first gonopod. 

Habitat, biology, and fisheries: 1/ Benthic crabs. 

Most eriphiids are only of minor importance to 

fisheries. The more commonly collected species in 

the area are Myomenippe hardwickii, Menippe 

rumphii, andHypothalassia armata. 


The Eriphiidae can easily be confused with the Xanthidae (likewise called “stone and mud crabs”) and 

Carpiliidae. These 3 groups were previously classified together in the Xanthidae. 

Xanthidae: adult males can be distinguished from eriphiids by having abdominal segments 3 to 5 fused 

and immovable (versus all segments freely movable), the male first gonopods slender and sinuous (rather 

than stout, cylindrical), and the male second gonopods very short (rather than very elongate, longer than 

first gonopod). 

Carpiliidae: can only be effectively distinguished from eriphiids by having male abdominal segments 3 to 

5 immovable, completely fused, and the sutures not discernible (versus all male abdominal segments freely 

movable, sutures clearly visible). 

male abdominal 

segments 3-5 fused, 

immovable 

Xanthidae male abdominal 

segments 3-5 fused, 

immovable 

Carpillidae 

anterolateral 

margin with a 

single tooth 

1/ The most important commercial species of this family is probably the Australian “Tasmanian giant crab”, also known 

as the “queen crab”, Pseudocarcinus gigas (Lamarck, 1818), which occurs just outside the boundaries of the Western 

Central Pacific. It grows up to 40 cm in carapace width, at a maximum weight in excess of 12 kg. Over 50 t of these 

giant crabs are caught annually in Australia, where it commands prices of US$6 per kg, but it is even more expensive 

when exported.

1104 Crabs 


1a. Surfaces of carapace, chelipeds and legs 

with numerous sharp, long, slender 

spines (Fig. 1) . . . . . . . . . Hypothalassia armata 

1b. Surfaces of carapace, chelipeds and legs 

smooth or armed at most with low granules 

or tubercles . . . . . . . . . . . . . . . . . . → 2 

2a. Front finely granulate; anterolateral margins 

with clearly defined lobes or teeth . . . . . . → 3 

2b. Front multilobed or multidentate; anterolateral 

margins denticulate but without 

distinct lobes or teeth . . . . . . . . . . . . . . . → 4 

3a. Outer surface of palm smooth (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . Eriphia sebana 

3b. Outer surface of palm granulose (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . Eriphia smithii 

outer surface 

of palm 

smooth 

Fig. 2 Eriphia sebana 

(after Garth and Alcala, 1977) 

outer surface of 

palm granulose 

4a. Anterolateral teeth not crested (Fig. 4a); base of movable finger of larger chela with 

large, gently curved tooth (Fig. 5a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

4b. Anterolateral teeth crested (Fig. 4b); base of movable finger of larger chela with large 

molariform tooth (Fig. 5b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

lateral 

teeth 

not 

crested 

a) 

lateral 

teeth with 

crests 

Fig. 4 anterolateral margin of carapace 

5a. Palm of smaller chela slender; 

fingers slender, forceps-like; 

fingers long 

palm and fingers either subequal 

and slender 

in length, or fingers longer 

(Fig. 6a) . . . . . . . . . Epixanthus dentatus 

5b. Palm of smaller chela moderately 

stout; fingers not forcepslike; 

palm and fingers either 

subequal in length, or (usually) 

fingers shorter (Fig. 6b). . . . . . . . . . → 6 

b) 

Fig. 1 Hypothalassia armata 

Fig. 3 Eriphia smithii 

sharp, slender 

spines 

curved tooth 

a) b) molariform 

tooth 

Fig. 5 outer surface of large chela 

fingers 

short 

a) Epixanthus dentatus b) others 

Fig. 6 outer surface of small chela


6a. Dorsal surface of carapace smooth or with numerous small, shallow pits (Fig. 7) . . .Ozius guttatus 

6b. Dorsal surface of carapace with numerous small but distinct rounded granules (Fig. 8) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ozius tuberculosus 

small shallow 

pits 

7a. Dorsal surface of carapace and chelipeds granulose (Fig. 9) . . . . . . . . Myomenippe hardwickii 

7b. Dorsal surface of carapace and chelipeds smooth . . . . . . . . . . . . . . . . . . . . . . . . → 8 

8a. Basis-ischium and merus of cheliped movable, with suture demarcating them still 

distinct (Fig. 10a); reddish brown in life; non-mangrove species . . . . . . . . . . Menippe rumphii 

8b. Basis-ischium and merus of cheliped fused, immovable, with suture demarcating them 

partially to almost completely reduced (Fig. 10b); dirty-brown in life; usually mangrove 

species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myomenippe fornasinii 

outer surface 

of palm 

smooth 

Fig. 9 Myomenippe hardwickii 

(from De Man, 1887) 



Epixanthus dentatus (White, 1847) 

Eriphia sebana (Shaw and Nodder, 1803) 

Eriphia smithii (MacLeay, 1838) 

Hypothalassia armata (De Haan, 1835) 

Menippe rumphii (Fabricius, 1798) 

Myomenippe hardwickii (Gray, 1831) 

Myomenippe fornasinii (Bianconi, 1851) 

Ozius guttatus H. Milne Edwards, 1834 

Ozius tuberculosus H. Milne Edwards, 1834 

 

Fig. 7 Ozius guttatus 

(from A. Milne Edwards, 1873) 

small rounded 

tubercles 

basis-ischium 

Fig. 8 Ozius tuberculosus 


suture 

distinct 

suture 

reduced 

a) Menippe rumphii b) Myomenippe fornasinii 

Fig. 10 basis-ischium and merus of cheliped 

References 

Alcock, A. 1898. Materials for a carcinological fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The family 




24:1-400. 

merus 

basis-ischium 

merus

1106 Crabs 

Eriphia smithii MacLeay, 1838 


Eriphia sebana (Shaw and Nodder, 1803). 

FAO name: En - Rough redeyed crab. 

Diagnostic characters: Carapace rectangular, 

anterior surface granulated; anterolateral margins 

with numerous spines, but without distinct teeth or 

lobes. Base of movable finger of larger claw with 

large molarifom tooth. Colour: dark reddish 

brown overall, with bright red eyes. 


Habitat, biology, and fisheries: A reef species, 

preferring intertidal areas. Both species of 

Eriphia included here have a low fishery value and are only collected locally, although frequently, by hand 

during low tide periods. Eriphia species are said to be poisonous in some areas, but those reports could 

not been confirmed biochemically. It may be possible that upon feeding on poisonous molluscs or material, 

the crabs become toxic for a short period as well. 


to Hawaii. 

Remarks: Can only be confused with 

Eriphia sebana, which is easily distinguished 

by the smooth outer surface of the 

palm (palm covered with numerous granules 

in E. smithii). E. sebana also has a 

lighter coloured, beige-brown carapace 

(dark-reddish brown in E. smithii). 

Hypothalassia armata (De Haan, 1835) 

Frequent synonyms / misidentifications: Acanthodes 

armatus De Haan, 1835 / None. 

FAO name: En - Champagne crab. 

Diagnostic characters: Carapace smooth; anterolateral 

margins with numerous very sharp spines of differing 

sizes. Surfaces of legs and chelae with 

numerous sharp, brown-black spines of differing sizes. 

Colour: carapace reddish brown to brown, especially 

on anterior part; spines black to brown; fingers black. 

Size: Maximum carapace width 15 cm for males, females 

generally smaller. 

Habitat, biology, and fisheries: The monotypic Hypothalassia armata prefers rocky and muddy substrates, 

at depths from 30 to 540 m. It is caught in deep-water lobster pots and sometimes in bottom trawls. 

Highly valuable for human consumption because of its large size and the enlarged chelae. The common 

name, “champagne crab”, refers to its flesh, which is considered to be of a very fine quality. Occasionally 

marketed in Australia. Larger markets are being sought for this crab which is exported to Taiwan Province 

of China and Singapore, where live specimens 

command premium prices of up to 

US$40 per kg. Outside the area, it is occasionally 

caught off Western Australia; 

sometimes also collected for food in southern 

Japan and Taiwan Province of China, 

but more frequently cleaned, dried, and 

mounted for the souvenir trade. 

Distribution: Australia, Guam, Fiji, Taiwan 

Province of China, and Japan.


Menippe rumphii (Fabricius, 1798) 


Myomenippe hardwickii (Gray, 1831). 

FAO name: En - Maroon stone crab. 

Diagnostic characters: Carapace ovate, smooth, regions 

well defined; 4 broad lobiform teeth on each anterolateral 

margin. Eyes red in life. A large molariform tooth at 

base of movable finger of larger chela. Colour: carapace 

and appendages reddish brown to pinkish brown and 

maroon in adults; young crabs maroon to reddish brown, 

longitudinally striped with white. 


Habitat, biology, and fisheries: Prefers shallow to intertidal 

waters, on sandy-muddy substrates, usually under rocks. 

Menippe rumphii is occasonally fished for food, collected by hand or with nets and fish traps. There are no 

targeted fisheries for this species, although it 

can be quite common in parts of the Sunda 

Shelf. Like Myomenippe hardwickii, usually 

only the chelae are retained for sale. 

Distribution: Malaysia, Singapore, Indonesia 

Thailand, southern China and Taiwan 

Province of China. 

Remarks: Can only be confused with 

Myomenippe hardwickii, which is easily 

distinguished by the dull brown coloration, 

green eyes, and rougher carapace surface. 

Myomenippe hardwickii (Gray, 1831) 

Frequent synonyms / misidentifications: Menippe 

granulosa De Man, 1888; Myomenippe granulosa 

(Gray, 1831) / Menippe rumphii (Fabricius, 1798); 

Myomenippe fornasinii (Bianconi, 1851). 

FAO name: En - Mangrove stone crab. 

Diagnostic characters: Carapace ovate, covered 

with numerous very small granules; regions well defined; 

4 broad lobiform teeth on each anterolateral 

margin. Eyes green in life. A large molariform tooth at 

base of movable finger of larger chela. Colour: carapace 

dirty-brown overall; eyes green, fingers black. 

(from De Man, 1887) 


Habitat, biology, and fisheries: Essentially a shallow-water to intertidal mangrove species, preferring 

rocky areas, or areas densely covered by bivalves, such as Perna spp. Caught using fish traps, drift nets, 

and also taken by hand. An abundant species, caught in large quantities for its massive chelae, but no 

targeted fisheries are known. Similar to practice in the Americas with certain crab species, the chelae are 

frequently broken off and the animal is thrown back into the water. 

Distribution: Throughout Southeast Asia, 

reaching the Philippines. 

Remarks: Can be confused with 

Myomenippe fornasinii and Menippe rumphii, 

but these 2 species have much smoother 

carapace and cheliped surfaces (not granulose 

as seen inMyomenippe hardwickii). 

Menippe rumphii additionally differs by its 

reddish brown coloration and its red eyes.

1108 Crabs 

Epixanthus dentatus (White, 1847) 

En - Longfingered peeler crab. 

Maximum carapace width 7 cm. Mainly along mangroves, usually under rocks or timber. A moderately 

large species, quite common in many areas and therefore very likely collected for food by local 

populations. Indo-West Pacific in distribution, southwards reaching northern Australia. 

Eriphia sebana (Shaw and Nodder, 1803) 

En - Smooth redeyed crab. 

Maximum carapace width 8 cm. A rocky-shore or reef-dwelling species. Occasionally collected for 

food, but never in large numbers. There have been reports that this species is occasionally mildly 

poisonous in some parts of its range (see also E. smithii). Throughout Indo-West Pacific, including 

Hawaii and various parts of Southeast Asia. 

(from Garth and Alcala, 1977)


Myomenippe fornasinii (Bianconi, 1851) 

En - Smooth stone crab. 

Maximum carapace width 9 cm. A littoral species, with preference for rocky shores with muddy-sand 

bottoms, commonly found under rocks and timber, and in crevices in mangroves. Probably occasionally 

collected for food by local populations for its large size. Occurs in parts of the Indian Ocean 


Ozius guttatus H. Milne Edwards, 1834 

En - Spottedbelly rock crab. 

Maximum carapace width 9 cm. Along rocky shores in intertidal to shallow subtidal waters, 

sometimes in estuaries, usually hiding in crevices. Occasionally collected by hand for human 

consumption. Indo-West Pacific, from the Indian Ocean to Southeast Asia, Japan, and New 

Caledonia. 


Ozius tuberculosus H. Milne Edwards, 1834 

En - Beaded rock crab. 

Maximum carapace width 9 cm. Usually along rocky shores in intertidal to shallow subtidal waters. 

Occasionally collected by hand or traps for human consumption. Indo-West Pacific, known from 

Mauritius and southern India to Southeast Asia, China, Japan, and New Caledonia. 

(from A. Milne Edwards, 1873)

1110 Crabs 

Carpiliidae CARPILIIDAE 

Reef crabs 

Diagnostic characters: Carapace transversely 

ovate; dorsal surface smooth, distinctly convex longitudinally 

and transversely; front entire; a single low, 

small tooth on each anterolateral margin of carapace. 

Legs simple. Longitudinal ridges which 

define efferent respiratory current usually 

absent or strongly developed on 

posterior part of endostome only; ridges 6 

not clearly visible on anterior part of 

5 

endostome when mouthparts pushed 

aside. Male abdominal segments 3 to 4 

5 immovable, completely fused. Male 3 

first gonopod stout, almost straight 2 

or gently curved; male second 1 

segments 3-5 


gonopod elongate, longer or subequal 

completely 

with a single tooth 

in length to male first gonopod. male abdomen fused 

Habitat, biology, and fisheries: Benthic reef crabs. A single genus of Carpiliidae, Carpilius (with only 2 

species in the Pacific), has fishery value. Both Pacific species of Carpilius, C. maculatus and C. convexus, 

are occasionally collected for food. 


Only the Xanthidae and Eriphiidae (both families known as “stone and mud crabs”) can easily be confused 

with carpiliids. These 3 taxa were all previously classified in a single family (Xanthidae). 

Xanthidae: can be distinguished from carpiliids by the shape of the male first gonopods, which are slender 

and sinuous (rather than stout, cylindrical), and the male second gonopods, which are very short (rather 

than very elongate, longer than first gonopod). 

Eriphiidae: can only be effectively distinguished from carpiliids by having all the male abdominal segments 

freely movable, with the sutures clearly visible (versus male abdominal segments 3 to 5 completely fused, 

sutures not discernible). 

anterolateral 

margin with more 

than 1 tooth 


1a. Cream to pink ground colour in life, carapace with 9 large violet to maroon spots 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carpilius maculatus 

1b. Uniform red to reddish brown colour in life . . . . . . . . . . . . . . . . . . . . Carpilius convexus 



Carpilius convexus (Forsskål, 1775) 

Carpilius maculatus (Linnaeus, 1758) 

 

Xanthidae 


segments freely movable 

Eriphiidae 

References 

Alcock, A. 1898. Materials for a carcinological fauna of India. No. 3. The Brachyura Cyclometopa. Part I. The family 




24:1-400.

Carpiliidae 1111 

Carpilius convexus (Forsskål, 1775) 


FAO name: En - Red reef crab. 

Diagnostic characters: Carapace ovate; dorsal 

surface very smooth and convex. Colour: uniform 

red to reddish brown, with irregular dark brown 

patches on the dorsal surface of carapace. 


Habitat, biology, and fisheries: A reef crab.Biology 

and fisheries similar to Carpilius maculatus (see 

below). 

Distribution: Indo-West Pacific, reaching Hawaii 

and French Polynesia. 

Remarks: C. convexus is easily distinguished from 

C. maculatus by its distinct coloration which remains 

even after preservation. 

Carpilius maculatus (Linnaeus, 1758) 


FAO name: En - Spotted reef crab. 

Diagnostic characters: Carapace ovate; dorsal 

surface very smooth and convex. Colour: cream 

to pink ground colour, with 9 large violet to maroon 

spots on dorsal surface of carapace: 3 on median 

region, 2 on posterior region, 2 on anterolateral 

region, and 2 around the orbits. 


Habitat, biology, and fisheries: A reef crab. Collected 

extensively for food, although never in large 

quantities. Frequently seen in markets of East Asia 

and parts of Indonesia, but only in small numbers. 

Usually collected by hand or with baited 

traps. There have been reports that this 

species is poisonous, but this could not 

been confirmed by biochemical tests. It is 

possible that after feeding on poisonous 

molluscs, the crabs become toxic for a 

short period as well. 


Hawaii and French Polynesia. 

Remarks: The only other species of Carpilius 

in the area is C. convexus, which can 

easily be distinguished by its very different 

coloration.

1112 Crabs 

Pilumnidae PILUMNIDAE 


transversely rectangular or 

transversely ovate; dorsal surface convex, 

smooth to granulated; frontal margin entire to 

multilobate; usually 1 to 4 teeth or lobes on 

each anterolateral margin. Longitudinal 

ridges defining efferent respiratory current 

usually well developed along entire 

endostome, ridges visible on anterior part of 

endostome when mouthparts pushed aside. 

Legs normal. Male abdominal segments 3 to 

5 freely movable. Male first gonopod slender, 

usually S-shaped, distal part simple; 

male second gonopod very short, sigmoid. 

Habitat, biology, and fisheries: Benthic 

crabs with diverse habits. Most species in this 

family are of no commercial value. The moderately 

large-sized Galene bispinosa has minor 

economic importance. 

Hairy crabs 

Remarks: Despite their common name, “hairy crabs”, many pilumnids (including Galene bispinosa) are 

actually not very setose (or “hairy”). 


The Pilumnidae is a very diverse group and its taxonomy 

remains unsettled. As their general (usually hexagonal) 

carapace shape is similar to those of xanthids, eriphiids, 

and goneplacids, the safest way to identify a pilumnid 

species is to examine the male abdomen and gonopods. 

All pilumnids share very similar male abdominal, male 

pleopodal, and larval characters. 

Xanthidae: male abdominal segments 3 to 5 fused (instead 

of freely movable). 

Eriphiidae: male first gonopods stout (rather than slender 

and sinuous); male second gonopods long (rather than 

very short). 


segments freely movable 

Xanthidae 

male abdominal 

segments fused, 

immovable 

Goneplacidae: generally have stouter male first gonopods and/or proportionately longer male second 

gonopods; some species with male abdominal segments 3 to 5 fused. 

Eriphiidae Goneplacidae 

References 

Balss, H. 1933. Beitrage zur Kenntnis der Gattungen Pilumnus (Crustacea Dekapoda) und verwandter Gattungen. 

Capita Zoologica, 4(3):1-47. 

Ng, P.K.L. 1987. The Indo-Pacific Pilumnidae II. A revision of the genus Rhizopa Stimpson, 1858 and the status of the 

Rhizopinae Stimpson, 1858 (Crustacea: Decapoda: Brachyura). Indo-Malayan Zoology, 4(1):69-111.

Pilumnidae 1113 


Galene bispinosa (Herbst, 1783) 

Frequent synonyms / misidentifications: Podopilumnus fittoni M’Coy, 1849; Gecarcinus trispinosus 

Desmarest, 1822; Galene granulosa Miers, 1884 / None. 

FAO name: En - Square-shelled crab. 

Diagnostic characters: Carapace subpentagonal, dorsal surface gently convex, lateral regions with 

numerous small, rounded granules; 2 or 3 distinct conical teeth on each lateral margin (2 teeth always 

distinct). Chelipeds stout, surfaces finely granulated. Colour: tan to purplish tan. 


Habitat, biology, and fisheries: In shallower waters, down to depths of about 100 m, living on muddy 

substrates. Caught mainly by bottom trawls, often in large quantities. Of low fishery value, although very 

abundant in some areas. Only occasionally seen in markets and sold for low prices. Fished mainly in 

Thailand and various parts of Indonesia. 

Distribution: India and Southeast Asia to Australia and Japan.

1114 Crabs 

Goneplacidae GONEPLACIDAE 

Rhomboid crabs 


transversely rectangular, trapezoidal, or 

transversely ovate; dorsal surface convex, usually 

smooth; frontal margin usually entire, sometimes 

multilobate; anterolateral margin usually armed with 

1 to 4 teeth or lobes, or entire. Male abdominal 

segments 3 to 5 distinct, movable or fused and 

immovable. Male first gonopod moderately stout, 

gently curved or sinuous; male second gonopod 

relatively short to elongate, but usually shorter 

than male first gonopod. 

Habitat, biology, and fisheries: Benthic crabs with 

diverse habits. Most species in this family are of little 

or no commercial value. The relatively common and 

large Carcinoplax longimanus has a minor importance 

to fisheries in the Western Central Pacific. 


The Goneplacidae is doubtless a very heterogeneous group.Obviously, the genera included here were assigned 

to this family due to lacking evidence to place them into any others of the known families. Although the angular 

carapace of most goneplacids readily separates them from species of other families, a clear definition of the 

Goneplacidae is not known. Accordingly, any comparisons with outside taxa are very difficult and must be done 

on a genus by genus basis. Carcinoplax, the only genus in the area that includes an edible species, resembles 

in general body shape some species of Xanthidae, Eriphiidae, and Carpiliidae. 

Xanthidae, Eriphiidae, and Carpiliidae: compared to species of these families, in Carcinoplax, the carapace 

is clearly more ovate in shape and the adult male chelipeds are extremely elongated. In addition, the male 

second gonopods of Carcinoplax are intermediate in relative length between xanthids (very short, as in 

pilumnids) and eriphiids (very long). 

References 

Guinot, D. 1989. Le genre Carcinoplax H. Milne Edwards, 1852 (Crustacea: Decapoda: Goneplacidae). In Résultats des 

Campagnes MUSORSTOM, 4, edited by J. Forest. Mém. Mus. natn. Hist. nat., 144:265-345. 

Tesch, J.J. 1918. The Decapoda Brachyura of the Siboga Expedition. II. Goneplacidae and Pinnotheridae. Siboga Exped. 

Monogr., 39c(1):149-295. 


Carcinoplax longimanus (De Haan, 1833) 

Frequent synonyms / misidentifications: Carcinoplax longimanus 

japonicus Doflein, 1904; Carcinoplax longimanus typicus 

Doflein, 1904 / None. 

FAO name: En - Long-armed crab. 

Diagnostic characters: Carapace ovoid; dorsal surface convex 

and smooth; lateral margin with 3 strong teeth in juveniles, 

becoming small to almost indiscernible in adults. Male chelipeds 

very elongate. Colour: red to pink overall. 


Habitat, biology, and fisheries: On muddy substrates, most 

commonly found in deeper waters from depths of 100 to 

800 m. Fished mainly in southern Japan and southern 

China. Taken incidentally in major fishery operations, 

mainly by bottom trawls and often in large numbers. Only 

larger specimens are sold, but have a low value in most 

markets. 

Distribution: Japan, Taiwan Province of China, China, 

Philippines, and Thailand. 

male chelipeds 

elongate 

carapace usually angular 



Portunidae 1115 

Portunidae PORTUNIDAE 

Swimming crabs 

Diagnostic characters: Carapace hexagonal, transversely ovate to transversely hexagonal,sometimes 

circular; dorsal surface relatively flat to gently convex, usually ridged or granulose; front broad, 

margin usually multidentate; usually 5 to 9 teeth on each anterolateral margin, posterolateral margins 

usually distinctly converging. Endopodite of second maxillipeds with strongly developed lobe on inner 

margin. Legs laterally flattened to varying degrees, last 2 segments of last pair paddle-like. Male 

abdominal segments 3 to 5 completely fused, immovable. 

5 

4 

3 

male abdomen 

segments 

3-5 fused, 

immovable 

male abdomen sedments 

3-5 immovable 

last pair of legs 

paddle-like 


with 5-9 teeth 

Habitat, biology, and fisheries: Benthic to semipelagic crabs with diverse habits. Many species of great 

fishery value, notably Scylla serrata, Portunus pelagicus, P. sanguinolentus, P. trituberculatus, and 

Charybdis feriatus. 


Portunids may be confused with spanner crabs (Raninidae) and moon crabs (Calappidae: Matutinae), 

which also possess paddle-like legs. They can be separated from portunids as follows: 

Raninidae: carapace usually longitudinally ovate; sternum very narrow; fingers of chela strongly bent; meri 

of third maxillipeds triangular (quadrate in portunids). 

Calappidae (subfamily Matutinae): carapace circular to subcircular; at least last 3 pairs of legs paddle-like, 

(not only the last pair); meri of third maxillipeds triangular in cross-section (quadrate in portunids). 

fingers 

strongly 

bent 

Raninidae 

carapace 

longitudinally ovate 

Calappidae (subfamily Matutinae) 

all legs 

paddle-like

1116 Crabs 


1a. Carapace with 2 anterolateral teeth; eyes very long, reaching lateral edge of carapace 

(Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Podophthalmus vigil 

1b. Carapace with more than 2 anterolateral teeth; eyes normal in size . . . . . . . . . . . . . . . → 2 

2a. Carapace rounded; ventral surface of palm with stridulatory (sound-producing) ridges 

(Fig. 2a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ovalipes punctatus 

2b. Carapace transversely ovate; palm without any stridulatory (sound-producing) ridges 

(Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2 anterolateral 

teeth 

Fig. 1 Podophthalmus vigil 

stridulatory ridges 

a) Ovalipes puntatus b) other species 

Fig. 2 chela in ventral view 

no stridulatory ridges 

3a. Five to 7 teeth on each anterolateral margin (Fig. 3a-c) . . . . . . . . . . . . . . . . . . . . . → 4 

3b. Nine teeth on each anterolateral margin (Fig. 3d) . . . . . . . . . . . . . . . . . . . . . . . . → 12 

5 teeth 

6 teeth 

a) b) 

7 teeth 9 teeth 

Fig. 3 lateral margin of carapace (dorsal view) 

4a. Width of frontal-orbital border not much less than greatest width of carapace; 5 teeth 

on each anterolateral margin (first tooth sometimes with accessory denticle) (Fig. 4a) . . . . . → 5 

4b. Width of frontal-orbital border distinctly less than greatest width of carapace; 6 or 7 teeth 

on each anterolateral margin (Fig. 4b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

5a. Basal antennal segment with a smooth or granulated ridge (Fig. 5a) . . . . . . .Thalamita crenata 

5b. Basal antennal segment with several sharp spines (Fig. 5b) . . . . . . . . . . Thalamita spinimana 

fronto-orbital 

border very wide 

fronto-orbital border 

moderately wide 

a) Thalamita b) Charybdis 


almost smooth 

c) 

a) Thalamita crenata b) Thalamita spinimana 

d) 

distinctly granular spiniform 

Fig. 5 basal antennal segment


6a. Posterior border of carapace forming an angular junction with posterolateral border 

(Fig. 6a); merus of cheliped without distal spine on posterior border . . . . . . . Charybdis truncata 

6b. Posterior border of carapace forming a curve with posterolateral border (Fig. 6b); merus 

of cheliped with distal spine on posterior border . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

7a. Carapace with distinct ridges or granular patches behind level of last pair of anterolateral 

teeth (Fig. 7a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Charybdis natator 

7b. Carapace without distinct ridges or granular patches behind level of last pair of 

anterolateral teeth (Fig. 7b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

angular 

a) Charybdis truncata 

rounded 

b) other species 

of Charybdis 

Fig. 6 left side of carapace (dorsal view) 

distinct 

ridges 

no 

ridges 

a) Charybdis natator b) others 

Fig. 7 left side of carapace (dorsal view) 

8a. Merus of cheliped with 2 spines on anterior border; palm with 2 spines on upper surface 

(Fig. 8a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Charybdis anisodon 

8b. Merus of cheliped with 3 or 4 spines on anterior border; palm with more than 2 spines 

on upper surface (Fig. 8b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

9a. First anterolateral tooth not truncate or notched (Fig. 9a) . . . . . . . . . . . . Charybdis annulata 

9b. First anterolateral tooth truncate or notched (Fig. 9b) . . . . . . . . . . . . . . . . . . . . . . → 10 

2 spines 

2 

spines 

4 spines 

3 

spines 

a) Charybdis anisodon b) others 

Fig. 8 right cheliped (dorsal view) 

1 st tooth simple 

1 st tooth notched 

a) Charybdis annulata b) others 

Fig. 9 lateral margin of carapace (dorsal view) 

10a. Palm of cheliped with 4 spines on upper surface (Fig. 10a); male abdominal segment 4 

keeled (Fig. 11a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Charybdis feriatus 

10b. Palm of cheliped with 5 spines on upper surface (Fig. 10b); male abdominal segment 4 

not keeled (Fig. 11b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11 

4 spines 5 spines 

a) b) 

Fig. 10 right cheliped (dorsal view) a) b) 

keel 

no keel 

Fig. 11 male abdomen

1118 Crabs 

11a. Palm with well-developed spines (Fig. 12a); male abdominal segment 6 with convex 

lateral borders (Fig. 13a); last anterolateral tooth smallest and spiniform, not projecting 

beyond preceding tooth (Fig. 14a) . . . . . . . . . . . . . . . . . . . . . . . . Charybdis japonica 

11b. Palm with poorly developed spines (Fig. 12b); male abdominal segment 6 with lateral 

borders parallel in proximal half (Fig. 13b); last anterolateral tooth elongate, projecting 

laterally beyond preceding tooth (Fig. 14b) . . . . . . . . . . . . . . . . . . . . . Charybdis affinis 

12a. Last anterolateral tooth subequal in size to others (Fig. 15a) . . . . . . . . . . . . . . . . . . → 13 

12b. Last anterolateral tooth at least 2 times larger than others (Fig. 15b) . . . . . . . . . . . . . . → 16 

strong spines 

a) Charybdis japonica 

weak spines 

b) Charybdis affinis 

Fig. 12 right cheliped 

gently 

convex 


almost 

straight 



last 2 teeth 

subequal 


last tooth 

larger than 

adjacent one 


Fig. 14 anterolateral teeth 

last a) Scylla 

anterolateral 

tooth enlarged 

b) Portunus 

Fig. 15 anterolateral teeth 

13a. Carpus of cheliped with only 1 low to very low granule on outer surface, never spiniform 

(Fig. 16a); colour of palm usually with at least some patches of orange or yellow in life . . . . → 14 

13b. Carpus of cheliped with 2 distinct spiniform or sharp granules or spines on outer surface 

(Fig. 16b); colour of palm in life green to purple . . . . . . . . . . . . . . . . . . . . . . . . . → 15 

14a. Frontal margin usually with sharp teeth (Fig. 17a); palm usually with distinct, sharp 

spines (Fig. 18a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scylla paramamosain 

14b. Frontal margin usually with rounded teeth (Fig. 17b); palm usually with reduced, blunt 

spines (Fig. 18b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scylla olivacea 

15a. Frontal margin usually with rounded teeth (Fig. 19a); sharp granules on palm and carpus 

never spiniform; colour in life: carapace usually very dark green to black, outer surface 

of palm purple and never with marbled pattern, last legs marbled only in males . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scylla tranquebarica 

15b. Frontal margin usually with sharp teeth (Fig. 19b); sharp granules on palm and carpus 

often spiniform; colour in life: carapace usually green to olive-green, outer surface of 

palm green and often with marbled pattern, last legs marbled both in males and females 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scylla serrata 

b) 

a) 

unarmed 

with 2 

spines 

Fig. 16 carpus 

of cheliped 

sharp teeth 

a) Scylla paramamosain 

rounded teeth 

b) Scylla olivacea 

Fig. 17 frontal margin of 

carapace (dorsal view) 

sharp 

spines 

a) Scylla paramamosain 

blunt spines 

b) Scylla olivacea 

Fig. 18 right cheliped 

rounded teeth 

a) Scylla tranquebarica 

sharp teeth 

b) Scylla serrata 

Fig. 19 frontal margin of 

carapace (dorsal view)


16a. Carapace with 3 purple to red spots on 

posterior half (Fig. 20) . . . Portunus sanguinolentus 

16b. Carapace marbled or with uniform coloration 

. . . . . . . . . . . . . . . . . . . . . . → 17 

17a. Front with 4 teeth (Fig. 21a); inner margin 

of merus of cheliped with 3 spines 

(Fig. 22a) . . . . . . . . . . . . . Portunus pelagicus 

17b. Front with 3 teeth (Fig. 21b); inner margin 

of merus of cheliped with 4 spines 

(Fig. 22b) . . . . . . . . . . Portunus trituberculatus 



Charybdis affinis Dana, 1852 

Charybdis anisodon (De Haan, 1850) 

Charybdis annulata (Fabricius, 1798) 

Charybdis feriatus (Linnaeus, 1758) 

Charybdis japonica (A. Milne Edwards, 1861) 

Charybdis natator (Herbst, 1794) 

Charybdis truncata (Fabricius, 1798) 

Ovalipes punctatus (De Haan, 1833) 

Podophthalmus vigil (Fabricius, 1798) 

Portunus pelagicus (Linnaeus, 1758) 

Portunus sanguinolentus (Herbst, 1783) 

Portunus trituberculatus (Miers, 1876) 

Scylla olivacea (Herbst, 1796) 

Scylla serrata (Forsskål, 1775) 

Scylla paramamosain Estampodor, 1949 

Scylla tranquebarica (Fabricius, 1798) 

Thalamita crenata (Latreille, 1829) 

Thalamita spinimana (Dana, 1852) 

 

4teeth 

3teeth 

a) b) 

Fig. 21 frontal margin of carapace (dorsal view) 

Fig. 20 Portunus sanguinolentus 

3 spines 

4spines 

a) b) 

Fig. 22 merus of cheliped (inner margin) 

References 

Keenan, C.P., P.J.F. Davie, and D.L. Mann. 1998. A revision of the genus Scylla De Haan (Crustacea: Decapoda: 

Brachyura: Portunidae). Raffles Bull. Zool., 46(1):in press. 

Leene, J.E. 1938. The Decapoda Brachyura of the Siboga Expedition. VII. Brachygnatha: Portunidae. Siboga Exped. 

Monogr., 39c(131):1-156. 

Stephenson, W. 1972. An annotated check list and key to the Indo-West Pacific swimming crabs (Crustacea: Decapoda: 

Portunidae). Royal Society of New Zealand Bulletin, 10:1-64.

1120 Crabs 

Charybdis feriatus (Linnaeus, 1758) 

Frequent synonyms / misidentifications: Charybdis crucifer (Fabricius, 1792); C. cruciata (Herbst, 

1794) / None. 

FAO name: En -Crucifixcrab. 

Diagnostic characters: Carapace ovate; 5 distinct teeth on each anterolateral margin. Colour: distinctive 

pattern of longitudinal stripes of maroon and white, usually with distinct white cross on median part of 

gastric region; legs and pincers with numerous scattered white spots. 


Habitat, biology, and fisheries: Prefers sandy to sandy-muddy substrates, at depths from 30 to 60 m. 

Collected mainly by bottom trawls, sometimes by traps and nets. The commercially most important species 

of Charybdis. LikePortunus spp., Charybdis feriatus is more delicate than Scylla, and is frequently sold 

frozen. The lucrative and booming live-seafood market, however, is seeing the increased use of aquaria to 

keep these species alive. The crucifix crab is especially important in markets in East Asia where it 

commands substantially higher premium prices than Portunus spp., being sold for US$8 to US$15 per kg. 

Distribution: Widely distributed in the Indo-West Pacific, reaching Japan and Australia. 

Remarks: There are several species ofCharybdis in the area which are also occasionally fished and infrequently 

appear in markets. These include C. affinis Dana, 1852, C. acuta (A. Milne Edwards, 1869), C. anisodon 

(De Haan, 1850), C. annulata (Fabricius, 1798), C. natator (Herbst, 1789) and C. truncata (Fabricius, 1798). 

They are all easily distinguished by carapace and cheliped armature features.


Charybdis japonica (A. Milne Edwards, 1861) 


None. 

FAO name: En - Japanese swimming crab. 

Diagnostic characters: Carapace without 

transverse ridges behind last anterolateral tooth; 

frontal teeth acutely triangular; anterolateral 

teeth all acutely triangular. Posterior border of 

propodus of legs serrated. Palm with 5 sharp 

spines, longitudinal ridges on palm granulated. 

Colour: carapace white with large greyish 

patches, tips of anterolateral teeth reddish 

brown; fingers red and white. 


Habitat, biology, and fisheries: Occurs just offshore 

on muddy, sandy, or stony substrates. 

Taken mainly by trawlers or in nets as incidental 

catch. Although locally common, there is 

no sustained fishery for this species. 

Fished mainly in Japanese and Chinese 

waters. 

Distribution: Japan, China, Taiwan Province 

of China, Thailand, and Malaysia. 

Remarks: See Charybdis feriatus. The 

above characters and coloration of C. japonica 

readily distinguish it from all other 

species of Charybdis. 

Charybdis natator (Herbst, 1789) 


None / None. 

FAO name: En - Ridged swimming crab. 

Diagnostic characters: Carapace with densely 

covered with very short pubescence which is 

absent on several distinct transverse granulated 

ridges in anterior half. Colour: orangish red 

overall, with ridges on carapace and legs dark 

reddish brown. 


Habitat, biology, and fisheries: Near or in 

rocky-sandy substrates, sometimes near reefs, 

from depths of 5 to 40 m. Charybdis natator 

is caught incidentally by trawlers, and 

has some commercial value because of its 

large size. 

Distribution: China, Taiwan Province of 

China, Philippines, Thailand, Indonesia, 

Malaysia, Singapore, and Australia. 

Remarks: See Charybdis feriatus.

1122 Crabs 

Ovalipes punctatus (De Haan, 1833) 


FAO name: En - Sand crab. 

(after Shen and Dai, 1964) 

Diagnostic characters: Carapace rounded, surfaces finely granular, appearing almost smooth; 4 

well-developed teeth on each anterolateral margin; stridulatory ridges present on ventral surface of palm. 

Colour: carapace reddish brown to maroon, margins lighter coloured, with scattered dirty-white and darker 

spots, white gastric depression, margins lighter coloured; dactylus of fourth walking leg bluish purple. 

Size: Maximum carapace width 9.5 cm. 

Habitat, biology, and fisheries: Found from depths of 30 to 50 m. Caught mainly by trawls or dredges, 

and fished intensively in southern Japan, Taiwan Province of China, and China (mainly off southern China 

and Japan). Usually, only the chelae are sold in markets, where they command premium prices. 

Distribution: Throughout the 

northern and southern hemispheres 

in the Pacific. In the 

area, it occurs in the northern 

part of the South China Sea 

and in Queensland (Australia). 

Remarks: Several species of 

Ovalipes of minor commercial 

importance are known, notably 

O. australiensis Stephenson 

and Mees, 1968, from 

Australia, which, however, 

does not occur in the Western 

Central Pacific. It can easily be 

separated from O. punctatus by the condition of the carapace surface (finely granular in O. punctatus, but 

coarsely granular in O. australiensis). In addition, O. australiensis has 2 large, clear pigmented ovate spots 

on the posterolateral region (absent in O. punctatus).


Podophthalmus vigil (Fabricius, 1798) 


FAO name: En - Sentinel crab. 

Diagnostic characters: Carapace distinctly broader than long; anterior margin much broader than 

posterior margin, with posterolateral margins converging strongly towards narrow posterior carapace 

margin; orbits very broad. Eyes very long, reaching to or extending beyond edge of carapace. Colour: 

carapace green; chelipeds and parts of legs violet to maroon in adults. 


Habitat, biology, and fisheries: On sandy to muddy substrates in offshore waters. Occasionally caught 

by offshore trawlers, although rarely in large numbers. When marketed, it commands prices similar to those 

for Portunus pelagicus. 

Distribution: Indo-West Pacific. 

Remarks: Three species of Podophthalmus are known. P. vigil is the only large species that shows the 

colour pattern described above, and the only species of the genus with commercial value.

1124 Crabs 

Portunus pelagicus (Linnaeus, 1758) SCD 

Frequent synonyms / misidentifications: Portunus mauritianus Ward, 1942 / Portunus trituberculatus 

(Miers, 1876). 

FAO name: En - Flower crab. 

 

Diagnostic characters: Carapace rough to granulose, regions discernible; front with 4 acutely triangular 

teeth; 9 teeth on each anterolateral margin, the last tooth 2 to 4 times larger than preceding teeth. Chelae 

elongate in males; larger chela with conical tooth at base of fingers; pollex ridged. Colour: males with blue 

markings, females dull green. 

Size: Maximum carapace width 20 cm for males (including lateral teeth). 

Habitat, biology, and fisheries: Prefers sandy to sandy-muddy substrates in shallow waters down to a 

depth of 50 m, including areas near reefs, mangroves, and in seagrass and algal beds. Juveniles tend to 

occur in shallow intertidal areas. The crabs mature at about 1 year. Collected mainly by artisanal traps, 

trawls, beach seines, cylindrical wire traps, folding traps, pots, hop nets, drop nets, and sunken crab gill 

nets. In shallow waters, beach seines, rakes, and dab nets are used. Although sold for lower prices than 

Scylla, crabs of Portunus are taken in much larger quantities. They are caught in enormous numbers for 

sale in local markets (frozen or fresh) and for the crab-flesh canning industry. Many species of Portunus 

are commercially collected in the area. Among the 3 more frequently collected species included here, P. 

pelagicus is most widely sold in markets in Southeast Asia, including the Philippines. The market price 

varies from US$3 to US$5 per kg for fresh crabs, and from about US$5 to US$8 for live crabs. The fisheries 

for this species is well managed in Australia. From 1990 to 1995, the reported yearly catch of P. pelagicus 

from the Western Central Pacific (Australia, Indonesia, and Thailand) ranged from around 36 700 to 

48 000 t (FAO Yearbook of Fishery Statistics). 

Distribution: Throughout 

Indo-West Pacific. 

Remarks: May be confused 

with P. trituberculatus, which 

resembles a large stocky female 

of P. pelagicus. P. trituberculatus, 

however, can 

easily be distinguished by 

having only 3 frontal teeth (4 

teeth in P. pelagicus), and by 

the presence of 4 spines on 

the inner margin of the chelipedal 

merus (only 3 spines in 

P. pelagicus).


Portunus sanguinolentus (Herbst, 1783) 


None / None. 

FAO name: En - Three-spot swimming crab. 

Diagnostic characters: Carapace finely granulose, 

regions just discernible; 9 teeth on each 

anterolateral margin, the last tooth 2 to 3 times 

larger than preceding teeth. Chelae elongated in 

males; larger chela with conical tooth at base of 

fingers; pollex ridged. Colour: olive to dark 

green, with 3 prominent maroon to red spots on 

posterior 1/3 of carapace. 


Habitat, biology, and fisheries: Occurs 

on sandy to sandy-muddy substrates, from 

the intertidal zone (especially juveniles) to 

depths of 30 m. Collected mainly by nets or 

seines. This species is less common than 

Portunus pelagicus, and appears only occasionally 

in markets. It is priced similarly 

to, or for slightly lower prices as, P. pelagicus. 


Remarks: This species can be easily separated 

from all other Portunus species by its 

very distinctive colour markings. 

Portunus trituberculatus (Miers, 1876) 


None / Portunus pelagicus (Linnaeus, 1758). 

FAO name: En - Horse crab. 

Diagnostic characters: Carapace rough to 

granulose, regions discernible; front with 3 

acutely triangular teeth; 9 teeth on each anterolateral 

margin, the last tooth much larger than 

preceding teeth. Larger chela with conical tooth 

at base of fingers; pollex ridged. Colour: carapace 

dull green to brown. 

Size: Maximum carapace width 15 cm (males). 

Habitat, biology, and fisheries: Prefers sandy to 

sandy-muddy substrates in shallow waters, up to 

depths of 50 m. Caught mainly by trawls. A commercially 

very important species in Japan and 

collected in large numbers in some areas. 


Remarks: This species is perhaps closest 

to Portunus pelagicus in its general appearance, 

resembling a large stocky female of 

that species. P. pelagicus, however, is easily 

distinguished by having 4 frontal teeth 

(only 3 teeth in P. trituberculatus) and by 

the presence of 3 spines on the inner margin 

of the chelipedal merus (4 spines in P. 

trituberculatus).

1126 Crabs 

Scylla serrata (Forsskål, 1775) MUD 

Frequent synonyms / misidentifications: Acheolus crassimanus MacLeay, 1838; Scylla serrata var. 

oceanica Dana, 1852 / see Remarks. 

FAO name: En - Giant mud crab. 


smooth, with strong transverse 

ridges; H-shaped gastric 

groove deep; relatively broad frontal 

lobes, all more or less in line 

with each other; broad anterolateral 

teeth, projecting obliquely outwards. 

Well-developed spines 

present on outer surface of chelipedal 

carpus and anterior and 

posterior dorsal parts of palm. 

Colour: carapace green to almost 

black; legs may be marbled. 


between 25 and 28 cm (males); 

(from Rüppell, 1830) 

maximum weight between 2 and 

3kg. 

Habitat, biology, and fisheries: Scylla serrata prefers more oceanic waters, usually found just offshore on 

soft muddy bottoms. Crabs can be caught up to 50 km offshore as they migrate there to spawn. The other 

3 species of Scylla included here prefer mangroves in continental shelves with less saline waters. All species 

of Scylla dig deep burrows in mangroves and soft substrates in shallow or intertidal waters. Species of Scylla 

are collected mainly using trawls, traps, baited wire mesh pots, hooking, and by hand throughout their 

ranges. From 1990 to 1995, the reported yearly catch of Scylla serrata from the Western Central Pacific 

ranged from around 6 150 to 18 600 t (FAO Yearbook of Fishery Statistics). It must be pointed out, however, 

that these figures cover all 4 species of Scylla recognized here (see remarks on that problem below). 

Species of Scylla are almost always marketed alive. At present, the main markets are Taiwan Province of 

China, Hong Kong (China), and Singapore, where large crabs (so-called “meat crabs”) and females with 

ripe ovaries (“roe crabs”) command premium prices. For both kinds of crabs, current demand still far 

exceeds the supply. They are always sold for high prices, ranging from US$5 to US$10 per kg. “Roe crabs” 

can even cost 25 to 50% more. In Australia, they are sold for an average price of US$6 per kg. Currently, 

the largest exporters of mud crabs in Asia are Indonesia, Sri Lanka, India, and Bangladesh, with the markets 

in Myanmar, Viet Nam, and Pakistan picking up rapidly. S. serrata and S. olivacea are also the main food 

species in Australia. There is no clear management for the 4 species of Scylla in Southeast and East Asia. 

In Australia, the fishery for S. serrata and S. olivacea is quite intense (700 t collected between 1989 to 1990, 

mainly from Queensland with over 400 t) but is reasonably well managed. There have been attempts to 

culture the crabs in captivity, but none of the closed-cycle enterprises have gone commercial. Many Scylla 

crab farms rely on the tide to bring megalopae or late zoeae into ponds, where the crabs grow out. 

Alternatively, many farms in Southeast Asia keep smaller or freshly moulted crabs (so-called “water crabs” 

because of the consistency of their flesh) caught from natural stocks to grow them out or add on more flesh. 

Similarly, female crabs of all 4 species are kept until their ovaries are full to improve their market prices. 

Distribution: All 4 species of 

Scylla apparently have a wide 

Indo-West Pacific distribution. 

Scylla serrata, has been introduced 

to Hawaii from Samoa 

55 years ago, and has become 

established there. 

Remarks: The taxonomy of 

the genus Scylla has been terribly 

confused and is still difficult. 

While generally a single, 

supposedly widely distributed 

species is recognized, namely 

S. serrata, it is now known that the genus includes 4 species. Recent research in Australia (Keenan et al., 

1998) has clearly shown, using morphological, DNA, and allozyme data, that there are 4 species of Scylla.


The differences in allozyme and mtDNA are substantial, but the morphological features which separate the 

4 species are rather subtle and sometimes difficult to recognize in smaller specimens. The distal parts of 

the male gonopods are also slightly but distinctly different (unpublished data). Given that all 4 species of 

Scylla are marketed throughout their range by the extensive export market, the existing catch figures and 

fishery management practices currently applied to a single species (S. serrata) obviously have to be 

revised. 

The very large species depicted above (often called the “Sri Lanka crab” in South, Southeast and East 

Asia) is the “real” Scylla serrata and has a wide distribution, with preference for more saline waters. It varies 

from green to almost black, has a smooth carapace with distinct transverse ridges, deep H-shaped gastric 

groove, relatively broad frontal lobes, all of which are more or less in line with each other, broad anterolateral 

teeth which project obliquely outwards, and has very well-developed spines on the outer surface of the 

chelipedal carpus and palm. 

Two of the species are smaller and more closely associated with mangroves than the real S. serrata, and 

occur more or less in the same region, but generally prefer less saline conditions and are more common 

in continental shelf waters. S. serrata is not known to occur inside the Sunda Shelf, but it is the only species 

of Scylla known from the Red Sea. Scylla olivacea (Herbst, 1796) is usually brownish to brownish green 

in colour (sometimes orangish). It has a smoother, more evenly convex carapace with very low transverse 

ridges, a shallow H-shaped gastric groove, the median pair of the frontal lobes more rounded and projecting 

slightly forwards of the lateral ones, the anterolateral teeth gently curving anteriorly, giving the carapace a 

less transverse appearance. It also has very low spines on both the outer surface of the chelipedal carpus 

and the dorsal surface of palm. (See abbreviated species account below). The second mangrove species, 

Scylla tranquebarica (Fabricius, 1798) (= Lupa lobifrons H. Milne Edwards, 1834) varies from brown to 

almost black in coloration, and has very well-developed spines on the outer surfaces of the chelipedal 

carpus and the palm (as seen in S. serrata). It differs from S. serrata, however, by having the frontal teeth 

more acutely triangular, the median pair projecting slightly forwards of the lateral pair, and the anterolateral 

teeth gently curving anteriorly, giving the carapace a less transverse appearance. (See abbreviated species 

account below). 

The fourth Scylla species, Scylla paramamosain Estampador, 1949, seems to prefer areas which are more 

rocky or near reefs, although it is also known from estuarine ponds and mangrove forests. It seems to be 

intermediate between S. serrata and S. olivacea both in morphology and coloration but can usually be 

distinguished by the form of its frontal margin and cheliped armature (see key). (See abbreviated species 

account below). 

Scylla olivacea (Herbst, 1796) 

En - Orange mud crab. 

Maximum carapace width 18 cm (males). Carapace brownish to brownish green in colour (sometimes 

orangish), palm orange to yellow. Inhabits mangroves. Collected in large numbers and 

probably the most common species of Scylla to be found in many markets in Sundaic Southeast 

Asia and Thailand. Known so far from the continental waters of the Sunda Shelf and various parts 

of the East Pacific. (See species account of S. serrata for further information).

1128 Crabs 

Scylla paramamosain Estampador, 1949 

En - Green mud crab. 

Maximum carapace width 20 cm (males), commonly between 15 and 18 cm; weight up to 2 kg. 

Carapace usually green to light green, palm green to greenish blue with lower surface and base of 

fingers usually pale yellow to yellowish orange. Rock areas, near reef, and mangroves. Common in 

northern parts of South China Sea and parts of Java, but less so elsewhere. Shelf species. (See 

species account of S. serrata for further information). 

Scylla tranquebarica (Fabricius, 1798) 

En - Purple mud crab. 

Maximum carapace width 20 cm (males); weight up to 2 kg. Carapace green to almost black, palm 

purple. Mainly in mangroves (down to sublittoral parts) and collected in large numbers. This is 

probably the second most common species seen in Sundaic Southeast Asian markets, but less 

common in Thailand and the Philippines. Known from various parts of the Indo-West Pacific, 

including shelf waters. (See species account of S. serrata for further information).


Thalamita crenata (Latreille, 1829) 


FAO name: En - Crenate swimming crab. 

Diagnostic characters: Surface of carapace 

smooth, ridges low but distinct; front with 6 

equal-sized, rounded lobes. Colour: dark to 

olive green overall. 


Habitat, biology, and fisheries: One of the 

most distinctive species of the genus, and 

one of the few found in shallow non-reef habitats 

with soft substrates. Prefers areas near 

mangroves or with muddy-rocky substrates. 

Frequently collected by traps, trawlers, 

seines, and nets. Fished mainly in Southeast 

and East Asian countries. Although it can 

be very common in some areas, T. crenata 

has a low value in markets as it grows 

smaller than other, more commercial portunids 

found in the area. 

Distribution: China, Indonesia, Malaysia, 

Singapore, Australia, Tuamotu, Tonga, 


Thalamita spinimana (Dana, 1852) 


FAO name: En - Spiny claw swimming crab. 

Diagnostic characters: Surface of carapace 

smooth, sometimes with low pubescence; ridges 

distinct; front with 6 lobes, median 4 lobes truncate, 

lateral 2 lobes rounded. Colour: usually bright red 

overall, but sometimes green, or with a mixture of 

red and green. 


Habitat, biology, and fisheries: Occasionally collected 

for food throughout its range, caught by traps 

and nets. It is sold for comparatively low prices in 

markets. 

Distribution: West Pacific. 

Remarks: A large number of coral reef 

species of Thalamita are found in the area. 

Most of these, however, are of small size 

and have no economic value. T. spinimana 

is one of the more common larger species 

in the genus, easily distinguished by its 

spinose palm and the bright red coloration. 

(from A. Milne Edwards, 1873)

1130 Crabs 

Charybdis affinis Dana, 1852 

En - Smoothshelled swimming crab. 

Maximum carapace width 6 cm. Prefers sandy to muddy substrates in subtidal waters. Fished 

sporadically and occasionally seen in local markets where it is sold for low prices, due to its small 

size. China and Japan to various parts of India and Southeast Asia. 

(from Leene, 1930) 

Charybdis anisodon (De Haan, 1850) 

En - Twospined arm swimming crab. 

Maximum carapace width 8 cm. Prefers muddy substrates at depths to 15 m. Occasionally collected 

by trawls and sold in local markets for its moderately large size. Indo-West Pacific in distribution, 

reaching Hawaii. 

(from Leene, 1930)


Charybdis annulata (Fabricius, 1798) 

En - Banded-legged swimming crab. 

Maximum carapace width 7 cm. Shows distinctive broad blue and white bands on the legs when 

alive. Prefers rocky areas and reefs, from the intertidal zone to a depth of about 20 m. Occasionally 

collected for food, but never abundant enough to be commercially important. Indo-West Pacific, from 

South Africa to Southeast Asia, Japan, and Tahiti. 


Charybdis truncata (Fabricius, 1798) 

En - Blunt-toothed crab. 

Maximum carapace width 5 cm. Prefers muddy bottoms at depths from 10 to 100 m. Locally 

abundant in some areas and obtained by trawls. Not commonly sold in markets. Indo-West Pacific, 

reaching Japan and Australia. 




1132 Crabs 

Geryonidae GERYONIDAE 

Diagnostic characters: Carapace hexagonal; 

dorsal surface relatively smooth to 

granular; frontal margin with 4 teeth; anterolateral 

margins distinctly convex, each with 3 to 5 

low, sometimes indistinct teeth. Dactylus of 

walking legs T-shaped in cross-section. Male 

abdominal segments 3 to 5 fused, functionally 

immovable, but sutures still visible. 

Habitat, biology, and fisheries: These are 

deep-sea crabs, normally occurring in depths 

below 100 m. Taken incidentally by trawls and 

traps. Crabs of the genus Chaceon are represented 

by numerous species of interest to fisheries. 

Although not all these species are 

harvested in large numbers, the fishery potential 

of geryons is quite great. The most widely 

exploited species is the Atlantic C. maritae.Five 

species of Chaceon are known from the West- 

Geryons 

ern Central Pacific so far. The more important species in the area are C. granulatus and C. bicolor. Further 

new species can be expected when more deep-water areas are sampled. 

Remarks: The golden crabs or geryons (genus Chaceon) are a very distinctive taxonomic group. However, 

the composition of the family is still not settled and some genera which have been assigned to the 

Geryonidae should probably be transferred to the Goneplacidae instead. The known species of Chaceon 

can easily be separated into 2 groups: among the species occurring in the area, C. bicolor, C. australis, 

and C. poupini belong to the group in which the dactylus of walking legs is not laterally flattened, and the 

height at midlength is greater than, or subequal to, the width at midlength. 1/ Only 2 species in the area, 

namely C. granulatus and C. karubar, belong to the other group, in which the dactylus of legs is laterally 

flattened, and the height at midlength is less than the width at midlength. 


Members of the genus Chaceon can only be confused with some members of the Goneplacidae which also 

have a squarish carapace (the only commercial species of Goneplacidae in the area, Carcinoplax 

longimanus, has an ovoid carapace). In addition, the large size of geryons (usually exceeding 14 cm 

carapace width), the relatively long legs, the T-shaped cross-section of the dactylus of walking legs, and 

their occurrence in deep waters (deeper than 200 m), easily separates them from the goneplacids (no 

geryons are known from shallow waters). 

Key to species of Chaceon occurring in the area 

1a. Dorsal surface of carapace smooth to gently rugose, not granulose; branchial and 

posterolateral regions not swollen; dorsoventrally flattened, height at midlength 0.8 

times or less than width at midlength . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Dorsal surface of carapace granulose; 

branchial and posterolateral regions 

raised; dactylus of legs laterally flattened, 

height at midlength subequal to or 

merus of legs 

without spine 

greater than width at midlength (0.9 

times and more) . . . . . . . . . . . . . . . . . . → 4 

2a. Merus of walking leg unarmed, without 

distinct dorsal distal spine or tooth (Fig. 1) 

. . . . . . . . . . . . . . . . . . . Chaceon poupini 

2b. Merus of walking leg with dorsal distal 

spine or tooth . . . . . . . . . . . . . . . . . . . → 3 

male abdominal segments 3-5 

immovable but sutures visible 

Fig. 1 Chaceon poupini 

carapace 

hexagonal 

dactylus T-shaped 

in cross-section 

1/ Three other species belonging to the same group have been reported from other places in the Pacific: Chaceon 

yaldwyni Manning, Dawson, and Webber, 1989 (New Zealand), C. imperialis Manning, 1992 (Emperor Seamount 

Chain), and C. manningi Ng, Lee, and Yu, 1994 (Tung-Sa Islands, South China Sea).

Geryonidae 1133 

3a. Anterolateral margin of adults with spiniform teeth (Fig. 2). . . . . . . . . . . . . Chaceon australis 

3b. Anterolateral margins of adults with low, lobiform teeth (Fig. 3) . . . . . . . . . . . Chaceon bicolor 

spiniform 

teeth 

4a. Outer surface of chelipedal carpus with spine or projection on outer surface; merus of 

legs with distinct dorsal distal spine or tooth (Fig. 4) . . . . . . . . . . . . . . . . Chaceon karubar 

4b. Outer surface of chelipedal carpus unarmed; merus of legs unarmed, without distinct 

dorsal distal spine or tooth (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . Chaceon granulatus 

carpus of 

cheliped 

with spine 



Chaceon australis Manning, 1993 

Chaceon bicolor Manning and Holthuis, 1989 

Chaceon granulatus (Sakai, 1978) 

Chaceon karubar Manning, 1993 

Chaceon poupini Manning, 1992 

 

Fig. 2 Chaceon australis Fig. 3 Chaceon bicolor 

Fig. 4 Chaceon karubar 

Fig. 5 Chaceon granulatus 

References 

Manning, R.B. and L.B. Holthuis. 1981. West African Brachyuran Crabs (Crustacea: Decapoda). Smithson. Contrib. 

Zool., 306:1-379. 

Manning, R.B. and L.B. Holthuis. 1989. Two new genera and nine new species of geryonid crabs (Crustacea: Decapoda: 

Geryonidae). Proc. Biol. Soc. Wash., 102:50-77. 

Ng, P.K.L. and R.B. Manning. 1998. A new deepwater crab from the Palau Islands, Micronesia (Decapoda: Brachyura: 

Geryonidae). Proc. Biol. Soc. Wash., 111:in press.

1134 Crabs 

Chaceon bicolor Manning and Holthuis, 1989 


FAO name: En - Pacific golden crab. 

Diagnostic characters: Carapace hexagonal; dorsal 

surface not inflated; anterolateral teeth low. Merus of legs 

long, slender. Dactylus of legs not laterally flattened, 

height at midlength greater than or subequal to width at 

midlength.Colour: reddish tan to purplish black (anterior 

half of carapace may be differently coloured than posterior 

half), sometimes cream-white throughout. 

Size: Maximum carapace width 18 cm (males) and 

15 cm (females). 

Habitat, biology, and fisheries: Like most 

of the known species of the genus, 

Chaceon bicolor lives on muddy substrates 

in deeper waters at depths between 

200 and 1 600 m, and apparently burrows 

sometimes into the sediment. Occasionally 

caught throughout its range, especially in 

Australia. Taken in bottom trawls and lobster 

pots, and is believed to be commercially 

valuable in the future, being 

supposedly quite abundant in some areas. 

Distribution: New Caledonia, northwest 

and eastern Australia, and Loyalty Islands. 

Chaceon granulatus (Sakai, 1978) 

Frequent synonyms / misidentifications: None / Chaceon affinis (A. Milne Edwards and Bouvier, 1894). 

FAO name: En - Japanese golden crab. 

Diagnostic characters: Carapace hexagonal; dorsal 

surface (especially branchial regions) distinctly inflated; 

anterolateral teeth low in adults. Merus of legs 

long, slender, with dorsal subdistal tooth; dactylus of 

legs laterally flattened, height at midlength less than 

width at midlength. Colour: uniform tan to cream 

colour overall. 


Habitat, biology, and fisheries: On soft substrates 

in deep waters from 300 to 1 500 m. Caught on a 

regular basis in Japan, but never in large numbers. 

When it does appear in markets, it commands high 

prices (up to US$60 to US$80 for a fresh crab of 

20 cm width). Also caught for sale in Palau. 

Distribution: Japan, China, Taiwan Province 

of China, and Palau. 

Remarks: Only 3 Pacific species of 

Chaceon, C. granulatus, C. karubar,andC. 

manningi belong to the group in which the 

dactylus of legs is laterally flattened, and the 

height at midlength is less than the width at 

midlength. Specimens from Saipan and 

Palau which have been identified with C. 

granulatus possibly belong to separate species, 

but studies on these are still ongoing. 

For the moment, crabs from these 2 islands are tentatively identified as C. granulatus. The Palau specimens 

were recently recognized as belonging to a new species.

Geryonidae 1135 

Chaceon australis Manning, 1993 

En - Austral golden crab. 

Maximum carapace width 10 cm. Carapace pink and tan in life. Occurs in deeper waters at depths 

of about 900 m. Not collected commercially for food so far, but has good fishery potential with the 

future development of deep-water fisheries, due to its large size. Known only from the Austral 

Islands. 

Chaceon karubar Manning, 1993 

En - Indonesian golden crab. 

Maximum carapace width 19 cm. In deeper waters at depths of about 550 m. Not collected 

commercially for food so far, but has a good fishery potential because of its large size. Known only 

from the Tanimbar Islands in Indonesia. 

Chaceon poupini Manning, 1992 

En - Polynesian golden crab. 

Maximum carapace width 14 cm. Colour uniform red in life. Occurs in deeper waters at depths from 

600 to 1 000 m. Not collected commercially for food so far, but has a good fishery potential because 

of its large size. Known only from the Marquesas Islands.

1136 Crabs 

Majidae MAJIDAE 

Spider crabs 

Diagnostic characters: Carapace pyriform 

(pear-shaped), circular to subovate, 

anterior 1/2 to 1/3 usually distinctly 

narrower than posterior part; dorsal surface 

gently convex, spinulose, granulose, and/or 

ridged; front narrow, often with 2 long horn-like 

projections (rostra); orbits poorly developed 

to absent; anterolateral margins of carapace 

often armed with well-developed spines. Legs 

spinulose and/or granulose, often with stiff setae. 

All male abdominal segments usually 

freely movable in most species. 

Habitat, biology, and fisheries: 1/ orbits incomplete 

Benthic 

crabs. Most species of minor or no importance 

to fisheries, with only the larger species of 

Schizophrys being occasionally collected in the 

Western Central Pacific. 


None. The generally pyriform (pear-like) shape of many majids easily distinguishes them from other families. 

Their orbits are usually poorly demarcated to incomplete and this feature is often quite diagnostic. In 

addition, many species have hook-like setae on their bodies and appendages with which they use to attach 

various materials for camouflage. 

The Hymenosomatidae (crown crabs, non-commercial) closely resemble many majids, but are easily 

distinguished by their very small size (small species at 2 mm adult carapace width, being contenders with 

the pinnotherids for the smallest crabs in the world), the absence of hook-like setae, and having only 5 

abdominal segments (excluding the telson). 


1a. Two accessory spines at base of 

each rostral horn (Fig. 1a) . . . 

. . . . . . . . . . . . . . Schizophrys dama 

1b. One accessory spine at base of 

each rostral horn (Fig. 1b) . . . 

. . . . . . . . . . . . . . Schizophrys aspera 



Schizophrys aspera (H. Milne Edwards, 1834) 

Schizophrys dama (Herbst, 1804) 

 

horn-like projections 

1 assessory 

spine 

2 assessory 

spines 


carapace 

a) Schizophrys dama b) Schizophrys aspera 

Fig. 1 rostral horn (dorsal view) 

References 

Griffin, D.J.G. 1966. A review of the Australian majid spider crabs (Crustacea, Brachyura). Australian Zoologist, 

13:259-298. 

Griffin, D.J.G. and H.A. Tranter. 1986. The Decapoda Brachyura of the Siboga Expedition. VII. Majidae. Siboga Exped. 

Monogr., 39c(4):1-335. 

1/ The most important majids in fisheries are the large crabs of the genus Chionoecetes which occur in cold northern 

waters only. Near the area, the “Japanese giant spider crab” (Macrocheira kaempferi), known only from Japan and 

Taiwan Province of China, is occasionally collected for food. This is the largest crab in the world, reaching a carapace 

width of 30 cm and with legs spanning 2.5 m from tip to tip. The only other majid crab of fishery importance near the 

area is the southern spider crab (Jacquinotia edwardsii) from New Zealand, with several tonnes a month being landed.

Majidae 1137 

Schizophrys aspera (H. Milne Edwards, 1834) 


FAO name: En - Common decorator crab. 

Diagnostic characters: Carapace pear-shaped, 

with 2 pronounced rostral horns. Colour: brown 

overall. 


Habitat, biology, and fisheries: Found on rocks, 

especially near reefs, from intertidal areas to a 

depth of 40 m. Usually caught incidentally by 

hand, sometimes in crab pots. Only of local commercial 

importance. 

Distribution: Indo-West Pacific, eastwards to 

Hawaii. 

Remarks: Schizophrys aspera can be confused 

with S. dama, which is easily distinguished by the 

2 accessory spines near the base of each rostral 

horns (only 1 spine in S. aspera). 

Schizophrys dama (Herbst, 1804) 


En - Pronghorn decorator crab. 

Maximum carapace width 6 cm. A subtidal coral reef species, occasionally invades rocky and sandy 

areas. Collected incidentally by hand or with nets, but nowhere commercially important. Southeast 

Asia to New Guinea and Australia. 

(from Alcock and Anderson, 1899)

1138 Crabs 

Grapsidae GRAPSIDAE 

Sally-light-foots, vinegar crabs, and paddler crabs 

Diagnostic characters: Carapace squarish, transversely rectangular, trapezoidal, or circular; 

dorsal surface flat to gently convex, with low oblique or transverse ridges; front much broader than 

orbits; orbits occupying almost entire anterior border (excluding front); antero- and posterolateral 

margins of carapace usually not clearly demarcated, lateral margins appearing almost straight or gently 

convex, usually armed with 1 or 2 teeth anteriorly, sometimes unarmed. Rhomboidal gap usually present 

between third maxillipeds, often with mandibles exposed. Dactylus of legs with distinct spines. Male 

abdominal segments 3 to 5 freely movable in most species. 

carapace squarish 

Grapsinae 

carapace 

circular 

Habitat, biology, and fisheries: Swimming, climbing, or terrestrial crabs, with the majority of species 

occurring in intertidal areas or semiterrestrial habitats. Many also occur in estuarine waters and a few 

species live exclusively in fresh water. Most species are of minor commercial importance, with species of 

the genera Episesarma and Varuna being most often collected for food. 1/ 



Ocypodidae: with a much narrower frontal margin 

of the carapace; always lack a rhomboidal gap 

between their third maxillipeds; crabs walk on the 

tip of dactyli of legs, not on the sides of dactyli (as 

seen in grapsids). 

Key to the subfamilies of Grapsidae 

1a. No distinct rhomboidal gap between 

third maxillipeds 

(Fig. 1a) . . . . . . . . . . . . . . . . . → 2 

1b. Distinct rhomboidal gap between 

third maxillipeds 

(Fig. 1b) . . . . . . . . . . . . . . . . . → 3 

no distinct 

rhomboidal 

gap, mandibles 

not visible 

Sesarminae 

Ocypodidae 

rhomboidal 

gap with 

mandibles 

exposed 

a) Varuninae, Plagusiinae b) Sesarminae, Grapsinae 


long orbits 

1/ The most important grapsids commercially are the “mitten crabs” of the genus Eriocheir (Varuninae) from China, 

Taiwan Province of China, Japan, and Korea (see Guo et al., 1997). Costing up to US$ 20 each, these relatively large 

grapsids are netted in enormous numbers when they migrate from fresh waters to the sea to spawn. They are much 

sought after for their ripe ovaries.

Grapsidae 1139 

2a. Carapace squarish; frontal margin entire, without lobes or teeth; third maxilliped with 

broad exopod and long flagellum (Fig. 2a); male abdomen with all segments movable 

(Fig. 3a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Varuninae 

2b. Carapace circular; third maxilliped with slender exopod and no flagellum (Fig. 2b); male 

abdomen with segments 3 to 6 immovable, with suture between segments 3 and 4 still 

evident (Fig. 3b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plagusiinae 

(a single species of interest to fisheries, Plagusia tuberculata, occurring in the area) 

broad 

exopod 

with 

flagellum 

a) Varuninae b) Plagusiinae 


narrow 

exopod 

without 

flagellum 

3a. Merus and ischium of third maxillipeds with ridge (Fig. 4a); pterygostomial region with 

network-like (reticulated) pattern of very short, stiff setae (Fig. 5a); carapace usually 

squarish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sesarminae 

3b. Merus and ischium of third maxillipeds without hairy oblique ridge (Fig. 4b); pterygostomial 

region may be setose but no network-like pattern discernible (Fig. 5b); carapace 

usually circular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grapsinae 

ridges on 

merus and 

ischium 

no clear 

ridges 

a) Sesarminae b) Grapsinae 


Key to food species of Grapsinae occurring in the area 

1a. Front gently deflexed, margin weakly serrated 

(Fig. 6a); inner angle of carpus of cheliped 

with acutely triangular spine (Fig. 7a); pectinated 

fingertips of chela longitudinally broad 

(Fig. 8a); adult male sixth abdominal segment 

shorter than fifth segment (Fig. 9a) . . . . . 

. . . . . . . . . . . . . . . . . . . .Grapsus albolineatus 

1b. Front strongly deflexed and appears almost 

vertical from frontal view, margin serrated 

(Fig. 6b); inner angle of carpus of cheliped 

with subtruncate base (Fig. 7b); pectinated 

fingertips of chela longitudinally narrow 

(Fig. 8b); adult male sixth abdominal segment 

distinctly longer than or subequal in length to 

fifth segment (Fig. 9b) . . . . . . . Grapsus tenuicrustatus 

6 

5 

4 

3 

segments 3-6 

movable 

6 

5 

4 

3 

a) Varuninae b) Plagusiinae 


dense 

network of 

hairs 

a) Sesarminae b) Grapsinae 

Fig. 5 body in frontal view 

weakly serrated 

a) Grapsus albolineatus 

distinctly serrated 

b) Grapsus tenuicrustatus 

segments 

3-6 

immovable 

scattered 

setae 

Fig. 6 frontal margin of carapace

1140 Crabs 


triangular 

spine 

subtruncate 

base 


Fig. 7 carpus of cheliped 



Fig. 8 chela 

broad 

pectinated tip 

narrow 

pectinated tip 


6 th 

segment 

shorter 

than 5 th 

6 th 

segment 

longer 

than 5 th 


Key to food species of Sesarminae occurring in the area 

1a. Tubercles on dorsal margin of dactylus of chela numbering 64 to 76 (Fig. 10a); male 

first gonopod with broad, truncate pectinated tip (Fig. 11a); most of outer surface of palm 

and proximal part of dactylus purple to reddish, dactylus white distally, pollex orangish 

red to pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Episesarma chengtongense 

1b. Tubercles on dorsal margin of dactylus of chela numbering less than 61; male first 

gonopod tip not as above; outer surface of palm not coloured as above . . . . . . . . . . . . . → 2 

2a. Tubercles on dorsal margin of dactylus of chela similarly sized throughout length, 

numbering 40 to 50 (Fig. 10b); male first gonopod with broad, rounded pectinated tip 

(Fig. 11b); outer surface of palm light brown with white fingertips . . . . . Episesarma palawanense 

2b. Tubercles on dorsal margin of dactylus of chela increasing in size towards finger tip; 

male first gonopod not as above; outer surface of palm light brown with white fingertips 

or coloured in other ways . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Outer surface of palm violet at the proximal part, distal part and fingers white; tubercles 

on dorsal margin of dactylus of chela 40 to 48 (Fig. 10c); male first gonopod with 

relatively narrow truncate pectinated tip (Fig. 11c) . . . . . . . . . . . . . . Episesarma versicolor 

3b. Outer surface of palm uniformly red or brown; tubercles on dorsal margin of dactylus of 

chela 35 to 60; male first gonopod tip not as above . . . . . . . . . . . . . . . . . . . . . . . . → 4 

4a. Tubercles on dorsal margin of dactylus of chela numbering 35 to 45 (Fig. 10d); male 

first gonopod with relatively broad, bent pectinated tip (Fig. 11d); outer surface of palm 

uniformly red to chestnut brown . . . . . . . . . . . . . . . . . . . . . . Episesarma singaporense 

4b. Tubercles on dorsal margin of dactylus of chela numbering 40 to 60 (Fig. 10e); male 

first gonopod with relatively narrow, bent pectinated tip (Fig. 11e); outer surface of palm 

light brown with white fingertips . . . . . . . . . . . . . . . . . . . . . . . . . . Episesarma mederi 

64-76 tubercles 40-50 tubercles 40-48 tubercles 35-45 tubercles 

a) Episesarma 

chengtongense 

b) Episesarma 

palawanense 

c) Episesarma 

versicolor 

Fig. 10 dactylus of chela (lateral view) 

d) Episesarma 

singaporense 

6 

5 

6 

5 


40-60 tubercles 

e) Episesarma 

mederi


Key to food species of Varuninae occurring in the area 

1a. Distal process of male first gonopod 

with 1 process distinctly longer than 

the other (Fig. 12a); occurs mainly in 

continental shelf waters . . . . . . . . Varuna yui 

1b. Distal process of male first gonopod 

with both processes subequal in size 

(Fig. 12b); occurs mainly in areas 

with more oceanic waters . . . . Varuna litterata 



Subfamily GRAPSINAE 

Grapsus albolineatus Lamarck, 1818 

Grapsus tenuicrustatus (Herbst, 1783) 

 

Subfamily PLAGUSIINAE 

Plagusia tuberculata Lamarck, 1818 

Subfamily SESARMINAE 

Episesarma mederi (A. Milne Edwards, 1854) 

Episesarma chengtongense (Serène and Soh, 1967) 

Episesarma palawanense (Rathbun, 1914) 

Episesarma singaporense (Tweedie, 1936) 

Episesarma versicolor (Tweedie, 1940) 

 

Subfamily VARUNINAE 

Varuna litterata (Fabricius, 1798) 

Varuna yui Hwang and Takeda, 1984 

 

a) Episesarma 

chengtongense 

broad 

truncate 

tip 

b) Episesarma 

palawanense 

broad 

rounded 

tip 

c) Episesarma 

versicolor 

relatively 

narrow 

truncate tip 


d) Episesarma 

singaporense 

relatively 

broad, 

bent tip 

unequal processes 

e) Episesarma 

mederi 

narrow, 

bent 

tip 

less unequal processe 

a) Varuna yui b) Varuna litterata 


References 

Alcock, A. 1900. Materials for a carcinological fauna of India. No. 6. The Brachyura Catometopa or Grapsoidea. J. Asiat. 

Soc. Bengal, 69, pt. 2(3):279-486. 

Crosnier, A. 1965. Crustacés Décapodes. Grapsidae et Ocypodidae. Faune de Madagascar, 18:1-143.

1142 Crabs 

Grapsus albolineatus Lamarck, 1818 

Frequent synonyms / misidentifications: Grapsus strigosus (Herbst, 1799) / None. 

FAO name: En - Mottled Sally-light-foot. 

Diagnostic characters: Carapace rounded; 

front straight, entire; anterolateral margins 

rounded, each with 1 tooth; lateral regions 

with numerous oblique striae. Fingertips 

strongly spooned. Colour: carapace with 

green and white transverse markings. 


Habitat, biology, and fisheries: Found out of 

the water, on rocks in the splash zone. Collected 

only incidentally and consumed by local 

residents only. Usually caught by hand or 

with special nets. 


Remarks: Several other Grapsus species 

live in the region, but the most common are 

G. albolineatus and G. tenuicrustatus (see 

below). These 2 species, however, can 

easily be separated (see key). 

Grapsus tenuicrustatus (Herbst, 1783) 


None / None. 

FAO name: En - Natal Sally-light-foot. 

Diagnostic characters: Carapace rounded; 

front straight, finely serrated; anterolateral 

margins rounded, each with 1 tooth; lateral 

regions with numerous oblique striae. Fingertips 

slightly spooned. Colour: carapace with 

green and white transverse markings. 


Habitat, biology, and fisheries: Found out 

of the water, on rocks in the splash zone. 

Collected only incidentally and consumed by 

local residents. Usually caught by hand or 

with special nets. Probably the most common 

representative of Grapsus among 

several species occurring in the area. 

Distribution: Indo-West Pacific, including 

French Polynesia and Hawaii.


Episesarma versicolor (Tweedie, 1940) 


FAO name: En - Violet vinegar crab. 


squarish, with 1 small anterolateral 

tooth; dorsal surface relatively flat; 

regions well defined, covered with 

short, stiff setae. Dorsal margin of 

dactylar finger with numerous (40 to 

48) tubercles, forming a stridulatory 

organ. Colour: carapace brown to 

brownish grey; outer surface of palm 

with proximal parts violet, distal parts 

and fingers white. 


5cm(forEpisesarma spp.). 

Habitat, biology, and fisheries: All 

members of Episesarma are mangrove 

crabs, digging burrows at the base of trees or at mud lobster 

(Thalassina) mounds, and are predominantly herbivorous. There 

are several species of Episesarma known from Southeast Asia, 

all of which are harvested to varying degrees. They are collected 

in large numbers for food in many Southeast Asian countries and 

some southern Chinese communities. The crabs are collected by 

hand, often pickled in vinegar and/or salt solutions and are then 

eaten as they are with rice, or deep fried. In northern Australia, an 

undescribed species of Episesarma is eaten by the aborigines (P. 

Davie, pers. comm.). 

Distribution: Southeast Asia and southern China. 

Remarks: Four other species 

of Episesarma in Southeast 

Asia are large and common 

enough to be collected for food. 

These are E. singaporense, E. 

mederi, E. chengtongense,and 

E. palawanense (see abbreviated 

species accounts below). 

They are easily distinguished 

by various carapace and cheliped 

characters, and on the basis 

of live colours. 

outer surface of chela

1144 Crabs 

Varuna litterata (Fabricius, 1798) 


FAO name: En - Oceanic paddler crab. 

Diagnostic characters: Carapace squarish, 

surface smooth; front straight; anterolateral 

margins each with 3 very broad, low but sharp 

teeth. Dactylus, propodus, and carpus of legs 

laterally flattened, fringed with long, closely 

packed setae. Colour: light brown to brownish 

grey on dorsal surfaces. 


Habitat, biology, and fisheries: Varuna litterata 

prefers areas faced by more oceanic 

waters whereas V. yui is only known from the 

continental shelf waters of Southeast Asia and 

neighbouring areas. Both species occur together in the Philippines. V. litterata and V. yui are generally 

estuarine crabs that usually prefer slow-moving or almost stagnant bodies of water. Both species, however, 

can be found up to 20 km from the sea in completely fresh water. Large or ovigerous specimens can be 

found in intertidal areas, frequently associated with floating clumps of brown algae, Sargassum spp. 

Collected in good numbers in most Southeast Asian countries for food. Usually collected by hand, but also 

caught by traps, seines, and fish corrals. Like Episesarma, the species of Varuna in the area are usually 

pickled in vinegar and/or salt solutions. They are then eaten as such or deeply fried. They are also collected 

for their tasty ovaries and are 

especially common in markets 

during their breeding period. 


in the Indo-West Pacific. 

Remarks: The 2 species of 

Varuna in the area, V. litterata 

and V. yui (see abbreviated 

species account below), are 

very similar externally, and 

can only be distinguished effectively 

by means of their 

male gonopods. 

Varuna yui Hwang and Takeda, 1984 

En - Sundaic paddler crab. 

Maximum carapace width 5 cm. In estuarine waters up to 20 km inland in completely fresh water. 

Collected in large numbers in most Southeast Asian countries for local consumption. Usually 

collected by hand, but also caught by traps, seines, and fish corrals. In continental shelf waters of 

the Sunda Shelf up to southern China and Philippines, occurring together with Varuna litterata in 

the latter 2 localities. V. yui is the dominant species of the genus in the Sunda Shelf.


Episesarma chengtongense (Serène and Soh, 1967) 

En - Pinkfingered vinegar crab. 

Maximum carapace width 5 cm. Found only in mangroves. Collected by hand for food, and usually 

sold fresh or pickled in vinegar. Occurs in southern China and various parts of Southeast Asia. 

Episesarma mederi (A. Milne Edwards, 1854) 

En - Thai vinegar crab. 

Maximum carapace width 4 cm. Inhabits mangroves and forested muddy habitats. Collected by hand 

in large numbers for food, and usually sold fresh or pickled in vinegar. Occurs in southern China 

and various parts of Southeast Asia. 

Episesarma palawanense (Rathbun, 1914) 

outer surface of chela 

En - Rathbun’s vinegar crab. 

Maximum carapace width 4 cm. Inhabits mangroves. Collected by hand for food, and usually sold 

fresh. Occurs in various parts of Southeast Asia.

1146 Crabs 

Episesarma singaporense (Tweedie, 1936) 

En - Singapore vinegar crab. 

Maximum carapace width 4 cm. Inhabits mangroves. Collected by hand in good numbers for food, 

and is usually sold fresh in markets or pickled in vinegar. Occurs in parts of Southeast Asia. 

Plagusia tuberculata Lamarck, 1818 

outer surface of chela 

En - Tuberculated Sally-light-foot. 

Maximum carapace width about 4 cm. On rocks just above the splash zone. Collected by hand and 

traps for food in rural communities, and usually pickled in vinegar or sauce as a food supplement. 

Indo Pacific in distribution, reaching New Zealand, Hawaii, French Polynesia, and California in the 

eastern Pacific. 

(after Tung et al., 1986) 



Gecarcinidae 1147 

Gecarcinidae GECARCINIDAE 

Land crabs 

Diagnostic characters: Carapace circular to 

transversely ovate; dorsal surface 

smooth, strongly convex longitudinally and 

transversely; frontal margin entire; anterolateral 

margins unarmed or each with a single tooth. 

Rhomboidal gap present between third maxillipeds. 

Legs stout, dactylus longitudinally ridged, 

often with dense, stiff setae, margins with spines. 

All male abdominal segments distinct, movable. 

Habitat, biology, and fisheries: Terrestrial crabs. 

Although gercarcinids can be found many kilometres 

inland, they must return to the sea to spawn 

and release their planktonic larvae. The most frequently 

collected food species in the area is Cardisoma 

carnifex. 


The swollen carapace and terrestrial habitats of the gecarcinids make them a very distinctive group. In this 

respect they may be confused with several fresh-water and terrestrial genera of Gecarcinucidae, Potamidae, 

and Parathelphusidae (species of the latter 2 are collected for food in many parts of Southeast Asia 

and Indo-China). 

Potamidae, Gecarcinucidae, Parathelphusidae (= Sundathelphusidae): several genera of these exclusive 

fresh-water families may resemble gecarcinids, but are readily distinguished by the lack of a rhomboidal 

gap between the third maxillipeds. 

Grapsidae: also have a rhomboidal gap between the third maxillipeds, but most species possess a much 

flatter dorsal carapace surface compared to gecarcinids. 

Ocypodidae: also have many terrestrial members, but lack a rhomboidal gap between the third maxillipeds 

and generally do not have as thick a shell as seen in gecarcinids. 


long orbits 

Grapsidae carapace generally 

flatter 

Ocypodidae 


1a. Exopod of third maxilliped without flagellum (Fig. 1a); orbits oblique, eyes small, never 

reaching first anterolateral tooth (Fig. 2) . . . . . . . . . . . . . . . . . . . . . Gecarcoidea lalandii 

1b. Exopod of third maxilliped with long flagellum (Fig. 1b); orbits horizontal, eyes well 

developed, reaching to just before first anterolateral tooth . . . . . . . . . . . . . . . . . . . . .→ 2 

exopod 

short, no 

flagellum 

a) Gecarcoidea lalandii 

b) Cardisoma spp. 


exopod 

with long 

flagellum 

eyes 

small 

Fig. 2 Gecarcoidea lalandii

1148 Crabs 

2a. Legs very long (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cardisoma longipes 

2b. Legs of normal length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Surface of carapace with numerous scattered flattened to rounded granules (Fig. 4) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cardisoma rotundum 

3b. Surface of carapace smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

4a. Carapace transversely ovate; face (pterygostomial and sub-branchial regions) with 

large setose area which covers anterior part of the branchiostegal region; stiff setae 

present on merus of legs; males with 2 equally enlarged chelae; carapace bluish brown 

to blue coloration in life (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . Cardisoma hirtipes 

4b. Carapace circular; face (pterygostomial and sub-branchial regions) with small setose 

area which does not reach branchiostegal region; short (or no) setae on merus of legs; 

males with 1 cheliped several times size of other; carapace brown in life (Fig. 6) . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cardisoma carnifex 



Cardisoma carnifex (Herbst, 1794) 

Cardisoma hirtipes Dana, 1852 

Cardisoma longipes (A. Milne Edwards, 1873) 

Cardisoma rotundum (Quoy and Gaimard, 1824) 

Gecarcoidea lalandii (H. Milne Edwards, 1837) 

 

Fig. 3 Cardisoma longipes 


flattened to 

rounded 

granules 

Fig. 4 Cardisoma rotundum 

Fig. 5 Cardisoma hirtipes Fig. 6 Cardisoma carnifex 

References 

Burggren, W.W. and B.R. McMahon (eds). 1988. The biology of land crabs. Cambridge University Press, 479 p. 

Türkay, M. 1974. Die Gecarcinidae Asiens und Ozeaniens (Crustacea: Decapoda). Senckenb. Biol., 55:223-259.


Cardisoma carnifex (Herbst, 1794) 

Frequent synonyms / misidentifications: Cardisoma urvillei H. Milne Edwards, 1853; C. obesum Dana, 

1851 / None. 

FAO name: En - Brown land crab. 

Diagnostic characters: Carapace subovate, swollen, surface smooth; setose area on pterygostomial and 

subbranchial regions small, not extending to branchiostegal region. Third maxilliped with well-developed 

flagellum on exopod, entirely covered by setae. Adult males with one chela greatly enlarged. Merus of legs 

not distinctly lined with stiff setae. Colour: brown to brownish grey. 


Habitat, biology, and fisheries: In back mangroves or similar brackish-water habitats. Completely 

terrestrial crabs, living near the sea and excavating deep burrows. The larvae are released into the sea 

and return back to land upon completion of their development. Cardisoma carnifex is collected in 

substantial numbers whenever it is common, but has a comparatively low market value. It is usually caught 

by hand at night, or with special traps placed at the mouth of its burrow. The crabs are sold live. There have 

been reports that some land crabs are poisonous but this is almost certainly associated with their diet, as 

these herbivorous crabs sometimes consume poisonous plants. Once they are kept for short periods and 

their guts are empty however, they are safe for human consumption. Three other species of Cardisoma are 

found in the area, C. hirtipes Dana, 1852, C. rotundum and C. longipes (A. Milne Edwards, 1873) (see 

abbreviated species accounts below), but these are only of minor food value, being much less common 

than C. carnifex. 

Distribution: Indo-West Pacific.

1150 Crabs 

Gecarcoidea lalandii (H. Milne Edwards, 1837) 

Frequent synonyms / misidentifications: Hylaeocarcinus humei Wood Mason, 1873; Pelocarcinus 

marchei A. Milne Edwards, 1890; P. cailloti A. Milne Edwards, 1890; Limnocarcinus intermedius De Man, 

1879 / None. 

FAO name: En - Purple land crab. 

Diagnostic characters: Carapace distinctly transversely ovate, swollen, surfaces smooth; orbits relatively 

small, eyes slanted obliquely in adults; third maxilliped without any flagellum on exopod. Colour: purplish 

brown to purple and reddish purple. 


Habitat, biology, and fisheries: Fully terrestrial, can be found many kilometres away from the sea. Normally 

inhabits shallow burrows or under trees and rocks in undisturbed forests in isolated islands, where they may be 

very common. The crabs release their larvae into the sea, which return to land upon completion of their 

development. Gecarcoidea lalandii is only occasionally collected for food, much the same as Cardisoma 

carnifex. Caught by hand or traps throughout their range, and sold live. Its market value, however, is 

comparatively low. 

Distribution: Throughout Indo-West Pacific. 

Remarks: Another species of the genus occurs in the area, G. natalis (Pocock, 1898), known only from 

Christmas and Cocos-Keeling islands. The 2 species are easily distinguished by their coloration: G. lalandii 

is purple whereas G. natalis is bright red.


Cardisoma hirtipes Dana, 1852 

En - Blue land crab. 

Maximum carapace width 12 cm. On more sandy areas behind beaches, usually near the sea. 

Occasionally collected for food by hand or traps, when common. Widely distributed in the Indo-West 

Pacific. 

Cardisoma longipes (A. Milne Edwards, 1873) 

En - Longlegged land crab. 

Maximum carapace width 6 cm. On small isolated islands, in caves or holes. Rarely collected for 

food, due to its small size and general scarcity. Guam, New Caledonia, Niue, Cook Islands, Tuamotu, 

Ocean Island, and Kandavu. 


Cardisoma rotundum (Quoy and Gaimard, 1824) 

En -Rugoselandcrab. 

Maximum carapace width 7.5 cm. On more sandy areas adjacent to reefs. A relatively small, not 

common species, rarely collected for food. Occurs mainly in islands of the South Seas, reaching to 

southern Taiwan Province of China and southern Japan, and the eastern Indian Ocean.

1152 Crabs 

Ocypodidae OCYPODIDAE 


squarish, transversely rectangular, 

trapezoidal or transversely ovate; dorsal 

surface gently convex, usually smooth or with 

grooves; frontal margin entire, relatively 

narrow; orbits broad, occupying almost 

entire anterior border (excluding the front), 

antero- and posterolateral margins of carapace 

usually not clearly demarcated, lateral 

margins appearing almost straight or gently 

convex, lateral margins unarmed. Eyestalks 

long, longer than width of orbit. No rhomboidal 

gap between third maxillipeds. Dactylus 

of legs with numerous stiff setae. Ventral 

surface of abdomen or base of legs may 

have tufts of fine setae. All male abdominal 

segments distinct, movable. 

Ghost crabs 

Habitat, biology, and fisheries: Terrestrial crabs. Most species are very small and are generally of minor 

importance to fisheries. Two larger species of Ocypode are occasionally caught for food, with O. ceratophthalma 

being most often collected. 


Gecarcinidae: a rhomboidal gap present between third maxillipeds (no rhomboidal gap in ocypodids). 

Grapsidae: a rhomboidal gap usually present between the third maxillipeds; frontal margin of carapace less 

narrow as in ocypodids; crabs walk on the sides of the dactyli of legs, not on the tips of the dactyli (as seen 

in ocypodids). 

front broad 


1a. Inner surface of palm without stridulating 

(sound-producing) ridge (Fig. 1a); eyes 

rounded . . . . . . . . . . . . . . Ocypode cordimanus 

1b. Inner surface of palm with stridulating ridge 

no ridge 

(Fig. 1b); eye with long process above tip 

. . . . . . . . . . . . . . . . . Ocypode ceratophthalma 



Ocypode ceratophthalma (Pallas, 1872) 

Ocypode cordimanus Desmarest, 1825 

 

Gecarcinidae 

very narrow front long 

orbits 

carapace 

squarish 

a) Ocypode 

cordimanus 

Grapsidae 

References 

Alcock, A. 1900. Materials for a carcinological fauna of India. No. 6. The Brachyura Catometopa or Grapsoidea. J. Asiat. 

Soc. Bengal, 69, pt. 2 (3):270-486. 

Crosnier, A. 1965. Crustacés Décapodes. Grapsidae et Ocypodidae. Faune de Madagascar, 18:1-143. 

ridge 

b) Ocypode ceratophthalma 

Fig. 1 distal end of cheliped 

(inner surface)

Ocypodidae 1153 

Ocypode ceratophthalma (Pallas, 1872) 


FAO name: En - Horned ghost crab. 

Diagnostic characters: Carapace squarish; 

anterolateral margins unarmed; eyes in 

adults with long projection above cornea; inner 

surface of palm with transverse stridulatory 

ridge. Colour: carapace bluish grey, with 

median parts brown; chelipeds and distal half 

of legs white. 


Habitat, biology, and fisheries: A terrestrial 

species that lives in the lower part of the 

supralittoral zone, on sandy beaches. Excavates 

deep burrows. Caught mainly by hand, 

often at night when the crabs are more active. 

It is sometimes dug up. Only of local economic 

importance, rarely sold in markets. 

Usually fried for human consumption. 


Remarks: The only other species 

in the area of similar size is 

Ocypode cordimanus Desmarest, 

1825 (see below), readily 

distinguishable from O. ceratophthalma 

by lacking the “horned” 

eyes, lacking the stridulatory 

ridge on the palm, and the generally 

paler coloration. 

Ocypode cordimanus Desmarest, 1825 

En - Common ghost crab. 

Maximum carapace width 4.5 cm. Inhabits higher supralittoral habitats not exposed to the sea, 

where it digs deep burrows. Collected by hand for human consumption in some places, but rarely 

seen in markets. Indo-West Pacfic, from South Africa to French Polynesia.

1154 Crabs 

Infraorder Anomura - Coenobitidae Infraorder ANOMURA 

COENOBITIDAE 

Land hermit crabs and coconut crabs 

Diagnostic characters: Carapace relatively well calcified; eyestalk laterally flattened; eyes usually 

held subparallel to each other. Antennae laterally flattened. Coxae of third maxillipeds close 

to each other, without distinct gap between them. Chelipeds short, stocky, equal or unequal; when 

unequal, left chela larger. First 2 pairs walking legs, last 2 pairs reduced, chelate. Abdomen bilaterally 

asymmetrical, not clearly divided into segments. Either hermit crabs or distinctly crab-like animals with 

abdomen tucked under carapace; uropods modified into a “rasp” used for clinging interior of 

gastropod shells (except in adult Birgus latro). 

antenna 

laterally 

flattened 

eye 

laterally 

flattened 

last leg strongly 

reduced 

carapace 


3 rd leg with 

chela 

Habitat, biology, and fisheries: All members of the Coenobitidae are fully terrestrial, but they must return 

to the sea to release their larvae. Most species live in gastropod shells. None of the species, other than 

the “coconut crab” Birgus latro, have any fishery value, although many species of land hermit crabs (genus 

Coenobita) are regularly collected for the pet trade. 


Only the hermit crabs of the families Diogenidae and Paguridae may be confused with coenobitids, but 

their habitats are exclusively aquatic, not terrestrial. In addition, the laterally flattened antennae are highly 

diagnostic for the Coenobitidae. 

References 

Brown, I. W. and D. R. Fielder (eds). 1991. The coconut crab: aspects of Birgus latro biology and ecology in Vanuatu. 

Canberra, Australian Centre for International Agricultural Research (ACIAR), Monograph Number 8:i-x, 128 p. 

Miyake, S. 1965. The Crustacea Anomura of Sagami Bay. Tokyo, Biological Laboratory of the Imperial Household, 161 p.

Infraorder Anomura - Coenobitidae 1155 


Birgus latro (Linnaeus, 1767) 


FAO name: En - Coconut crab. 

Diagnostic characters: Surface of carapace with 

numerous scale-like ridges. Antennae laterally flattened. 

Eyes laterally flattened. Abdomen large, 

tucked underneath carapace. First 2 pairs legs; third 

pair of legs short, chelate; last pair of legs very 

small, chelate, tucked underneath the swollen abdomen; 

surfaces of chelipeds and legs with numerous 

distinct transverse ridges. Colour: carapace 

bluish grey to purplish brown. 

Size: Maximum carapace length 15 cm (or 35 cm, 

if the outstretched chelipeds are included in the 

measurement); weight up to 2.5 kg. 

Habitat, biology, and fisheries: Birgus latro is 

actually a highly modified, fully terrestrial hermit 

crab, and young crabs do in fact inhabit gastropod 

shells, as seen in other members of the family 

Coenobitidae. As they grow, they discard the shell 

and tuck their hardended abdomen (normally soft 

in hermit crabs) under their carapace. Adults are 

fully terrestrial and breathe by means of a special 

“pseudo-lung”. They must, however, return to the 

sea to release their planktonic zoeae. The coconut 

crab tends to occur on small isolated islands, in areas washed by oceanic waters.It is generally a scavenger, 

but also has a preference for coconuts and fruits. One of its common names (“palm thief”) is derived from 

its habit of stealing shiny objects from human habitations. Birgus latro is the largest known land arthropod. 

The coconut crab is a very valuable species and is sold live in markets, where large specimens may 

command prices of up to US$100. It has always been eaten throughout its wide Indo-West Pacific range, 

but its population has nowadays seen a very sharp decline where demand for the species skyrocketed in 

countries like Taiwan and Hong Kong. The very high price it commands in these markets has contributed 

to the serious decline of the species, which has been exterminated from many islands. It is now gradually 

becoming protected throughout much of its range. The species is collected by hand or with baited traps. 

Distribution: Indo-West Pacific, 

reaching eastwards to French 

Polynesia. The only widely distributed 

species in the genus. 



HOLOTHURIANS 

(Sea cucumbers, Class Holothuroidea) 

by C. Conand

1158 Holothurians 


Notes on the phylum Echinodermata 

The holothurians or sea cucumbers belong to the Echinodermata which form a very distinct phylum in 

the animal kingdom. Echinoderms are characterized by the possession of a radial symmetry (generally 

pentamerous), an intradermic skeleton consisting of closely fitted plates, articulated plates, or ossicles, 

and a peculiar water vascular system of tubes filled with fluid. The phylum is divided into 5 classes of very 

different appearance (Fig. 1): the crinoids (or feather stars), holothuroids (or sea cucumbers), echinoids 

(or sea urchins), asteroids (or sea stars), and ophiuroids (or brittle stars). Echinoderms are almost 

exclusively marine, although a few species are found in brackish water. 

pinnules 

madreporite 

podia and spines 

mouth pedicellaria 

anus 

radial 

shield 

ambulacral 

groove 

aboral surface oral surface 

asteroids (sea stars) 

disc 

aboral surface 

arm 

buccal 

shield 

mouth 

bursal 

slit madreporite 

oral surface 

The body is of variable shape, rounded to cylindrical, or 

star-like, and subdivided into 10 areas (Fig. 2): 5 radii (or 

ambulacra) alternating with 5 interradii (or 

interambulacra). The radii correspond to the arms of the 

asteroids, ophiuroids, and crinoids, and to the rows of 

podia or papillae found in sea urchins and sea 

cucumbers. Some groups show a secondary bilateral 

symmetry. The holothurians are elongate orally-aborally 

and lie upon one side. Asteroids, echinoids, and ophiurids 

have the oral surface on the underside of body. 

echinoids (sea urchins) 

mouth 

tentacles 

rows of podia 

ophiurids (brittle stars) 

holothuroids (sea cucumbers) 

Fig. 1 the five classes of echinoderms 

D 

E 

interradius 

EA 

radius A 

madreporite 

arm 

disc 

stem 

A B 

crinoids (feather 

stars) 

bilateral 

symmetry 

anus 

C 

mouth 

(oral surface) 

anus 

(aboral surface) 

Fig. 2 schematic presentation of an echinoderm 

(following the Carpenter system for orientation)


The endoskeleton is produced by the dermis. It may be composed of closely fitted plates forming a rigid 

test or shell as in most echinoids, or articulated plates giving flexibility as in asteroids, ophiuroids, and 

crinoids, or consisting of calcareous spicules dispersed in the body wall, as seen in holothuroids. Spines 

and tubercules are parts of the endoskeleton and are covered by the epidermis. In addition, the body wall 

is composed of extracellular collagen-based material, the viscosity of which is controlled and provides 

rigidity or flexibility. 

The general cavity (or coelomic cavity) contains the viscera. The water vascular system or aquiferous 

system, consists of a circumoral ring, from which arise radially water canals forming many extensions, the 

tube feet or podia. Themadreporite is a plate with tiny pores which allows the communication with the 

environment. It is always situated in an interradial position and serves for the orientation: the opposite radius 

is called A and is followed, in a clockwise direction, by radii named B, C, D, and E (according to the 

“Carpenter system” for orientation, see Fig. 2). 

Despite the differences between the classes, there are several common features in the digestive system 

of echinoderms. This system is conspicuously developed and attached by mesenteries to the body wall. 

Its shape is simple, either bag-like as in ophiuroids, or tube-like, looping in the general cavity as in crinoids, 

echinoids, and holothuroids. In asteroids, it forms conspicuous specialized organs. Structural variations 

appear progressively along the gut. A hemal system is associated with the digestive system. Its importance 

varies among the different classes. 

The respiratory exchange partly occurs by means of the podia, and partly by specialized organs that differ 

among the classes. 

The genital system is composed of a single gonad in holothuroids, a genital cord sending extensions to 

the pinnules of the arms in crinoids, and 5 gonads in the other classes. Sexes are generally separated, but 

cannot be distinguished externally. In most species, mature gametes are released into the sea. After 

fertilization, the development often passes planktonic larval stages with a bilateral symmetry, until the larvae 

metamorphose into benthic juveniles. 

Species identification is done by examination of preserved specimens, mostly based on characters of the 

skeleton. Echinoderms should be preserved in alcohol, as formalin may dissolve the calcareous skeleton. 

In addition to the morphological characters, the colour, size, and ecology of live specimens can be useful 

for identification. 

To date, approximately 6 000 described echinoderm species are known worldwide, living in all kinds of 

marine bottoms where they represent an important component of the benthic biomass. About 1 000 littoral 

species are known to occur in the Indo-West Pacific. 

Among the 5 classes of echinoderms, only echinoids and holothuroids are of interest to fisheries. The 

holothurian fishery of the Western Central Pacific is the largest of the world (Conand, 1997). 

External morphology of holothurians 

Holothurians have an orally-aborally elongated body (Fig. 1).The body is formed like a short or long cylinder, 

with the mouth (at the anterior end) encircled by tentacles, and the anus (at the posterior end) often edged 

by papillae. The pentamerous symmetry is sometimes recognizable by the presence of 5 meridional 

ambulacra bearing podia. Holothurians often lay on the substrate with their ventral surface or trivium, 

formed by the radii A, B, and E in the Carpenter system for orientation. This creeping sole bears the 

locomotory podia, while on the dorsal surface, or bivium, the podia are often represented by papillae. 

Consequently, a secondary bilateral symmetry is evident. The body shape is different in the orders Apoda, 

members of which are vermiform, and Molpadida, members of which have a tail-like region bearing the 

anus. In the order Elasipoda, some extraordinary forms are found, with modified papillae making up sails. 

The mouth is terminal or displaced dorsally, surrounded by a thin buccal membrane, and generally 

bordered by a circle of tentacles (Fig. 1). Tentacles are buccal podia containing extensions from the water 

vascular system. Their number varies between 10 and 30, generally being a multiple of 5. In the 

Aspidochirotida all tentacles are of the same size, but in the Dendrochirotida some tentacles are 

generally smaller. The shape of the tentacles differs among the various orders and is used as a key 

character (Fig. 3). In the Dendrochirotida they are dendritic (branching in an arborescent manner) and can 

reach a large size when extended. The Aspidochirotida and most Elasipoda have peltate tentacles, each 

with a central stalk. The Apoda have pinnate tentacles, with a central axis bearing series of digitations. 

The Molpadida have digitate tentacles, consisting of short projections with small terminal fingers. In all 

cases they are very retractile, particularly in the Dendrochirotida which have an introvert where the 

tentacles insert. The tentacles and the introvert can be contracted into the interior by a set of retractor 

muscles. These muscles also occur in a few Apoda and Molpadida, but not in the other orders.


dentritic peltate pinnate digitate 

Fig. 3 basic types of tentacles 

The body surface is thick, slimy in many species and wears warts, tubercules, or papillae. Podia appear 

on the body wall in 3 orders, but they are lacking in the Apoda and rare in the Molpadida. They typically 

have the form of locomotory tube feet: hollow tubular projections from the body wall form a stem, which 

allows the podium to lengthen, flex, or retract. It contains a branch of the water-vascular system and 

generally does not terminate in a concave sucker (as it is still often called), but in a flat disc, which allows 

the podium to adhere to the substratum during locomotion. Epidermal cells produce adhesive secretions. 

Internally the disc is supported by a large skeletal ossicle. Podia also can have the shape of papillae. The 

tube feet are rarely arranged in 5 regular rows, but generally they loose the discs on the dorsal surface and 

spread into the interradial areas. The anus is often displaced dorsally, encircled by small papillae or anal 

teeth. The coloration varies between species and sometimes also between individuals of the same species. 

The creeping sole is often brighter and lighter than the dorsal surface. 

Body wall 

The body wall is thin in Apodida and Molpadida, but thicker in the other orders, particularly in the 

Aspidochirota. It constitutes the part of the body that is processed for human consumption and therefore 

commercial species are characterized by a thick body wall. Its structure consists of a thin cuticle over the 

epidermis and a thick dermis underneath. The dermis is composed of connective tissue, enclosing the 

endoskeletal spicules or ossicles (see next paragraph). Below the dermis, a layer of circular muscles form 

a cylinder generally interrupted by 5 longitudinal muscle bands situated in the radial positions. 

Spicules 

Also called ossicles or deposits, spicules (Fig. 4) are characteristic of the class and of primary importance 

for identification. These are fenestrated (or perforated) calcareous bits of microscopic size. There is a wide 

variety of simple to complex shapes. Rods can be simple or branching, smooth, warty, or spiny. They can 

bear knobs at their ends, or are a characteristic C- or S-shape. Fenestrated plates also come in various 

forms. Buttons are oval ossicles, perforated with a varying number of holes arranged in 2 rows. Tables 

are more complicated. They appear as a perforated disc, bearing an erect spire (or tower) and show many 

variations according to the arrangement of its constituents. Rosettes are short rods subdivided into short 

branches. Baskets are concave, perforated plates. Anchors are peculiar of the family Synaptidae (order 

Apoda). They are orientated in the body wall, so that they support the attachment to the substrate during 

crawling, in the absence of podia. They are attached to an accompanying perforated plate. Wheels are 

characteristic of the family Chirodotidae (order Apoda) and are also found in the Elasipoda, which are best 

characterized by the presence of special spiny branched spicules. Miliary bodies (grains) are very tiny 

spicules found in some Stichopodidae. Apart from the body wall, spicules are found in the tentacles, the 

podia, and also in the mesenteries or other internal organs. Their developmental stages can differ from the 

definitive shapes in the adults and thus can make species identification difficult.


crown 

cross-bridge 

pillar 

smooth 

C-shaped 

spiracle 

nodose 

spire 

disc 

side view top view pseudo-table 

ellipsoid 

tables 

buttons grains 

rosettes 

plate 

rods 

anchor wheel 

Fig. 4 basic types of spicules 

Calcareous ring 

A ring of usually 10 calcified plates encircles the pharynx. It is composed of alternating larger radial plates, 

opposite to the ambulacra, and smaller interradial plates. The plates may be simple or composed of smaller 

pieces. Longitudinal muscles attach to the radial plates. 

Digestive system and connected organs 

The gut is composed of a pharynx, an esophagus, a stomach, all of which are short structures, and a very 

long intestine (Fig. 5). The intestine consists of 3 portions, a descending, an ascending, and finally a 

descending loop which connects to both the rectum and the cloaca opening outwards through the anus. 

When present, respiratory trees are connected to the cloaca. The oxygenated water enters the body by 

these water lungs, which are found in all orders except the Apoda. Cuvierian tubules, present in several 

species of Aspidochirota, are generally considered as defensive structures. They are composed of sticky 

tubules attached to the base of the respiratory trees and can be expelled in some Holothuria and 

Bohadschia species through the cloaca towards the source of irritation. 

crown 

disc


oral tentacles 

genital orifice 

peripharyngeal 

calcareous ring 

madreporite 

vesicles of oral podia 

Polian vesicle 

genital stolon 

genital gland 

intestine 

transverse vessel 

intestinal cavity 

rete mirabile 

(from Conand, 1986) 

rectum 

cloaca 

anus 

Fig. 5 anatomy of a holothuroid 

(Holothuria nobilis) 

water ring canal 

pharyngeal bulb 

dorsal mesentery 

radial muscular band 

right respiratory tree 

body wall 

papillae 

Cuvierian organs 

muscle fibres 

Reproductive system 

In contrast to other echinoderms, the reproductive system of holothurians consists of a single gonad or 

genital gland (Fig. 5). The gonad is situated dorsal (interambulacrum CD) and composed of either 2 tufts 

of tubules, or only 1 tuft in many species of the family Holothuriidae. The sexes are generally separated 

and show little dimorphism unless in the period of maturing. The gonad is attached to the dorsal mesentery 

through which the gonoduct or genital stolon opening passes, leading to the outside by the gonopore 

(genital orifice) or a genital papilla. In most species, the mature gametes are freely released into the sea 

water. The spawning behaviour, observed in many Aspidochirota species, involves an upright posture of 

males and females followed by a swaying back and forth, while the gametes are being released. 

Water vascular system, perivisceral coelom, and hemal system 

The water vascular system (Fig. 5) is a coelomic space bordered by a mesothelium. It consists of the 

lumen of the buccal tentacles and the tube feet, a water ring around the esophagus, the radial canals, 

the madreporic canal, and the Polian vesicles. The perivisceral coelom is a large cavity containing watery 

proteinaceous coelomic fluid and different forms of cells (coelomocytes). The hemal system is well 

developed and composed of large hemal vessels along the gut, sinus, and lacunae. The hemal vessels 

associated with the gut can form a complex meshwork with the left respiratory tree, the rete mirabile, 

suggesting different functions of nutrient and gas transfers. 

Habitat and biology 

Holothurians are found throughout all oceans, at all latitudes, from the shore down to abyssal plains. They 

are usually benthic (living on the bottom); some species live on hard substrates, rocks, coral reefs, or as 

epizoites on plants or invertebrates; most of the species inhabit soft bottoms, on their surface or in the 

sediment. Among the commercial coastal holothurians, the Aspidochirota are predominant in the tropics, 

while the Dendrochirota are more common in temperate areas. The Aspidochirota from the tropical western 

teats

General Remarks/Glossary of Technical Terms 1163 

Pacific generally show a distribution reflecting the organization of the reef and lagoonal systems. Six 

different categories of species have therefore been defined, being characteristic of the main biotopes in 

which they occur: coral slopes (and passages), inner lagoons, inner reef flats, outer reef flats, outer lagoons, 

and coastal bays. The density and biomass increase from the outer reef slopes to the inner reef flats and 

coastal areas. Although there is much variation between the different sites, coral-slope species generally 

show lower densities and relatively large individual sizes, while species occurring on inner reef flats and 

inner lagoons show intermediate values, and those living on outer reef flats come in higher densities of 

smaller individuals. Most Aspidochirota species have comparatively few animal predators and their major 

effects in the reef communities are related to their deposit-feeding habits, as they are able to rework large 

amounts of sediments (bioturbation). 

Fisheries 

Holothurians have been harvested commercially for at least a thousand years, occasionally for the raw 

body wall or viscera, but mostly in order to be processed into a dry product called bêche-de-mer, trépang, 

or hai-sum, which is considered a delicacy by the Chinese. Harvesting in the tropics is usually done by 

hand, while collecting at low tide or by free-diving from small boats. The processing methods for 

bêche-de-mer include different stages of boiling, gutting, and drying, with variable procedures according 

to the species. Bêche-de-mer is then exported from the producer country to a central market such as Hong 

Kong, and then re-exported to the Chinese consumers. The economic significance of these artisanal 

fisheries is particularly important in less developed countries. The recent developments of these activities 

have led to a global increase of the catches, especially with a strong increase of the Hong Kong and Chinese 

markets, and the participation of new producer countries, with a shift of the exploited species, probably 

due to overcollecting. 

Around 300 shallow-water species of holothurians are known to occur in the area, but only a few of them 

are of commercial interest. From 1990 to 1995, the reported yearly production of sea cucumbers in the 

Western Central Pacific ranged from around 6 800 to 9 000 t (FAO Yearbook of Fishery Statistics). 


As in other echinoderms, species identification is mostly done by examination of the skeletal parts of 

preserved specimens. Holothurians should be preserved in alcohol (70 %), as formalin may dissolve the 

calcareous skeleton. The calcareous ring can be readily observed after dissection. 

The spicules, which are deeply hidden in the body wall, can be obtained by the following method: 

1. Small pieces of body-wall tissue are removed from the bivium and the trivium, as well as the oral 

tentacles and podia, and dissected and macerated in sodium hypochlorite (bleach), or sodium 

hydroxide, in order to dissolve the organic material. 

2. After washing in distilled water, the spicules are rinsed in alcohol and can be processed with a drop 

of a mountant (Canada balsam). 

3. After processing, the spicules can be observed either on permanent slides with a light microscope, 

or prepared for scanning electron microscope. 

General Remarks/Glossary of Technical GLOSSARY Terms OF TECHNICAL TERMS 

Anal teeth - radial calcareous papillae encircling the anus. 

Bêche-de-mer - term used in the tropical Pacific for the processed product of sea cucumbers (see also 

trepang). 

Bivium - the dorsal part of the body in the pentaradiate symmetry, with 2 radii. 

Calacareous ring - internal collar of plates, generally 10, surrounding the pharynx. 

Cloaca - anal cavity where the intestine ends. 

Cuvierian tubules - threads becoming sticky when thrown out of the anus; used as a defense mechanism. 

Dendritic - branching in an arborescent manner; used as descriptive term for the shape of tentacles in 

Dendrochirotida. 

Digitate - finger-like. 

Digitations - finger-like structures. 

Fenestrated - having small window-like openings or holes.


Interradii (or interambulacra) - in the pentaradiate symmetry, the 5 areas between the rows of podia or 

papillae (Fig. 2). 

Papillae - conical lumps on the surface of the body wall. 

Pentamerous - having 5 radiating parts, resulting in a pentaradiate symmetry. 

Peltate - with a central stalk. 

Pinnate - feather-like. 

Podia (or tube feet) - water-filled tubes used for locomotion. 

Radii (or ambulacra) - in the pentaradiate symmetry, the 5 areas with podia or papillae. 

Respiratory tree - arborescent organ (1 pair), opening in the cloaca. 

Spicules - or ossicles, microscopic carbonate skeleton particles in the body wall, useful for species 

identification; they come in various shapes (Fig. 4). 

Teats - large papillae. 

Tentacles - buccal podia (Fig. 3). 

Trépang - Malaysian name for sea cucumber, also used for the processed product (see also 

bêche-de-mer) 

Trivium - the ventral surface of body in the pentaradiate symmetry, with 3 radii. 

KEY TO THE SHALLOW-WATER ORDERS OF THE CLASS HOLOTHUROIDEA 

1a. Podia absent; body vermiform; body wall thin, often translucent; dominant spicules in 

form of anchors with associated anchor plates tentacles pinnate; pharynx without 

retractor muscle; no respiratory tree . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apodida 

1b. Podia present; body-wall moderately thick; body wall with dominant spicules in form of 

tables, perforated plates, buttons, rods, or rosettes . . . . . . . . . . . . . . . . . . . . . . . . → 2 

2a. Tentacles peltate or pelyo-digitate; anterior end of body not introverted and associated 

with retractor muscles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aspidochirotida 

(the only order with commercial species in the area) 

2b. Tentacles branched (dendritic); anterior end of body introverted, associated with retractor 

muscles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrochirotida 

Order dae Aspidochirotida 

ORDER ASPIDOCHIROTIDA 

Diagnostic characters: Tentacles peltate, constituting a short stem ending in a disc covered by small 

digitations. Body wall bearing podia and/or papillae. Calcareous ring without posterior prolongation. 

Pharynx without special retractor muscles, but can be retracted within a fold of skin. Tentacle ampullae 

present. Respiratory trees well developed. Cuvierian organs present or absent. Dominant spicules in form 

of tables, buttons, rods, rosettes, or grains. 

Key to the shallow-water families of Aspidochirotida occurring in the area 

1a. Body with trivium (sole) usually flattened and dorsal bivium convex; gonads forming a 

single tuft appended to the left dorsal mesentery; Cuvierian organs present or absent; 

dominant spicules of form of tables, buttons (simple or modified), and rods (excluding 

C- and S-shaped rods) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holothuriidae (p. 1165) 

1b. Body square-shaped or trapezoidal in cross-section; Cuvierian organs always absent; 

gonads forming 2 tufts appended on each side of the dorsal mesentery; dominant 

spicules in form of branched rods and C-and S-shaped rods . . . . . . . . Stichopodidae (p. 1185) 



Order Aspidochirotida - Holothuriidae 1165 

Order Aspidochirotida - Holothuriidae HOLOTHURIIDAE 

Diagnostic characters: Body dome-shaped in cross-section, with trivium (or sole) usually flattened 

and dorsal bivium convex and covered with papillae. Gonads forming a single tuft appended to the left 

dorsal mesentery. Tentacular ampullae present, long, and slender. Cuvierian organs present or absent. 

Dominant spicules in form of tables, buttons (simple or modified), and rods (excluding C-and S-shaped 

rods). 

Key to the genera and subgenera of Holothuriidae occurring in the area (after Clark and Rowe, 1971) 

1a. Body wall very thick; podia and papillae short, more or less regularly arranged on bivium 

and trivium; spicules in form of rods, ovules, rosettes, but never as tables or buttons . . . . . . → 2 

1b. Body wall thin to thick; podia irregularly arranged on the bivium and scattered papillae 

on the trivium; spicules in various forms, with tables and/or buttons present . . . (Holothuria) → 4 

2a. Tentacles 20 to 30; podia ventral, irregularly arranged on the interradii or more regularly 

on the radii; 5 calcified anal teeth around anus; spicules in form of spinose rods and 

rosettes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Actinopyga 

2b. Tentacles 20 to 25; podia ventral, usually irregularly arranged, rarely on the radii; no 

calcified anal teeth around anus, occasionally 5 groups of papillae; spicules in form of 

spinose and/or branched rods and rosettes . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Podia on bivium arranged in 3 rows; spicules comprise rocket-shaped forms . . . . Pearsonothuria 

3b. Podia on bivium not arranged in 3 rows; spicules not comprising rocket-shaped forms . . . Bohadschia 

4a. Spicules in form of well-developed tables, rods and perforated plates, never as buttons . . . . . → 5 

4b. Spicules in form of tables, occurring alone or along with buttons, rods, or rosettes . . . . . . . → 7 

5a. Spicules never in form of rosettes;tables with no or reduced disc, bearing spines terminating 

in form of a maltese cross; body cylindrical; body wall soft . . . . . . . . Holothuria (Semperothuria) 

5b. Spicules in form of rosettes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

6a. Podia in 3 rows on trivium; small papillae dispersed on bivium; spicules in form of 

rosettes, and tables with reduced disc, bearing a spine ending in a maltese cross (when 

viewed from above) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holothuria (Halodeima) 

6b. Podia not very numerous, irregularly arranged in 10 rows (radial and interradial); papillae 

on bivium irregularly arranged in 10 rows; spicules in form of rosettes, and tables with 

disc in a “cup and saucer form”, bearing a low, wide spire ending in a spinose crown 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holothuria (Acanthotrapeza) 

7a. Spicules in form of smooth buttons or pseudo-buttons; tables variously developed . . . . . . . → 8 

7b. Spicules in form of buttons, always knobby, occasionally in form of ellipsoids; tables 

strongly developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

8a. Tables well developed, their disc usually squarish with smooth rim and 8 holes, 

occasionally strongly perforated . . . . . . . . . . . . . . . . . . . . . . Holothuria (Thymiosycia) 

8b. Tables moderately developed, their disc with usually notched rim; buttons of variable 

shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

9a. Tentacles 16 to 30; tables with notched rim and a low spire terminating in a ring of 

irregular spines; pseudo-buttons abundant, irregular; collar of papillae present around 

base of tentacles and anus . . . . . . . . . . . . . . . . . . . . . . . . Holothuria (Lessonothuria) 

9b. Tentacles 18 to 20; some tables often with narrow disc, spire reduced; other tables 

having a well-developed disc and spire with 4 columns; buttons irregular; no collar of 

papillae around base of tentacles . . . . . . . . . . . . . . . . . . . Holothuria (Mertensiothuria) 

10a. Buttons never modified to form fenestrated ellipsoids . . . . . . . . . . . . . . . . . . . . . . → 11 

10b. Buttons modified to form fenestrated ellipsoids, and buttons . . . . . . . . . . . . . . . . . . → 12


11a. Tables with wavy disc, bearing a small spire ending in small spines, giving the table a 

tack-like shape, with a large central hole surrounded by peripheral holes variable in 

number; presence of buttons, pseudo-buttons, and small irregular C-, S-, or O-shaped 

spicules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holothuria (Stichothuria) 

11b. Tables well developed, with smooth disc, spire ending in a cluster of small spines, never 

giving the table a tack-like shape; presence of buttons with irregular knobs . . Holothuria (Metriatyla) 

12a. Calcified anal papillae absent; collar of papillae around base of tentacles present or 

absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 13 

12b. Calcified anal papillae present; no collar of papillae around base of tentacles; body wall 

very thick and rigid; tables with wavy disc, bearing a low and massive spire ending in a 

very spinose crown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holothuria (Microthele) 

13a. Collar of papillae around base of tentacles; anal teeth present or absent; tables with 

notched disc and a low to moderate spire, ending in a cluster of very small, or numerous 

spines; a fringe of rather large papillae between bivium and trivium . . . Holothuria (Theelothuria) 

13b. No collar of papillae around base of tentacles; anal teeth absent; tables with knobby 

disc and low spire, ending in a cluster of spines; buttons knobby, occasionally modified 

to fenestrated ellipsoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holothuria (Cystipus) 



Actinopyga echinites (Jaeger, 1833) 

Actinopyga mauritiana (Quoy and Gaimard, 1833) 

Actinopyga miliaris (Quoy and Gaimard, 1833) 

Actinopyga palauensis Panning, 1944 

Actinopyga spinea Cherbonnier, 1980 

Bohadschia argus Jaeger, 1833 

Bohadschia similis (Semper,1868) 

Bohadschia vitiensis (Semper, 1868) 

Holothuria (Acanthotrapeza) coluber Semper, 1868 

Holothuria (Halodeima) atra Jaeger, 1833 

Holothuria (Halodeima) edulis Lesson, 1830 

Holothuria (Mertensiothuria) leucospilota (Brandt, 1835) 

Holothuria (Metriatyla) scabra Jaeger, 1833 [including H. (M.) scabra var. versicolor (Conand, 1986)] 

Holothuria (Microthele) fuscogilva Cherbonnier, 1980 

Holothuria (Microthele) fuscopunctata Jaeger, 1833 

Holothuria (Microthele) nobilis (Selenka, 1867) 

Pearsonothuria graeffei (Semper, 1868) 

References 

Cannon, L.R.G and H. Silver. 1986. Sea cucumbers of Northern Australia. Brisbane, Queensland Museum, 60 p. 

Clark, A.M. and F.W. Rowe. 1971. Holothurioidea. In Monograph of shallow water Indo-West Pacific echinoderms. 

London, Trustees of the British Museum, pp. 171-210. 

Conand, C. 1989. Les holothuries aspidochirotes du lagon de Nouvelle-Calédonie: biologie, écologie et exploitation. 

Paris, Etudes et thèses ORSTOM, 393 p. 

Conand, C. 1990. The fishery resources of Pacific island countries. Part 2: Holothurians. Rome, FAO Fish. Tech. Pap., 

(272.2):143 p. 

Conand, C. 1997. Are holothurian fisheries for export sustainable? Intern. Cong. Reefs Panama, 2:2021-2026. 

Feral, J.P. and G. Cherbonnier. 1986. Les holothurides. In Guide des étoiles de mer, oursins et autres échinodermes du 

lagon de Nouvelle-Calédonie, edited by Guille, Laboute, and Menou. Paris, ORSTOM, pp. 56-107. 

Tan Tiu, A. 1981. The intertidal Holothurian fauna (Echinodermata: Holothuroidea) of Mactan and the neighbouring 

islands, Central Philippines. Philipp. Sci., 18:45-119.


Actinopyga echinites (Jaeger, 1833) 

Frequent synonyms / misidentifications: None / Actinopyga mauritiana (Quoy and Gaimard, 1833). 

FAO names: En - Deep-water redfish. 

radial 

anus with 

interradial 

5 teeth 

bivium with slender papillae 

spicules of 

tentacles 


spicules of 

ventral podia 

mouth ventral, 

with 20 short and 

stout tentacles 

spicules of ventral tegument 

(after Féral and Cherbonnier, 1986) 

spicules of dorsal tegument 

Diagnostic characters: Body elongate, arched dorsally (bivium) and flattened ventrally (trivium). Bivium 

sometimes wrinkled and covered by sediment, wider in the middle and tapering towards both ends. Papillae 

on bivium long and slender; podia on trivium arranged more or less regularly in 3 rows; calcareous disc 

of podia around 350 µm in diameter. Mouth ventral, surrounded by 20 short, stout tentacles. Anus 

surrounded by 5 anal teeth. Calcareous ring with large radial pieces and narrow interradials. Cuvierian 

tubules few (10), short, pink, and never expelled. Colour: bivium brown, varying from light to dark among 

specimens; trivium lighter, sometimes orange, with yellow podia. Spicules: ventral tegument with a variety 

of small rosettes and rods, occasionally perforated; dorsal tegument with larger rosettes and branched 

plates; ventral podia with some straight, spinous rods; tentacles with rods large, straight, or curved, and 

very spiny at their extremities. 

Size: Maximum length about 35 cm, commonly to about 20 cm; mean live weight 0.3 kg (up to 1 kg); 

body-wall thickness about 7 mm. 

Habitat, biology, and fisheries: Dwelling in moderately shallow waters (despite its English common 

name), rarely found in depths of more than 12 m; mostly on reef flats of fringing and lagoon-islet reefs. 

Abundant in seagrass beds, on rubble reef flats, and compact flats where populations can reach high 

densities (up to 1/m 2 ). Sexual reproduction during the warm season. A species with a high potential 

fecundity and early sexual maturity. Harvested in artisanal fisheries throughout the area. Collected by hand 

while wading on the reefs at 

low tide. The processed 

product is not distinguished 

from those of other Actinopyga 

species and has a moderate 

commercial value. 


the tropical Indo-Pacific, 

including the Red Sea, but 

excluding the Persian Gulf and 

Hawaii.


Actinopyga mauritiana (Quoy and Gaimard, 1833) 

Frequent synonyms / misidentifications: None / Actinopyga echinites (Jaeger, 1833). 

FAO names: En - Surf redfish; Fr - Holothurie brune des brisants. 

interradial radial 

white spots 


spicules of tentacles 


mouth ventral, with 

25 tentacles 

spicules of dorsal podia 


anus with 

5 stout teeth 


Diagnostic characters: Body elongate, arched dorsally (bivium) and flattened ventrally (trivium). Bivium 

sometimes wrinkled, wider in the middle and tapering towards both ends. Papillae on bivium long and 

slender; podia numerous on trivium, stout and crowded on the radii and interradii; calcareous disc of podia 

around 500 µm in diameter. Mouth ventral, surrounded by 25 short and stout tentacles, with a large 

collar of long papillae at their base. Anus surrounded by 5 stout anal teeth. Calcareous ring showing large 

radial pieces and interradials with a triangular top. Cuvierian tubules few (10), short, pink and never 

expelled. Colour: bivium variable, usually dark brown, with more or less distinct whitish spots; trivium 

lighter, occasionally white to pink, with yellow podia. Spicules: ventral tegument with round and biscuit-like 

grains, rods bearing small spines, and denticulate pseudo-plates; dorsal tegument without grains, but with 

rosettes and denticulate rods; podia with some spinous rods and large rosettes; tentacles with large rods 

bearing small spines. 

Size: Maximum length about 35 cm, commonly to 20 cm; mean live weight about 0.3 kg (up to 1 kg); 


Habitat, biology, and fisheries: Dwelling in very shallow waters, rarely found in depths of more than 20 m; 

mostly on outer reef flats of barrier reefs and fringing reefs exposed to strong hydrodynamism. Most abundant 

in the surf zone where populations can reach high densities (more than 1/m 2 ). Sexual reproduction takes place 

during the warm season. A species with a high potential fecundity and early sexual maturity. Harvested in 

artisanal fisheries throughout 

the area, but probably less 

frequently than other species 

of Actinopyga. Collected by 

hand while wading on the reef at 

low tide. The processed product 

is not distinguished from those of 

other Actinopyga species and 

has a moderate commercial 

value. 


the tropical Indo-Pacific.


Actinopyga miliaris (Quoy and Gaimard, 1833) 

Frequent synonyms / misidentifications: None / Actinopyga palauensis Panning, 1944; A. spinea 

Cherbonnier, 1980; other black species of Actinopyga. 

FAO names: En - Hairy blackfish. 

long and numerous papillae 


spicule of tentacles 

spicules of ventral 

podia 


20 stout tentacles 

spicules of dorsal 

podia 


tegument 


anus with 

5 teeth 


tegument 

Diagnostic characters: Body elongate, cylindrical, slightly arched dorsally (bivium) and flattened ventrally 

(trivium). Bivium generally covered by mucus and fine sediment. Papillae on bivium numerous, long and 

slender; long and thick podia on trivium arranged more or less regularly in tight rows on the radii; calcareous 

disc of podia around 700 µm in diameter. Mouth ventral, surrounded by 20 stout tentacles. Anus 

surrounded by 5 strong, triangular anal teeth. Calcareous ring with large radial pieces and narrow 

interradials. Cuvierian tubules absent. Colour: bivium black; trivium lighter, dark brown. Spicules: ventral 

tegument with a variety simple or more complicated rosettes; dorsal tegument with rosettes; ventral and 

dorsal podia with some rosettes; podia and papillae containing large spicules; tentacles with large rods. 

Size: Maximum length about 35 cm, commonly to about 25 cm; mean live weight about 0.4 kg (up to 1 kg); 


Habitat, biology, and fisheries: Dwelling in moderately shallow waters, rarely found in depths of more 

than 10 m; mostly on reef flats of fringing and lagoon-islet reefs, never on barrier reefs. Abundant in 

seagrass beds and on rubble reef flats where populations can reach high densities (up to 1/m 2 ). Known to 

feed on epiphytes and seagrass leaves. Biology poorly known. Harvested in artisanal fisheries throughout 

the area. Collected by hand while wading on the reefs at low tide, or by divers. The processed product is 

not distinguished from those of 


and has a moderate 



the tropical Indo-Pacific, 


excluding the Persian Gulf and 

Hawaii.


Actinopyga palauensis Panning, 1944 

Frequent synonyms / misidentifications: None / Actinopyga miliaris (Quoy and Gaimard, 1833); 

A. spinea Cherbonnier, 1980; other black species of Actinopyga. 

FAO names: En - Panning’s blackfish. 




podia 

spicule of dorsal 

podia 


20 stout tentacles 


arched bivium 

spicules of tegument 

anus with 5 

strong anal 

teeth 

Diagnostic characters: Body elongate, cylindrical, slightly arched dorsally (bivium) and flattened 

ventrally (trivium). Bivium generally covered by mucus and fine sediment. Papillae on bivium small and 

conical; long and thick podia on bivium arranged more or less regularly in tight rows on the radii; calcareous 

disc of podia around 350 µm in diameter. Mouth ventral, surrounded by 20 stout tentacles. Anus 

surrounded by 5 strong, triangular anal teeth. Calcareous ring with large radial pieces showing an arch 

at their base, and narrow interradials. Colour: bivium black; trivium lighter, dark brown. Spicules: ventral 

and dorsal tegument with a variety of shredded rods and X-shaped spicules; ventral podia and dorsal 

papillae with spiny rods; tentacles with large rods (up to 700 µm). 

Size: Maximum length about 40 cm, commonly to about 30 cm; mean live weight 0.5 kg (up to 1.2 kg); 


Habitat, biology, and fisheries: Occurs in moderately shallow waters, rarely in depths of more than 25 m; 

never found on reef flats, but on flagstones of reef slopes. Populations reach medium densities (up to 

0.1/m 2 ). A poorly known species. Probably collected by divers for artisanal fisheries, but not intensively so, 

as it is found in lower densities 

and has a deeper distribution 

than other “blackfish” species. 

The processed product is not 

distinguished from those of 


has a moderate commercial 

value. 

Distribution: Known only from 

Palau and New Caledonia.


Actinopyga spinea Cherbonnier, 1980 

Frequent synonyms / misidentifications: None / Actinopyga miliaris (Quoy and Gaimard, 1833); 

A. palauensis Panning, 1944; other black species of Actinopyga. 

FAO names: En - New Caledonia blackfish. 



spicules of anal tegument 

anus subdorsal, with 

5 nodose teeth 

spicules of 

peristome 


layer of sediment on bivium 

spicules of 

podia 


with 20 stout 

tentacles 


podia 

Diagnostic characters: Body elongate, cylindrical, slightly arched dorsally (bivium) and flattened ventrally 

(trivium). Bivium generally covered by fine sediment. Papillae on bivium small, slender, and conical; 

cylindrical and thick podia on trivium arranged irregularly on the radii and interradii; calcareous disc of 

podia around 700 µm in diameter. Mouth ventral, surrounded by 20 stout tentacles. Anus subdorsal, 

surrounded by 5 strong, triangular, and nodose anal teeth. Calcareous ring thick, with large radial pieces 

and narrow interradials. Cuvierian tubules absent. Colour: bivium black; trivium dark brown. Spicules: 

sparse in the ventral and dorsal tegument, abundant only near the mouth, anus, and tentacles; a few rods 

in the dorsal tegument give rise to numerous rosettes, rods, and various plates; small to large plates 

provided with holes and spines near the anus; tentacles with rods straight or arched, bearing small spines. 

Size: Maximum length about 38 cm, commonly to about 27 cm; mean live weight about 0.7 kg (up to 1.2 kg); 


Habitat, biology, and fisheries: In moderately shallow water from depths of 5 to 30 m, on the lagoon floor, 

sometimes burrowed in sandy-muddy sediments. Populations reach medium densities (up to 0.1/m 2 ). A 

poorly known species. May be collected by divers for artisanal fisheries along with other “blackfish” species, 

but not intensively so, since it is found in comparatively low densities and has a deeper distribution. The 

processed product is not 

distinguished from those of 


probably has a moderate 


Distribution: Known only from 

New Caledonia.


Bohadschia argus Jaeger, 1833 


FAO names: En - Leopard fish; Fr - Holothurie léopard. 




20 dark tentacles 


tegument 



tegument 

Diagnostic characters: Body cylindrical, arched dorsally (bivium) and flattened ventrally (trivium). Bivium 

smooth. Podia on bivium small, conical, and irregularly arranged; podia on trivium numerous, short, and 

arranged on the radii and interradii; calcareous disc of podia around 400 µm in diameter. Mouth ventral, 

surrounded by 20 short, dark tentacles. Anus nearly dorsal, surrounded by 5 groups of papillae. 

Calcareous ring with large radial pieces and narrow interradials. Cuvierian tubules numerous and large. 

Colour: bivium whitish to brown, variable among specimens and showing characteristic dark eye-like 

spots, with a podia placed in their middle and encircled with a light colour; trivium lighter, yellow to brown; 

some specimens uniformly brown with yellow spots. Spicules: ventral tegument with various simple 

rosettes and biscuit-like nodules, occasionally with small holes; dorsal tegument with rosettes only; ventral 

podia with short rods; dorsal podia with X-shaped rods; tentacles with spiny rods and large irregular plates. 



Habitat, biology, and fisheries: Found in shallow waters, rarely in depths of more than 30 m; a typical 

reef species, generally on barrier reef flats and slopes, or outer lagoons on white sand. Populations never 

reach high densities (generally between 0.001 and 0.01/m 2 ). Symbiotic pearlfish (Carapidae, Ophidiiformes) 

are often found in the respiratory tree or the general cavity. Biology poorly known. Sexual reproduction 

probably takes place during the warm season. Occasionally collected in artisanal fisheries, although the 

sticky Cuvierian tubules make 

the collection and processing 

disagreeable. Collected by 

divers. The processed product 

is not distinguished from those 

of other Bohadschia species 

and is of little commercial 

value. 

Distribution: Widespread 

in the tropical Indo-Pacific, 


excluding Hawaii. 

spots 

anus subdorsal, 

with 5 groups of 

papillae 

spicules of podia


Bohadschia similis (Semper,1868) 


FAO names: En - Brownspotted sandfish. 

bivium with 

brown spots 



spicules of 

dorsal podia 

spicules of ventral podia (after Féral and Cherbonnier, 1986) 

mouth ventral with 

20 short tentacles 


tegument 

podia long 

and slender 

anus dorsal 


tegument 

Diagnostic characters: Body cylindrical, flattened ventrally (trivium). Bivium smooth and rigid. Podia on 

bivium long, slender, conical, and irregularly arranged; podia on trivium more numerous, shorter and 

irregularly arranged on the radii and interradii; calcareous disc of podia around 400 µm in diameter. Mouth 

ventral, surrounded by 20 short tentacles. Anus nearly dorsal, surrounded by 5 groups of radial 

papillae. Calcareous ring with large radial pieces and narrow interradials having sharp tops. Cuvierian 

tubules long and large. Colour: variable among specimens and localities; bivium beige to light brown, 

showing characteristic brown spots; trivium lighter, yellowish. Spicules: ventral tegument with simple 

nodules with or without holes and sparse rods giving rise to rosettes; dorsal tegument with rosettes and 

small biscuit-like plates; ventral podia with variable rods; dorsal podia with X-shaped rods; tops of tentacles 

with spiny rods. 



Habitat,biology,and fisheries:Foundinveryshallowwaters,rarelyindepthsofmorethan3m;occursincoastal 

lagoons or inner reef flats, generally burrowing in sandy-muddy bottoms. Populations can reach densities of 

0.03/m 2 . Biology poorly known. May be harvested along with other reef flat species, such as Holothuria scabra, 

when they occur together, but the sticky Cuvierian tubules make the collection and processing disagreeable. 

Collectedbydivers.Theprocessed 

product is not distinguished 

from those of other Bohadschia 

species and is of little commercial 

value. 

Distribution: Widespread in the 

tropical Pacific, excluding Hawaii; 

reported from the Mascarene 

Islands, but not from elsewhere 

in the Indian Ocean.


Bohadschia vitiensis (Semper, 1867) 

Frequent synonyms / misidentifications: None / Bohadschia marmorata Jaeger, 1833; B. bivittata 

(Mitsukuri, 1912). 

FAO names: En - Brown sandfish; Fr - Holothurie brune. 



mouth ventral 

with 20 short 

tentacles 


tegument 



podia 

Diagnostic characters: Body cylindrical, arched dorsally (bivium) and flattened ventrally (trivium); 

calcareous disc of podia around 300 µm in diameter. Mouth ventral, surrounded by 20 small, short and 

yellowish tentacles. Anus nearly dorsal. Calcareous ring with large radial pieces and narrow interradials. 

Cuvierian tubules numerous and thick. Colour: bivium yellow with brown bands; trivium lighter; podia 

on bivium and trivium encircled by a characteristic brown spot. Spicules: ventral tegument with a 

variety of pseudo-rosettes and biscuit-like nodules; dorsal tegument with rosettes and a few rods; ventral 

podia with various rods; tentacles with rods, straight or slightly curved and very spiny. 



Habitat, biology, and fisheries: Found in moderately shallow waters, rarely in depths of more than 20 m; 

mostly on coastal lagoons and inner reef flats; abundant in sandy-muddy sediments where it burrows most 

of the time. Population densities generally less than 0.02/m 2 . May be harvested with other reef flat species, 

such as Bohadschia similis and Holothuria scabra, when they occur together, but the sticky Cuvierian 

tubules make the collection and processing disagreeable. Collected by hand at low tide, or by divers. The 

processed product is not distinguished from those of other Bohadschia species and is of little commercial 

value. 


the tropical Indo-Pacific. 

podia 


tegument 

anus 

subdorsal


Holothuria (Acanthotrapeza) coluber Semper, 1868 


FAO names: En - Snake fish; Fr - Holothurie serpent. 


spicules of 

podia 

spicules of 

tentacles 

anus with 

papillae 

trivium with conical yellow podia 




with 20 long, 

yellow tentacles 

Diagnostic characters: Body cylindrical, very elongate, larger near the posterior end. Tegument very 

tough. Short papillae on bivium emerging from warts; podia on trivium conical, stout, yellow, arranged 

in 10 rows on the radii and interradii; calcareous disc of podia around 400 µm in diameter. Mouth ventral, 

surrounded by 20 very long, stout tentacles of characteristic yellow colour; mouth also surrounded 

by a collar of small papillae. Anus surrounded by 5 groups of 3 small papillae. Calcareous ring with 

large and high radial pieces and narrow interradials. Cuvierian tubules absent. Colour: black, with yellow 

podia. Spicules: tegument with tables and pseudo-buttons; tables with circular disc showing 8 holes, 

bearing a spire of 4 pillars, with spiny crown, hollow in the middle; pseudo-buttons plate-like with denticulate 

border; rods in the podia and papillae smooth, with holes at their ends; tentacles with nodulous rods. 



Habitat, biology, and fisheries: Dwelling in moderately shallow water, rarely found in depths of more than 

15 m; mostly on inner reef flats of fringing and lagoon-islet reefs and shallow coastal lagoons; abundant 

on sandy-muddy grounds with rubble or coral patches, where it hides the posterior part of body. Population 

densities between 0.01 and 0.2/m 2 . Biology very poorly known. Not known to be traditionally harvested, 

but in recent times, due to increasing demand, this species also appears in the processed products of 

some Pacific Islands. Collected by hand while wading on the reefs at low tide, or by divers. The processed 

product is of low commercial 

value. 


the tropical Pacific, excluding 

Hawaii; not recorded in the 

Indian Ocean.


Holothuria (Halodeima) atra Jaeger, 1833 

Frequent synonyms / misidentifications: None / Holothuria leucospilota (Brandt, 1835). 

FAO names: En -Lollyfish. 


spicule of 

ventral podia 

spicule of 

tentacles 


with 20 tentacles 

black patches 

anus 

terminal 

Diagnostic characters: Body cylindrical, elongate, with rounded ends. Tegument smooth, often covered 

by sand, but also showing round patches lacking sand. A red toxic fluid is secreted upon rubbing the 

body surface vigorously. Podia on bivium sparsely distributed, ending in a small disc around 150 µm in 

diameter; podia on trivium numerous, short and stout, distributed on the radii and the interradii, their 

calcareous disc around 500 µm in diameter. Mouth ventral, surrounded by 20 black tentacles. Anus 

terminal. Calcareous ring with large radial pieces and narrow interradials. Cuvierian tubules absent. 

Colour: entirely black. Spicules: tegument with tables and rosettes; tables with circular disc showing 8 

holes (4 central and 4 smaller holes in between) and a spire of 4 pillars, ending in a maltese crown; rosettes 

small and simple, more abundant in ventral tegument; ventral podia without rods, but with pseudo-plates; 

dorsal podia and papillae with short rods, showing denticulate borders. 



Habitat, biology, and fisheries: The most common shallow-water species in the area, rarely found in 

depths of more than 20 m; mostly on inner and outer reef flats and back reefs or shallow coastal lagoons; 

abundant on sandy-muddy grounds with rubble or coral patches and in seagrass beds.The mean population 

density is around 0.5/m 2 , but can exceed 4/m 2 . Inshore shallow-water populations are denser, composed 

of smaller individuals, and reproduce mostly by transversal fission, while in deeper or outer reef populations 

the individuals are more scattered, larger, and reproduce sexually. Traditionally harvested, but the 

processed product is of low 

commercial value. In recent 

times, due to increasing 

demand, this species also 

appears in the processed 

products of many Pacific 

Islands. Collected by hand at 

low tide while wading on the 

reefs, or by divers. 


the tropical Indo-Pacific. 

sand 


(after Féral and Cherbonnier, 1986)


Holothuria (Halodeima) edulis Lesson, 1830 


FAO names: En - Pinkfish; Fr - Trépang rose. 



spicule of podia 


with 20 grey 

tentacles 


dark dorsal band 


anus 

subdorsal 

Diagnostic characters: Body cylindrical, elongate, with rounded ends. Tegument rough. Podia sparse on 

bivium, ending in a small disc of around 100 µm diameter; podia on trivium numerous, short and stout, 

distributed on the radii and interradii, their calcareous disc around 460 µm in diameter. Mouth ventral, 

surrounded by 20 grey tentacles. Anus subdorsal. Calcareous ring with large radial pieces and narrow 

interradials. Cuvierian tubules absent. Colour: a characteristic black or dark medio-dorsal band; 

coloration lighter laterally; trivium lighter, with small, dark dots. Spicules: tegument with tables and rosettes; 

tables with circular small disc, having a single hole in the middle, and bearing a spire ending in spines 

which may form a maltese crown; presence of rosettes with 4 holes (2 small and 2 larger holes); a second 

form of rosettes with 6 holes and small perforated plates mostly found in the dorsal tegument; ventral podia 

with large plates and rods; dorsal podia with nodose rods or showing holes at their ends. 



Habitat, biology, and fisheries: A common shallow-water species in the area, rarely found in depths of 

more than 30 m; mostly on inner and outer flats of coastal reefs, back reefs, or shallow coastal lagoons. 

Specimens from barrier reefs have been reported to have a brown bivium and a whitish trivium. Most 

abundant on sandy-muddy grounds with rubble or coral patches and in seagrass beds. Mean population 

density around 0.01/m 2 (not 

exceeding 0.1/m 2 ). Not 

traditionally harvested. The 

processed product looks 

similar to that of Holothuria 

atra and is of low commercial 

value. 




Holothuria (Mertensiothuria) leucospilota (Brandt, 1835) 

Frequent synonyms / misidentifications: None / Holothuria atra Jaeger, 1833; H. coluber Semper, 1868. 

FAO names: En - White threads fish; Fr - Trépang à canaux blancs. 

bivium with slender 

papillae 



spicule of dorsal podia 

spicule of ventral podia 



mouth with 20 

black tentacles 

anus 

subdorsal 

Diagnostic characters: Body very elongate, narrower anteriorly than posteriorly. Tegument very smooth. 

Podia and papillae randomly distributed on bivium, the podia ending in a disc of around 480 µm diameter; 

podia on bivium numerous, short and stout, distributed on the radii and interradii, their calcareous disc 

around 700 µm in diameter. Mouth ventral, surrounded by 20 black tentacles. Anus subdorsal. 

Calcareous ring with large radial pieces and triangular interradials. Cuvierian tubules very thin and long. 

Colour: entirely black. Spicules: dorsal and ventral tegument with tables and buttons; tables with circular 

large disc, having 8 holes (or more), spire with 4 pillars, and ending in a crown with large central hole; 

buttons regular, with 6 or 8 holes, or irregular; plates large in ventral podia, with many holes; dorsal podia 

also with long rods; tentacles containing few rods. 



Habitat, biology, and fisheries: A common shallow-water species in the area, rarely found in depths of more 

than 10 m; mostly on outer and inner reef flats, back reefs, and shallow coastal lagoons. Abundant in seagrass 

beds, sandy-muddy grounds with rubble or coral patches where it hides the posterior part of body. Mean 

population density around 0.05/m 2 (can exceed 0.5/m 2 ). Inshore, shallow-water populations are denser, 

composed of smaller individuals and reproduce mostly by transversal fission, while in deeper or outer reef 

populations the individuals are 

more scattered, larger, and 

reproduce sexually. Not 

traditionally harvested, due to 

the thin tegument and the 

presence of Cuvierian tubules, 

but may be confused with other 

commercial black species of 

Holothuria. 




Holothuria (Metriatyla) scabra Jaeger, 1833 

Frequent synonyms / misidentifications: None / Holothuria (Metriatyla) scabra var. versicolor 

(Conand, 1986). 

wrinkles 

FAO names: En -Sandfish. 

anus terminal 

with 20 papillae 


spicule of podia 


20 short tentacles 


Diagnostic characters: Body oval, arched dorsally (bivium) and flattened ventrally (trivium). Bivium with 

characteristic wrinkles, covered by sediment when the animal is coming out of the bottom. Bivium with 

small papillae within black dots, and black podia ending in a disc of around 220 µm diameter; podia on 

trivium arranged irregularly, their calcareous disc around 350 µm in diameter. Mouth ventral, surrounded 

by a collar of papillae and 20 grey, short and stout tentacles. Anus terminal, surrounded by a circle 

of 5 groups of radial papillae. Calcareous ring with a large medio-ventral radial piece. Cuvierian tubules 

absent. Colour: highly variable; bivium whitish to dark brown, occasionally with dark transverse markings; 

trivium lighter, generally whitish. Spicules: very numerous; ventral tegument with tables and buttons, the 

tables having a moderately small disc, perforated by a varying number of holes of variable size, the spire 

with 4 pillars and a cross-like bridge, ending in a spiny crown with a hole in the middle; knobby buttons 

generally have 6 holes, occasionally more; small rods and denticulate plates also present; tables and 

buttons in dorsal tegument: tables with a nodose disc, much larger than in ventral tegument; buttons 

variable, larger, and more perforated than those in ventral tegument; ventral and dorsal podia with long and 

large spinose rods; tentacles with long spiny rods, small smooth rods, and large denticulate plates. 



Habitat, biology, and fisheries: Found in shallow waters, rarely in depths of more than 10 m; mostly on inner 

reef flats of fringing and lagoon-islet reefs, coastal areas under terrigenous influence, and near mangroves. 

Burrows in mud and sandy-muddy bottoms where the populations can reach high densities (up to 1/m2 spicules of tentacles (after Féral and Cherbonnier, 1986) 


).Sexual 

reproduction takes place during the warm season. A species with a high potential fecundity and early sexual 

maturity. Provides the principal share of the tropical Indo-Pacific production of bêche-de-mer and is 

harvested throughout the area in artisanal 

fisheries. Collected by hand while wading 

on the reefs at low tide. The processing 

method is particular of this species: the 

sea cucumbers are buried overnight, and 

the next day the numerous spicules are 

removed by brushing the tegument. The 

processed product is of major commercial 

value and at present highly demanded. 

Distribution: Widespread in the tropical 

Indo-Pacific, excluding Hawaii.


Holothuria (Metriatyla) scabra var. versicolor (Conand, 1986) 

Frequent synonyms / misidentifications: None / Holothuria albiventer Semper, 1868; H. acculeata 

Semper, 1868; H. (Metriatyla) scabra Jaeger, 1833. 

FAO names: En - Golden sandfish. 


spicules of podia 


mouth with 20 

short tentacles 



bivium with more or less dark 

dots, large papillae, no wrinkles 

anus terminal 

with papillae 

Diagnostic characters: Body oval, arched dorsally (bivium) and flattened ventrally (trivium). Bivium without 

characteristic wrinkles (as in the typical Holothuria scabra), sometimes covered by sediment when the animal 

is coming out of the bottom. Bivium with large papillae as well as black podia ending in a disc of around 

220 µm diameter; podia on trivium arranged irregularly, their calcareous disc around 350 µm in diameter. 

Mouth ventral, surrounded by a collar of papillae and 20 grey, short and stout tentacles. Anus 

terminal, surrounded by a circle of 5 groups of radial papillae. Calcareous ring with a large medio-ventral 

radial piece. Cuvierian tubules absent. Colour: variable; 3 main patterns can be recognized: speckled, with 

moderate black areas, or black overall. Spicules: very numerous; ventral tegument with tables and buttons, 

the tables having a moderately small disc, perforated by a various number of holes of variable size, the 

spire with 4 pillars and a cross-like bridge, ending in a spiny crown with a hole in the middle; knobby buttons 

generally have 6 holes, occasionally more; small rods and denticulate plates also present; tables and 

buttons in dorsal tegument: tables with a nodose disc, much larger than in ventral tegument; buttons 

variable, larger, and more perforated than those in ventral tegument; ventral and dorsal podia with long and 

large spinose rods; tentacles with long spiny rods, small smooth rods, and large denticulate plates. 



Habitat, biology, and fisheries: Generally inhabits shallow waters, but often found in depths of more than 

20 m (deeper than the typical H. scabra); on inner reef flats and coastal lagoons, frequently on flagstones; 

burrows in mud and sandy-muddy bottoms. Population densities are less than in the typical H. scabra, witha 

mean of around 0.01/m 2 . Sexual reproduction takes place during the warm season. It has a high potential 

fecundity. Harvested in some artisanal fisheries throughout the area, in places where its habitat occurs. 

Collected by hand at low tide while wading on the reefs. Processed with the same method used with the 

typical H. scabra: the sea cucumbers are buried overnight and the next day the numerous spicules are 

removed by brushing the tegument.The processed product is different in appearance from the typical H. scabra, 

having a golden colour. Of major commercial value, the processed product is at present highly demanded. 

Distribution: Widespread in the tropical Pacific, 

excluding coral reef islands. 

Remarks: Despite distinct differences in its 

ecology and biology, when compared to the typical 

H. scabra, this form is currently considered a 

variety, due to the lack of known differences in the 

spicules and internal anatomy. The name of this 

variety, versicolor, is due to the dorsal tegument 

which shows various colour patterns. Generally 

grows larger than the typical H. scabra.


Holothuria (Microthele) fuscogilva Cherbonnier, 1980 

Frequent synonyms / misidentifications: None / Holothuria nobilis (Selenka, 1867); H. maculata. 

FAO names: En - White teatfish; Fr - Holothurie blanche à mamelles. 

anus terminal, with 5 

bivium with slender papillae 

stout calcareous teeth 




tegument 


tegument 

mouth ventral, with papillae 

and 20 stout tentacles 



lateral teats 


Diagnostic characters: Body suboval, stout, firm and rigid, arched dorsally (bivium) and flattened ventrally 

(trivium). Bivium with characteristic large lateral papillae (teats) and often covered by sand. Bivium with 

small papillae as well as podia ending in a disc of around 450 µm diameter; podia on trivium stout, arranged 

irregularly, their calcareous disc around 600 µm in diameter. Mouth ventral, surrounded by a collar of long 

yellowish papillae and 20 grey, stout tentacles.Anus surrounded by 5 stout calcareous teeth.Calcareous 

ring with large radial pieces (slightly different from Holothuria nobilis). Cuvierian tubules absent. Colour: 

variable; bivium brown with more or less distinct whitish spots, becoming larger on sides; trivium lighter, 

generally whitish. Spicules: in form of tables and buttons; dorsal and ventral tegument with 2 kinds of tables; 

one form of table having an undulated disc with 10 to 15 irregular holes, supporting a massive spire with 4 pillars, 

ending in a very large, spiny, perforated crown; the other form of table, more frequent in the dorsal tegument, 

have a larger disc, with more holes and a spire ending in a crown composed of 2 or 3 rows of stout spines and 

5 or 6 spiny pillars; ventraltegument buttons simple, with 8 holes or more complicated as fenestrated ellipsoids; 

dorsal-tegument buttons only as fenestrated ellipsoids; ventral and dorsal podia with large plates, 

multiperforated; tentacles with spiny ending rods (up to 700 µm). 



Habitat, biology, and fisheries: With a deeper distribution than H. nobilis, mostly found in depths between 10 

and 40 m; generally occurs on outer barrier reefs and passes, but also known to inhabit shallow seagrass beds. 

Populations do not reach densities as high as H. nobilis, with medium densities around 0.001/m 2 .Sexual 

reproduction takes place during the warm season. A species with a low potential fecundity and late sexual 

maturity. Harvested in artisanal fisheries throughout the area, in places where its habitat occurs. Collected by 

skin diving or using diving gear (if not banned), making the populations vulnerable due to overexploitation. 

The processed product is of major commercial value and very highly demanded at present, even though 

the stocks have declined within the area. 

Distribution: Widespread in the tropical Indo-Pacific. 

Remarks: Although described only some years ago, 

this species, previously considered to be identical to 

H. nobilis. However, for a long time it was well known 

by fishermen from various Pacific islands, which give 

it a different name than H. nobilis, based on the colour. 

It differs from H. nobilis by the colour of the tegument, 

the absence of Cuvierian tubules, and the shape of 

spicules. Their habitats are also different.


Holothuria (Microthele) fuscopunctata Jaeger, 1833 

Frequent synonyms / misidentifications: Holothuria axiologa H.L. Clark, 1921 / None. 

FAO names: En - Elephant trunkfish; Fr - Holothurie trompe d’éléphant. 


tegument 




tegument 


with 20 brown 

tentacles 



bivium with brown wrinkles 

brown podia 


anus with 

papillae 

Diagnostic characters: Body suboval, stout, firm and rigid, arched dorsally (bivium) and flattened ventrally 

(trivium); bivium with characteristic brown wrinkles. Podia on bivium small, brown, withadiscof 

around 200 µm diameter; podia on trivium more densely distributed on the median radius and more 

sparsely distributed on the other radii and interradii, with their calcareous disc around 300 µm in diameter. 

Mouth ventral, with 20 brown, stout tentacles. Anus surrounded by 5 radial groups of 3 or 4 papillae 

and a black band. Calcareous ring with large radial pieces and narrower interradials. Cuvierian tubules 

absent. Cloaca large and black. Colour: bivium golden brown, with numerous brown spots; trivium 

whitish; mature gonads bright yellow. Spicules: in form of tables and buttons; tables of dorsal and ventral 

tegument similar in shape and moderately sparse, disc small, irregular, with 4 to 6 irregular holes, 

supporting a short spire, ending in an irregular crown of smooth spines; ventral-tegument buttons elongated 

ellipsoids, with 12 to 16 holes and median nodules; dorsal-tegument buttons generally more regular and 

less perforated; ventral and dorsal podia with large plates, multiperforated and indentated; tentacles 

containing both large very spiny rods and short nodose rods. 



Habitat, biology, and fisheries: Commonly found in reef slopes and lagoons, also seen in shallow 

seagrass beds; mostly in depths of less than 25 m. Population densities around 0.005/m 2 .Sexual 

reproduction takes place during the warm season. A species with a low potential fecundity and late sexual 

maturity. Occasionally harvested in artisanal fisheries throughout the area. Collected by skin diving or using 

diving gear (if not banned), 

making the populations very 

vulnerable, due to overexploitation. 

The processed product 

is of minor commercial value. 

Distribution: In the tropical 

Indian Ocean known only from 

Madagascar; in the tropical 

Pacific, in the west from Australia 

and Sulawesi eastwards to 

Marianna Islands, Palau, and 

New Caledonia.


Holothuria (Microthele) nobilis (Selenka, 1867) 

Frequent synonyms / misidentifications: Holothuria guamensis Quoy and Gaimard, 1833 / Holothuria 

fuscogilva Cherbonnier, 1980. 

FAO names: En - Black teatfish; Fr - Holothurie noire à mamelles. 



mouth ventral, with 20 papillae 

and 20 stout tentacles 

lateral teats 

black bivium 




tegument 


anus with 

5 stout 

calcareous 

teeth 


tegument 

Diagnostic characters: Body suboval, stout, firm and rigid, arched dorsally (bivium), and flattened 

ventrally (trivium). Bivium with characteristic large lateral papillae (teats) and often covered by sand. 

Bivium with small papillae as well as podia ending in a disc around 220 µm in diameter; podia on trivium 

stout, arranged irregularly, their calcareous disc around 700 µm in diameter. Mouth ventral, surrounded 

by 2 rows of black papillae and 20 grey, stout tentacles. Anus surrounded by short papillae and 5 

stout calcareous teeth. Calcareous ring with large wing-shaped radial pieces and narrow sharp 

interradials (slightly different from Holothuria fuscogilva). Cuvierian tubules present, but never expelled. 

Colour: less variable than in H. fuscogilva; bivium dark brown to black; trivium lighter, generally greyish. 

Spicules: in form of tables and buttons; only one kind of tables in the dorsal and ventral tegument; tables 

with undulated, circular disc with 12 to 16 holes, supporting a massive spire with 4 pillars and a cross 

bridge, ending in a large, spiny crown; ventral-tegument buttons in form of fenestrated ellipsoids; 

dorsal-tegument buttons shorter, larger, and less fenestrated; ventral podia with numerous spiny rods and 

large multiperforated plates; tentacles with spiny rods, large plates, small plates, and pseudo-plates. 



Habitat, biology, and fisheries: Inhabits shallower waters than H. fuscogilva, mostly found from the 

surface to a depth of 20 m; generally occurs on reef flats, slopes, and in shallow seagrass beds. Populations 

reach higher densities than H. fuscogilva, with medium densities of around 0.002/m 2 . Characterized by a 

sexual reproduction that takes place during the cold season. It has a medium potential fecundity. Harvested 

in artisanal fisheries throughout the area, in places were its habitat occurs. Collected by hand at low tide, 

by skin diving or using diving gear (if not banned), making the populations very vulnerable, due to 

overexploitation. The processed product is 

of major commercial value and very highly 

demanded at present, even though the 

stocks have declined within the area. 


Indo-Pacific. 

Remarks: Differs from H. fuscogilva by the 

colour of the tegument, the presence of 

Cuvierian tubules, and the shape of 

spicules. Their habitats are also different.


Pearsonothuria graeffei (Semper, 1868) 

Frequent synonyms / misidentifications: Bohadschia graeffei (Semper, 1868) / None. 

FAO names: En - Blackspotted sea cucumber. 



spicules of anal tegument 



spicules of dorsal and ventral tegument 

black spots 

white 

papillae on 

bivium 

mouth with 25 

large black 

tentacles 

Diagnostic characters: Body subcylindrical, arched dorsally (bivium) and slightly flattened ventrally 

(trivium). White conical papillae sparsely distributed on bivium; podia on trivium long and large, their 

calcareous disc around 240 µm in diameter. Mouth ventral, surrounded by 25 large, black tentacles. 

Anus nearly dorsal. Calcareous ring with irregular pieces. Cuvierian tubules numerous, but never expelled. 

Colour: bivium whitish with large, brown dots and numerous small black spots; trivium grey, also with 

small black spots. Spicules: dorsal and ventral tegument with pseudo-tables and rosettes; pseudo-tables 

consisting of large base with a large hole, and a spiny spire; rosettes numerous, those in the tegument 

simple, but more complicated in the dorsal papillae and ventral podia; also present around the anus are 

rods, small plates, and tables with a very spiny crown; tentacles containing a few rods and large plates 

derived from rosettes. 



Habitat, biology, and fisheries: A coral reef species, rarely found in depths of more than 25 m; mostly 

found on reef slopes, close to 

the coast; abundant on corals 

mixed with calcareous red 

algae. Population densities 

generally less than 0.005/m 2 . 

Not harvested for bêche-demer 

production. 


tropical Indo-Pacific, excluding 

the Persian Gulf and Hawaii. 

anus 



Order Aspidochirotida - Stichopodidae 1185 

Order Aspidochirotida - Stichopodidae STICHOPODIDAE 

Diagnostic characters: Body square-shaped or trapezoidal in cross-section. Cuvierian organs absent. 

Gonads forming 2 tufts appended on each side of the dorsal mesentery. Dominant spicules in form of 

branched rods and C-and S-shaped rods. 

Key to the genera of Stichopodidae occurring in the area (after Clark and Rowe, 1971) 

1a. Bivium covered with large papillae, leaf-shaped, simple or branched, and without podia 

regularly arranged longitudinally; spicules never developod as tables, but numerous 

grains, dichotomously branched rods . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelenota 

1b. Bivium covered with tubercules and papillae, at least on sides; trivium more or less 

covered by podia; spicules developod as tables, branched rods, and C-and S-shaped 

rods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stichopus 



Stichopus chloronotus Brandt, 1835 

Stichopus horrens Selenka, 1867 

Stichopus variegatus Semper, 1868 

Thelenota ananas (Jaeger, 1833) 

Thelenota anax Clark, 1921


Stichopus chloronotus Brandt, 1835 


FAO names: En - Greenfish; Fr - Trépang vert. 





with papillae and 

20 tentacles 


row of large papillae 


anus 

terminal 

Diagnostic characters: Body firm, rigid with quadrangular section, flattened ventrally (trivium); body wall 

easily disintegrates outside sea water. Radii of bivium with characteristic double row of large papillae, 

each radius ending in a small red or orange papilla. Trivium delimited by characteristic double row of large 

papillae; stout podia arranged regularly on 3 radial bands, with 10 rows in the medio-ventral band and 5 in 

the lateral. Mouth ventral, surrounded by a row of papillae and 20 green, stout tentacles. Anus 

terminal. Calcareous ring with large radial pieces and narrow interradials. Cuvierian tubules absent. Cloaca 

large and greenish. Colour: bivium dark green to black; trivium lighter. Spicules: presence of very 

characteristic tables with a narrow disc showing 4 holes, bearing a spire generally ending in a moderately 

spiny crown; tables with larger disc and simpler crown also present; presence of many C-shaped spicules; 

rosettes absent; tables in dorsal with have a large multiperforated disc, bearing a large spire; ventral papillae 

with long, smooth rods; ventral podia with denticulate rods and large multiperforated plates; tentacles with 

smooth and granular rods. 



Habitat, biology, and fisheries: A reef species, mostly found in shallow areas from near the surface to a 

depth of 15 m; generally occurs on reef flats and upper slopes. Populations reaching high densities on hard 

substrates, with a mean of around 0.04/m 2. Shallow-water populations are generally smaller in size than 

those found in deeper waters. 

Biology poorly known. 

Probably harvested in some 

artisanal fisheries of the area. 

Collected by hand at low tide, 

or by divers. The processed 

product is of low commercial 

value. 



the Persian Gulf and Hawaii.


Stichopus horrens Selenka, 1867 

Frequent synonyms / misidentifications: Stichopus godeffroyi Semper, 1868 / None. 

FAO names: En - Selenka’s sea cucumber. 





18 tentacles 

irregular warts 

spicules of podia spicules of papillae spicules of tegument 


anus 

terminal 

Diagnostic characters: Body firm, rigid, squarish in cross-section, flattened ventrally (trivium); body wall 

easily disintegrates outside sea water. Bivium covered with irregular warts, arranged in 10 longitudinal 

rows; warts larger near mouth. Trivium delimited by a characteristic double row of large papillae (4 to 5 

mm); stout podia arranged in 4 rows, on 3 radial bands, their disc about 350 µm in diameter. Mouth ventral, 

surrounded by a half row of papillae and 18 brown, short tentacles. Anus terminal. Calcareous ring 

with a deeply indented radial pieces and triangular interradials. Cuvierian tubules absent. Colour: bivium 

whitish to grey, with brown irregular dots; trivium lighter. Spicules: very characteristic rosettes, X-shaped, 

or elongate, numerous in the tegument; presence of numerous C-shaped spicules of 3 sizes; ventral 

tegument with tables of 2 sizes; some tables have a circular disc with 4 central and about 15 peripheral 

holes, bearing a spire with 4 pillars ending in a moderately spiny and cross-shaped crown; tables with a 

larger disc also present, more perforated, with a higher, spiny crown; only the first kind of table found in 

the dorsal tegument; tables in papillae provided with large multiperforated disc, with a long, conical, smooth 

spire, ending in a single point; ventral papillae containing long, smooth rods; ventral podia have long rods, 

with a central apophysis, and large, elongate, multiperforated plates; tentacles with large rods, either curved 

with few spines, or straight and very spiny. 



Habitat, biology, and fisheries: A reef species, mostly found in shallow areas from near the surface to a 

depth of 15 m; generally occurs in rubbles, or hidden in reef flats. A nocturnal species. Populations not 

reaching high densities, with a mean of around 0.007/m 2. Biology poorly known. Probably not harvested, 

as the tegument disintegrates too quickly. 

An aquaculture programme for this 

species is presently being carried out, for 

stock enhancement purposes. 


Indo-Pacific.


Stichopus variegatus Semper, 1868 

FAO names: En - Curryfish; Fr - Trépang curry. 



spicules of papillae 




20 tentacles 



conical warts 


papillae 

anus 

terminal 

Diagnostic characters: Body firm, rigid, squarish in cross-section, flattened ventrally (trivium). Body wall 

easily disintegrates outside sea water. Bivium covered with irregular conical warts, arranged in 8 

longitudinal rows, with smaller papillae in between. Trivium with yellow to pink podia, arranged in rows 

on the radii, their disc about 380 µm in diameter.Mouth ventral, surrounded by a circle of conical papillae 

and 20 yellowish tentacles. Anus terminal. Calcareous ring with a deeply indented radial pieces and 

small interradials. Cuvierian tubules absent. Colour: variable on bivium, yellow to greenish, with black 

spots; trivium lighter. Spicules: tables in the tegument with 2 forms of discs, some being undulated, with 

8 to 20 holes, while others form a denticulate disc with 4 central holes and a variable number of peripheral 

holes; spire of tables with 4 pillars ending in a moderately spiny and perforated crown; some tables with 

large disc and irregular crown also present; tables in papillae have a very large multiperforated disc; rosettes 

cross-shaped, branching; X-shaped spicules occur in 3 different sizes; ventral podia with spiny rods and 

large multiperforated plates showing pentagonal holes; tentacles with long, narrow, and spiny rods and X-, 

S-, and C-shaped small spicules. 

Size: Maximum length about 50 cm, commonly to about 35 cm; mean live weight about 1 kg (up to 2.5 kg); 


Habitat, biology, and fisheries: A shallow-water species, found in coastal reefs and lagoons, mostly from 

near the surface to a depth of 25 m. Generally occurs in seagrass beds, rubbles, and muddy-sand bottoms. 

Populations not reaching high densities, with a mean of around 0.005/m 2. Sexual reproduction takes place 

during the warm season. A 

species with a low potential 

fecundity and late sexual 

maturity. Probably rarely 

collected as the tegument 

disintegrates very easily, 

resulting in a low commercial 

value of the species. 



Hawaii.


Thelenota ananas (Jaeger, 1833) 


FAO names: En - Prickly redfish; Fr - Holothurie ananas. 



lobate papillae 

mouth ventral, with a circle of 

papillae and 20 large tentacles 



anus 

terminal 

trivium bordered by large papillae 


Diagnostic characters: Body firm, rigid, flattened ventrally (trivium). Bivium entirely covered with 

characteristic, large, leaf-shaped, lobate papillae. Trivium with brown to pink podia, more numerous on 

the radii, their disc about 400 µm in diameter. Mouth ventral, surrounded by a circle of conical papillae 

which are larger on dorsal side, and 20 large, brown tentacles. Anus terminal. Calcareous ring with large 

radial pieces and narrow interradials. Cuvierian tubules absent. Colour: variable on bivium, reddish orange 

to brown; trivium generally red; mature gonads deep purple. Spicules: tegument with cross-shaped 

spicules, spiny branched spicules (sometimes with median pillar), perforated plates, pseudo-tables, and 

grains; branched spicules larger in dorsal tegument and papillae; ventral podia with branched spicules, 

long rods, and large, multiperforated plates; tentacles with characteristic branched spicules, in form of a 

“rose-window”. 



Habitat, biology, and fisheries: A common reef species, mostly found from near the surface to a depth 

of 25 m; generally occurs on hard bottoms, large rubble and coral patches, on reef slopes and near passes. 

Populations not reaching very high densities, with a mean of around 0.003/m 2. Sexual reproduction takes place 

during the warm season. A species with a low potential fecundity and late sexual maturity. Symbiotic pearlfish 

(Carapidae, Ophidiiformes) are often found in its general cavity. Harvested by hand. Collected by skin diving or 

using diving gear (if not not 

banned), making the populations 

very vulnerable, due to 

overexploitation. The processed 

product is of major 

quality and the demand is still 

high. 



Hawaii.


Thelenota anax Clark, 1921 


FAO names: En - Amber fish. 


spicule of papillae 

anus with 

5 teeth 

spicules of tegument spicules of podia 


conical papillae 



with 18 brown 

tentacles 

large row of papillae between 

bivium and trivium 

Diagnostic characters: Body firm, rigid, squarish in cross-section, flattened ventrally (trivium). Bivium 

entirely covered with numerous characteristic, conical papillae and minute podia; bivium demarcated 

from the trivium by a row of large papillae. Podia numerous on trivium, with large disc, about 600 µm in 

diameter. Mouth ventral, surrounded by a circle of 18 large brown tentacles. Anus terminal. Calcareous 

ring with large radial pieces and narrow interradials. Cuvierian tubules absent. Cloaca large. Colour: bivium 

cream, with large beige dots; trivium generally beige; mature gonads deep purple. Spicules: tegument with 

branched spicules showing polygonal holes, and spicules in form of a “rose window”, mostly abundant in 

the tentacles; tentacles also with straight, curved, or X-shaped rods; ventral podia with short, smooth rods; 

dorsal papillae with sparse, very long, spiny rods. 



Habitat, biology, and fisheries: A rare reef species, mostly found at depths between 10 and 30 m; 

generally occurs on hard grounds, large rubbles and sand patches, on reef slopes, outer lagoon and near 

passes. Populations not reaching high densities, with a mean of around 0.001/m 2. Biology poorly known. 

Rarely harvested until few years ago, being generally found in low densities. Collected by skin diving or 

using diving gear (if not banned), making the populations presently very vulnerable, due to overexploitation. 

The processed product is probably of low to moderate commercial value and the exploitation of this species 

should be avoided. 


Indian Ocean known from the 

Glorieuses Islands; in the 

tropical Pacific, from northern 

Australia to Enewetok, Guam, 

the China Sea, and the Ryukyu 

Islands southwards to New 

Caledonia, Fiji, and the Society 

Islands. 



HAGFISHES 

by B. Fernholm and J.R. Paxton

1192 Hagfishes 

MYXINIDAE 

Hagfishes 

Diagnostic characters: Moderate-sized to large (to 100 cm), very elongate with eel-shaped body. 

Mouth with laterally biting horny teeth; no jaws. Eye reduced. Anterior single nostril surrounded 

by 4 tentacles. No operculum; 1 to 16 pairs of external gill openings. Two ventrolateral 

rows of slime glands. No paired fins; median fins without rays. No scales. Skeleton cartilaginous. 

Colour: pink to brown. 

Habitat, biology, and fisheries: Benthic fishes, often burrowing in mud, from inshore to deepsea. Feed 

as scavengers, mostly on dead or disabled fishes. Rare to common, most efficiently taken in baited traps, 

of recent commercial interest for skin (eelskin) industry based in Korea. 

Remarks: Six genera with about 60 species throughout the world’s oceans in tropical and temperate 

latitudes; tropical species occur in deep water. A revision of the family is needed; the best recent overview 

is that of Fernholm (1998). 


None. The laterally biting horny teeth, the separate external gill openings, and the absence of fins with rays 

are not found, either singly or in combination, in any other fish in the area. 


Eptatretus cirrhatus 

Eptatretus carlhubbsi McMillan and Wisner, 1984 

Eptatretus cirrhatus (Forster, 1801) 

Eptatretus strahani McMillan and Wisner, 1984 

Eptatretus spp. nov. (to be described by Fernholm from Papua New Guinea and the Philippines) 

Reference 

Fernholm, B. 1998. Hagfish systematics. In The biology of hagfishes, edited by J.M. Jorgensen et al. London, Chapman 

and Hall, pp. 33-44. 



General Remarks 

SHARKS

1194 Sharks 


snout 

snout 

nostril 

spiracle 

nostril 

prepectoralfin 

length 

mouth 

pectoral fin 

gill slits 

dorsal-fin spine 

(if present) 

trunk 

(distance in straight line) 

1 st dorsal fin 

vent 

anal fin 

preanal ridges 

pelvic fin 

(female, no claspers) 

ventral view 

spiracle 

head (dorsal view) 

precaudal tail 

interorbital 

space 

2 nd dorsal fin 

clasper 

(male sex 

organ) 

internasal 

distance 

precaudal 

pit 

keel 

labial 

furrows gill 

slits 

pelvic 

fin 

caudal 

peduncle 

anal 

fin 

pectoral fin 

preoral 

length 

head trunk 

tail 

total length 

(caudal fin depressed to body axis) 

eye 

diameter 

head (lateral view) 

interdorsal space 

preoral 

length 

caudal 

fin 

caudal fin 

mouth 

width 

head (ventral view) 

subterminal 

notch


apex 

upper (dorsal) lobe 

terminal lobe 

posterior tip 

spine 

posterior 

margin 

dorsal margin 

upper origin 

terminal 

margin 

subterminal margin 

subterminal notch 

anterior 

margin 

base 

free 

rear 

tip 

lower origin 

upper postventral margin 

posterior notch 

lower postventral margin 

fin origin insertion 

dorsal fin 

inner 

margin 

preventral margin 

ventral tip 

lower (ventral) lobe 

caudal fin 

fin 

origin 

fin insertion inner margin 

anterior 

margin 

base 

pectoral fin 

mouth corner 

labial furrow 

labial fold 

free rear tip 

apex 

posterior 

margin 

incurrent 

aperture 

anterior nasal flap lifted 

excurrent aperture 

circumnarial groove 

excurrent aperture 

nostril 

nasoral groove 

mouth 

circumnarial fold 

upper labial furrow 

barbel 

head of an orectoloboid shark 


incurrent 

aperture 

anterior 

nasal flap 

posterior 

nasal 

flap 

secondary 

lower eyelid 

symphyseal groove 

anterior nasal flap 

lower labial furrow 

upper eyelid 

eye 

notch 

nictitating 

lower 

eyelid 

subocular pocket

1196 Sharks 


by L.J.V. Compagno 

Sharks include a variety of usually cylindrical, elongated, or moderately depressed fishes which differ 

from the closely related rays or batoids in having lateral gill openings (or gill slits) and pectoral fins not 

fused to the sides of the head over the gill openings. The greatly depressed angelsharks (family Squatinidae) 

might be mistaken for rays at first sight; they have large, broad, ray-like pectoraI fins that extend as 

triangular lobes alongside the gill openings, but are not connected to the head above them. Sharks have 

eyes on the dorsal surface or sides of the head and spiracles (when present) on its dorsal or dorsolateral 

surfaces. The tail and caudal fin are always well developed and serve to propel the animal by lateral 

undulations; the pectoral fins are mostly not used for propulsion through the water but aid in stabilizing and 

steering the shark. There are usually 5 gill openings on each side of the head, rarely 6 or 7. The mouth is 

usually ventral or subterminal on the head, but terminal or nearly so in a few species. Most sharks have 2 

(rarely 1) dorsal fins, sometimes with spines on their front edges; an anal fin is usually present, but missing 

in several families. The teeth on the jaws are set in numerous transverse rows and are constantly replaced 

from inside the mouth.All shark species are more or less covered by small (occasionally enlarged) tooth-like 

placoid scales or dermal denticles. 

Male sharks have cylindrical copulatory organs or claspers on their pelvic fins, used for internal fertilization 

of eggs in females; about 1/3 of the species of sharks have females that deposit eggs in rectangular or 

conical capsules, formed of a horn-like material (oviparity); the remainder are livebearers. Some livebearing 

sharks, including many houndsharks (Triakidae), most requiem sharks (Carcharhinidae), and all weasel 

sharks (Hemigaleidae) and hammerheads (Sphyrnidae) are viviparous (placental viviparous), with yolk 

sacs of fetuses forming a placenta with the maternal uterus for nutrient transfer; other livebearinq sharks 

are ovoviviparous (aplacental viviparous), without a placenta. Ovoviviparous lamnoid sharks of the families 

Odontaspididae, Alopiidae, Lamnidae, and Pseudocarchariidae practice uterine cannibalism, in which one 

or more fetuses in each uterus resorb their yolk sacs and then devour eggs passed down the oviducts for 

nutriment (oophagy) and grow to considerable size with massive yolk stomachs before birth. In the 

Odontaspididae (Carcharias taurus) the largest fetus kills and eats its siblings (adelphophagy) and only 1 

fetus survives in utero, while several young may cohabit the uterus in the other families. Members of 2 

families of carcharhinoid sharks (Proscylliidae and Pseudotriakidae) practice oophagy, but fill their yolk 

sacs with yolk that they consume. Mature sharks vary in total length from about 15 to 19 cm (dwarf species 

of Squalidae and Proscylliidae) to 12.1 m or more (whale shark, family Rhincodontidae) and range in weight 

from between 10 and 20 g to several metric tons. Most sharks are of small or moderate size; about 50% 

are small, between 15 cm and 1 m; 32% between 1 and 2 m; 14% between 2 and 4 m; and only 4% are 

over 4 m in total length. 

All sharks are predators, with their prey ranging widely, from planktonic crustaceans and benthic invertebrates 

to pelagic cephalopods, small to large bony fishes, other sharks and rays, marine mammals, and 

other marine and terrestrial vertebrates. Sharks are primarily marine, but a few requiem sharks (Carcharhinidae, 

members of the genera Carcharhinus and Glyphis) have broad salinity tolerances, and one 

species (bull shark, Carcharhinus leucas) is wide-ranging in tropical lakes and rivers with sea access as 

well as shallow inshore waters. No sharks are known to be confined to fresh water, unlike several species 

of stingrays (families Dasyatidae and Potamotrygonidae). Sharks are widely distributed in all oceans, from 

the Arctic to subantarctic islands, and from close inshore on reefs, off beaches, and in shallow, enclosed 

bays to the lower continental slopes, the abyssal plains, sea mounts and ridges, and the high seas. They 

are most diverse in continental waters of tropical and warm-temperate seas, from inshore waters down to 

upper continental slopes, but are less so in colder waters, at great depths (below 1 500 to 2 000 m), in the 

open ocean and off oceanic islands. The richest shark faunas occur in the Indo-West Pacific from South 

Africa and the Red Sea to Australia and Japan. The Western Central Pacific (Fishing Area 71 and the 

southwestern part of Fishing Area 77) has one of the most diverse shark faunas in the world, including 

approximately 23 families, 69 genera, and between 164 and 188 species. Worldwide there are 33 families, 

101 genera, and between 379 to 478 species of sharks (estimate as of 8 August 1995). Several genera 

and families are poorly known and require further taxonomic study. Many species of sharks are endemic 

to the area and have restricted ranges within it, several species (including inshore species) are known from 

1 or a few museum specimens only, and a wealth of new species have been revealed in deep water, offshore 

continental, and even inshore habitats in the past forty years (many of which are still undescribed). 

Undoubtedly more new species and many records of described species will be discovered with further 

collecting in poorly known parts of the area. Knowledge of the coastal shark fauna of Area 71 beyond 

Australia is very sketchy, and many maritime countries need further surveys to determine which species 

occur there. The deep-water shark fauna is very poorly known in the area, except for off northern Australia 

and a few other localities (such as New Caledonia) where systematic deep-water exploration for fisheries 

resources is proceeding apace. Basic knowledge of the biology of many species is often very deficient or 

entirely lacking, and can be supplemented by new information gathered by fisheries workers in the area.


The shark attack hazard has been grossly exaggerated in recent years. Large carcharhinids, sphyrnids 

and lamnids, and less frequently other sharks, pose a potential threat to people in the water or boats. Large 

gill nets have been regularly set in the vicinity of popular bathing beaches off Queensland, Australia during 

the past 3 decades to reduce the number of potentially dangerous sharks. This ‘shark meshing’ has 

presumably reduced shark attacks there although few attacks were recorded off Queensland prior to the 

onset of meshing (unlike New South Wales, where the practice originated, and off South Africa). About 9% 

of known shark species are definitely known to be dangerous (that is, are known to have been implicated 

in at least 1 shark attack worldwide), and about 10% more are large enough and sufficiently well-armed to 

be potentially so; the rest are mostly too small and poorly armed to be a hazard to people. ‘Dangerous’ is 

highly relative; perhaps less than 100 shark attacks (and less than 20 fatalities) occur worldwide each year. 

Sharks are not very dangerous compared to any number of other causes of death or injury to people, 

including drownings and near-drownings and large terrestrial predators. The 3 shark species most 

frequently implicated in shark attacks (white shark, tiger shark, and bull shark) do not automatically attack 

when confronted by people in the water. Great white sharks usually do not attack in such situations. And 

if biting does occur it is usually restricted to single bites delivered with minimal force. ‘Man-eating’ does not 

appear to be an important source of nutrients for any shark. Unfortunately, the shark attack issue has tended 

to obscure the ‘human attack’ problem and its implications for shark conservation in the face of burgeoning 

fisheries driven by the expanding world human population and enormous markets for shark products. It 

was recognized over the past 4 decades that aspects of the life history strategy of sharks (long lives, long 

maturation times, and low fecundity, plus relatively large size) made them very vulnerable to overexploitation, 

and that several targeted shark fisheries had suddenly collapsed after recruitment had been impaired 

by overexploitation of the breeding stocks. However, only in the past 5 years has there been widespread 

concern about world trends in fisheries for sharks and other cartilaginous fishes. After the second world 

war world fisheries for chondrichthyan fishes essentially tripled in reported catches to FAO, which has not 

kept pace with the approximately fourfold increases in total fisheries worldwide. Much of the catch is as 

bycatch in fisheries driven by larger catches of exploitation-resistant bony fishes with far higher fecundity. 

World catches of cartilaginous fishes reported to FAO have leveled off in the 5-year period 1988 to 1992 

to about 690 thousand metric tonnes, which may indicate that there is little scope for further increases in 

catches. Some sharks have been accorded limited or total protection in a number of developed countries, 

yet on a world basis shark exploitation is mostly unregulated and out of control nationally and regionally. 

In the next decade international agreements, including CITES listings, will likely occur to protect a variety 

of sharks and other cartilaginous fishes from excessive exploitation. 

In the Western Central Pacific, sharks are used mainly for human food; shark meat is marketed fresh, 

frozen, and especially dried-salted. Sharks are utilized on the oriental market for fins; also for liver oil, 

fishmeal, and possibly for leather, although details of utilization in the area are sketchy. The total catch of 

sharks reported from Fishing Area 71 is uncertain; total catches of cartilaginous fishes in the area was 

approximately 119 000 t in 1995, of which about 59 000 t were reported as rays (batoids), 52 000 t mixed 

sharks and rays and about 8 000 t were sharks. If the mixed sharks and rays included 55% sharks the 1995 

shark catch is roughly 37 000 t; the actual landings of sharks in the area are doubtlessly much higher. 

Catches in the section of Area 77 included in this work were relatively small and may add roughly 6 000 t 

of chondrichthyans to the 1995 total. The present area had the second highest catches of cartilaginous 

fishes worldwide in 1995, being surpassed only by FAO Area 51 (Western Indian Ocean, with 145 000 t). 

The present area includes Indonesia, which in 1995 had the second highest cartilaginous fish catch of any 

nation (75 000 t, compared to India with 86 000 t), the next highest countries being Pakistan, Taiwan 

Province of China, and the USA. Malaysia had a catch of about 19 000 t, Thailand and Philippines had 

catches of about 9 000 t each, and the Korean Republic took about 10 000 t in the area in 1995. Data on 

gear used in the area is sketchy, but line gear (including pelagic longlines), fixed and floating gill nets, 

bottom trawls, fixed fish traps, and purse seines are used to target sharks or take sharks as a bycatch. 

Sharks are taken in artisanal fisheries, by local inshore and offshore commercial fisheries, and by large 

fishing fleets in offshore waters. Requiem sharks (Carcharhinidae) are especially important, but considerable 

numbers of threshers (Alopiidae) and makos (Lamnidae, genus Isurus are fished offshore, and a 

number of other families, including longtailed carpetsharks (Hemiscylliidae), zebra sharks (Stegostomatidae), 

nurse sharks (Ginglymostomatidae), weasel sharks (Hemigaleidae), and hammerheads (Sphyrnidae) 

are commonly taken in inshore fisheries. Dogfish (family Squalidae) are important in offshore deep-set 

longline fisheries targeting sharks for liver oil.

1198 Sharks 

Key to Families KEY TO FAMILIES OCCURRING IN THE AREA 

1a. No anal fin (Figs 1 to 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Anal fin present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

2a. Body strongly depressed and ray-like; pectoral fins greatly enlarged, with anterior 

triangular lobes that overlap gill slits; mouth terminal (Fig. 1) . . . . . . . . . Squatinidae (p. 1235) 

2b. Body cylindrical, compressed, or slightly depressed, not ray-like; pectoral fins small, 

without anterior lobes; mouth ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

pectoralfin 

lobe 

mouth 

terminal 

ventral view dorsal view 

Fig. 1 Squatinidae Fig. 2 Pristiophoridae 

3a. Snout flattened and elongated, saw-like (Fig. 2) . . . . . . . . . . . . . Pristiophoridae (p. 1233) 

3b. Snout normal, not saw-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

4a. First dorsal fin behind pelvic-fin origins; dermal denticles moderately large or very large, 

thorn-like (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Echinorhinidae (p. 1211) 

4b. First dorsal fin partially or entirely in front of pelvic-fin origins (Fig. 4); dermal denticles 

small to moderately large, variable in shape . . . . . . . . . . . . . . . . . . . Squalidae (p.1213) 

dermal denticles 

Fig. 3 Echinorhinidae Fig. 4 Squalidae 

5a. Only 1 dorsal fin, far posterior on back; 6 or 7 gill slits on each side (Fig. 5) . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hexanchidae (p. 1208) 

5b. Two dorsal fins (except the scyliorhinid Pentanchus profundicolus with 1 dorsal fin); 5 

gill slits on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

6a. A strong spine on each dorsal fin (Fig. 6) . . . . . . . . . . . . . . . . . Heterodontidae (p. 1238) 

6b. Dorsal fins without spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

6-7 slits 

Fig. 5 Hexanchidae 

1 dorsal fin 

spine 

spine 

Fig. 6 Heterodontidae

Key to Families 1199 

7a. Head with lateral expansions or blades, like a double-edged axe (Fig. 7) . . . Sphyrnidae (p. 1361) 

7b. Head normal, not expanded laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

8a. Eyes behind mouth; deep nasoral grooves connecting nostrils and mouth (Fig. 8a) . . . . . . . → 9 

8b. Eyes partly or entirely over mouth; nasoral grooves usually absent (Fig. 8b), when 

present (Atelomycterus in family Scyliorhinidae) broad and shallow . . . . . . . . . . . . . . → 15 

enlarged 

underside 

of head Fig. 7 Sphyrnidae 

nostrils 

a) Ginglymostoma sp. b) Carcharhinus sp. 

Fig. 8 underside of head 

9a. Mouth huge and nearly terminal; external gill slits very large, internal gill slits inside 

mouth cavity with filter screens; caudal peduncle with strong lateral keels; caudal fin 

with a strong ventral lobe, but without a strong terminal lobe and subterminal notch 

(Fig. 9) . . . . . . . . . . . . . . . . . . . . . . . Rhincodontidae (= Rhiniodontidae) (p. 1263) 

9b. Mouth smaller and subterminal; external gill slits small, internal gill slits without filter 

screens; caudal peduncle without strong lateral keels; caudal fin with a weak ventral 

lobe or none, but with a strong terminal lobe and subterminal notch (Fig. 10) . . . . . . . . . → 10 

ridges 

Fig. 9 Rhincodontidae 

10a. Caudal fin about as long as rest of shark (Fig. 11) . . . . . . . . . . . . Stegostomatidae (p. 1262) 

10b. Caudal fin much shorter than rest of shark . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11 

11a. Head and body greatly flattened, head with skin flaps on sides; 2 rows of large, fang-like 

teeth at symphysis of upper jaw and 3 in lower jaw (Fig. 12) . . . . . . . . Orectolobidae (p. 1245) 

11b. Head and body cylindrical or moderately flattened, head without skin flaps; teeth small, 

not enlarged and fang-like at symphysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 12 

nostril 

labial 

folds 

barbels 

level of eyes 

caudal fin 

Fig. 10 

nasoral 

groove 

Fig. 11 Stegostomatidae Fig. 12 Orectolobidae 

strong terminal lobe 

subterminal 

notch

1200 Sharks 

12a. No lobe and groove around outer edges of nostrils (Fig. 13) . . . . Ginglymostomatidae (p. 1260) 

12b. A lobe and groove around outer edges of nostrils (Fig. 14) . . . . . . . . . . . . . . . . . . . → 13 

Fig. 13 Ginglymostomatidae 

13a. Spiracles minute; origin of anal fin 

well in front of second dorsal-fin 

origin, separated from lower caudal-fin 

origin by space equal or 

greater than its base length 

(Fig. 15) . . . . . . . Parascylliidae (p. 1241) 

13b. Spiracles large; origin of anal fin 

well behind second dorsal-fin origin, 

separated from lower caudalfin 

origin by space less than its 

base length . . . . . . . . . . . . . . . . → 14 

Fig. 14 

Fig. 15 Parascylliidae 

lobe and 

groove 

14a. Nasal barbels very large; anal fin high and angular; distance from vent to lower 

caudal-fin origin shorter than distance from snout to vent (Fig. 16) . . . . Brachaeluridae (p. 1243) 

14b. Nasal barbels short; anal fin low, rounded and keel-like; distance from vent to lower 

caudal-fin origin longer than distance from snout to vent (Fig. 17) . . . . . Hemiscylliidae (p. 1249) 

Fig. 16 Brachaeluridae Fig. 17 Hemiscylliidae 

15a. A strong keel present on each side of caudal peduncle; caudal fin crescentic and nearly 

symmetrical, with a long lower lobe (Fig. 18) . . . . . . . . . . . . . . . . . . Lamnidae (p. 1274) 

15b. No keels on caudal peduncle, or weak ones (Pseudocarcharias in Pseudocarchariidae, 

Galeocerdo and Prionace in Carcharhinidae); caudal fin asymmetrical, not crescentic, 

with ventral lobe relatively short or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . → 16 

16a. Caudal fin about as long as rest of shark (Fig. 19) . . . . . . . . . . . . . . . . Alopiidae (p. 1269) 

16b. Caudal fin less than half the length of rest of shark . . . . . . . . . . . . . . . . . . . . . . → 17 

keel 

Fig. 18 Lamnidae Fig. 19 Alopiidae

Key to Families 1201 

17a. No nictitating eyelids (Fig. 20a), largest teeth in mouth are 2 or 3 rows of anteriors on 

either side of lower jaw symphysis; upper anteriors separated from large lateral teeth 

at sides of jaw by a gap that may have 1 or more rows of small intermediate teeth 

(Fig. 20b); all gill slits in front of pectoral fins (Figs 22 and 23) . . . . . . . . . . . . . . . . . → 18 

17b. Nictitating eyelids present (Fig. 21a); largest teeth in mouth are well lateral on dental 

band, not on either side of symphysis; no gap or intermediate teeth separating large 

anterior teeth from still larger teeth in upper jaw (Fig. 21b); last 1 or 2 gill slits over 

pectoral-fin bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 19 

no nictitating 

eyelid 

a) eye level of 

symphysis 

intermediate 

anterior 

gap 

lateral 

b) upper and lower teeth of one side 

Fig. 20 

nictitating 

eyelid 

a) eyes 

level of 

symphysis 

anterior 

18a. Eyes very large; gill slits extending onto upper surface of head; both upper and lower 

precaudal pits present; a low keel on each side of caudal peduncle (Fig. 22) . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudocarchariidae (p. 1268) 

18b. Eyes smaller; gill slits not extending onto upper surface of head; lower precaudal pit 

absent; no keels on caudal peduncle (Fig. 23) . . . . . . . . . . . . . . Odontaspididae (p. 1264) 

gill slits high 

keel 

eye large lower precaudal pit 

Fig. 22 Pseudocarchariidae 

gill slits lower 

eye smaller 

19a. Origin of first dorsal fin over or 

behind pelvic-fin bases (Fig. 24) 

. . . . . . . . . . . Scyliorhinidae (p. 1279) 

19b. Origin of first dorsal fin well ahead 

of pelvic-fin bases . . . . . . . . . . . . → 20 

Fig. 24 Scyliorhinidae 

20a. No precaudal pits, dorsal caudal-fin margin smooth (Fig. 25) . . . . . . . . . . . . . . . . . . → 21 

20b. Precaudal pits and rippled dorsal caudal margin present (ripples sometimes irregular 

in Scoliodon and Triaenodon of family Carcharhinidae) (Fig. 26) . . . . . . . . . . . . . . . . → 23 

margin smooth 

margin rippled 

no precaudal 

precaudal pits 

pits 

Fig. 25 

no gap 

lateral 

b) upper and lower teeth of one 

side far back 

Fig. 21 

Fig. 23 Odontaspididae 

Fig. 26

1202 Sharks 

21a. First dorsal fin long, about the 

length of caudal fin, and formed as 

a low, rounded keel; adults with 

over 200 rows of teeth in each jaw; 

spiracles nearly or quite as long as 

eyes (Fig. 27) . . . Pseudotriakidae (p. 1296) 

21b. First dorsal fin short, about 2/3 of 

caudal fin or less; subtriangular in 

shape; adults with less than 110 

rows of teeth in each jaw; spiracles 

much smaller than eyes . . . . . . . . . → 22 

spiracles 

very large 

22a. Labial furrows very short or absent, confined to extreme mouth corners; posterior teeth 

comb-like; base of first dorsal fin closer to pelvic-fin bases than to pectoral-fin bases 

(Fig. 28) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proscylliidae (p. 1293) 

22b. Labial furrows longer, extending anteriorly for a greater or lesser distance on lips; 

posterior teeth not comb-like; base of first dorsal fin either equidistant between pectoral 

and pelvic-fin bases or closer to pectoral-fin bases (Fig. 29) . . . . . . . . . . Triakidae (p. 1297) 

ventral view 

of head 

labial furrow 

short or absent 

Fig. 28 Proscylliidae 

ventral view 

of head 

23a. Intestine with a spiral valve (Fig. 30a) having 4 to 6 turns 

. . . . . . . . . . . . . . . . . . . . . . . Hemigaleidae (p. 1305) 

23b. Intestine with a scroll valve (Figs 30b and 32) 

. . . . . . . . . . . . . . . . . . . . . . Carcharhinidae (p. 1312) 

Fig. 31 Hemigaleidae 

Fig. 32 Carcharhinidae 

low, keel-like 

Fig. 27 Pseudotriakidae 

labial 

furrow 

larger 

Fig. 29 Triakidae 

a) spiral valve 

rolled 

unrolled 

b) scroll valve 

Fig. 30


List of Families and LIST Species OF FAMILIES AND SPECIES OCCURRING IN THE AREA 

The symbol is given when species accounts are included. A question mark indicates that presence in 

the area is uncertain. 

HEXANCHIDAE: Sixgill and sevengill sharks, cow sharks 

Heptranchias perlo (Bonnaterre, 1788) 

Hexanchus griseus (Bonnaterre, 1788) 

Hexanchus nakamurai Teng, 1962 

ECHINORHINIDAE: Bramble sharks 

Echinorhinus brucus (Bonnaterre, 1788) 

Echinorhinus cookei Pietschmann, 1928 

SQUALIDAE: Dogfish sharks 

Centrophorus atromarginatus Garman, 1906 

Centrophorus granulosus (Bloch and Schneider, 1801) 

? Centrophorus isodon (Chu, Meng, and Liu, 1981) 

? Centrophorus lusitanicus Bocage and Capello, 1864 

Centrophorus moluccensis Bleeker, 1860 

Centrophorus niaukang Teng, 1959 

Centrophorus squamosus (Bonnaterre, 1788) 

Centrophorus sp. [New Caledonia] 

Centroscyllium cf. kamoharai Abe, 1966 

Centroscymnus coelolepis Bocage and Capello, 1864 

Cirrhigaleus barbifer Tanaka, 1912 

Dalatias licha (Bonnaterre, 1788) 

? Deania calcea (Lowe, 1839) 

Deania profundorum (Smith and Radcliffe, 1912) 

Deania quadrispinosa (McCulloch, 1915) 

? Etmopterus baxteri Garrick, 1957 

Etmopterus brachyurus Smith and Radcliffe, 1912 

? Etmopterus decacuspidatus Chan, 1966 

? Etmopterus granulosus (Günther, 1880) 

Etmopterus lucifer Jordan and Snyder, 1902 

Etmopterus molleri Whitley, 1939 

? Etmopterus princeps Collett, 1904 

Etmopterus splendidus Yano, 1988 

? Etmopterus unicolor (Engelhardt, 1912) 

? Etmopterus sp. C [Last and Stevens, 1994] 

Etmopterus sp. D [Last and Stevens, 1994] 

Etmopterus sp. F [Last and Stevens, 1994] 

Euprotomicrus bispinatus (Quoy and Gaimard, 1824) 

Isistius brasiliensis (Quoy and Gaimard, 1824) 

Scymnodon squamulosus (Günther, 1877) 

? Somniosus pacificus Bigelow and Schroeder, 1944 

Squaliolus aliae Teng, 1959 

Squaliolus laticaudus Smith and Radcliffe, 1912 

Squalus japonicus Ishikawa, 1908 

Squalus megalops (Macleay, 1881) 

Squalus melanurus Fourmanoir and Rivaton, 1979 

Squalus mitsukurii Jordan and Snyder, 1903 

Squalus rancureli Fourmanoir and Rivanton, 1979 

Squalus sp. A [Last and Stevens, 1994] 

Squalus sp. B [Last and Stevens, 1994] 

Squalus sp. F [Last and Stevens, 1994]

1204 Sharks 

PRISTIOPHORIDAE: Sawsharks 

Pristiophorus sp. B [Last and Stevens, 1994] (Australia) 

Pristiophorus sp. (Philippines) 

SQUATINIDAE: Angelsharks 

Squatina australis Regan, 1906 

Squatina japonica Bleeker, 1858 

Squatina sp. A [Last and Stevens, 1994] 

HETERODONTIDAE: Bullhead sharks 

Heterodontus galeatus (Günther, 1870) 

Heterodontus portusjacksoni (Meyer, 1793) 

Heterodontus zebra (Gray, 1831) 

PARASCYLLIIDAE: Collared carpetsharks 

Cirrhoscyllium expolitum Smith and Radcliffe, 1913 

Parascyllium collare Ramsay and Ogilby, 1888 

BRACHAELURIDAE: Blind sharks 

Brachaelurus waddi (Bloch and Schneider, 1801) 

Heteroscyllium colcloughi (Ogilby, 1908) 

ORECTOLOBIDAE: Wobbegongs 

Eucrossorhinus dasypogon (Bleeker, 1867) 

Orectolobus japonicus Regan, 1906 

Orectolobus maculatus (Bonnaterre, 1788) 

Orectolobus ornatus (de Vis, 1883) 

Orectolobus wardi Whitley, 1939 

HEMISCYLLIIDAE: Longtail carpetsharks 

Chiloscyllium griseum Müller and Henle, 1839 

Chiloscyllium hasselti Bleeker, 1852 

Chiloscyllium indicum (Gmelin, 1789) 

Chiloscyllium plagiosum (Bennett, 1830) 

Chiloscyllium punctatum Müller and Henle, 1838 

Hemiscyllium freycineti (Quoy and Gaimard, 1824) 

Hemiscyllium hallstromi Whitley, 1967 

Hemiscyllium ocellatum (Bonnaterre, 1788) 

Hemiscyllium strahani Whitley, 1967 

Hemiscyllium trispeculare Richardson, 1845 

GINGLYMOSTOMATIDAE: Nurse sharks 

Nebrius ferrugineus (Lesson, 1830) 

STEGOSTOMATIDAE: Zebra sharks 

Stegostoma fasciatum (Hermann, 1783) 

RHINCODONTIDAE: Whale sharks 

Rhincodon typus Smith, 1828 

ODONTASPIDIDAE: Sand tiger sharks 

Carcharias taurus Rafinesque, 1810 

? Odontaspis ferox (Risso, 1810) 

PSEUDOCARCHARIIDAE: Crocodile sharks 

Pseudocarcharias kamoharai (Matsubara, 1936) 

ALOPIIDAE: Thresher sharks 

Alopias pelagicus Nakamura, 1935 

Alopias superciliosus (Lowe, 1839) 

Alopias vulpinus (Bonnaterra, 1788)


LAMNIDAE: Mackerel sharks 

Carcharodon carcharias (Linnaeus, 1758) 

Isurus oxyrinchus Rafinesque, 1810 

Isurus paucus Guitart Manday, 1966 

SCYLIORHINIDAE: Catsharks 

? Apristurus acanutus Chu, Meng, and Li in Meng, Chu, and Li, 1985 

? Apristurus gibbosus Meng, Chu, and Li, 1985 

Apristurus herklotsi (Fowler, 1934) 

Apristurus longicephalus Nakaya, 1975 

? Apristurus macrostomus Meng, Chu, and Li, 1985 

? Apristurus micropterygeus Meng, Chu, and Li in Chu, Meng, and Li, 1986 

Apristurus sibogae (Weber, 1913) 

? Apristurus sinensis Chu and Hu in Chu, Meng, Hu, and Li, 1981 

Apristurus spongiceps (Gilbert, 1895) 

Apristurus verweyi (Fowler, 1934) 

Apristurus sp. A [Last and Stevens, 1994] 

Apristurus sp. B [Last and Stevens, 1994] 

Apristurus sp. G [Last and Stevens, 1994] 

Apristurus sp. [Seret] (New Caledonia) 

Apristurus sp. [Seret] (Philippines) 

Apristurus sp. [Seret] (Indonesia) 

Asymbolus sp. E [Last and Stevens, 1994] 

Asymbolus sp. [Seret] (New Caledonia) 

Atelomycterus fasciatus Compagno and Stevens, 1993 

Atelomycterus macleayi Whitley, 1939 

Atelomycterus marmoratus (Bennett, 1830) 

Aulohalaelurus kanakorum Seret, 1990 

Cephaloscyllium fasciatum Chan, 1966 

Cephaloscyllium sp. [Compagno, 1984, 1988] 

Cephaloscyllium sp. [J.Randall, pers. comm. 1994] (Papua New Guinea) 

Cephaloscyllium sp. [Seret] (New Caledonia) 

Cephaloscyllium sp. B [Last and Stevens, 1994] 

Cephaloscyllium sp. C [Last and Stevens, 1994] 

Cephaloscyllium sp. D [Last and Stevens, 1994] 

Cephaloscyllium sp. E [Last and Stevens, 1994] 

Galeus boardmani (Whitley, 1928) 

Galeus eastmani (Jordan and Snyder, 1904) 

Galeus gracilis Compagno and Stevens, 1993 

Galeus sauteri (Jordan and Richardson, 1909) 

Galeus schultzi Springer, 1979 

Galeus sp. B. [Last and Stevens, 1994] 

? Halaelurus immaculatus Chu and Meng, 1982 

Halaelurus boesemani Springer and D’Aubrey, 1972 

? Halaelurus buergeri (Müller and Henle, 1838) 

Parmaturus melanobranchius (Chan, 1966) 

Parmaturus sp. A [Last and Stevens, 1994] 

? Parmaturus sp. [Seret] (Indonesia) 

Pentanchus profundicolus Smith and Radcliffe, 1912 

Scyliorhinus garmani (Fowler, 1934) 

Scyliorhinus torazame (Tanaka, 1908) 

PROSCYLLIIDAE: Finback catsharks 

Eridacnis radcliffei Smith, 1913 

Gollum attenuatus (Garrick, 1954) 

Proscyllium habereri Hilgendorf, 1904

1206 Sharks 

PSEUDOTRIAKIDAE: False catsharks 

Pseudotriakis microdon Capello, 1968 

TRIAKIDAE: Houndsharks 

Galeorhinus galeus (Linnaeus, 1758) 

Gogolia filewoodi Compagno, 1973 

Hemitriakis abdita Compagno and Stevens, 1993 

? Hemitriakis japanica (Müller and Henle, 1839) 

Hemitriakis leucoperiptera Herre, 1923 

Hemitriakis sp. [Compagno, 1988] (Philippines) 

Hypogaleus hyugaensis (Miyosi, 1939) 

Iago garricki Fourmanoir, 1979 

Mustelus antarcticus Günther, 1870 

Mustelus griseus Pitschmann, 1908 

Mustelus manazo Bleeker, 1854 

Mustelus cf. manazo [Seret, pers. comm. 1994] 

Mustelus sp. A [Last and Stevens, 1994] 

Mustelus sp. B [Last and Stevens, 1994] 

? Triakis scyllium Müller and Henle, 1839 

HEMIGALEIDAE: Weasel sharks 

Chaenogaleus macrostoma (Bleeker, 1852) 

Hemigaleus microstoma Bleeker, 1852 

Hemigaleus sp. aff. “microstoma” 

Hemipristis elongata (Klunzinger, 1871) 

Paragaleus tengi (Chen, 1963) 

CARCHARHINIDAE: Requiem sharks 

Carcharhinus albimarginatus (Rüppell, 1837) 

Carcharhinus altimus (Springer, 1950) 

Carcharhinus amblyrhynchos (Bleeker, 1856) 

Carcharhinus amboinensis (Müller and Henle, 1839) 

Carcharhinus borneensis (Bleeker, 1859) 

Carcharhinus brachyurus (Günther, 1870) 

Carcharhinus brevipinna (Müller and Henle, 1839) 

Carcharhinus cautus (Whitley, 1945) 

Carcharhinus dussumieri (Valenciennes in Müller and Henle, 1839) 

Carcharhinus falciformis (Bibron in Müller and Henle, 1839) 

Carcharhinus fitzroyensis (Whitley, 1943) 

Carcharhinus galapagensis (Snodgrass and Heller, 1905) 

Carcharhinus hemiodon (Valenciennes in Müller and Henle, 1839) 

Carcharhinus leucas (Valenciennes in Müller and Henle, 1839) 

Carcharhinus limbatus (Valenciennes in Müller and Henle, 1839) 

Carcharhinus longimanus Poey, 1861) 

Carcharhinus melanopterus (Quoy and Gaimard, 1824) 

Carcharhinus obscurus (LeSueur, 1818) 

Carcharhinus plumbeus (Nardo, 1827) 

Carcharhinus sealei (Pietschmann, 1916) 

Carcharhinus sorrah (Valenciennes in Müller and Henle, 1839) 

Carcharhinus tilstoni (Whitley, 1950) 

Carcharhinus sp. (= “Carcharhinus porosus”) 

Galeocerdo cuvier (Peron and LeSueur in LeSueur, 1822) 

Glyphis sp. A [Last and Stevens, 1994] (Queensland) 

Glyphis sp. B [Compagno] (Borneo) 

Glyphis sp. C [Compagno] (New Guinea, Australia) 

Lamiopsis temmincki (Müller and Henle, 1839)


Loxodon macrorhinus Müller and Henle, 1839 

Negaprion acutidens (Rüppell, 1837) 

Prionace glauca (Linnaeus, 1758) 

Rhizoprionodon acutus (Rüppell, 1837) 

Rhizoprionodon oligolinx Springer, 1964 

Rhizoprionodon taylori (Ogilby, 1915) 

Scoliodon laticaudus Müller and Henle, 1838 

Triaenodon obesus (Rüppell, 1837) 

SPHYRNIDAE: Hammerhead sharks 

Eusphyra blochii (Cuvier, 1817) 

Sphyrna lewini (Griffith and Smith in Cuvier, Griffith and Smith, 1834) 

Sphyrna mokarran (Rüppell, 1837) 

Sphyrna zygaena (Linnaeus, 1758) 

References 

Bigelow, H.B. and W.C. Schroeder. 1948. Sharks. Mem. Sears Fnd. Mar. Res., (1):56-575. 

Chu Y.-T. (Zhu Yuanding) (ed.). 1963. Fishes of the South China Sea. People’s Republic of China, 1184 p. 

Compagno, L.J.V. 1984. FAO species catalogue. Vol. 4. Sharks of the world. An annotated and illustrated catalogue of 

shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fish. Synop., (125)Vol.4,Pt.1:249 p. 


shark species known to date. Part 2. Carcharhiniformes. FAO Fish. Synop., (125)Vol.4,Pt.2:251-655. 

Compagno. L.J.V. 1988. Sharks of the order Carcharhiniformes. Princeton, New Jersey, Princeton University Press, 

572 p. 

Fowler, H.W. 1941. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostariophysi obtained by 

United States Bureau of Fisheries Steamer ALBATROSS in 1907 to 1910, chiefly in the Philippine Islands and 

adjacent seas. Bull. U. S. Natl. Mus., 100(13):879 p. 

Garman, S. 1913. The Plagiostomia. Mem. Mus. Comp. Zool. Harv. Univ., 36:515 p. 

Gubanov, Y.P., V.V. Kondyurin, and N.A. Myagkov. 1986. Sharks of the World Ocean. Identification Handbook. Moscow, 

Agropromizdat, 272 p. 

Last, P.R. and J.D. Stevens. 1993. Sharks and rays of Australia. Australia, CSIRO, 513 p. 

Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino, K. M. Muzik (Eds). 1984. The fishes of the Japanese 

Archipelago. Tokai, Japan, Tokai University Press, 2 vols., 435 p. 

Monkolprasit, S. 1984. The cartilaginous fishes (Class Elasmobranchii) found in Thai waters and adjacent areas. Dept. 

Fish. Biol., Fac. Fish., Kasetsart Univ., Bangkok, 175 p. 

Shen Shih-Chieh, C.T. Chen, H.M. Chen, L.W. Chen, W.E. Eschmeyer, S.J. Joung, S.C. Lee, H.K. Mok, K.T. Shao, and 

C.S. Tzeng. 1995. Fishes of Taiwan, 960 p. 



1208 Sharks 

HEXANCHIDAE 

Cowsharks, sixgill, and sevengill sharks 

by L.J.V. Compagno and V.H. Niem 

Diagnostic characters: Small to large sharks with slender to stout bodies. Head with 6 or 7 pairs of 

long gill slits, the last pair in front of pectoral-fin origins, the first pair not connected across throat; 

short dermal gill rakers present on inner gill slits; spiracles present, small; nostrils without barbels or nasoral 

grooves; no nictitating lower eyelids; snout short, acutely to bluntly pointed; mouth very long and extending 

far behind eyes; teeth of upper and lower jaws unlike at sides of mouth, uppers small, narrow, with a 

main cusp and often smaller cusplets, lowers very large, broad, compressed, and saw-like, with a series 

of cusps or large cusplets. A single dorsal fin, posterior to pelvic fins; anal fin present; caudal fin much less 

than 1/2 the total length, strongly asymmetrical, with a pronounced subterminal notch but lower lobe very 

short. Caudal peduncle not depressed, without keels; no precaudal pits. Intestinal valve of spiral type. 

Colour: grey, blackish, or brown above, lighter below. 

6-7 gill 

slits 

a single dorsal fin 

no precaudal pits 

Habitat, biology, and fisheries: These are moderately abundant, inshore to deep-water sharks, found in 

shallow bays down to the continental slopes and submarine canyons, near the bottom or well above it. They 

feed on a wide variety of bony fishes, other sharks, batoid fishes, marine mammals, cephalopods, and 

crustaceans. Cow sharks are comparatively unimportant but regular components of shark fisheries and 

bycatches of other fisheries and are incidentally caught in trawls. They may bite aggressively during capture 

but only the larger species are potentially dangerous, particularly when provoked. 


None. No other sharks in the area have a single dorsal fin and 6 or 7 gill slits. 

subterminal 

notch 

teeth of left side (Hexanchus griseus) intestinal valve of spiral type 

Key to the species of Hexanchidae occurring in the area 

1a. Seven gill slits (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heptranchias perlo 

1b. Six gill slits (Figs 2 and 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

7 gill 

slits 

Fig. 1 Heptranchias perlo

Hexanchidae 1209 

2a. Lower jaw with 6 rows of large comb-like teeth on each side; dorsal-fin base separated 

from upper caudal-fin origin by a distance about equal to, or slightly greater than its 

length (Fig. 2); size very large, up to 4.8 m . . . . . . . . . . . . . . . . . . . . Hexanchus griseus 

2b. Lower jaw with 5 rows of large comb-like teeth on each side; dorsal-fin base separated 

from upper caudal-fin origin by a distance much greater that its length (Fig. 3); size 

smaller, up to 1.8 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hexanchus nakamurai 

6 gill 

slits 

Fig. 2 Hexanchus griseus 






Fig. 3 Hexanchus nakamurai 

(after Last and Stevens, 1994) 

References 


shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fish. Synop., (125)Vol.4,Pt.1: 249 p. 

Ebert, D.A. 1990. The taxonomy, biogeography and biology of cow and frilled sharks (Chondrichthyes:Hexanchiformes). 

Unpublished Ph.D. thesis. Grahamstown, Rhodes University, 308 pp. 


Springer, S. and R.A. Waller. 1969. “Hexanchus vitulus”, a new sixgill shark from the Bahamas. Bull. Mar. Sci., 

19(1):159-174. 

6 gill 

slits

1210 Sharks 


En - Sharpnose sevengill shark; Fr - Requin perlon; Sp - Cañabota bocadulce. 

Maximum total length about 1.37 m. On the bottom of the outer continental and insular shelves and 

upper slopes in depths of 100 to 400 m, also inshore and down to 1000 m. Feeds on bony fish and 

squid. Caught as bycatch in bottom trawls, but of minor importance to fisheries. Almost circumglobal 

in tropical and temperate seas, except for the eastern North Pacific, with a primarily continental 

distribution. 


En - Bluntnose sixgill shark; Fr - Requin grise; Sp - Cañabota gris (Cañabota). 

Maximum total length about 4.8 m. On the outer continental and insular shelves and upper slopes 

down to at least 1 875 m. Benthic or pelagic, sluggish, bottom-dwelling; feeds on a wide range of 

marine organisms. Caught in bottom trawls and with hook-and-line; utilized fresh, frozen, and dried 

salted, also for fishmeal and oil. Almost circumglobal in tropical and temperate seas, found off 

continents, oceanic islands, and on sea mounts in the Atlantic, Mediterranean Sea, and Indo-Pacific. 


En - Bigeye sixgill shark. 

Maximum total length about 1.8 m. On the continental and insular shelves and upper slopes, usually 

near the bottom at depths from 90 to 600 m. Probably feeds on bony fish and crustaceans. Eastern 

Central America and Caribbean Sea, eastern Atlantic from France to Morocco, including the 

Mediterranean Sea, Indo-West Pacific off eastern and southern Africa and Japan, Taiwan Province 

of China, Philippines, New Caledonia, and Australia. 

(after Last and Stevens, 1994)

Echinorhinidae 1211 

ECHINORHINIDAE 

Bramble sharks 


Diagnostic characters: Large sharks with cylindrical bodies without abdominal ridges. Head moderately 

depressed; last (fifth) gill slits abruptly expanded in width; spiracles present, very small, well 

behind eyes; nostrils far apart from each other; snout short; mouth broadly arched, with very short labial 

furrows that do not encircle mouth; teeth alike in both jaws, strongly compressed and blade-like, with 

a cusp and up to 3 side cusplets in adults, but with a cusp only in young. Two small spineless posterior 

dorsal fins, both smaller than the pelvic fins, situated close together, the origin of the first dorsal fin behind 

the pelvic-fin origin; anal fin absent; caudal fin without a subterminal notch. Dermal denticles moderately 

large to very large, thorn-like. Intestine valve of spiral type. Colour: greyish brown dorsally, pale to white 

below. 

2 posterior dorsal fins 

Habitat, biology, and fisheries: These are large, poorly known wide-ranging, deep-water sharks, with a 

spotty but virtually circumglobal distribution on continental and insular shelves and slopes, on or near the 

bottom. They are ovoviviparous and feed on a variety of benthic and neritic fishes, as well as crabs, 

octopuses, and squids. The 2 known species are uncommon to rare in most areas where they occur and 

hence are of minimal interest to fisheries. They generally form a bycatch of other fisheries, including those 

for other sharks, and are taken on line gear, deepset gill nets, and more commonly in bottom trawls, 

sometimes caught on rod and reel by sports anglers. Very sluggish harmless sharks, never recorded as 

attacking people. 

Remarks: This family is sometimes placed as a synonym of Squalidae, but morphological studies indicate 

that it is very distinct from the Squalidae. It has a single living genus, Echinorhinus. 


Squalidae and Pristiophoridae: fifth gill slits not abruptly larger than first to fourth; spiracles larger; first 

dorsal-fin origin well anterior to pelvic-fin origins; pelvic fins usually about as large as second dorsal fin or 

smaller; Pristiophoridae also with rostral saw and barbels. 

Squatinidae: trunk much flattened dorsoventrally; mouth terminal; eyes on upper surface of head; teeth not 

blade-like, with a single cusp and no cusplets; origin of first dorsal fin posterior to pelvic-fin bases; anterior 

margins of pectoral fins expanded as triangular lobes past the gill slits and partly concealing them; both 

the pectoral and pelvic fins very large and wing-like; caudal fin nearly symmetrical, but with lower lobe 

longer than upper lobe. 

All other shark families: anal fin present. 

pelvic fins usually smaller 

than 1 st dorsal fin 

Squalidae 

body with large 


terminal 

mouth 

anterior 

lobe 

no anal fin 

pectoral fin 

enlarged 

Squatinidae 

pelvic fin 


behind pelvic fin 

long lower 

lobe

1212 Sharks 

Key to the species of Echinorhinidae occurring in the area 

1a. Denticles on body few, irregulary distributed, 

relatively large, not stellate, some 

fused into compound plates with multiple 

cusps (Fig. 1a) . . . . . . . Echinorhinus brucus 

1b. Denticles on body numerous, regulary 

distributed, relatively small, stellate, not 

fused into plates with multiple cusps 

(Fig. 1b) . . . . . . . . . . . . Echinorhinus cookei a) Echinorhinus brucus b) Echinorhinus cookei 





References 


shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fish. Synop., (125)Vol.4,Pt.1: 249 p. 

Garrick, J.A.F. 1960. Studies on New Zealand Elasmobranchii. Part 10. The genus “Echinorhinus”, with an account of 

a second species, “E. cookei”. Trans. R. Soc. N.Z., 88(1):105-117. 

Springer, S. and R.A. Waller. 1969. “Hexanchus vitulus”, a new sixgill shark from the Bahamas. Bull. Mar. Sci., 

19(1):159-174. 


En - Bramble shark; Fr - Squale bouclé; Sp - Tiburón de clavos. 

Maximum total length about 3.1 m. A large, sluggish, primarily deep-water shark of the continental 

and insular shelves and upper slopes at depths from the intertidal to 900 m. Bottom-dwelling; feeds 

on small bony fishes, other sharks, and crabs. Of minor importance to fisheries in the area; caught 

in bottom trawls. Wide-ranging in tropical and temperate areas in the Atlantic, Mediterranean Sea, 

Indian Ocean, and western Pacific. 


Fig. 1 dermal denticles 

En - Pickly shark; Fr - Squale bouclé du Pacifique; Sp - Tiburón negro espinoso. 

Maximum total length about 4 m. A large, sluggish bottom-living shark, occurring on continental and 

insular shelves and upper slopes at depths from 11 to least 424 m. Feeds on a variety of fishes, 

including other sharks, also octopuses and squids. Of minor importance to fisheries; occasionally 

taken by line gear, gill nets and bottom trawls. Tropical and temperate areas of the western and 

Central Pacific, also occurring in the eastern Pacific from Oregon to the Gulf of California and in 

Peru and Chile. 



Squalidae 1213 

SQUALIDAE 

Dogfish sharks 


Diagnostic characters: Small to moderately large sharks, with cylindrical or slightly compressed 

bodies, either without ridges between pectoral and pelvic fins or with inconspicuous ridges 

(Centroscymnus, Dalatias, Scymnodon). Head with 5 gill slits, all anterior to pectoral fins, the fifth not 

abruptly longer than the others; spiracles always present, moderately large; eyes on sides of head, 

without nictitating eyelids; nostrils usually well apart from each other; snout short to moderately long, 

not formed as a rostral saw; no barbels on snout; mouth arched or transverse; teeth strong-cusped, alike 

or dissimilar in both jaws, with or without cusplets. Two dorsal fins with a long to very short spine 

sometimes present (tip of latter may be concealed by skin), on their anterior margins; origin of first dorsal 

fin varying in position from a little (Isistius) in front of pelvic-fin origins to over pectoral-fin bases; pelvic fins 

equal to or smaller than second dorsal fin; no anal fin; caudal fin strongly asymmetrical to nearly 

symmetrical, with a lower lobe varying from virtually absent to very strong. Dermal denticles usually 

close-set, not greatly enlarged and plate-like. Intestinal valve of spiral type. Colour: back greyish in 

shallow-water species, dark to black in those from deep water; several species have light organs (in the 

area, members of the genera Etmopterus, Euprotomicrus, Squaliolus, and Isistius). 

dorsal-fin 

spines often 

present 

pelvic fin 

anal fin absent 

Habitat, biology, and fisheries: Dogfish sharks occurring in warm-temperate and tropical areas are mostly 

confined to deeper water (50 m and more); those occurring in cold-temperate water are usually shallowwater 

forms. Dogfish sharks often form schools; they feed mainly on fishes, and may cause damage to 

fishing gear when preying on the catch. One species in the area, the “cookiecutter shark” (Isistius 

brasiliensis) is semiparasitic, attaching to large fishes, whales and dolphins with its suctorial lips and 

gouging conical plugs of flesh out of its victims. In the western Pacific, dogfish sharks support important 

deep-water line fisheries, for their squalene-rich livers. The family has mainly potential importance as a 

fishery resource for food and liver oil. 

Remarks: Ongoing research on the systematics of this family suggests that it should be divided into several 

families. The traditional arrangement is retained here as a temporary expedient. 


Echinorhinidae: body set with sparse, large, plate-like denticles; spiracles small; fifth pair of gill slits abruptly 

longer than others; first dorsal-fin origin over or posterior to pelvic-fin origins; pelvic fins much larger than 

second dorsal fin. 

Pristiophoridae: snout elongated into a flattened blade, with lateral teeth; barbels present in front of nostrils. 

Echinorhinidae Pristiophoridae

1214 Sharks 

Squatinidae: trunk much flattened dorsoventrally; eyes on 

upper side of head; anterior margins of pectoral fins extending 

forward past gill openings and partly concealing 

them; pelvic fins also very broad, wing-like. 

All other shark families: anal fin present. 

Key to the species of Squalidae occurring in the area 

1a. Second dorsal fin, and usually first dorsal 

fin (except in Squaliolus), without a 

spine . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Spines present on both dorsal fins . . . . . . . → 7 

2a. Lips fringed (Fig. 1a); edges of lower teeth serrated (Fig. 2a) . . . . . . . . . . . . . Dalatias licha 

2b. Lips not fringed; edges of lower teeth smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

a) ventral view of head 

b) fringed lips 

Fig. 1 Dalatias licha 

ventral view 

Squatinidae 

c) lateral view 

dorsal view 

3a. Cusps of lower teeth erect, distal edges not notched (Fig. 2b); lips expanded and 

suctorial; rear end of first dorsal-fin base about over pelvic-fin origins (Fig. 3) . . Isistius brasiliensis 

3b. Cusps of lower teeth oblique, distal edges notched (Fig. 2c); lips not expanded and 

suctorial; rear end of first dorsal-fin base well in front of pelvic-fin origins . . . . . . . . . . . . → 4 

serrated 

smooth 

not 

notched 

a) Dalatias licha b) Isistius 

brasiliensis 

Fig. 2 lower tooth 

notched 

c) Euprotomicrus 

bispinatus 


Fig. 3 Isistius brasiliensis 

pelvic fin 

4a. Second dorsal-fin base as long as first dorsal-fin base; upper caudal-fin lobe not 

shortened, caudal fin not paddle-shaped (Fig. 4); giant benthic sharks, adults to over 

4 m total length and possibly born at a size over 35 cm . . . . . . . . . . . . . Somniosus pacificus 

(occurrence in the area uncertain) 

4b. Second dorsal-fin base at least twice as long as first dorsal-fin base; upper caudal-fin 

lobe shortened, caudal fin paddle-shaped (Fig. 5); dwarf oceanic sharks, adults not 

exceeding 15 to 27 cm total length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5 

Fig. 4 Somniosus pacificus Fig. 5


5a. First dorsal fin without spine; fin length less than 1/3 the length of second dorsal fin; first 

dorsal-fin base closer to pelvic fins than to pectoral fins; snout bulbously conical, snout 

length about 2/5 of head length (Fig. 6) . . . . . . . . . . . . . . . . . . Euprotomicrus bispinatus 

5b. First dorsal fin with a spine, sometimes partly hidden by skin; fin length about 1/2 the 

length of second dorsal fin; first dorsal-fin base closer to pectoral fins than to pelvic fins; 

snout pointed, snout length about 1/2 length of head (Fig. 7) . . . . . . . . . . . . (Squaliolus) → 6 


spine 1 st dorsal fin 

pectoral fin 

pelvic fin 

Fig. 6 Euprotomicrus bispinatus 

Fig. 7 Squaliolus 

6a. Eye large, diameter 73 to 86% of interorbital width; upper margin of eyelid nearly straight; 

upper lip without papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . Squaliolus laticaudus 

6b. Eye smaller, diameter 46 to 70 % of interorbital width; upper margin of eye angular, 

chevron-shaped; upper lip with a pair of prominent lateral papillae (rarely indistinct) . . Squaliolus aliae 

7a. Teeth comb-like in both jaws, with a cusp and 1 or more cusplets, not blade-like; skin 

and muscle of body extremely soft, scabby, semi-gelatinous, and delicate, usually badly 

damaged by capture . . . . . . . . . . . . . . . . . . . . . . . . . . Centroscyllium cf. kamoharai 

7b. Teeth compressed and blade-like in lower jaw, with a single erect to oblique cusp, upper 

teeth comb-like (Etmopterus), blade-like (Squalus), or cuspidate and without cusplets; 

skin and muscle of body more or less firm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

8a. Teeth nearly alike in both jaws, the lowers not greatly enlarged; 

both with strongly oblique nearly horizontal cusps 

(Fig. 8); fin spines without grooves on sides, no subterminal 

notch on caudal fin; caudal peduncle with a strong keel and 

usually an upper precaudal pit . . . . . . . . . . . . . . . (Squalus) → 9 

8b. Teeth more or less unlike in both jaws, the lowers much larger 

than uppers, the latter with erect to oblique cusps; fin spines 

with grooves on sides; subterminal notch on caudal fin; 

caudal peduncle without keels or precaudal pits . . . . . . . . . . → 16 

9a. Preoral snout greatly elongated, about 2.1 to 2.7 times 

mouth width (Fig. 9), preorbital snout 2 to 2.5 times eye 

length in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

9b. Preoral snout short to moderately elongated, 1.7 times 

mouth width or less, preorbital snout less than 2 times eye 

length in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11 

Fig. 8 upper and lower 

tooth (Squalus) 

10a. Lower caudal-fin lobe with a conspicuous black tip; mouth 

larger, 2.1 in preoral snout and equal to internarial space 

(Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . Squalus melanurus 

10b. Lower caudal-fin lobe with a conspicuous white tip; mouth 

smaller, 2.7 in preoral snout and about 3/4 of internarial 

space (Fig. 11) . . . . . . . . . . . . . . . . . . . . . Squalus rancureli 

Fig. 9 ventral view of head 

Fig. 10 Squalus melanurus Fig. 11 Squalus rancureli

1216 Sharks 

11a. Diagonal distance from centre of snout tip to inner edge of nostril greater than distance 

from nostril to upper labial furrow (Fig. 12a) . . . . . . . . . . . . . . . . . . . . . . . . . . . → 12 

11b. Diagonal distance from centre of snout tip to inner edge of nostril less than or about 

equal to distance from nostril to upper labial furrow (Fig. 12b) . . . . . . . . . . . . . . . . . → 14 

12a. Snout acutely pointed; eye closer to first gill opening than snout tip (Fig. 13) . . . Squalus japonicus 

12b. Snout narrowly parabolic; eye slightly closer to snout tip than first gill opening . . . . . . . . . → 13 

a) b) 


13a. Head relatively narrow, direct preorbital distance longer than interorbital distance; 

monospondylous vertebrae 38 to 42 or more (Fig. 14) . . . . . . . . . . . . . . . . . Squalus sp. F 

13b. Head relatively broad, direct preorbital distance shorter than interorbital distance; 

monospondylous vertebrae 43 to 46 (Fig. 15) . . . . . . . . . . . . . . . . . . . Squalus mitsukurii 

14a. First dorsal fin raked backwards 

slightly (Fig. 16a); denticles lanceolate 

(Fig. 17a); precaudal vertebrae 

78 to 82 (Fig. 18) . . . . . . . . Squalus megalops 

14b. First dorsal fin more upright (Fig.16b); 

denticles with 3 posterior cusps 

(Figs 17b, c); more than 82 precaudal 

vertebrae . . . . . . . . . . . . . . . . . . → 15 

Fig. 13 Squalus japonicus 

Fig. 14 Squalus sp. F Fig. 15 Squalus mitsukurii 

a) b) 

Fig. 16 first dorsal fin 

a) b) lateral keels c) 

Fig. 17 Fig. 18 Squalus megalops


15a. Dorsal-fin spines slender (Fig. 19); a dark bar along base of lower caudal-fin lobe 

(Fig.19), more prominent in juveniles; denticle crowns without lateral keels (Fig. 17b); 

precaudal vertebrae mostly 94 to 96 . . . . . . . . . . . . . . . . . . . . . . . . . . Squalussp. A 

15b. Dorsal-fin spines robust (Fig. 20); no dark bar on base of lower caudal-fin lobe (Fig. 20); 

denticle crowns with lateral keels (Fig. 17c); precaudal vertebrae 90 to 93 . . . . . . Squalus sp. B 

Fig. 19 Squalus sp. A Fig. 20 Squalus sp. B 

16a. Upper teeth with slender primary cusp and 1 or more cusplets on each side (Fig. 21b); 

second dorsal fin noticably larger than first . . . . . . . . . . . . . . . . . . . . (Etmopterus) → 28 

16b. Upper teeth with slender to thick primary cusps but with no cusplets; second dorsal fin 

as large or noticably smaller than first . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 17 

17a. Snout greatly elongated, its length greater than distance from centre of mouth to 

pectoral-fin origins (Fig. 22a); dermal denticles of back pitchfork-shaped, crowns on tall, 

slender pedicels (Fig. 22b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Deania) → 18 

17b. Snout short to moderately elongated, its length equal or usually less than distance from 

centre of mouth to pectoral-fin origins; dermal denticles with short pedicels and broad 

crowns, not pitchfork-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 20 

cusplets 

primary 

a) upper teeth b) lower teeth 

cusp 

Fig. 21 Etmopterus 

18a. A subcaudal keel on the lower 

surface of the caudal peduncle 

(Fig. 23) . . . . . . . . . Deania profundorum 

18b. No subcaudal keel on the lower 

caudal peduncle (Figs 24 and 25) . . . . . →19 


b) dermal denticle 

19a. First dorsal fin rather high, angular, and short, distance from its spine origin to its free tip 

about 2/3 of distance from rear origin of second dorsal-fin spine to free rear tip of its fin 

(Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Deania quadrispinosa 

19b. First dorsal fin rather low, rounded, and long, distance from its spine origin to its free rear 

tip greater than distance from origin of second dorsal-fin spine to free rear tip of its fin 

(Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Deania calcea 

a 

b 

Fig. 22 Deania 

Fig. 23 Deania profundorum 

Fig. 24 Deania quadrispinosum Fig. 25 Deania calcae 

keel

1218 Sharks 

20a. Upper teeth relatively broad and low-cusped, the lowers low and wide (Fig. 26); 

dorsal-fin spines prominent and strong; origin of first dorsal-fin spine over or just 

posterior to inner margins of pectoral fins; inner corners of pectoral fins angular or 

greatly elongated (Fig. 26b) . . . . . . . . . . . . . . . . . . . . . . . . . . (Centrophorus) → 21 

20b. Upper teeth lanceolate and high; dorsal-fin spines very small, the first dorsal spine well 

posterior to pectoral fin tips; inner corners of pectoral fins short and broadly rounded 

on their posterior ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 27 

21a. Dermal denticles on sides of body with leaf-like, overlapping crowns on elevated narrow 

pedicels extending above the denticle bases, and with 3 or more medial and lateral cusps 

on their posterior ends. (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . Centrophorus squamosus 

21b. Dermal denticles on sides of body with flat sessile, not overlapping crowns atop the 

denticle bases, without separate pedicels and with or without a posterior medial cusp 

(Fig. 28) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 22 

upper and lower teeth 


high 

pedicel 

Fig. 26 Centrophorus Fig. 27 Centrophorus squamosus 

no high 

pedicel 

a) Centrophorus moluccensis b) Centrophorus lusitanicus 

Fig. 28 dermal denticles 

c) Centrophorus granulosus 

22a. Second dorsal fin very small, 1/2 height of first dorsal fin or less, with spine origin usually 

well posterior to pelvic fin rear tips (Fig. 29). . . . . . . . . . . . . . . . Centrophorus moluccensis 

22b. Second dorsal fin larger, nearly or quite as high than first dorsal fin, with spine origin 

usually over pelvic fin inner margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 23 

23a. Snout elongated and narrow, preoral length greater than width of head at mouth (Fig. 30) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrophorus isodon 

23b. Snout shorter and broadly parabolic, preoral length less than width of head at mouth . . . . . → 24 


Fig. 29 Centrophorus moluccensis Fig. 30 Centrophorus isodon


24a. Free rear tips of pectoral fins moderately elongated and attenuated, not extending 

behind first dorsal-fin spine; postdorsal space short, about 6% of total length; cusps of 

lateral trunk denticles acutely angular in adults; body colour dark grey (Fig. 31) . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrophorus niaukang 

24b. Free rear tips of pectoral fins greatly elongated, usually extending well behind first dorsal 

spine; postdorsal space longer, about 7.5 to 8% of total length; cusps of lateral trunk 

denticles obtusely angular to absent in adults; body colour medium to light grey . . . . . . . . → 25 

25a. First dorsal fin greatly elongated, base about 1.6 to 2 times base of second dorsal fin 

and over 16% of total length; postventral caudal margin nearly straight in adults (Fig. 32) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrophorus lusitanicus 


25b. First dorsal fin moderately elongated, base about 1.3 times base of second dorsal fin 

and less than 16% of total length; postventral caudal margin deeply notched in adults 

(Figs 33 and 34) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 26 

Fig. 31 Centrophorus niaukang 

26a. Adults with tips of dorsal fins dusky, not prominently marked (Fig. 33) . . Centrophorus granulosus 

26b. Adults with tips of dorsal fins black, prominently marked from base of fins (Fig. 34) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centrophorus atromarginatus 

27a. Lower teeth with comparatively low, more or less oblique cusps; denticles on sides of 

body smooth, without ridges (Fig. 35) . . . . . . . . . . . . . . . . . . . Centroscymnus coelolepis 

27b. Lower teeth with comparatively high, more or less erect cusps; denticles on sides of 

body with cross-ridges as well as transverse ridges (Fig. 36) . . . . . . . Scymnodon squamulosus 

28a. Upper teeth with 4 or 5 pairs of 

cusplets on each side (Fig. 37) 

. . . . . . . . . . Etmopterus decacuspidatus 


28b. Upper teeth usually with 3 or 

fewer pairs of cusplets on each 

side . . . . . . . . . . . . . . . . . . . . → 29 

long 

Fig. 32 Centrophorus lusitanicus 

Fig. 33 Centrophorus granulosus Fig. 34 Centrophorus atromarginatus 

Fig. 35 Centrophorus coelolepis Fig. 36 Scymnodon squamulosus 

upper 

tooth 

Fig. 37 Etmopterus decacuspidatus

1220 Sharks 

29a. Denticles on sides of body in regular lines . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 30 

29b. Denticles on sides of body randomly arranged, not in regular lines . . . . . . . . . . . . . . . → 33 

30a. First dorsal-fin origin well anterior to pectoral fin free rear tip; black flank marking on 

side of tail base with short, truncated posterior lobe; additional oval dark marking near 

base of caudal fin, but no transverse marking across midlength of caudal fin (Fig. 38) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etmopterus splendidus and Etmopterus sp. C 

30b. First dorsal-fin origin over or posterior to pectoral fin free rear tip; black flank marking 

on side of tail base with long, narrow posterior lobe with pointed tip; no oval dark marking 

near base of caudal fin, but with a transverse marking across midlength of caudal fin . . . . . → 31 

31a. Second dorsal fin with bluntly rounded apex and shallowly concave posterior margin; 

black flank marking on side of tail with base under second dorsal-fin spine base; 

posterior branch of flank marking rather broad and short, shorter than anterior branch 

(Fig. 39) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etmopterus lucifer 

31b. Second dorsal fin with angular apex and deeply concave posterior margin; black flank 

marking on side of tail with base anterior to second dorsal-fin spine base; posterior 

branch of flank marking very narrow and greatly expanded, longer than anterior branch . . . . → 32 

Fig. 38 Etmopterus splendidus Fig. 39 Etmopterus lucifer 

32a. Dermal denticles usually present on outer web of second dorsal fin; black longitudinal 

marking on caudal-fin base longer than black longitudinal marking along midlength of 

caudal fin, basal marking with angular rear tip (Fig. 40) . . . . . . . . . . . Etmopterus brachyurus 

32b. Dermal denticles absent on outer web of second dorsal fin; black longitudinal marking 

on caudal-fin base shorter than black longitudinal marking along midlength of caudal 

fin, basal marking with rounded rear tip (Fig. 41) . . . . . . . . . . . . . . . . . Etmopterus molleri 

Fig. 40 Etmopterus brachyurus Fig. 41 Etmopterus molleri 

33a. Caudal peduncle long, distance from pelvic-fin insertions to lower caudal-fin origin as 

long as head (snout to fifth gill openings); flanks with prominent rows of dark dashes 

(Fig. 42) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etmopterus sp. F 

33b. Caudal peduncle shorter, distance from pelvic-fin insertions to lower caudal-fin origin 

shorter than prebranchial head (snout to first gill openings); flanks with or without rows 

of dark dashes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 34 

34a. Upper surface pale, sharply delineated from dark under surface, flank and caudal 

markings distinct (Fig. 43) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etmopterus sp. D 

34b. Upper surface dark, not sharply delineated from dark under surface, flank and caudal 

markings indistinct or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 35 

Fig. 42 Etmopterus sp. F Fig. 43 Etmopterus sp. D


35a. Interdorsal space longer than distance from snout tip to pectoral-fin insertions; distance 

from second dorsal-fin insertion to upper caudal-fin origin about 1/3 as long as 

interdorsal space . . . . . . . . . . . . . . . . . Etmopterus baxteri and Etmopterus “granulosus” 

35b. Interdorsal space about equal to head length; distance from second dorsal-fin insertion 

to upper caudal-fin origin about 1/2 the length of interdorsal space . . . . . . . . . . . . . . . → 36 

36a. Dermal denticles on sides with fairly thick cusps (Fig. 44) . . . . . . . . . . . Etmopterus princeps 

36b. Dermal denticles on sides with slender, bristle-like cusps (Fig. 45) . . . . . . . Etmopterus unicolor 







? Centrophorus isodon (Chu, Meng, and Liu, 1981) 1/ 

? Centrophorus lusitanicus Bocage and Capello, 1864 2/ 




Centrophorus sp. [New Caledonia] 3/ 

Centroscyllium cf. kamoharai Abe, 1966 4/ 

Centroscymnus coelolepis Bocage and Capello, 1864 



? Deania calcea (Lowe, 1839) 



? Etmopterus baxteri Garrick, 1957 5/ 


? Etmopterus decacuspidatus Chan, 1966 6/ 

? Etmopterus granulosus (Günther, 1880) 7/ 



Fig. 44 Etmopterus princeps Fig. 45 Etmopterus unicolor 

1/ Described from Xisha (Paracel) Islands and off the Zujiang River mouth, Hong Kong, in the South China Sea just 

adjacent to the area; a similar species, possibly identical, occurs off the Maldive Islands, Sri Lanka, and the Philippines. 

2/ Uncertain in the area but should be watched for. Often confused with Centrophorus niaukang and C. granulosus. 

3/ This species could not be placed in the present key. Status uncertain. Described from the upper insular slopes of New 

Zealand somewhat south of the area. 

4/ Recently collected in deep water (1 037 to 1 100 m) off Luzon, Philippines; close to this species but possibly distinct. 

5/ This species has been synonymized with Etmopterus granulosus, but this is possibly incorrect. 

6/ Known from the South China Sea just north of the area between the Viet Nam coast and Hainan Island, China. 

7/ Presence in the western Pacific uncertain.

1222 Sharks 


? Etmopterus princeps Collett, 1904 8/ 


? Etmopterus unicolor (Engelhardt, 1912) 9/ 

? Etmopterus sp. C [Last and Stevens, 1994] 





Scymnodon squamulosus (Günther, 1877) 10/ 

? Somniosus pacificus Bigelow and Schroeder, 1944 11/ 










Squalus sp. F [Last and Stevens, 1994] 

References 

Bigelow, H.B. and W.C. Schroeder. 1957. A study of the sharks of the suborder Squaloidea. Bull. Mus. Comp. Zool. 

Harv. Univ., 117(1):150 p. 



Last P.R. and J.D. Stevens. 1993. Sharks and rays of Australia. Australia, CSIRO, 513 p. 

Shirai, S. 1992. Squalean phylogeny. A new framework of “squaloid” sharks and related taxa. Sapporo, Hokkaido 

Univ. Press, 151 p. 

8/ This species was nominally recorded from New Caledonia, but its status there needs clarification. Other large species 

of lanternsharks, including Etmopterus baxteri and E. unicolor may be involved instead. 

9/ Described from Japan; a large lanternshark from the south coast of Australia termed Etmopterus sp. B by Last and 

Stevens (1994) and presently known to extend to just south of the area may be synonymous with E. unicolor. 

10/ Placed in a separate genus, Zameus, by some authors. 

11/ Some records in the South Pacific (Tasmania, off New Zealand and somewhat south of the area, and possibly 

Macquarie Island). Assignment of large southern hemisphere Somniosus to the North Pacific S. pacificus is tentative. 





En - Blackfin gulper shark. 

Maximum total length at least 87 cm. A little-known deep-water dogfish, often confused with 

Centrophorus granulosus, from the upper continental slopes from 183 to at least 450 m. Probably 

taken in bottom trawls. Described from Japan (Suruga Gulf); also known from Taiwan Province of 

China, northern Papua New Guinea, and the Gulf of Aden. 


En -Gulpershark;Fr - Squale-chagrin commun; Sp - Quelvacho. 

Maximum total length at least 96 cm. On the outer continental shelves and slopes near the bottom 

in depths from 100 to 1 200 m. Feeds mainly on bony fishes. Mode of utilization and fishing gear 

uncertain. Western North Atlantic (Gulf of Mexico), eastern Atlantic from France to South Africa, the 

Mediterranean Sea, western Indian Ocean (South Africa, Mozambique, and Aldabra Islands) and 

the western Pacific from Japan, Papua New Guinea, and tropical Australia. 


En - Smallfin gulper shark; Fr - Squale-chagrin cagaou; Sp - Quelvacho de aleta corta. 

Maximum total length about 1 m. On the outer continental shelves and upper slopes at depths from 

130 to 820 m. Bottom-dwelling; feeds primarily on bony fishes, as well as other dogfish sharks, 

cephalopods, and shrimps. Probably taken in bottom trawls; utilized at least for fishmeal. Western 

Indian Ocean from South Africa to Mozambique, India, and western Pacific from Japan to Indonesia 

(Amboina), the Philippines, New Caledonia, and Australia.

1224 Sharks 


En - Taiwan gulper shark; Fr - Squale-chagrin quelvacho; Sp - Quelvacho chino. 

Maximum total length at least 1.6 m, probably the largest gulper shark. On the outer continental 

shelves and upper slopes at depths from 250 to 720 m and probably deeper. Bottom-dwelling; little 

known. Taken in bottom trawls and on deep-set longlines; utilized for fishmeal and for human 

consumption. North Atlantic, southwestern Indian Ocean from South Africa and Mozambique, 

possibly the Maldives, and western Pacific from Japan, South China Sea in the northwestern part 

of the area, and probably Australia. Often confused with Centrophorus granulosus and C. lusitanicus. 


En - Leafscale gulper shark; Fr - Squale-chagrin de l’Atlantique; Sp - Quelvacho negro. 

Maximum total length about 1.6 m. On the continental slopes at depths from 230 to 2 400 m near 

the bottom, also pelagically in the upper 1 250 m of water 4 000 m deep. Caught with bottom trawls, 

line gear, and fixed bottom nets; dried and salted for human consumption, also used for fishmeal. 

Eastern Atlantic from Iceland to South Africa, western Indian Ocean (South Africa, Aldabra Islands) 

and western Pacific from Japan, the Philippines, New Zealand, and southeastern Australia. 


En - Mandarin dogfish; Fr - Squale moustache; Sp - Tollo mandarín. 

Maximum total length about 1.26 m. On or near the bottom of the uppermost continental and insular 

slopes, and probably the outer continental-insular shelves at depths of 146 to 640 m. Probably feeds 

mostly on bottom fishes and some invertebrates. High in squalene oil, but at present not utilized 

commercially. Western Pacific from Japan, Torres Island, New Zealand, and Australia (New South 

Wales).



En - Kitefin shark; Fr - Squale liche; Sp -Carocho. 

Maximum total length least 1.6 m. Occurs on the bottom and in the midwater of the outer continental 

and insular shelves from depths of 40 to 1 800 m. Feeds on bony fish, as well as sharks, skates, 

cephalopods, and crustaceans. Caught for its squalene-rich liver, leather and meat, also for fishmeal. 

Western Atlantic (Georges Bank and Gulf of Mexico), eastern Atlantic from Scotland to Cameroon, 

the Mediterranean, western Indian Ocean (southern Africa), and western and Central Pacific from 

Japan, Australia, New Zealand, and Hawaii. 


En - Arrowhead dogfish; Fr - Squale-savate lutin; Sp - Tollo flecha. 

Maximum total length about 76 cm. On the upper continental and insular slopes, found on or near 

the bottom at depths from 280 to 1 790 m. Feeds on small bony fishes, including lanternfish, squid, 

and crustaceans. Interest to fisheries unknown. Western North Atlantic (North Carolina), eastern 

Atlantic from West Sahara to Namibia, western Indian Ocean from South Africa and the western 

Pacific (Philippines). 


En - Longsnout dogfish; Fr - Squale-savate à long nez; Sp - Tollo trompalarga. 

Maximum total length about 1.15 m. On the outer continental shelves and upper slopes at depths 

of 150 to 820 m, mostly below 400 m. Feeds on small bony fishes. Taken in bottom trawls, but with 

minor importance to fisheries. Southern Africa from Namibia to Mozambique, western Pacific off 

western and southern Australia, and New Zealand.

1226 Sharks 


En - Shorttail lanternshark; Fr - Sagre porte-feu à queue courte; Sp - Tollo lucero mocho. 

Maximum total length about 50 cm. Occurs on or near the bottom at depths of 400 to 610 m. Without 

interest to fisheries at present. Japan, the Philippines, and probably elsewhere in the western Pacific. 

Records from southern Africa possibly based on other species. Often confused with Etmopterus 

molleri and E. lucifer. 



En - Blackbelly lanternshark; Fr - Sagre lucifer; Sp - Tollo lucero diablo. 

Maximum total length about 47 cm. On the outer continental and insular shelves and upper slopes on 

or near the bottom, at depths of 183 to 1 000 m. Feeds mostly on squids and small bony fishes, including 

lanternfish, and also shrimps. Interest to fisheries unknown at present. South Atlantic from Uruguay, 

Argentina, and possibly Namibia, also southern Africa and the western Pacific from Japan, New 

Caledonia, southern Australia, and New Zealand. Some records probably based on Etmopterus molleri 

and E. brachyurus. 


En - Slendertail lanternshark. 

Maximum total length about 46 cm. Probably demersal on the upper continental slope in depths of 

about 250 to 480 m. Only recently recognized as distinct from Etmopterus lucifer. Biology and 

distribution poorly known. Without interest to fisheries. Known from Japan, eastern Australia, and 

New Zealand. 




En - Splendid lanternshark. 

Maximum total length about 30 cm. Probably demersal on the outer continental shelves and upper 

slopes at depths of 120 to 210 m. Biology little known, feeds on squid. Known from Japan, Taiwan 

Province of China, Java, and possibly northwestern Australia if Etmopterus sp. C [Last and Stevens, 

1994] is identical to it. 


En - Pink lanternshark. 

Maximum total length at least 41 cm. Occurs near the bottom on the upper continental slope in 

depths of 800 to 880 m. Biology and distribution almost unknown. Without interest to fisheries at 

present. So far recorded only from off Cairns, northern Queensland. 



En - Lined lanternshark. 

Maximum total length at least 45 cm. On or near the bottom of the upper continental slope at depths 

of 590 to 700 m. Its biology is poorly known. The few known specimens were collected off northern 

Queensland between Cairns and Rockhampton. Interest to fisheries unknown. 


?

1228 Sharks 


En - Pygmy shark; Fr - Squale pygmée; Sp - Tollo pigmeo. 

Maximum total length about 27 cm. Occurs at or near the surface at night and apparently descends 

to below 400 m (possibly as deep as 1 800 m) during the day. Feeds on squid, bony fishes, and 

crustaceans. Without interest to fisheries. Oceanic and circumglobal in the tropical and temperate 

oceans. 


En - Cookiecutter shark; Fr - Squalelet féroce; Sp - Tollo cigarro. 

Maximum total length about 50 cm. Makes diurnal vertical migrations probably from below 1 000 m 

in the day to or near the surface at night. Feeds on free-living deep-water prey, but is also a facultative 

ectoparasite on larger marine organisms. Minor importance to fisheries in the area. Widespread 

oceanic in temperate and tropical oceans. 

Scymnodon squamulosus (Günther, 1877) 

En - Velvet dogfish; Fr - Squale-grogneur velouté; Sp - Bruja terciopelo. 

Maximum total length at least 84 cm. Demersal or pelagic near continental slopes and seamounts 

in depths of 550 to 2 000 m. Without interest to fisheries. Western Atlantic (Gulf of Mexico, Surinam, 

Brazil), eastern Atlantic from Iceland to Senegal, southern Africa and the western Pacific from 

Japan, South China Sea, Australia, and New Zealand. 

? 

?



En - Smalleye pigmy shark. 

Maximum total length about 22 cm.Together with the following species possibly the smallest living shark. 

Epipelagic or mesopelagic near continental and island land masses; makes diurnal migrations probably 

from within 200 m of the surface at night down to about 2 000 m during the day. Feeds on cephalopods 

and small bony fish. Without interest to fisheries. Western Pacific from Japan to Australia. 



En - Spined pygmy shark; Fr - Squale nain; Sp - Tollo pigmeo espinudo. 

Maximum total length about 25 cm. Epipelagic near continental and island land masses, usually 

over the slopes at depths of 200 to 500 m. Feeds on deep-water squid and bony fish. Without interest 

to fisheries. Oceanic and nearly circumtropical. 


En - Japanese spurdog; Fr - Aiguillat togari; Sp - Galludo japones. 

Maximum total length about 91 cm. On the outer continental and insular shelves and uppermost 

slopes at depths of 150 to 300 m, presumably on or near bottom. Interest to fisheries unknown. 

Southeastern Japan to the East China Sea, including Korea and the Philippines.

1230 Sharks 


En - Shortnose spurdog; Fr - Aiquillat nez court; Sp - Galludo ñato. 

Maximum total length about 71 cm. On the outer continental shelves and slopes on or near the 

bottom at depths of 50 to 730 m. Schooling; feeds on bony fish, also on cephalopods, crustaceans 

and other elasmobranchs. Taken in bottom trawls, and by hook-and-line (sports catches); consumed 

fresh, dried salted, or smoked. Eastern Atlantic from Guinea to South Africa, in the Pacific from 

South Africa to Mozambique, from Japan to (possibly) Viet Nam and off Australia, and possibly New 

Caledonia and Vanuatu. Possibly a species complex. Western North Pacific representatives often 

recognized as Squalus brevirostris. 


En - Blacktailed spurdog; Fr - Aiguillat à queue noire; Sp - Galludo cola negra. 

Maximum total length 75 cm (adult females). Occurs on the insular slopes of New Caledonia, at 

depths of 320 to 340 m. Feeds on lanternfishes, boarfishes, barracudinas, and flatheads. Without 

interest to fisheries at present. Known only from New Caledonia, from the Ad and Bulari passes. 


En - Shortspine spurdog; Fr - Aiguillat épinette; Sp - Galludo espinilla. 

Maximum total length about 1.1 m; commonly to about 76 cm. On the continental and insular slopes 

and shelves and upper slopes at depths from 50 to 740 m. Feeds on bony fishes, cephalopods, and 

crustaceans. Caught in bottom trawls, but without importance to fisheries in the area. Considered 

to be widely distributed in temperate and subtropical parts of most oceans but possibly consisting 

of a species complex. 

? 

?



En - Cyrano spurdog; Fr - Aiguillat cyrano; Sp - Galludo cirann. 

Maximum total length at least 77 cm. Occurs on the insular slopes of Vanuatu, at depths of 320 and 

400 m. Without interest to fisheries at present. Known only from the vicinity of Vate, Vanuatu. 


En - Bartail spurdog. 

Maximum total length at least 62 cm. Known only from a few specimens collected off Queensland 

between Cairns and Rockhampton in 220 to 450 m. Interest to fisheries unknown. 



En - Eastern highfin spurdog. 

Maximum total length at least 65 cm.On the upper continental slopes in depths to 240 to 450 cm.Biology 

almost entirely unknown. Interest to fisheries unknown. Eastern Australia from the Queensland Plateau 

to Byron Bay. A similar and probably identical spurdog occurs off northern Papua New Guinea.

1232 Sharks 

Squalus sp. F [Last and Stevens, 1994] 

En - Eastern longnose spurdog. 

Maximum total length about 64 cm. On the continental slope off Queensland between Cape York 

and Rockhampton in depths from 220 to 500 m. A similar small, long-nosed spurdog occurs off north 

and eastern Luzon, Philippines, from coastal waters less than a depth of 40 m to 385 m. Interest to 

fisheries limited. The Philippines spurdog is fished locally. 



Pristiophoridae 1233 

PRISTIOPHORIDAE 

Saw sharks 


Diagnostic characters: Small sharks with cylindrical to somewhat depressed bodies, without lateral 

ridges but tail with long lateral folds reaching caudal fin; precaudal tail about as long as trunk. Head 

not expanded laterally, considerably depressed; 5 or 6 small gill slits present, all in front of pectoral-fin 

origins, their upper ends not expanded onto upper surface of head; no gill sieves or complex rakers on 

internal gill slits; spiracles present and very large, behind eyes; nostrils without barbels, nasoral grooves 

or circumnarial grooves, far anterior to mouth; eyes dorsal on head, without nictitating eyelids; snout 

extremely long, depressed and blade-like, with lateral teeth and unique rostral barbels in front of 

nostrils; mouth small, short, transversely arched, and well behind eyes; labial furrows very short, 

confined to mouth corners; teeth small, not blade-like, with a single low cusp, similar in upper and lower 

jaws and weakly differentiated along the jaws. Two dorsal fins, without spines, the first dorsal fin 

moderately large, high and angular, much shorter than caudal fin, and with its base nearly equidistant 

between pectoral- and pelvic-fin bases; second dorsal fin about as large as first; anal fin absent; caudal 

fin strongly asymmetrical, much less than 1/2 of total length, without a rippled or undulated dorsal margin 

but with a strong subterminal notch; lower lobe not present or very short; vertebral axis of caudal fin slightly 

raised above body axis. Caudal peduncle depressed, without precaudal pits but with low lateral folds 

continuing from precaudal tail. Intestinal valve of spiral type. Colour: uniform or mottled grey, brown or 

yellowish above, lighter below, fins dusky. 

rostral saw 

barbels 

nostrils 

Habitat, biology, and fisheries: These are moderately abundant, primarily deep-water sharks, found on 

the outer continental shelves and upper slopes down to 915 m, sometimes inshore in shallow water. All 

species are ovoviviparous. They probably use their rostral saws to injure and kill small fishes and 

crustaceans, much as do the batoid sawfishes (Pristidae). They have a disjunct distribution at present from 

the western Pacific, western Indian Ocean, and western North Atlantic, but were formerly almost worldwide. 

Saw sharks are taken in bottom trawls, and are used fresh for human consumption, but are only of minor 

importance to fisheries in the area.Considerable fisheries exist in southern Australia, but also in the western 

North Pacific. Harmless sharks, not exceeding 1.4 m total length. 


No other sharks have a rostral saw with barbels. 

Sawfishes (Pristidae, a family of batoid fishes) are (or 

formerly were) common in the area and also have a rostral 

saw, but differ from the sawsharks in having the pectoral fins 

expanded anteriorly over the gill openings and fused to the 

sides of the head, so that the head and pectoral fins form a 

distinct pectoral disc with the gill openings ventral (as in other 

batoids); additionally, the trunk is shorter and more 

depressed, the first dorsal fin is partially or entirely above the 

pelvic-fin bases, the rostral saw has relatively few, uniformly 

large, continuously growing teeth (small, varying in size 

along the rostrum, and not growing, but periodically replaced 

in Pristiophoridae) and no barbels. Furthermore, the species 

of sawfishes are much larger, reaching 6 m or more. 

teeth larger 

gill slits ventral 

no barbels 

no anal fin 

ventral view of head teeth intestinal valve of spiral type 

Pristidae (sawfishes)

1234 Sharks 

Key to the species of Pristiophoridae occurring in thea area 

1a. Barbels closer to rostral tip than mouth or approximately equidistant, prebarbel snout 

45 to 51% of preoral snout; spiracles moderately large, width less than 0.5 of eye 

diameter; distance from nostrils to mouth more than 1.3 times internarial space; colour 

uniform pale yellowish brown above; maximum total length 80 cm, males mature at 

62 cm (Australia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pristiophorus sp. B 

1b. Barbels slightly closer to mouth than rostral tip, prebarbel snout 51 to 55% of preoral 

snout. Spiracles large, width almost 0.75 of eye diameter; distance from nostrils to mouth 

about 1.1 to 1.2 times internarial space; colour uniform dark brown above; maximum 

total length at least 73 cm (females) (Philippines) . . . . . . . . . . . . . . . . . Pristiophorus sp. 



Pristiophorus sp. B [Last and Stevens, 1994] (Australia) 

Pristiophorus sp. (Philippines) 

References 

Compagno, L.J.V. 1984. FAO species catalogue. Vol.4. Sharks of the world. An annotated and illustrated catalogue of 


Last, P. and J.D. Stevens. 1993. Sharks and rays of Australia. Australia, CSIRO, 513 p. 

Springer, S. and H.R. Bullis, Jr. 1960. A new species of sawsharks, Pristiophorus schroederi, from the Bahamas. Bull. 

Mar. Sci. Gulf Caribb., 10(2):241-254. 

Pristiophorus sp. B [Last and Stevens, 1994] 

En - Tropical sawshark. 

Maximum total length at least 84 cm. So far only known from the continental slope off tropical 

northeastern Australia between Rockhampton and Cairns in depths of 300 to 400 m. Interest to 

fisheries unknown. 

Pristiophorus sp. 


En - Philippine sawshark. 

Maximum total length at least 73 cm. A little-known deep-water sawshark, so far only known from 

the upper continental slope off southern Luzon (Balayan Bay, Ragay Gulf) and between Negros and 

Siquijor in the Philippines at depths of 229 to 593 m. Interest to fisheries unknown. Previously 

confused with Pristiophorus cirratus and P. japonicus, which apparently do not occur in the area.

Squatinidae 1235 

SQUATINIDAE 

Angelsharks, sand devils 


Diagnostic characters: Moderately large, flattened, ray-like sharks. Head transversely oval or round, 

with a distinct neck at the pectoral-fin bases; 5 pairs of moderately long gill slits situated 

ventrolaterally and not visible dorsally; no gill rakers; nostrils at tip of snout, with anterior flaps shaped as 

elaborate barbels; eyes on dorsal surface of head, without nictitating eyelids; mouth terminal, short and 

angular, extending under front of eyes when jaws are not protruded; teeth small, similar in both jaws, with 

a single, strong, needle-sharp cusp and no cusplets. Two equally small, spineless dorsal fins located 

far rearward on tail, the first originating behind the pelvic-fin bases; pectoral fins greatly enlarged, with 

a broad triangular lobe extending forward from their bases on either side of gill slits (but not fused 

to sides of head as in rays); pelvic fins enlarged and wing-like; anal fin absent; caudal fin very short, nearly 

symmetrical but not lunate, its lower lobe slightly longer than the upper. Caudal peduncle moderately 

depressed, with a short, low, longitudinal keel on each side, but without precaudal pits. Intestine with an 

auger- or corkscrew-like spiral valve. Colour: grey or brownish above, white below, with irregular darker 

markings or light ocelli. 

nostril 

mouth 

terminal 

gill slits 

pectoral-fin 

lobe 

spiracle 

neck 

Habitat, biology, and fisheries: Angelsharks are widely distributed and often abundant in cool temperate 

to tropical seas, ranging in depth from shallow inshore waters down to the upper continental slope. They 

are bottom-dwelling sharks, often burying themselves in sand or mud, and feed on small fishes and bottom 

invertebrates. Ordinarily harmless, but aggressive when provoked and capable of causing serious cuts with 

their small but needle-sharp teeth and strong jaws. Angelsharks are commonly caught in trawls but their 

use varies from region to region; some are utilized for food and fishmeal; their skin makes good leather 

and shagreen for sanding wood. 


Orectolobidae: also with considerably depressed 

bodies, but pectoral fins much smaller and anal fin 

present. 

The combination of characters in boldface readily 

distinguishes the angelsharks from all other shark 

families in the area. 

pectoral fin 

ventral view dorsal view 

dermal 

flaps 

pelvic fin 

1 st dorsal 

fin 

pectoral fin smaller 

Orectolobidae 

2 nd 

dorsal fin 

lower lobe 

of caudal fin 

anal fin present

1236 Sharks 

Rays (Batoidea): pectoral fins fused to head over 

ventral gill slits, no neck at pectoral-fin bases, 

usually a ventral mouth, and lower lobe of caudal fin 

(when present) much shorter than the upper. 

Key to the species of Squatinidae occurring in the area 

1a. Anterior nasal barbels simple, with narrow spatulate tip . . . . . . . . . . . . . Squatina japonica 

1b. Anterior nasal barbels strongly fringed (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

2a. Interorbital space flat or convex, orbital thorns absent (Fig. 1a); numerous dark spots 

on lower lobe of caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Squatina australis 

2b. Interorbital space concave, orbital thorns usually present (Fig. 1b) more pale spots than 

dark spots on lower lobe of caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . Squatina sp. A 

interorbital space 

flat or convex 

Fig. 1 frontal view of head 







ventral 

mouth 

gill slits 

ventral view 

interorbital 

space concave 

Batoidea 

orbital thorns 

a) Squatina australis 

anterior nasal 

barbels 

b) Squatina sp. A 

dorsal view 

pectoral fin 

References 



Last, P. and J.D. Stevens. 1993. Sharks and rays of Australia. Australia, CSIRO, 513 p.

Squatinidae 1237 


En - Australian angelshark; Fr - Ange de mer australien; Sp - Angelote australiano. 

Maximum total length about 1.52 m. A common but little-known angelshark of the continental shelf 

and uppermost slope, on or near bottom from close inshore to a depth of 130 m. Taken by bottom 

trawlers in Australia for human consumption. Southern Australia from Rottnest Island (western 

Australia) to New South Wales, including Tasmania. 


En - Japanese angelshark; Fr - Ange de mer Kasuzame; Sp - Angelote japones. 

Maximum total length about 2 m. A little known angelshark, found on or near the bottom mainly of 

temperate western North Pacific waters. Used for human consumption, but importance to fisheries 

in the area uncertain.Known from Japan, the Yellow Sea, Korea, northern China, and the Philippines. 


En - Eastern angelshark. 

Maximum total length at least 63 cm. On the outer continental shelf and upper slope off eastern 

Australia between Cairns (Queensland) and Lakes Entrance (Victoria) in depths of 130 to 315 m. 

Utilized for human consumption, but at present of minor importance to fisheries. 


1238 Sharks 

HETERODONTIDAE 

Bullhead sharks, horn sharks 


Diagnostic characters: Small to medium-sized sharks, with cylindrical or slightly compressed 

bodies. Head conical and slightly elevated; 5 pairs of gill slits present on sides of head, the last 3 

above the pectoral-fin bases; spiracles present and small, behind and below eyes; nostrils without barbels 

but with strong circumnarial grooves and with prominent nasoral grooves connecting nostrils to mouth; 

anterior nasal flaps elongated posteriorly and reaching mouth; eyes on dorsolateral surface of head, 

without nictitating lower eyelids; snout very short and bluntly rounded; mouth moderate, arched and 

short, well in front of eyes; labial furrows very large, present on both jaws; teeth strongly differentiated 

along jaws, with anterior teeth small and cuspidate and posteriors enlarged, cuspidate and 

molariform; no small intermediate teeth or a gap between anterior and lateroposterior teeth in upper jaw. 

Two dorsal fins, each with a stout fin spine, the first with its origin over the pectoral-fin bases or inner 

margins; pectoral fins moderately large, not ray-like and without triangular anterior lobes; pelvic fins 

moderately large, with vent continuous with their inner margins; anal fin present; caudal fin with a 

moderately long dorsal lobe and moderately long ventral lobe, the latter shorter than the dorsal lobe. 

Vertebral axis raised into caudal-fin lobe; intestinal valve of spiral type. Colour: brownish to greyish, with 

colour patterns of dark bars, stripes, or saddles in species of the area. 

dorsal fins with stout spine 

teeth strongly 

differentiated 

anal fin present 

Habitat, biology, and fisheries: These are common, sluggish, warm-temperate and tropical bottom sharks 

of the continental and insular shelves and uppermost slopes of the western and eastern Pacific and western 

Indian Ocean. They are apparently night-active sharks and occur on or near the bottom from the intertidal 

to 275 m depth, but mostly in shallower water than 100 m. All species are oviparous. Bullhead sharks 

primarily feed on benthic invertebrates. They are of minimal interest to fisheries, being caught as a bycatch 

of bottom trawl and line fisheries and utilized for human consumption and for fishmeal. They are commonly 

caught by divers and in sport fisheries. These sharks can snap when provoked and occasionally pursue 

and bite their tormentors. 


None. No other living sharks combine fin spines in the dorsal fins with the presence of an anal fin. The 

tooth morphology of bullhead sharks is unique among sharks of the area.

Heterodontidae 1239 

Key to the species of Heterodontidae occurring in the area 

1a. Supraorbital ridges very high, 

abruptly ending behind eyes 

(Fig. 1); body with broad blackish 

bands or saddle-markings . . . 

. . . . . . . . . . . . . Heterodontus galeatus 

1b. Supraorbital ridges relatively low, 

not abruptly ending behind eyes; 

body with narrow bands or stripes . . . . . → 2 

2a. Colour pattern with narrow, dis- 

Fig. 1 Heterodontus galeatus 

crete brown or black vertical 

2b. 

bands on a pale background (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . Heterodontus zebra 

Colour pattern with a set of harness-like narrow dark stripes on the back (Fig. 3) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heterodontus portusjacksoni 

vertical bands 

narrow dark stripes 

Fig. 2 Heterodontus zebra 


The symbol is are given when species accounts are included. 




supraorbital 

ridges high 

bands and saddle markings 

Fig. 3 Heterodontus portusjacksoni 

References 


shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fish. Synop., (125)Vol.4.Pt.1:249 p. 


Regan, C.T. 1908. A synopsis of the sharks of the family Cestraciontidae. Ann. Mag. Nat. Hist. (Ser.8), 1(69):493-497. 

Taylor, L.R., Jr. 1972. A revision of the shark family Heterodontidae (Heterodontiformes, Selachii). Universityof 

California, San Diego, Ph.D. Thesis, 176 p. Available from University Microfilms International, Ann Arbor, Michigan.

1240 Sharks 


En - Crested bullhead shark; Fr - Requin dormeur à crête; Sp - Dormilón carenado. 

Maximum total length about 1.3 m. A moderately common benthic and epibenthic shark of the 

continental shelves from close inshore to a depth of about 90 m. Feeds primarily on sea urchins 

(echinoids), but also on crustaceans, molluscs, and small fishes. Of minor interest to fisheries. 

Western South Pacific of Australia from southern Queensland to New South Wales. 


En - Port Jackson shark; Fr - Requin dormeur taureau; Sp - Dormilón toro. 

Maximum total length about 1.65 m. A common shark of the continental shelves from close inshore 

to depths of at least 275 m. Feeds on benthic invertebrates, primarily echinoderms. Taken by bottom 

trawls, shrimp nets, beach seines, bottom longlines, and by rod and reel, but probably little utilized. 

Southern Australia from the Houtman Abrolhos (western Australia) to New South Wales, including 

Tasmania; a single record from New Zealand. 




En - Zebra bullhead shark; Fr - Requin dormeur zèbre; Sp - Dormilón acebrado. 

Maximum total length about 1.22 m. A common but little-known bottom shark of the continental and 

insular shelves in depths down to at least 50 m. Probably feeds on bottom invertebrates. Of minor 

interest to fisheries. Distributed from Japan, Korea, China, and Viet Nam to Indonesia; also known 

from northern Australia. 


Parascylliidae 1241 

PARASCYLLIIDAE 

Collared carpetsharks 


Diagnostic characters: Small sharks with cylindrical or slightly depressed bodies, without ridges on sides. 

Head narrow and slightly flattened, without lateral flaps of skin;gill slits small, fifth overlapping fourth;internal 

gill slits without filter screens; spiracles minute, much smaller than eyes and not below them; nostrils with 

short, pointed barbels and distinct circumnarial folds and grooves around outer edges of incurrent 

apertures;eyes dorsolaterally on head, with subocular pockets;snout broadly rounded to slightly pointed;mouth 

small, entirely in front of eyes, and arched, without a symphyseal groove on chin; teeth not strongly 

differentiated in jaws, with a medial cusp, lateral cusplets and relatively strong labial root lobes; tooth rows 27 

to 54/25 to 49. Dorsal fins equal sized, first dorsal fin with origin and insertion well behind the pelvic-fin 

bases; pectoral fins small, broad, and rounded, as large as pelvic fins or slightly larger; pelvic fins about as 

large as dorsal fins but slightly greater than anal fin; anal fin somewhat smaller than second dorsal fin, with its 

origin well ahead of second dorsal-fin origin; anal fin with broad base and angular apex, separated by 

a space greater than its base length from lower caudal-fin origin; caudal fin with its upper lobe not elevated 

above the body axis, less than a quarter as long as the entire shark, with a strong terminal lobe and subterminal 

notch but no ventral lobe. Caudal peduncle without lateral keels or precaudal pits. Supraorbital crests absent 

from cranium. Intestinal valve of spiral type. Colour: pattern of dark and light spots and saddle markings, in 

some species also a dark collar around gills. 

spiracle minute 

mouth well in 

front of eyes 

Habitat, biology, and fisheries: These are rare to common, harmless bottom sharks of often deepish 

temperate and tropical, continental waters of the western Pacific, occurring from close inshore down to at 

least 183 m offshore. They are found on muddy, sandy, or rocky bottom, and apparently can change colour 

somewhat to match the bottom type. All species are small, less than a metre long when mature. At least 

some of the species are ovoviviparous. They feed probably on small fish, crustaceans, and other bottom 

invertebrates. Several species are taken in bottom trawls, but utilization is probably minimal. 


Scyliorhinidae: mouth not entirely in front of 

eyes; no circumnarial folds and grooves around 

the nostrils. 

Their mouth and nostril structures, 2 spineless 

dorsal fins and an anal fin, anal-fin origin well 

ahead of second dorsal-fin origin, and minute 

spiracles distinguish the Parascylliidae from all 

other sharks. 

anal-fin origin well ahead 

of 2 nd dorsal-fin origin 

without 

circumnarial 

grooves 

Key to the species of Parascylliidae occurring in the area 

1a. A pair of barbels on throat 

(Fig. 1a); gill region without collar 

marking . . . . Cirrhoscyllium expolitum 

1b. No barbels on throat (Fig. 1b); 

gill region with a prominent dark 

brownish collar . . . . . Parascyllium collare 

mouth not in 

front of eyes Scyliorhinidae 

2 barbels 

a) Cirrhoscyllium expolitum 


subterminal 

notch 

no barbels 

b) Parascyllium collare

1242 Sharks 





References 



Goto, T. and K. Nakaya. 1996. Revision of the genus Cirrhoscyllium, with the designation of a neotype for C. japonicum 

(Elasmobranchii, Parascylliidae). Ichthyol. Res., 43(3):199-209. 


Ogilby, J.D. and A.R. McCulloch. 1908. A revision of the Australian Orectolobidae. J. Proc. R. Soc. N.S.W., 42:264-299. 

Regan, C.T. 1908. A revision of the sharks of the family Orectolobidae. Proc. Zool. Soc. Lond., 1908:347-364. 


En - Barbelthroat carpetshark; Fr - Requin carpette à moustache; Sp - Alfombrera barbuda. 

Maximum total length at least 33.5 cm. A little-known tropical shark of the continental shelf, offshore 

on or near the bottom at a depth of 180 m. Probably oviparous, but eggs are not known; food habits 

unknown. Interest to fisheries unknown. In the China Sea between Luzon, Philippines, and China. 


En - Collared carpetshark; Fr - Requin carpette à collarette; Sp - Alfombrera collareja. 

Maximum total length about 86 cm. A common but little-known temperate-water shark of the 

continental shelf, on or near rock reefs and on firm bottom at depths from 20 to 160 m. Oviparous. 

Probably of minor interest to fisheries, although commonly taken by bottom trawlers and sometimes 

with line gear. Western South Australia from Victoria to southern Queensland. 


Brachaeluridae 1243 

BRACHAELURIDAE 

Blind sharks 


Diagnostic characters: Small sharks. Trunk cylindrical or moderately depressed, precaudal tail 

shorter than head and trunk, lateral ridges on sides of trunk and tail absent. Head broad and 

somewhat flattened, without lateral flaps of skin; gill slits small, fifth close to fourth but not overlapping it; 

internal gill slits without filter screens; spiracles very large, subequal or larger than eyes and somewhat 

below them; nostrils with long, pointed barbels and distinct circumnarial folds and grooves around outer 

edges of incurrent apertures; eyes dorsolaterally situated on head, with subocular pockets; snout broadly 

rounded; mouth small, subterminal on head, and nearly transverse, with a symphyseal groove on 

chin; teeth not strongly differentiated in jaws, with a medial cusp, lateral cusplets and weak labial root lobes; 

tooth rows 32/21. Dorsal fins equal sized, first dorsal fin with origin over the pelvic-fin bases and 

insertion well behind the pelvic fin rear tips; pectoral fins moderate sized, broad and rounded, as large as 

pelvic fins or slightly larger, with fin radials not expanded into fin web; pelvic fins about as large as dorsal 

fins but slightly greater than anal fin; anal fin as large as or somewhat smaller than second dorsal fin,with 

its origin about opposite midbase of second dorsal fin or its insertion; anal fin with broad base and 

angular apex, separated by a space or narrow notch much less than base length from lower caudal-fin 

origin; caudal fin with its upper lobe at a low angle above the body axis, less than 1/3 as long as the entire 

shark, with a strong terminal lobe and subterminal notch but no ventral lobe. Caudal peduncle without 

lateral keels or precaudal pits. Supraorbital crests present on cranium, not laterally expanded. Valvular 

intestine of spiral-ring type. Colour: colour pattern of dark saddles and light spots present, or colour plain. 

large spiracles 

barbels 

long 

length of head and trunk 

Habitat, biology, and fisheries: Blind sharks are common, harmless, inshore bottom sharks confined to 

temperate and tropical continental waters of Australia, in depths from the intertidal down to 110 m. They 

occur on rocky reefs or on coral close inshore, sometimes in water just sufficient to cover them. They are 

known to feed on small fishes, crustaceans, cuttlefish, and sea anemones. They are captured in bottom 

trawls but are not generally utilized; Brachaelurus is captured by sports fishermen. 

Remark: The name “blind shark” stems not from lack of vision but because these sharks close their eyelids 

when removed from the water. 


Ginglymostomatidae: spiracles smaller than eyes, 

nostrils without circumnarial grooves, no symphyseal 

groove on chin, fins angular, second dorsal fin smaller 

than first. 

Hemiscylliidae: nasal barbels shorter; no symphyseal 

groove on chin; precaudal tail greatly elongated, 

somewhat longer than head and trunk. 

barbels 

short 

precaudal tail 

Ginglymostomatidae 

Hemiscylliidae

1244 Sharks 

Key to the species of Brachaeluridae occurring in the area 

1a. Nostrils inferior on snout, nasal barbel 

bifurcated (Fig. 1); anal-fin origin 

below or in front of the second dorsalfin 

origin; no white spots on body 

. . . . . . . . . . . . . Heteroscyllium colcloughi 

1b. Nostrils nearly terminal on snout, nasal 

barbel single lobed (Fig. 2); anal- 

fin origin just behind second 

dorsal-fin origin; mostly with white 

spots on body . . . . . . . . Brachaelurus waddi 





References 



Regan, C.T. 1908. A revision of the sharks of the family Orectolobidae. Proc. Zool. Soc. Lond., 1908:347-364. 


En - Blind shark; Fr - Requin aveugle des roches; Sp - Tiburón ciego de roca. 

Maximum total length to about 1 m; commonly to 60 cm. A common bottom shark of the continental 

shelf from the intertidal zone to about 140 m depth; favours rocky shoreline areas and coral reefs. 

Feeds on small reef invertebrates and small fish. Taken in bottom trawls but not used commercially. 

Western South Australia from southern Queensland to New South Wales. 


a) Heteroscyllium 

colcloughi 

nostrils 

inferior 

Fig. 1 underside of head 


b) Brachaelurus 

waddi 

En - Bluegray carpetshark; Fr - Requin aveugle gris-bleu; Sp - Tiburón ciego gris. 

Maximum total length to about 76 cm. A little-known tropical or subtropical inshore bottom shark of 

the Queensland continental shelf, and off York Peninsula and the Great Barrier Reef. Of minor 

interest to fisheries. 

nostrils 

terminal

Orectolobidae 1245 

ORECTOLOBIDAE 

Wobbegongs 


Diagnostic characters: Small to large sharks with considerably depressed bodies, without ridges on 

sides.Head very broad and flattened, with unique lateral flaps of skin; gill slits small, fifth well separated 

from fourth or close to it but not overlapping; internal gill slits without filter screens; spiracles very large, larger 

than eyes and somewhat below and lateral to them; nostrils with long, pointed or branched barbels and 

distinct circumnarial folds and grooves around outer edges of incurrent apertures; snout truncated; eyes 

dorsolaterally situated on head, with subocular pockets; mouth fairly large, nearly terminal on head, and 

nearly transverse, with a symphyseal groove on chin; teeth strongly differentiated in jaws, with 3rowsof 

fang-like teeth at the upper symphysis and 2 rows at the lower; teeth with a median cusp, lateral cusplets 

variably present or absent, and weak labial root lobes; tooth rows 23 to 26/19.Dorsal fins equal sized, first dorsal 

fin with origin over or slightly behind the pelvic-fin insertions and insertion far behind rear tips of pelvic fins; 

pectoral fins moderate sized or large, broad and rounded, slightly larger than pelvic fins, with fin radials not 

expanded into fin web; pelvic fins larger than dorsal and anal fins; anal fin somewhat smaller than second dorsal 

fin, with its origin about opposite rear 1/3 of second dorsal-fin base or insertion; anal fin with broad base 

and subrectangular apex,separated by a narrow notch much less than base length from lower caudal-fin 

origin; caudal fin with its upper lobe hardly elevated above the body axis, less than 1/4 as long as the entire 

shark, with a strong terminal lobe and subterminal notch but without a ventral lobe.Caudal peduncle without 

lateral keels or precaudal pits. Supraorbital crests present on cranium, not laterally expanded. Intestine valve of 

ring type. Colour: colour pattern highly developed, including dark and light spots, dark saddles, rings, and 

reticulations on back. 

dermal flaps 

depressed body 

Habitat, biology, and fisheries: These are common bottom 

sharks of warm-temperate to tropical continental waters of the 

western Pacific, occurring from the intertidal down to at least 

110 m. They are often found on rocky and coral reefs or on sandy 

bottom, where they lurk and are concealed in part by their cryptic 

coloration and dermal lobes on their heads. All species are 

ovoviviparous, with large litters of 20 or more young. They are 

sluggish sharks and known to feed on bottom fishes and 

invertebrates. They are utilized for food and for their colourful 

skins which are sometimes used for leather. Wobbegongs of all 

sizes, but especially the larger individuals, should be regarded as 

potentially dangerous and should be treated with due respect. 

anal fin present 

fang-like teeth 

anteriormost part of head 

(ventral view)

1246 Sharks 


Squatinidae: also with considerably depressed bodies, 

but pectoral fins much larger and anal fin absent. 

The distinctive flattened body and the narrow dermal 

flaps of skin around mouth and head distinguish the 

wobbegongs from other shark families in the area. 

pectoral fin enlarged 

anal fin absent 

Squatinidae 

Key to the species of Orectolobidae occurring in the area 

1a. Dermal lobes highly branched, present on sides of head and on chin (Fig. 1a); body with 

a reticular pattern of narrow dark lines. . . . . . . . . . . . . . . . . . . Eucrossorhinus dasypogon 

1b. Dermal lobes weakly branched, present on sides of head but absent from chin (Fig. 1b); 

colour pattern variable, but without a reticular pattern of narrow dark lines . . . . . . . . . . . . → 2 

2a. Nasal barbels not branched (Fig. 2a); dermal lobes of head very broad-based, only 2 

or 3 in front of eyes; colour pattern simple, dark rounded saddles with light outlining 

widely spaced by dusky areas and with a few dark spots; saddles on head and trunk 

forming conspicuous eyespots . . . . . . . . . . . . . . . . . . . . . . . . . . . Orectolobus wardi 

2b. Nasal barbels branched (Fig. 2b); dermal flaps narrow-based and more numerous, 5 or 

more in front of eyes; colour pattern with elaborate variegated spots and saddles . . . . . . . . → 2 

3a. Back dark, with light O-shaped markings obscuring darker saddles; about 6 to 10 dermal 

flaps below and in front of eyes (Fig. 2b) . . . . . . . . . . . . . . . . . . . Orectolobus maculatus 

3b. Back with dark colour variegated with light blotches and prominent saddle markings; 

about 5 or 6 dermal flaps below and in front of eyes . . . . . . . . . . . . . . . . . . . . . . . → 3 

4a. Back with light areas between dark saddles marked with broad reticulated dark lines 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orectolobus japonicus 

4b. Back with light areas between dark saddles marked with dark, light-centred blotches 

and spots, not reticulated lines . . . . . . . . . . . . . . . . . . . . . . . . . Orectolobus ornatus 

dermal lobes 

highly branched 

a) Eucrossrhinus 

dasypogon 

dermal lobes 

weakly branched 

b) Orectolobus 

ornatus 

nasal barbels 

not branched 

a) Orectolobus 

wardi 

6-10 

dermal 

flaps 

b) Orectolobus 

maculatus 

nasal barbels 

branched 

Fig. 1 head (ventral view) 

Fig. 2 head (ventral view) 









References 





Regan, C.T. 1908. A revision of the sharks of the family Orectolobidae. Proc. Zool. Soc. Lond., (1908):347-364.

Orectolobidae 1247 


En - Tasselled wobbegong; Fr - Requin-tapis barbu; Sp - Tapicero barbudo. 

Maximum total length about 1.25 m. A little-known inshore bottom shark, present on coral reefs. 

Probably feeds on bottom invertebrates and fishes. Of minor importance to fisheries; the tough skin 

is sometimes used for leather. Known from Indonesia, Papua New Guinea, and northern Australia. 



En - Japanese wobbegong; Fr - Requin-tapis moustache; Sp - Tapicero japonés. 

Maximum total length at least 1 m. A little-known inshore bottom shark, nocturnal in habits. Feeds 

on fish, and presumably on bottom invertebrates. Interest to fisheries probably limited; caught in set 

nets in Japan and used for human consumption; also taken in China, Korea, and Viet Nam. 

Distributed from Japan and Korea southward to Viet Nam and the Philippines. 


En - Spotted wobbegong; Fr - Requin-tapis tacheté; Sp - Tapicero manchado. 

Maximum total length about 3.2 m; commonly between 1.5 and 1.8 m. An abundant, mostly inshore 

bottom shark but taken in depths to at least 110 m. Nocturnal, feeds on bottom invertebrates and 

fishes. Interest to fisheries limited, sometimes utilized for its meat and leather; commonly caught in 

trawls, beach seines, trammel nets, in lobster pots and traps, and with line gear. Western Australia 

to southern Queensland; possibly Japan and South China Sea. 


?

1248 Sharks 


En - Ornate wobbegong; Fr - Requin-tapis paste; Sp - Tapicero ornamentado. 

Maximum total length to about 2.9 m. A common bottom-shark of continental waters, found on 

algal-covered rocky areas and coral reefs to depths of at least 100 m. Nocturnal, probably feeds 

on bottom invertebrates and fishes. Interest to fisheries limited; skin very tough and attractively 

patterned, and making a good leather. Western Pacific from Indonesia, Papua New Guinea and 

Australia (Queensland, New South Wales, Victoria, South Australia). 



En - Northern wobbegong; Fr - Requin-tapis savetier; Sp - Tapicero zapatilla. 

Maximum total length at least 63 cm. A little-known, but possibly common tropical inshore bottom 

shark of the continental shelf. Presumably feeds on bottom invertebrates and fishes, but diet 

unrecorded. Of minor interest to fisheries at present. Northern Australia from Queensland to Onslow 

(western Australia). 




Hemiscylliidae 1249 

HEMISCYLLIIDAE 

Longtail carpetsharks 


Diagnostic characters: Small sharks. Trunk cylindrical or moderately depressed, precaudal tail 

cylindrical and somewhat longer than head and trunk, lateral ridges on sides of trunk and tail present 

or absent. Head not expanded laterally, cylindrical or moderately depressed; 5 small gill slits present, the 

last 3 over the pectoral-fin base, their upper ends not expanded onto upper surface of head; no gill sieves 

or rakers on internal gill slits; spiracles very large and located behind and below eyes; nostrils with 

barbels, nasoral grooves, and circumnarial grooves, close in front of mouth; eyes above and medial to 

sides of head, without nictitating eyelids; snout short to moderately long, slightly depressed, parabolic to 

broadly rounded, not greatly flattened and blade-like and without lateral teeth or barbels; mouth small, 

nearly transverse, and well in front of eyes, without a symphyseal groove on chin; labial furrows 

present on both jaws and relatively large, with upper furrows extending in front of mouth; teeth small, not 

blade-like, with a single cusp on upper and lower teeth and with cusplets small or absent; teeth similar in 

both jaws, not differentiated into medials, anteriors, intermediates, laterals, or posteriors. Two dorsal fins 

without spines, the first moderate sized, subangular, much shorter than the caudal fin, and with its origin 

over or behind the pelvic-fin bases; second dorsal fin about as large as the first and of similar shape; 

anal fin moderately large, very low, broad and rounded, with its origin well behind the second 

dorsal-fin base and its base separated by a notch from the caudal fin; caudal fin strongly asymmetrical, 

much less than 1/2 of total length, without a rippled dorsal margin or lower lobe but with a strong subterminal 

notch; vertebral axis of caudal fin hardly raised above body axis. Caudal peduncle cylindrical, without 

precaudal pits or keels. Intestinal valve of ring type. Colour: back yellowish, brownish or grey-brown, lighter 

below, with dark or light spots or dark saddles, sometimes absent in adults. 

spiracles large 

length of head and body precaudal tail 

Habitat, biology, and fisheries: Longtail carpetsharks are a small 

group of inshore tropical sharks of the Indian Ocean and western 

Pacific, being confined to continental waters and continental 

islands. They are slow-swimming bottom-dwellers, often intestinal valve of ring type 

clambering with their muscular paired fins on coral and rocky reefs. 

At least some of the species are oviparous. They feed on 

invertebrates and small fishes and are harmless to people. Hemiscyllium species are little utilized for 

fisheries, but Chiloscyllium species are commonly caught in small-scale artisanal fisheries and by bottom 

trawlers in the western and eastern Pacific and eastern central Indian Ocean. 


Brachaeluridae: nasal barbels longer; symphyseal 

groove present on chin; precaudal tail not greatly 

elongated, shorter than head and trunk. 

nasal 

barbels 

longer Brachaeluridae 

caudal fin strongly 

asymmetrical

1250 Sharks 

Ginglymostomatidae: precaudal tail not greatly 

elongated, shorter than head and trunk; no 

circumnarial grooves around nostrils; head more 

depressed and flattened; spiracles minute; labial 

furrows not connected across chin by a dermal 

flap (present in Hemiscylliidae); anal fin higher, 

more angular, and separated from the lower 

caudal-fin origin by a space; origin of anal fin 

under second dorsal-fin base. 

Stegostomatidae: precaudal tail not greatly 

elongated, shorter than head and trunk; no 

circumnarial grooves around nostrils, labial 

furrows not connected across chin by a dermal 

flap; first dorsal-fin origin far anterior to pelvic-fin 

bases, its insertion over or slightly anterior to 

pelvic-fin insertions (far posterior to pelvic-fin 

insertions in Hemiscylliidae), second dorsal fin 

much smaller than first dorsal fin; pelvic fins 

much smaller than pectoral fins; anal-fin origin 

under second dorsal-fin base; caudal fin about as 

long as rest of shark. 

Stegostomatidae 

no 

circumnarial 

groove 

barbels 

labial furrows 

not connected 


nasoral 

groove 


circumnarial 

groove 

(ventral view of head) 

labial furrows 

connected by 

dermal flap 

Hemiscylliidae 

Key to the species of Hemiscylliidae occurring in the area 

1a. Nostrils subterminal on snout (Fig. 1a); eyes and supraorbital ridges hardly elevated; 

preoral snout long, mouth closer to eyes than snout tip; no black hood on head or large 

dark spot or spots on sides of body above pectoral fins (Fig. 1b) . . . . . . . . (Chiloscyllium) → 2 

1b. Nostrils terminal on snout (Fig. 2a); eyes and supraorbital ridges prominently elevated; 

preoral snout short, mouth closer to snout tip than eyes; a large spot or spots on sides 

of body above pectoral fins, or a black hood on head (Fig. 2b) . . . . . . . . . (Hemiscyllium) → 6 

nostrils subterminal on snout 


nostrils terminal on snout 


Fig. 1 Chiloscyllium 

Fig. 2 Hemiscyllium 


b) lateral view


2a. Body and tail very slender; length of anal fin from origin to free tip subequal to length 

of hypural caudal-fin lobe from lower caudal-fin origin to subterminal notch (Fig. 3) . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chiloscyllium indicum 

2b. Body and tail moderately slender to relatively stout; length of anal fin considerably 

shorter than hypural caudal-fin lobe (Figs 4 and 5) . . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Body with lateral dermal ridges; young and adults with transverse dark bands and 

numerous white spots (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . .Chiloscyllium plagiosum 

3b. Body without lateral dermal ridges; adults usually without colour pattern, dark transverse 

bands in young only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4 

Fig. 3 Chiloscyllium indicum Fig. 4 Chiloscyllium plagiosum 

4a. Dorsal fins larger than pelvic fins, 

with projecting free rear tips 

(Fig. 5) . . . . . . . Chiloscyllium punctatum 

4b. Dorsal fins smaller than pelvic 

fins, without projecting free rear 

tips . . . . . . . . . . . . . . . . . . . . → 5 

5a. First to second dorsal-fin distance usually more than 9.3% of total length; first dorsal-fin 

height more than 6.6% of total length; second dorsal-fin height usually more than 5.8% 

of total length; dark bands of juveniles not outlined in black (Fig. 6) . . . . . Chiloscyllium griseum 

5b. First to second dorsal-fin distance less than 9.3% of total length; first dorsal-fin height 

less than 6.6% of total length; second dorsal-fin height usually less than 5.8% of total 

length; dark bands of juveniles outlined in black (Fig. 7) . . . . . . . . . . . . Chiloscyllium hasselti 

juvenile 

Fig. 6 Chiloscyllium griseum 

6a. Head and snout with an abrupt 

black hood; body with conspicuous 

large white spots (Fig. 8) . . 

. . . . . . . . . . . . . Hemiscyllium strahani 

6b. Head and snout light, without a 

black hood but with conspicuous 

black spots above pectoral fins; 

body with light spots inconspicuous 

or absent . . . . . . . . . . . . . . . → 7 

Fig. 5 Chiloscyllium punctatum 

juvenile 

Fig. 7 Chiloscyllium hasselti 

Fig. 8 Hemiscyllium strahani

1252 Sharks 

7a. Black spot behind gills small, not in the form of a conspicuous ocellus (Fig. 9) . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiscyllium freycineti 

7b. Black spot behind gills large, in the form of a conspicuous ocellus, ringed with white 

(Figs 10 to 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

8a. Body covered with numerous, densely clustered dark small and large spots that form a 

reticular network of light ground colour between them; dark crossbands strong on ventral 

surface of tail (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiscyllium trispeculare 

8b. Body with sparse, large spots that do not form a reticular network of light ground colour 

between them; dark crossbands not reaching ventral surface of tail . . . . . . . . . . . . . . . → 9 

Fig. 9 Hemiscyllium freycineti Fig. 10 Hemiscyllium trispeculare 

9a. Lateral ocellus not surrounded by large spots; spots present on head in front and below 

eyes (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiscyllium ocellatum 

9b. Lateral ocellus surrounded by large black spots; spots absent from head in front and 

below eyes (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiscyllium hallstromi 

Fig. 11 Hemiscyllium ocellatum Fig. 12 Hemiscyllium hallstromi 






Chiloscyllium plagiosum (Bennett, 1830) 







Reference 



Dingerkus, G. and T.C. DeFino. 1983. A revision of the orectolobiform shark family Hemiscylliidae (Chondrichthyes, 

Selachii). Bull. Am. Mus. Nat. Hist., 176(1):1-93. 




Frequent synonyms / misidentifications: None / Chiloscyllium hasselti Bleeker, 1852; C. plagiosum 

(Bennett, 1830); C. punctatum Müller and Henle, 1838. 

FAO names: En - Grey bambooshark; Fr - Requin-chabot gris; Sp - Bamboa gris. 

Diagnostic characters: A small shark; body moderately stout, without lateral ridges; precaudal tail 

moderately stout, longer than head and trunk. Snout broadly rounded; 5 small gill slits; spiracles large and 

below eyes; eyes hardly elevated, with a low supraorbital ridge above them, without nictitating eyelids; 

nostrils subterminal, with short barbels, nasoral grooves, and circumnarial grooves; mouth small, 

transverse, and well in front of eyes; teeth small, similar in both jaws, with a single small cusp. Two dorsal 

fins, slightly smaller than pelvic fins and without attenuated, projecting free rear tips; origin of first 

dorsal fin varying from over last 1/3 of pelvic-fin bases to over pelvic-fin insertions, second dorsal 

fin almost as large as first; anal fin long, low and broadly rounded, with its origin behind free rear tip of 

second dorsal fin and with its insertion at lower caudal-fin origin; caudal fin strongly asymmetrical, with 

a pronounced caudal subterminal notch but without a ventral lobe, its length less than 1/3 the length of rest 

of shark. Caudal peduncle cylindrical, without keels or precaudal pits. Intestinal valve of ring type. Colour: 

light brown, yellow-brown or grey-brown above, cream below, with 12 or 13 prominent saddle marks 

in young, fading out with growth and absent in adults. 

Size: Maximum total length at least 74 cm. 

Habitat, biology, and fisheries: A common, sluggish inshore bottom dweller. Oviparous, deposits eggs 

in small, oval egg cases on the bottom. Probably feeds mainly on invertebrates. Caught in bottom trawls 

and in fixed bottom gill nets, drifting bottom gill nets, and occasionally pelagic gill nets; in the area regularly 

taken in inshore fisheries off Thailand, and utilized for human consumption. 

Distribution: Western Indian 

Ocean from the Arabian Sea 

eastward to Pakistan, India, 

and probably Sri Lanka; in the 

eastern Indian Ocean and 

western Central Pacific 

extending eastward to 

Malaysia, Thailand, Indonesia, 

Viet Nam, South China, Japan, 

Philippines, and New Guinea, 

but many records need 

confirmation. 

adult 

juvenile

1254 Sharks 


Frequent synonyms / misidentifications: None / Chiloscyllium griseum Müller and Henle, 1839; 

C. plagiosum (Bennett, 1830); C. punctatum Müller and Henle, 1838. 

FAO names: En - Hasselt’s bambooshark. 

(after Dingerkus and DeFino, 1983) 

Diagnostic characters: A small shark, body moderately slender, without lateral ridges, precaudal tail 

moderately slender, longer than head and trunk. Snout broadly rounded; 5 small gill slits; spiracles large 

and below eyes; eyes hardly elevated, with a low supraorbital ridge above them, without nictitating eyelids; 

nostrils subterminal, with short barbels, nasoral grooves and circumnarial grooves; mouth small, 


fins, smaller than pelvic fins and without attenuated, projecting free rear tips; first dorsal-fin origin 

over pelvic-fin bases; second dorsal fin almost as large as first; anal fin long, low, and broadly rounded, 

with its origin behind free rear tip of second dorsal fin and with its insertion at lower caudal-fin origin; 

caudal fin strongly asymmetrical with a pronounced subterminal notch but without a ventral lobe; caudal 

fin less than 1/3 the length of rest of shark. Caudal peduncle cylindrical, without keels or precaudal pits. 

Intestinal valve of ring type. Colour: juveniles with dark grey-brown bands outlined in black, adult 

specimens with uniform medium- to dark-brown colour, the black edgings being the last parts of the 

colour pattern to disappear. 

Size: Maximum total length about 60 cm. 

Habitat, biology, and fisheries: A common inshore bottom-dweller. Biology little known. Presumably 

oviparous and primarily an invertebrate-feeder as with other Chiloscyllium species. Caught in bottom trawls 

and in fixed bottom gill nets, possibly traps and with line gear. Utilized for human consumption in the area. 


from Thailand, Malaysia, and 

Indonesia (Sumatra, Java, 

and Moluccas). 

adult 

juvenile



Frequent synonyms / misidentifications: Chiloscyllium colax (Meuschen, 1781) / Chiloscyllium 

plagiosum (Bennett, 1980). 

FAO names: En - Slender bambooshark; Fr - Requin-chabot élégant; Sp - Bamboa elegante. 

Diagnostic characters: Small sharks, body slender, with low lateral 

ridges, precaudal tail slender, longer than head and trunk. Snout narrowly 

rounded or almost pointed; 5 small gill slits; spiracles large and below eyes; 

eyes hardly elevated, nostrils subterminal, with short barbels, nasoral 

grooves and circumnarial grooves; mouth small, transverse, and well in 

front of eyes, teeth small, similar in both jaws, with a single small cusp. Two 

dorsal fins, about as large as pelvic fins and without attenuated, 

projecting free rear tips; first dorsal-fin origin over inner margins of 

pelvic tins and behind pelvic-fin insertions; second dorsal fin almost as 

large as first; anal fin long, low, and broadly rounded, with its origin far 

behind free rear tip of second dorsal fin and with its insertion at lower 

caudal-fin origin, caudal fin strongly asymmetrical with a pronounced 

subterminal notch but without a ventral lobe; caudal fin less than 1/3 the 

length of rest of shark. Caudal peduncle cylindrical, without keels or 

precaudal pits. Intestinal valve of ring type. Colour: light brown above, cream below, with numerous 

dark spots on body, tail, and fins, these often forming indistinct vertical bars and saddles. 


Habitat, biology, and fisheries: A common, but little-known inshore sluggish bottom dweller. Oviparous, 

deposits eggs in small, oval egg cases on bottom. Probably feeds mainly on invertebrates. Caught in bottom 

trawls and in fixed bottom gill nets, drifting bottom gill nets, and occasionally pelagic gill nets; utilized fresh 

for human consumption, but relatively unimportant to fisheries in the area. 


Pacific from the Arabian Sea 

eastward to India, Sri Lanka, 

Singapore, Thailand, 

Indonesia, Viet Nam, Taiwan 


Philippines, Solomon Islands, 

and possibly Korea and 

Japan. 

? 

? 

? ? 

ventral view of head

1256 Sharks 

Chiloscyllium plagiosum (Bennett, 1830) | 

Frequent synonyms / misidentifications: Scyllium ornatum Gray, 1832 / Chiloscyllium griseum Müller 

and Henle, 1839; C. hasselti Bleeker, 1852; C. indicum (Gmelin, 1789); C. punctatum Müller and Henle, 

1839. 

FAO names: En - Whitespotted bambooshark; Fr - Requin-chabot à taches blanches; Sp - Bamboa 

punteada. 

Diagnostic characters: A small shark, body fairly stout, with lateral ridges, precaudal tail stout, longer 

than head and trunk. Snout rounded anteriorly; 5 small gill slits; spiracles large and below eyes; eyes hardly 

elevated, with a low supraorbital ridge above them, without nictitating eyelids; nostrils subterminal, with 

short barbels, nasoral grooves and circumnarial grooves; mouth small, transverse, and well in front of eyes; 

teeth small, similar in both jaws, with a single small cusp. Two dorsal fins, about equal in size to pelvic 

fins and without attenuated, projecting free rear tips; first dorsal-fin origin over or behind pelvic-fin 

bases; second dorsal fin almost as large as first; anal fin long, low and broadly rounded, with its origin 

somewhat behind free rear tip of second dorsal fin and with its insertion at lower caudal-fin origin; 

caudal fin strongly asymmetrical with a pronounced subterminal notch but without a ventral lobe; caudal 

fin less than 1/3 the length of rest of shark. Caudal peduncle cylindrical, without keels or precaudal pits. 

Intestinal valve of ring type. Colour: a prominent colour pattern of numerous white spots on a dark brown 

background, with darker brown or blackish transverse bands. 

Size: Maximum total length about 95 cm; adult males 67 to 69 cm, an adult female 95 cm. 

Habitat, biology, and fisheries: A common but little-known inshore bottom shark. Oviparous. Regularly 

taken in inshore fisheries in India, Thailand, China, and probably elsewhere where it occurs, and utilized 


Distribution: Indo-West Pacific: 

India, Sri Lanka, Singapore, 

possibly Malaysia, Thailand, 

Indonesia, Viet Nam, China 

(including Taiwan Province), 

Japan, and the Philippines.



Frequent synonyms / misidentifications: Chiloscyllium margaritiferum Bleeker, 1964 / Chiloscyllium 

griseum Müller and Henle, 1839; C. hasselti Bleeker, 1852; C. plagiosum (Bennett, 1830). 

FAO names: En - Brownbanded bambooshark; Fr - Requin-chabot bambou; Sp - Bamboa estriada. 


juvenile 

Diagnostic characters: A small shark, body moderately slender, without lateral ridges, precaudal tail 

moderately slender, longer than head and trunk. Snout rounded anteriorly; 5 small gill slits; spiracles large 

and below eyes; eyes hardly elevated, with a low supraorbital ridge above them, without nictitating eyelids; 

nostrils subterminal, with short barbels, nasoral grooves and circumnarial grooves; mouth small, 


fins, somewhat larger than pelvic fins and with attenuated, projecting free rear tips; first dorsal-fin 

origin over anterior halves of pelvic-fin bases; second dorsal fin almost as large as first; anal fin long, 

low, and broadly rounded, with its origin somewhat behind free rear tip of second dorsal fin and with 

its insertion at lower caudal-fin origin; caudal fin strongly asymmetrical with a pronounced subterminal 

notch but without a ventral lobe; caudal fin less than 1/3 the length of rest of shark. Caudal peduncle 

cylindrical, without keels or precaudal pits. Intestinal valve of ring type. Colour: young with dark 

transverse bands and usually a scattering of a few dark spots; adults light-brown, usually without a 

colour pattern. 

Size: Maximum total length about 1.04 m. 

Habitat, biology, and fisheries: A common inshore bottom shark found on coral reefs, often in tidepools. 

Very tenacious of life, can survive out of water for a long period (half a day). Oviparous, deposited in rounded 

egg cases. Gills sometimes infested by larval isopods (Praniza-larva of the isopod Gnathia). Regularly 

taken in inshore fisheries in India and Thailand, and utilized for human food. In Australia it is taken in beach 

seines and on hook-and-line and is said to prefer squid bait; it is little utilized by Australians but regarded 

as good eating. 

Distribution: Indo-West Pacific: 

India, Thailand, Malaysia, 

Singapore, Indonesia (Java, 

Sumatra, Komodo, Sulawesi), 

Philippines, New Guinea, and 

northern Australia (Northern 

Territory, western Australia, 

Queensland); also Viet Nam, 

China (including Taiwan 

Province), and Japan. 

adult

1258 Sharks 


En - Indonesian speckled carpetshark; Fr - Requin-chabot grivelé; Sp - Bamboa jaspeada. 

Maximum total length at least 46 cm. A little-known bottom shark, probably common on coral reefs. 

Of minor interest to fisheries at present. Western South Pacific from Indonesia (Irian Jaya, Waigeo) 

and Papua New Guinea. 


En - Papuan epaulette shark; Fr - Requin-chabot épaulette; Sp - Bamboa hombrera. 

Maximum total length at least 75 cm. A little-known inshore bottom shark, probably on coral reefs. 

Of minor interest to fisheries at present. Western South Pacific from Papua New Guinea and 

Indonesia (Irian Jaya). 



En - Epaulette shark; Fr - Requin-chabot ocellé; Sp - Bamboa ocelada. 

Maximum total length about 1 m. An abundant, small, harmless tropical shark found on coral reefs 

in shallow water, often in tidepools. Oviparous, feeding on benthic invertebrates. Survives well in 

aquaria, but otherwise not used commercially. Known from New Guinea and Australia (Northern 

Territory, western Australia, Queensland, and New South Wales); possibly also Malaysia, Indonesia 

(Sumatra), and Solomon Islands. 

? 

?



En - Hooded carpetshark; Fr - Requin-chabot moine; Sp - Bamboa capuchona. 

Maximum total length about 75 cm. A little-known inshore bottom shark of singular and unique 

appearance, probably on coral reefs. Of minor importance to fisheries at present. Western South 

Pacific from Papua New Guinea and Indonesia (Irian Jaya). 



En - Speckled carpetshark; Fr - Requin-chabot marqueterie; Sp - Bamboa moteada. 

Maximum total length about 64 cm. A common, small, harmless tropical continental shelf shark that 

is found on coral reefs in shallow water. Oviparous, probably mainly feeding on benthic invertebrates. 

Of minor interest to fisheries. Australia (Northern Territory, Western Australia, and Queensland) and 

possibly Indonesia (Moluccas). 

(after Dingerkus and DeFino, 1983)

1260 Sharks 

GINGLYMOSTOMATIDAE 

Nurse sharks 



Nebrius ferrugineus (Lesson, 1830) 

Frequent synonyms / misidentifications: Ginglymostoma ferrugineum (Lesson, 1830); Nebrius 

concolor Rüppell, 1837; N. doldi Smith, 1953 / None. 

FAO names: En - Tawny nurse shark; Fr - Requin-nourrice fauve; Sp - Gata nodriza atezada. 

spiracles much smaller than eye 

dorsal and anal fins 

with angular apex 

no 

circumnarial 

groove 

barbels 

mouth 

ventral view of head 

nasoral 

groove 

lower tooth intestinal valve of ring type 

Diagnostic characters: A large, relatively stout-bodied shark, without lateral ridges; precaudal tail 

shorter than trunk. Head with 5 moderate gill slits, the last 2 behind pectoral-fin origin and very close to 

each other, no gill rakers; spiracles much smaller than eyes; nostrils close to front of snout, with short 

barbels and nasoral grooves connecting them with the mouth but without circumnarial grooves and 

folds; no nictitating lower eyelids; snout very short, broad, and very broadly rounded or truncated; mouth 

moderately large, nearly transverse and far forward on head, well in front of eyes; teeth small, weakly 

differentiated in different regions of the mouth, somewhat compressed, with short medial cusps and short 

cusplets on sides; tooth rows 24 to 38/22 to 32. Two dorsal fins, both with angular apices, the origin of 

the first about over the pelvic-fin origins and its insertion slightly behind the pelvic-fin insertions; 

second dorsal fin slightly smaller than first; anal fin present, high and with an angular apex, and with 

its origin about under the midbase of the second dorsal fin; caudal fin about 1/3 of total length, strongly 

asymmetrical, with a strong subterminal notch but with ventral lobe weak to short. Caudal peduncle not 

strongly depressed, without lateral keels or precaudal pits. Supraorbital crests present on cranium, these 

laterally expanded. Valvular intestine of ring type. Colour: no colour pattern, tan above, lighter below, fins 

slightly dusky.

Ginglymostomatidae 1261 



Brachaeluridae: spiracles very large, subequal or 

larger than eyes, nostrils with circumnarial grooves, a 

symphyseal groove present on chin, fins broadly 

rounded, dorsal fins equal sized. 

Hemiscylliidae: precaudal tail longer than trunk; 

spiracles large, nearly or quite eye length; nostrils 

Brachaeluridae 

with circumnarial grooves; anal fin very low and 

arcuate. 

Stegostomatidae: body with lateral ridges; spiracles as large as eyes; first dorsal fin with origin far anterior 

to pelvic-fin origins; caudal fin about 1/2 of total length. 



Hemiscyllidae 


caudal fin very elongate 

The combination of characters such as the nasoral grooves, the presence of barbels, the anterior mouth, 

the posterior position of the first dorsal fin, the absence of nictitating lower eyelids, the absence of body 

ridges, caudal keels and precaudal pits, and the asymmetrical caudal fin with ventral lobe weak or absent 

readily distinguishes this family from all others in the area. 

Size: Maximum total length about 3.2 m; commonly to 2.5 m. 

Habitat, biology, and fisheries: A sluggish, nocturnal and sometimes diurnal shallow-water bottom shark 

common on coral and rocky reefs, in lagoons and on sand flats, at depths from the intertidal zone to at 

least 70 m. Ovoviviparous, size at birth about 60 cm. Feeds on a wide variety of bottom invertebrates and 

small fishes; capable of capturing small reef fishes with its powerful suction feeding mechanism. Caught 

inshore in Pakistan, India, Thailand, Philippines, and probably elsewhere where it occurs; taken in bottom 

trawls, in floating and fixed 

bottom gill nets, and with 

longlines; utilized fresh and 

dried-salted for human food; 

livers are processed for 

vitamins; fins dried for the 

oriental sharkfin trade; also 

processed for fishmeal. 

Distribution: In the Indian 

Ocean and western Pacific 

from southeastern Africa and 

the Red Sea eastward to 

Japan, Australia, and Tahiti. 

References 



Dingerkus, G. 1986. Interrelationships of orectolobiform sharks (Chondrichthyes: Selachii). Proc. 2nd Int. Conf. 

Indo-Pacific Fish., 1986:227-245. 


1262 Sharks 

A single species in this family. 

Stegostoma fasciatum (Hermann, 1783) 

STEGOSTOMATIDAE 

Zebra sharks 


Frequent synonyms / misidentifications: Stegostoma varium (Seba, 1758);S. tygrinus (Bonnaterre, 1788) / None. 

FAO names: En - Zebra shark; Fr - Requin zèbre; Sp - Tiburón acebrado. 

adult 

juvenile 

ventral view 

of head 

lower 

tooth 

Diagnostic characters: A large, moderately stout-bodied shark with prominent ridges on side. Head with 5 small 

gill slits, the last 3 behind pectoral-fin origin and the last 2 very close to each other; no gill rakers; spiracles 

subequal in size to eyes; nostrils close to front of snout, with short barbels and nasoral grooves 

connecting them with the mouth but without circumnarial grooves and folds; no nictitating lower eyelids; 

snout very short, broad and bluntly rounded; mouth short, nearly transverse, and far forward on head, well in 

front of eyes; teeth small, poorly differentiated in different regions of the mouth, with moderately long medial 

cusps and short cusplets on sides; tooth rows 28 to 33/22 to 32. Two dorsal fins, the base of the first 

extending forward of pelvic-fin origins as a low keel that reaches level of pectoral-fin bases but with 

insertion posterior to pelvic-fin origins; second dorsal fin 1/2 the size of first or less; anal fin present, rounded 

but not keel-shaped with its origin under rear 1/3 of second dorsal-fin base; caudal fin nearly or quite 1/2 of 

total length, strongly asymmetrical, with a deep subterminal notch but with the lower lobe hardly developed. 

Caudal peduncle not strongly depressed, without lateral keels or precaudal pits, but with dermal ridges 

extending forward onto sides. Supraorbital crests present on cranium, these laterally expanded.Intestinal valve 

of ring type. Colour: young below 60 cm with the back dark brown or blackish, with vertical yellow bars, 

spots and reticulations, and the underside of the head, abdomen and tail whitish; in subadults and adults the 

dark areas break up into scattered dark spots on a yellowish background, shading into the whitish ventral surface. 


None. The barbels, nasoral grooves, anterior mouth, teeth, anteriorly elongated dorsal fin, lateral ridges 

on the sides, greatly elongated caudal fin about 1/2 the total length, and distinctive colour patterns of young 

and adults separate this shark from all others in the area. 

Size: Maximum total length at least 2.35 m; possibly to 3.54 m. 

Habitat, biology, and fisheries: Common in inshore waters of the continental and insular shelves, often 

found on coral reefs, on or near the bottom. Oviparous, depositing eggs in rounded oblong egg cases 10 

to 17 cm long; size at birth between 20 and 36 cm. Probably nocturnal, feeds primarily on molluscs but 

also takes small fishes. Caught in bottom trawls, in floating 

and fixed bottom gill nets, and with longlines; utilized 

fresh and dried-salted for human consumption; livers are 

? 

processed for vitamins; fins dried for the oriental sharkfin 

? 

trade; also processed for fishmeal. 

Distribution: In the Indian Ocean and western Pacific from 

South Africa and the Red Sea eastward to Japan, Palau, 

western and northern Australia, and New Caledonia. 

References 



Dingerkus, G. 1986. Interrelationships of orectolobiform sharks (Chondrichthyes: Selachii). Proc. 2nd Int. Conf. Indo- 

Pacific Fish., 1986:227-245. 


Rhincodontidae 1263 


Rhincodon typus Smith, 1828 

RHINCODONTIDAE 

Whale sharks 


Frequent synonyms / misidentifications: Rhiniodon typus Smith, 1828 / None. 

FAO names: En - Whale shark; Fr - Requin baleine; Sp - Tiburón ballena. 

Diagnostic characters: Averylarge 

shark with cylindrical or moderately 

depressed body. Head very broad 

and flattened, with 5 large gill slits, the 

posterior 3 over the pectoral-fin bases; 

no gill rakers but filter grids of 

transverse bars and lobes across the 

internal gill slits; spiracles much 

smaller than eyes; nostrils with short, 

quadrate anterior nasal flaps, minute 

barbels, and shallow nasoral grooves; no nictitating eyelids; snout extremely short, 

truncated ; mouth nearly subterminal, very wide, transverse and short, not reaching 

backward to eyes; teeth very small and extremely numerous, similar in both jaws, not 

bladelike and with hooked cusps. Two dorsal fins, the first with rear 1/3 of base over 

pelvic-fin bases, the second less than half the size of first; anal fin present; caudal fin 

asymmetrical, crescentic, with a strong lower lobe but no subterminal notch. Caudal 

peduncle depressed, with a strong keel on each side continuing forward onto the barbel 

back and over the gill slits as a small ridge and flanked by 2 additional ridges above; head 

upper precaudal pit present. Supraorbital crests present on cranium, these laterally 

expanded. Valvular intestine of ring type. Colour: dark grey, reddish, or greenish grey above, with white or 

yellow spots and transverse stripes; white or yellowish below. 


None. The combination of characters such as the truncated snout, the transverse mouth in front of eyes, 

the numerous small teeth, the lateral ridges, the precaudal keels and the colour pattern distinguishes the 

whale shark from all other sharks in the area. 

Size: Maximum total length at least 12 m; possibly to 21.4 m. 

Habitat, biology, and fisheries: This huge pelagic filter feeder occurs singly or in schools, often at or near the 

surface, near shore or on the open sea. Ovoviviparous, can have as many as 300 fetuses. Feeds on small pelagic 

crustaceans, schooling fishes including anchovies, sardines, and even albacores, and squids. Often seen in a 

vertical position with head at or near the surface when feeding. Usually harmless, and permitting close approach 

by divers; rarely ramming small boats, possibly when excited by fish hooked from the boats, but more often struck 

by ships while basking at the surface. The whale shark was formerly of limited interest to fisheries worldwide, but 

recently became the subject of a high value fishery off Taiwan Province of China and the Pilippines for fins, flesh, 

and other products. Captured in gill nets and sometimes in trawls, and often harpooned. Utilized fresh and 

dried-salted for human consumption, liver processed for oil, fins used for soup base, and offal probably used for 

fishmeal. These sharks are increasingly popular as the subject of ecotouristic dive tours, as they migrate 

seasonally along coasts and concentrate in inshore tropical areas during part of the year. This shark is listed on 

the IUCN Red List of Threatened Animals (data deficient). 

Distribution: Circumglobal in the tropical and warm 

temperate Pacific and Atlantic Oceans, oceanic and coastal. 

References 

Compagno, L.J.V. 1984. FAO species catalogue. Vol. 4. Sharks 

of the world. An annotated and illustrated catalogue of 

shark species known to date. Part 1. Hexanchiformes to 

Lamniformes. FAO Fish. Synop., (125)Vol.4.Pt.1:249 p. 

Wolfson, F.H. 1986. Occurrences of the whale shark, 

Rhincodon typus Smith. Proc. 2nd Int. conf. Indo-Pacific Fish., 1986:208-226. 

Wolfson, F.H. and G. Notarabartolo di Sciara. 1981. The whale shark, Rhiniodon typus Smith, 1828; an annotated 

bibliography (Selachii Rhiniodontidae). Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano, 122(3-4):171-203.

1264 Sharks 

Odontaspididae ODONTASPIDIDAE 

Sand tiger sharks 

by L.J.V. Compagno and V.H Niem 

Diagnostic characters: Large sharks. Head with 5 medium-sized gill slits, all in front of pectoral-fin 

bases, their upper ends not extending onto dorsal surface of head; gill arches without rakers; 

spiracles present but very small; no nasal barbels or nasoral grooves; eyes small or moderately large, 

without nictitating eyelids; snout conical or moderately depressed, not blade-like; mouth very long and 

angular, extending well behind eyes when jaws are not protruded; lower labial furrows usually present 

at mouth corners; anterior teeth enlarged, with long, narrow, sharp-edged but unserrated cusps and small 

basal cusplets (absent in young of at least 1 species), the upper anteriors separated from the laterals by 

a gap and tiny intermediate teeth. Two moderately large, high dorsal fins, the first originating well in 

advance of the pelvic fins, the second as large as or somewhat smaller than the first; anal fin as large 

as second dorsal fin or slightly smaller; caudal fin short, asymmetrical, with a strong subterminal notch and 

a short but well-marked ventral lobe. Caudal peduncle not depressed, without keels; a deep upper 

precaudal pit present but no lower pit. Intestinal valve of ring type, with turns closely packed like a stack 

of washers. Colour: grey or grey-brown above, white or lighter below, with round or oval spots on at least 

1 species. 

eys small to 

moderately large 

gill slits not extending 

high on head 2 nd dorsal fin 

large 

gill slits in front of 

pectoral-fin origin 

Habitat, biology and fisheries: These are wide-ranging, tropical to cool-temperate sharks, found inshore 

and down to moderate depths on the edge of the continental shelves and around some oceanic islands, 

but not oceanic. Development is ovoviviparous. They feed on small bony fishes other sharks, squids, and 

occasionally bottom crustaceans. Normally inoffensive, but potentially dangerous if provoked. In the area, 

at least 1 species is regularly caught for food, liver oil, and processed for fishmeal. 


Pseudocarchariidae: body slimmer, gill slits higher 

and reaching onto dorsal sides of head, eyes larger, 

no true labial furrows, dorsal and anal fins lower, a 

weak lateral keel on caudal peduncle and both 

upper and lower precaudal pits present. 

Proscylliidae, Triakidae, Hemigaleidae, and 

Carcharhinidae: nictitating eyelids present, anterior 

teeth not greatly enlarged, no intermediate teeth 

between anteriors and laterals, intestinal valve of 

spiral or scroll type. 

anal fin 

large 

intestinal valve of ring type 

upper 

precaudal pit 

no lower 

precaudal pit 

large higher lower 

Pseudocarchariidae 

keel 

lower precaudal pit

Odontaspididae 1265 

Key to the species of Odontaspididae occurring in the area 

1a. Snout short and somewhat flattened (Fig. 1a); eyes very small; 3 rows of anterior teeth 

on either side of upper symphysis (Fig. 2b); dorsal and anal fins about equal in size, first 

dorsal fin closer to pelvic fin than to pectoral-fin bases (Fig. 3) . . . . . . . . . . Carcharias taurus 

1b. Snout longer, bulbous and conical (Fig. 1b); eyes relatively large; 2 rows of large anterior 

teeth on either side of upper symphysis (Fig. 2b); first dorsal fin markedly larger than 

the second, closer to pectoral than to pelvic-fin bases; second dorsal fin considerably 

larger than anal fin (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Odontaspis ferox 

intermediate 

anterior lateral 

a) Carcharias taurus b) Odontaspis feros 


about equal 




? Odontaspis ferox (Risso, 1810) 

References 

Compagno. L.J.V. 1984. FAO Species catalogue. Vol. 4. Sharks of the world. An annotated and illustrated catalogue of 

shark species known to date. Part 1. Hexanchiformes to Lamniformes. FAO Fish Synop., (125)Vol.4, Pt.1:249 p. 


gap 

a) Carcharias taurus 

anterior intermediate lateral 

b) Odontaspis ferox 

Fig. 2 upper teeth of left side 

Fig. 3 Carcharius taurus Fig. 4 Odontaspis ferox

1266 Sharks 


Frequent synonyms / misidentifications: Odontaspis taurus (Rafinesque, 1810); Eugomphodus taurus 

(Rafinesque, 1810) / Odontaspix ferox (Risso, 1810). 

FAO names: En - Sandtiger shark; Fr - Requin taureau (= Requin sable tacheté, Fishing Area 31); Sp -Toro 

bacota (= Pez toro). 

Diagnostic characters: A large shark. Head 

with 5 medium to large gill slits, all in front of 

pectoral-fin bases, no gill rakers; snout very 

short, moderately flattened; no nasal 

barbels or nasoral grooves; eyes small, 

without nictitating eyelids; mouth very 

long and angular, extending well behind 

eyes; anterior teeth in 3 rows on either side 

of symphysis, large, with long, narrow, 

hooked, sharp-edged but non-serrated 

cusps and usually 1 short cusplet on each side; upper anteriors separated from the smaller laterals by a 

single row of tiny intermediate teeth (lacking in lower jaw); lower anteriors separated at front by 2 rows of 

small symphyseal teeth (generally lacking in upper jaw). Two dorsal fins, the base of first just in front of 

pelvic-fin bases and well posterior to pectoral fins; second dorsal fin about as large as first dorsal and 

anal fins; caudal fin short, strongly asymmetrical, with a pronounced subterminal notch and a short ventral 

lobe. No keels on caudal peduncle, but with a strong upper precaudal pit. Intestinal valve of ring type. 

Colour: light grey-brown above, white below, often with round or oval, yellow, or yellow-brown spots. 

Size: Maximum total length about 3.18 m; adults between 2.2 to 3 m. 

Habitat, biology, and fisheries: A common littoral shark found inshore from the surf zone and in shallow bays 

to at least 191 m on the outer continental shelves in tropical and (mostly) temperate waters. It commonly lives 

near or on the bottom but occurs at midwater and at the surface. It is a slow but strong swimmer that can readily 

halt and hover motionless in midwater, and is the only known shark to gulp and store air in its stomach to maintain 

neutral buoyancy while swimming. Ovoviviparous, with litters of 2 young recorded. Feeds on a wide variety of 

bony fishes, small sharks, rays, squids, crabs, and lobsters. This species previously had a bad reputation as a 

“maneater” in Australian waters (“gray nurse shark”), probably in large part by confusion with certain requiem 

sharks (Carcharhinidae) and with the white shark. It is apparently mostly inoffensive, but occasionally can 

become aggressive and nips divers without attempting to feed. It is caught by a large variety of fishing gear 

including line gear, bottom gill nets, and in pelagic and bottom trawls and is utilized for its flesh, liver oil, fins, 

and hides for leather. It is now protected in Australian waters after suffering local declines due to divers killing 

this easily-approached shark for “sport”with powerheads.This shark is listed on the IUCN Red List ofThreatened 

Animals (vulnerable). 

Distribution: Found in all warm seas 

except perhaps the eastern Pacific. In the 

Indo-West Pacific, off South Africa and in 

the Red Sea westwards to Japan, Korea, 

and Australia. In the area, along the entire 

northern coast of Australia, off the south 

coast of Viet Nam, and Indonesia (Aru and 

Obi Islands, possibly more widespread). 

Nominally recorded as Carcharias 

tricuspidatus from the Philippines. 

? 

intermediate 

anterior lateral posterior 

symphyseal 

gap 

upper and lower teeth of left side

Odontaspididae 1267 

Odontaspis ferox (Risso, 1810) 

Frequent synonyms / misidentifications: Odontaspis herbsti Whitley, 1950 / Carcharias taurus 

Rafinesque, 1810. 

FAO names: En - Smalltooth sand tiger; Fr - Requin féroce; Sp - Solrayo. 

Diagnostic characters: A large shark. Head 

with 5 medium to large gill slits, all in front of 

pectoral-fin bases; no gill rakers; snout 

moderately elongated, bulbously conical; 

no nasal barbels or nasoral grooves eyes 

moderately large, without nictitatinq 

eyelids; mouth very long and angular, 

extending well behind eyes; anterior teeth 

moderately large. with long, narrow, hooked, 

sharp-edged but non-serrated cusps and 2or 

3 moderately long cusplets on each side, separated in front by 2 rows of small symphyseal teeth in both 

jaws; upper anteriors set in 2 rows on either side of symphysis and separated from the smaller laterals by 

3 or 4 rows of tiny intermediate teeth; lower anteriors set in 3 rows on either side of symphysis and not followed 

by small intermediate teeth. Two dorsal fins, the first large and situated closer to the pectoral fins than to 

the pelvic fins, its free rear tip well ahead of pelvic-fin origins, the second dorsal fin smaller than the first 

and usually slightly larger than anal fin; caudal fin short, strongly asymmetrical, with a pronounced subterminal 

notch and a short ventral lobe. No keels on caudal peduncle, but a strong upper precaudal pit. Intestinal valve 

of ring type. Colour: grey above, paler below, tips of dorsal, anal, pectoral, and pelvic fins may be dark-tipped 

in young; dark spots present on sides in some individuals. 


Habitat, biology, and fisheries: An uncommon, little-known, primarily deep-water species found at depths 

between 15 and 420 m from inshore waters to the upper continental and insular slopes. Probably ovoviviparous. 

Bottom-dwelling, feeds on small bony fishes, squids, and crustaceans. The large oily liver presumably has a 

hydrostatic function. This species is not implicated in attacks on people. Taken with bottom gill nets, line gear and 

bottom trawls, primarily in the Mediterranean Sea and off Japan; used for its meat and for its squalene-rich liver. 

Distribution: Known from the Mediterranean Sea and eastern Atlantic; Indo-West Pacific off South Africa and 

Maldives, Madagascar, southern Japan, Australia (New South Wales and northwestern Australia just adjacent 

to the area), and New Zealand; Central Pacific off Hawaii; and eastern Pacific off southern California and Baja 

California. Probably has a wider range than is known and is to be expected inside the area. 

Remarks:Unspotted individuals 

have been distinguished as 

Odontaspis herbsti, but 

apparently presence of spots is 

a matter of individual variation in 

1 species. Carcharias taurus is 

also variable in having or lacking 

spots. A somewhat similar 

deepwater and possibly oceanic 

species, Odontaspis noronhai, 

has been recorded off Hawaii in 

deep water and may eventually 

be recorded for the area. 

anterior intermediate 

lower 

symphyseal 

anterior 

? 

? 

lateral 

? 

posterior 

lateral 

posterior 

upper and lower teeth of left side

1268 Sharks 


Pseudocarcharias kamoharai (Matsubara, 1936) 

PSEUDOCARCHARIIDAE 

Crocodile sharks 


Frequent synonyms / misidentifications: Odontaspis kamoharai (Matsubara, 1936) / None. 

FAO names: En - Crocodile shark; Fr - Requin crocodile; Sp - Tiburón cocodrilo. 

Diagnostic characters: A small relatively slender 

anterior 

shark. Head with 5 large gill slits, all in front of pectoral-fin 

bases, their upper ends extending onto dorsal surface of 

head; no gill rakers; spiracles usually present but very 

small; no nasal barbels or nasoral grooves; eyes very 

large, without nictitating eyelids; snout conical (not 

greatly elongated or flattened and blade-like); mouth very 

long and angular, extending well behind eyes; notrue 

labial furrows; anterior teeth very large, with long, narrow, 

hooked, sharp-edged but unserrated cusps and no cusplets, set in 2 rows on either side of symphysis in 

both jaws, and not separated in front by small symphyseal teeth; upper anteriors separated from the smaller 

laterals by a gap and tiny intermediate teeth. Two low dorsal fins, the first about midway between the pectoral 

and pelvic fins, and well in front of pelvic fin bases, the second somewhat smaller than the first, but larger 

than anal fin; caudal fin short, strongly asymmetrical, with a pronounced subterminal notch and a short 

ventral lobe. Caudal peduncle slightly depressed, with a low keel on each side and upper as well as lower 

precaudal pits. Intestinal valve of ring type, with close-set turns resembling a stack of washers. Colour: 

light or dark grey above, lighter below, fins white-edged, sometimes small white spots on body and a white 

blotch between the mouth and gill slits. 

intermediate 

ventral view 

of head 

lateral 

posterior 

anterior 

lateral 

teeth of left side 

posterior 


None. The combination of the characters described above separates this species from all other sharks. 

Size: Maximum total length to about 1.1 m; commonly between 75 cm and 1 m. 

Habitat, biology, and fisheries: A rare to locally abundant oceanic and possibly mesopelagic shark, 

usually found offshore from the surface to at least 300 m. Its habits are little known. Ovoviviparous. Feeds 

on oceanic fishes, cephalopods, and crustaceans. Most frequently caught by pelagic longline fisheries, but 

usually discarded due to its small size; utilized for its large, squalene-rich liver. 

Distribution: Possibly circumtropical; known from the 

eastern and southwestern Atlantic, southwestern and 

perhaps northeastern Indian Ocean, northwestern, 

central and eastern Pacific. In the Indo-West Pacific, off 

South Africa and the Mozambique Channel near 

Madagascar, possibly the Bay of Bengal, New Zealand, 

Indonesia (Java, Sumatra), Taiwan Province of China, 

Korea, Japan, Australia (Queensland), and Coral Sea. 

References 






Alopiidae 1269 

ALOPIIDAE 

Thresher sharks 


Diagnostic characters: Large sharks. Trunk and precaudal tail cylindrical, not depressed and without 

lateral ridges; precaudal tail much shorter than trunk. Head not expanded laterally, not depressed; 

5 small to medium-sized gill slits present, the last 2 behind pectoral-fin origins, their upper ends not 

expanded onto upper surface of head; no gill rakers or sieves on internal gill slits; spiracles present and 

minute; nostrils without barbels, nasoral grooves, or circumnarial grooves, well separated from mouth; eyes 

on sides of head, without nictitating lower eyelids; snout moderately long, bluntly conical, not flattened and 

without lateral teeth or barbels; mouth small but arched and elongate, extending well behind eyes; labial 

furrows present on lower jaw only or absent, when present not reaching front of mouth; teeth small, 

blade-like and compressed, with erect to oblique cusps and cusplets very small or absent; anterior teeth 

in upper jaw slightly larger than lateral teeth and sometimes separated from them by a row of smaller 

intermediate teeth on each side. Two dorsal fins, without spines, the first moderately large, high and angular, 

much shorter than the caudal fin, and with its base located over the interspace between pelvic and 

pectoral-fin bases; second dorsal fin low, minute, and less than 1/10 the size of the first dorsal fin; anal fin 

present, very small, with its origin under or behind the second dorsal-fin insertion; caudal fin strongly 

asymmetrical, the upper lobe enormously enlarged, about 1/2 the total length and with a subterminal 

notch, and an undulated or rippled dorsal margin, the lower lobe short but strong; vertebral axis of caudal 

fin raised above body axis. Caudal peduncle not depressed, without keels; precaudal pits present. Intestinal 

valve of ring type. Colour: bluish, blackish, grey, or brown above, shading to white or grey below. 

caudal fin very long 

terminal lobe 


minute 

trunk 

anal fin 

precaudal 

tail 

Habitat, biology, and fisheries: These are active, strong-swimming, pelagic, coastal and deep-water 

sharks, with the young of 1 species occurring close inshore and inside bays. They feed mainly on small to 

moderately large schooling fishes and squid, which may be herded and stunned by the long, strap-like tail. 

Threshers are circumtemperate and tropical in all warm oceans. This monogeneric family comprises only 

3 or 4 species worldwide, 3 of which occur in the area. Thresher sharks form an important component of 

the oceanic shark fishery, particularly because of their high-quality meat which is utilized fresh, frozen, 

smoked, and dried-salted. Their fins are used for shark-fin soup, livers for vitamin extraction, and hides for 

leather. Thresher sharks are primarily captured by offshore longline fisheries but also offshore and near 

shore with line gear (including rod and reel) and fixed bottom gill nets. 


Stegostomatidae: this is the only other family of sharks 

with the caudal fin about as long as the body; it differs 

from Alopiidae in numerous characters, including its 

striped or barred colour pattern, nasal barbels, 

transverse mouth in front of eyes, small tricuspid teeth, 

broad rounded pectoral fins, first dorsal fin over pelvic-fin 

bases, larger second dorsal and anal fins, broad upper 

lobe on caudal fin, no ventral caudal-fin lobe, and axis of 

caudal fin not raised. 

No other sharks in the area have the caudal fin about 1/2 

the total length. 

caudal fin 


subterminal 

notch

1270 Sharks 

Key to the species of Alopiidae occurring in the area 

1a. Head nearly flat between eyes; a deep horizontal groove on nape of each side above 

gills; eyes very large, with orbits expanded onto dorsal surface of head; teeth larger, 

less than 25 rows in each jaw; first dorsal-fin base closer to pelvic-fin bases than 

pectoral-fin bases (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alopias superciliosus 

1b. Head strongly arched between eyes; no horizontal groove or an inconspicuous one on nape 

of each side; eyes smaller, with orbits not expanded onto dorsal surface of head; teeth 

smaller, 29 or (usually) more rows in each jaw; first dorsal-fin base about equidistant 

between pectoral and pelvic-fin bases or closer to pectoral-fin bases (Figs 2 and 3) . . . . . . . . → 2 

eyes directed upward 

groove 

a) dorsal view of head 

grooves 

Fig. 1 Alopias superciliosus 

2a. Sides above pectoral-fin bases dark, without an extension of the white abdominal area; 

head narrow, snout more elongated, forehead nearly straight; labial furrows absent; 

pectoral fins nearly straight and broad-tipped; distance between pelvic and caudal-fin 

bases shorter than prebranchial length; terminal lobe of caudal fin shorter, its length 

from subterminal notch to caudal tip about equal to second dorsal-fin base (Fig. 2) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Alopias pelagicus 

2b. Sides above pectoral-fin bases marked with a white patch extending forward from the 

abdominal area; head broad, snout shorter, forehead strongly arched; labial furrows 

present; pectoral fins falcate and narrow-tipped; distance between pelvic and caudal-fin 

bases greater than prebranchial length; terminal lobe of caudal fin longer, its length from 

subterminal notch to caudal tip over twice second dorsal-fin base (Fig. 3) . . . . . Alopias vulpinus 

white here 

falcate and 

narrow-tipped 

Fig. 2 Alopias vulpinus 





Alopias vulpinus (Bonnaterre, 1788) 

dark here 

straight and 

broad tipped 


Fig. 3 Alopias pelagicus 

References 

Compagno, L.J.V. 1984. FAO Species catalogue. Vol. 4. Sharks of the world. An annotated and illustrated catalogue of 


Last, P.R. and J.D. Stevens. 1993. Sharks and rays of Australia. Australia, CSIRO, 513 p.



Frequent synonyms / misidentifications: None / Alopias superciliosus (Lowe, 1839); A. vulpinus 

(Bonnaterre, 1788). 

FAO names: En - Pelagic thresher; Fr - Renard pélagique; Sp - Zorro pelágico. 

Diagnostic characters: A large shark. Head with 5 medium-sized gill slits, the last 2 above pectoral-fin 

bases; a weak horizontal groove on nape on each side from level of mouth to pectoral fins; no nasal barbels 

or nasoral grooves on nostrils; snout moderately long and conical; forehead nearly straight in lateral 

view, broadly arched between eyes; head narrow; no nictitating eyelids; eyes moderately enlarged in 

adults and subadults, but greatly enlarged in young, not expanded onto dorsal furface of head; mouth 

moderately long and semicircular, placed below eyes, with labial furrows rudimentary or absent; teeth small, 

more than 29 rows in each jaw, sharp-edged, with a single, narrow, nearly erect or distally oblique cusp 

and often a distal cusplet; anterior teeth not greatly enlarged, uppers separated from the large laterals by 

smaller intermediate teeth. Two dorsal fins, the first moderately large and located about equidistant 

between the pectoral and pelvic-fin bases or slightly closer to the pectoral-fin bases; second dorsal 

fin minute and positioned well ahead of the small anal fin; pectoral fins narrow, long and nearly straight, 

broad-tipped, and not falcate; upper lobe of caudal fin very long and strap-like, about as long as the rest 

of the shark; lower lobe short but strong; terminal lobe very small. Upper precaudal pit present but caudal 

keels absent. Intestinal valve of ring type. Colour: bluish or grey above, white below, with a silvery sheen 

in gill region; white colour from belly not expanded over pectoral-fin bases. 

Size: Maximum total length at least 3.3 m (adult females). 

Habitat, biology, and fisheries: A little-known species, primarily oceanic and epipelagic, but sometimes 

caught near-shore, ranging from the surface to a depth of at least 150 m. An active, strong-swimming 

species. Ovoviviparous, with at least 2 young; apparently a uterine cannibal like other species of Alopias. 

Presumably feeds on small fishes and squid, but no details are known. Harmless to people. Formerly 

exploited by the longline fishery in the northwestern Indian Ocean (primarily by Russia), but is also fished 

in the Central and eastern Pacific. Utilized for its meat (for human consumption), liver oil for vitamin 

extraction, hides for leather, and fins for shark-fin soup. 

Distribution: Wide-ranging in 

the tropical and subtropical 

Indo-Pacific.

1272 Sharks 


Frequent synonyms / misidentifications: Alopias profundus Nakamura, 1935 / Alopias pelagicus 

Nakamura, 1935; A. vulpinus (Bonnaterre, 1788). 

FAO names: En - Bigeye thresher; Fr - Renard à gros yeux; Sp - Zorro ojón. 

eyes directed upward 

grooves 



bases; a deep horizontal groove on nape on each side from the level of mouth to pectoral fins; no nasal 

barbels or nasoral grooves on nostrils; snout moderately long and conical; profile of forehead distinctly 

indented over eyes; interorbital space nearly flat; no nictitating eyelids; eyes very large, expanding 

onto dorsal surface of head, permitting upward vision; mouth moderately long and semicircular, placed 

below the eyes, with rudimentary labial furrows; teeth moderately large, less than 25 rows in upper or 

lower jaws, sharp-edged, with a single, broad, straight or posteriorly curved cusp and no cusplets; anterior 

teeth not greatly enlarged, uppers not separated from the large laterals by smaller intermediate teeth. Two 

dorsal fins, the first moderately large and located just in front of the pelvic-fin origins, closer to the 

pelvic fins than to the pectoral fins; second dorsal fin minute and positioned well ahead of the small anal 

fin; pectoral fins very narrow, long and falcate, broad-tipped; upper lobe of caudal fin very long and strap-like, 

almost or quite equal to the length of rest of shark; lower lobe short but well developed. Upper precaudal 

pit present but caudal keels absent. Intestinal valve of ring type. Colour: purplish grey above, cream below, 

posterior edges of pectoral and pelvic fins and sometimes first dorsal fin dusky; light colour of abdomen 

not expanded over pectoral-fin bases. 

Size: Maximum total length about 4.6 m; commonly between 3 and 4 m. 

Habitat, biology, and fisheries: Found in coastal waters over the continental shelves, sometimes close 

inshore in shallow waters, and on the high seas far from land, in deep water down to at least 500 m. 

Apparently strong-swimming. Ovoviviparous, with uterine cannibalism, number of young usually 2 per litter, 

but sometimes up to 4. Feeds on pelagic fishes (lancetfishes, clupeoids, scombroids, and small billfishes) 

and bottom fishes (hakes); also squids. Apparently stuns its prey with its long caudal fin, as individuals are 

often tail-hooked on longlines. Apparently harmless to people. Caught in oceanic longline fisheries; 

especially important areas for these fisheries are the North Atlantic, northwestern Indian Ocean, and the 

Central and eastern Pacific. The species is also taken in fixed bottom and pelagic gill nets, in trawls, and 

with sportsfishing gear (rod 

and reel). Its meat is utilized 

fresh, smoked, and driedsalted 

for human consumption, 

its liver oil is processed for 

vitamins, its skin for leather, 

and fins for shark-fin soup. 

Distribution: Virtually circumglobal 

in tropical and warm 

temperate seas.


Alopias vulpinus (Bonnaterre, 1788) 

Frequent synonyms / misidentifications: None / Alopias pelagicus Nakamura, 1935; A. superciliosus 

(Lowe, 1839). 

FAO names: En - Thresher shark; Fr - Renard; Sp - Zorro. 


bases; no grooves on nape; no gill rakers; no nasal barbels or nasoral grooves on nostrils; snout short 

and conical; forehead broadly convex in lateral view, not indented at nape; no nictitating eyelids; eyes 

moderately large, not expanded onto dorsal surface of head; mouth short and semicircular, below 

eyes, with short lower labial furrows; teeth small, usually over 29 rows in upper and lower jaws, sharpedged, 

with a single, broad, straight or posteriorly curved cusp and usually no cusplets; anterior teeth not 

greatly enlarged, uppers usually separated from the laterals by a small intermediate tooth. Two dorsal 

fins, the first moderately large, with its base well ahead of the pelvic-fin bases and farther from them 

than from the pectoral-fin bases; second dorsal fin minute and positioned just in front of the small anal 

fin; pectoral fins very long and falcate, with narrowly rounded (small juveniles) to acutely pointed, 

narrow tips; upper lobe of caudal fin very long and strap-like, about as long as, or longer than, rest of 

shark; lower lobe short but well-developed. Upper precaudal pit present but caudal keels absent. Intestinal 

valve of ring type. Colour: brown, grey, blue-grey, or blackish on back and underside of snout, lighter on 

sides, and abruptly white below; a white area extends from the abdomen over the pectoral-fin bases; 

pectoral-, pelvic-, and dorsal fins blackish, white dots sometimes present on pectoral-, pelvic-, and 

caudal-fin tips. 

Size: Maximum total length about 5.5 m; commonly between 4.3 and 4.9 m; apparently larger than Alopias 

superciliosus and A. pelagicus. 

Habitat, biology, and fisheries: Coastal over the continental and insular shelves and epipelagic far from 

land in temperate to tropical waters; young often close inshore and in shallow bays, from the surface down 

to 370 m. An active, strong-swimming shark, sometimes leaping out of the water. Ovoviviparous and 

apparently a uterine cannibal, number of young 2 to 4 per litter (usually 2). Feeds mostly on small schooling 

fishes, including mackerels, bluefishes, clupeids, needlefishes, lancetfishes, and lanternfishes; also squids, 

octopuses and pelagic crustaceans, and rarely seabirds. Herds and stuns its prey with its long, whip-like 

caudal fin, and is often caught on longlines by being tail-hooked. Apparently harmless to people, though 

the size of adults of this species should invite respect. Caught in oceanic longline fisheries; especially 

important areas for these fisheries are or were the northwestern Indian Ocean and the Central Pacific. Also 

fished with anchored bottom and surface gill nets, floating gill nets and sportfishing gear (rod and reel). 

The meat is highly prized fresh for human consumption but is also eaten smoked and dried-salted; the fins 

are valuable for shark-fin soup; 

the hide is usable for leather 

and the liver oil can be 

processed for vitamins. 

Distribution: Virtually circumglobal 

in temperate to tropical 

waters.

1274 Sharks 

LAMNIDAE 

Mackerel sharks, makos, white sharks, porbeagles 


Diagnostic characters: Large-sized sharks with a fusiform body. Head with 5 gill slits, all in front of 

pectoral-fin origins; gill arches without rakers; no nictitating eyelids; teeth long and few in number, 

awl-or blade-like, with a single cusp. Two dorsal fins, the first much shorter at base than caudal fin and 

far in advance of pelvic fins; second dorsal fin and anal fin much smaller than first dorsal fin; caudal 

fin lunate, less than 1/3 of total length. Caudal peduncle strongly depressed dorsoventrally and 

expanded laterally, with a prominent keel on each side, extending well out on caudal fin. Intestinal 

valve of ring type. Colour: back greyish blue to black, or brownish; belly white. 

Habitat, biology, and fisheries: Mackerel sharks inhabit temperate and tropical waters (oceanic as well 

as coastal) throughout the world. They are very fast swimmers and voracious predators, feeding mainly on 

fish and squid, but also other sharks, batoids, marine mammals, sea birds, and carrion; some species are 

dangerous to man. Mackerel sharks are often used for food or for production of liver oil, fishmeal and other 

shark products. 


Cetorhinidae: the basking shark, Cetorhinus 

maximus (Gunnerus, 1765) is known to occur close 

to the area (western North Pacific southwards to 

Taiwan Province of China as well as southwestern 

Australia) and may eventually be found in the area. It 

can be distinguished from members of Lamnidae by 

having much longer gill openings, extending from 

upper surface of head to throat; gill rakers well 

developed on internal gill openings; teeth minute and 

Cetorhinidae 

hooked, not blade-like; anal fin and second dorsal fin 

larger; and size of adults larger (9 m or more). 

Rhincodontidae: body with several prominent dermal ridges on either side; last gill slit well behind 

pectoral-fin origin; snout squared off anteriorly; mouth nearly terminal; at least half of first dorsal-fin base 

posterior to pelvic-fin origins; gill arches connected by masses of spongy tissue; and a spotted and striped 


All other shark families: caudal fin strongly asymmetrical and not lunate, the upper lobe extending far 

beyond lower lobe; caudal peduncle not greatly flattened dorsoventrally. 

Rhincodontidae 

other shark families (e.g. Carcharhinidae)

Lamnidae 1275 

Key to the species of Lamnidae occurring in the area 

1a. Upper teeth triangular with serrate edges (Fig. 1a); origin of first dorsal fin opposite or 

slightly anterior to inner corners of pectoral fins when the latter are laid back; anal-fin 

origin posterior to second dorsal-fin base (Fig. 2) . . . . . . . . . . . . . Carcharodon carcharias 

1b. Upper teeth with smooth-edged cusps (Fig. 1b, c); origin of first dorsal fin posterior to 

inner corners of pectoral fins when the latter are laid back; anal-fin origin below midbase 

or insertion of second dorsal-fin base (Figs 3 and 4) . . . . . . . . . . . . . . . . . . . . . . . → 3 

serrations 

smoothedged 

a) Carcharodon 

carcharias 

b) Isurus 

paucus 

c) Isurus 

oxyrinchus 

Fig. 1 upper tooth Fig. 2 Carcharodon carcharias 

2a. Snout usually acutely pointed (Fig. 5a); cusps of upper and lower anterior teeth recurved 

at bases but with tips reversed and curving outward; pectoral fins considerably shorter 

than head, relatively narrow-tipped in young, acutely pointed in adults; origin of anal fin 

about under midbase of second dorsal fin (Fig. 3); underside of snout and mouth white 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isurus oxyrinchus 

2b. Snout narrowly to bluntly (usually not acutely) pointed (Fig. 5b); cusps of upper and 

lower anterior teeth straighter, with tips not reversed; pectoral fins about as long as head, 

relatively broad-tipped in young and adults; origin of anal fin about under insertion of 

second dorsal fin (Fig. 4); underside of snout and mouth dusky . . . . . . . . . . . . Isurus paucus 

Fig. 3 Isurus oxyrinchus Fig. 4 Isurus paucus 






pointed 

broader, less 

pointed 

a) Isurus oxyrinchus b) Isurus paucus 


References 

Bigelow, H.B. and W.C. Schroeder. 1948. Sharks. Mem. Sears Found. Mar. Res., (1):56-576 



Garrick J.A.F. 1967. Revision of sharks of genus Isurus with description of a new species (Galeoidea, Lamnidae). Proc. 

U.S. Natl. Mus., 118(3537):663-690. 

Last, P.R. and J.D. Stevens. 1993. Sharks and rays of Australia. Australia, CSIRO, 513 p.

1276 Sharks 



FAO names: En - Great white shark; Fr - Grand requin blanc; Sp - Jaquentón blanco (= Jaquetón). 


Diagnostic characters: A very large shark with a 

fusiform, usually heavy body and a moderately long, 

bluntly pointed snout. Head with 5 long gill slits, all in 

front of pectoral-fin origins; gill arches without rakers; 

spiracles very small; mouth long and broadly rounded; 

teeth very large and relatively few, narrower in the lower 

than in the upper jaw, pointed backwards, with a single 

broad cusp and strong serrations at most sizes 

(irregular in individuals below 1.5 m length, and with 

intermediate 

anterior 

anterior 

lateral 


posterior 

posterior 

cusplets present up to about 2 m length, but lost in larger individuals); anterior teeth greatly enlarged in 

both jaws, in 2 rows on either side of symphysis, broadly triangular and compressed, not recurved; intermediate 

and first few lateral teeth a little smaller, the intermediate ones less differentiated from the anterior and lateral 

teeth than in other members of the family.Two dorsal fins, the first large, originating over inner margins of pectoral 

fins, the second very small; pectoral fins shorter than head and falcate; anal-fin origin posterior to rear end 

of second dorsal-fin base; caudal fin lunate, its lower lobe strongly developed. Caudal peduncle very much 

flattened dorsoventrally, expanded laterally, with a prominent keel on either side extending well out on 

caudal fin but with no secondary keel on the fin. Colour: grey-brown, dark grey, blue-grey, blackish, light grey 

or grey-white above, white below, fins with dusky margins below, black tips on underside of pectoral fins, a black 

spot present or lacking on pectoral-fin axils. 

Size: Maximum total length possibly 6.4 to 7.2 m or more (a record of 10.98 m later proved incorrect); 

commonly between 5 and 6 m. 

Habitat, biology, and fisheries: An inshore, offshore, and oceanic species, often occurring on the continental 

shelves off island and reefs, in enclosed shallow bays and off beaches; recorded from the surface and the 

intertidal down to 1 280 m. A powerful, strong swimmer. Ovoviviparous, possibly up to 10 fetuses in a litter. A 

powerful, highly opportunistic apical predator, feeding on a wide variety of marine animals, including other 

sharks, rays, chimaeras, bony fishes, seals, sea lions, dolphins, porpoises, sea birds, squid, crustaceans, and 

carrion. One of the most dangerous sharks, responsible for a number of unprovoked attacks on swimmers, 

divers, surfers, and boats.Of limited interest to fisheries, mostly taken as a bycatch with longlines, hook-and-line, 

fixed bottom gill nets, fish traps, herring weirs, and trammel nets, harpoons, and even bottom and pelagic trawls, 

as well as purse seines.Utilized fresh, dried-salted, and smoked for human consumption;the liver oil is extracted 

for vitamins; the carcass used for fishmeal; the skin for leather; the fins for shark-fin soup; and the teeth and 

jaws for decorations, with properly prepared 

large jaws bringing a high price. Protected 

in South Africa, Namibia, Israel, the USA, 

and Australia, threatened by targeted and 

bycatch fisheries. This shark is listed on the 

IUCN Red List of Threatened Animals 

(vulnerable). 

Distribution: Cosmopolitan in boreal to 

tropical seas, but apparently more 

abundant in cool to warm-temperate 

waters. 

lateral

Lamnidae 1277 


Frequent synonyms / misidentifications: Isurus glaucus (Müller and Henle, 1839) / Isurus paucus Guitart 

Manday, 1966. 

FAO names: En - Shortfin mako; Fr - Taupe bleu; Sp - Marrajo dientuso (= Marrajo). 


upper and lower teeth in 

natural position at tip of 

mouth (lateral view) 

anterior 

intermediate 

anterior lateral posterior 

lateral 


posterior 

Diagnostic characters: A large shark with a fusiform and moderately slender body and a long and acutely 

pointed snout. Head with 5 long gill slits, all in front of pectoral-fin origins; gill arches without rakers; 

spiracles very small; mouth broadly rounded and notably long; teeth strong and relatively few, alike in 

both jaws, backward-pointing, somewhat flexuous in outline, smooth-edged, with a single cusp; the first 

2 in each jaw much the largest, recurved at base but curve-reversed at tips. Two very unequal dorsal fins, 

the first comparatively large, its origin posterior to inner corners of pectoral fins when latter are 

laid back, its apex bluntly rounded (young) to acutely pointed (adults); pectoral fins moderately long 

(shorter than head) and falcate; anal-fin origin below about middle of second dorsal-fin base; caudal 

fin lunate, its lower lobe strongly developed. Caudal peduncle very much flattened dorsoventrally, but 

expanded laterally, with a prominent keel on each side extending well out on caudal fin. Colour: 

back grey-blue, occasionally deep blue; belly white. 

Size: Maximum total length to about 4 m; commonly to 2.7 m. 

Habitat, biology, and fisheries: An oceanic and coastal species, usually in surface waters, approaching 

close inshore, but also in deeper water to at least 150 m. Perhaps the most active and strong-swimming 

of sharks, renowned for leaping out of the water, especially when hooked. Ovoviviparous, number of young 

in a litter 1 to 6, rarely 10. Feeds heavily on schooling fishes (mackerels, jacks, herrings, etc.), also small 

sharks and attacks larger species such as tunas and swordfishes. An aggressive, dangerous shark, 

responsible for unprovoked attacks on swimmers and boats; hooked individuals fight very hard and may 

leap into the boats of anglers. An important species for longline fisheries, because of its high-quality meat, 

and also it is also famed as one of the finest game fishes, highly prized by sport anglers. Caught in gill 

nets, and on pelagic longlines, and 

hook-and-line. The meat is utilized fresh, 

frozen, smoked, and dried-salted for 

human consumption; the oil is extracted for 

vitamins; the fins used for shark-fin soup; 

the hides processed into leather and the 

jaws and teeth used for ornaments. 

Distribution: Cosmopolitan in 

warm-temperate and tropical seas.

1278 Sharks 


Frequent synonyms / misidentifications: Isurus alatus Garrick, 1966 / Isurus oxyrinchus Rafinesque, 1810. 

FAO names: En - Longfin mako; Fr - Petit taupe; Sp - Marrajo carite. 

Diagnostic characters: A large shark with a 

fusiform and moderately slender body and 

a long, pointed snout. Head with 5 long gill 

slits, all in front of pectoral-fin origins; gill 

arches without rakers; spiracles very small; 

mouth long and broadly rounded; teeth large 

and relatively few, alike in both jaws, pointed 

backward, with a single cusp, but without 

cusplets or serrations; anterior teeth greatly enlarged in both jaws, in 2 rows on each side, cusps 

recurved at bases but not reversed at tips. Two dorsal fins, the first large, originating posterior to free 

rear tips of pectoral fins, with a bluntly rounded apex, the second very small; anal fin very small, 

originating about under rear end of second dorsal-fin base; pectoral fins about as long as head, 

straight to falcate, and broad-tipped; caudal fin lunate, with a very long lower lobe. Caudal peduncle 

strongly flattened dorsoventrally and expanded laterally, with a prominent keel on each side 

extending well onto caudal fin. Colour: back and sides intense blue in life, fading to blackish after death, 

abdomen white; underside of snout and mouth partly to entirely dusky; undersides of pectoral fins with 

dark blotches in larger individuals, pelvic fins dark with white posterior ends above, white- or dark-blotched 

below; anal fin with dark blotches or white with an anterior dark blotch. 

Size: Maximum total length at least 4.17 m. 

Habitat, biology, and fisheries: A little-known oceanic shark, possibly approaching land to give birth. 

Ovoviviparous, number of young 2. Probably feeds on oceanic schooling fishes as does Isurus oxyrinchus, 

but its large broad fins and slender body suggest that it is a slower, less active shark than that species. Not 

known to have attacked people or boats, but potentially dangerous because of its size and large teeth. 

Taken with longlines, hook-and-line, and anchored gill nets. It is utilized fresh, frozen, and dried-salted for 

human consumption. 

Distribution: Western North 

Atlantic from eastern USA to 

Cuba and southern Brazil, 

eastern Atlantic from Guinea, 

Ghana, and possibly the Cape 

Verde Islands, western Indian 

Ocean from Madagascar, 

western Pacific off Taiwan 

Province of China and Central 

Pacific near Phoenix Island 

and north of Hawaii. 

? 

intermediate 

anterior 

anterior 

? 

? 

lateral 

lateral 


? 

posterior 

? 

posterior 



Scyliorhinidae 1279 

SCYLIORHINIDAE 

Catsharks 


Diagnostic characters: small sharks with slender and elongate to moderately stout bodies. Head with 

5 gill slits, the last 2 posterior to pectoral-fin origins; gill arches with or without small papillose 

rakers; nostrils with or without barbels and lacking deep nasoral or circumnarial grooves; eyes horizontally 

oval, elongate, with weakly differentiated nictitating lower eyelids delimited below by a variably 

developed subocular pouch; mouth moderately large, with rear corners behind front margins of eyes; labial 

furrows present or absent (in species from the area); teeth very small, numerous, with a single medial 

cusp and usually 1 or more cusplets on each side near the centre of mouth, the rear teeth often 

comb-like. Two dorsal fins (only 1 dorsal fin in Pentanchus), the first originating over or posterior to 

pelvic-fin bases, the second dorsal fin smaller, as large, or larger than the first dorsal fin, but never greatly 

reduced; anal fin usually considerably longer than, and originating in advance of, second dorsal fin; caudal 

fin strongly asymmetrical, with a subterminal notch, its lower lobe absent or only weakly indicated, its 

upper edge unrippled or with a denticulated crest. Caudal peduncle not flattened dorsoventrally, 

without lateral keels or precaudal pits. Intestine with a corkscrew or auger-like spiral valve, with 5 to 22 

turns. Colour: grey, brown, yellowish, or black, often with light or dark spots and dark blotches, bars, and 

saddles. 

eyes horizontally oval, 

weak nictitating eyelids 


anal fin 

comparatively 

large in most 

species 

intestinal valve of spiral type 

no keels caudal fin 

strongly 

asymmetrical 

examples of teeth 

(with a central cusp and 1 or more 

pairs of lateral cusplets) 

Habitat, biology, and fisheries: This is by far the largest family of sharks, with small to moderate-sized 

species (rarely reaching to 1 m total length) from tropical and temperate latitudes, ranging from shallow 

coastal waters to depths greater than 2 000 m. They are generally poor swimmers and do not migrate over 

great distances. Most species live on or near the bottom, feeding chiefly on invertebrates and small fishes. 

Some species are rather common and regularly taken as bycatch in trawl fisheries, and are used for 

fishmeal, oil, and lobster bait. Many are deep-water sharks, and are not known to be utilized to a great 

extent, although they may be a minor component of the catch of large, deep-fishing offshore trawlers. 


None. The catsharks are easily distinguished from superficially similar families by the combination of 

characters such as their small size, the location of the last 2 gill slits behind the pectoral-fin origins, the 

posterior position of the first dorsal fin, the comparatively large anal fin, the strongly asymmetrical caudal 

fin, the absence of keels or precaudal pits on the caudal peduncle and the presence of a spiral intestinal 

valve.

1280 Sharks 

Key to the species of Scyliorhinidae occurring in the area 

Note: this key does not include various undescribed species of the large and problematical genus 

Apristurus that occur in the area but which need further study to determine their validity and the means of 

separating them from described species. There are also several Apristurus species recently described 

from the South China Sea that may occur in the area but were not published with precise localities given 

and which are omitted from the key. There is an undescribed species of Parmaturus from Indonesia that 

could not be placed in the key. 

1a. Supraorbital crests present on cranium, above eyes (Fig. 1a) . . . . . . . . . . . . . . . . . . → 2 

1b. Supraorbital crests absent from cranium (Fig. 1b) . . . . . . . . . . . . . . . . . . . . . . . . → 14 

2a. Second dorsal fin about as large as first . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

2b. Second dorsal fin considerably smaller than first . . . . . . . . . . . . . . . . . . . . . . . . . → 6 

3a. Anterior nasal flaps not expanded and not reaching mouth; nasoral grooves absent 

(Figs 2a and 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulohalaelurus kanakorum 

3b. Anterior nasal flaps greatly expanded, reaching mouth; nasoral grooves present 

(Fig. 2b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Atelomycterus) → 4 

crest 

a) Cephaloscyllium b) Galeus 

Fig. 1 cranium (dorsal view) 

no crest 

nasal flaps not 

reaching mouth 

a) Aulohalaelurus 

kanakorum 

nasoral 

grooves 

nasal flaps 

reaching mouth 

b) Atelomycterus 


4a. Dorsal fins not angled rearwards, posterior margins sloping posteroventrally from fin 

apices; black spots and markings relatively few, small, and scattered, colour pattern 

dominated by greyish saddles and bands on light background (Fig. 4) . . . Atelomycterus fasciatus 

4b. Dorsal fins angled rearwards, posterior margins sloping anteroventrally from fin apices; 

black spots and markings numerous and dominating colour pattern . . . . . . . . . . . . . . . → 5 

Fig. 3 Aulohalaelurus kanakorum Fig. 4 Atelomycterus fasciatus 

5a. Colour pattern of grey saddles separated by light areas and outlined by numerous small 

black spots (hatchlings have a simpler pattern of dusky saddles, remarkably similar to 

the coolie loach, Acanthophthalmus semicinctus) (Fig.5) . . . . . . . . . . Atelomycterus macleayi 

5b. Saddle markings obsolete, light grey and white spots outlined by large black spots, bars 

and lines (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atelomycterus marmoratus 

Fig. 5 Atelomycterus macleayi Fig. 6 Atelomycterus marmoratus


6a. Labial furrows present on 1 or both jaws 

(Fig. 7a) . . . . . . . . . . . . . . . (Scyliorhinus) → 7 

6b. Labial furrows absent or rudimentary 

(Fig. 7b) . . . . . . . . . . . . . (Cephaloscyllium) → 8 

7a. Colour pattern of dark spots on fins and 

body, with 7 dusky saddle marks, interspersed 

with dark and light spots (Fig. 8) . . 

. . . . . . . . . . . . . . . . . . . Scyliorhinus garmani 

7b. Colour pattern of 6 to 9 distinct dusky saddle 

marks, interspersed with dark and light 

spots; fins plain (Fig. 9) . . . . . . Scyliorhinus torazame 

8a. Colour pale with darker pattern of narrow lines . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

8b. Body coloration not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10 

9a. Colour pattern of very narrow lines forming open-centred saddles, blotches and reticulations 

on back and sides (Fig. 10) . . . . . . . . . . . . . . . . . . . . Cephaloscyllium fasciatum 

9b. Colour pattern of narrow, transverse bars on back and sides, not connected to form 

open-centred saddles, blotches and reticulations (northeastern Australia) (Fig. 11) . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloscyllium sp. D 

Fig. 10 Cephaloscyllium fasciatum 

Fig. 11 Cephaloscyllium sp. D 


10a. A simple colour pattern, consisting of only a few broad dark saddles on back and sides, 

and without white spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11 

10b. A strong colour pattern, mostly with scattered blotches and dark and white spots on 

body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 12 

11a. First dorsal-fin origin about opposite 

pelvic-fin insertions; pectoral 

fin wide, posterior margin greater 

than mouth width; pelvic-anal 

space greater than anal-fin length 

in adults; a moderate-sized species, 

to at least 70 cm total length, 

males maturing at about 55 cm 

(northeastern Australia) (Fig. 12) 

. . . . . . . . . . . . . Cephaloscyllium sp. B 

a) Scyliorhinus b) Cephaloscyllium 


Fig. 8 Scyliorhinus garmani Fig. 9 Scyliorhinus torazame 

Fig. 12 Cephaloscyllium sp. B 


11b. First dorsal-fin origin just behind pelvic-fin origins; pectoral fin narrow, posterior margin 

less than mouth width; pelvic-anal space less than anal-fin length in adults; a dwarf 

species, to 44 cm total length, males mature at 39 cm, females maturing at 36 cm (South 

China Sea to Viet Nam) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloscyllium sp. 

12a. Ventral surface (including pectoral fins) prominently spotted with darker and light spots 

on grey background; dark blotches and saddles on tail and caudal fin outlined and dotted 

with large bright white spots; anal fin with prominent dark blotch (northeastern Papua 

New Guinea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloscyllium sp. 

12b. Ventral surface plain grey or white; posterior dark blotches and saddles on tail and 

caudal fin with scattered outlined and dotted with bright white spots; anal fin plain . . . . . . . → 13

1282 Sharks 

13a. Pectoral fin broader than mouth width; upper surface of body with dark saddles, 

interspersed with light blotches and flecks that extend onto the fins (temperate eastern 

Australia) (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaloscyllium sp. C 

13b. Pectoral-fin width about equal to mouth width; upper surface of body heavily mottled, 

with saddles (tropical Australia) (Fig. 14) . . . . . . . . . . . . . . . . . . . Cephaloscyllium sp. E 


Fig. 13 Cephaloscyllium sp. C 

14a. Head broadly flattened and spatulate, 

snout elongated and usually 

longer than mouth width; labial furrows 

very long, uppers reaching upper 

symphysis (Fig. 15a) . . . . . . . . . . . → 15 

14b. Head moderately or little-flattened, 

not spatulate, snout equal or usually 

shorter than mouth width; labial furrows 

shorter or absent, when present 

not reaching upper symphysis 

(Fig. 15b) . . . . . . . . . . . . . . . . . . . → 20 

Fig. 14 Cephaloscyllium sp. E 

a) Apristurus 

b) Galeus 

longicephalus boardmani 


15a. One dorsal fin (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . Pentanchus profundicolus 

15b. Two dorsal fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Apristurus) →16 

16a. First dorsal fin much smaller than second, about 1/2 its area or less, with its origin usually 

behind pelvic-fin insertions but over last 1/4 of pelvic-fin bases in some species . . . . . . . . → 17 

16b. First dorsal fin nearly or quite as large as second, 2/3 to equal its area, with its origin 

about opposite pelvic-fin midbases or more posterior and about opposite last 1/3 of 

pelvic-fin bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 19 

17a. Origin of first dorsal fin somewhat in front of pelvic-fin insertions; distance between 

pectoral-and pelvic-fin bases extremely short, less than preoral snout, rear tips of 

pectoral fins about opposite or just in front of pelvic-fin origins (Fig. 17) . . . . Apristurus herklotsi 

17b. Origin of first dorsal fin near or behind pelvic-fin insertions; distance between pectoraland 

pelvic-fin bases long, at least length of preoral snout, rear tips of pectoral fins far 

in front of pelvic-fin origins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 18 

1 dorsal fin 

Fig. 16 Pentanchus profundicolus Fig. 17 Apristurus herklotsi 

18a. Colour white or reddish white; snout relatively narrow and pointed; mouth extending well 

in front of eyes; eyes very small, about equal to longest gill slit (Fig. 18) . . . . . Apristurus sibogae 

18b. Colour black, brown, or grey; snout broad and rounded; mouth below eyes; eyes larger, 

their length much greater than widest gill slit (Fig. 19) . . . . . . . . . . . . . .Apristurus verweyi 

Fig. 18 Apristurus sibogae Fig. 19 Apristurus verweyi


19a. Gill slits covered with grooves and pleats that extend to the epibranchial area and to the 

entire throat region behind the jaws; snout shorter, preoral length about 9% of total 

length (Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Apristurus spongiceps 

19b. Gill slits not covered with grooves and pleats; snout extremely long, preoral length about 

12% of total length (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . .Apristurus longicephalus 

grooves 

and pleats Fig. 20 Apristurus spongiceps Fig. 21 Apristurus longicephalus 

20a. Dorsal caudal-fin margin, and sometimes preventral 

margin, with a crest of enlarged denticles 

(Fig. 22a) . . . . . . . . . . . . . . . . . . . . . . . . . . → 21 

20b. No caudal-fin crests of denticles (Fig. 22b) . . . . . . . . → 28 

21a. Pectoral fins relatively small, width of their posterior 

margins usually smaller than mouth width; subocular 

ridges well-developed, eyes dorsolateral; body soft; 

colour plain, no pattern (Figs 23 and 24) . . . (Parmaturus) → 22 

21b. Pectoral fins relatively large, width of their posterior 

margins usually larger than mouth width; subocular 

ridges obsolete or nearly so, eye lateral; body firm; 

colour pattern of blotches and spots often present 

(Figs 26 to 31) . . . . . . . . . . . . . . . . . . (Galeus) → 23 

22a. Colour light brown with distal parts and anterior margins of fins, snout, nostrils, and gills 

blackish brown in young, possibly uniform blackish brown in adults; second dorsal fin 

about as large as anal fin (South China Sea, just adjacent to the area) (Fig. 23) . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parmaturus melanobranchius 

22b. Colour uniform pale yellowish brown; second dorsal fin noticably smaller than anal fin 

(Australia) (Fig. 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parmaturus sp. A 

Fig. 23 Parmaturus melanobranchius 

Fig. 24 Parmaturus sp. A 


23a. A crest of denticles present on the preventral caudal-fin 

margin (Fig. 25) . . . . . . . . . . . . . . . . . . . . . . . . → 24 

23b. No crest of denticles on the preventral caudal-fin 

margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 25 

crest of denticles 

24a. Three broad dark saddles in front of first dorsal fin, 

Fig. 25 caudal peduncle 

about as wide as eye diameter or wider (northeastern 

Australia) (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galeus boardmani 

24b. Ten to 16 narrow dark bands and saddles in front of first dorsal fin, about as wide as 

eye diameter (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galeus sp. B 

3 broad saddles 

Fig. 26 Galeus boardmani 

a) 

b) 

Fig. 22 caudal fin 

Fig. 27 Galeus sp. B 


crest of denticles 

no crest

1284 Sharks 

25a. Labial furrows very short, confined to mouth corners; 

snout broadly rounded, usually considerably 

less than mouth width (Fig. 28) . . . . . . . . Galeus schultzi 

25b. Labial furrows more elongated, extending well beyond 

mouth corners; snout more angular and 

pointed, usually nearly equal to, or about equal to 

mouth width (Fig. 29) . . . . . . . . . . . . . . . . . . . → 26 

26a. Dorsal fins and sometimes upper and lower caudal-fin 

lobes with prominent black tips . . . . . Galeus sauteri 

26b. Dorsal and caudal fins without black tips, usually 

edged with white (Figs 30 and 31) . . . . . . . . . . . . → 27 

ventral view 

of head 

Fig. 28 Galeus 

schultzi 

ventral view 

of head 

Fig. 29 Galeus 

sauteri 

27a. Eyes smaller and dorsolateral on head, length 3.2 to 3.3% total length; colour pattern 

of bold saddle markings on body and precaudal tail, but without dark markings on the 

terminal and hypural caudal-fin lobes (Fig. 30) . . . . . . . . . . . . . . . . . . . . Galeus gracilis 

27b. Eyes larger and lateral on head, length 3.5 to 4.2% total length; colour pattern of obscure 

saddle markings on body and precaudal tail, and with dark markings on the terminal 

and hypural caudal-fin lobes (Fig. 31) . . . . . . . . . . . . . . . . . . . . . . . . Galeus eastmani 

Fig. 30 Galeus gracilis Fig. 31 Galeus eastmani 

28a. Adult males with inner margins of pelvic fins fused over claspers, forming an “apron”; a 

colour pattern of spots present, but gill slits not elevated and snout rounded (Fig. 32) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Asymbolus) → 31 

28b. Adult males without inner margins of pelvic fins fused over claspers; either no colour 

pattern or, if pattern of dark spots is present, gill slits elevated above level of mouth and 

snout pointed (Figs 33 to 35) . . . . . . . . . . . . . . . . . . . . . . . . . . . (Halaelurus) → 29 

29a. Snout bluntly rounded; gill slits not elevated above mouth level, lateral in position; body 

soft, skin thin with erect denticles that gives it a velvety texture; no colour pattern 

(Fig. 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halaelurus immaculatus 


29b. Snout more or less pointed and wedge-shaped; gill slits elevated above level of mouth 

and dorsolateral in position; body firm, skin thick with low, flat, smooth denticles; colour 

pattern of dark spots, with saddles or vertical bars indistinct or absent (Figs 34 and 35) . . . . → 30 

Fig. 32 Asymbolus Fig. 33 Halaelurus immaculatus 

30a. Dark spots few and mostly much larger than spiracles, sometimes in clusters around 

vague saddle blotches; labial furrows reduced or absent, lower furrows 2 mm long or 

less (Fig. 34) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halaelurus buergeri 


30b. Dark spots small and very numerous, usually not much larger than spiracles, over or 

between weak saddles or bars; labial furrows moderately strong, lower furrows 5 mm 

long or more (Fig. 35) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halaelurus boesemani 

Fig. 34 Halaelurus buergeri Fig. 35 Halaelurus boesemani 

31a. Colour pattern of scattered dark brown spots on pale yellow-brown background (northeastern 

Australia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Asymbolus sp. E 

31b. Colour pattern of numerous small light spots and scattered small and large dark spots 

and blotches on a dark background (New Caledonia) . . . . . . . . . . . . . . . . .Asymbolus sp.




? Apristurus acanutus Chu, Meng, and Li, In Meng, Chu, and Li, 1985 

? Apristurus gibbosus Meng, Chu, and Li, 1985 



? Apristurus macrostomus Meng, Chu, and Li, 1985 

? Apristurus micropterygeus Meng, Chu, and Li, In Chu, Meng, and Li, 1986 


? Apristurus sinensis Chu and Hu, In Chu, Meng, Hu, and Li, 1981 



Apristurus sp. A. [Last and Stevens, 1994] 

Apristurus sp. B. [Last and Stevens, 1994] 

Apristurus sp. G. [Last and Stevens, 1994] 

Apristurus sp. [Seret] (New Caledonia) 

Apristurus sp. [Seret] (Philippines) 

Apristurus sp. [Seret] (Indonesia) 

Asymbolus sp. E. [Last and Stevens, 1994] 

Asymbolus sp. [Seret] (New Caledonia) 






Cephaloscyllium sp. [Compagno, 1984, 1988] 

Cephaloscyllium sp. [J.Randall, pers. comm. 1994] (Papua New Guinea) 

Cephaloscyllium sp. [Seret] (New Caledonia) 

Cephaloscyllium sp. B. [Last and Stevens, 1994] 

Cephaloscyllium sp. C. [Last and Stevens, 1994] 

Cephaloscyllium sp. D. [Last and Stevens, 1994] 

Cephaloscyllium sp. E. [Last and Stevens, 1994] 






Galeus sp. B. [Last and Stevens, 1994] 

? Halaelurus immaculatus Chu and Meng, 1982 1/ 


? Halaelurus buergeri (Mueller and Henle, 1838) 2/ 

Parmaturus melanobranchius (Chan, 1966) 3/ 

Parmaturus sp. A [Last and Stevens, 1994] 

? Parmaturus sp. [Seret] (Indonesia) 




References 

Compagno, L.J.V. 1988. Sharks of the order Carcharhiniformes. Princeton, New Jersey, Princeton University Press, 572 p. 


Regan, C.T. 1908. A synopsis of the sharks of the family Scyliorhinidae. Ann. Mag. Nat. Hist. (Ser.8), 1(6):453-65. 

Springer, S. 1979. A revision of the catsharks, family Scyliorhinidae. NOAA Tech. Rep., NMFS Circ., (422):152 p. 

1/ Occurrence in the area uncertain, marginal in the South China Sea. 

2/ This species was collected less than a degree west of the northwestern boundary of the area. 

3/ This species was collected less than a degree north of the northwestern boundary of the area.

1286 Sharks 


En - Longfin catshark; Fr - Holbiche à longues nageoires; Sp - Pejegato aletón. 

Maximum total length at least 48.5 cm. A poorly known deep-water bottom shark. Of minor interest 

to fisheries. Known from Japan, the East China Sea, and the Philippines. 


En - Longhead catshark; Fr - Holbiche à grande tête; Sp - Pejegato cabezón. 

Maximum total length at least 50 cm. Inhabits deep water, probably near the bottom at depths to 

900 m. Biology poorly known. Of minor interest to fisheries. Known from Japan, the East China Sea, 

Seychelles, the Philippines, and northern Australia. 


En - Pale catshark; Fr - Holbiche pâle; Sp - Pejegato paliducho. 

Maximum total length over 21 cm (juvenile). A rare catshark. Known only from the holotype taken 

at the Makassar Straits slope (between Borneo and Sulawesi) at a depth of 655 m. Without interest 

to fisheries.



En - Spongehead catshark; Fr - Holbiche tête molle; Sp - Pejegato esponjoso. 

Maximum size at least 50 cm. A rare shark known from 2 specimens taken on the insular slopes, 

on or near the bottom at depths of 572 to 1482 m. Probably oviparous; the holotype is a gravid 

female. Without interest to fisheries. Central Pacific off Hawaii and western South Pacific in the 

Banda Sea off southern Sulawesi. 


En - Borneo catshark; Fr - Holbiche malaise; Sp - Pejegato de Borneo. 

Maximum total length at least 30 cm. A rare deep-water catshark, known only from the holotype. 

Without interest to fisheries. Known only from the type locality (southeastern Sabah, Borneo, 

Malaysia). 


En - Banded sand catshark. 

Maximum total length about 45 cm, with size at maturity between 30 to 39 cm. A common offshore 

catshark off northern Australia, on mud, sand, or shelly-sand bottom at depths of 27 to 122 m with 

most records shallower than 60 m. Oviparous. Confined to the northwestern shelf of Western 

Australia, and the Northern Territory and northern Queensland in the area. Western Australian 

specimens lack the white spots and are lighter coloured.

1288 Sharks 


En - Australian marbled catshark; Fr - Chien marbré; Sp - Pejegato jaspeado. 

Maximum total length about 60 cm. A little-known inshore, shallow-water catshark, found on sandy 

and rocky bottom at depths of 0.5 to 3.5 m and presumably deeper. Oviparous. Of minor interest to 

fisheries at present. In the western South Pacific off northwestern Australia, and possibly Queensland. 


En - Coral catshark; Fr - Chien corail; Sp - Pintarroja coral. 

Maximum total length about 70 cm. A common but little-known, harmless inshore species, found on 

coral reefs, and thought to inhabit crevices and holes on reefs, Oviparous. Relatively unimportant 

for fisheries, forming a minor catch of inshore artisanal fisheries. From Pakistan and India eastward 

to Malaysia, Singapore, Indonesia, New Guinea, Thailand, Viet Nam, Philippines, South China, and 

Taiwan Province of China. 


En - New Caledonia catshark. 

Maximum total length about 79 cm. A rare inshore catshark, found on coral reefs at a depth of 49 m. 

Mode of reproduction unknown. Known only from a single specimen, collected off southwestern 

New Caledonia. 

?



En - Reticulated swellshark; Fr - Holbiche bouffie; Sp - Pejegato mallero. 

Maximum total length at least 42 cm. A tropical swellshark found in fairly deep water on or near the 

bottom on the outer continental shelf and uppermost slope, at depths of 220 to 450 m. Can expand 

itself with air or water. Oviparous. Of minor interest to fisheries at present, caught by commercial 

bottom trawlers. In the western Pacific off Viet Nam, China (Hainan Island), and northwestern 

Australia. 


En - Australian sawtail catshark; Fr - Chien égoïne; Sp - Pintarroja australiana. 

Maximum total length about 61 cm, males mature at 54 cm. A little-known but common Australian 

catshark of temperate and subtropical waters, from the outer continental shelf and upper slope, 

presumably on or near bottom at depths from 128 to 823 m. Frequently found in the demersal trawl 

bycatch, but of minor interest to fisheries at present. Southern coasts of Australia off western 

Australia (including Tasmania) to southern Queensland. 


En - Gecko catshark; Fr - Chien gecko; Sp - Pintarroja salamanquesa. 

Maximum total length at least 40 cm; possibly to 50 cm. A little-known, but very common shark, 

found in deep water near the bottom. Oviparous. In Japanese waters, this species shows sexual 

segregation, with reported schools of mostly females. Of minor interest to fisheries at present. 

Western North Pacific off Japan, the East China Sea, and possibly Viet Nam. 

?

1290 Sharks 


En - Slender sawtail catshark. 

Maximum total length about 34 cm. A little-known bottom-dwelling catshark of the uppermost 

continental slopes at depths of 290 to 470 m. Mode of reproduction unknown. Rare and of no 

commercial interest at present. Confined to the tropics of Australia, from Western Australia to 

northern Queensland. 


En - Blacktip sawtail catshark; Fr - Chien lime; Sp - Pintarroja rabonegro. 

Maximum total length 45 cm. A little-known bottom-dwelling catshark of the continental shelves, 

offshore at depths of 60 to 90 m in the Taiwan Straits, but possibly deeper elsewhere. Apparently 

oviparous. Taken by bottom trawls in Taiwan Straits, but of limited interest to fisheries. Western North 

Pacific off Taiwan Province of China, the Philippines, and Japan. 


En - Dwarf sawtail catshark; Fr - Chien nain; Sp - Pintarroja enana. 

Maximum total length about 27 cm; one of the smallest sharks. A little-known bottom-dwelling shark 

of the continental slopes at depths of 329 to 431 m. Of minor interest to fisheries. Known from Luzon 

(Philippines).



En - Speckled catshark; Fr - Holbiche mouchetée; Sp - Pejegato pintado. 

Maximum total length about 48 cm. A little-known but wide-ranging bottom-dwelling shark of the 

continental and insular shelves, at depths of 37 to 91 m. Of minor interest to fisheries at present. In 

the Indo-West Pacific off Somalia, the Gulf of Aden, Western Australia, Indonesia, the Philippines, 

and Viet Nam. 

Parmaturus melanobranchius (Chan, 1966) 

En - Blackgill catshark; Fr - Holbiche à joues noires; Sp - Pejegato de agallas negras. 

Maximum total length about 85 cm. A poorly known deep-water bottom-dwelling shark from the 

upper continental slopes off China, on mud bottom at depths of 549 to 810 m. Without interest to 

fisheries. Known only from 3 specimens taken in the South China Sea. 


En - Onefin catshark; Fr - Holbiche voile; Sp - Pejegato velero. 

Maximum total length at least 50 cm. A poorly known deep-water bottom-dwelling shark inhabiting 

the insular slopes. Without interest to fisheries. The holotype and only known specimen was taken 

in the Mindanao Sea (Philippines).

1292 Sharks 


En - Brownspotted catshark; Fr - Roussette à taches brunes; Sp - Alitán manchado. 

Maximum total length possibly 38 cm; adult size unknown. Without interest to fisheries. Recorded 

from the indefinite type locality (”East Indies” = Indonesia) and from the Philippines (Dumaguete, 

Negros). 


En - Cloudy catshark; Fr - Roussette nuageuse; Sp - Alitán nubarrado. 

Maximum total length to about 48 cm. A common catshark of the western Pacific continental shelf, 

from close inshore down to a depth of at least 100 m. Oviparous. Interest to fisheries unknown. 

Western North Pacific from Japan, Korea to Taiwan Province of China and the Philippines (the 

Philippine record needs confirmation). 

? ? 

? 

? 

? 



Proscylliidae 1293 

PROSCYLLIIDAE 

Finback catsharks 


Diagnostic characters: Small sharks. Trunk and precaudal tail cylindrical or somewhat compressed, 

not depressed and without lateral ridges; precaudal tail much shorter than head and trunk. Head not 

expanded laterally, moderately depressed; 5 small gill slits present, the last 2 or 3 over the pectoral fin 

bases, their upper ends not expanded onto upper surface of head; small gill raker papillae on internal 

gill slits (except in genus Gollum); spiracles moderately large and behind eyes; nostrils without 

barbels, nasoral grooves, or circumnarial grooves, well separated from mouth; eyes dorsolateral on 

head, with weakly differentiated nictitating lower eyelids; snout short to moderately long, depressed and 

parabolic or narrowly rounded, not greatly flattened and blade-like and without lateral teeth and barbels; 

mouth moderately large, arched and elongated, and extending behind front margins of eyes; very short 

labial furrows present on both jaws or absent; teeth similar in upper and lower jaws, not enlarged toward 

front of mouth, small, with a sharp primary cusp and 1 or more cusplets on either side of it, posterior teeth 

comb-shaped. Two dorsal fins, without spines, small, moderately high and angular or subangular, much 

shorter than caudal fin; first dorsal-fin base located over the interspace between pectoral- and pelvic-fin 

bases, but closer to pelvic fins than to pectoral fins; second dorsal fin about at large as first dorsal fin; 

anal fin moderately large, with its origin slightly in front or slightly behind second dorsal-fin origin but well 

in front of midpoint of second dorsal-fin base; caudal fin asymmetrical, much less than 1/2 of total 

length, without a rippled dorsal margin and without ventral lobe but with a strong subterminal notch; 

vertebral axis of caudal fin little raised above body axis. Caudal peduncle cylindrical or compressed, 

without keels or precaudal pits. Intestinal valve of spiral type. Colour: grey or brown above, white or 

lighter below, either plain, with dark stripes on the caudal fin, or with a spotted or blotched colour pattern. 


This is a small family of poorly 

known, deep-water sharks with a 

disjunct distribution in tropical to 

warm temperate waters of the 

western North Atlantic and 

Indo-West Pacific. Finback 

catsharks live on the outer 


upper and lower tooth 

continental and insular shelves and upper slopes, on or near the bottom, at depths of 50 to 713 m. Most 

of the species are ovoviviparous, except for the oviparous Proscyllium habereri. Food of these harmless 

sharks consists of small fishes and invertebrates. Their interest to fisheries is minimal, a few species are 

taken by commercial bottom trawlers and longliners, but their small size makes them unsuitable for fisheries 

utilization other than for fishmeal. 


Pseudotriakidae: first dorsal fin long, low, and keel-shaped, as long as caudal fin; spiracles about as large 

as eyes. 

Scyliorhinidae: first dorsal fin over or behind pelvic-fin bases. 

Pseudotriakidae 

Scyliorhinidae (Atelomycterus) 

ventral view 

of head

1294 Sharks 

Triakidae: no gill raker papillae on internal gill 

openings; nictitating lower eyelids better 

differentiated, with a deeper subocular pocket and a 

well-developed secondary lower eyelid edge; labial 

furrows long; teeth stouter, with heavier cusps or no 

cusps, posterior teeth not comb-like; first dorsal-fin 

base in species from the area more anterior, closer to 

the pectoral-fin bases than to the pelvic fins or about 

equidistant between the 2. 

No other sharks in the area combine 

the following characteristics: 

nictitating lower eyelids, small, 

cuspidate teeth in both jaws, mouth 

located under eyes, intestinal valve 

of spiral type, no precaudal pits, and 

no rippled dorsal caudal-fin margin. 

teeth 

Key to the species of Proscylliidae occurring in the area 

long 


1a. Head and snout bell-shaped in dorsoventral view (Fig. 1a); no oral papillae or gill rakers 

in mouth (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gollum attenuatus 

1b. Head and snout narrowly rounded in dorsoventral view (Fig. 1b, c); oral papillae and gill 

rakers present in mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

a) Gollum 

attennuatus 

b) Proscyllium 

habereri 

Triakidae 

(Mustelus) 

c) Eridacnis 

radcliffei 

Proscylliidae 

(Eridacnis) 

2a. Caudal fin narrow and ribbon-like; colour dark brown with blackish markings on dorsal 

fins (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eridacnis radcliffei 

2b. Caudal fin broad and not ribbon-like; colour pattern of round dark brown spots and 

indistinct saddles (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Proscyllium habereri 





Proscyllium habereri Hilgendorf, 1904 

Triakidae (Mustelus) 

Triakidae 

Fig. 1 ventral view of head Fig. 2 Gollum attenuatus 

Fig. 3 Eridacnis radcliffei Fig. 4 Proscyllium haberi 

short or 

absent 

Proscylliidae 

References 

Compagno. L.J.V. 1970. Systematics of the genus Hemitriakis (Selachii: Carcharhinidae), and related genera. Proc. 

Calif. Acad. Sci., (38)4:63-98. 

Compagno. L.J.V. 1973. Ctenacis and Hemitriakis, two genera of sharks (Selachii: Carcharhinidae). Proc. Calif. Acad. 

Sci., (39)4:257-272. 

Compagno. L.J.V. 1973. Gogolia filewoodi, a new genus and species of shark from New Guinea (Carchariniformes: 

Triakidae), with a redefinition of the family Triakidae and a key to the genera. Proc. Calif. Acad. Sci., 

(39)4:383-410. 

Compagno. L.J.V. 1988. Sharks of the order Carcharhiniformes. Princeton, New Jersey, Princeton University Press, 572 p.

Proscylliidae 1295 


En - Pygmy ribbontail catshark; Fr - Requin chat pygmé; Sp - Tollo coludo pigmeo. 

Maximum total length about 24 cm. One of the smallest living sharks. Often occurs on mud bottoms, 

on the upper continental and insular slopes and the outer shelves at depths from 71 to 766 m. Feeds 

primarily on small bony fishes and crustaceans. Taken in bottom trawls in the Philippines, but 

utilization not known. Wide-ranging in the Indo-West Pacific, but with only spotty records from 

Tanzania, the Gulf of Aden, India, the Andaman Islands, Viet Nam, and the Philippines. 


En - Slender smooth-hound; Fr - Requin chat golloum; Sp - Tollo coludo elegante. 

Maximum total length about 1.1 m, maturing at about 70 cm, with females growing slightly larger 

than males. An uncommon to common deep-water bottom-dwelling shark of the outermost continental 

shelf and upper slope of New Zealand and on adjacent seamounts and submarine banks, 

found at depths of 220 to 660 m, but most commonly between 400 and 600 m. Probably occurs in 

schools. Feeds on a wide variety of mostly small pelagic and benthic bony fishes, deep-water sharks, 

cephalopods (including pelagic squid and octopuses), sea snails, isopods, crabs and shrimps, and 

brittle stars. Ovoviviparous, with usually 2 young per litter, fetuses eat unfertilized eggs (uterine 

cannibalism) and store consumed yolk in their yolk sacs. Without interest to fisheries at present, 

although taken in small numbers by bottom trawlers off New Zealand and collected in moderate 

numbers by experimental longliners fishing in deep water on seamounts and banks. Occurs in the 

western South Pacific, off New Zealand and on rises between New Zealand and the east coast of 

Australia, New Caledonia, and Fiji just south of the area. Placement in Proscylliidae provisional, 

probably will be relocated in Pseudotriakidae. 

Proscyllium habereri Hilgendorf, 1904 

En - Graceful catshark; Fr - Requin chat gracile; Sp - Tollo coludo grácil. 

Maximum total length about 65 cm. A little-known, uncommon bottom-dwelling shark of tropical and 

warm-temperate continental and insular waters, found on the shelves at depths from 50 to 100 m. 

Food habits little-known. Taken by bottom trawlers in the Taiwan Straits and elsewhere in its range, 

utilisation unknown. In the western Pacific from southeastern Japan southwards to Viet Nam, also 

known from northwestern Java.

1296 Sharks 


Pseudotriakis microdon Capello, 1868 

PSEUDOTRIAKIDAE 

False catsharks 



FAO names: En - False catshark; Fr - Requin à longue dorsale; Sp - Musolón aleta larga. 

Diagnostic characters: A large, soft-bodied shark. Head with 5 small gill 

slits, the last 2 over the pectoral-fin bases; no dermal gill rakers; spiracles 

very large, about as long as eyes; nostrils without barbels or nasoral 

grooves; eyes above sides of head, horizontally elongated, with weakly 

differentiated nictitating lower eyelids that are delimited below the eyes by 

shallow pouches; snout moderately long, narrowly rounded; mouth very 

wide and long, extending behind front of eyes, angular in shape; labial 

furrows present but short, not extending forward to front of mouth; teeth 

extremely small and numerous, similar in both jaws and not bladelike, 

with a small primary cusp and 1 or more cusplets, becoming comblike in 

the rear of mouth; upper anterior teeth small and grading into the laterals, 

not separated from these by small intermediate teeth. Two dorsal fins, 

ventral view 

of head 

dermal 

denticle 

the first greatly elongated, low, keel-like, and broadly rounded above, its base just ahead of pelvic-fin 

origins and as long as caudal fin; second dorsal fin short but higher than the first and larger than the anal 

fin; anal-fin base under second dorsal-fin base; caudal fin greatly asymmetrical, its lower lobe hardly 

developed, its upper edge not rippled and a subterminal notch present. Caudal peduncle not depressed, 

without lateral keels or precaudal pits. Intestinal valve of spiral type. Colour: dark brownish grey above and 

below, darker on posterior edges of pelvic, dorsal, anal and caudal fins. 


None. No other sharks in the area combine the presence of a low, keel-like first dorsal fin equal in length 

to the caudal fin, no anal-fin and no dorsal-fin spines. 

Size: Maximum total length about 2.95 m; females mature at about 2.1 m. 

Habitat, biology, and fisheries: A deep-water bottom-dwelling shark, normally occurring on the upper 

continental and insular slopes at depths between 200 and 1500 m; occasionally wandering onto continental 

shelves, even in shallow water. Ovoviviparous; practices uterine cannibalism, with fetuses eating unfertilized 

eggs; 2 young per litter. Apparently somewhat inactive and sluggish. Feeding habits little known, once 

photographed in deep water eating a bony fish; probably feeds on a variety of deep-water bony fishes, 

elasmobranchs, and invertebrates. Taken incidentally offshore on deep-set longlines and less commonly 

on bottom trawls. Utilization not recorded. 

Distribution: Western North Atlantic from 

New York and New Jersey, eastern North 

Atlantic from Iceland to Senegal, western 

Indian Ocean from the Aldabra Islands 

group and western Pacific from Japan, Taiwan 

Province of China, New Zealand, 

Western Australia, and Hawaii. 

Reference 

Compagno. L.J.V. 1988. Sharks of the order 

Carcharhiniformes. Princeton, New Jersey, 

Princeton University Press, 572 p.

Triakidae 1297 

TRIAKIDAE 

Houndsharks, smoothhounds, topes 

by L.J.V Compagno and V.H. Niem 

Diagnostic characters: Body elongate and slender to moderately stout. Head with 5 gill slits, the last 

pair posterior to pectoral-fin origins; small spiracles present; gill arches without rakers; eyes 

horizontally oval, situated on or above sides of head, with a nictitating eyelid partly or entirely within the 

eye opening; anterior nasal flaps of nostrils either broadly to narrowly expanded or greatly reduced, but 

not in the form of slender barbels; mouth ending below or behind eyes; labial furrows moderately long; 

teeth usually similar in both jaws, but differentiated in a few species; in Mustelus, they are numerous, small, 

cuspless (or weak-cusped), and arranged in a pavement, in Galeorhinus they are larger, blade-like, with 

a strong cusp, and small cusplets but no serrations, and in Triakis they are of intermediate structure. Two 

dorsal fins, the first much shorter than caudal fin and with its base entirely anterior to pelvic fins; second 

dorsal fin somewhat smaller than the first, originating ahead of anal fin; anal fin as large as or smaller 

than second dorsal fin; caudal fin asymmetrical, its lower lobe varying from virtually absent to strong, its 

upper edge not rippled. Caudal peduncle not flattened dorsoventrally or expanded laterally, without keels 

or precaudal pits. Intestine with a corkscrew or auger-like spiral valve, with6to10turns.Colour: 

back usually greyish brown, belly white; some species are capable of undergoing slow colour changes. 

Mustelus 

teeth 

Galeorhinus 


Habitat, biology, and fisheries: Houndsharks are widely distributed in tropical and warm temperate to 

cold seas, ranging from shallow to moderately deep water (300 m or more). The species are variably 

ovoviviparous or viviparous, and lack or have a yolk sac placenta. They feed on bottom-dwelling invertebrates 

(especially crustaceans, but also molluscs and worms), and on small bony fishes and fish eggs. 

None of the species are particulary dangerous to people. Smoothhounds (Mustelus) and topes or soupfin 

sharks (Galeorhinus) are important commercial species. 


Carcharhinidae and Hemigaleidae: upper edge of caudal fin with a rippled or undulating margin; precaudal 

pits present. Furthermore, Carcharhinidae have an intestinal valve of scroll type. 

precaudal pit 

dorsal margin 

viewed from above 

Carcharhinidae 

notch 

well-developed 

lower lobe 

rolled 

unrolled 


(intestinal valve of scroll type)

1298 Sharks 

Proscylliidae: gill raker papillae 

present on internal gill 

openings (lacking in Gollum); 

nictitating lower eyelids not 

well developed, of rudimentary 

type, with a weaker 

subocular pocket and a 

poorly differentiated secondary 

lower eyelid edge; teeth 

with slender cusps, comb-like 

at ends of dental bands. 

Other families: the combination of characters such as nictitating lower eyelids, small spiracles, mouth under 

or behind the eyes, well-developed labial furrows, 2 spineless dorsal fins with the first over the interspace 

between the pectoral and pelvic fins, absence of precaudal pits, and presence of an anal fin separates this 

family from all other sharks occurring in the area. 

Key to the species of Triakidae occurring in the area 

1a. First dorsal-fin base about as 

long as caudal fin and 2.3 to 3.2 

times the first dorsal height; preoral 

length about 1.6 to 1.7 times 

mouth width (Fig. 1) . . . . Gogolia filewoodi 

1b. First dorsal-fin base 0.7 times or 

less in length of caudal fin and 2 

times or less the first dorsal-fin 

height; preoral length 0.7 to 1.4 

times mouth width . . . . . . . . . . . . → 2 

2a. Ventral caudal-fin lobe very long 

at all stages; second dorsal fin 

markedly smaller than first, 1/2 

itsareaorless(Figs2and3) . . . . . . → 3 

2b. Ventral caudal-fin lobe absent to 

short in adults, weak or absent 

in young; second dorsal fin 

nearly or quite as large as first, 

2/3 to about equal its area . . . . . . . . → 4 

3a. Mouth angular; second dorsal 

fin considerably larger than anal 

fin; terminal lobe of caudal fin 

about 1/3 of total dorsal caudalfin 

margin length (Fig. 2) . . . . 

. . . . . . . . . . . Hypogaleus hyugaensis 

3b. Mouth arcuate; second dorsal 

fin about as large as anal fin; 

terminal lobe of caudal fin about 

1/2 of total dorsal caudal-fin 

margin length (Fig. 3) . . Galeorhinus galeus 

nasal flap 

a) ventral view 

of head 


of head 


of head 

Proscylliidae 


Fig. 1 Gogolia filewoodi 


Fig. 2 Hypogaleus hyugaensis 

lower teeth comb-like 

rear teeth 


Fig. 3 Galeorhinus galeus 

ventral 

caudal-fin 

lobe 

ventral 

caudal-fin 

lobe


4a. Eyes lateral, subocular ridges obsolete (Fig. 4a); origin of first dorsal fin far anterior, over 

pectoral-fin bases (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iago garricki 

4b. Eyes dorsolateral, subocular ridges strong (Fig. 4b); origin of first dorsal fin more posterior, 

over or behind inner margins of pectoral fins (Figs 6 to14) . . . . . . . . . . . . . . . . . . . . → 5 

eyes 

lateral 

eyes 

dorsolateral 

b) Hemitriakis, 

a) Iago garricki Triakis, Mustelus 

Fig. 4 dorsal view of head Fig. 5 Iago garricki 

5a. Internarial width about 2.5 times the nostril width (Figs 6 and 7); teeth strongly 

compressed and blade-like, differentiated into medials at symphyses of both jaws and 

antero-posteriors adjacent to them . . . . . . . . . . . . . . . . . . . . . . . . . (Hemitriakis) → 6 

5b. Internarial width 1 to 2 times the nostril width (Figs 9,10 and12); teeth broad and blunt 

to semi-blade-like, not strongly compressed, not differentiated into medials and anteroposteriors 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9 

6a. Eyes relatively low and 

slit-like; first dorsal-fin origin 

over or behind pectoral 

fin free rear tips; fins 

not strongly falcate in 

adults (Fig. 6) . . Hemitriakis japanica 


6b. Eyes relatively high and 

horizontally oval; first dorsal-fin 

origin anterior to 

pectoral fin free rear tips; 

fins strongly falcate in 

adults . . . . . . . . . . . . . . . → 7 

subocular 

ridge 

present 


of head 


Fig. 6 Hemitriakis japanica 

7a. Precaudal vertebral counts 94 to 96, monospondylous precaudal vertebral counts 34 

to 35; no dusky bar on underside of snout; young with dark bars on caudal fin but not 

elsewhere (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemitriakis leucoperiptera 

7b. Precaudal vertebral counts 116 to 132, monospondylous precaudal counts 44 to 57; 

dusky bar present on underside of snout; young with prominent dark bars and spots on 

fins and body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8 

8a. Young with light spots in centres of saddle markings, light tip on caudal fin; precaudal 

vertebral counts 116, monospondylous precaudal counts 44, diplospondylous precaudal 

counts 72 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemitriakis sp. 

8b. Young with solid saddle markings, without light central spots, dark tip on caudal fin; 

precaudal vertebral counts 126 to 132, monospondylous precaudal counts 48 to 57, 

diplospondylous precaudal counts 75 to 78 (Fig. 8) . . . . . . . . . . . . . . . . Hemitriakis abdita 

ventral view 

of head upper and lower tooth 

Fig. 7 Hemitriakis leucoperiptera Fig. 8 Hemitriakis abdita

1300 Sharks 

9a. Snout bluntly rounded in dorsoventral 

view; mouth arcuate, lower jaw with 

convex edges (Fig. 9) . . . . . Triakis scyllium 


9b. Snout parabolic to subangular in dorsoventral 

view;mouth angular, lower jaw 

with straight or nearly straight edges 

(Figs 10 and 12) . . . . . . . (Mustelus) → 10 

10a. No white spots on body; upper labial 

furrows longer than lower furrows 

(Fig. 10) . . . . . . . . . . . . . . . → 11 


Fig. 9 Triakis scyllium 

10b. Numerous small white spots on dorsal surface of body; upper labial furrows subequal 

to or shorter than lower furrows (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 12 

11a. Interorbital space 3.6 to 4.5% of total length; teeth in both jaws with low cusps; precaudal 

centra fewer, 73 to 80 (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mustelus griseus 

11b. Interorbital space 5.6 to 6.0% of total length; teeth in both jaws with high cusps; 

precaudal centra more numerous, 90 to 92 (Fig. 11) . . . . . . . . . . . . . . . . . Mustelus sp. A 


of head 


Fig. 10 Mustelus griseus 

12a. Interorbital space relatively narrow, 

3.7 to 4.5% of total length 

(Fig. 12) . . . . . . . . . . . Mustelus manazo 

12b. Interorbital space relatively broad, 

6.1 to 7.1% of total length 

(Mustelus antarcticus species 

complex) . . . . . . . . . . . . . . . . . → 13 


of head 

Fig. 11 Mustelus sp. A 



of head 


13a. Total vertebrae 125 to 133; mono- 

Fig. 12 Mustelus manazo 

spondylous precaudal centra 34 to 

37; precaudal centra 79 to 86; size 

at maturity over 80 to 85 cm and reaching 1.75 m total length; temperate species 

occurring from inshore to 350 m (Fig. 13) . . . . . . . . . . . . . . . . . . . . Mustelus antarcticus 

13b. Total vertebrae 135 to 143; monospondylous centra 37 to 39; precaudal centra 88 to 95; 

size at maturity over 70 cm and reaching 1.17 m total length; tropical species in deeper 

water, 120 to 400 m (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . Mustelus sp. B (eastern form) 

13c. Total vertebrae 119 to 128, mostly less than 125; monospondylous centra 33 to 35; 

precaudal centra 76 to 80; size at maturity about 60 cm and reaching 1.03 m total length; 

tropical species in deeper water, 120 to 400 m (Fig. 14) . . . . . . . Mustelus sp. B (western form) 

Fig. 13 Mustelus antarcticus 

Fig. 14 Mustelus sp. B 







Hemitriakis abdita Compagno and Stevens, 1993 1/ 

 

 

 

 

 

 

? Hemitriakis japanica (Müller and Henle, 1839) 2/ 


Hemitriakis sp. [Compagno, 1988] 3/ 


Iago garricki Fourmanoir, 19794/ 


Mustelus griseus Pitschmann, 19085/ 


Mustelus cf. manazo [Seret, pers. comm. 1994] 

Mustelus sp. A. [Last and Stevens, 1994] 6/ 

Mustelus sp. B. [Last and Stevens, 1994] 7/ 

? Triakis scyllium Müller and Henle, 1839 8/ 

References 

Compagno, L.J.V. 1970. Systematics of the genus Hemitriakis (Selachii: Carcharhinidae), and related genera. Proc. 

Calif. Acad. Sci., 38(4):63-98. 

Compagno, L.J.V. 1973. Gogolia filewoodi, a new genus and species of shark from New Guinea (Carcharhiniformes; 

Triakidae), with a redefinition of the family Triakidae and a key to the genera.Proc. Calif. Acad.Sci., 39(4):383-410. 

Compagno, L.J.V. 1988. Sharks of the order Carcharhiniformes. Princeton, New Jersey, Princeton University Press, 

572 p. 


1/ Including specimens from the Coral Sea off Queensland, Australia, and a specimen from New Caledonia that agrees 

with the Coral Sea material in most particulars. 

2/ Nominal from Amboina and New Caledonia; unconfirmed from the former locality, and based on a specimen close 

to if not identical to Hemitriakis abdita from New Caledonia. 

3/ Based on 4 late fetuses from the Philippines in the Stanford University collections, originally misidentified as Triakis 

scyllium. Postnatal material has not been examined. 

4/ A Philippines’ Iago is similar to I. garricki but may be distinct. 

5/ A Philippines’ Mustelus is similar to M. griseus and Mustelus sp. A, particularly the former, but needs further 

investigation to determine its identity. 

6/ An unspotted smoothhound known from isolated localities off northern Australia, but presumably more widely 

distributed. 

7/ Known from few isolated localities off northern Australia. It is not clear if the western and eastern Australian 

populations represent 1 or 2 species or are just variations or subspecies of Mustelus antarcticus. 

8/ Nominal records from Philippines, apparently based in part on Hemitriakis sp. Otherwise a temperate coastal 

species occurring from China to Korea and Japan.

1302 Sharks 


En - Tope shark; Fr - Requin-hâ; Sp - Cazón. 

Maximum total length about 1.95 m. An active, strong-swimming, abundant, coastal-pelagic species 

of temperate continental and insular waters, near or on the bottom, from inshore waters down to a 

depth of 300 m. Ovoviviparous, without a yolk-sac placenta. Feeds on small schooling fish, other 

bottom fishes, crustaceans, and echinoderms. Caught with bottom trawls, on longlines, and in 

pelagic trawls; utilized fresh, dried salted and processed for oil (Vitamin A) and fishmeal. In the 

western South Atlantic, eastern North and South Atlantic, also off southern Australia, New Zealand, 

the Laysan Islands, Hawaii, and the eastern North and South Pacific. 

dorsal view of head ventral view of head 



En - Sailback houndshark; En - Requin-hâ voile; Sp - Cazón velero. 

Maximum total length about 74 cm. A little-known shark of the northern New Guinea continental 

shelf, the only specimen of which was taken at a depth of 73 m, probably near the bottom. 

Ovoviviparous. Of minor interest to fisheries. 

dorsal view of head ventral view of head 


En - Whitefin topeshark; Fr - Requin-hâ aile blanche; Sp - Cazón de aleta blanca. 

Maximum total length about 96 cm. A little-known inshore tropical shark, in Philippine coastal waters 

down to a depth of 48 m. Viviparous, but it is not known if a yolk-sac placenta is formed; presumably 

caught by local fisheries in the Philippines, but details are unknown. 




En - Blacktip tope; Fr - Requin-hâ elegant; Sp - Cazón elegant. 

Maximum total length at least 1.3 m. A bottom-living shark of fairly deep continental waters, ranging 

in depths from 40 to 230 m. Viviparous, with a yolk-sac placenta. Feeds on bony fishes. Caught in 

bottom trawls and on hook-and-line; a minor fisheries catch off southern Australia and Japan, but 

uncommon and apparently little utilized elsewhere. Occurs off South Africa, Kenya, Zanzibar, the 

Persian Gulf, and in the western Pacific off southern Japan, Taiwan Province of China, and Australia. 



Iago garricki Fourmanoir and Rivaton, 1979 

En - Longnose houndshark; Fr - Requin-hâ long nez; Sp - Cazón picudo. 

Maximum total length about 75 cm. A little-known, deep-water tropical shark found at depths of 250 

to 475 m. Viviparous, with a yolk-sac placenta. Feeds on cephalopods. Of minor interest to fisheries 

at present. Known from the Vanuatu and tropical Australia; also, a shark that is somewhat similar to 

this species, but may be distinct, occurs in the Philippines (Batangas, Luzon) in depths of 124 to 

441 m. 


1304 Sharks 


En - Gummy shark; Fr - Emissole gommée; Sp - Musola austral. 

Maximum total length about 1.75 m. An abundant inshore and offshore shark of temperate waters, 

found on or near the bottom and from the intertidal to a depth of 183 m. Ovoviviparous, without a 

yolk-sac placenta. Feeds on crustaceans (including crabs), marine worms, and small fishes. This 

small shark is widely fished in Australia, and utilized fresh for human consumption. Western Australia 

to southern Queensland, also Tasmania. 


Mustelus griseus Pietschmann, 1908 

En - Spotless smooth-hound; Fr - Emissole cotiere; Sp - Musola gris. 

Maximum size at least 1 m. A common western Pacific temperate and tropical inshore bottom-dwelling 

shark, found down to a depth of at least 51 m. Viviparous, with a yolk-sac placenta. Probably 

feeds on bottom-dwelling invertebrates, especially crustaceans. Regularly fished off Japan, China 

and Taiwan Province of China, presumably also caught by local fisheries in the area, but details are 

unknown. In the western North Pacific off Japan, Korea, China, Taiwan Province of China, and Viet 

Nam, possibly also the Philippines. 



En - Starspotted smooth-hound; Fr - Emissole etoilée; Sp - Musola celestrial. 

Maximum total length to about 1.17 m. An abundant, bottom-living shark found in continental waters, 

ranging from the intertidal zone to a depth of at least 360 m. Ovoviviparous, without a yolk-sac 

placenta. Feeds mostly on bottom invertebrates. Caught in bottom trawls and on hook-and-line 

(sports catches). In the western Pacific southwards to Viet Nam, also recorded from Kenya. 


?

Hemigaleidae 1305 

HEMIGALEIDAE 

Weasel sharks 


Diagnostic characters: Small to medium-sized sharks with cylindrical or slightly compressed bodies 

without lateral ridges; precaudal tail much shorter than trunk. Head not expanded laterally, moderately 

depressed; 5 small to medium-sized gill slits present, the last 2 or 3 over or behind pectoral-fin origins, 

their upper ends expanded partway onto upper surface of head in some species; no gill sieves or gill 

rakers; spiracles minute, and behind but not below eyes; nostrils without barbels, nasoral grooves, or 

circumnarial grooves, well-separated from mouth; eyes on sides of head, with a well-developed nictitating 

lower eyelid; snout moderately long, depressed, and parabolic to broadly rounded, not greatly flattened 

and blade-like and without lateral teeth and barbels; mouth moderately large, arched, and elongated, and 

extending well behind eyes; labial furrows present on both jaws and moderately large, reaching front of 

mouth or ending well behind it; teeth small to large, blade-like, and with a single cusp on teeth of both 

jaws, cusplets or strong serrations present on upper teeth, and cusplets variably present or absent on lower 

teeth; anterior teeth in upper jaw smaller than lateral teeth and not separated from them by smaller 

intermediate teeth on each side. Two dorsal fins without spines, the first dorsal fin moderately large, high 

and angular or subangular, much shorter than the caudal fin, with its base located over the interspace 

between, the pectoral- and pelvic-fin bases and entirely anterior to origins of pelvic fins; second dorsal fin 

moderately large, about 2/3 the size of first dorsal fin; anal fin moderately large, slightly smaller than second 

dorsal fin, with its origin slightly behind second dorsal-fin origin but in front of second dorsal-fin midbase; 

caudal fin strongly asymmetrical, much less than half of total length, with a rippled or undulated dorsal 

margin, a well-marked subterminal notch, and a short, but well-defined lower lobe; vertebral axis of caudal 

fin raised above body axis.Caudal peduncle cylindrical, without keels but with well-developed precaudal 

pits. Intestinal valve of spiral type. Colour: grey, grey-brown or dark grey above, white or cream below, 

fins sometimes with dusky tips or white posterior margins; sometimes a few white spots but no elaborate 


ventral view of head upper and lower teeth 

Habitat, biology, and fisheries: The Hemigaleidae is a small family of small to medium-sized, coastal 

sharks with a primary diversity (about 7 to 9 species) in the continental and insular tropical waters of the 

Indo-West Pacific (but not extending into the Central Pacific); a single additional species occurs in the 

Atlantic. It is closely related to the large family Carcharhinidae. These sharks feed on small fishes, 

octopuses, and probably other invertebrates, and are not known to have attacked people. All species in 

the area are fished for human consumption, but due to their modest abundance they form only a small 

fraction of the shark catch in the area.

1306 Sharks 


Carcharhinidae: intestine with a scroll valve; also, no carcharhinids in the area show the character 

combination of a long snout, spiracles, upper teeth with cusplets, lower teeth well differentiated from uppers, 

long labial furrows, and second dorsal fin about 2/3 as large as first dorsal fin and with its origin anterior 

to that of the slightly smaller anal fin. 

rolled 

unrolled 


Hemigaleidae 

and many other families 

scroll valves spiral valve 


Proscylliidae and Triakidae: no precaudal pits or undulated dorsal caudal-fin margin, teeth not strongly 

differentiated in upper and lower jaws, spiracles usually larger. 

upper and lower teeth Proscylliidae 

Key to the species of Hemigaleidae occurring in the area 

1a. Lower teeth near symphysis with short, straight or weakly hooked cusps that are 

concealed or barely protrude when mouth is closed (Fig. 1a, b); gill slits small, less than 

twice the eye length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2 

1b. Lower teeth near symphysis with long, strongly hooked cusps that prominently protrude 

from mouth when it is closed (Fig. 1c, d); gill slits large, over twice the eye length . . . . . . . . → 4 

wedgedshaped 

root 

short 

slightly 

arched 

a) Paragaleus tengi 

long 

smooth 

cusps 

short 

cusp 

long 

very short 

root 

highly 

arched 

b) Hemigaleus microstoma 

rounded 

c) Chaenogaleus macrostoma 

d) Hemipristis elongatus 

Fig. 1 teeth and ventral view of head 

long 

serrated 

cusps 

short 

cusp 

long


2a. Lower teeth near symphysis with mostly erect cusps and slightly arched roots, giving 

them an inverted T-shape; cusplets present on lower teeth; mouth longer, narrowly 

arched (Fig. 1a); pelvic, dorsal and caudal fins not falcate (Fig. 2) . . . . . . . . . Paragaleus tengi 

2b. Lower teeth near symphysis with erect cusps and highly arched roots, giving them an 

inverted Y-shape; no cusplets on lower teeth; mouth very short, broadly arched (Fig. 1b); 

pelvic fins, both dorsal fins, and ventral caudal fin lobe strongly falcate (Fig. 3) . . . . . . . . . → 3 

Fig. 2 Paragaleus tengi Fig. 3 Hemigaleus microstoma 

3a. Fins dark-tipped; tooth counts 28-34/43-54; total vertebral counts 111 to 118 (Australia) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemigaleus sp. aff. “microstoma” 

3b. Fins light-tipped; tooth counts 25-32/37-43; total vertebral counts 137 to 150 (Java, 

Singapore, and Thailand) . . . . . . . . . . . . . . . . . . . . . . . . . . Hemigaleus microstoma 

4a. Snout obtusely wedge-shaped in dorsoventral view; teeth present at symphysis of lower 

jaw; mesial edges of upper teeth smooth at all sizes (Fig. 1c); fins not falcate, posterior 

margins of pelvic and pectoral fins straight or slightly concave (Fig. 4) . . . Chaenogaleus macrostoma 

4b. Snout bluntly rounded in dorsoventral view; teeth absent at symphysis of lower jaw, 

mesial edges of upper teeth serrated (but smooth in young below 60 cm) (Fig. 1d); fins 

strongly falcate, posterior margins of pelvic and pectoral fins deeply concave (Fig. 5) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemipristis elongatus 

Fig. 4 Chaenogaleus macrostoma Fig. 5 Hemipristis elongatus 





Hemigaleus sp. aff. “microstoma” 

Hemipristis elongatus (Klunzinger, 1871) 


References 


572 p. 

Compagno, L.J.V. and M.J. Smale. 1985. Paragaleus leucolomatus, a new shark fro South Africa, with notes on the 

systematics of hemigaleid sharks (Carcharhiniformes: Hemigaleidae). Spec. Publ. J.L.B. Smith Inst. Ichthyol., 

(37):391-401. 

Compagno, L.J.V., F. Krupp, and K.E. Carpenter. 1996. A new weasel shark of the genus Paragaleus from the 

northwestern Indian Ocean and the Arabian Gulf. Fauna of Saudi Arabia, 15:391-401.

1308 Sharks 


Frequent synonyms / misidentifications: Hemigaleus macrostoma Bleeker, 1852; H. balfouri Day, 1878 / 

Hemigaleus microstoma Bleeker, 1852; Hemipristis elongatus (Klunzinger, 1871). 

FAO names: En - Hooktooth shark; Fr - Milandre harpon; Sp - Comadreja ganchuda. 

Diagnostic characters: A small shark. Body moderately slender. Snout long, its length slightly greater 

than mouth width, obtusely wedge-shaped toward tip; labial furrows moderately long and easily seen, 

the uppers ending well behind symphysis of lower jaw; anterior nasal flaps with a short, broad, triangular 

lobe; mouth long and parabolic, its length over 2/3 of the width; spiracles small; gill slits very long, 

the longest over twice the eye length; teeth in upper jaw with narrow, erect to oblique, high cusps and 

distal cusplets (except for those at symphysis), entirely smooth-edged; teeth in lower jaw with arched 

roots and long, hooked, slender mostly erect cusps that prominently protrude when mouth is 

closed, without cusplets or serrations. First dorsal fin moderately large, with a pointed or narrowly rounded 

apex and short inner margin, its origin slightly posterior to free rear tips of pectoral fins and the free rear 

tip anterior to pelvic-fin origins; second dorsal fin high, about 2/3 of length of first, with a short inner 

margin less than fin height, and its origin slightly anterior to anal-fin origin; anal fin slightly smaller than 

second dorsal fin, without long preanal ridges; upper precaudal pit transverse and crescentic, no keels 

on caudal peduncle; pectoral and pelvic fins with straight or slightly concave inner margins. Intestine 

with a spiral valve. Colour: bronzy-grey above, white below when fresh, fading to greyish or greyish brown 

in preservation, dorsal fins often with dusky or black tips. 

Size: Maximum total length about 1 m. 

Habitat, biology, and fisheries: A small, common, coastal, inshore and offshore shark of the continental 

and insular shelves, caught at depths down to 59 m. Viviparous, number of young 4; size at birth 20 cm. 

Probably feeds on small fishes and invertebrates; harmless to people. Commonly caught in inshore and 

offshore artisanal fisheries probably everywhere where it occurs. Caught in drifting and bottom gill nets 

and on longlines and other line gear. Utilized fresh for human consumption; offal processed into fishmeal. 

Distribution: Indo-Pacific 

from the Persian Gulf to India 

and Sri Lanka, and off 

Singapore, Thailand, Viet 

Nam, China (including Taiwan 

Province), Java, and Sulawesi.



Frequent synonyms / misidentifications: Negogaleus microstoma (Bleeker, 1852) / Chaenogaleus 

macrostoma (Bleeker, 1852); Hemipristis elongatus (Klunzinger, 1871). 

FAO names: En - Sicklefin weasel shark; Fr - Milandre faucille; Sp - Comadreja segadora. 

Diagnostic characters: A small shark. Body moderately slender. Snout long, its length somewhat greater 

than mouth width, parabolic toward tip; labial furrows moderately long and easily seen, the upper 

furrows nearly reaching symphysis of lower law, anterior nasal flaps with a short, broad, triangular lobe; 

mouth very short and broadly arched, its length about 1/3 of the width; spiracles small; gill openings 

short, the longest slightly longer than eye length in adults, slightly shorter in young; teeth in upper jaw 

with very narrow, short, oblique cusps and prominent distal cusplets (except for those at symphysis), 

entirely smooth-edged; teeth in lower jaw very small, with arched roots and short, mostly erect, 

slender, straight cusps that do not protrude when mouth is closed, and no cusplets or serrations. First 

dorsal fin moderately large, with a pointed apex and short inner margin, its origin slightly posterior to free 

rear tips of pectoral fins and the free rear tip anterior to pelvic-fin origins; second dorsal fin high, about 

2/3 of length of first dorsal fin, with a short inner margin less than fin height, and its origin slightly anterior 

to anal-fin origin; anal fin slightly smaller than second dorsal fin, without long preanal ridges; upper 

precaudal pit transverse and crescentic, no keels on caudal peduncle; pectoral and pelvic fins strongly 

falcate, with deeply concave posterior margins. Intestine with a spiral valve. Colour: grey-brown above, 

lighter below, dorsal fins with white tips and posterior margins: sometimes white spots on sides of body. 


Habitat, biology, and fisheries: A small coastal, inshore and offshore shark of continental tropical waters, 

at depths down to 170 m. Viviparous, 2 fetuses in a litter.Feeds mainly on cephalopods, particularly octopus. 

Harmless to humans. Taken regulary in inshore artisanal fisheries in the Indo-Pacific, but of no commercial 

importance locally. Caught with floating and bottom gill nets, longlines and hook-and-line. Utilized fresh for 

human consumption; offal used for fishmeal. 

Distribution: Indo-West Pacific off southern India, Sri Lanka, Thailand, Singapore, Java, China, Taiwan 

Province of China, northern Viet Nam, the Philippines, New Guinea, and eastern, northern, and western 

Australia. 

Remarks: Differences in 

coloration and meristics 

suggest that the Australian 

? 

and possibly the northern 

Indian Ocean representatives 

of this species may be 

separated as species or 

subspecies, as indicated in 

the key.

1310 Sharks 

Hemipristis elongatus (Klunzinger, 1871) 

Frequent synonyms / misidentifications: Carcharhinus ellioti (Day, 1878) / Chaenogaleus macrostoma 

(Bleeker, 1852); Hemigaleus microstoma Bleeker, 1852. 

FAO names: En - Snaggletooth shark; Fr - Milandre chicor; Sp - Comadreja sobrediente. 

Diagnostic characters: A medium-sized shark. Body moderately slender. Snout long, its length slightly 

greater than mouth width, bluntly rounded toward tip; labial furrows moderately long and easily seen, 

the upper furrows falling well behind symphysis of lower jaw; anterior nasal flaps with a short triangular 

lobe; mouth long and semiparabolic, with a truncated lower symphysis, its length about 2/3 of the 

width; spiracles small; gill openings long, the longest over twice the eye length; teeth in upper jaw 

with strong, distally curved, broad and oblique cusps (except for erect-cusped symphyseal teeth), 

prominent distal cusplets and mesial serrations; teeth in lower jaw large but considerably narrower than 

uppers, with arched roots and long, strong, hooked, erect to oblique cusps that prominently protrude 

when mouth is closed, and basal serrations or small cusplets on more distal teeth. First dorsal fin 

moderately large, with a pointed apex and short inner margin, its origin slightly posterior to free rear tips 

of pectoral fins and the free rear tip anterior to pelvic-fin origins; second dorsal fin high, about 2/3 of 

length of first, with a short inner margin less than fin height and its origin somewhat anterior to anal-fin 

origin; anal fin slightly smaller than second dorsal fin, without long preanal ridges; upper precaudal pit 

transverse and crescentic; no keels on caudal peduncle; pectoral and pelvic fins strongly falcate, with 

deeply concave posterior margins. Intestine with a spiral valve. Colour: grey or grey-brown above, lighter 

below, no prominent markings. 

Size: Maximum total length about 2.4 m; commonly to 2 m. 

Habitat, biology, and fisheries: A rare to common tropical coastal shark, inshore and offshore at depths 

down to 130 m. Viviparous, number of young 6 to a per litter; size at birth about 45 cm. Feeds on a variety 

of fish species, also on cephalopods. Potentially dangerous because of its size and large teeth, but never 

recorded as attacking people. Regularly taken in artisanal fisheries, in the area especially off Thailand. 

Caught with bottom gill nets, floating longlines and probably on hook-and-line. Utilized fresh for human 

consumption; liver processed for vitamins; fins used in the oriental sharkfin trade, and offal for fishmeal. 


Pacific off South Africa, 

Madagascar, Mozambique, 

Tanzania, Aden, Red Sea, the 

Persian Gulf, Pakistan, India, 

Thailand, Viet Nam, China, 

the Philippines, and Australia 

(Queensland, and western 

Australia).



Frequent synonyms / misidentifications: Negogaleus tengi Chen, 1963; N. longicaudatus Bessednov, 

1964 / None. 

FAO names: En - Straight-tooth weasel shark; Fr - Milandre belette; Sp - Comadreja coluda. 

upper teeth of right side 

Diagnostic characters: A small shark. Body slender. Snout long, its length slightly 

greater than mouth width, rounded toward tip; labial furrows moderately long and 

easily seen, the upper furrows falling well behind symphysis of lower jaw; anterior 


nasal flaps with a short triangular lobe; mouth rather short and arched, its length 

about 2/3 of the width; spiracles small; gill openings moderate sized, the longest slightly longer than 

eye length in adults, slightly shorter in young; teeth in upper jaw with narrow, moderately long, 

semierect to oblique cusps and distal cusplets (except for erect-cusped symphyseal teeth), entirely 

smooth-edged; teeth in lower jaw with slightly arched roots and moderately long, mostly erect cusps 

that do not protrude when mouth is closed, and with low cusplets but no serrations. First dorsal fin 

moderately large, with a narrowly rounded apex and short inner margin, its origin slightly posterior to free 

rear tips of pectoral fins and the free rear tip anterior to pelvic-fin origins, second dorsal fin high, about 

2/3 of length of first dorsal fin, with a short inner margin less than fin height, and its origin slightly anterior 

to anal-fin origin; anal fin slightly smaller than second dorsal fin, without preanal ridges; upper precaudal 

pit transverse and crescentic, no keels on caudal peduncle; pectoral fins weakly falcate and pelvic fins 

not falcate. Intestine with a spiral valve. Colour: body grey or grey-brown above, light below, no prominent 

markings on body and fins. 


Habitat, biology, and fisheries: A little-known inshore shark, depth range not reported. Taken in fisheries 

in Thailand, and elsewhere where it occurs. 


from Japan, Taiwan Province 

of China, Viet Nam, southern 

China, and Thailand. 



1312 Sharks 

Carcharhinidae CARCHARHINIDAE 

Requiem sharks 

(also, ground sharks, blue sharks, sharpnose sharks) 


Diagnostic characters: Small to large sharks. Trunk and precaudal tail cylindrical, not depressed and 

without lateral ridges: precaudal tail much shorter than trunk. Head not expanded laterally, conical to 

moderately depressed; 5 small- to medium-sized gill slits present, the last 1 to 3 over or behind pectoral-fin 

origins, their upper ends not expanded onto dorsal surface of head; no gill sieves and usually no gill rakers 

on internal gill slits (short dermal gill rakers present in Prionace); spiracles usually absent (but always 

present in Galeocerdo); nostrils well separated from mouth, without barbels, nasoral grooves, or circumnarial 

grooves; eyes on sides of head, with a well-developed nictitating lower eyelid; snout short to 

moderately long, conical and slightly pointed to depressed and broadly rounded, never greatly flattened 

and blade-like and without lateral teeth and barbels; mouth usually large, arched and elongated, and 

extending well behind eyes; labial furrows usually present on both jaws but generally greatly reduced, 

confined to mouth corners, and barely visible when mouth is closed (but Galeocerdo and Rhizoprionodon 

species have well-developed labial furrows); upper labial furrows usually not reaching front of mouth 

(except in Galeocerdo); teeth small to large, blade-like, with a single cusp and cusplets variably 

developed; anterior teeth in upper jaw smaller than lateral teeth and not separated from them by 

smaller intermediate teeth on each side. Two dorsal fins, without spines, the first dorsal fin moderately 

large, high and angular or subangular, much shorter than the caudal fin, its base located over the interspace 

between pectoral and pelvic-fin bases and entirely anterior to origins of pelvic fins (free rear tip of dorsal 

fin may reach or extend posterior to pelvic-fin origins in Scoliodon, Negaprion, Rhizoprionodon, and 

Triaenodon); second dorsal fin varying from less than 1/5 the height of the first dorsal fin to about as high 

as the first (Lamiopsis and Negaprion); anal fin present, moderately large, with its origin varying from 

somewhat anterior to the second dorsal-fin origin to under the first half of second dorsal-fin base; caudal 

fin strongly asymmetrical, much less than 1/2 of total length, with a rippled or undulated dorsal margin, 

a well-marked subterminal notch, and a short but well-defined lower lobe; vertebral axis of caudal fin 

raised above body axis. Caudal peduncle not strongly depressed dorsoventrally or widely expanded 

laterally with weak longitudinal keels (Prionace, Galeocerdo) or none; precaudal pits present and well 

developed. Intestinal valve of scroll type. Colour: brown, grey, yellowish or bluish above, white to cream 

or yellowish below, some species with prominent dark or light markings on fins; body usually without a 

prominent colour pattern (except for Galeocerdo). 

posterior margin 

apex of 1 st dorsal fin 

inner margin 

nictitating membrane 

nostril 

internasal 

width 

anterior 

nasal flap 

labial folds 

length of snout 

width of 

mouth 

inner margin 

of pectoral fin 

position of interdorsal 

ridge when present 

rolled 

unrolled 

intestinal valve of scroll type 

second dorsal fin 

precaudal pit 

free tip of 

posterior lobe 

subterminal 

notch 

dorsal view of caudal fin 

example of upper and lower teeth 

(blade-like, with a single cusp, often serrated)

Carcharhinidae 1313 

Habitat, biology, and fisheries: The Carcharhinidae are one of the largest families of sharks and are the 

dominant sharks in tropical waters, often both in variety and in abundance and biomass. Small to very large 

species often occur close inshore, but most large ones are more abundant well offshore, but still near or over 

the continental or insular shelves. A few species, including the blue, silky, and oceanic whitetip sharks, are truly 

oceanic. Requiem sharks are active, strong swimmers, occurring singly or in small to large schools. Some 

species are continually active, while others are capable of resting motionless for extended periods on the bottom. 

Many are more active at night or dawn and dusk than the daytime. Except for the ovoviviparous tiger shark 

(Galeocerdo cuvier), all species are viviparous, with a yolk sac placenta, and have litters of young from 1 or 2 

to 135. All are voracious predators, feeding heavily on bony fishes, other sharks, rays, squid, octopuses, 

cuttlefishes, crabs, lobsters, and shrimp, but also sea birds, turtles, sea snakes, marine mammals, gastropods, 

bivalves, and carrion. The larger carcharhinids are dangerous to people, and they make up an important fraction 

of the shark species known to have attacked people. This is by far the most important shark family for fisheries 

in the tropics, and various species figure prominently in catches within the area. Most are utilized for human 

food, but also for the preparation of various subproducts, including oil and Vitamin A from the liver, fishmeal, 

and fins for the oriental soupfin markets. 

cusplets 


Hemigaleidae: intestinal valve of spiral type; also, no 

carcharhinids in the area combine the characters of 

long snout, spiracles, upper teeth with strong distal 

cusplets, long labial furrows, and second dorsal fin 

large, about 2/3 as large as first dorsal fin, with a very 

short inner margin, and with its origin anterior to that 

of the slightly smaller anal fin. 

Proscylliidae and Triakidae: no precaudal pits, dorsal 

margin not undulated, intestinal valve of spiral type, 

eyes usually dorsolateral on head (except for 

Hemigaleidae 

Hypogaleus and Galeorhinus). 

Proscylliidae 

Triakidae 

no precaudal pit 

Scyliorhinidae: first dorsal-fin base over or posterior to pelvic-fin bases (anterior to pelvic-fin bases in 

Carcharhinidae). 

Ginglymostomatidae: origin of first dorsal fin over, or only slightly anterior to pelvic-fin bases; nostrils 

connected with mouth by deep nasoral grooves, their anterior margins with a long, cylindrical barbel; eyes 

well behind mouth (over mouth in Carcharhinidae). 

Scyliorhinidae Ginglymostomatidae

1314 Sharks 

Odontaspididae: fifth gill opening in front of pectoral-fin origin; eyes without 

nictitating folds; largest teeth in front part of jaws (on either side of 

symphysis), in upper jaw separated from large teeth at sides by a gap, 

usually with 1 or 2 rows of intermediate teeth (largest teeth as sides of 

jaws and no gap in teeth row of upper jaw in Carcharhinidae). 

Other shark families: either caudal fin very long (Alopiidae), or head with 

“hammer-like” lateral projections (Sphyrnidae), or caudal fin lunate 

(Lamnidae), or size of adults much larger (Rhincodontidae), or a single 

dorsal fin and 6 or 7 gill slits (Hexanchidae), or anal fin absent (Squalidae 

and Squatinidae). 

Key to the species of Carcharhinidae occurring in the area 

1a. Upper labial furrows very long, extending to front of eyes; spiracles present and 

relatively large; prominent lateral keels on caudal peduncle (Fig. 1); vertical black or 

dusky bars on back, obscure or absent on adults . . . . . . . . . . . . . . . . . Galeocerdo cuvier 

1b. Upper labial furrows long to very short, not extending in front of eyes; spiracles usually 

absent; lateral keels usually absent (except for weak ones in Prionace glauca) (Fig. 2) . . . . . → 2 

2a. High proximal and distal cusplets present on most teeth in both jaws; expanded anterior 

nasal and mesonarial flaps forming a tube for the excurrent aperture (Fig. 3) . . Triaenodon obesus 

2b. Cusplets usually absent on lower teeth, low or absent on uppers (Fig. 4); nasal flaps 

not forming a tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3 

high cusplets 

ventral view 

of head 

Odontaspididae 

labial furrows very long 

Fig. 1 Galeocerdo cuvier 

nostril 

lateral keel 

teeth 

spiral 

ring 

scroll 

(only Carcharhinidae and Sphyrnidae) 

types of intestinal valves 

head caudal fin 

Fig. 2 other species 

Fig. 3 Triaenodon obesus Fig. 4 teeth


3a. Second dorsal fin nearly or quite as large as first (Fig. 5). . . . . . . . . . . . . . . . . . . . . → 4 

3b. Second dorsal fin considerably smaller than first (Fig. 6) . . . . . . . . . . . . . . . . . . . . . → 5 

Fig. 5 dorsal fins Fig. 6 dorsal fins 

4a. Snout short, preoral length much less than mouth width; upper and lower teeth with 

narrow, unserrated cusps (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . Negaprion acutidens 

4b. Snout longer, preoral length about equal to mouth width; upper teeth with broad, 

triangular, serrated cusps, lowers with narrow, smooth cusps (Fig. 8). . . . . Lamiopsis temmincki 

ventral view 

of head 

Fig. 7 Negaprion acutidens 

teeth 

ventral view 

of head 

5a. Head greatly depressed and trowel-shaped; pectoral fins broadly triangular, their length 

from origins to free rear tips about equal to their anterior margins; free rear tip of first 

dorsal fin about over midbases of pelvic fins; postventral margin of caudal fin usually 

only shallowly concave (Fig. 9) . . . . . . . . . . . . . . . . . . . . . . . . . Scoliodon laticaudus 

5b. Head varying from conical to slightly depressed; pectoral fins narrower, length 4/5 or 

less of anterior margin (usually less); free rear tip of first dorsal fin over or (usually) 

anterior to pelvic-fin origins; postventral margin of caudal fin deeply incised (Fig. 10). . . . . . → 6 

ventral view 

of head 

Fig. 9 Scoliodon laticaudus 

shallowly 

concave 

teeth 

Fig. 8 Lamiopsis temmincki 

lateral view of head 

Fig. 10 


1316 Sharks 

6a. Second dorsal-fin origin well behind anal-fin origin, usually over or slightly anterior to 

anal-fin insertion (Fig. 11a); preanal ridges very long and prominent, subequal to or 

greater in length than anal-fin base (Fig. 11b); posterior margin of anal fin straight or 

shallowly concave (Fig. 11b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7 

6b. Second dorsal-fin origin usually near anal-fin origin, in some species posterior to it 

(Fig. 12a); but usually well anterior to anal-fin insertion (Fig. 12b) and midbase of anal 

fin; preanal ridges variably developed, short, up to 1/2 the anal-fin base length or less 

(Fig. 12c); posterior margin of anal fin deeply concave or deeply notched (Fig. 12c) . . . . . . → 10 

preanal 

ridges 

long 

a) 

b) ventrolateral view of anal fin 

Fig. 11 

a) b) 

preanal 

ridges 

short 

c) ventrolateral view of anal fin 

Fig. 12 

7a. Posterior notches present on eyes; first dorsal-fin base 2 to 3 times in distance between 

pectoral and pelvic-fin bases (Fig. 13). . . . . . . . . . . . . . . . . . . . . Loxodon macrorhinus 

7b. No eye notches; first dorsal-fin base usually less than 2 times in distance between 

pectoral to pelvic-fin bases (up to 2 times in adult R. acutus) (Fig. 14). . . . (Rhizoprionodon) → 8 


eye 

Fig. 13 Loxodon macrorhinus 

notch 

ventral view 

of head 

Fig. 14 Rhizoprionodon 

8a. Upper labial furrows long and rather prominent, 1.4 to 2% of total length; uppers usually 

longer than lower furrows (Fig. 15a); tooth rows more numerous in average, counts 

usually 25/24 (Fig. 15b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhizoprionodon acutus 

8b. Upper labial furrows reduced and often inconspicuous, generally less than 1% of total 

length and rarely up to 1.3%; uppers usually shorter than lower furrows (Fig. 16); tooth 

rows averaging fewer, counts 23-25/21-24 but mostly below 25/24 . . . . . . . . . . . . . . . . → 9 

a) ventral view of head b) body in lateral view 

Fig. 15 Rhizoprionodon acutus 

eye 


Fig. 16


9a. Total number of enlarged hyomandibular lateral-line pores just behind mouth corners 

on both sides of head fewer, 7 to 16 and rarely above 14; precaudal vertebral centra 84 

to 91 (Fig. 17). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhizoprionodon oligolinx 

9b. Total number of enlarged hyomandibular pores greater, 15 to 22; precaudal vertebral 

centra 73 to 80 (Fig. 18). . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhizoprionodon taylori 

10a. Papillose gill rakers present on gill arches (Fig. 19a); weak lateral keels present on 

caudal peduncle; first dorsal-fin base much closer to pelvic- than to pectoral-fin bases 

(Fig. 20); colour brilliant dark blue above in life . . . . . . . . . . . . . . . . . . . Prionace glauca 

10b. No papillose gill rakers on gill arches (Fig. 19b); no lateral keels on caudal peduncle; first 

dorsal-fin base equidistant between pectoral- and pelvic-fin bases or (usually) closer to 

pectoral fins (Figs 21 and 22); colour light to dark grey, grey-brown, brown, or grey-black 

above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11 

papillose 

gill 

rakers 

Fig. 17 Rhizoprionodon oligolinx Fig. 18 Rhizoprionodon taylori 

a) 

11a. Second dorsal fin 1/2/ to 3/5 height of first dorsal fin; precaudal pits longitudinal and not 

crescentic (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Glyphis) → 12 

11b. Second dorsal fin 2/5 height of first dorsal fin or less; precaudal pits transverse and 

crescentic (Fig. 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Carcharhinus) → 14 

ventral view 

of head 

Fig. 19 

gill 

arches 

Fig. 21 Glyphis 

precaudal pit 

longitudinal 

b) 

ventral view 

of head 

Fig. 20 Prionace glauca 

lateral keel 

precaudal pit 

transverse 

Fig. 22 Carcharhinus

1318 Sharks 

12a. Head very flat and narrowly wedgeshaped 

in lateral view; total vertebral 

counts 147 to 148, diplospondylous 

caudal centra 65 to 68 (New Guinea 

and northern Australia) (Fig. 23) . . Glyphis sp. C 

12b. Head higher and broader in lateral 

view; total vertebral counts 196 to 

217, diplospondylous caudal centra 

85 to 93 . . . . . . . . . . . . . . . . . . . → 13 

Fig. 23 Glyphis sp. C 

13a. Lower anterior teeth enlarged, with cusps smooth basally but with a serrated, spear-like 

(hastate) expanded tip; total tooth row counts 55; free rear tip of first dorsal fin about 

opposite pelvic-fin origins; total vertebral counts 217, monospondylous precaudal count 

70, diplospondylous precaudal count 54; all fins with black or dusky edges and dusky 

webs in young (Queensland, Australia) (Fig. 24) . . . . . . . . . . . . . . . . . . . . Glyphissp. A 

13b. Lower anterior teeth with cusps entirely serrate and without a spear-like expanded tip; total 

tooth row counts 60 to 63; free rear tip of first dorsal fin somewhat anterior to pelvic-fin 

origins; total vertebral counts 196 to 205, monospondylous precaudal count 63 to 67, 

diplospondylous precaudal count 43 to 51; fins plain and light, except for dark patch on 

pectoral-fin bases and dusky tip on ventral caudal-fin lobe (Borneo) (Fig. 25) . . . . . . . Glyphis sp. B 

Fig. 24 Glyphis sp. A Fig. 25 Glyphis sp. B 

14a. Pectoral and first dorsal fins very broad distally and broadly rounded apically, only 

slightly tapering toward their apices; most fin tips mottled white in adults, also blacktipped 

and with black dorsal saddle-marks on the caudal peduncle in juveniles (Fig. 26) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus longimanus 

14b. Pectoral and first dorsal fins tapering distally and usually pointed or narrowly rounded; fins 

not mottled white, often black tipped but without black saddles on the caudal peduncle . . . . . . → 15 

15a. First dorsal, pectoral, pelvic, and caudal fins with extremely conspicuous white tips and 

posterior edges (Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus albimarginatus 

15b. Fins not conspicuously tipped and edged with white . . . . . . . . . . . . . . . . . . . . . . → 16 

Fig. 26 Carcharhinus longimanus Fig. 27 Carcharhinus albimarginatus 

16a. Second dorsal fin with a conspicuous black tip but other fins plain (Figs 28 and 29) . . . . . . → 17 

16b. Second dorsal fin plain, white or black-tipped but never the only fin with markings . . . . . . . → 18


17a. First dorsal fin triangular, erect, and with a posteroventrally sloping posterior margin; 

usually 13/13-14 rows of anteroposterior teeth, and 28/27 to 29 total rows of teeth; 

distal cusplets serrated on upper anterolateral teeth; pectoral-fin length 1.4 to 1.8 in 

anterior margin length; mouth width 6.4 to 8.3% of total length; precaudal centra 54 to 

74 (Fig. 28) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus dussumieri 

17b. First dorsal fin falcate, with almost vertical posterior margin (apart from free rear tip); 

usually 12/12 rows of anteroposterior teeth, and 26/25 total rows of teeth; distal cusplets 

smooth on upper anterolateral teeth; pectoral length 1.7 to 2 in anterior margin length; 

mouth width 4.2 to 6.6% of total length; precaudal centra 74 to 85 (Fig. 29) . . Carcharhinus sealei 

upper 

tooth 

Fig. 28 Carcharhinus dussumieri Fig. 29 Carcharhinus sealei 

18a. Caudal fin prominently edged with black along entire posterior edge; first dorsal fin plain 

or white-tipped, never black-tipped (Fig. 30) . . . . . . . . . . . . . Carcharhinus amblyrhynchos 

18b. Caudal fin either plain or prominently edged with black, but if black, first dorsal fin also 

prominently black-tipped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 19 

19a. Upper anterolateral teeth with bent, hooked, narrow cusps (Fig. 31). . . Carcharhinus brachyurus 

19b. Upper anterolateral teeth variably shaped, and broad or narrow, but with cusps nearly 

straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 20 

Fig. 30 Carcharhinus amblyrhynchos 

20a. Dermal interdorsal ridge present 

(Fig. 32). . . . . . . . . . . . . . . . . → 21 

20b. Dermal interdorsal ridge absent. . . . . → 27 

21a. Second dorsal fin, pectoral fin, 

and ventral caudal-fin lobe strikingly 

black-tipped . . . . . . . . . . . . → 22 

21b. Fins plain or dusky-tipped but not 

strongly black-tipped. . . . . . . . . . . → 23 

upper 

tooth 

Fig. 31 Carcharhinus brachyurus 

Fig. 32 

interorbital ridge

1320 Sharks 

22a. Second dorsal fin low, with very elongated inner margin over twice fin height; upper 

anterolateral teeth with strongly serrated cusps; usually only 12 rows of upper anteroposterior 

teeth (Fig. 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus sorrah 

22b. Second dorsal fin higher, with shorter inner margin 1.4 to 1.6 times fin height; upper 

anterolateral teeth with smooth or weakly serrated cusps; 14 or 15 rows of upper 

anteroposterior teeth (Fig. 34) . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus hemiodon 

upper 

tooth 

Fig. 33 Carcharhinus sorrah 

23a. First dorsal-fin origin well behind 

free rear tips of pectoral fins; very 

coarse serrations or small cusplets 

on feet of upper anterolateral teeth; 

inner margin of second dorsal fin 

very long, usually over twice (but 

exceptionally down to 1.6) times the 

fin height (Fig. 35) . . . Carcharhinus falciformis 

23b. First dorsal-fin origin over or anterior 

upper 

tooth 

Fig. 34 Carcharhinus hemiodon 

upper 

tooth 

Fig. 35 Carcharhinus falciformis 

to free rear tips of pectoral fins; serrations on feet of upper anterolateral teeth small and 

not very coarse; inner margin of second dorsal fin shorter and generally less than twice 

the fin height (but up to 2.1 times the fin height in Carcharhinus obscurus) . . . . . . . . . . . → 24 

24a. First dorsal-fin origin in front or over pectoral-fin insertions or at least nearer to them 

than to free rear tips of pectoral fins (Figs 36 and 37) . . . . . . . . . . . . . . . . . . . . . . → 25 

24b. First dorsal-fin origin opposite or somewhat in front of free rear tips of pectoral fin but 

closer to them than pectoral-fin insertions (Figs 38 and 39) . . . . . . . . . . . . . . . . . . . → 26 

25a. Anterior nasal flaps usually low and inconspicuous; distance from nostrils to mouth more 

than 2.4 times in mouth width; upper anterolateral teeth moderately high, usually in 14 

rows; first dorsal fin very high, its height about 1/2 of predorsal length (Fig. 36); 

interdorsal ridge low . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus plumbeus 

25b. Anterior nasal flaps usually high and triangular; distance from nostrils to mouth less 

than 2.4 times in mouth width; upper anterolateral teeth very high, usually in 15 rows; 

first dorsal fin lower, its height much less than 1/2 of predorsal length (Fig. 37); 

interdorsal ridge high . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus altimus 

upper tooth 

very high 

Fig. 36 Carcharhinus plumbeus 

upper tooth 

Fig. 37 Carcharhinus altimus


26a. Upper anterolateral teeth relatively high and narrow; pectoral fins nearly straight; first 

dorsal fin higher and with a nearly straight anterior margin; height of second dorsal fin 

2.1 to 3.3% of total length and 1.3 to 1.7 times in inner margin length; precaudal centra 

103 to 109 (Fig. 38) . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus galapagensis 

26b. Upper anterolateral teeth relatively low and broad; pectoral fins more falcate; first dorsal 

fin lower and with a rounded anterior margin; height of second dorsal fin 1.5 to 2.3% of 

total length and 1.6 to 2.1 times in inner margin length; precaudal centra 89 to 95 

(Fig. 39). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus obscurus 

upper 

tooth 

Fig. 38 Carcharhinus galapagensis 

upper 

tooth 

Fig. 39 Carcharhinus obscurus 

27a. Entire posterior margin of caudal fin with a narrow but obvious black edge; pectoral, 

second dorsal, and caudal fins with obvious black tips . . . . . . . . . . . . . . . . . . . . . → 28 

27b. Posterior margin of caudal not black or only partly dusky or black; fins black-tipped or not . . . . → 29 

28a. First dorsal fin with a broad black blotch at its apex, highlighted below with white (Fig. 40) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus melanopterus 

28b. First dorsal fin with a narrow black edge on its anterior margin but without a black blotch 

at its apex (Fig. 41) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus cautus 

Fig. 40 Carcharhinus melanopterus Fig. 41 Carcharhinus cautus 

29a. Snout very short and broadly rounded, internarial space usually less than preoral length; 

upper anterolateral teeth with very broad, triangular cusps and straight to concave distal 

margins; lower anterolaterals with strongly arched roots (Fig. 42) . . . . . . . . . . . . . . . → 30 

29b. Snout longer and parabolic or wedge-shaped to pointed, internarial space equal to, or 

greater than preoral length; upper anterolateral teeth with narrow cusps and strongly 

notched distal margins; lower anterolaterals with nearly transverse roots (Fig. 43) . . . . . . → 31 

internasal 

space 

snout 

length 


Fig. 42 

arched 

root 

lower tooth 

internasal 

space 

snout 

length 


Fig. 43 

transverse 

root 

lower tooth

1322 Sharks 

30a. Usually 11 lower anteroposterior teeth, with extremely broad cusps; first dorsal-fin 

height more than 3.1 times the second dorsal-fin height; second dorsal-fin margin 

usually nearly straight; angle of notch in anal fin posterior margin more acute, usually 

less than a right angle; precaudal centra 89 to 95 (Fig. 44) . . . . . . . Carcharhinus amboinensis 

30b. Usually 12 lower anteroposterior teeth, with moderately broad cusps; first dorsal-fin 

height more than 3.1 times the second dorsal-fin height or less; second dorsal-fin margin 

usually concave; angle of notch in anal fin posterior margin more obtuse, usually a right 

angle or more; precaudal centra 101 to 123 (Fig. 45) . . . . . . . . . . . . . . Carcharhinus leucas 

lower 

tooth 

Fig. 44 Carcharhinus amboinensis 

31a. Origin of second dorsal fin well behind anal-fin origin, about opposite its midbase . . . . . . . → 32 

31b. Origin of second dorsal fin about over anal-fin origin . . . . . . . . . . . . . . . . . . . . . . → 34 

32a. Upper anterolateral teeth with 

large mesial and distal cusplets 

and no serrations; inner margin of 

first dorsal fin extremely long, 

about 2/3 of fin base; rostrum expanded 

as a hypercalcified, hardened 

mass, easily detected by 

pinching or cutting into the snout 

(Fig. 46) . . . . . . . . Carcharhinus macloti 

32b. Upper anterolateral teeth with 

distal cusplets and serrations; inner 

margin of first dorsal fin 

shorter, 1/2 fin base or less; rostrum 

not hypercalcified . . . . . . . . . . → 33 

lower 

tooth 

upper tooth 

Fig. 45 Carcharhinus leucas 

Fig. 46 Carcharhinus macloti 

33a. Hyomandibular pores conspicuously enlarged alongside mouth corners; anteroposterior 

teeth 11-12/11-12; second dorsal fin lower, height 2.2 to 2.5 times in inner margin 

(Fig. 47) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus borneensis 

33b. Hyomandibular pores not enlarged alongside mouth corners; anteroposterior teeth 

13-15/13-14; second dorsal fin higher, height 1.5 to 1.9 times in inner margin (Fig. 48) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus sp. (= “Carcharhinus porosus”) 

upper 

tooth 

Fig. 47 Carcharhinus borneensis Fig. 48 Carcharhinus sp.


34a. Upper anterolateral teeth with 

semioblique cusps and strong cusplets; 

gill slits shorter, longest 3% of 

total length; pectoral fins rather broad 

and triangular, their lengths 1.5 in 

anterior margin length; fins not blacktipped 

(Fig. 49). . . . .Carcharhinus fitzroyensis 

34b. Upper anterolateral teeth with erect 

or nearly erect cusps and no cusplets 

(Fig. 50); gill slits longer, longest usu- 

ally at least 4% of total length; pectoral fins narrower and falcate, their lengths 1.8 or 

more in anterior margin length; fins often black-tipped . . . . . . . . . . . . . . . . . . . . . → 35 

35a. Upper labial furrows noticeably elongated and prominent; usually at least 16 rows of 

upper anteroposterior teeth; first dorsal fin lower, its height over 2.2 times in the 

interdorsal space; first dorsal-fin origin over or just behind rear tips of pectoral fins 

(Fig. 51). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus brevipinna 

35b. Upper labial furrows shorter and less noticeable; usually 15 or fewer rows of upper 

anteroposterior teeth; first dorsal fin higher, its height 2.2 times or less in interdorsal 

space; first dorsal-fin origin over or just behind insertions of pectoral fins . . . . . . . . . . . → 36 

no cusplets 

Fig. 50 upper tooth 

ventral view 

of head 

upper 

tooth 

Fig. 49 Carcharhinus fitzroyensis 

Fig. 51 Carcharhinus brevipinna 

36a. Snout rather short and wedge-shaped, internarial space 1 to 1.2 times in preoral snout; 

second dorsal height 1 to 1.2 times in inner margin length; precaudal centra usually less 

than 82 (Fig. 52) . . . . . . . . . . . . . . . . . . . . . . . . . . Carcharhinus amblyrhynchoides 

36b. Snout longer and pointed, internarial space 1.3 to 1.7 times in preoral snout (Fig. 53); 

second dorsal height 1.1 to 1.6 times in inner margin length; precaudal centra usually 

more than 83 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 37 


Fig. 52 Carcharhinus amblyrhynchoides Fig. 53 ventral view of head 

37a. Precaudal centra 94 to 102 (Fig. 54) . . . . . . . . . . . . . . . . . . . . . Carcharhinus limbatus 

37b. Precaudal centra 84 to 91 (Fig. 55). . . . . . . . . . . . . . . . . . . . . . . Carcharhinus tilstoni 

Fig. 54 Carcharhinus limbatus Fig. 55 Carcharhinus tilstoni

1324 Sharks 





Carcharhinus amblyrhynchoides (Whitley, 1934) 








Carcharhinus falciformis (Bibron in Müller and Henle, 1839). 





Carcharhinus limbatus (Valenciennes in Müller and Henle, 1839 

Carcharhinus longimanus (Poey, 1861) 

Carcharhinus macloti (Müller and Henle, 1839) 





Carcharhinus sorrah (Valenciennes in Müller and Henle, 1839) 

Carcharhinus tilstoni (Whitley, 1950) 

Carcharhinus sp. [Compagno, 1988] (= “Carcharhinus porosus“ of Garrick, 1982 for western Pacific 

specimens) 


Glyphis sp. A [Last and Stevens, 1994] (Queensland) 

Glyphis sp. B [Compagno] (Borneo) 

Glyphis sp. C [Compagno] (New Guinea, Australia) 

Lamiopsis temmincki (Müller and Henle, 1839) 









References 

Compagno. L.J.V. 1988. Sharks of the order Carcharhiniformes. Princeton, New Jersey, Princeton University Press, 

572 p. 

Garrick, J.A.F. 1982. Sharks of the genus Carcharhinus. NOAA Tech. Rep. NMFS Circ., (445)8:194 p. 

Garrick, J.A.F. 1985. Additions to a revision of the shark genus Carcharhinus: synonymy of Aprionodon and Hypoprion, 

and description of a new species of Carcharhinus. NOAA Tech. Rep., NMFS Circ., (34):26 p. 


Springer, V.G. 1964. A revision of the carcharhinid shark genera Scoliodon, Loxodon, andRhizoprionodon. Proc. U.S. 

Natl. Mus., 115:559-632. 





Frequent synonyms / misidentifications: None / Carcharhinus longimanus (Poey, 1861); Triaenodon 

obesus (Rüppell, 1837). 

FAO names: En - Silvertip shark; Fr - Requin pointe blanche; Sp - Tiburón de puntas blancas. 

Diagnostic characters: A large, slender to 

moderately stout shark. Snout moderately long and 

broadly parabolic, its length subequal to or slightly 

shorter than mouth width and equal to or greater than 

internasal space; labial furrows very short; anterior 

nasal flaps very low; spiracles absent; teeth with 

serrated edges, upper teeth broadly triangular and 

erect at front of mouth, progressively oblique 

posteriorly, without conspicuous cusplets; teeth in 

lower jaw erect and stout-cusped, serrated. First dorsal 

fin moderately high, with a narrowly rounded apex, its 

origin over inner margins of pectoral fins; second 

dorsal fin moderately high, its origin about opposite that of anal fin, its inner margin less than twice its 

height, and its posterior margin nearly straight; pectoral fins long and slightly falcate, with narrow, pointed 

tips. Interdorsal ridge present. Colour: dorsal surface dark grey or grey-brown, ventral surface white; all 

fins have conspicuous white tips and posterior margins. 

Size: Maximum total length about 3 m; adults mature at 1.6 to 1.99 m; size at birth about 63 to 68 cm. 

Habitat, biology, and fisheries: A continental and insular species occurring from the surface to a depth 

of 800 m, close inshore in lagoons and near island dropoffs or well offshore, but not oceanic. Viviparous, 

number of embryos 1 to 11. Feeds on both bottom and pelagic fish, including rays and octopi. Can be 

aggressive to divers and is potentially dangerous. Specific information on fisheries for this species is 

lacking, but it is presumably taken in areas where it occurs. Probably used fresh and dried-salted for human 

consumption. 

Distribution: In the western 

Indian Ocean recorded from 

East Africa, Madagascar and 

the Red Sea; in the western 

Pacific off southern Japan, 

from Taiwan Province of 

China southwards to 

Indonesia, and off northern 

Australia, eastern New 

Guinea and the Solomon 

Islands; also known from the 

eastern Central Pacific. 

? 


upper and lower 

tooth near centre

1326 Sharks 


Frequent synonyms / misidentifications: Carcharhinus radamae Fourmanoir, 1961 / Carcharhinus 

albimarginatus (Rüppell, 1837); C. galapagensis (Snodgrass and Heller, 1905); C. obscurus (Lesueur, 

1818). 

FAO names: En - Bignose shark; Fr - Requin babosse (= Réquiem babosse, Area 31); Sp - Tiburón 

baboso. 

Diagnostic characters: Body slender. Snout 

rounded and moderately long, its length about 

equal to, or greater than, mouth width, and greater 

than internasal space; labial furrows very short; 

anterior nasal flaps expanded as low, broadly 

triangular lobes; spiracles absent; teeth with 

serrated edges, upper teeth broadly triangular and 

erect in front of mouth, with very high cusps, 

progressively oblique posteriorly; teeth in lower jaw 

erect and narrow-cusped. First dorsal fin 

moderately high with a narrowly rounded apex, its 

origin over inner margins of pectoral fins; second 


dorsal fin high, its origin about opposite that of anal fin, its inner margin less than 1.5 times the fin-height; 

pectoral fins long and not strongly falcate, broad-tipped but with angular apices. A high interdorsal 

ridge present. Colour: back greyish; belly whitish; inner corners of pectoral fins blackish. 

Size: Maximum total length to about 3 m; commonly to 2.4 m; size at birth probably 70 to 90 cm. 

Habitat, biology, and fisheries: Usually found in the edges of the continental shelves and uppermost 

slopes near the bottom, ranging from a depth of 30 to 430 m, but commonly between 80 and 220 m; rare 

in shallow waters. Viviparous. Bottom-dwelling; feeds chiefly on fishes and cephalopods. Apparently taken 

on deep-set longlines, also in bottom trawls and probably on hook-and-line and with gill nets. Utilized for 

fishmeal, liver oil, and shagreen. 

Distribution: Circumglobal, 

with patchy records in tropical 

and warm seas. 

? 

? 


tooth near centre


Carcharhinus amblyrhynchoides (Whitley, 1934) 

Frequent synonyms / misidentifications: Carcharhinus pleurotaenia (Bleeker, 1852) / Carcharhinus 

limbatus (Valenciennes in Müller and Henle, 1839), C. brevipinna (Müller and Henle, 1839). 

FAO names: En - Graceful shark; Fr - Requin gracile; Sp - Tiburón grácil. 

Diagnostic characters: A medium-sized stout 

shark. Snout pointed but short, its length less 

than mouth width, 1.0 to 1.2 times internasal 

space; labial furrows very short; anterior nasal 

flaps very low; no spiracles; upper and lower 

teeth with serrated edges, including cusps, 

upper teeth with narrow cusps, and no 

prominent cusplets, not broadly triangular, 

cusps of upper anterior teeth erect, laterals 

erect to oblique; lower teeth erect and 

narrow-cusped. First dorsal fin moderately 

high, with an angular or narrowly rounded apex 

and short inner margin, its origin over inner margins of pectoral fins; origin of second dorsal fin about 

opposite anal-fin origin; second dorsal fin moderately high, its inner margin less than twice its height, 

its posterior margin concave; pectoral fins moderately long and falcate, with narrow, pointed tips. No dermal 

ridge between dorsal fins. Colour: grey or grey-brown on dorsal surface, white or cream below, with a 

conspicuous band of white on sides from pelvic fins to first dorsal fin; pectoral, dorsal, and pelvic fins, 

and ventral lobe of caudal fin black or dusky-tipped, sometimes inconspicuously so. 

Size: Maximum total length about 1.8 m; size at birth about 52 to 55 cm. 

Habitat, biology, and fisheries: A poorly known, inshore, coastal pelagic species. Viviparous. Feeds 

mainly on fish, also crustaceans and cephalopods. Taken on longlines and drifting gill nets; in the area it 

is caught in the Gulf of Thailand, northern Australia and probably elsewhere where it occurs. Utilized fresh 

and dried salted for human 

consumption; fins used in the 

oriental sharkfin trade; liver oil 

is processed for vitamins. 

Distribution: In the western 

Indian Ocean from the Gulf of 

Aden, southern India and Sri 

Lanka, in the western Pacific 

from Viet Nam, Thailand, the 

Philippines, Java, Borneo, 

Australia, and New Guinea. 

? 

? 



tooth near centre

1328 Sharks 


Frequent synonyms / misidentifications: Carcharhinus menisorrah (Valenciennes in Müller and Henle, 

1839); ? Carcharhinus wheeleri Garrick, 1982 / None. 

FAO names: En -Greyreefshark;Fr - Requin dagsit; Sp - Tiburón de arrecifes. 

Diagnostic characters: A medium-sized shark. 

Body rather stout. Snout broadly rounded, its 

length less than mouth width, equal to or 

somewhat greater than internasal space; labial 

furrows very short; anterior nasal flaps very low, 

no spiracles; teeth with serrated edges but with 

cusplets low or absent on upper teeth, always 

absent on lowers; upper teeth narrowly 



triangular, high, moderately narrow and 

tooth near centre 

erect-cusped in front of mouth, progressively oblique posteriorly; teeth in lower jaw mostly erect and 

narrow-cusped. First dorsal fin moderately high and with narrowly rounded apex, its origin over inner 

margins of pectoral fins; origin of second dorsal fin about opposite anal-fin origin; second dorsal fin 

moderately high, its inner margin less than 1.5 times the fin height and its posterior margin deeply notched; 

pectoral fins long and not strongly falcate, with narrow, angular apices. A weak interdorsal ridge present 

between dorsal fins, or no ridge. Colour: dark grey or bronze-grey above, white below; caudal fin with 

a conspicuous wide black posterior margin; undersides of pectoral and pelvic fins with black tips and 

posterior margins, but fins otherwise not conspicuously black, or white-tipped except for white-tipped first 

dorsal fin in some individuals. 

Size: Maximum total length about 2.55 m; commonly to 1.8 m; size at birth about 50 to 60 cm 

Habitat, biology, and fisheries: An inshore shark, most common over coral reefs, often near the bottom. 

Viviparous, number of young per litter 1 to 6. A bottom-feeding shark, eating small reef fishes and 

octopuses. Aggressive, particularly when attracted by spearfishing or when cornered by divers: when 

provoked, gives an exaggerated swimming display, with back arched and pectoral fins thrust downward, 

followed by a swift attack and bite if the provocation continues. Several divers have been bitten by this shark 

in the western Pacific, none fatally, and even small diver-operated submarines have been attacked. Fished 

in Thailand and elsewhere mainly for the flesh and fins. 

Distribution: Known from 

Madagascar and the 

Mauritius-Seychelles area, 

possibly also from India; also 

Red Sea to South Africa if 

Carcharhinus wheeleri is 

synonymized with this 

species; in the western 

Central Pacific from Sumatra 

eastward to the Philippines, 

Australia, New Guinea, 

Hawaii, and the Tuamotu 

Archipelago.



Frequent synonyms / misidentifications: Triaenodon obtusus Day, 1878 / Carcharhinus leucas 

(Valenciennes in Müller and Henle, 1839); Glyphis gangeticus (Müller and Henle, 1839). 

FAO names: En - Pigeye shark; Fr - Requin balestrine; Sp - Tiburón baleta. 

Diagnostic characters: A medium to large, 

stout-bodied shark. Snout extremely short (usually 

shorter than distance between nostrils, and much 

shorter than mouth width), very broadly rounded; labial 

furrows very short; spiracles absent; nostrils with a low, 

broadly triangular anterior nasal flap; teeth in upper jaw 

triangular, with broad, heavy, serrated cusps, their 

outer edges nearly straight in anterior teeth but 

becoming increasingly concave in lateral teeth; cusps of 

lower teeth heavy, erect to slightly oblique with serrated 

edges, their bases strongly arched.First dorsal fin very 

high (its height 3.2 or more times that of second dorsal fin) with a pointed or slightly rounded apex, its origin a 

little in advance of insertions of pectoral fins; second dorsal fin low, with its inner margin about equal to fin 

height, its posterior margin nearly straight, and its origin slightly in front of anal fin; pectoral fins large, broad, 

with narrow, pointed tips. No dermal ridge between dorsal fins. Colour: grey above, light below, tips of fins 

darker in young, fading in adults. 

Size: Maximum total length about 2.8 m, maturing at about 2.1 to 2.2 m; size at birth between about 43 to 

53 cm length. 

Habitat, biology, and fisheries: An inshore as well as offshore continental species occurring from the 

surfline to a depth of 150 m. Found in shallow bays and estuaries as well as off open coast but apparently 

not ascending rivers like Carcharhinus leucas or Glyphis spp. Viviparous, number of young 3 to 13 per 

litter. Feeds on a wide variety of demersal and pelagic bony fishes (which are its most important prey), 

sharks and rays, squid, cuttlefish and octopuses, and lobsters and shrimp. Uncommonly scavenges prey 

and rarely feeds on marine mammals. Potentially dangerous to people, but not recorded in shark attacks 

to date and perhaps less 

inclined to attack people than 

C. leucas because of its 

narrower prey spectrum. 

Caught on longlines and in gill 

nets and utilized fresh and 

dried-salted. 

Distribution: Known from off 

South Africa, Madagascar, the 

Gulf of Aden, Pakistan, and Sri 

Lanka; elsewhere from the 

eastern Atlantic (Nigeria) and 

western South Pacific. 

? 

? 

? 

ventral view 

of head 


tooth near centre

1330 Sharks 



FAO names: En - Borneo shark; Fr - Requin-tigre houareau; Sp - Tiburón de Borneo. 

Diagnostic characters: A small shark. Body 

relatively slender. Snout very long and 

pointed, its length greater than mouth width and 

distance between nostrils; labial furrows very 

short; anterior nasal flaps high and narrow, 

nipple-shaped; spiracles absent; teeth with 

serrated edges, those in upper jaw with 

narrow, oblique cusps and large cusplets on 

each side of cusp; teeth in lower jaw with erect 

to oblique, narrow serrated cusps, weak 

cusplets or large serrations and transverse 

roots. First dorsal fin moderately large, with a 

ventral view 

of head 


tooth 

blunty pointed apex, its inner margin moderately long, the free rear tip attenuated, and its origin slightly 

anterior inner margins of pectoral fins; second dorsal fin very low, the inner margin over twice the fin 

height, fin origin over or slightly behind midbase of anal fin, pectoral fins relatively short, with narrowly 

rounded or angular tips. No dermal ridge between dorsal fins. Colour: brown above, white below, tip of 

first dorsal fin and dorsal caudal-fin margin dusky, paired fins and anal fin with light edges, but markings 

not conspicuous. 


Habitat, biology, and fisheries: A rare coastal, inshore, tropical shark, with biology virtually unknown. 

Apparently rare, but undoubtedly taken in local fisheries. 

Distribution: Recorded in 

the Indo-West Pacific from 

China, Borneo, and possibly 

from Java and the Philippines.



Frequent synonyms / misidentifications: Carcharhinus remotus (Dumeril, 1865) / None. 

FAO names: En - Copper shark; Fr - Requin cuivre; Sp - Tiburón cobrizo. 

Diagnostic characters: A large shark. Body slender to 

moderately stout. Snout rounded or broadly angular, its 

length about equal to, or somewhat smaller than width 

of mouth, but greater than internasal space; labial 

furrows short; anterior nasal flaps very short to 

rudimentary; upper teeth with narrow, mostly oblique, 

somewhat flexed cusps, well delimited from the 

tooth bases and finely serrated; lower teeth with 

moderately high, narrow, erect to semioblique, weakly 

serrated, cusps; gill slits relatively short. First dorsal 

fin moderately high, with a nearly straight anterior margin 

and a narrowly rounded or pointed apex, its origin over 

inner margins of pectoral fins; second dorsal fin moderately high, with a slightly concave posterior margin 

and an inner margin much shorter than half the height of fin; its origin over that of anal fin; pectoral fins not 

strongly falcate, apically pointed. Usually no interdorsal ridge (occasionally a weak ridge present). Colour: 

dark brownish grey above, white below; fins mostly plain, except for dusky tips on pelvic fins, as well as 

dusky to black tips and rear edges on pectoral fins. 

Size: Maximum total length about 2.9 m; matures at between 2 to 2.5 m, with females somewhat larger 

than males; size at birth about 60 to 70 cm. 

Habitat, biology, and fisheries: A coastal and offshore shark, preferring temperate to tropical waters. 

Viviparous, number of fetuses 13 to 20. Feeds on bottom-dwelling bony fishes, including gurnards, 

flatfishes, hakes, puffers, sea catfishes, jacks, and mullets; also on rays, small sharks, squids, and 

cuttlefishes. Dangerous or potentially dangerous to man, known to have been implicated in shark attacks 

on people in other areas. Little is recorded on the use of this species, but it is undoubtedly caught and used 

for human consumption. It is 

taken in bottom trawls, by line 

gear, and by sports anglers. 

Distribution: Nearly circumglobal 

in temperate, subtropical 

and some tropical 

seas (California, South 

America, West and South 

Africa, the Mediterranean, 

Japan, China, New Zealand, 

Australia). 

? 

ventral view 

of head 


lateral teeth

1332 Sharks 


Frequent synonyms / misidentifications: Aprionodon brevipinna (Müller and Henle, 1839); 

Carcharhinus johnsoni Smith, 1951 / Carcharhinus limbatus (Valenciennes in Müller and Henle, 1839); 

C. amblyrhynchoides (Whitley, 1934). 

FAO names: En - Spinner shark; Fr - Requin tisserand (= Requiem tisserand, Area 31); Sp - Tiburón aleta 

negra. 

Diagnostic characters: A slender-bodied medium 

to large-sized shark. Snout pointed and long, its 

length equal to or greater than mouth width and 

greater than internasal space; labial folds short, 

but usually the upper pair longer and more 

prominent than in other Carcharhinus species 

from the area; anterior nasal flaps rudimentary, very 

low; upper and lower teeth nearly symmetrical 

and very similar, with mostly erect, very narrow 

cusps; uppers with entirely or partly serrated 

edges, lowers smooth; gill slits relatively long. 

First dorsal fin with a narrowly rounded apex, its 

origin above or slightly behind free rear tips of pectoral fins; second dorsal fin high, its inner margin 

less than twice the height of fin, its origin about over that of anal fin; pectoral fins falcate and with pointed 

tips. No dermal ridge between dorsal fins. Colour: grey on back, white below, with a conspicuous white 

band on sides; second dorsal, anal, undersides of pectorals and lower caudal-fin lobe black or dark 

grey-tipped in subadults and adults, but unmarked or nearly so in small individuals (below 1 m). 

Size: Maximum total length about 2.8 m (mature adults); commonly to 2.5 m; size at birth about 60 to 80 cm. 

Habitat, biology, and fisheries: An active, fast-swimming shark, often making vertical spinning leaps out of 

the water, as a feeding technique in which the shark spins through a school of small fish with open mouth and 

then breaks the surface. Viviparous, number of embryos about 6 to 15. Feeds mostly on small schooling fishes, 

also squid, small sharks, and rays. Caught with drifting gill nets and longlines. Utilized fresh and dried salted for 

human consumption, fins probably used in the oriental sharkfin trade, and livers for vitamin oil production. 

Distribution: Known in the western Indian Ocean from South Africa to Madagascar and the 

Mauritius-Seychelles area, the 

Red Sea, Gulf of Oman, and 

southern India; in the western 

Pacific from southern Japan, 

Viet Nam, Indonesia, 

Australia, and possibly the 

Philippines; also found in the 

eastern Atlantic and the 

Mediterranean, but apparently 

absent from the eastern 

Pacific. 

? 

ventral view 

of head 


tooth near centre



Frequent synonyms / misidentifications: None / Carcharhinus melanopterus (Quoy and Gaimard, 

1824). 

FAO Names: En - Nervous shark; Fr - Requin nerveux; Sp - Tiburón nervioso. 

Diagnostic characters: A small to medium-sized shark. Body moderately 

stout. Snout very short and broadly rounded, its length less than mouth 

width and about equal to distance between nostrils; labial furrows very 

short; anterior nasal flaps with a stout, broad lobe; spiracles absent; teeth with 

serrated edges, those in upper jaw with narrow semioblique to oblique 

cusps and low basal cusplets; teeth in lower jaw with erect or semierect 

narrow cusps and serrated edges. First dorsal fin moderately large, with a 

narrowly rounded or pointed apex, its origin over inner margins of pectoral fins, 

its free rear tip short; second dorsal fin high, its inner margin much less than 

twice the fin height, its origin over or slightly anterior to anal-fin origin; pectoral 

fins moderately long, with narrowly rounded or pointed tips; rear tip of anal fin 

ending well in front of lower caudal-fin origin. No dermal ridge between dorsal 

ventral view 

of head 

fins. Colour: grey or light brown above, white below; dorsal, caudal and pectoral fins with black margins, 

expanded apically to black tips on caudal-fin lobes and pectoral fins; probably a conspicuous white band 

on flank. 

Size: Maximum total length about 1.5 m, adult females 1.2 to 1.5 m; size at birth between 35 and 39 cm. 

Habitat, biology, and fisheries: The nervous shark is a little-known South Pacific reef shark that may have 

a wider distribution. It apparently lives in shallow water on the continental and insular shelves, but may 

range in deeper water. Skittish and timid when approached by people, hence the vernacular name “nervous 

shark”. Presumably viviparous. Feeds on small fishes, including lizardfish and smelt-whiting (Sillago), and 

crabs. Probably harmless or minimally hazardous to people. Taken in small numbers in northern Australia 

for its meat. 

Distribution: In the eastern Indian Ocean and western South Pacific off Australia (Queensland, western 

and northern Australia), Ugi 

and the Solomon Islands. 

Remarks: This species is 

similar to the blacktip reef 

shark, Carcharinus melanopterus, 

but lacks the conspicuous 

highlighted black blotch 

on its first dorsal fin, and has 

lower vertebral counts.

1334 Sharks 



1839); C. tjutjot (Bleeker, 1852) / Carcharhinus sealei (Pietschmann, 1916). 

FAO names: En - Whitecheek shark; Fr - Requin à joues blanches; Sp - Tiburón cariblanco. 


moderately stout. Snout moderately long and 

broadly parabolic or wedge-shaped, its length 

usually shorter than mouth width but subequal to the 

internasal space; labial furrows very short; anterior 

nasal flaps expanded; spiracles absent;gill slits short; 

teeth with serrated edges, upper teeth with 

narrow-based,strongly oblique cusps and strong, 

serrated cusplets;teeth in lower jaw erect to oblique, 

without cusplets, serrated and narrow-cusped; first 

dorsal fin moderately high, with an angular apex, 

posteroventrally sloping, straight posterior margin, 

ventral view 

of head 



and short inner margin, not falcate; origin of first dorsal fin over pectoral fin inner margins; origin of second 

dorsal fin about opposite that of anal fin; second dorsal fin high, its inner margin less than 1.5 times the fin height 

and its posterior margin concave; pectoral fins short and not strongly falcate, with narrow, angular apexes. An 

interdorsal ridge present or occasionally absent on back. Colour: back greyish or grey-brown, belly whitish; 

a black spot on the second dorsal fin is the only conspicuous marking. 

Size: Maximum total length about 90 cm; size at birth about 35 to 40 cm. 

Habitat, biology, and fisheries: A common, but little-known shark of the continental and insular inshore 

waters. Viviparous, number of young 1 to 4, usually 2. Feeds primarily on fish, also crustaceans and 

cephalopods; harmless to people. Caught with drifting gill nets and longlines in artisanal and smallscale 

industrial fisheries and is commonly marketed for its meat for human consumption. 

Distribution: Occurs from the Persian Gulf eastward to Thailand, China, southern Japan, Java, Borneo, and 

probably New Guinea and 

northern Australia. 

Remarks: This species is very 

similar to the blackspot shark, 

Carcharhinus sealei but has a 

triangular rather than falcate 

first dorsal fin, more numerous 

upper teeth, a broader mouth, 

broader pectoral fins, and less 

numerous vertebrae.


Carcharhinus falciformis (Bibron in Müller and Henle, 1839) 


FAO names: En - Silky shark; Fr - Requin soyeux (= Réquiem soie, Area 31); Sp - Tiburón jaquetón. 

Diagnostic characters: A large shark, with an 

elongate and slender body. Snout narrowly 

rounded, moderately long, equal to or slightly 

shorter than mouth width, but longer than internasal 

space; labial furrows very short; anterior nasal flaps 

low, rudimentary; spiracles absent; upper teeth 

with relatively narrow cusps well delimited from 

the heavy, serrated bases, their outer edges 

notched; teeth in lower jaw erect, their edges only 

slightly serrated. First dorsal fin moderately high, its 

apex rounded, its origin behind the free rear tips 

of pectoral fins; second dorsal fin very low, its 

ventral view 

of head 



posterior lobe noticeably long and slender, its inner margin twice the height of fin, its origin about over that 

of anal fin; pectoral fins long and falcate, more so in adults than in young; interdorsal ridge present. Colour: 

back dark grey, greyish brown or bluish black (in life); belly greyish or white. 

Size: Maximum total length 3.5 m; commonly to 2.5 m; size at birth 57 to 87 cm. 

Habitat, biology, and fisheries: Inhabits oceanic waters near and beyond the continental slopes, but also found 

in coastal waters. Lives usually near the surface, but occurs sometimes at considerable depths (to 500 m). 

Viviparous, number of young 2 to 14 per litter. Feeds chiefly on fishes, including tunas, squids, and pelagic 

octopuses. Very quick in its movements, it often causes damage to the catch and gear in tuna fisheries. Reported 

to be dangerous to humans. This species is very commonly taken by pelagic longline fisheries, but is also taken 

in fixed bottom nets and probably also drifting gill nets. Its meat is used fresh or dried-salted for human 

consumption, its hide for leather, its fins for shark-fin soup, and its liver is extracted for oil, which has a high 

Vitamin A content. 

Distribution: Circumglobal in 

tropical and subtropical seas; 

in the western Atlantic from 

Massachusetts to southern 

Brazil, in the eastern Atlantic 

from Madeira to northern 

Angola, in the Indian Ocean 

off East Africa, Red Sea, Gulf 

of Oman and South India, 

widespread in the western 

Pacific (including Japan and 

New Zealand), and in the 

eastern Central Pacific from 

southern Baja California to 

Peru.

1336 Sharks 



FAO names: En - Creek whaler; Fr - Requin baleinier; Sp - Tiburón ballenero. 


shark. Body moderately stout. Snout long 

and parabolic, its length greater than mouth 

width and distance between nostrils; labial 

furrows very short; anterior nasal flaps 

moderately low, nipple-shaped; spiracles 

absent; teeth with serrated edges, those in 

upper jaw with narrow, semierect and oblique 

high cusps, and crown feet with coarse 

serrations and small cusplets; teeth in lower jaw 

with erect, narrow, partly serrated cusps, and 

transverse roots. First dorsal fin moderately 

ventral view 

of head 


tooth 

large, with a pointed or narrowly rounded apex, its inner margin short, and its origin over or usually 

slightly anterior to inner margins of pectoral fins; second dorsal fin large and high, the inner margin 

1.5 times the fin height, fin origin about over anal-fin origin, pectoral fins moderately large, broad and 

triangular, with narrowly rounded apices. No dermal ridge between dorsal fins. Colour: bronze above, 

fading to greyish brown after death and in preserved specimens; light below, without conspicuous markings 

on fins; no conspicuous white band on flanks. 

Size: Maximum total length about 1.35 m; matures at 80 to 90 cm; size at birth about 50 cm. 

Habitat, biology, and fisheries: A little-known tropical shark of the Australian littoral, found inshore and 

offshore on the continental shelves from the intertidal to a depth of at least 40 m. Feeds primarily on teleost 

fishes, also on crustaceans. Taken in small numbers off northern Australia, and is used for human 

consumption. 


off Australia (Queensland, 

northern and northwestern 

Australia).




FAO names: En - Galapagos shark; Fr - Requin de Galapagos; Sp - Tiburón de Galápagos. 

Diagnostic characters: A large shark. Body 

slender to moderately stout. Snout rounded and 

short, its length equal to or less than mouth 

width and about equal to or greater than 

internarial space; labial furrows short; anterior 

nasal flaps rudimentary; upper teeth broadly 

triangular, erect to moderately oblique, the 

anterior ones strongly serrated and with 

higher, broad cusps not delimited from the 

bases; lower teeth with high, narrow cusps and 

serrations; gill slits relatively short. First dorsal 

fin rather high, nearly straight anteriorly, with 

ventral view 

of head 


lateral tooth 

a narrowly rounded or pointed apex, its origin over inner margins of pectoral fins; second dorsal fin 

moderately high, with a concave posterior margin, its inner margin less than twice the fin height and 

its origin over or slightly anterior to that of anal fin; pectoral fins nearly straight and apically pointed. A 

low interdorsal ridge present. Colour: dark grey above, light below, fins plain except for slightly dusky 

tips in some individuals. 

Size: Maximum total length about 3.7 m; commonly to 3 m; size at birth about 57 to 78 cm. 

Habitat, biology, and fisheries: A wide-ranging, inshore and offshore shark often preferring the waters 

around islands to those of the continental shelf. Viviparous, number of fetuses 6 to 16. Feeds on bottom 

fishes, including basses, flatheads, eels, and flatfishes; also on cephalopods and bivalves. An aggressive 

species, dangerous to people. No information on utilization or fishing methods are available, but likely to 

figure in shark fisheries because of its abundance in habitats if prefers. 


in tropical and subtropical 

seas, but of spotty 

occurrence in the Pacific and 

Atlantic, primarily off island 

groups but offshore in 

continental waters in the 

eastern Pacific.

1338 Sharks 


Frequent synonyms / misidentifications: Hypoprion hemiodon (Valenciennes in Müller and Henle, 

1839) / None. 

FAO names: En - Pondicherry shark; Fr - Requin baliai; Sp - Tiburón de Pondicherry. 

Diagnostic characters: A small shark (maximum 

size unknown). Body rather stout. Snout 

moderately pointed and parabolic, its length 

equal to or slightly less than mouth width and 

greater than internasal space;labial furrows short; 

anterior nasal flaps with a short, slender, 

narrow lobe; upper teeth with oblique or 

semioblique, narrow, unserrated or weakly 

serrated cusps and strong distal cusplets, 

lowers with erect cusps, no cusplets and smooth 

edges; gill slits relatively short. First dorsal fin with 

a narrowly rounded apex, its origin just posterior 

ventral view 

of head 



to pectoral-fin base insertions and over inner margins of pectoral fins, its inner margin and free rear tip 

moderately long; second dorsal fin moderately high, its inner margin attenuated and elongated but less 

than twice the height of the fin, its origin slightly behind that of anal fin; pectoral fins weakly falcate and with 

narrowly rounded tips. A dermal ridge present between dorsal fins. Colour: grey above, white below, with 

the tips of the pectorals, and upper and lower caudal-fin lobes black; other fins dusky. 

Size: Maximum total length uncertain, probably not over 1.5 to 2 m; immature specimens up to 60 cm long, 

size at maturity unknown. 

Habitat, biology, and fisheries: A little-known, wide-ranging, possibly common grey shark of the continental 

shelves. Has been reported from both river mouths and rivers, including the Saigon River in Viet Nam, but these 

old records require confirmation. Presumably viviparous. Diet unknown, presumably small fishes, cephalopods, 

and crustaceans. Not known to be dangerous to humans. Caught in bottom-set gill nets and presumably on 

long gear. Utilized fresh for 

human consumption. 

Distribution: Occurs from the 

Gulf of Oman to Pakistan, 

India and possibly Sri Lanka; 

also known in the eastern 

Indian Ocean and western 

Pacific from scattered 

localities from India eastward 

to Viet Nam, the Philippines, 

China, and Indonesia.



Frequent synonyms / misidentifications: Carcharhinus zambezensis (Peters, 1852); C. vanrooyeni Smith, 

1958 / Carcharhinus amboinensis (Müller and Henle, 1839). 

FAO names: En - Bull shark; Fr - Requin bouledogue (= Réquiem taureau, Area 31); Sp - Tiburón sarda. 

Diagnostic characters: A large, stout shark. Snout very 

broadly rounded and extremely short, its length less than 

distance between nostrils, and much less than mouth width; 

labial furrows very short; spiracles absent; nostrils with a low, 

broadly triangular anterior nasal flap; teeth in upper jaw 

triangular, with broad, heavy, serrated cusps, their outer 

edges nearly straight in anterior teeth, but becoming increasingly 

concave to the sides; lower teeth with erect to slightly oblique, 

heavy cusps with serrated edges and strongly arched bases. 

First dorsal fin high and broad with a pointed or slightly 

rounded apex, its origin a little in advance of insertion of 

ventral view 

of head 



pectoral fins; second dorsal fin high with a short posterior lobe, its inner margin less than the fin 

height, and its origin slightly in front of that of anal fin; pectoral fins broad, with narrow pointed tips. No 

interdorsal ridge. Colour: back greyish, belly white; tips of fins dark, especially in young individuals. 

Size: Maximum total length about 3.4 m; commonly to 2.6 m; size at birth about 60 cm. 

Habitat, biology, and fisheries: Predominantly a coastal and fresh-water species inhabiting shallow waters, 

especially in bays, river estuaries, rivers, and lakes. It tolerates a wide range of salinities, readily penetrates 

far up rivers and also into hypersaline bays. Usually slow-swimming if active while cruising, this 

bottom-living shark may develop great speed when chasing its prey. Viviparous, number of embryos up to 

12. The young readily tolerate low salinities, and some are born in fresh water. An opportunistic predator 

with a very wide food spectrum that includes bony fishes, sharks, rays, invertebrates (crabs, shrimps, sea 

urchins, etc.), marine and freshwater turtles, birds, marine and terrestrial mammals, and carrion. It has 

large strong jaws and large stout teeth for its size, which enable it to dismember and feed on relatively large 

prey. Known to be dangerous to people, and possibly one of the most dangerous sharks because of its 

inshore and fresh-water habitat, large size, powerful feeding structures, and omnivorous habits. Caught 

mainly with longlines and gill 

nets and used for its meat, 

hide, fins, liver oil, and as 

fishmeal. 

Distribution: Widespread 

along the continental coasts of 

all tropical and subtropical 

seas; also, the most 

wide-ranging cartilaginous fish 

in fresh water.

1340 Sharks 

Carcharhinus limbatus (Valenciennes in Müller and Henle, 1839) 

Frequent synonyms / misidentifications: None / Carcharhinus tilstoni (Whitley, 1950); C. brevipinna 

(Müller and Henle, 1839); C. amblyrhynchoides (Whitley, 1934). 

FAO names: En - Blacktip shark; Fr - Requin bordé (= Réquiem macuire, Area 31); Sp - Tiburón macuira. 

Diagnostic characters: Body fusiform, 

moderately slender. Snout long, about equal to, 

or slightly less than mouth width, greater than 

distance between nostrils, its tip narrowly 

rounded to pointed; labial furrows short; spiracles 

absent; upper and lower teeth nearly 

symmetrical and similar, with erect, narrow 

cusps and serrated edges; gill slits moderately 

long. First dorsal fin with a pointed or very 

narrowly rounded apex, its origin above, or 

slightly posterior to insertion of pectoral fins; 

second dorsal fin high, its inner margin less than twice the height of fin, and its origin over or slightly in 

front of that of anal fin; pectoral fins falcate. No interdorsal ridge. Colour: back dark grey, ashy blue or 

dusky bronze; belly white or yellowish white; a dark band extending rearward along each side to about 

over origin of pelvic fin; tips of pelvic fins with a persistent black spot; tips of dorsals, anal, pectorals 

and the lower lobe of caudal fin usually black or dusky in young individuals, but these markings fade with 

growth. 

Size: Maximum total length about 2.5 m; commonly to 1.5 m; size at birth about 60 cm. 

Habitat, biology, and fisheries: Inhabits coastal as well as offshore surface waters. A fast-moving shark 

that sometimes leaps out of the water. Occasionally enters brackish waters, but without a tolerance for 

fresh water. Viviparous, number of embryos ranging from 1 to 10. Feeds mainly on small schooling fishes; 

also on rays and squids. Apparently minimally dangerous to people, but can be aggressive when divers 

are spearing fish. Caught with floating longlines, floating gill nets, and probably other gear. Marketed fresh 

for human consumption, oil valuable for Vitamin A. 



continental waters. 

? 

ventral view 

of head 


tooth near centre


Carcharhinus longimanus (Poey, 1861) 

Frequent synonyms / misidentifications: Carcharhinus maou (Lesson, 1830) / None. 

FAO names: En - Oceanic whitetip shark; Fr - Requin océanique (= Réquiem océanique, Area 31); 

Sp - Tiburón oceánico. 

Diagnostic characters: A large, moderately stout shark. 

Snout short and broadly rounded, its length equal to, 

or somewhat less than, mouth width, and greater than 

distance between nostrils; labial furrows very short; 

anterior nasal flaps very low, rudimentary; spiracles 

absent; teeth with serrated edges, those in upper jaw 

triangular with broad, heavy, mostly erect, cusps 

nearly symmetrical anteriorly, but becoming increasingly 

oblique at sides; teeth in lower jaw with erect, heavy 

cusps and serrated edges. First dorsal fin noticeably 

ventral view 

of head 



large, with a very broadly rounded apex, its origin slightly behind insertion of pectoral fins; second dorsal 

fin high, its inner margin less than twice the fin height, its origin over, or slightly in front of that of anal fin; 

pectoral fins very long (as long as, or even longer than, head) with broadly rounded, wide tips; rear tip 

of anal fin extending nearly to origin of caudal fin. An interdorsal ridge present. Colour: back usually 

dark grey with a bronze tinge, but sometimes brown or bluish; belly whitish, sometimes with a yellow tinge; 

tips of first dorsal fin, pectoral fins and lower lobe of caudal fin often white or with white spots 

(sometimes absent); ventral surface of pelvic fins, apices of anal and second dorsal fins, and ventral lobe 

of caudal fin often with black spots; also black or dusky saddle-marks in front of second dorsal fin, upper 

margin of caudal fin and between dorsal fins (especially in young). 

Size: Maximum total length about 3.5 m; commonly to 2.7 m or less; size at birth 60 to 65 cm. 

Habitat, biology, and fisheries: Along with the silky shark (Carcharhinus falciformis), this is one of the most 

abundant sharks in warm oceanic waters. It occasionally enters coastal waters, but is more typically found from 

the edges of continental or insular shelves to far beyond land. This is a slow-swimming species while cruising, 

but it can be fast in pursuit of prey. Viviparous, number of embryos ranging from 6 to 9. Feeds mainly on fishes 

(especially scombrids and carangids) and squids; also crustaceans (especially portunid crabs), turtles, and 

carrion. This species causes much damage to the catch in tuna fisheries, and formerly also to dead whales that 

were inflated and buoyed after harpooning by the whaling ships. Reported to be dangerous to humans, and 

prone to investigate divers and 

swimmers that venture into its 

offshore habitat. Caught with 

floating longlines, also drifting 

gillnets and handlines. Utilized 

fresh for human consumption, 

also processed for liver oil; fins 

probably used for the oriental 

sharkfin trade. 



waters.

1342 Sharks 

Carcharhinus macloti (Müller and Henle, 1839) 

Frequent synonyms / misidentifications: Hypoprion macloti (Müller and Henle, 1839) / None. 

FAO names: En - Hardnose shark; Fr - Requin à nez rude; Sp - Tiburón trompudo. 

Diagnostic characters: A small shark. Body relatively 

slender. Snout very long and narrowly rounded or 

pointed, its length greater than mouth width and distance 

between nostrils; labial furrows very short; anterior nasal 

flaps with a slender elongate lobe; spiracles absent; teeth 

with smooth edges, those in upper jaw with narrow, 

oblique or nearly erect cusps and strong cusplets on 

each side of cusp; teeth in lower jaw with erect to oblique, 

smooth cusps and no cusplets. First dorsal fin moderately 

large, with a narrowly rounded or pointed apex, its inner 

margin greatly elongated, the free rear tip attenuated, 

and its origin over inner margins of pectoral fins; second dorsal fin very low, the inner margin over twice 

the fin height, fin origin slightly behind that of anal fin, pectoral fins relatively short, with narrowly rounded 

or angular tips; elongated rear tip of anal fin extending nearly to lower precaudal pit. No dermal ridge 

between dorsal fins. Colour: back greyish or grey-brown, belly white; posterior margin of pectoral fins 

and ventral caudal-fin lobe with an inconspicuous white edge; posterioventral and dorsal margins of caudal 

fin with a narrow black edge. 

Size: Maximum total length below 1 m; size at birth 45 to 50 cm. 

Habitat, biology, and fisheries: A common shark of continental waters inshore and offshore down to a 

depth of 170 m. Viviparous, number of embryos usually 2 (one per uterus). Feeds mainly on fishes, also 

on cephalopods and crustaceans. Not known to be dangerous to humans. Caught with floating gill nets, 

also bottom gill nets and longlines. One of the most abundant sharks taken in gill nets (also caught by line 

gear), but of limited interest to fisheries because of its small size. Utilized fresh and probably dried salted 


Distribution: Occurs off 

Kenya and Tanzania, possibly 

the Gulf of Aden, and from 

Pakistan eastward to New 

Guinea, northern Australia, 

China, Taiwan Province of 

China and southern Korea 

and Japan. 

? 

? 

? 

ventral view 

of head 


tooth near centre



Frequent synonyms / misidentifications: Hypoprion playfairi (Günther, 1870) / None. 

FAO names: En - Blacktip reef shark; Fr - Requin pointes noires; Sp - Tiburón de puntas negras. 

Diagnostic characters: A small to medium-sized 

shark. Body moderately stout. Snout very short and 

broadly rounded, its length less than mouth 

width and about equal to distance between 

nostrils; labial furrows very short; anterior nasal 

flaps with a stout, broad lobe; spiracles absent; teeth 

with serrated edges, those in upper jaw with 

narrow semioblique to oblique cusps and low 

basal cusplets; teeth in lower jaw with erect or 

semierect narrow cusps and serrated edges. First 

dorsal fin moderately large, with a narrowly rounded 

or pointed apex, its origin over inner margins of 

ventral view 

of head 



pectoral fins, its free rear tip short; second dorsal fin high, its inner margin much less than twice the fin 

height, its origin over or slightly anterior to anal-fin origin; pectoral fins moderately long, with narrowly 

rounded or pointed tips; rear tip of anal fin ending well in front of lower caudal-fin origin. No dermal ridge 

between dorsal fins. Colour: yellow-brown on dorsal surface, underside white; all fins conspicuous 

with black or dark brown tips also anterior and posterior dark edging on pectoral fins and upper lobe of 

caudal fin; a prominent black tip of first dorsal fin set off abruptly by a light band below it; a 

conspicuous dark band on flanks, extending rearward to pelvic fins. 

Size: Maximum total length about 2 m; commonly to 1.6 m; size at birth between 33 to 50 cm. 

Habitat, biology, and fisheries: A common inshore and sometimes offshore shark, on continental and insular 

shelves; prefers shallow water on and around coral reefs. May occur in brackish and even fresh water, but does 

not occur in tropical lakes and rivers far from the sea.Viviparous, number of young 2 to 4 (commonly 4).A bottom 

and midwater feeding shark that feeds on small bony fishes (including mullets), octopuses and small sharks. It 

has been definitely recorded as having attacked humans without provocation, but it should not be regarded as 

particularly dangerous because of its small size. May be aggressive when divers are spearfishing. Apparently 

regulary caught in fisheries where it occurs, including off of Thailand. The meat is used fresh and dried salted 

for human consumption, and for 

its liver oil. 

Distribution: Wide-ranging 

from South Africa, the Red Sea, 

Pakistan, India eastward to the 

western Central Pacific; also in 

the eastern Mediterranean 

Sea as an invader from the 

Red Sea through the Suez 

Canal.

1344 Sharks 


Frequent synonyms / misidentifications: Carcharhinus iranzae Fourmanoir, 1961 / None. 

FAO names: En - Dusky shark; Fr - Requin sombre (= Réquiem de sable, Area 31); Sp - Tiburón arenero. 

Diagnostic characters: Body slender to moderately stout. 

Snout rounded and short, its length equal to or less than 

mouth width and greater than or about equal to internasal 

space; labial furrows short; anterior nasal flaps 

rudimentary; upper teeth broadly triangular, erectto 

moderately oblique, anterior teeth with strongly serrated 

broad cusps not delimited from the bases; lower teeth with 

low, narrow, serrated cusps; gill slits relatively short. First 

dorsal fin relatively low, with a broadly arched anterior 

margin and a narrowly rounded or pointed apex, its origin 

about over free rear tips of pectoral fins; second 

dorsal fin also moderately low, with a nearly straight 

ventral view 

of head 



posterior margin, an inner margin nearly or quite twice the fin height, and its origin about over that of 

anal fin; pectoral fins falcate and apically pointed. A low interdorsal ridge present. Colour: blue-grey, 

lead-grey above, white below; tips of pectoral and pelvic fins, as well as lower lobe of caudal fin and dorsal 

fins often dusky in young, plain in adults. 

Size: Maximum total length about 3.64 m; matures at about 2.8 m; size at birth about 70 cm to 1 m. 

Habitat, biology, and fisheries: A semi-pelagic shark occurring from inshore waters to the outer 

continental shelf. Viviparous, number of embryos 6 to 14. Feeds chiefly on fishes, including scombrids, 

clupeids, serranids, trichiurids, bluefish, wrasses, anchovies, grunts, barracudas, sharks and rays, also 

squids, octopi, gastropods, shrimps, crabs, and carrion. Reported to be dangerous to humans, but attacks 

in the area are unverified. Regularly caught with longlines and probably gill nets, also hook-and-line and 

set bottom nets; utilized fresh, dried-salted, frozen and smoked for human consumption; hides used for 

leather; fins for shark-fin soup; liver oil extracted for vitamins. 

Distribution: Wide-ranging, 

but with a patchy distribution 


seas; in the western Central 

Pacific more confined to the 

marginal parts of the area 

(Japan, China, Viet Nam, 

Australia).



Frequent synonyms / misidentifications: Carcharhinus milberti (Valenciennes in Müller and Henle, 

1841) / None. 

FAO names: En - Sandbar shark; Fr - Requin gris (= Réquiem plombe, Area 31); Sp - Tiburón trozo 

(= Tiburón de Milberto). 

Diagnostic characters: A medium-sized, comparatively 

stout shark. Snout broadly rounded and short, 

its length less than width of mouth but greater than 

distance between nostrils; spiracles absent; teeth finely 

serrate, those in upper jaw broadly triangular and 

erect to slightly oblique, with broad, heavy cusps; lower 

teeth with narrow, erect cusps. First dorsal fin 

triangular, very high (height of fin twice the length of 

snout in adults), with a pointed or narrowly rounded 

apex, its origin over insertions of pectoral fins;origin 

of second dorsal fin about opposite that of anal fin, its 

inner margin less than twice the fin height; pectoral fins 

long and broad, their corners narrowly rounded or pointed. Interdorsal ridge present. Dermal denticles 

widely spaced, their free edges without definite teeth. Colour: back grey, or rarely brown; belly whitish. 

Size: Maximum total length about 2.4 m, records to 3 m uncertain; size at birth 60 to 75 cm. 

Habitat, biology, and fisheries: A coastal species usually found over sandy or muddy bottoms; often 

coming near estuaries but sometimes occurring in oceanic waters to depths of 280 m. Viviparous, number 

of young 1 to 14. Feeds mainly on bottom-dwelling animals, including flatfishes, rays, crabs, and snails; 

also on schooling fishes and squids. Not known to be dangerous to humans. Caught with longlines, 

hook-and-line, and set-bottom nets and is also fished with rod and reel by sports anglers as a game fish. 

It is utilized fresh, fresh-frozen, smoked, and dried-salted for human consumption; the hides are prized for 

leather and other products; the fins are prepared as the base for shark-fin soup; the liver is extracted for 

oil (rich in vitamins). 

Distribution: Wide-ranging 


areas of the eastern and 

western Atlantic, from the 

Mediterranean Sea, Indian 

Ocean and western Central 

Pacific to Hawaii; records 

from the eastern Central 

Pacific are uncertain. ? 

? 

? 

ventral view 

of head 


tooth near centre

1346 Sharks 



1839) / Carcharhinus dussumieri. 

FAO names: En - Blackspot shark; Fr - Requin à tache noire; Sp - Tiburón alinegro. 

Diagnostic characters: A small, stout to 

slender-bodied shark. Snout moderately long 

and narrowly parabolic or wedge-shaped, its 

length usually shorter than mouth width but 

subequal to the internarial space; labial furrows 

very short; anterior nasal flaps expanded; 

spiracles absent; gill slits short; teeth with 

serrated edges, upper teeth with 

narrow-based, strongly oblique serrated 

cusps and strong, smooth-edged cusplets; 

teeth in lower jaw erect to oblique, without 

cusplets, serrated and narrow-cusped; first 

ventral view 

of head 



dorsal fin moderately high, with an angular apex, notched posterior margin, and short inner margin, 

strongly falcate; origin of first dorsal fin over pectoral inner margins; origin of second dorsal fin about 

opposite or slightly behind that of anal fin; second dorsal fin high, its inner margin less than 1.5 times the 

fin height and its posterior margin concave; pectoral fins short and strongly falcate, with narrow, angular 

apexes. An interdorsal ridge present or occasionally absent on back. Colour: back greyish or 

grey-brown, belly whitish; a black spot on the second dorsal fin is the only conspicuous marking. 

Size: Maximum total length about 95 cm; maturing at 70 to 80 cm; size at birth 33 to 45 cm. 

Habitat, biology, and fisheries: A common coastal shark on the continental and insular shelves, from the 

surf line and intertidal to a depth of 40 m, usually in shallow water. Viviparous, number of young 1 or 2. 

Feeds on small fish (including sea horses), prawns, and squid. Not known to be dangerous to people. 

Commonly caught by artisanal and smallscale commercial fisheries as well as sport anglers fishing from 

the shore. Commonly fished with line gear and gill nets and utilized for human consumption. 


South Africa northward to 

Kenya, Madagascar, the 

Seychelles, Mauritius, and 

the southwest coast of India; 

in the eastern Indian Ocean 

and western Pacific eastward 

to China, the Philippines, and 

New Guinea.


Carcharhinus sorrah (Valenciennes in Müller and Henle, 1839 

Frequent synonyms / misidentifications: Carcharhinus bleekeri (Dumeril, 1865) / None. 

FAO names: En - Spottail shark; Fr - Requin tachete; Sp - Tiburón rabo manchado. 

Diagnostic characters: A small to medium-sized shark. 

Body slender to moderately stout. Snout moderately 

pointed, parabolic, and long, its length equal to or 

slightly less than mouth width and greater than 

internasal space; labial furrows short; anterior nasal 

flaps with a short, slender, narrow lobe; upper teeth 

with oblique or semioblique, narrow, serrated cusps 

and strong distal cusplets; lowers with semierect or 

oblique serrated cusps and no cusplets; gill slits 

relatively short. First dorsal fin with a narrowly rounded 

apex, its origin usually over the pectoral inner margins, 

its inner margin and free rear tip moderately long; 

second dorsal fin low, with a long, attenuated free rear tip and inner margin over twice fin height; 

origin of second dorsal fin over or slightly behind origin of anal fin; pectoral fins weakly falcate and with 

narrowly rounded tips; a dermal ridge present between dorsal-fin bases. Colour: grey or grey-brown 

above, white on belly, with a golden-brown sheen on the area between eyes and gill slits in fresh specimens; 

pectoral fins, second dorsal fin, and lower caudal-fin lobe with conspicuous black tips, first dorsal 

and upper caudal-fin lobe with black edging. A dark band on flank extending rearwards to pelvic fins. 

Sie: Maximum total length to about 1.5 to 1.6 m, possibly to 2.3 m but this is dubious; adults commonly 

1.06 to 1.5 cm; size at birth about 50 to 60 cm. 

Habitat, biology, and fisheries: A common inshore and sometimes offshore shark, on continental and 

insular shelves from close inshore and the surface down to a depth of at least 140 m. Often on and around 

coral reefs, but apparently occurring on other bottom habitats. Viviparous, number of young 2 to 6. Feeds 

on small bony fishes (including serranids and scombrids) and octopuses. Not known to have attacked 

people, and probably not particularly dangerous because of its small size. Caught in floating gill nets and 

on longlines. Utilized fresh for human consumption; fins of large individuals may be used in the oriental 

sharkfin trade, livers for vitamin oil, and offal for fishmeal. 


Madagascar, possibly South 

Africa, the Mauritius- 

Seychelles area, possibly the 

Gulf of Aden and Oman, 

western India, and probably 

Sri Lanka; occurs in the 

western Central Pacific from 

China southward to Indonesia 

and Australia, but without a 

wide distribution in Oceania. 

? 

? 

? 

ventral view 

of head 

? 


tooth near centre

1348 Sharks 

Carcharhinus tilstoni (Whitley, 1950 

Frequent synonyms / misidentifications: None / Carcharhinus limbatus (Valenciennes in Müller and 

Henle, 1839); C. brevipinna (Müller and Henel, 1839); C. amblyrhinchoides (Whitley, 1934). 

FAO names: En - Australian blacktip shark. 

Diagnostic characters: Body fusiform, moderately 

slender. Snout long, about equal to, or slightly less than 

mouth width, greater than distance between nostrils, its 

tip narrowly rounded to pointed; labial furrows short; 

spiracles absent; upper and lower teeth nearly 

symmetrical and similar, with erect, narrow cusps 

and serrated edges; gill slits moderately long. First 

dorsal fin with a pointed or very narrowly rounded 

apex, its origin above, or slightly posterior to 

insertion of pectoral fins; second dorsal fin high, its 

inner margin less than twice the height of fin, and its 

origin over or slightly in front of that of anal fin; pectoral 


ventral view 

of head 

5 th tooth from symphysis 

(upper jaw) 

fins falcate. No interdorsal ridge. Colour: back dark grey, ashy blue or dusky bronze; belly white or 

yellowish white; a dark band extending rearward along each side to about over origin of pelvic fin; 

tips of pelvic fins with a persistent black spot. 

Size: Maximum total length about 2 m; size at birth about 60 cm. 

Habitat, biology, and fisheries: On the continental shelf from close inshore to depths of about 150 m, 

mainly in midwater or near the surface. Feeds on teleost fishes, also on cephalopods. Until recently, the 

species was caught with gill nets and longlines for its meat by Taiwanese fisheries in northern Australia. It 

currently forms the basis of a small Australian gill net fishery (up to 500 t annually). 

Distribution: So far only known 

from northern Australia. 

Remarks: This species has 

only recently been separated 

from Carcharhinus limbatus. 

At present, these 2 species 

can be reliably distinguished 

only on vertebral counts and 

enzyme systems. 





Frequent synonyms / misidentifications: Galeocerdo arcticus (Faber, 1829); G. rayneri McDonald and 

Barron, 1868 / None. 

FAO names: En - Tiger shark; Fr - Requin tigre commun (= Requin tigre, Area 31); Sp - Tintorera. 

Diagnostic characters: A large, fusiform shark. 

Snout very short and bluntly rounded, its 

length much less than width of mouth; 

spiracles small, slit-like, but easily visible; upper 

labial furrows about as long as snout, 

reaching to front of eyes; teeth coarsely 

serrated, their outer edges deeply notched 

and the tips directed obliquely outward, their 

inner edges broadly convex. Second dorsal fin 

much smaller than first. A low rounded keel on 

each side of caudal peduncle. Colour: back 

dark grey or greyish brown with dark brown or 

ventral view 

of head 

upper 

labial 

furrow 



black rectangular spots often forming bars on sides and fins, but fading with growth. 

Size: Maximum total length at least 6.5 m; commonly to 4 m; size at birth between 60 cm and 1.04 m. 

Habitat, biology, and fisheries: Inhabits coastal, and offshore waters, near the surface and bottom; often 

found in shallow waters close inshore, including river estuaries. Ovoviviparous and very prolific with 10 to 

82 young in a litter. A voracious, indiscriminate predator feeding on all kinds of fish (including other sharks 

and rays), marine mammals, turtles, seabirds, sea snakes, squids, conchs, and crabs. Often swallows a 

variety of undigestible and non-nutritive items, and readily feeds on carrion. Considered among the most 

dangerous of sharks because of its shallow-water habitat, large jaws and teeth, indiscriminate appetite, 

and large size; several attacks on people have been recorded for this species. Caught in floating and bottom 

gill nets and with line gear (including pelagic longlines). Utilized for its high-quality hide, for its fins, liver oil 

and flesh, and offal for fishmeal. 


most tropical seas, with 

seasonal migrations into 

warm-temperate to temperate 

seas.

1350 Sharks 

Lamiopsis temmincki (Müller and Henle, 1839) 

Frequent synonyms / misidentifications: Carcharhinus temmincki (Müller and Henle, 1839); Eulamia 

temmincki (Müller and Henle, 1839) / Negaprion acutidens (Rüppell, 1837). 

FAO names: En - Broadfin shark; Fr - Requin grandes ailes; Sp - Tiburón aleton. 


shark. Body moderately stout. Snout moderately 

long, parabolic in shape, its length about equal 

to mouth width and greater than distance between 

nostrils; labial furrows short; anterior nasal flaps 

with a short, broad lobe; spiracles absent; teeth 

in upper jaw with high, broadly triangular, erect 

to semioblique, serrated cusps, and no cusplets; 

teeth in lower jaw with erect, high, hooked, 

smooth-edged narrow cusps, and no cusplets. 

First dorsal fin moderately large, with a narrowly 

rounded apex, its origin over inner margins of 

ventral view 

of head 



pectoral fins, its free rear tip moderately long; second dorsal fin very large, nearly or quite as large as 

first dorsal, its inner margin shorter than fin height, its origin anterior to anal-fin origin; pectoral fins 

moderately long, basally very broad and not falcate with narrowly rounded tips; anal fin with posterior 

margin slightly concave; upper precaudal pit a shallow longitudinal depression, not transverse and 

crescentic. No dermal ridge between dorsal fins, and no keels on caudal peduncle. Colour: grey or 

yellow-grey above, lighter below; no conspicuous markings. 

Size: Maximum total length about 1.7 m; size at birth between 40 and 60 cm. 

Habitat, biology, and fisheries: A little-known coastal, inshore, tropical shark. Viviparous, number of young 

4 to 8 per litter. Probably feeds on small fishes and invertebrates. Not known to be dangerous to people. 

Caught in bottom and floating gill nets and with line gear. Meat utilized fresh for human consumption; livers 

used for vitamin oil. 


scattered localities in the 

Indian Ocean and western 

Pacific off Pakistan, India, 

Burma, Indonesia (Makassar 

Straits), Sarawak, and China.



Frequent synonyms / misidentifications: Scoliodon acutus (Rüppell, 1837); S. ceylonensis Setna and 

Sarangdhar, 1946 / Scoliodon laticaudus Müller and Henle, 1838; Carcharhinus macloti (Müller and Henle, 

1838). 

FAO names: En - Sliteye shark; Fr - Requin sagrin; Sp - Tiburón ojuelo. 

Diagnostic characters: A small, very slender 

shark. Snout very long, parabolic in shape, 

its length greater than mouth width and 

distance between nostrils; labial furrows very 

short; anterior nasal flaps with a short, broadly 

triangular lobe; eyes large, with a posterior 

notch; spiracles absent; teeth in both jaws 

with low, narrow, oblique, smooth-edged 

cusps, and no cusplets. First dorsal fin 

small, its origin behind free rear tips of pectoral 

fins by a distance greater than length of fourth 

gill opening, its base 2 or 3 times in distance 

ventral view 

of head 



between pectoral and pelvic-fin bases, its free rear tip moderately long and not reaching backward to 

pelvic-fin origins; second dorsal fin very small, its height less than 1/3 of that of first dorsal fin, the inner 

margin elongated and over twice the fin height and the fin origin usually just behind anal-fin insertion 

(occasionally over or slightly in front of it, but far behind anal midbase); pectoral fins small, narrow and 

slightly falcate; anal fin with a slightly concave posterior margin and long preanal ridges. Upper 

precaudal pit transverse and crescentic; no keels on caudal peduncle; interdorsal ridge usually absent. 

Colour: grey above, pale below, fins with pale edges (transparent in life), caudal and first dorsal fins with 

narrow dark margin, first dorsal fin also with a dusky tip. 

Size: Maximum total length about 91 cm; maturing at 73 to 85 cm; size at birth about 40 to 43 cm. 

Habitat, biology, and fisheries: Occurs in tropical, coastal, clear waters, near the surface and bottom, 

inshore and offshore at depths from 7 to 80 m. Viviparous number of young usually 2 in a litter. Feeds on 

small bony fishes, including anchovies and croakers, and shrimp and cuttlefish. Harmless to people. Caught 

in artisanal and small scale commercial fisheries with floating and bottom gill nets and with line gear 

(including pelagic longlines). 

Utilized fresh for human 

consumption. 

Distribution: In the Indo-West 

Pacific from South Africa, 

southern Mozambique, 

Madagascar, Seychelles and 

the Red Sea eastward to India, 

Sri Lanka, Indonesia, China, 

Taiwan Province of China, the 

Philippines, and Australia.

1352 Sharks 


Frequent synonyms / misidentifications: None / Lamiopsis temmincki (Müller and Henle, 1839). 

FAO names: En - Sicklefin lemon shark; Fr - Requin citron faucille; Sp - Tiburón segador. 

Diagnostic characters: A large, stout shark. 

Snout short (shorter than width of mouth) 

and broad, rounded or obtusely 

wedge-shaped; labial folds short; spiracles 

usually absent; teeth narrow, their cusps 

smooth-edged, erect in anterior part of jaws, 

but becoming progressively oblique toward the 

sides; bases of upper teeth smooth or weakly 

serrated. Origin of first dorsal fin over or behind 

free rear tips of pectoral fins, closer to these fins 

than to the pelvic fins; second dorsal fin nearly 

ventral view 

of head 



as large as the first (its base more than 3/4 of first dorsal-fin base); pectoral fins broad and strongly falcate, 

pelvic fins falcate. No dermal ridge between dorsal fins. Colour: yellowish brown above, paler below. 

Size: Maximum total length about 3 m; maturing at about 2.2 m; size at birth about 50 to 70 cm. 

Habitat, biology, and fisheries: Occurs in tropical, shallow inshore and offshore waters near the bottom; 

often found on and around coral reefs and on sandy plateaus near coral, at depths down to at least 30 m. 

Viviparous, 12 or 13 young in a litter. A fish-eating shark, but few details of its diet are available from the 

area. Potentially dangerous because of its large size, powerful jaws and dagger-like teeth; normally 

inoffensive and sluggish but very aggressive when provoked. Caught in floating and bottom gill nets and 

on line gear (including floating longlines). Used fresh and dried-salted for human consumption; livers 

processed for vitamin oil; offal processed for fishmeal; and fins are processed for shark-fin soup base. 


the Indian Ocean and western 

Central Pacific, extending 

from South Africa to the 

Australian region and 

Oceania.




FAO names: En - Blue shark; Fr - Peau bleue; Sp - Tiburón azul. 

Diagnostic characters: very slender, fusiform 

shark. Snout long, (its length greater than mouth 

width) and narrowly rounded; upper labial furrows 

very short; spiracles absent; nictitating eyelids 

present; teeth serrated, broadly triangular and curved 

in upper jaw, narrower in lower jaw; upper medial 

tooth very large, nearly the size of teeth on either 

side of it (but sometimes absent); inner gill arches 

with gill-raker papillae (visible through open mouth). 

First dorsal-fin origin well posterior to free rear tips of ventral view 


pectoral fins, the midpoint of its base closer to of head 


pelvic fin than to pectoral-fin origins; second dorsal fin much smaller than first; pectoral fins very long, 

narrow and somewhat falcate. A weak keel present of sides of caudal peduncle. No interdorsal ridge. 

Colour: in life, dark blue above, bright blue on sides, white below, fading to purple blackish after death; 

tips of pectoral fins and anal fin dusky. 

Size: Maximum total length about 3.8 m, though larger specimens (up to 4.8 to 6.5 m) are mentioned on 

poor evidence in the literature; most specimens below 3.35 m. 

Habitat, biology, and fisheries: A slow-cruising, very common oceanic species capable of bursts of speed 

when excited. Usually well offshore and in the open sea near the surface, but sometimes penetrating coastal 

waters. Viviparous, litters usually large, ranging from 4 to 63 young. Feeds on a wide variety of bony fishes, 

small sharks, squids, pelagic crustaceans, and occasionally sea birds and carrion. Sometimes aggressive 

to people in the water, and considered a dangerous species although attacks on people are relatively 

uncommon. Usually caught with pelagic longlines and gill nets but also hook-and-lines, pelagic trawls, and 

even bottom trawls near coasts. It is utilized fresh, smoked, and dried-salted for human consumption; its 

hides are used for leather; fins for shark-fin soup base; and also for fishmeal and liver oil. This shark is also 

considered a game fish and 

taken by sports anglers with 

rod and reel. 


all tropical and temperate 

seas.

1354 Sharks 


Frequent synonyms / misidentifications: Scoliodon acutus (Rüppell, 1837); S. palasorra (Bleeker, 

1853); S. walbeehmi (Bleeker, 1856) / Rhizoprionodon oligolinx Springer, 1964; R. taylori (Ogilby, 1915); 

Loxodon macrorhinus Müller and Henle, 1839; Scoliodon laticaudus Müller and Henle, 1838; 

Carcharhinus macloti Müller and Henle, 1839. 

FAO names: En - Milk shark; Fr - Requin à museau pointu; Sp - Cazón picudo. 

Diagnostic characters: A small, slender shark. 

Snout long and depressed its length usually greater 

than width of mouth, its tip narrowly rounded; eyes 

without a posterior notch; no spiracles; labial furrows 

well developed and moderately long, the upper 

ones about equal in length to eye diameter and 

ending well behind eyes; teeth similar in both jaws, 

low-crowned, oblique and narrow-cusped, with the 

outer edges deeply notched and without cusplets, 

smooth-edged in young but often finely serrated in 

adults. Origin of first dorsal fin over or posterior to 

ventral view 

of head 


lateral teeth 

inner corners of pectoral fins, base length of first dorsal fin twice or less in distance between pectoral and 

pelvic-fin bases, free rear tip usually anterior to pelvic-fin origins; second dorsal fin smaller than anal 

fin, its origin far posterior to midlength of anal-fin base; anal fin with slightly concave posterior 

margin and a pair of long preanal ridges. Colour: grey or grey-brown above, white below, dorsal and 

anal fins with dusky or blackish edges, fins slightly darker than back. 

Size: Maximum total length about 1 m; a single record of a 1.78 m specimen off Africa (possibly based on 

some other species); adults maturing at about 75 cm; size at birth about 35 cm. 

Habitat, biology, and fisheries: An extremely abundant, small, inshore and offshore shark of the tropics, 

ranging from the surfline down to a depth of 200 m, and occurring near the surface, as well as near the 

bottom. Viviparous, with 2 to 8 fetuses in a litter, gestation period about one year. Feeds on small bony 

fishes and small crustaceans; harmless to people. Very commonly caught inshore in artisanal and 

smallscale fisheries as well as offshore in fishing fleets. Caught on line gear (including floating longlines 

set near the coasts), and 

especially floating and bottom 

gill nets. Utilized fresh and 

possibly dried salted for food 

and for fishmeal. 

Distribution: In all tropical 

and subtropical areas of the 

Indian Ocean and the western 

Central Pacific (but absent in 

Oceania); also in the eastern 

Atlantic off Madeira and from 

Mauretania to Angola.



Frequent synonyms / misidentifications: Scoliodon palasorra (Bleeker, 1853) / Rhizoprionodon acutus 

(Rüppell, 1837); R. taylori (Ogilby, 1915); Loxodon macrorhinus Müller and Henle, 1839; Scoliodon 

laticaudus Müller and Henle, 1839; Carcharhinus macloti (Müller and Henle, 1839). 

FAO names: En - Grey sharpnose shark; Fr - Requin aiguille gris; Sp - Cazón picudo gris. 


Snout long and depressed, its length usually greater 

than width of mouth, its tip narrowly rounded; eyes 

without a posterior notch; no spiracles; labial 

furrows very short, much less than eye length, 

ending well behind eyes; teeth similar in both jaws, 

low-crowned, oblique and narrow-cusped, with the 

outer edges deeply notched and without cusplets, 

smooth edged in young but often finely serrated in 

adults. Origin of first dorsal fin over or posterior to 

inner corners of pectoral fins, its base length less 

than 2 times in distance between pectoral and 

pelvic-fin bases, its free rear tip usually anterior to pelvic-fin origins but occasionally over them; second 

dorsal fin smaller than anal fin, its origin far posterior to midlength of anal-fin base; anal fin with 

slightly concave posterior margin and a pair of long preanal ridges. Colour: grey dorsal and anal fins 

with dusky slightly darker than back. 

Size: Maximum total length about 70 cm; males may mature at 35 cm; size at birth about 20 to 30 cm. 

Habitat, biology, and fisheries: A common but little-known littoral, inshore and offshore tropical shark of 

coastal waters, ranging down to at least depths of 36 m from close inshore. Viviparous, with a yolk-sac 

placenta; number of young 3 to 5 per litter. Probably feeds on small fishes and invertebrates; harmless to 

people. Caught with floating and bottom gill nets, and line gear. Utilized fresh and probably dried salted for 

human consumption, also for fishmeal. 


Indo-West Pacific from the 

Persian Gulf eastward to 

Thailand, Indonesia, China, 

and Japan; also recorded 

from Australia (Gulf of 

Carpentaria). 

? 

ventral view 

of head 


lateral teeth

1356 Sharks 


Frequent synonyms / misidentifications: Protozygaena taylori (Ogilby, 1915) / Rhizoprionodon acutus 

(Rüppell, 1837); R. oligolinx Springer, 1964; Loxodon macrorhinus Müller and Henle, 1839; Scoliodon 

laticaudus Müller and Henle, 1838; Carcharhinus macloti (Müller and Henle, 1839). 

FAO Names: En - Australian sharpnose shark; Fr - Requin aiguille réchine; Sp - Cazón picudo australiano. 


Snout long and depressed, its length usually 

greater than width of mouth, its tip narrowly 

rounded; eyes without a posterior notch; no 

spiracles; labial furrows very short, much 

less than eye length, ending well behind 

eyes; teeth similar in both jaws, low-crowned, 

oblique and narrow-cusped, with the outer 

edges deeply notched and without cusplets, 

smooth-edged in young but often finely serrated 

in adults. Origin of first dorsal fin just anterior or 

posterior to inner corners of pectoral fins, base 

ventral view 

of head 


length of first dorsal fin twice or less in distance between pectoral and pelvic-fin bases, free rear tip usually 

anterior to pelvic-fin origins; second dorsal fin smaller than anal fin, its origin over past 1/3 of anal-fin 

base; anal fin with slightly concave posterior margin and a pair of long preanal ridges. Colour: 

brownish grey above, white below, fins light-edged but not conspicuously marked. 

Size: Maximum total length about 67 cm, males adolescent at about 41 cm; size at birth about 45 cm. 

Habitat, biology, and fisheries: A little-known tropical inshore shark of the Australian continental shelf. 

Viviparous, with a yolk-sac placenta; number of young 2 per litter. Feeds mainly on fish but also on 

cephalopods and crustaceans. Locally very common and taken incidentally in mackerel nets, but not used 

commercially because of its small size. 

Distribution: Western South 

Pacific off Papua New Guinea 

and from northwestern 

Australia to southern 

Queensland. 

?



Frequent synonyms / misidentifications: Physodon mülleri (Valenciennes in Müller and Henle, 1839); 

Scoliodon palasorra (Bleeker, 1853); S. sorrakowa (Bleeker, 1853) / None. 

FAO names: En - Spadenose shark; Fr - Requin epée; Sp - Cazón espadachin. 


moderately stout and markedly compressed. 

Head and snout strongly depressed; snout 

long, narrowly rounded, its length greater than 

mouth width; labial furrows very short; anterior 

nasal flaps with a short, narrowly triangular lobe; 

eyes moderately large, without a posterior 

notch; spiracles absent; teeth similar in both 

jaws, oblique and narrow-cusped, with the distal 

edges deeply notched and without cusplets or 

serrations. First dorsal fin moderately large, its 

ventral view 

of head 



origin well behind pectoral free rear tips, its base closer to pelvic than to pectoral-fin bases, and its free 

rear tip over or behind middle of pelvic-fin bases; second dorsal fin very small, its height less than 

1/3 of that of first dorsal, its inner margin elongated and over twice the fin height, fin origin over or 

slightly anterior to anal-fin insertion; pectoral fins small broad, triangular, and not falcate, originating 

under or slightly anterior to fifth gill openings, anal fin with a slightly concave posterior margin and 

relatively short preanal ridges. Upper precaudal pit transverse and crescentic; no keels on caudal peduncle; 

no interdorsal ridge. Colour: bronzy grey above, white below, fins sometimes darker than body; no 

conspicuous markings. 

Size: Maximum total length about 74 cm, but most individuals smaller; size at birth about 13 to 15 cm. 

Habitat, biology, and fisheries: A common tropical shark in coastal waters, often near the bottom in rocky 

areas. Viviparous, number of young 5 to 14. A small harmless shark, very abundant where it occurs in the area, 

and forming large schools. Feeds on shrimps, cuttlefishes, and small schooling fishes including anchovies, 

bregmacerotids, tripauchenids, and Bombay ducks (Harpodon nehereus). Caught with hook-and-line, longlines, 

floating and bottom gill nets, set bottom sets, and traps.Utilized for human consumption, processed into fishmeal, 

and used for bait for other sharks and bony fishes. 


Indo-West Pacific from the 

Persian Gulf eastward to 

Thailand, Indonesia, China, 

and Japan; also recorded 

from Tanzania, but absent 

from Oceania and the 

Australasian region.

1358 Sharks 



FAO names: En - Whitetip reef shark; Fr - Requin corail; Sp - Cazón coralero trompacorta. 


shark. Body moderately stout. Snout very short, 

broadly rounded, its length much less than mouth 

width and equal to or less than distance between 

nostrils; labial furrows very short; anterior nasal flaps 

with a short, truncate, prominent lobe, formed into 

a partial tube; spiracles usually absent (small ones 

present in a few specimens); teeth in upper and lower 

jaws with high, narrow, smooth-edged cusps with 

strong cusplets on each side, no serrations. First 

dorsal fin moderately large, with a narrowly rounded 

apex, its origin well posterior to free rear tips of 

ventral view 

of head 

nostril 



pectoral fins, the midpoint of its base closer to pelvic fins than pectoral fins, and its free rear tip about 

over pelvic-fin origins; second dorsal fin very large, about 1/2 the surface of first dorsal fin and over 

half its height, its inner margin shorter than fin height, its origin over or slightly anterior to anal-fin origin; 

pectoral fins moderately long, moderately narrow, slightly falcate, and with narrow tips; anal fin with 

posterior margin deeply notched; upper precaudal pit transverse and crescentic. No interdorsal ridge, and 

no keels on caudal peduncle. Colour: grey-brown above, sometimes with a few or several dark spots on 

sides, first dorsal-fin lobe and dorsal caudal-fin lobe with conspicuous white tips, second dorsal-fin 

lobe and ventral caudal-fin lobe often white-tipped; ventral surface cream-white. 

Size: Maximum total length 1.7 m; size at birth about 52 to 60 cm. 

Habitat, biology, and fisheries: A common shark in tropical, coastal clear waters, usually on or around 

coral reefs; commonly in holes and crevices, often in shallow water near the bottom, but exceptionally at 

considerable depths down to 330 m. Viviparous, number of young 1 to 5 in a litter. A common reef shark, 

feeding on a wide variety of reef fishes including moray eels, squirrelfishes, snappers, damselfishes, 

parrotfishes, surgeonfish, triggerfishes, goatfishes; also octopuses, lobsters, and crabs. A relatively 

non-aggressive shark to people in the water, and generally considered as not particularly dangerous. In 

response to exciting stimuli, especially speared fish, this shark has been known to attack divers, but never 

with serious results. Caught in 

floating and bottom gill nets 

and with line gear, including 

floating longlines. Utilized 

fresh for human consumption. 


the Indo-Pacific with an 

extensive distribution among 

islands of the tropical Pacific.


Carcharhinus sp. 

En - False smalltail shark. 

Maximum total length at least 43 cm, probably attains a maximum length of about 1 m as a term 

fetus is 34 cm total length and a freeliving individual with an umbilical scar is 37 cm total length. A 

rare, little-known inshore tropical shark endemic to the area, previously confused with the American 

smalltail shark, Carcharhinus porosus (Ranzani, 1839) but closer to (but distinct from) C. borneensis. 

Presumably viviparous. Probably occurs in local fisheries, but of minor interest. Known only from 3 

specimens from Viet Nam (Ho Chi Minh City), Borneo (Baram, Sarawak), and Thailand (Bangkok). 

Conservation status needs investigation. 

ventral view 

of head 

Glyphis sp. A [Last and Stevens, 1994] 

En - Queensland river shark. 

Maximum total length at least 75 cm (newborn specimen with umbilical scar), probably attains a 

much larger size. A rare, little-known tropical riverine shark. Probably occurs in local fisheries, but 

of minor interest. Known from 2 specimens from the lower reaches of the Bizant River in Queensland, 

Australia, probably in brackish rather than fresh water. Possibly identical with the speartooth shark, 

Glyphis glyphis (Müller and Henle, 1839) which was described from a single stuffed specimen 

without locality. Conservation status needs investigation. 


1360 Sharks 

Glyphis sp. B 

En - Borneo river shark. 

Maximum total length about 81 cm, probably attains a much larger size. A rare, little-known tropical 

shark. Occurs in local fisheries, but of little interest. Known from Borneo. Conservation status needs 

investigation. 

ventral view 

of head 

Glyphis sp. C 

En - New Guinea river shark. 

Maximum total length at least 1.31 m probably larger. A rare, little-known riverine tropical shark. 

Probably occurs in local fisheries, but of minor interest. A species including 3 specimens from Papua 

New Guinea and from fresh water in the Adelaide River, Northern Territory, Australia. Jaws of Glyphis 

from Papua New Guinea in estuaries or fresh water near Port Romilly and Bainuru and from fresh 

water at Alligator Island in the Fly River may be this species or Glyphis sp. A. Conservation status 

needs investigation. 

ventral view 

of head 



Sphyrnidae 1361 

SPHYRNIDAE 

Hammerhead and bonnethead sharks 


Diagnostic characters: Medium- to large-sized sharks. Body elongate and moderately slender. Anterior 

portion of head much flattened dorsoventrally and widely expanded laterally in “hammer” 

form, with the eyes at its outer edges; well-developed nictitating lower eyelids; teeth blade-like, with a 

single cusp. Two dorsal fins, the first high and pointed, its base much shorter than caudal fin and wholly 

anterior to pelvic-fin origin; caudal fin strongly asymmetrical, with a well-marked subterminal notch and a 

small, but well-defined lower lobe. Caudal peduncle not strongly flattened dorsoventrally or widely 

expanded laterally, without longitudinal ridges but with precaudal pits. Colour: back predominantly grey or 

brassy; belly white. 

Habitat, biology, and fisheries: Hammerhead sharks inhabit surface waters in 

tropical and warm-temperate areas. Small species are confined to coastal 

waters; juveniles of large species are coastal, while adults are primarily semioceanic, 

although they often approach the coast in search of food. They are 

voracious predators, feeding mainly on fishes, sharks, rays, and bottom-dwelling 

animals (some crustaceans and molluscs). A few species are reported dangerous 

to bathers. Hammerhead sharks are important for fisheries in the area and 

are used as food and also for the preparation of various subproducts, especially 

Vitamin A from the liver and soup base from the fins. 

nare 

eye 


None. No other shark family has the characteristic hammer-shaped head of the 

Sphyrnidae. 

Key to the species of Sphyrnidae occurring in the area 

1a. Expanded lateral blades of head 

very narrow and wing-like, with a 

series of small bumps along 

edges in front of nostrils; width 

across head 40 or 50% of total 

length; nostrils enormously expanded, 

each nearly 2 times the 

mouth width (Fig.1) . . . . . Eusphyra blochii 

1b. Expanded lateral blades of head 

relatively broad, not wing-like, and 

without small bumps along edges 

in front of nostrils; width across 

head less than 31% of total length; 

nostrils narrow, less than 1/2 the 

mouth width (Figs 2 to 4) . . . (Sphyrna) → 2 

posterior lobe 

of 1 st dorsal fin 

pectoral fin anal fin 

pelvic fin 

ventral view 

of head 

posterior lobe of 


Fig. 1 Eusphyra blochii 

corner of 

mouth 

gill 

slits 


1362 Sharks 

2a. Anterior margin of head nearly 

straight in adults, moderately 

convex in young; prenarial 

grooves hardly developed; teeth 

strongly serrated at all sizes; first 

dorsal fin markedly falcate; second 

dorsal fin about 1/3 as high 

as first, with a short inner margin; 

posterior margins of second dorsal 

and pelvic fins deeply concave 

(Fig. 2) . . . . . . . . Sphyrna mokarran 

2b. Anterior margin of head moderately 

convex in adults, strongly so 

in young; prenarial grooves well 

developed; teeth generally 

smooth, but may be finely ser- 

rated in adults; first dorsal fin erect or slightly falcate; second dorsal fin less than 1/3 the 

height of first, with a long inner margin; posterior margins of second dorsal and pelvic 

fins slightly concave to nearly straight (Figs 3 and 4) . . . . . . . . . . . . . . . . . . . . . . . → 3 

3a. Median indentation present on anterior margin of head; free rear tip of second dorsal 

fin nearly reaching upper caudal-fin origin; anal-fin base noticeably larger than that of 

second dorsal fin (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphyrna lewini 

3b. Median indentation absent from anterior margin of head; free rear tip of second dorsal 

fin well ahead of upper caudal-fin origin; anal-fin base about as large as that of second 

dorsal fin (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sphyrna zygaena 

median indentation 

prenarial groove 




Sphyrna lewini (Griffith and Smith, 1834) 



 

ventral view 

of head 

Fig. 3 Sphyrna lewini 

median indentation 

ventral view 

of head 

ventral view 

of head 

prenarial groove weak 

Fig. 2 Sphyrna mokarran 

Fig. 4 Sphyrna zygaena 

References 


572 p. 

Gilbert, C.R. 1967. A revision of the hammerhead sharks (family Sphyrnidae). Proc. U.S. Natl. Mus., 119:88 p. 

Gilbert, C.R. 1967. A taxonomic synopsis of the hammerhead sharks (family Sphyrnidae). In Sharks, skates and rays, 

edited by P.W. Gilbert, R.F. Mathewson, and D.P. Rall. Baltimore, Johns Hopkins Press, pp. 69-76. 

nostril 

nostril



Frequent synonyms / misidentifications: Sphyrna blochii (Cuvier, 1817) / None. 

FAO names: En - Winghead shark; Fr - Requin-marteau planeur; Sp - Cornuda planeadora. 

Diagnostic characters: Body elongate and 

compressed. Head shaped like a broad arrowhead 

or a pair of aircraft wings in dorsal 

or ventral view, tremendously expanded laterally 

and relatively narrow from front to back, 

with width across head 2/5 to 1/2 of total length; 

anterior contour of head almost V-shaped in 

young but with lateral wings of head becoming 

transverse in adults; a shallow, but distinct ventral view of head 

indentation, at the midline of head and a 

very broad, shallow indentation opposite 


each nostril, the edge of which has a row of low bumps; nostrils greatly elongated, wider than mouth, 

with strong prenarial grooves anteromedial to their incurrent apertures; posterior margins of eyes opposite 

or behind front of mouth; teeth triangular, deeply notched distally, with relatively narrow oblique cusps and 

unserrated edges. First dorsal fin very high, strongly falcate; second dorsal fin small and low, less than 

1/3 of height of first, with a greatly elongated inner margin nearly or quite twice the fin height, a free rear 

tip that nearly or quite reaches upper caudal-fin origin, and a shallowly concave posterior margin; pelvic 

fins with posterior margin nearly straight; anal-fin base about 1/3 longer than second dorsal fin. Colour: 

grey or grey-brown above, paler below. 

Size: Maximum total length about 18.6 m; size at birth between 32 and 45 cm. 

Habitat, biology, and fisheries: Found in shallow water on the continental and insular shelves. Viviparous, 

litters from 6 to 25 young. Feeds mainly on small bony fishes, but also on cephalopods and crustaceans. 

A small species, probably harmless to people. A common fisheries species in India, Pakistan, Malaysia, 

and Thailand, and probably elsewhere in its range. Caught with floating gill nets, probably fixed bottom gill 

nets, with floating longlines, and probably on hook-and-line. Utilized fresh for human consumption; livers 

yield a high-potency vitamin oil; 

and offal is probably used for 

fishmeal. 

Distribution: An Indo-West 

Pacific coastal species distributed 

from the Persian Gulf 

eastward to Pakistan, India, Sri 

Lanka, Thailand, Borneo, 

China, the Philippines, and 

northern Australia.

1364 Sharks 

Sphyrna lewini (Griffith and Smith, 1834) 

Frequent synonyms / misidentifications: Sphyrna diplana Springer, 1941 / Sphyrna mokarran (Rüppell, 

1837); S. zygaena (Linnaeus, 1758). 

FAO names: En - Scalloped hammerhead; Fr - Requin-marteau halicorne; Sp - Cornuda común. 

Diagnostic characters: Body elongate and laterally compressed. 

Head “hammer”-shaped, its anterior contour broadly arched in 

young, but moderately so in adults, with a shallow but distinct 

indentation at the midline and a deep rounded depression opposite 

each nostril; lateral expansions of head very prominent, broad transversely 

and narrow from front to back; nostrils with strong prenarial 

grooves; hind margins of eyes slightly posterior to or nearly 

opposite front of mouth; mouth broadly arched; teeth triangular, 

deeply notched posteriorly, with smooth or finely serrated edges. 


First dorsal fin high, moderately falcate; second dorsal fin small, less 

than 1/4 of height of first, with a greatly elongated free rear tip extending backward nearly to upper 

caudal-fin origin, an inner margin about twice as long as the anterior fin margin and a shallowly 

concave posterior margin; pectoral fins short and broad; pelvic fins with a nearly straight posterior 

margin; second dorsal-fin base about 3/5 to 4/5 the length of anal-fin base. Colour: uniform grey, greyish 

brown, or olivaceous above, shading to white below; pectoral fins tipped grey or black ventrally. 

Size: Maximum total length about 4.2 m; commonly to 3.6 m; size at birth between 45 and 50 cm. 

Habitat, biology, and fisheries: Estuarine and inshore to well offshore and semi-oceanic, with young 

mostly in coastal waters. Adults solitary, in pairs, or in schools while the young form huge schools. 

Viviparous, number of young up to 30. Feeds on pelagic fishes, other sharks and rays, squids, lobsters, 

shrimps, and crabs. Adults considered potentially dangerous but often unaggressive when approached by 

divers. Probably the most abundant tropical hammerhead, readily available to inshore artisanal and small 

commercial fisheries as well as to offshore operations. Caught with pelagic longlines, fixed bottom 

longlines, fixed bottom nets, and even bottom and pelagic trawls; the young are easily caught on light 

longline gear. The meat is utilized 

fresh, fresh-frozen, driedsalted, 

and smoked for human 

consumption; the fins are used 

to prepare shark-fin soup 

base; the hides for leather, the 

oil for vitamins, and carcasses 

for fishmeal. 

Distribution: Essentially circumglobal 

in coastal warm 

temperate and tropical seas.



Frequent synonyms / misidentifications: Sphyrna tudes (Valenciennes, 1822) / Sphyrna lewini (Griffith 

and Smith, 1834); S. zygaena (Linnaeus, 1758). 

FAO names: En - Great hammerhead; Fr - Grand requin-marteau; Sp - Cornuda gigante. 

Diagnostic characters: Body elongate and laterally compressed. 

Head “hammer”-shaped, its anterior contour moderately arched in 

young but nearly straight in adults, with a shallow but distinct 

indentation at the midline and a shallow rounded depression opposite 

each nostril; lateral expansions of head very prominent, broad transversely 

and narrow from front to back; nostrils with weak prenarial 

grooves; posterior margins of eyes well anterior to mouth; mouth 

broadly arched; teeth triangular, deeply notched posteriorly, with 

strongly serrated edges. First dorsal fin very high, strongly falcate; 


second dorsal fin very large, with a moderately short inner margin 

(about equal to anterior fin margin), a free rear tip ending well anterior to upper caudal-fin origin, and a 

deeply concave posterior margin; anal-fin base about as long as second dorsal-fin base; pectoral fins 

short and broad; pelvic fins with a deeply concave posterior margin; Colour: grey or grey-brown above, 

paler below; fins with dusky tips in young. 

Size: Maximum total length between 5.5 and 6 m, and possibly more; commonly between 2.4 and 3.7 m; 

size at birth between 60 and 70 cm. 

Habitat, biology, and fisheries: A powerful coastal and semi-oceanic species coming close inshore, often 

around and on coral reefs; also occurring near the surface over deep water not far from land. Viviparous, 

litters from 18 to 38 fetuses. Feeds on bony fishes (including sparids), other sharks, rays, squids, and 

lobsters. Potentially dangerous to people in the water. Although less abundant than Sphyrna lewini, this 

species is regularly caught in the tropics, with longlines, fixed bottom nets, hook-and-line, and possibly 

with pelagic and bottom trawls. Utilized for its meat, fresh, fresh-frozen, dried-salted, and smoked for human 

consumption; for hides, processed into leather; for fins used for shark-fin soup base; for liver oil, processed 

for vitamins; and carcasses for fishmeal. 

Distribution: Essentially circumglobal 

in coastal warm 

temperate and tropical seas.

1366 Sharks 


Frequent synonyms / misidentifications: None / Sphyrna lewini (Griffith and Smith, 1834); S. mokarran 

(Rüppell, 1837). 

FAO names: En - Smooth hammerhead; Fr - Requin-marteau commun; Sp - Cornuda cruz. 

Diagnostic characters: Body elongated and 

laterally compressed. Head “hammer”-shaped, 

its anterior contour strongly arched in 

young but moderately rounded in adults, 

without a median indentation but with a deep 

rounded depression opposite each nostril; lateral 

expansions of head very prominent, broad 

transversely and narrow from front to back; nostrils 

with strong prenarial grooves; eyes 

large, their horizontal diameter greater than ventral view of head 


length of shortest (fifth) gill slit, their posterior 

margins about opposite mouth or just anterior to it; mouth broadly arched; teeth triangular, deeply 

notched posteriorly, with smooth or finely serrated edges. First dorsal fin high, moderately falcate; 

second dorsal fin small, with a very long inner margin (almost twice the anterior fin margin), afree 

rear tip ending well anterior to upper caudal-fin origin, and a nearly straight to shallowly concave 

posterior margin; anal-fin base slightly longer than second dorsal-fin base; pectoral fins short and broad; 

pelvic fins with posterior margins straight to shallowly concave. Colour: brownish olive, or plain grey above, 

white or grey-white below; fins nearly plain, dusky or blackish tipped. 

Size: Maximum total length probably between 3.7 and 4 m, commonly between 2.75 and 3.35 m; size at 

birth between 50 and 60 cm. 

Habitat, biology, and fisheries: A common to abundant coastal and semi-oceanic species, living close 

inshore (especially the young) and near the surface in deep water not far offshore. A strong-swimming 

shark, migrating northward in summer; young often found in large aggregations of hundreds of individuals. 

Viviparous, litters from 29 to 37 fetuses. Feeds on bony fishes, other sharks, rays, crustaceans, and squids. 

Potentially dangerous to people. Caught with pelagic longlines, handlines, and even pelagic and bottom 

trawls. Utilized fresh, dried-salted, and possibly smoked for human consumption; hides are processed for 

leather; liver oil is extracted for 

vitamins; fins are processed 

into shark-fin soup base; and 

carcasses utilized for fishmeal. 


temperate and tropical seas 

(western and eastern Atlantic, 

Mediterranean, western and 

eastern Pacific); occurs in the 

western Pacific from southern 

Siberia to Viet Nam, also 

southern Australia and New 

Zealand. 



Index of Scientific and Vernacular Names 1367 

INDEX OF SCIENTIFIC AND VERNACULAR NAMES 

Explanation of the System 

Italics : Valid scientific names (genera and species). 

Italics : Synonyms (genera and species), misidentifications. 

ROMAN : Family names. 

ROMAN : Names of divisions, classes, subclasses, orders, suborders, and subfamilies. 

Roman : FAO and local names.

1368 The Living Marine Resources of the Western Central Pacific 

A 

abaculus, Octopus . . . . . . . . . . . . . . . 800, 822 

Abralia . . . . . . . . . . . . . . . . . . . . . . . . 781 

Abraliopsis . . . . . . . . . . . . . . . . . . . . . . 781 

Acanthacaris . . . . . . . . . . . . . . . . . . . . . 977 

Acanthacaris opipara . . . . . . . . . . . . . . . . 988 

Acanthacaris tenuimana . . . . . . . . . . . . . . 988 

Acanthodes armatus . . . . . . . . . . . . . . . . 1106 

Acanthosepion pageorum . . . . . . . . . . . . . . 753 

Acanthosepion whitleyanum . . . . . . . . . . . . . 755 

acculeata, Holothuria . . . . . . . . . . . . . . . 1180 

Acetes . . . . . . . . . . . . . . . . . . . . . 855, 858 

Acetes erythraeus . . . . . . . . . . . . . . . . . . 862 

Acetes indicus . . . . . . . . . . . . . . . . . . . . 862 

Acetes intermedius . . . . . . . . . . . . . . . . . 863 

Acetes japonicus . . . . . . . . . . . . . . . . . . . 863 

Acetes serrulatus . . . . . . . . . . . . . . . . . . 864 

Acetes sibogae . . . . . . . . . . . . . . . . . . . . 864 

Acetes spp. . . . . . . . . . . . . . . . . . . . . . . 966 

Acetes vulgaris . . . . . . . . . . . . . . . . . . . 865 

Acheolus crassimanus . . . . . . . . . . . . . . . 1126 

Actinopyga echinites . . . . . . . . . . . . 1167-1168 

Actinopyga mauritiana . . . . . . . . . . . 1167-1168 

Actinopyga miliaris . . . . . . . . . . . . . 1169-1171 

Actinopyga palauensis . . . . . . . . 1169-1170-1171 

Actinopyga sp. . . . . . . . . . . . . . . . . 1167-1171 

Actinopyga spinea . . . . . . . . . . . . . . 1169-1171 

aculeata, Sepia . . . . . . . . . . . . . . . . . . . 736 

aculeatus, Octopusi . . . . . . . . . . . . . . 800, 822 

acuminata, Sepia . . . . . . . . . . . . . . . . . . 759 

acuta, Charybdis . . . . . . . . . . . . . . . . . . 1120 

acutidens, Negaprion . . . . . . . . . . . . 1350, 1352 

acutus, Rhizoprionodon . . . . . . . . . . 1354-1356 

acutus, Scoliodon . . . . . . . . . . . . . . 1351, 1354 

adhaesa, Solitosepia plangon . . . . . . . . . . . . 749 

Adonis shrimp . . . . . . . . . . . . . . . . . . . . 946 

aegina, Octopus . . . . . . . . . . . . . 811, 815, 819 

aeneus, Zosimus . . . . . . . . . . . . . . . 1049, 1098 

Aesop slipper lobster . . . . . . . . . . . . . . . . 1038 

affinis, Chaceon . . . . . . . . . . . . . . . . . . 1134 

affinis, Charybdis . . . . . . . . . . . . . . 1120, 1130 

affinis, Loliolus . . . . . . . . . . . . . . . . . . . 780 

affinis, Metapenaeus . . . . . . . . . . . . . . . . 933 

affinis, Sepia . . . . . . . . . . . . . . . . . . . . . 756 

Aiguillat à queue noire . . . . . . . . . . . . . . . 1230 

Aiguillat cyrano . . . . . . . . . . . . . . . . . . . 1231 

Aiguillat epinette . . . . . . . . . . . . . . . . . . 1230 

Aiguillat togari . . . . . . . . . . . . . . . . . . . 1229 

Aiquillat nez court . . . . . . . . . . . . . . . . . 1230 

Akiami paste shrimp . . . . . . . . . . . . . . . . . 863 

Alamang shrimp . . . . . . . . . . . . . . . . . . . 864 

Alaskan king crab . . . . . . . . . . . . . . 1048, 1055 

alatus, Isurus . . . . . . . . . . . . . . . . . . . . 1278 

albatrossi, Sepia kobiensis var. . . . . . . . . . . . 743 

albiflagellum, Panulirus . . . . . . . . . . 1015, 1017 

albimarginatus, Carcharhinus . . . . . . . 1325-1326 

albiventer, Holothuria . . . . . . . . . . . . . . . 1180 

albolineatus, Grapsus . . . . . . . . . . . . . . . 1142 

Alepisaurus ferox . . . . . . . . . . . . . . . . . . 799 

Alfombrera barbuda . . . . . . . . . . . . . . . . 1242 

Alfombrera collareja . . . . . . . . . . . . . . . . 1242 

alfonso, Solenocera . . . . . . . . . . . . . . 884-885 

aliae, Squaliolus . . . . . . . . . . . . . . . . . . 1229 

Alitán manchado . . . . . . . . . . . . . . . . . . 1292 

Alitán nubarrado . . . . . . . . . . . . . . . . . . 1292 

ALLOPOSIDAE . . . . . . . . . . . . . . . . . . . 802 

Alloposus pacificus . . . . . . . . . . . . . . . . . 802 

Aloha prawn . . . . . . . . . . . . . . . . . . . . . 920 

Alopias . . . . . . . . . . . . . . . . . . . . . . . 1271 

Alopias pelagicus . . . . . . . . . . . . . . 1271-1273 

Alopias profundus . . . . . . . . . . . . . . . . . 1272 

Alopias superciliosus . . . . . . . . . 1271-1272-1273 

Alopias vulpinus . . . . . . . . . . . . . . 1271-1273 

ALOPIIDAE . . . . . . . . . . 1196-1197, 1269, 1314 

ALPHEIDAE . . . . . . . . . . . . . . . . . . . . . 958 

alpheus, Octopus . . . . . . . . . . . . . . . 801, 823 

alticarinata, Solenocera . . . . . . . . 882, 885, 887 

altimus, Carcharhinus . . . . . . . . . . . . . . 1326 

Amber fish . . . . . . . . . . . . . . . . . . . . . 1190 

amblyrhinchoides, Carcharhinus . . . . . . . . . 1348 

amblyrhynchoides, Carcharhinus . 1327, 1332, 1340 

amblyrhynchos, Carcharhinus . . . . . . . . . . 1328 

amboinensis, Carcharhinus . . . . . . . . 1329, 1339 

amboinensis, Lysmata . . . . . . . . . . . . . . . 961 

Ameloctopus litoralis . . . . . . . . . . . . . . . . 820 

AMPHIONIDACEA . . . . . . . . . . . . . . . . . . 955 

AMPHITRETIDAE . . . . . . . . . . . . . . . . . . 802 

Amplisepia parysatis . . . . . . . . . . . . . . . . 738 

Amplisepia verreaux . . . . . . . . . . . . . . . . . 738 

ananas, Thelenota . . . . . . . . . . . . . . . . 1189 

anax, Thelenota . . . . . . . . . . . . . . . . . . 1190 

Anchisquilla fasciata . . . . . . . . . . . . . . . . 842 

anchistus, Metapenaeus . . . . . . . . . . . 909, 938 

anchoralis, Trachypenaeus . . . . . . . . . . . . . 948 

Ancistoteuthis . . . . . . . . . . . . . . . . . . . . 784 

Ancistrocheirus . . . . . . . . . . . . . . . . 781-782 

Ancistrocheirus lesueuri . . . . . . . . . . . . . . . 797 

Andaman lobster . . . . . . . . . . . . . . . . . . . 992 

andamanicus, Metanephrops . . . . . . . . . . . . 992 

Andrea cuttlefish . . . . . . . . . . . . . . . . . . . 737 

andreana, Sepia . . . . . . . . . . . . . . . . 737, 743 

andreanoides, Sepia . . . . . . . . . . . . . . . . . 743 

andreanoides, Sepia kobiensis var. . . . . . . . . . 743 

Ange de mer australien . . . . . . . . . . . . . . 1237 

Ange de mer Kasuzame . . . . . . . . . . . . . . 1237 

Angelote australiano . . . . . . . . . . . . . . . . 1237 

Angelote japones . . . . . . . . . . . . . . . . . . 1237 

Angelsharks . . . . . . . . . . . . . . . . . 1196, 1235 

angulatus, Puerulus . . . . . . . . . . . . . . . 1027 

anisodon, Charybdis . . . . . . . . . 1120, 1130-1131 

annulata, Charybdis . . . . . . . . . . . . 1120, 1131 

anomala, Thalassina . . . . . . . . . . . . . . . 1048 

ANOMURA . . . . . . . . . . . . . . 1048, 1077, 1154 

Anomuran crabs . . . . . . . . . . . . . . . . . . 1048 

Anomurans . . . . . . . . . . . . . . . . . . . . . 1048 

antarcticus, Mustelus . . . . . . . . . . . . . . . 1304 

antarcticus, Parribacus . . . . . . . . . . . . . . 1037


apama, Sepia . . . . . . . . . . . . . . . . . . . . 738 

APODA . . . . . . . . . . . . . . . . . . . . 1159-1161 

APODIDA . . . . . . . . . . . . . . . . . . . . . . 1160 

Aprionodon brevipinna . . . . . . . . . . . . . . 1332 

Apristurus . . . . . . . . . . . . . . . . . . . . . 1280 

Apristurus herklotsi . . . . . . . . . . . . . . . . 1286 

Apristurus longicephalus . . . . . . . . . . . . . 1286 

Apristurus sibogae . . . . . . . . . . . . . . . . 1286 

Apristurus spongiceps . . . . . . . . . . . . . . 1287 

Apristurus verweyi . . . . . . . . . . . . . . . . 1287 

Arafura shrimp . . . . . . . . . . . . . . . . . . . . 941 

arafurica, Parapenaeopsis . . . . . . . . . . . . . 941 

arakawai, Callistoctopus . . . . . . . . . . . . . . 817 

arakawai, Octopus . . . . . . . . . . . . . . . . . . 817 

arcticus, Galeocerdo . . . . . . . . . . . . . . . . 1349 

Arctides . . . . . . . . . . . . . . . . . . . . . . 1028 

Arctides regalis . . . . . . . . . . . . . . . . . . 1041 

arctipinnis, Sepioteuthis . . . . . . . . . . . . . . . 778 

Arctosepia braggi xera . . . . . . . . . . . . . . . 739 

Arctosepia limata . . . . . . . . . . . . . . . . . . 739 

Arctosepia rhoda . . . . . . . . . . . . . . . . . . . 739 

Arctosepia treba . . . . . . . . . . . . . . . . . . . 739 

Arctosepia versuta . . . . . . . . . . . . . . . . . . 739 

Argonauta . . . . . . . . . . . . . . . . . . . . . . 801 

ARGONAUTIDAE . . . . . . . . . . . . . . . 709, 801 

Argonauts . . . . . . . . . . . . . . . . . . . 689, 801 

argus, Bohadschia . . . . . . . . . . . . . . . . 1172 

Aristaeomorpha . . . . . . . . . . . . . . . . . . . 868 

Aristaeomorpha foliacea . . . . . . . . . . . . . . 872 

Aristaeomorpha rostridentata . . . . . . . . . . . . 872 

Aristaeopsis edwardsiana . . . . . . . . . . . . . . 874 

Aristeid shrimps . . . . . . . . . . . . . . . . . . . 868 

ARISTEIDAE . . . . . . . . . . 855, 868, 876, 890, 952 

ARISTEINAE . . . . . . . . . . . . . . . . . . . . . 868 

Aristeus virilis . . . . . . . . . . . . . . . . . . . . 873 

Arm-band ocellate octopus . . . . . . . . . . . . . 826 

armata, Hypothalassia . . . . . . . . . . . 1103, 1106 

armatus, Acanthodes . . . . . . . . . . . . . . . 1106 

Arrow squids . . . . . . . . . . . . . . . . . . . . . 788 

Arrowhead dogfish . . . . . . . . . . . . . . . . . 1225 

ashiaka, Penaeus . . . . . . . . . . . . . . . . . . . 925 

Ashtoret lunaris . . . . . . . . . . . 1091, 1095-1096 

asper, Trachypenaeus . . . . . . . . . . . . . 927, 950 

aspera, Schizophrys . . . . . . . . . . . . . . . . 1137 

ASPIDOCHIROTA . . . . . . . . . . . . . . 1160-1163 

ASPIDOCHIROTIDA . . . . . . . . . . . . . 1159, 1164 

aspilosomatis, Octopus . . . . . . . . . . . . 801, 823 

ASTEROIDEA . . . . . . . . . . . . . . . . . . . 1085 

Asteroids . . . . . . . . . . . . . . . . . . . 1158-1159 

ATELECYLIDAE . . . . . . . . . . . . . . . . . . 1050 

Atelomycterus fasciatus . . . . . . . . . . . . . . 1287 

Atelomycterus macleayi . . . . . . . . . . . . . . 1288 

Atelomycterus marmoratus . . . . . . . . . . . . 1288 

Atergatis floridus . . . . . . . . . . . . . . 1049, 1098 

Atergatis integerrimus . . . . . . . . . . . . . . . 1049 

Atergatis spp. . . . . . . . . . . . . . . . . . . . 1049 

Atergatopsis . . . . . . . . . . . . . . . . . . . . 1098 

Atergatopsis signatus . . . . . . . . . . . . . . . 1102 

atlanticus, Haliphron . . . . . . . . . . . . . . . . 802 

atra, Holothuria . . . . . . . . . . . . . . . 1177-1178 

atra, Holothuria (Halodeima) . . . . . . . . . . 1176 

atromarginatus, Centrophorus . . . . . . . . . . 1223 

attenuatus, Gollum . . . . . . . . . . . . . . . . 1295 

Atyid shrimps . . . . . . . . . . . . . . . . . . . . . 960 

ATYIDAE . . . . . . . . . . . . . . . . . . . . . . . 960 

Atyopsis moluccensis . . . . . . . . . . . . . . . . 960 

Atyopsis spinipes . . . . . . . . . . . . . . . . . . 960 

Atypopenaeus . . . . . . . . . . . . . . . . . . . . 890 

Atypopenaeus formosus . . . . . . . . . . . . . . 929 

Atypopenaeus stenodactylus . . . . . . . . . . . . 929 

Aulohalaelurus kanakorum . . . . . . . . . . . 1288 

Austral golden crab . . . . . . . . . . . . . . . . . 1135 

Australia mud shrimp . . . . . . . . . . . . . . . . . 886 

Australian angelshark . . . . . . . . . . . . . . . 1237 

Australian blacktip shark . . . . . . . . . . . . . . 1348 

Australian marbled catshark . . . . . . . . . . . . 1288 

Australian sawtail catshark . . . . . . . . . . . . . 1289 

Australian sharpnose shark . . . . . . . . . . . . 1356 

australiana, Solenocera . . . . . . . . . . . . 884, 886 

australiensis, Haliporoides sibogae . . . . . . . . . 881 

australiensis, Metanephrops . . . . . . . . . . . . 992 

australiensis, Ovalipes . . . . . . . . . . . . . . 1122 

australis, Chaceon . . . . . . . . . . . . . 1132, 1135 

australis, Loligo . . . . . . . . . . . . . . . . . . . 774 

australis, Octopus . . . . . . . . . . . . . . . . . . 824 

australis, Rossia . . . . . . . . . . . . . . . . . . . 715 

australis, Squatina . . . . . . . . . . . . . . . . 1237 

axiologa, Holothuria . . . . . . . . . . . . . . . 1182 

B 

Baby octopus . . . . . . . . . . . . . . . . . . 811, 819 

balfouri, Hemigaleus . . . . . . . . . . . . . . . 1308 

Bamboa capuchona . . . . . . . . . . . . . . . . 1259 

Bamboa elegante . . . . . . . . . . . . . . . . . 1255 

Bamboa estriada . . . . . . . . . . . . . . . . . . 1257 

Bamboa gris . . . . . . . . . . . . . . . . . . . . 1253 

Bamboa hombrera . . . . . . . . . . . . . . . . . 1258 

Bamboa jaspeada . . . . . . . . . . . . . . . . . 1258 

Bamboa moteada . . . . . . . . . . . . . . . . . 1259 

Bamboa ocelada . . . . . . . . . . . . . . . . . . 1258 

Bamboa punteada . . . . . . . . . . . . . . . . . 1256 

Banana prawn . . . . . . . . . . . . . . . . . . . . 921 

Banded long-arm octopus . . . . . . . . . . . . . . 820 

Banded mantis shrimps . . . . . . . . . . . . 829, 835 

Banded sand catshark . . . . . . . . . . . . . . . 1287 

Banded whip lobster . . . . . . . . . . . . . . . . 1027 

Banded-legged swimming crab . . . . . . . . . . 1131 

bandensis, Sepia . . . . . . . . . . . . . . . . . . . 757 

banksii, Matuta . . . . . . . . . . . . . . . . . . 1095 

baramensis, Metapenaeus ensis . . . . . . . . . . . 910 

barbata, Metapenaeopsis . . . . . . . . . . . . . . 907 

barbeensis, Metapenaeopsis . . . . . . . . . . . . 908 

Barbelthroat carpetshark . . . . . . . . . . . . . . 1242 

barbifer, Cirrhigaleus . . . . . . . . . . . . . . . 1224 

Bartail spurdog . . . . . . . . . . . . . . . . . . . 1231 

Bartlett’s cuttlefish . . . . . . . . . . . . . . . . . . 758 

bartletti, Sepia . . . . . . . . . . . . . . . . . . . . 758 

bartramii, Ommastrephes . . . . . . . . . . . . . . 793


Bartsch’s squid . . . . . . . . . . . . . . . . . . . . 779 

bartschi, Uroteuthis . . . . . . . . . . . . . . . . . 779 

Basking shark . . . . . . . . . . . . . . . . . . . 1274 

Batoid sawfishes . . . . . . . . . . . . . . . . . . 1233 

BATOIDEA . . . . . . . . . . . . . . . . . . . . . 1236 

Beaded rock crab . . . . . . . . . . . . . . . . . 1109 

Bêche-de-mer . . . . . . . . . 1163-1164, 1179, 1184 

Beka squid . . . . . . . . . . . . . . . . . . . . . . 772 

beka, Nipponololigo . . . . . . . . . . . . . . . . . 772 

belauensis, Nautilus . . . . . . . . . . . . . . . . . 711 

Bellybutton nautilus . . . . . . . . . . . . . . . . . 711 

benettae, Metapenaeus . . . . . . . . . . . . . . . 934 

BENTHESICYMINAE . . . . . . . . . . . . . . . . 868 

Benthic crabs . . . . . . . . . . . . . . . . . . . . 1085 

Benthic octopuses . . . . . . . . . . . . . . . . . . 800 

Benthoctopus . . . . . . . . . . . . . . . . . . . . . 801 

beppauna, Sepia kobiensis var. . . . . . . . . . . . 743 

Berrya . . . . . . . . . . . . . . . . . . . . . . . . 801 

bertholdii, Scyllarus . . . . . . . . . . . . . . . . 1043 

bicolor, Chaceon . . . . . . . . . . . . . . 1132, 1134 

Big blue octopus . . . . . . . . . . . . . . . . . . . 812 

Big bottom bobtail squid . . . . . . . . . . . . . . . 715 

Big-eyed bobtail . . . . . . . . . . . . . . . . . . . 716 

Bigeye sixgill shark . . . . . . . . . . . . . . . . . 1210 

Bigeye thresher . . . . . . . . . . . . . . . . . . . 1272 

Bigfin reef squid . . . . . . . . . . . . . . . . . . . 778 

Bignose shark . . . . . . . . . . . . . . . . . . . 1326 

bimaculatus, Octopus . . . . . . . . . . . . . . . . 812 

bipapillata, Rossia . . . . . . . . . . . . . . . . . . 716 

Bird shrimp . . . . . . . . . . . . . . . . . . . . . . 939 

Birgus latro . . . . . . . . . . 1048, 1061, 1154-1155 

birostrata, Sepiola . . . . . . . . . . . . . . . 716-717 

bispinatus, Euprotomicrus . . . . . . . . . . . . 1228 

bispinosa, Galene . . . . . . . . . . . . . . 1112-1113 

bivittata, Bohadschia . . . . . . . . . . . . . . . 1174 

Black teatfish . . . . . . . . . . . . . . . . . . . . 1183 

Blackbelly lanternshark . . . . . . . . . . . . . . 1226 

Blackfin gulper shark . . . . . . . . . . . . . . . . 1223 

Blackfish . . . . . . . . . . . . . . . . . . . 1170-1171 

Blackgill catshark . . . . . . . . . . . . . . . . . . 1291 

Blackspot shark . . . . . . . . . . . . . . . 1334, 1346 

Blackspotted sea cucumber . . . . . . . . . . . . 1184 

Blacktailed spurdog . . . . . . . . . . . . . . . . 1230 

Blacktip reef shark . . . . . . . . . . . . . . 1333, 1343 

Blacktip sawtail catshark . . . . . . . . . . . . . . 1290 

Blacktip shark . . . . . . . . . . . . . . . . . . . 1340 

Blacktip tope . . . . . . . . . . . . . . . . . . . . 1303 

Blacktipped reef shark . . . . . . . . . . . . . . . 1333 

Blanket octopuses . . . . . . . . . . . . . . . . . . 802 

bleekeri, Carcharhinus . . . . . . . . . . . . . . 1347 

Blind lobsters . . . . . . . . . . . . . . . . . . . . . 978 

Blind shark . . . . . . . . . . . . . . . . . . . . . 1244 

Blind sharks . . . . . . . . . . . . . . . . . . . . 1243 

blochii, Eusphyra . . . . . . . . . . . . . . . . . 1363 

blochii, Sphyrna . . . . . . . . . . . . . . . . . . 1363 

Blue land crab . . . . . . . . . . . . . . . . . . . 1151 

Blue shark . . . . . . . . . . . . . . . . . . . . . 1353 

Blue sharks . . . . . . . . . . . . . . . . . . . . . 1312 

Blue-lined octopus . . . . . . . . . . . . . . . . . . 820 

Bluegray carpetshark . . . . . . . . . . . . . . . . 1244 

Blunt slipper lobster . . . . . . . . . . . . . . . . 1039 

Blunt-toothed crab . . . . . . . . . . . . . . . . . 1131 

Bluntnose sixgill shark . . . . . . . . . . . . . . . 1210 

boardmani, Galeus . . . . . . . . . . . . . . . . 1289 

Bobtail squids . . . . . . . . . 688-689, 694, 712, 719 

bocki, Octopus . . . . . . . . . . . . . . . . . . . . 801 

boesemani, Halaelurus . . . . . . . . . . . . . . 1291 

Bohadschia . . . . . . . . . . . . . . . . . . . . 1161 

Bohadschia argus . . . . . . . . . . . . . . . . . 1172 

Bohadschia bivittata . . . . . . . . . . . . . . . . 1174 

Bohadschia graeffei . . . . . . . . . . . . . . . . 1184 

Bohadschia marmorata . . . . . . . . . . . . . . 1174 

Bohadschia similis . . . . . . . . . . . . . 1173-1174 

Bohadschia sp. . . . . . . . . . . . . . 1161, 1172-1174 

Bohadschia vitiensis . . . . . . . . . . . . . . . 1174 

BOLITAENIDAE . . . . . . . . . . . . . . . . . . . 802 

Bombay duck sharks . . . . . . . . . . . . . . . . 1357 

Bombay prawn . . . . . . . . . . . . . . . . . . . . 966 

Bonnethead sharks . . . . . . . . . . . . . . . . 1361 

borneensis, Carcharhinus . . . . . . . . . 1330, 1359 

Borneo catshark . . . . . . . . . . . . . . . . . . 1287 

Borneo river shark . . . . . . . . . . . . . . . . . 1360 

Borneo shark . . . . . . . . . . . . . . . . . . . . 1330 

Bottle squids . . . . . . . . . . . . . . . 688, 712, 719 

Bottletail squids . . . . . . . . . . . . . . . . . . . 719 

Boucot chevalier . . . . . . . . . . . . . . . . . . . 954 

Bouqet araignée . . . . . . . . . . . . . . . . . . . 968 

Bouqet bombay . . . . . . . . . . . . . . . . . . . 966 

Bouqet chagrin . . . . . . . . . . . . . . . . . . . . 967 

Bouqet géant . . . . . . . . . . . . . . . . . . . . . 964 

Bouqet mangrove . . . . . . . . . . . . . . . . . . 969 

Bouqet rosna . . . . . . . . . . . . . . . . . . . . . 965 

Bouqet singe . . . . . . . . . . . . . . . . . . . . . 967 

Bouqet tipattes . . . . . . . . . . . . . . . . . . . . 968 

Box crabs . . . . . . . . . . . . . . . . . . . 1057, 1091 

BRACHAELURIDAE . . . . . . . . . 1243, 1249, 1261 

Brachaelurus . . . . . . . . . . . . . . . . . . . . 1243 

Brachaelurus waddi . . . . . . . . . . . . . . . . 1244 

BRACHYURA . . . . . . . . . . . . 1048, 1050, 1083 

Brachyuran crabs . . . . . . . . . . . . . . 1048, 1061 

Brachyurans . . . . . . . . . . . . . . . . . 1048, 1050 

brachyurus, Carcharhinus . . . . . . . . . . . . 1331 

brachyurus, Etmopterus . . . . . . . . . . . . . 1226 

braggi xera, Arctosepia . . . . . . . . . . . . . . . 739 

braggi, Sepia . . . . . . . . . . . . . . . . . . . . 739 

Bramble shark . . . . . . . . . . . . . . . . . . . 1212 

Bramble sharks . . . . . . . . . . . . . . . . . . . 1211 

brasiliensis, Isistius . . . . . . . . . . . . . 1213, 1228 

brevicornis, Metapenaeus . . . . . . . . . . . . . 934 

brevimana, Sepia . . . . . . . . . . . . 740, 754, 761 

brevipinna, Aprionodon . . . . . . . . . . . . . . 1332 

brevipinna, Carcharhinus . . 1327, 1332, 1340, 1348 

brevirostris, Phoberus . . . . . . . . . . . . . . . . 988 

brevirostris, Squalus . . . . . . . . . . . . . . . . 1230 

Brittle stars . . . . . . . . . . . . . . . . . . . . . 1158 

Broadclub cuttlefish . . . . . . . . . . . . . . . . . 744 

Broadfin shark . . . . . . . . . . . . . . . . . . . 1350 

Brown land crab . . . . . . . . . . . . . . . . . . 1149 

Brown sandfish . . . . . . . . . . . . . . . . . . . 1174 

Brown tiger prawn . . . . . . . . . . . . . . . . . . 915


Brownbanded bambooshark . . . . . . . . . . . . 1257 

Brownspotted catshark . . . . . . . . . . . . . . . 1292 

Brownspotted sandfish . . . . . . . . . . . . . . . 1173 

brucus, Echinorhinus . . . . . . . . . . . . . . . 1212 

Bruja terciopelo . . . . . . . . . . . . . . . . . . . 1228 

bubulus, Penaeus . . . . . . . . . . . . . . . . . . . 922 

budo, Loligo . . . . . . . . . . . . . . . . . . . . . 776 

Bull shark . . . . . . . . . . . . . . . 1196-1197, 1339 

Bullhead sharks . . . . . . . . . . . . . . . . . . 1238 

Bullseye reef lobster . . . . . . . . . . . . . . . . 1000 

burgeri, Panulirus . . . . . . . . . . . . . . . . . 1016 

burkenroadi, Metapenaeus . . . . . . . . . . . . . 912 

Butterfly bobtail . . . . . . . . . . . . . . . . . . . . 716 

C 

Cacahouète indopacifique . . . . . . . . . . . . . 1004 

caecus sublevis, Phoberus . . . . . . . . . . . . . . 988 

caesius, Penaeus . . . . . . . . . . . . . . . . . . . 919 

cailloti, Pelocarcinus . . . . . . . . . . . . . . . 1150 

Calamar beka . . . . . . . . . . . . . . . . . . . . 772 

Calamar buril . . . . . . . . . . . . . . . . . . . . . 777 

Calamar espada . . . . . . . . . . . . . . . . . . . 776 

Calamar índico . . . . . . . . . . . . . . . . . . . . 775 

Calamar kobí . . . . . . . . . . . . . . . . . . . . . 773 

Calamar manopla . . . . . . . . . . . . . . . . . . 778 

Calamar mitrado . . . . . . . . . . . . . . . . . . . 774 

Calamarete . . . . . . . . . . . . . . . . . . . . . . 779 

Calappa . . . . . . . . . . . . . . . . . . . . . . 1094 

Calappa calappa . . . . . . . . . . . . . . . 1094, 1096 

Calappa guerini . . . . . . . . . . . . . . . . . . 1094 

Calappa hepatica . . . . . . . . . . . . . . 1094, 1097 

Calappa lophos . . . . . . . . . . . . . . . 1091, 1094 

Calappa philargius . . . . . . . . . 1091, 1094, 1097 

calappa, Calappa . . . . . . . . . . . . . . 1094, 1096 

CALAPPIDAE . . . . . . . . . 1050, 1057, 1091, 1115 

CALAPPINAE . . . . . . . . . . . . . . . . . . . 1091 

Caledonian mitten lobster . . . . . . . . . . . . . 1041 

caledonicus, Parribacus . . . . . . . . . . . . . 1041 

CALLIANASSIDAE . . . . . . . . . . . . . . . . . 976 

Callistoctopus arakawai . . . . . . . . . . . . . . . 817 

Calmar baril . . . . . . . . . . . . . . . . . . . . . 777 

Calmar cracheur . . . . . . . . . . . . . . . . . . . 772 

Calmar épée . . . . . . . . . . . . . . . . . . . . . 776 

Calmar indien . . . . . . . . . . . . . . . . . . . . 775 

Calmar kobi . . . . . . . . . . . . . . . . . . . . . . 773 

Calmar mitre . . . . . . . . . . . . . . . . . . . . . 774 

Calmar tépo . . . . . . . . . . . . . . . . . . . . . 779 

Calmar tonnelet . . . . . . . . . . . . . . . . . . . 778 

Camarón adonis . . . . . . . . . . . . . . . . . . . 946 

Camarón aguji . . . . . . . . . . . . . . . . . . . . 948 

Camarón aloha . . . . . . . . . . . . . . . . . . . . 920 

Camarón amarillo . . . . . . . . . . . . . . . . . . 934 

Camarón arafura . . . . . . . . . . . . . . . . . . . 941 

Camarón araña . . . . . . . . . . . . . . . . . . . . 968 

Camarón arco iris . . . . . . . . . . . . . . . . . . 944 

Camarón banana . . . . . . . . . . . . . . . . . . . 921 

Camarón blanco de la India . . . . . . . . . . . . . 916 

Camarón brujo . . . . . . . . . . . . . . . . . . . . 914 

Camarón cigueña . . . . . . . . . . . . . . . . . . 941 

Camarón cintura . . . . . . . . . . . . . . . . . . . 911 

Camarón colorado . . . . . . . . . . . . . . . . . . 923 

Camarón coral . . . . . . . . . . . . . . . . . . . . 942 

Camarón coromandel . . . . . . . . . . . . . . . . 942 

Camarón cortapluma. . . . . . . . . . . . . . . . . 881 

Camarón costurón . . . . . . . . . . . . . . . . . . 882 

Camarón dalí . . . . . . . . . . . . . . . . . . . . . 935 

Camarón de Bombay . . . . . . . . . . . . . . . . 966 

Camarón de manglar . . . . . . . . . . . . . . . . . 969 

Camarón de piedra lanzón . . . . . . . . . . . . . . 954 

Camarón devo . . . . . . . . . . . . . . . . . . . . 938 

Camarón diablo . . . . . . . . . . . . . . . . . . . 936 

Camarón esmeralda . . . . . . . . . . . . . . . . . 938 

Camarón fanguero chino . . . . . . . . . . . . . . . 887 

Camarón fanguero de orilla . . . . . . . . . . . . . 883 

Camarón fijador arquero . . . . . . . . . . . . . . . 927 

Camarón fijador de granos . . . . . . . . . . . . . . 949 

Camarón fijador malayo . . . . . . . . . . . . . . . 928 

Camarón fijador norteño . . . . . . . . . . . . . . . 949 

Camarón fino . . . . . . . . . . . . . . . . . . . . . 937 

Camarón flamenco . . . . . . . . . . . . . . . . . . 947 

Camarón gamuza barbudo . . . . . . . . . . . . . . 907 

Camarón gamuza mogi . . . . . . . . . . . . . . . 930 

Camarón gamuza norteño . . . . . . . . . . . . . . 931 

Camarón gamuza rosado . . . . . . . . . . . . . . 931 

Camarón gamuza sureño . . . . . . . . . . . . . . 908 

Camarón gamuza tolo . . . . . . . . . . . . . . . . 932 

Camarón gigante . . . . . . . . . . . . . . . . . . . 964 

Camarón huesudo . . . . . . . . . . . . . . . . . . 948 

Camarón jinga . . . . . . . . . . . . . . . . . . . . 933 

Camarón jorobado . . . . . . . . . . . . . . . . . . 930 

Camarón kadal . . . . . . . . . . . . . . . . . . . . 936 

Camarón kuruma . . . . . . . . . . . . . . . . . . . 917 

Camarón lanzón . . . . . . . . . . . . . . . . . . . 913 

Camarón leña . . . . . . . . . . . . . . . . . . . . 935 

Camarón lija . . . . . . . . . . . . . . . . . . . . . 967 

Camarón liso . . . . . . . . . . . . . . . . . . . . . 945 

Camarón maclayo . . . . . . . . . . . . . . . . . . 939 

Camarón mono . . . . . . . . . . . . . . . . . . . . 967 

Camarón moyebi . . . . . . . . . . . . . . . . . . . 912 

Camarón nailón mino . . . . . . . . . . . . . . . . 970 

Camarón naranji. . . . . . . . . . . . . . . . . . . . 929 

Camarón ñata . . . . . . . . . . . . . . . . . . . . 961 

Camarón papuense . . . . . . . . . . . . . . . . . 940 

Camarón parancero . . . . . . . . . . . . . . . . . 939 

Camarón patojo . . . . . . . . . . . . . . . . . . . 968 

Camarón peine . . . . . . . . . . . . . . . . . . . . 887 

Camarón periscopio . . . . . . . . . . . . . . . . . 929 

Camarón perro . . . . . . . . . . . . . . . . . . . . 943 

Camarón rabo verde . . . . . . . . . . . . . . . . . 934 

Camarón real . . . . . . . . . . . . . . . . . . . . . 918 

Camarón real manchado . . . . . . . . . . . . . . . 919 

Camarón real oriental . . . . . . . . . . . . . . . . 924 

Camarón resbaloso . . . . . . . . . . . . . . . . . 910 

Camarón rosna . . . . . . . . . . . . . . . . . . . . 965 

Camarón soldado . . . . . . . . . . . . . . . . . . 960 

Camarón tigre gigante . . . . . . . . . . . . . . . . 922 

Camarón tigre marrón . . . . . . . . . . . . . . . . 915 

Camarón tigre verde . . . . . . . . . . . . . . . . . 925 

Camarón torpedo . . . . . . . . . . . . . . . . . . 944 

Camarón uncta . . . . . . . . . . . . . . . . . . . . 945


Camarón violinista . . . . . . . . . . . . . . . . . . 932 

Camarón york . . . . . . . . . . . . . . . . . . . . 937 

Camaroncillo akiami . . . . . . . . . . . . . . . . . 863 

Camaroncillo alamang . . . . . . . . . . . . . . . . 864 

Camaroncillo javlá . . . . . . . . . . . . . . . . . . 862 

Camaroncillo jembre . . . . . . . . . . . . . . . . . 865 

Camaroncillo mauxia dè Formosa . . . . . . . . . . 863 

Camaroncillo mauxia sureño . . . . . . . . . . . . . 864 

Camaroncillo tsivakihini . . . . . . . . . . . . . . . 862 

CAMPTANDRIIDAE . . . . . . . . . . . . . . . . 1050 

camtschaticus, Paralithodes . . . . . . . . . 1048, 1055 

Cañabota . . . . . . . . . . . . . . . . . . . . . . 1210 

Cañabota bocadulce . . . . . . . . . . . . . . . . 1210 

Cañabota gris . . . . . . . . . . . . . . . . . . . 1210 

canaliculatus, Penaeus . . . . . . . . . . . . . . . 914 

Cancer dormitator . . . . . . . . . . . . . . . . . 1087 

Cancer lunaris . . . . . . . . . . . . . . . . . . . 1095 

Cannonball sponge crab . . . . . . . . . . . . . . 1088 

Capricorn night octopus . . . . . . . . . . . . . . . 823 

CARAPIDAE . . . . . . . . . . . . . . . . . . . . 1189 

Carcharhinid sharks . . . . . . . . . . . . . . . . 1197 

CARCHARHINIDAE . . . . . . 1196-1197, 1264, 1297, 

1305-1306, 1312 

Carcharhinoid sharks . . . . . . . . . . . . . . . 1196 

Carcharhinus . . . . . . . . . . . . . . . . . . . 1196 

Carcharhinus albimarginatus . . . . . . . 1325-1326 

Carcharhinus altimus . . . . . . . . . . . . . . . 1326 

Carcharhinus amblyrhinchoides . . . . . . . . . 1348 

Carcharhinus amblyrhynchoides . 1327, 1332, 1340 

Carcharhinus amblyrhynchos . . . . . . . . . . 1328 

Carcharhinus amboinensis . . . . . . . . . 1329, 1339 

Carcharhinus bleekeri . . . . . . . . . . . . . . . 1347 

Carcharhinus borneensis . . . . . . . . . . 1330, 1359 

Carcharhinus brachyurus . . . . . . . . . . . . 1331 

Carcharhinus brevipinna . . . 1327, 1332, 1340, 1348 

Carcharhinus cautus . . . . . . . . . . . . . . . 1333 

Carcharhinus dussumieri . . . . . . . . . . 1334, 1346 

Carcharhinus elliot . . . . . . . . . . . . . . . . 1310 

Carcharhinus falciformis . . . . . . . . . . 1335, 1341 

Carcharhinus fitzroyensis . . . . . . . . . . . . 1336 

Carcharhinus galapagensis . . . . . . . . 1326, 1337 

Carcharhinus hemiodon . . . . . . . . . . . . . 1338 

Carcharhinus iranzae . . . . . . . . . . . . . . . 1344 

Carcharhinus johnsoni . . . . . . . . . . . . . . 1332 

Carcharhinus leucas . . . . . . . . 1196, 1329, 1339 

Carcharhinus limbatus . . . . 1327, 1332, 1340, 1348 

Carcharhinus longimanus . . . . . . . . . 1325, 1341 

Carcharhinus macloti . . . . . 1342, 1351, 1354-1356 

Carcharhinus maou . . . . . . . . . . . . . . . . 1341 

Carcharhinus melanopterus . . . . . . . . 1333, 1343 

Carcharhinus menisorrah . . . . . . 1328, 1334, 1346 

Carcharhinus milberti . . . . . . . . . . . . . . . 1345 

Carcharhinus obscurus . . . . . . . . . . . 1326, 1344 

Carcharhinus pleurotaenia . . . . . . . . . . . . 1327 

Carcharhinus plumbeus . . . . . . . . . . . . . 1345 

Carcharhinus porosus . . . . . . . . . . . . . . . 1359 

Carcharhinus radamae . . . . . . . . . . . . . . 1326 

Carcharhinus remotus . . . . . . . . . . . . . . . 1331 

Carcharhinus sealei . . . . . . . . . . . . . 1334, 1346 

Carcharhinus sorrah . . . . . . . . . . . . . . . 1347 

Carcharhinus sp. . . . . . . . . . . . . . . . . . 1359 

Carcharhinus spp. . . . . . . . . . . . . . . . . . 1332 

Carcharhinus temmincki . . . . . . . . . . . . . 1350 

Carcharhinus tilstoni . . . . . . . . . . . . 1340, 1348 

Carcharhinus tjutjot . . . . . . . . . . . . . . . . 1334 

Carcharhinus vanrooyeni . . . . . . . . . . . . . 1339 

Carcharhinus wheeleri . . . . . . . . . . . . . . 1328 

Carcharhinus zambezensis . . . . . . . . . . . . 1339 

Carcharias taurus . . . . . . . . . . . 1196, 1266-1267 

Carcharias tricuspidatus . . . . . . . . . . . . . 1266 

carcharias, Carcharodon . . . . . . . . . . . . . 1276 

Carcharodon carcharias . . . . . . . . . . . . . 1276 

Carcinoplax . . . . . . . . . . . . . . . . . . . . 1114 

Carcinoplax longimana . . . . . . . . . . . 1114, 1132 

Carcinoplax longimanus japonicus . . . . . . . . 1114 

Carcinoplax longimanus typicus . . . . . . . . . 1114 

carcinus, Macrobrachium . . . . . . . . . . . . . . 964 

Cardinal shrimp . . . . . . . . . . . . . . . . . . . 962 

Cardisoma . . . . . . . . . . . . . . . . . . . . . 1049 

Cardisoma carnifex . . . . . . . . . . 1147, 1149-1150 

Cardisoma hirtipes . . . . . . . . . . . . . 1149, 1151 

Cardisoma longipes . . . . . . . . . . . . . 1149, 1151 

Cardisoma obesum . . . . . . . . . . . . . . . . 1149 

Cardisoma rotundum . . . . . . . . . . . . 1149, 1151 

Cardisoma spp. . . . . . . . . . . . . . . . . . . 1049 

Cardisoma urvillei . . . . . . . . . . . . . . . . . 1149 

CARIDEA . . . . 855-856, 859, 867, 870, 877, 891, 953, 

956-957 

Caridean shrimps . . . . . . . . . . 855-856, 957-958 

Carideans . . . . . . . . . . . . . . . . . . . . . . 855 

Caridina weberi . . . . . . . . . . . . . . . . . . . 961 

carinatus, Penaeus . . . . . . . . . . . . . . . . . . 922 

carnifex, Cardisoma . . . . . . . . . 1147, 1149-1150 

Carocho . . . . . . . . . . . . . . . . . . . . . . 1225 

caroli stenodactyla, Ommastrephes . . . . . . . . . 793 

CARPILIIDAE . . 1049, 1058, 1098, 1103, 1110, 1114 

Carpilius . . . . . . . . . . . . . . . . . . . . . . 1110 

Carpilius convexus . . . . . . . . . . . . . 1110-1111 

Carpilius maculatus . . . . . . . . . . . . . 1110-1111 

Carpilius spp. . . . . . . . . . . . . . . . . . . . 1049 

Catsharks . . . . . . . . . . . . . . . . . . . . . 1279 

cautus, Carcharhinus . . . . . . . . . . . . . . . 1333 

Cazón . . . . . . . . . . . . . . . . . . . . . . . 1302 

Cazón coralero trompacorta . . . . . . . . . . . . 1358 

Cazón de aleta blanca . . . . . . . . . . . . . . . 1302 

Cazón elegant . . . . . . . . . . . . . . . . . . . 1303 

Cazón espadachin . . . . . . . . . . . . . . . . . 1357 

Cazón picudo . . . . . . . . . . . . . . . . . 1303, 1354 

Cazón picudo australiano . . . . . . . . . . . . . 1356 

Cazón picudo gris . . . . . . . . . . . . . . . . . 1355 

Cazón velero . . . . . . . . . . . . . . . . . . . . 1302 

celetaria pacifica, Histioteuthis . . . . . . . . . . 787 

Centrophorus atromarginatus . . . . . . . . . . 1223 

Centrophorus granulosus . . . . . . . . . 1223-1224 

Centrophorus lusitanicus . . . . . . . . . . . . . 1224 

Centrophorus moluccensis . . . . . . . . . . . . 1223 

Centrophorus niaukang . . . . . . . . . . . . . 1224 

Centrophorus squamosus . . . . . . . . . . . . . 1224 

Centroscymnus . . . . . . . . . . . . . . . . . . . 1213


Cephaloscyllium fasciatum . . . . . . . . . . . . 1289 

ceratophthalma, Ocypode . . . . . . . . . . 1152-1153 

CETORHINIDAE . . . . . . . . . . . . . . . . . . 1274 

Cetorhinus maximus . . . . . . . . . . . . . . . . 1274 

ceylonensis, Scoliodon . . . . . . . . . . . . . . . 1351 

Chaceon . . . . . . . . . . . . . . . . . . . 1132, 1134 

Chaceon affinis . . . . . . . . . . . . . . . . . . 1134 

Chaceon australis . . . . . . . . . . . . . . 1132, 1135 

Chaceon bicolor . . . . . . . . . . . . . . . 1132, 1134 

Chaceon granulatus . . . . . . . . . . . . 1132, 1134 

Chaceon karubar . . . . . . . . . . 1132, 1134-1135 

Chaceon manningi . . . . . . . . . . . . . . . . . 1134 

Chaceon maritae . . . . . . . . . . . . . . . . . . 1132 

Chaceon poupini . . . . . . . . . . . . . . 1132, 1135 

Chaenogaleus macrostoma . . . . . . . . . 1308-1310 

Chambered nautiluses . . . . . . . . . . . . . . . . 709 

Champagne crab . . . . . . . . . . . . . . . . . . 1106 

chani, Justitia . . . . . . . . . . . . . . . . . . . 1022 

Charybdis . . . . . . . . . . . . . . . . . . 1120-1121 

Charybdis acuta . . . . . . . . . . . . . . . . . . 1120 

Charybdis affinis . . . . . . . . . . . . . . 1120, 1130 

Charybdis anisodon . . . . . . . . . . . . . 1120, 1130 

Charybdis annulata . . . . . . . . . . . . . 1120, 1131 

Charybdis cruciata . . . . . . . . . . . . . . . . 1120 

Charybdis crucifer . . . . . . . . . . . . . . . . . 1120 

Charybdis feriatus . . . . . . . . . . 1115, 1120-1121 

Charybdis japonica . . . . . . . . . . . . . . . . 1121 

Charybdis natator . . . . . . . . . . . . . . 1120-1121 

Charybdis truncata . . . . . . . . . . . . . 1120, 1131 

CHEIRAGONIDAE . . . . . . . . . . . . . . . . . 1050 

chengtongense, Episesarma . . . . . . . . 1143, 1145 

Chevrette akiami . . . . . . . . . . . . . . . . . . . 863 

Chevrette alamang . . . . . . . . . . . . . . . . . . 864 

Chevrette jawla . . . . . . . . . . . . . . . . . . . . 862 

Chevrette jembre . . . . . . . . . . . . . . . . . . . 865 

Chevrette mauxia de Formose . . . . . . . . . . . . 863 

Chevrette mauxia méridionale . . . . . . . . . . . . 864 

Chevrette tsivakihini . . . . . . . . . . . . . . . . . 862 

Chien corail . . . . . . . . . . . . . . . . . . . . . 1288 

Chien égoïne . . . . . . . . . . . . . . . . . . . . 1289 

Chien gecko . . . . . . . . . . . . . . . . . . . . 1289 

Chien lime . . . . . . . . . . . . . . . . . . . . . 1290 

Chien marbré . . . . . . . . . . . . . . . . . . . . 1288 

Chien nain . . . . . . . . . . . . . . . . . . . . . 1290 

Chiloscyllium colax . . . . . . . . . . . . . . . . 1255 

Chiloscyllium griseum . . . . 1253-1254, 1256-1257 

Chiloscyllium hasselti . . . . . 1253-1254, 1256-1257 

Chiloscyllium indicum . . . . . . . . . . . 1255-1256 

Chiloscyllium margaritiferum . . . . . . . . . . . 1257 

Chiloscyllium plagiosum . . . 1253-1254, 1256-1257 

Chiloscyllium punctatum . . . 1253-1254, 1256-1257 

Chiloscyllium sp. . . . . . . . . . . . . . . 1249, 1254 

China lobster . . . . . . . . . . . . . . . . . . . . . 993 

chinensis, Loligo . . . . . . . . . . . . . . . . . . . 774 

chinensis, Photololigo . . . . 765-766, 774-775, 777 

Chinese mitten crabs . . . . . . . . . . . . . . . . 1055 

Chinese mud shrimp . . . . . . . . . . . . . . . . . 887 

Chinese spiny lobster . . . . . . . . . . . . . . . 1026 

Chionoecetes japonicus . . . . . . . . . . . . . . 1055 

Chionoecetes opilio . . . . . . . . . . . . . . . . 1055 

Chipirón volantín . . . . . . . . . . . . . . . . . . . 797 

Chipliloua commun . . . . . . . . . . . . . . . . . . 797 

CHIRODOTIDAE . . . . . . . . . . . . . . . . . . 1160 

Chirostylids . . . . . . . . . . . . . . . . . . . . . 1048 

Chiroteuthid squids . . . . . . . . . . . . . . . . . . 798 

CHIROTEUTHIDAE . . . . . . . . . . . . . . 798-799 

Chiroteuthis . . . . . . . . . . . . . . . . . . . . . 798 

Chiroteuthis imperator . . . . . . . . . . . . . . . 798 

chloronotus, Stichopus . . . . . . . . . . . . . . 1186 

choprai, Solenocera . . . . . . . . . . . 882, 885, 887 

Cigala de Andamán . . . . . . . . . . . . . . . . . 992 

Cigala del Oceano Indico . . . . . . . . . . . . . . 991 

Cigala raspa . . . . . . . . . . . . . . . . . . . . . 988 

Cigale glabre . . . . . . . . . . . . . . . . . . . . 1035 

Cigale grenue . . . . . . . . . . . . . . . . . . . 1039 

Cigale raquette . . . . . . . . . . . . . . . . . . . 1040 

Cigale savate . . . . . . . . . . . . . . . . . . . . 1037 

Cigarra chata . . . . . . . . . . . . . . . . . . . . 1040 

Cigarra chinesa . . . . . . . . . . . . . . . . . . 1037 

Cigarra liso . . . . . . . . . . . . . . . . . . . . . 1035 

Cigarra ñato . . . . . . . . . . . . . . . . . . . . 1039 

ciliatus pubescens, Ibacus . . . . . . . . . . . . 1036 

ciliatus, Ibacus . . . . . . . . . . . . . . . 1034-1036 

CIRRATA . . . . . . . . . . . . . . . . . . . . 689, 801 

Cirrate octopods . . . . . . . . . . . . . . . . . . . 694 

Cirrate octopuses . . . . . . . . . . . . . . . 689, 801 

cirratus, Pristiophorus . . . . . . . . . . . . . . 1234 

cirrhatus, Eptatretus . . . . . . . . . . . . . . . . 1192 

Cirrhigaleus barbifer . . . . . . . . . . . . . . . 1224 

Cirrhoscyllium expolitum . . . . . . . . . . . . . 1242 

CIRROTEUTHIDAE . . . . . . . . . . . . . . . . . 801 

Cistopus indicus . . . . . . . . . . . . . . . . . . . 810 

Cleaner shrimps . . . . . . . . . . . . . . . . . . . 955 

Cloridopsis . . . . . . . . . . . . . . . . . . . 842, 846 

Cloridopsis scorpio . . . . . . . . . . . . . . 842, 846 

Cloudy catshark . . . . . . . . . . . . . . . . . . 1292 

Coarse shrimp . . . . . . . . . . . . . . . . . . . . 949 

Coastal mud shrimp . . . . . . . . . . . . . . . . . 883 

Coconut crab . . . . . . . . . . . . . . . . . 1048, 1155 

Coconut crabs . . . . . . . . . . . . . . . . 1061, 1154 

Coenobita . . . . . . . . . . . . . . . . . . 1048, 1154 

COENOBITIDAE . . . . . . . . . . . . . . . 1061, 1154 

Coenobitids . . . . . . . . . . . . . . . . . . . . 1154 

colax, Chiloscyllium . . . . . . . . . . . . . . . . 1255 

colcloughi, Heteroscyllium . . . . . . . . . . . . 1244 

collare, Parascyllium . . . . . . . . . . . . . . . 1242 

Collared carpetshark . . . . . . . . . . . . . . . . 1242 

Collared carpetsharks . . . . . . . . . . . . . . . 1241 

coluber, Holothuria . . . . . . . . . . . . . . . . 1178 

coluber, Holothuria (Acanthotrapeza) . . . . . 1175 

Comadreja coluda . . . . . . . . . . . . . . . . . 1311 

Comadreja ganchuda . . . . . . . . . . . . . . . 1308 

Comadreja segadora . . . . . . . . . . . . . . . . 1309 

Comadreja sobrediente . . . . . . . . . . . . . . 1310 

Comb shrimp . . . . . . . . . . . . . . . . . . . . . 887 

Common banded mantis shrimp . . . . . . . . . . . 837 

Common box crab . . . . . . . . . . . . . . . . . 1094 

Common cleaner shrimp . . . . . . . . . . . . . . . 961 

Common decorator crab . . . . . . . . . . . . . . 1137


Common ghost crab . . . . . . . . . . . . . . . . 1153 

Common moon crab . . . . . . . . . . . . . . . . 1095 

Common sponge crab . . . . . . . . . . . . . . . 1087 

Common squillid mantis shrimp . . . . . . . . . . . 848 

Common Sydney octopus . . . . . . . . . . . . . . 818 

concinnus, Palaemon . . . . . . . . . . . . . . . . 969 

concolor, Nebrius . . . . . . . . . . . . . . . . . 1260 

conjunctus, Metapenaeus . . . . . . . . . . . . . . 935 

convexus, Carpilius . . . . . . . . . . . . . 1110-1111 

convexus, Etisus . . . . . . . . . . . . . . . . . . 1100 

cookei, Echinorhinus . . . . . . . . . . . . . . . 1212 

Cookiecutter shark . . . . . . . . . . . . . . 1213, 1228 

Copper shark . . . . . . . . . . . . . . . . . . . . 1331 

Coral catshark . . . . . . . . . . . . . . . . . . . 1288 

Coral reef crabs . . . . . . . . . . . . . . . . . . 1049 

Coral shrimp . . . . . . . . . . . . . . . . . . . . . 942 

cordiformis, Mastigoteuthis . . . . . . . . . . . . . 799 

cordimanus, Ocypode . . . . . . . . . . . . . . . 1153 

Cornuda común . . . . . . . . . . . . . . . . . . 1364 

Cornuda cruz . . . . . . . . . . . . . . . . . . . . 1366 

Cornuda gigante . . . . . . . . . . . . . . . . . . 1365 

Cornuda planeadora . . . . . . . . . . . . . . . . 1363 

cornuta, Parapenaeopsis . . . . . . . . . . . 942, 944 

Coromandel shrimp . . . . . . . . . . . . . . . . . 942 

coromandelica, Parapenaeopsis . . . . . . . . . . 942 

CORYSTIDAE . . . . . . . . . . . . . . . . . . . 1089 

cottlesloensis, Decorisepia . . . . . . . . . . . . . 751 

Cotton’s cuttlefish . . . . . . . . . . . . . . . . . . 758 

cottoni, Sepia . . . . . . . . . . . . . . . . . . . . 758 

Cowsharks . . . . . . . . . . . . . . . . . . . . . 1208 

Crabs . . . . . . . . . . . . . . . . . . . . . . . . 1333 

Cranchiid squids . . . . . . . . . . . . . . . . . . . 694 

cranioides, Dromidiopsis . . . . . . . . . . . . . 1088 

crassa, Sepia kobiensis var. . . . . . . . . . . . . . 743 

crassicornis, Solenocera . . . . . . . . . . . . . . 883 

crassimanus, Acheolus . . . . . . . . . . . . . . . 1126 

crebripunctata, Matuta . . . . . . . . . . . . . . 1095 

Creek whaler . . . . . . . . . . . . . . . . . . . . 1336 

crenata, Thalamita . . . . . . . . . . . . . . . . 1129 

Crenate swimming crab . . . . . . . . . . . . . . 1129 

Crested bullhead shark . . . . . . . . . . . . . . 1240 

Crested reef crab . . . . . . . . . . . . . . . . . . 1049 

Crevette adonis . . . . . . . . . . . . . . . . . . . . 946 

Crevette aiguille . . . . . . . . . . . . . . . . . . . 948 

Crevette aloha . . . . . . . . . . . . . . . . . . . . 920 

Crevette arafura . . . . . . . . . . . . . . . . . . . 941 

Crevette arc-en-ciel . . . . . . . . . . . . . . . . . 944 

Crevette banane . . . . . . . . . . . . . . . . . . . 921 

Crevette bois . . . . . . . . . . . . . . . . . . . . . 935 

Crevette bossue . . . . . . . . . . . . . . . . . . . 930 

Crevette ceinture . . . . . . . . . . . . . . . . . . . 911 

Crevette chamois barbulée . . . . . . . . . . . . . . 907 

Crevette chamois méridionale . . . . . . . . . . . . 908 

Crevette chamois mogi . . . . . . . . . . . . . . . . 930 

Crevette chamois nordique . . . . . . . . . . . . . . 931 

Crevette chamois rosée . . . . . . . . . . . . . . . 931 

Crevette chamois tolo . . . . . . . . . . . . . . . . 932 

Crevette chien . . . . . . . . . . . . . . . . . . . . 943 

Crevette cigogne . . . . . . . . . . . . . . . . . . . 941 

Crevette corail . . . . . . . . . . . . . . . . . . . . 942 

Crevette coromandel . . . . . . . . . . . . . . . . . 942 

Crevette dali . . . . . . . . . . . . . . . . . . . . . 935 

Crevette de maclay . . . . . . . . . . . . . . . . . . 939 

Crevette devo . . . . . . . . . . . . . . . . . . . . 938 

Crevette diable . . . . . . . . . . . . . . . . . . . . 936 

Crevette élégante . . . . . . . . . . . . . . . . . . 937 

Crevette émeraude . . . . . . . . . . . . . . . . . . 938 

Crevette flamand . . . . . . . . . . . . . . . . . . . 947 

Crevette gambri archée . . . . . . . . . . . . . . . 927 

Crevette gambri grenue . . . . . . . . . . . . . . . 949 

Crevette gambri malaise . . . . . . . . . . . . . . . 928 

Crevette gambri nordique . . . . . . . . . . . . . . 949 

Crevette géante tigrée . . . . . . . . . . . . . . . . 922 

Crevette glabre . . . . . . . . . . . . . . . . . . . . 945 

Crevette glissante . . . . . . . . . . . . . . . . . . 910 

Crevette jaune . . . . . . . . . . . . . . . . . . . . 934 

Crevette javelot . . . . . . . . . . . . . . . . . . . . 913 

Crevette jinga . . . . . . . . . . . . . . . . . . . . 933 

Crevette kadal . . . . . . . . . . . . . . . . . . . . 936 

Crevette kuruma . . . . . . . . . . . . . . . . . . . 917 

Crevette moyebi . . . . . . . . . . . . . . . . . . . 912 

Crevette nylon mino . . . . . . . . . . . . . . . . . 970 

Crevette oiseau . . . . . . . . . . . . . . . . . . . 939 

Crevette orange . . . . . . . . . . . . . . . . . . . 929 

Crevette os . . . . . . . . . . . . . . . . . . . . . . 948 

Crevette papou . . . . . . . . . . . . . . . . . . . . 940 

Crevette périscope . . . . . . . . . . . . . . . . . . 929 

Crevette queue rouge . . . . . . . . . . . . . . . . 923 

Crevette queue verte . . . . . . . . . . . . . . . . . 934 

Crevette royale à taches rouges . . . . . . . . . . . 919 

Crevette royale blanche (des Indes) . . . . . . . . . 916 

Crevette royale occidentale . . . . . . . . . . . . . 918 

Crevette royale orientale . . . . . . . . . . . . . . . 924 

Crevette soricère . . . . . . . . . . . . . . . . . . . 914 

Crevette tigrée brune . . . . . . . . . . . . . . . . . 915 

Crevette tigrée verte . . . . . . . . . . . . . . . . . 925 

Crevette torpille . . . . . . . . . . . . . . . . . . . 944 

Crevette uncta . . . . . . . . . . . . . . . . . . . . 945 

Crevette violoneux . . . . . . . . . . . . . . . . . . 932 

Crevette york . . . . . . . . . . . . . . . . . . . . . 937 

Crinoids . . . . . . . . . . . . . . . . . . . 1158-1159 

cristata, Sicyonia . . . . . . . . . . . . . . . . . . 954 

Crocodile shark . . . . . . . . . . . . . . . . . . 1268 

Crocodile sharks . . . . . . . . . . . . . . . . . . 1268 

Crown crabs . . . . . . . . . . . . . . . . . . . . 1136 

cruciata, Charybdis . . . . . . . . . . . . . . . . 1120 

crucifer, Charybdis . . . . . . . . . . . . . . . . 1120 

Crucifix crab . . . . . . . . . . . . . . . . . . . . 1120 

Crumenasepia hulliana . . . . . . . . . . . . . . . 748 

Crumenasepia ursulae . . . . . . . . . . . . . . . . 748 

CRUSTACEA . . . . . . . . . . . . . . . . . . . . . 958 

cultrata, Sepia . . . . . . . . . . . . . . . . . . . . 759 

cultrifer, Raoulius . . . . . . . . . . . . . . . . . . 834 

cultrifer, Scyllarus . . . . . . . . . . . . . . . . . 1028 

cultrirostris, Parapenaeopsis . . . . . . . . . . . . 913 

Curryfish . . . . . . . . . . . . . . . . . . . . . . 1188 

Curvespine cuttlefish . . . . . . . . . . . . . . . . . 750 

curvirostris, Trachypenaeus . . . . . . . 927-928, 950 

Cuttlefish . . . . . . . . . . . . . . . . . 692, 694, 696 

Cuttlefishes . . . . . . . . 688-689, 696, 698, 723, 765


cuvier, Galeocerdo . . . . . . . . . . . . . 1313, 1349 

cyanea, Octopus . . . . . . . . . . . 696, 812, 817-818 

cyaneus, Octopus . . . . . . . . . . . . . . . . . . 818 

CYCLODORIPPIDAE . . . . . . . . . . . . 1083, 1085 

CYMONOMIDAE . . . . . . . . . . . . . . . 1083, 1085 

Cyrano spurdog . . . . . . . . . . . . . . . . . . 1231 

D 

dacqueti, Macrobrachium rosenbergii . . . . . . . 964 

Dalatias . . . . . . . . . . . . . . . . . . . . . . 1213 

Dalatias licha . . . . . . . . . . . . . . . . . . . 1225 

dalli, Metapenaeus . . . . . . . . . . . . . . 912, 935 

dama, Schizophrys . . . . . . . . . . . . . . . . 1137 

DASYATIDAE . . . . . . . . . . . . . . . . . . . . 1196 

dasypogon, Eucrossorhinus . . . . . . . . . . . 1247 

dasypus, Panulirus . . . . . . . . . . . . . . . . . 1016 

daumi, Enoplometopus . . . . . . . . . . . . . . . 999 

Day octopus . . . . . . . . . . . . . . . . . . . . . 812 

Deania profundorum . . . . . . . . . . . . . . . 1225 

Deania quadrispinosa . . . . . . . . . . . . . . . 1225 

debelius, Enoplometopus . . . . . . . . . . . 995, 999 

debelius, Lysmata . . . . . . . . . . . . . . . . . . 962 

DECAPODA . . . . . . . . . . . . . . . . . 958, 1048 

Decorisepia cottlesloensis . . . . . . . . . . . . . . 751 

Decorisepia jaenschi . . . . . . . . . . . . . . . . 751 

Decorisepia rex . . . . . . . . . . . . . . . . . . . 751 

Deep-water carrier crabs . . . . . . . . . . . 1056, 1083 

Deep-water hedgehog crabs . . . . . . . . . . . . 1083 

Deep-water mud shrimp . . . . . . . . . . . . . . . 885 

Deep-water porter crabs . . . . . . . . . . . 1050, 1083 

Deep-water redfish . . . . . . . . . . . . . . . . . 1167 

Deep-water sponge crabs . . . . . . . . . . . . . 1083 

Deep-water squat lobsters . . . . . . . . . . . . . 1048 

defillipi, Octopus . . . . . . . . . . . . . . . . . . . 801 

dehaani, Lauridromia . . . . . . . . . . . . . . . 1088 

demani, Metapenaeus . . . . . . . . . . . . . . . . 936 

Demania . . . . . . . . . . . . . . . . . . . . . . 1049 

Demon crabs . . . . . . . . . . . . . . . . . . . . 1049 

Demon shrimp . . . . . . . . . . . . . . . . . . . . 936 

DENDROCHIROTA . . . . . . . . . . . . . . . . . 1162 

DENDROCHIROTIDA . . . . . . . . . . . . 1159, 1163 

dentata, Ranina . . . . . . . . . . . . . . . . . . 1090 

dentatus, Epixanthus . . . . . . . . . . . . . . . 1108 

dentatus, Etisus . . . . . . . . . . . . . . . 1101-1102 

Diamondback squid . . . . . . . . . . . . . . . . . 797 

Diamondback squids . . . . . . . . . . . . . . . . . 797 

dierythraeus, Octopus . . . . . . . . . . . . . 801, 824 

DIOGENIDAE . . . . . . . . . . . . . . . . 1048, 1154 

diplana, Sphyrna . . . . . . . . . . . . . . . . . . 1364 

dobsoni, Metapenaeus . . . . . . . . . . . . . . . 936 

Dog shrimp . . . . . . . . . . . . . . . . . . . . . . 943 

Dogfish sharks . . . . . . . . . . . . . . . . 1197, 1213 

doldi, Neabrius . . . . . . . . . . . . . . . . . . 1260 

dolfeini, Histioteuthis . . . . . . . . . . . . . . . . 787 

dollfusi, Octopus . . . . . . . . . . . . . . . . . . . 811 

DORIPPIDAE . . . . . . . . . . . . . 1083, 1085, 1091 

dormia, Dromia . . . . . . . . . . . . . . . . . . 1087 

Dormilón acebrado . . . . . . . . . . . . . . . . . 1240 

Dormilón carenado . . . . . . . . . . . . . . . . . 1240 

Dormilón toro . . . . . . . . . . . . . . . . . . . . 1240 

dormitator, Cancer . . . . . . . . . . . . . . . . 1087 

Doryteuthis kensak . . . . . . . . . . . . . . . . . 776 

Doryteuthis sibogae . . . . . . . . . . . . . . 776-777 

Doryteuthis singhalensis . . . . . . . . . . . . . . 774 

Dosidicus gigas . . . . . . . . . . . . . . . . . . . 793 

Dotilla spp. . . . . . . . . . . . . . . . . . . . . 1048 

Double-ear bobtail squid . . . . . . . . . . . . . . . 714 

Dromia . . . . . . . . . . . . . . . . . . . 1085, 1087 

Dromia dormia . . . . . . . . . . . . . . . . . . 1087 

Dromia hirsutissima . . . . . . . . . . . . . . . . 1087 

Dromia orientalis . . . . . . . . . . . . . . . . . 1088 

Dromia rumphii . . . . . . . . . . . . . . . . . . 1087 

Dromidiopsis cranioides . . . . . . . . . . . . . 1088 

DROMIIDAE . . . . . . . . . . . . . 1056, 1083, 1085 

Dromiids . . . . . . . . . . . . . . . . . . . . . . 1085 

Drop-arm octopuses . . . . . . . . . . . . . . . . . 800 

durbanensis, Rhynchocinetes . . . . . . . . . . . 971 

Dusky shark . . . . . . . . . . . . . . . . . . . . 1344 

dussumieri, Carcharhinus . . . . . . . . . 1334, 1346 

duvauceli, Loligo . . . . . . . . . . . . . . . . . . 775 

duvaucelii, Photololigo . . . . . . . . . . . . 774-775 

Dwarf sawtail catshark . . . . . . . . . . . . . . . 1290 

Dwarf sharks . . . . . . . . . . . . . . . . . . . . 1196 

DYNOMENIDAE . . . . . . . . . . . . . . . . . . 1048 

E 

Easter Island spiny lobster . . . . . . . . . . . . . 1026 

Eastern angelshark . . . . . . . . . . . . . . . . 1237 

Eastern highfin spurdog . . . . . . . . . . . . . . 1231 

Eastern king prawn . . . . . . . . . . . . . . . . . . 924 

Eastern longnose spurdog . . . . . . . . . . . . . 1232 

Eastern school shrimp . . . . . . . . . . . . . . . . 939 

eastmani, Galeus . . . . . . . . . . . . . . . . . 1289 

eblanae, Todaropsis . . . . . . . . . . . . . . . . . 796 

eboracensis, Metapenaeus . . . . . . . . . . . . . 937 

echinites, Actinopyga . . . . . . . . . . . . 1167-1168 

ECHINODERMATA . . . . . . . . . . . . . . . . . 1158 

Echinoderms . . . . . . . . . . . . . . 1050, 1158-1159 

Echinoids . . . . . . . . . . . . . . . . . . . 1158-1159 

ECHINORHINIDAE . . . . . . . . . . . . . 1211, 1213 

Echinorhinus . . . . . . . . . . . . . . . . . . . . 1211 

Echinorhinus brucus . . . . . . . . . . . . . . . 1212 

Echinorhinus cookei . . . . . . . . . . . . . . . 1212 

eclogaria, Ponderisepia . . . . . . . . . . . . . . . 744 

eduardoi, Penaeopsis . . . . . . . . . . . . . 947-948 

edulis, Holothuria (Halodeima) . . . . . . . . . 1177 

edulis, Loligo . . . . . . . . . . . . . . . . . . . . 765 

edulis, Photololigo . . . . . . . . . 765-766, 774, 776 

edwardsiana, Aristaeopsis . . . . . . . . . . . . . . 874 

edwardsianus, Plesiopenaeus . . . . . . . . . . . 874 

Egg crabs . . . . . . . . . . . . . . . . . . . . . 1098 

ELASIPODA . . . . . . . . . . . . . . . . . 1159-1160 

Eledone palari . . . . . . . . . . . . . . . . . . . . 801 

elegans, Metapenaeus . . . . . . . . . . . . . 937, 940 

Elephant trunkfish . . . . . . . . . . . . . . . . . 1182 

elliot, Carcharhinus . . . . . . . . . . . . . . . . 1310 

elliptica, Sepia . . . . . . . . . . . . . . . . . 741-742 

elongatus, Hemipristis . . . . . . . . . . . 1308-1310 

Emerald shrimp . . . . . . . . . . . . . . . . . . . 938


Emissole cotiere . . . . . . . . . . . . . . . . . . 1304 

Emissole etoilée . . . . . . . . . . . . . . . . . . 1304 

Emissole gommée . . . . . . . . . . . . . . . . . 1304 

Emperor nautilus . . . . . . . . . . . . . . . . . . . 711 

Encornet bande violette . . . . . . . . . . . . . . . 794 

Encornet bouquet . . . . . . . . . . . . . . . . . . 792 

Encornet volant . . . . . . . . . . . . . . . . . . . . 793 

Endeavour shrimp . . . . . . . . . . . . . . . . . . 938 

endeavouri, Metapenaeus . . . . . . . . . 909-910, 938 

Enope squids . . . . . . . . . . . . . . . . . . . . . 781 

ENOPLOMETOPIDAE . 977-978, 983, 995, 1002, 1007 

Enoplometopus . . . . . . . . . . . . . . . . . . . 975 

Enoplometopus daumi . . . . . . . . . . . . . . . 999 

Enoplometopus debelius . . . . . . . . . . . 995, 999 

Enoplometopus holthuisi . . . . . . . . . . . . . 1000 

Enoplometopus occidentalis . . . . . . . . . . . 1000 

ENOPLOTEUTHIDAE . . . . . . . . . . . . . 781, 797 

ENOPLOTEUTHINAE . . . . . . . . . . . . . . . . 696 

ensifer, Heterocarpus . . . . . . . . . . . . . 969-970 

ensis baramensis, Metapenaeus . . . . . . . . . . . 910 

ensis, Metapenaeus . 910, 912, 933, 935, 937-938, 940 

Epaulette shark . . . . . . . . . . . . . . . . . . . 1258 

Episesarma . . . . . . . . . . . . . . 1138, 1143-1144 

Episesarma chengtongense . . . . . . . . . 1143, 1145 

Episesarma mederi . . . . . . . . . . . . . 1143, 1145 

Episesarma palawanense . . . . . . . . . . 1143, 1145 

Episesarma singaporense . . . . . . . . . . 1143, 1146 

Episesarma spp. . . . . . . . . . . . . . . . . . . 1143 

Episesarma versicolor . . . . . . . . . . . . . . . 1143 

Epixanthus dentatus . . . . . . . . . . . . . . . 1108 

Eptatretus cirrhatus . . . . . . . . . . . . . . . . 1192 

equidens, Macrobrachium . . . . . . . . . . . . . 967 

Eridacnis radcliffei . . . . . . . . . . . . . . . . 1295 

Erimacrus isenbeckii . . . . . . . . . . . . . . . 1055 

Eriocheir hepuensis . . . . . . . . . . . . . . . . 1055 

Eriocheir japonicus . . . . . . . . . . . . . . . . 1055 

Eriocheir sinensis . . . . . . . . . . . . . . . . . 1055 

Eriphia sebana . . . . . . . . . . . . . . . 1106, 1108 

Eriphia smithii . . . . . . . . . . . . . . . 1106, 1108 

Eriphia spp. . . . . . . . . . . . . . . . . . 1049, 1106 

Eriphiid stone . . . . . . . . . . . . . . . . . . . . 1103 

ERIPHIIDAE . . . 1048-1050, 1057, 1098, 1103, 1110, 

1112, 1114 

Eriphiids . . . . . . . . . . . . . . . . . . . 1112, 1114 

Erugosquilla . . . . . . . . . . . . . . . . . . . . . 842 

Erugosquilla woodmasoni . . . . . . . . . . 842, 846 

erythraeus, Acetes . . . . . . . . . . . . . . . . . . 862 

esculenta, Sepia . . . . . . . . . . . . . . 740-742, 748 

esculentus, Penaeus . . . . . . . . . . . . . . 915, 925 

etheridgei, Loligo . . . . . . . . . . . . . . . 765, 774 

Etisus . . . . . . . . . . . . . . . . . . . . . . . 1098 

Etisus convexus . . . . . . . . . . . . . . . . . . 1100 

Etisus dentatus . . . . . . . . . . . . . . . 1101-1102 

Etisus laevimanus . . . . . . . . . . . . . . . . . 1100 

Etisus macrodactylus . . . . . . . . . . . . . . . 1100 

Etisus maculatus . . . . . . . . . . . . . . . . . . 1100 

Etisus splendidus . . . . . . . . . . . . . . . . . 1101 

Etisus spp. . . . . . . . . . . . . . . . . . . . . . 1049 

Etisus utilis . . . . . . . . . . . . . . . . . . . . 1101 

Etmopterus . . . . . . . . . . . . . . . . . . . . . 1213 

Etmopterus brachyurus . . . . . . . . . . . . . . 1226 

Etmopterus lucifer . . . . . . . . . . . . . . . . 1226 

Etmopterus molleri . . . . . . . . . . . . . . . . 1226 

Etmopterus sp. C . . . . . . . . . . . . . . . . . 1227 

Etmopterus sp. D . . . . . . . . . . . . . . . . . 1227 

Etmopterus sp. F . . . . . . . . . . . . . . . . . 1227 

Etmopterus splendidus . . . . . . . . . . . . . . 1227 

Eucrossorhinus dasypogon . . . . . . . . . . . . 1247 

Eugomphodus taurus . . . . . . . . . . . . . . . 1266 

Eulamia temmincki . . . . . . . . . . . . . . . . 1350 

EUMEDONIDAE . . . . . . . . . . . . . . . . . . 1050 

Euprotomicrus . . . . . . . . . . . . . . . . . . . 1213 

Euprotomicrus bispinatus . . . . . . . . . . . . 1228 

Eupryma morsei . . . . . . . . . . . . . . . . . . . 714 

Eupryma tasmanica . . . . . . . . . . . . . . . . . 714 

Eusphyra blochii . . . . . . . . . . . . . . . . . 1363 

exannulatus, Octopus . . . . . . . . . . . . . . . . 825 

Exopalaemon styliferus . . . . . . . . . . . . . . . 965 

Exopalaemon vietnamicus . . . . . . . . . . . . . 965 

expolitum, Cirrhoscyllium . . . . . . . . . . . . 1242 

exsulcatus, Penaeus semisulcatus . . . . . . . . . . 922 

Eye-bar ocellate octopus . . . . . . . . . . . . . . . 819 

F 

falciformis, Carcharhinus . . . . . . . . . 1335, 1341 

False catshark . . . . . . . . . . . . . . . . . . . 1296 

False catsharks . . . . . . . . . . . . . . . . . . 1296 

False comb shrimp . . . . . . . . . . . . . . . . . . 888 

False crabs . . . . . . . . . . . . . . . . . . . . . 1048 

False rose shrimp . . . . . . . . . . . . . . . . . . 946 

False smalltail shark . . . . . . . . . . . . . . . . 1359 

False white prawn . . . . . . . . . . . . . . . . . . 926 

fasciata, Anchisquilla . . . . . . . . . . . . . . . . 842 

fasciata, Hapalochlaena . . . . . . . . . . . . . . 820 

fasciatum, Cephaloscyllium . . . . . . . . . . . 1289 

fasciatum, Stegostoma . . . . . . . . . . . . . . 1262 

fasciatus, Atelomycterus . . . . . . . . . . . . . 1287 

fasciatus, Panulirus . . . . . . . . . . . . . 1020-1021 

Fe’e motot . . . . . . . . . . . . . . . . . . . . . . 825 

Feather stars . . . . . . . . . . . . . . . . . . . . 1158 

femoristriga, Panulirus longipes . . . . . . 1015, 1017 

Fenix lobsters . . . . . . . . . . . . . . . . . . . . 978 

feriatus, Charybdis . . . . . . . . . . 1115, 1120-1121 

ferox, Alepisaurus . . . . . . . . . . . . . . . . . . 799 

ferox, Odontaspis . . . . . . . . . . . . . . . . . 1267 

ferox, Odontaspix . . . . . . . . . . . . . . . . . 1266 

ferrugineum, Ginglymostoma . . . . . . . . . . . 1260 

ferrugineus, Nebrius . . . . . . . . . . . . . . . 1260 

Fiddler crabs . . . . . . . . . . . . . . . . . . . . 1048 

Fiddler shrimp . . . . . . . . . . . . . . . . . . . . 932 

filewoodi, Gogolia . . . . . . . . . . . . . . . . . 1302 

Finback catsharks . . . . . . . . . . . . . . . . . 1293 

Fine shrimp . . . . . . . . . . . . . . . . . . . . . . 937 

Finned cirrate octopuses . . . . . . . . . . . . . . . 801 

Firefly squids . . . . . . . . . . . . . . . . . . . . . 781 

fissuroides, Parapenaeus . . . . . . . . . . . . . . 946 

fissurus, Parapenaeus . . . . . . . . . . . . . . . . 946 

fittoni, Podopilumnus . . . . . . . . . . . . . . . 1113 

fitzroyensis, Carcharhinus . . . . . . . . . . . . 1336


Fivespined squillid mantis shrimp . . . . . . . . . . 849 

Flamingo shrimp . . . . . . . . . . . . . . . . . . . 947 

Flathead lobster . . . . . . . . . . . . . . . . . . 1040 

floridus, Atergatis . . . . . . . . . . . . . . 1049, 1098 

Flower crab . . . . . . . . . . . . . . . . . . . . . 1124 

Flower moon crab . . . . . . . . . . . . . . . . . 1094 

Flying squids . . . . . . . . . . . . . . . . . . . . . 788 

foliacea, Aristaeomorpha . . . . . . . . . . . . . . 872 

formosana, Loligo . . . . . . . . . . . . . . . . . . 774 

formosana, Sepia . . . . . . . . . . . . . . . . . . 748 

formosus, Atypopenaeus . . . . . . . . . . . . . . 929 

fornasinii, Myomenippe . . . . . . . . . . 1107, 1109 

Four-spined needle shrimp . . . . . . . . . . . . . . 947 

freycineti, Hemiscyllium . . . . . . . . . . . . . 1258 

fulvus, Trachypenaeus . . . . . . . . . . . . . . . . 928 

Funchalia . . . . . . . . . . . . . . . . . . . 889-890 

Furry lobsters . . . . . . . . . . . . . . . . . 979, 1001 

fuscogilva, Holothuria . . . . . . . . . . . . . . . 1183 

fuscogilva, Holothuria (Microthele) . . . . . . . 1181 

fuscopunctata, Holothuria (Microthele) . . . . 1182 

G 

galapagensis, Carcharhinus . . . . . . . . 1326, 1337 

Galapagos shark . . . . . . . . . . . . . . . . . . 1337 

GALATHEIDAE . . . . . . . . . . . . . . . . . . 1048 

Galatheids . . . . . . . . . . . . . . . . . . . . . 1048 

GALATHEOIDEA . . . . . . . . . . . . . . . . . . . 976 

galeatus, Heterodontus . . . . . . . . . . . . . . 1240 

galei, Sepia . . . . . . . . . . . . . . . . . . . . . . 747 

Galene bispinosa . . . . . . . . . . . . . . 1112-1113 

Galene granulosa . . . . . . . . . . . . . . . . . 1113 

Galeocerdo . . . . . . . . . . . . . . . . . . . . . 1312 

Galeocerdo arcticus . . . . . . . . . . . . . . . . 1349 

Galeocerdo cuvier . . . . . . . . . . . . . . 1313, 1349 

Galeocerdo rayneri . . . . . . . . . . . . . . . . 1349 

Galeorhinus . . . . . . . . . . . . . . . . . 1297, 1313 

Galeorhinus galeus . . . . . . . . . . . . . . . . 1302 

Galeus boardmani . . . . . . . . . . . . . . . . . 1289 

Galeus eastmani . . . . . . . . . . . . . . . . . . 1289 

Galeus gracilis . . . . . . . . . . . . . . . . . . . 1290 

Galeus sauteri . . . . . . . . . . . . . . . . . . . 1290 

Galeus schultzi . . . . . . . . . . . . . . . . . . 1290 

galeus, Galeorhinus . . . . . . . . . . . . . . . . 1302 

Galludo cirann . . . . . . . . . . . . . . . . . . . 1231 

Galludo cola negra . . . . . . . . . . . . . . . . . 1230 

Galludo espinilla . . . . . . . . . . . . . . . . . . 1230 

Galludo japones . . . . . . . . . . . . . . . . . . 1229 

Galludo ñato . . . . . . . . . . . . . . . . . . . . 1230 

Gamba carabinero . . . . . . . . . . . . . . . . . . 874 

Gamba española . . . . . . . . . . . . . . . . . . . 872 

Gambón colorado . . . . . . . . . . . . . . . . . . 973 

Gambon écarlat . . . . . . . . . . . . . . . . . . . 874 

Gambon gaillard . . . . . . . . . . . . . . . . . . . 873 

Gambon rouge . . . . . . . . . . . . . . . . . . . . 872 

gangeticus, Glyphis . . . . . . . . . . . . . . . . 1329 

garmani, Scyliorhinus . . . . . . . . . . . . . . 1292 

garricki, Iago . . . . . . . . . . . . . . . . . . . 1303 

Gata nodriza atezada . . . . . . . . . . . . . . . 1260 

GECARCINIDAE . . . . . . . . 1049, 1060, 1147, 1152 

Gecarcinids . . . . . . . . . . . . . . . . . . . . . 1147 

GECARCINUCIDAE . . . . . . . . . . . . . 1050, 1147 

Gecarcinus trispinosus . . . . . . . . . . . . . . 1113 

Gecarcoidea lalandii . . . . . . . . . . . . . . . 1150 

Gecarcoidea natalis . . . . . . . . . . . . . . . . 1150 

Gecko catshark . . . . . . . . . . . . . . . . . . . 1289 

genista, Solitosepia . . . . . . . . . . . . . . . . . 747 

GERYONIDAE . . . . . . . . . . . . . . . . 1059, 1132 

Geryons . . . . . . . . . . . . . . . . . . . . . . 1132 

Ghost crabs . . . . . . . . . . . . . . . . . 1060, 1152 

Ghost shrimps . . . . . . . . . . . . . . . . . . . . 976 

Giant Australian cuttlefish . . . . . . . . . . . . . . 738 

Giant box crab . . . . . . . . . . . . . . . . . . . 1096 

Giant egg crab . . . . . . . . . . . . . . . . . . . 1102 

Giant harpiosquillid mantis shrimp . . . . . . . . . . 841 

Giant mud crab . . . . . . . . . . . . . . . . . . . 1126 

Giant red shrimp . . . . . . . . . . . . . . . . . . . 872 

Giant river prawn . . . . . . . . . . . . . . . . . . . 964 

Giant Tasmanian crab . . . . . . . . . . . . . . . 1055 

Giant tiger prawn . . . . . . . . . . . . . . . . . . . 922 

gigas, Dosidicus . . . . . . . . . . . . . . . . . . . 793 

gigas, Pseudocarcinus . . . . . . . . . . . . . . . 1055 

Ginglymostoma ferrugineum . . . . . . . . . . . 1260 

GINGLYMOSTOMATIDAE 1197, 1243, 1250, 1260, 1313 

glauca, Prionace . . . . . . . . . . . . . . . . . 1353 

glaucus, Isurus . . . . . . . . . . . . . . . . . . . 1277 

Globigerina . . . . . . . . . . . . . . . . . . . . . 992 

GLYPHEIDAE . . . . . 977-978, 983, 996, 1002, 1007 

Glyphis . . . . . . . . . . . . . . . . . . . . . . . 1196 

Glyphis gangeticus . . . . . . . . . . . . . . . . 1329 

Glyphis glyphis . . . . . . . . . . . . . . . . . . 1359 

Glyphis sp. A . . . . . . . . . . . . . . . . 1359-1360 

Glyphis sp. B . . . . . . . . . . . . . . . . . . . 1360 

Glyphis sp. C . . . . . . . . . . . . . . . . . . . 1360 

Glyphis spp. . . . . . . . . . . . . . . . . . . . . 1329 

glyphis, Glyphis . . . . . . . . . . . . . . . . . . 1359 

Glyptosepia opipara . . . . . . . . . . . . . . . . . 746 

Gnathia . . . . . . . . . . . . . . . . . . . . . . 1257 

GNATHOPHYLLIDAE . . . . . . . . . . . . . . . . 958 

Go home prawn . . . . . . . . . . . . . . . . . . . 929 

godeffroyi, Stichopus . . . . . . . . . . . . . . . 1187 

Gogolia filewoodi . . . . . . . . . . . . . . . . . 1302 

Golden crabs . . . . . . . . . . . . . . . . . . . . 1059 

Golden cuttlefish . . . . . . . . . . . . . . . . . . . 742 

Golden sandfish . . . . . . . . . . . . . . . . . . 1180 

Gollum . . . . . . . . . . . . . . . . . . . . . . . 1293 

Gollum attenuatus . . . . . . . . . . . . . . . . 1295 

GONEPLACIDAE . . . . . . . 1058, 1061, 1112, 1114 

Goneplacids . . . . . . . . . . . . . . . . . . . . 1112 

GONODACTYLIDAE . . . . . . . . . . . . . . . . . 832 

gonospinifer, Trachypenaeus . . . . . . . . . . . . 949 

gouldi, Nototodarus . . . . . . . . . . . . . . . . . 792 

Graceful catshark . . . . . . . . . . . . . . . . . 1295 

Graceful shark . . . . . . . . . . . . . . . . . . . 1327 

gracilis, Galeus . . . . . . . . . . . . . . . . . . 1290 

gracillima, Parapenaeopsis . . . . . . . . . . . . . 943 

graeffei, Bohadschia . . . . . . . . . . . . . . . . 1184 

graeffei, Pearsonothuria . . . . . . . . . . . . . 1184 

Grand requin blanc . . . . . . . . . . . . . . . . . 1276 

Grand requin-marteau . . . . . . . . . . . . . . . 1365


granulatus, Chaceon . . . . . . . . . . . . 1132, 1134 

granulosa, Galene . . . . . . . . . . . . . . . . . 1113 

granulosa, Menippe . . . . . . . . . . . . . . . . 1107 

granulosa, Myomenippe . . . . . . . . . . . . . . 1107 

granulosa, Platypodia . . . . . . . . . . . . . . . 1049 

granulosus, Centrophorus . . . . . . . . . 1223-1224 

granulosus, Trachypenaeus . . . . . . . . . . . . 949 

GRAPSIDAE . . . . . . 1048, 1060, 1138, 1147, 1152 

Grapsids . . . . . . . . . . . . . . . . . . . . . . 1138 

Grapsus albolineatus . . . . . . . . . . . . . . . 1142 

Grapsus strigosus . . . . . . . . . . . . . . . . . 1142 

Grapsus tenuicrustatus . . . . . . . . . . . . . . 1142 

graptus, Octopus . . . . . . . . . . . . . . . 801, 813 

gravieri, Oratosquillina . . . . . . . . . . . . 843, 848 

Gray nurse shark . . . . . . . . . . . . . . . . . . 1266 

Greasyback shrimp . . . . . . . . . . . . . . . . . . 910 

Great hammerhead . . . . . . . . . . . . . . . . 1365 

Great white shark . . . . . . . . . . . . . . . . . 1276 

Greater blue-ringed octopus . . . . . . . . . . . . . 821 

Greater drop-arm octopus . . . . . . . . . . . . . . 822 

Green egg crab . . . . . . . . . . . . . . . . . . . 1049 

Green mud crab . . . . . . . . . . . . . . . . . . 1128 

Green tiger prawn . . . . . . . . . . . . . . . . . . 925 

Greenfish . . . . . . . . . . . . . . . . . . . . . . 1186 

Greentail shrimp . . . . . . . . . . . . . . . . . . . 934 

Grey bambooshark . . . . . . . . . . . . . . . . . 1253 

Grey reef shark . . . . . . . . . . . . . . . . . . . 1328 

Grey sharpnose shark . . . . . . . . . . . . . . . 1355 

griseum, Chiloscyllium . . . . 1253-1254, 1256-1257 

griseus, Hexanchus . . . . . . . . . . . . . . . . 1210 

griseus, Mustelus . . . . . . . . . . . . . . . . . 1304 

Grooved cuttlefish . . . . . . . . . . . . . . . . . . 761 

Ground sharks . . . . . . . . . . . . . . . . . . . 1312 

guamensis, Holothuria . . . . . . . . . . . . . . 1183 

guerini, Calappa . . . . . . . . . . . . . . . . . . 1094 

Gulper shark . . . . . . . . . . . . . . . . . . . . 1223 

Gummy shark . . . . . . . . . . . . . . . . . . . 1304 

guttatus, Ozius . . . . . . . . . . . . . . . . . . . 1109 

H 

haanii, Scyllarides . . . . . . . . . . . . . . . . . 1038 

habereri, Proscyllium . . . . . . . . . . . . 1293, 1295 

Hagfishes . . . . . . . . . . . . . . . . . . . . . . 1192 

Hai-sum . . . . . . . . . . . . . . . . . . . . . . 1163 

Hairy blackfish . . . . . . . . . . . . . . . . . . . 1169 

Hairy crabs . . . . . . . . . . . . . . 1055, 1058, 1112 

Hairy fan lobster . . . . . . . . . . . . . . . . . . 1036 

Halaelurus boesemani . . . . . . . . . . . . . . 1291 

Haliphron atlanticus . . . . . . . . . . . . . . . . . 802 

Haliporoides sibogae . . . . . . . . . . . . . . . . 881 

Haliporoides sibogae australiensis . . . . . . . . . 881 

Haliporoides sibogae madagascariensis . . . . . . 881 

halli, Solenocera . . . . . . . . . . . . . . . 884, 886 

hallstromi, Hemiscyllium . . . . . . . . . . . . . 1258 

Hammer octopus . . . . . . . . . . . . . . . . . . . 824 

Hammerhead sharks . . . . . . . . . 1196-1197, 1361 

Hapalochlaena cf. maculosa . . . . . . . . . . . . 821 

Hapalochlaena fasciata . . . . . . . . . . . . . . . 820 

Hapalochlaena lunulata . . . . . . . . . . . . . . 821 

Hardback shrimp . . . . . . . . . . . . . . . . . . . 948 

Hardnose shark . . . . . . . . . . . . . . . . . . 1342 

hardwickei, Octopus . . . . . . . . . . . . . . . . . 811 

hardwickii, Myomenippe . . . . . . . . . . 1103, 1107 

hardwickii, Parapenaeopsis . . . . . . . . . . . . 913 

harpax, Harpiosquilla . . . . . . . . . . . . . . . 841 

Harpiosquilla . . . . . . . . . . . . . . . . . . . . 830 

Harpiosquilla harpax . . . . . . . . . . . . . . . . 841 

Harpiosquilla raphidea . . . . . . . . . . . . . . . 841 

Harpiosquillid mantis shrimps . . . . . . 830, 834, 838 

HARPIOSQUILLIDAE . . . . . . . . . . 830, 838, 842 

Harpodon nehereus . . . . . . . . . . . . . . . . 1357 

Hasselt’s bambooshark . . . . . . . . . . . . . . 1254 

hasselti, Chiloscyllium . . . . 1253-1254, 1256-1257 

hathor, Penaeus latisulcatus . . . . . . . . . . . . 918 

Hawaiian flying squid . . . . . . . . . . . . . . . . . 792 

hawaiiensis, Latreillopsis . . . . . . . . . . . . . 1084 

hawaiiensis, Nototodarus . . . . . . 788, 792, 795-796 

hayashii, Heterocarpus . . . . . . . . . . . . 969-970 

hedleyi, Sepia . . . . . . . . . . . . . . . . . . . . 751 

HEMIGALEIDAE . 1196-1197, 1264, 1297, 1305, 1313 

Hemigaleus balfouri . . . . . . . . . . . . . . . . 1308 

Hemigaleus macrostoma . . . . . . . . . . . . . 1308 

Hemigaleus microstoma . . . . . . . 1308-1309-1310 

hemiodon, Carcharhinus . . . . . . . . . . . . . 1338 

hemiodon, Hypoprion . . . . . . . . . . . . . . . 1338 

Hemipristis elongatus . . . . . . . . . . . . 1308-1310 

HEMISCYLLIIDAE . . . . 1197, 1243, 1249-1250, 1261 

Hemiscyllium freycineti . . . . . . . . . . . . . . 1258 

Hemiscyllium hallstromi . . . . . . . . . . . . . 1258 

Hemiscyllium ocellatum . . . . . . . . . . . . . 1258 

Hemiscyllium sp. . . . . . . . . . . . . . . . . . . 1249 

Hemiscyllium strahani . . . . . . . . . . . . . . 1259 

Hemiscyllium trispeculare . . . . . . . . . . . . 1259 

Hemitriakis leucoperiptera . . . . . . . . . . . . 1302 

hepatica, Calappa . . . . . . . . . . . . . . 1094, 1097 

Heptranchias perlo . . . . . . . . . . . . . . . . 1210 

hepuensis, Eriocheir . . . . . . . . . . . . . . . . 1055 

herbsti, Odontaspis . . . . . . . . . . . . . . . . 1267 

hercules, Sepia . . . . . . . . . . . . . . . . . . . . 744 

herklotsi, Apristurus . . . . . . . . . . . . . . . 1286 

Hermit crabs . . . . . . . . . . . . . . 1048, 1154-1155 

Heterocarpus . . . . . . . . . . . . . . . . . . . . 958 

Heterocarpus ensifer . . . . . . . . . . . . . 969-970 

Heterocarpus hayashii . . . . . . . . . . . . 969-970 

Heterocarpus parvispina . . . . . . . . . . . 969-970 

Heterocarpus sibogae . . . . . . . . . . . . . 969-970 

HETERODONTIDAE . . . . . . . . . . . . . . . . 1238 

Heterodontus galeatus . . . . . . . . . . . . . . 1240 

Heterodontus portusjacksoni . . . . . . . . . . . 1240 

Heterodontus zebra . . . . . . . . . . . . . . . . 1240 

Heteropenaeus . . . . . . . . . . . . . . . . . 889-890 

Heteroscyllium colcloughi . . . . . . . . . . . . 1244 

Heteroteuthis weberi . . . . . . . . . . . . . . . . 715 

HEXANCHIDAE . . . . . . . . . . . . . . . 1208, 1314 

Hexanchus griseus . . . . . . . . . . . . . . . . 1210 

Hexanchus nakamurai . . . . . . . . . . . . . . 1210 

HEXAPODIDAE . . . . . . . . . . . . . . . . . . 1048 

High ridge mud shrimp . . . . . . . . . . . . . . . . 885


HIPPOLYTIDAE . . . . . . . . . . . . . . . . 958, 961 

hirsutissima, Dromia . . . . . . . . . . . . . . . 1087 

hirtipes, Cardisoma . . . . . . . . . . . . . 1149, 1151 

hispidus, Stenopus . . . . . . . . . . . . . . . . . . 955 

HISTIOTEUTHIDAE . . . . . . . . . . . . . . . . . 787 

Histioteuthids . . . . . . . . . . . . . . . . . . . . . 787 

Histioteuthis . . . . . . . . . . . . . . . . . . . . . 787 

Histioteuthis celetaria pacifica . . . . . . . . . . . 787 

Histioteuthis dofleini . . . . . . . . . . . . . . . . 787 

Histioteuthis miranda . . . . . . . . . . . . . . . . 787 

Holbiche à grande tête . . . . . . . . . . . . . . . 1286 

Holbiche à joues noires . . . . . . . . . . . . . . 1291 

Holbiche à longues nageoires . . . . . . . . . . . 1286 

Holbiche bouffie . . . . . . . . . . . . . . . . . . 1289 

Holbiche malaise . . . . . . . . . . . . . . . . . . 1287 

Holbiche mouchetée . . . . . . . . . . . . . . . . 1291 

Holbiche pâle . . . . . . . . . . . . . . . . . . . . 1286 

Holbiche tête molle . . . . . . . . . . . . . . . . . 1287 

Holbiche voile . . . . . . . . . . . . . . . . . . . 1291 

Holothuria . . . . . . . . . . . . . . . . . . 1161, 1178 

Holothuria (Acanthotrapeza) coluber . . . . . . 1175 

Holothuria (Halodeima) atra . . . . . . . . . . . 1176 

Holothuria (Halodeima) edulis . . . . . . . . . . 1177 

Holothuria (Mertensiothuria) leucospilota . . . 1178 

Holothuria (Metriatyla) scabra . . . . . . . 1179-1180 

Holothuria (Metriatyla) scabra var. versicolor 1179-1180 

Holothuria (Microthele) fuscogilva . . . . . . . 1181 

Holothuria (Microthele) fuscopunctata . . . . . 1182 

Holothuria (Microthele) nobilis . . . . . . . . . 1183 

Holothuria acculeata . . . . . . . . . . . . . . . 1180 

Holothuria albiventer . . . . . . . . . . . . . . . 1180 

Holothuria atra . . . . . . . . . . . . . . . 1177-1178 

Holothuria axiologa . . . . . . . . . . . . . . . . 1182 

Holothuria coluber . . . . . . . . . . . . . . . . 1178 

Holothuria fuscogilva . . . . . . . . . . . . . . . 1183 

Holothuria guamensis . . . . . . . . . . . . . . . 1183 

Holothuria leucospilota . . . . . . . . . . . . . . 1176 

Holothuria maculata . . . . . . . . . . . . . . . 1181 

Holothuria nobilis . . . . . . . . . . . . . . . . . 1181 

Holothuria scabra . . . . . . . . . . 1173-1174, 1180 

Holothuria versicolor . . . . . . . . . . . . . . . 1180 

Holothurian . . . . . . . . . . . . . . . . . . . . . 1159 

Holothurians . . . . . . . . . . 1158-1159, 1162-1163 

Holothurie ananas . . . . . . . . . . . . . . . . . 1189 

Holothurie blanche à mamelles . . . . . . . . . . 1181 

Holothurie brune . . . . . . . . . . . . . . . . . . 1174 

Holothurie brune des brisants . . . . . . . . . . . 1168 

Holothurie léopard . . . . . . . . . . . . . . . . . 1172 

Holothurie noire à mamelles . . . . . . . . . . . . 1183 

Holothurie serpent . . . . . . . . . . . . . . . . . 1175 

Holothurie trompe d’éléphant . . . . . . . . . . . 1182 

HOLOTHURIIDAE . . . . . . . . . . . . . . 1162, 1165 

Holothuroids . . . . . . . . . . . . . . . . . 1158-1159 

holthuisi, Enoplometopus . . . . . . . . . . . . 1000 

holthuisi, Parribacus . . . . . . . . . . . . . . . 1042 

homarus, Panulirus . . . . . . . . . . . . . . . . 1016 

HOMOLIDAE . . . . . . . . . . . . . 1056, 1083, 1089 

Homolids . . . . . . . . . . . . . . . . . . . . . . 1050 

HOMOLODROMIIDAE . . . . . . . . . . . . 1083, 1085 

Homolodromiids . . . . . . . . . . . . . . . . . . 1085 

Hooded carpetshark . . . . . . . . . . . . . . . . 1259 

Hooked squids . . . . . . . . . . . . . . . . . . . . 784 

Hooktooth shark . . . . . . . . . . . . . . . . . . 1308 

Horn sharks . . . . . . . . . . . . . . . . . . . . 1238 

Horned ghost crab . . . . . . . . . . . . . . . . . 1153 

horrens, Stichopus . . . . . . . . . . . . . . . . 1187 

horridus, Octopus . . . . . . . . . . . . . . . 800-801 

Horse crab . . . . . . . . . . . . . . . . . . . . . 1125 

Houndsharks . . . . . . . . . . . . . . . . . 1196, 1297 

hulliana, Crumenasepia . . . . . . . . . . . . . . . 748 

humei, Hylaeocarcinus . . . . . . . . . . . . . . 1150 

Humpback shrimp . . . . . . . . . . . . . . . . . . 930 

Hunchback locust lobster . . . . . . . . . . . . . 1043 

hungerfordi, Parapenaeopsis . . . . . . . . . . . . 943 

Hylaeocarcinus humei . . . . . . . . . . . . . . . 1150 

Hymenocera picta . . . . . . . . . . . . . . . . . . 963 

HYMENOCERIDAE . . . . . . . . . . . . . . 958, 963 

Hymenopenaeus sibogae . . . . . . . . . . . . . . 881 

HYMENOSOMATIDAE . . . . . . . . . . . . . . 1136 

Hypogaleus . . . . . . . . . . . . . . . . . . . . 1313 

Hypogaleus hyugaensis . . . . . . . . . . . . . . 1303 

Hypoprion hemiodon . . . . . . . . . . . . . . . 1338 

Hypoprion macloti . . . . . . . . . . . . . . . . . 1342 

Hypoprion playfairi . . . . . . . . . . . . . . . . 1343 

Hypothalassia armata . . . . . . . . . . . . 1103, 1106 

hyugaensis, Hypogaleus . . . . . . . . . . . . . 1303 

I 

Iago garricki . . . . . . . . . . . . . . . . . . . . 1303 

Ibacus . . . . . . . . . . . . . . . . . 977, 1028, 1040 

Ibacus ciliatus . . . . . . . . . . . . . . . . 1034-1036 

Ibacus ciliatus pubescens . . . . . . . . . . . . . 1036 

Ibacus novemdentatus . . . . . . . . . . . . . . 1035 

Ibacus peronii . . . . . . . . . . . . . . . . . . . 1035 

Ibacus pubescens . . . . . . . . . . . . . . 1035-1036 

IDIOSEPIIDAE . . . . . . . . . . . . . . . . . . . . 721 

Idiosepius . . . . . . . . . . . . . . . . . . . . . . 721 

Idiosepius pygmaeus . . . . . . . . . . . . . . . . . 721 

imperator, Chiroteuthis . . . . . . . . . . . . . . . 798 

INCIRRATA . . . . . . . . . . . . . . . . . 689, 801-802 

Incirrate octopuses . . . . . . . . . . . . . . . 689, 801 

incisipes, Metapenaeus . . . . . . . . . . . . . . . 910 

Indian Ocean lobsterette . . . . . . . . . . . . . . . 991 

Indian small prawn . . . . . . . . . . . . . . . . . . 966 

Indian squid . . . . . . . . . . . . . . . . . . . . . 775 

Indian white prawn . . . . . . . . . . . . . . . . . . 916 

indica, Lauridromia . . . . . . . . . . . . . . . . 1088 

indica, Loligo . . . . . . . . . . . . . . . . . 774-775 

indica, Sepia . . . . . . . . . . . . . . . . . . . . . 736 

indicum, Chiloscyllium . . . . . . . . . . . 1255-1256 

indicus, Cistopus . . . . . . . . . . . . . . . . . . 810 

indicus, Leandrites . . . . . . . . . . . . . . . . . 966 

indicus, Penaeus . . . . . . . . . . 916, 921, 923, 926 

indicus, Solenocera . . . . . . . . . . . . . . . . . 883 

Indo-Pacific furry lobster . . . . . . . . . . . . . . 1004 

Indonesian golden crab . . . . . . . . . . . . . . 1135 

Indonesian speckled carpetshark . . . . . . . . . 1258 

inermis, Sepiella . . . . . . . . . . . . . . . . . . . 756 

Inshore squids . . . . . . . . . . . . . . . . . . . . 764 

insolitus, Metapenaeus . . . . . . . . . . . . . . . 938


integerrimus, Atergatis . . . . . . . . . . . . . . 1049 

intermedius, Acetes . . . . . . . . . . . . . . . . . 863 

intermedius, Limnocarcinus . . . . . . . . . . . . 1150 

intermedius, Metapenaeus . . . . . . . . . . 909, 911 

interrupta, Oratosquillina . . . . . . . . . . . . . . 843 

iranzae, Carcharhinus . . . . . . . . . . . . . . . 1344 

isenbeckii, Erimacrus . . . . . . . . . . . . . . . 1055 

Isistius . . . . . . . . . . . . . . . . . . . . . . . 1213 

Isistius brasiliensis . . . . . . . . . . . . . 1213, 1228 

ISOLAPOTAMIDAE . . . . . . . . . . . . . . . . 1050 

Isurus . . . . . . . . . . . . . . . . . . . . . . . . 1197 

Isurus alatus . . . . . . . . . . . . . . . . . . . . 1278 

Isurus glaucus . . . . . . . . . . . . . . . . . . . 1277 

Isurus oxyrinchus . . . . . . . . . . . . . . 1277-1278 

Isurus paucus . . . . . . . . . . . . . . . . 1277-1278 

J 

Jack-knife shrimp . . . . . . . . . . . . . . . . . . . 881 

jaenschi, Decorisepia . . . . . . . . . . . . . . . . 751 

Japanese angelshark . . . . . . . . . . . . . . . 1237 

Japanese blunthorn lobster . . . . . . . . . . . . 1025 

Japanese bobtail . . . . . . . . . . . . . . . . . . . 718 

Japanese deepwater carrier crab . . . . . . . . . 1084 

Japanese fan lobster . . . . . . . . . . . . . . . . 1034 

Japanese flying squid . . . . . . . . . . . . . . . . 795 

Japanese furrow lobster . . . . . . . . . . . . . . 1022 

Japanese golden crab . . . . . . . . . . . . . . . 1134 

Japanese mitten crab . . . . . . . . . . . . . . . 1055 

Japanese nylon shrimp . . . . . . . . . . . . . . . . 969 

Japanese spear lobster . . . . . . . . . . . . . . 1024 

Japanese sponge crab . . . . . . . . . . . . . . . 1088 

Japanese spurdog . . . . . . . . . . . . . . . . . 1229 

Japanese squillid mantis shrimp . . . . . . . . . . . 847 

Japanese swimming crab . . . . . . . . . . . . . 1121 

Japanese wobbegong . . . . . . . . . . . . . . . 1247 

japonica, Charybdis . . . . . . . . . . . . . . . . 1121 

japonica, Justitia . . . . . . . . . . . . . . . . . 1022 

japonica, Paromola . . . . . . . . . . . . . 1083-1084 

japonica, Squatina . . . . . . . . . . . . . . . . 1237 

japonicus, Penaeus . . . . . . . . . . . . . . 914, 917 

japonicus, Acetes . . . . . . . . . . . . . . . . . . 863 

japonicus, Carcinoplax longimanus . . . . . . . 1114 

japonicus, Chionoecetes . . . . . . . . . . . . . . 1055 

japonicus, Eriocheir . . . . . . . . . . . . . . . . 1055 

japonicus, Orectolobus . . . . . . . . . . . . . . 1247 

japonicus, Panulirus . . . . . . . . . . . . . . . . 1017 

japonicus, Penaeus . . . . . . . . . . . . . . 914, 917 

japonicus, Pristiophorus . . . . . . . . . . . . . 1234 

japonicus, Squalus . . . . . . . . . . . . . . . . 1229 

Jaquentón blanco . . . . . . . . . . . . . . . . . 1276 

Jaquetón . . . . . . . . . . . . . . . . . . . . . . 1276 

Jawla paste shrimp . . . . . . . . . . . . . . . . . . 862 

jejunus, Penaeus . . . . . . . . . . . . . . . . . . . 919 

Jembret shrimp . . . . . . . . . . . . . . . . . . . . 865 

Jewel crab . . . . . . . . . . . . . . . . . . . . . 1049 

Jewel squids . . . . . . . . . . . . . . . . . . . . . 787 

Jinga shrimp . . . . . . . . . . . . . . . . . . . . . 933 

johnsoni, Carcharhinus . . . . . . . . . . . . . . 1332 

Justitia . . 975, 979, 984, 997, 1002, 1005-1006, 1010 

Justitia chani . . . . . . . . . . . . . . . . . . . 1022 

Justitia japonica . . . . . . . . . . . . . . . . . . 1022 

Justitia longimanus . . . . . . . . . . . . . 1005, 1023 

Justitia mauritiana . . . . . . . . . . . . . . . . . 1023 

Justitia vericeli . . . . . . . . . . . . . . . . . . 1023 

K 

Kadal shrimp . . . . . . . . . . . . . . . . . . . . . 936 

kagoshimensis, Octopus . . . . . . . . . . . . . . . 815 

kamoharai, Odontaspis . . . . . . . . . . . . . . 1268 

kamoharai, Pseudocarcharias . . . . . . . . . . 1268 

kanakorum, Aulohalaelurus . . . . . . . . . . . 1288 

karubar, Chaceon . . . . . . . . . . . 1132, 1134-1135 

kensak, Doryteuthis . . . . . . . . . . . . . . . . . 776 

King cuttlefish . . . . . . . . . . . . . . . . . . . . 751 

Kisslip cuttlefish . . . . . . . . . . . . . . . . . . . 745 

Kitefin shark . . . . . . . . . . . . . . . . . . . . 1225 

Knifebone cuttlefish . . . . . . . . . . . . . . . . . 759 

Knight rock shrimp . . . . . . . . . . . . . . . . . . 954 

Knobby bobtail squid . . . . . . . . . . . . . . . . . 717 

Kobi cuttlefish . . . . . . . . . . . . . . . . . . . . 743 

Kobi squid . . . . . . . . . . . . . . . . . . . . . . 773 

kobiensis var. albatrossi, Sepia . . . . . . . . . . . 743 

kobiensis var. andreanoides, Sepia . . . . . . . . . 743 

kobiensis var. beppauna, Sepia . . . . . . . . . . . 743 

kobiensis var. crassa, Sepia . . . . . . . . . . . . . 743 

kobiensis, Loligo . . . . . . . . . . . . . . . . . . . 773 

kobiensis, Sepia . . . . . . . . . . . . . . . . 737, 743 

Koch’s bottle squid . . . . . . . . . . . . . . . . . . 720 

kochii, Sepiadarium . . . . . . . . . . . . . . 694, 720 

koelbeli, Solenocera . . . . . . . . . . . . . . 884, 887 

kuboi, Solenocera . . . . . . . . . . . . . . . . . . 883 

Kuruma prawn . . . . . . . . . . . . . . . . . . . . 917 

L 

Laai Por . . . . . . . . . . . . . . . . . . . . . . . 810 

laevimanus, Etisus . . . . . . . . . . . . . . . . 1100 

lalandii, Gecarcoidea . . . . . . . . . . . . . . . 1150 

lamellata, Metapenaeopsis . . . . . . . . . . . . . 930 

Lamiopsis . . . . . . . . . . . . . . . . . . . . . 1312 

Lamiopsis temmincki . . . . . . . . . . . . 1350, 1352 

Lamnid sharks . . . . . . . . . . . . . . . . . . . 1197 

LAMNIDAE . . . . . . . . . . . 1196-1197, 1274, 1314 

Lamnoid sharks . . . . . . . . . . . . . . . . . . 1196 

lana, Solitosepia . . . . . . . . . . . . . . . . . . . 747 

lancifera, Sicyonia . . . . . . . . . . . . . . 952, 954 

Land crabs . . . . . . . . . . . . . . 1049, 1060, 1147 

Land hermit crabs . . . . . . . . . . 1048, 1061, 1154 

Langosta colorete . . . . . . . . . . . . . . . . . 1021 

Langosta de muelas . . . . . . . . . . . . . . . . 1023 

Langosta del Indo-Pacifico . . . . . . . . . . . . . 1004 

Langosta duende . . . . . . . . . . . . . . . . . . 1017 

Langosta fanguera . . . . . . . . . . . . . . . . . 1020 

Langosta festoneada . . . . . . . . . . . . . . . . 1016 

Langosta horquilla . . . . . . . . . . . . . . . . . 1019 

Langosta ornamentada . . . . . . . . . . . . . . 1018 

Langouste barriolée . . . . . . . . . . . . . . . . 1021 

Langouste de vase . . . . . . . . . . . . . . . . . 1020 

Langouste diablotin . . . . . . . . . . . . . . . . 1017 

Langouste festonée . . . . . . . . . . . . . . . . 1016 

Langouste fourchette . . . . . . . . . . . . . . . . 1019


Langouste gibbon . . . . . . . . . . . . . . . . . 1023 

Langouste ornée . . . . . . . . . . . . . . . . . . 1018 

Langoustes . . . . . . . . . . . . . . . . . . 979, 1005 

Langoustine andamane . . . . . . . . . . . . . . . 992 

Langoustine indienne . . . . . . . . . . . . . . . . 991 

Langoustine spinuleuse . . . . . . . . . . . . . . . 988 

lar, Macrobrachium . . . . . . . . . . . . . . . . . 967 

laticaudus, Scoliodon . . . . . . . . 1351, 1354-1357 

laticaudus, Squaliolus . . . . . . . . . . . . . . 1229 

latimanus, Sepia . . . . . . . . . . . . . . . . . . . 744 

latisulcatus hathor, Penaeus . . . . . . . . . . . . 918 

latisulcatus, Penaeus . . . . . . . . . . . 914, 918-919 

LATREILLIDAE . . . . . . . . . . . . . . . . . . 1083 

Latreillopsis hawaiiensis . . . . . . . . . . . . . 1084 

latro, Birgus . . . . . . . . . . 1048, 1061, 1154, 1155 

Lauridromia . . . . . . . . . . . . . . . . . . . . 1085 

Lauridromia dehaani . . . . . . . . . . . . . . . 1088 

Lauridromia indica . . . . . . . . . . . . . . . . 1088 

Leafscale gulper shark . . . . . . . . . . . . . . . 1224 

Leandrites indicus . . . . . . . . . . . . . . . . . . 966 

Leopard fish . . . . . . . . . . . . . . . . . . . . 1172 

Leptocarpus potamiscus . . . . . . . . . . . . . . 966 

Lesser blue-ringed octopus . . . . . . . . . . . . . 821 

Lesser flying squid . . . . . . . . . . . . . . . . . . 796 

lessoniana, Sepioteuthis . . . . . . 688, 765, 778, 797 

lesuerii, Matuta . . . . . . . . . . . . . . . . . . 1095 

lesueuri, Ancistrocheirus . . . . . . . . . . . . . . 797 

leucas, Carcharhinus . . . . . . . . 1196, 1329, 1339 

leucoperiptera, Hemitriakis . . . . . . . . . . . . 1302 

LEUCOSIIDAE . . . . . . . . . . . . . . . . . . 1092 

leucospilota, Holothuria . . . . . . . . . . . . . . 1176 

leucospilota, Holothuria (Mertensiothuria) . . . 1178 

lewini, Sphyrna . . . . . . . . . . . . . . . 1364-1366 

licha, Dalatias . . . . . . . . . . . . . . . . . . . 1225 

limata, Arctosepia . . . . . . . . . . . . . . . . . . 739 

limbatus, Carcharhinus . . . . 1327, 1332, 1340, 1348 

Limnocarcinus intermedius . . . . . . . . . . . . 1150 

Lined lanternshark . . . . . . . . . . . . . . . . . 1227 

lineolata, Sepioloidea . . . . . . . . . . . . . . . . 720 

Linuparus . . . . . . . . . . . . . . . 977, 1006, 1011 

Linuparus sordidus . . . . . . . . . . . . . . . . 1024 

Linuparus trigonus . . . . . . . . . . . . . . . . 1024 

LITHODIDAE . . . . . . . . . . . . . . . . . . . . 1048 

litoralis, Ameloctopus . . . . . . . . . . . . . . . . 820 

litterata, Varuna . . . . . . . . . . . . . . . . . . 1144 

Little cuttlefish . . . . . . . . . . . . . . . . . . . . 760 

Lizardfish . . . . . . . . . . . . . . . . . . . . . . 1333 

lobifrons, Lupa . . . . . . . . . . . . . . . . . . . 1127 

Lobsterettes . . . . . . . . . . . . . . . . . . 977, 982 

Lobsters . . . . . . . . . . . . . . . . . . . . . . . 976 

LOLIGINIDAE . . . . . . . . . . . . 688, 721, 764, 797 

Loliginids . . . . . . . . . . . . . . . . . . . . 688, 765 

Loligo . . . . . . . . . . . . . . . . . . . . . . . . 765 

Loligo australis . . . . . . . . . . . . . . . . . . . 774 

Loligo budo . . . . . . . . . . . . . . . . . . . . . 776 

Loligo chinensis . . . . . . . . . . . . . . . . . . . 774 

Loligo duvauceli . . . . . . . . . . . . . . . . . . . 775 

Loligo edulis . . . . . . . . . . . . . . . . . . . . . 765 

Loligo etheridgei . . . . . . . . . . . . . . . . 765, 774 

Loligo formosana . . . . . . . . . . . . . . . . . . 774 

Loligo indica . . . . . . . . . . . . . . . . . . 774-775 

Loligo kobiensis . . . . . . . . . . . . . . . . . . . 773 

Loligo n.sp. . . . . . . . . . . . . . . . . . . . . . 766 

Loligo oshimai . . . . . . . . . . . . . . . . . . . . 775 

Loligo reesi . . . . . . . . . . . . . . . . . . . . . 766 

Loligo rhomboidalis . . . . . . . . . . . . . . . . . 773 

Loligo sibogae . . . . . . . . . . . . . . . . . . . . 777 

Loligo vossi . . . . . . . . . . . . . . . . . . 766, 774 

Loligo yokoyae . . . . . . . . . . . . . . . . . . . . 773 

Loliolus . . . . . . . . . . . . . . . . . . . . . . . 765 

Loliolus spp. . . . . . . . . . . . . . . . . . . . . . 688 

Loliolus affinis . . . . . . . . . . . . . . . . . . . 780 

Loliolus noctiluca . . . . . . . . . . . . . . . . . . 780 

Lollyfish . . . . . . . . . . . . . . . . . . . . . . . 1176 

LOMIDAE . . . . . . . . . . . . . . . . . . . . . 1077 

Long barrel squid . . . . . . . . . . . . . . . . . . . 777 

Long-armed crab . . . . . . . . . . . . . . . . . . 1114 

Longarm furrow lobster . . . . . . . . . . . . . . 1023 

Longfin catshark . . . . . . . . . . . . . . . . . . 1286 

Longfin mako . . . . . . . . . . . . . . . . . . . . 1278 

Longfingered peeler crab . . . . . . . . . . . . . 1108 

Longhead catshark . . . . . . . . . . . . . . . . . 1286 

longicaudatus, Negogaleus . . . . . . . . . . . . 1311 

longicephalus, Apristurus . . . . . . . . . . . . 1286 

longimanus japonicus, Carcinoplax . . . . . . . 1114 

longimanus typicus, Carcinoplax . . . . . . . . . 1114 

longimanus, Carcharhinus . . . . . . . . . 1325, 1341 

longimanus, Carcinoplax . . . . . . . . . . 1114, 1132 

longimanus, Justitia . . . . . . . . . . . . 1005, 1023 

longipes femoristriga, Panulirus . . . . . . 1015, 1017 

longipes longipes, Panulirus . . . . . . . . . . . 1017 

longipes, Cardisoma . . . . . . . . . . . . 1149, 1151 

longipes, Panulirus . . . . . . . . . . . . . 1015, 1017 

longipes, Panulirus longipes . . . . . . . . . . . 1017 

longipes, Parapenaeus . . . . . . . . . . . . . . . 947 

longipes, Trachypenaeus . . . . . . . . . . . 927, 950 

longistylus, Penaeus . . . . . . . . . . . . . 918-919 

Longlegged land crab . . . . . . . . . . . . . . . 1151 

Longlegged rough shrimp . . . . . . . . . . . . . . 950 

Longlegged spiny lobster . . . . . . . . . . . . . 1017 

Longnose houndshark . . . . . . . . . . . . . . . 1303 

Longsnout dogfish . . . . . . . . . . . . . . . . . 1225 

Longtail carpetsharks . . . . . . . . . . . . . . . 1249 

Longtailed carpetsharks . . . . . . . . . . . . . . 1197 

lophos, Calappa . . . . . . . . . . . . . . . 1091, 1094 

Lophozozymus . . . . . . . . . . . . . . . . . . . 1049 

Lophozozymus pictor . . . . . . . . . . . . 1049, 1098 

Loxodon macrorhinus . . . . . . . . 1351, 1354-1356 

lucifer, Etmopterus . . . . . . . . . . . . . . . . 1226 

LUCIFERIDAE . . . . . . . . . . . . . . . . . 856, 858 

Luminous bay squid . . . . . . . . . . . . . . . . . 780 

lunaris, Ashtoret . . . . . . . . . . . . 1091, 1095-1096 

lunaris, Cancer . . . . . . . . . . . . . . . . . . 1095 

lunaris, Matuta . . . . . . . . . . . . . . . . . . 1095 

lunulata, Hapalochlaena . . . . . . . . . . . . . . 821 

Lupa lobifrons . . . . . . . . . . . . . . . . . . . 1127 

lusitanicus, Centrophorus . . . . . . . . . . . . . 1224 

luteus, Octopus . . . . . . . . . . . . . . . . 801, 814 

luteus, Octopus cf. . . . . . . . . . . . . . . . . . 814 

lycidas, Sepia . . . . . . . . . . . . . . . . . . . . 745


lysianassa, Metapenaeus . . . . . . . . . . . . . . 939 

Lysiosquilla . . . . . . . . . . . . . . . . . . 829, 835 

LYSIOSQUILLIDAE . . . . . . . . . . . . . . 829, 835 

Lysiosquillids . . . . . . . . . . . . . . . . . . . . . 835 

Lysiosquillina . . . . . . . . . . . . . . . . . 829, 835 

Lysiosquillina maculata . . . . . . . . . 829, 835, 837 

Lysmata amboinensis . . . . . . . . . . . . . . . . 961 

Lysmata debelius . . . . . . . . . . . . . . . . . . 962 

M 

Mackerel sharks . . . . . . . . . . . . . . . . . . 1274 

macleayi, Atelomycterus . . . . . . . . . . . . . 1288 

macleayi, Metapenaeus . . . . . . . . . . . . . . . 939 

macloti, Carcharhinus . . . . 1342, 1351, 1354-1356 

macloti, Hypoprion . . . . . . . . . . . . . . . . 1342 

Macrobrachium sp. . . . . . . . . . . . . . . . . . 969 

Macrobrachium carcinus . . . . . . . . . . . . . . 964 

Macrobrachium equidens . . . . . . . . . . . . . 967 

Macrobrachium lar . . . . . . . . . . . . . . . . . 967 

Macrobrachium mirabile . . . . . . . . . . . . . . 968 

Macrobrachium rosenbergii . 855, 957, 964, 966-967 

Macrobrachium rosenbergii dacqueti . . . . . . . . 964 

macrochira, Paromola . . . . . . . . . . . . 1083-1084 

macrodactylus, Etisus . . . . . . . . . . . . . . . 1100 

macromphalus, Nautilus . . . . . . . . . . . . . . 711 

Macrophtalmus spp. . . . . . . . . . . . . . . . 1048 

macropus, Octopus . . . 689, 800-801, 810, 814, 816 

macrorhinus, Loxodon . . . . . . . 1351, 1354-1356 

macrostoma, Chaenogaleus . . . . . . . . 1308-1310 

macrostoma, Hemigaleus . . . . . . . . . . . . . 1308 

MACRURA NATANTIA . . . . . . . . . . . . . . . . 854 

MACRURA REPTANTIA . . . . . . . . . . . . . . . 854 

maculata, Holothuria . . . . . . . . . . . . . . . 1181 

maculata, Lysiosquillina . . . . . . . . 829, 835, 837 

maculatus, Carpilius . . . . . . . . . . . . 1110-1111 

maculatus, Etisus . . . . . . . . . . . . . . . . . 1100 

maculatus, Orectolobus . . . . . . . . . . . . . . 1247 

maculosa, Hapalochlaena cf. . . . . . . . . . . . 821 

madagascariensis, Haliporoides sibogae . . . . . 881 

madokai, Sepia . . . . . . . . . . . . . . . . 751, 759 

Madokai’s cuttlefish . . . . . . . . . . . . . . . . . 759 

magnificus, Palibythus . . . . . . . . . . . . . . 1004 

magnocellatus, Octopus . . . . . . . . . . . . . . . 812 

MAJIDAE . . . . . . . . . . . . . . . 1059, 1083, 1136 

Majids . . . . . . . . . . . . . . . . . . . . 1050-1051 

major, Parribacus ursus . . . . . . . . . . . . . . 1037 

Mako sharks . . . . . . . . . . . . . . . . . . . . 1197 

Makos . . . . . . . . . . . . . . . . . . . . . . . 1274 

malaiana, Trachypenaeus . . . . . . . . 927-928, 950 

Malayan mud shrimp . . . . . . . . . . . . . . . . . 886 

Malayan rough shrimp . . . . . . . . . . . . . . . . 928 

malayense, Sepiadarium . . . . . . . . . . . . . . . 720 

manazo, Mustelus . . . . . . . . . . . . . . . . . 1304 

Mandarin dogfish . . . . . . . . . . . . . . . . . . 1224 

mandroni, Sepiella . . . . . . . . . . . . . . . . . . 756 

Mangrove prawn . . . . . . . . . . . . . . . . . . . 969 

Mangrove stone crab . . . . . . . . . . . . . . . . 1107 

manillensis, Penaeus monodon . . . . . . . . . . . 925 

manningi, Chaceon . . . . . . . . . . . . . . . . 1134 

Mantis shrimps . . . . . . . . . . . . . . . . . 829, 855 

maou, Carcharhinus . . . . . . . . . . . . . . . . 1341 

Marbled mitten lobster . . . . . . . . . . . . . . . 1042 

Marbled octopus . . . . . . . . . . . . . . . . . . . 811 

marchei, Pelocarcinus . . . . . . . . . . . . . . . 1150 

margaritiferum, Chiloscyllium . . . . . . . . . . 1257 

marginatus, Octopus . . . . . . . . . . . . . . . . 815 

marginatus, Penaeus . . . . . . . . . . . . . . . . 920 

maritae, Chaceon . . . . . . . . . . . . . . . . . 1132 

marmorata, Bohadschia . . . . . . . . . . . . . . 1174 

marmoratus, Atelomycterus . . . . . . . . . . . 1288 

marmoratus, Octopus . . . . . . . . . . . . . . . . 812 

marmoratus, Saron . . . . . . . . . . . . . . . . . 962 

Maroon stone crab . . . . . . . . . . . . . . . . . 1107 

Marrajo . . . . . . . . . . . . . . . . . . . . . . . 1277 

Marrajo carite . . . . . . . . . . . . . . . . . . . 1278 

Marrajo dientuso . . . . . . . . . . . . . . . . . . 1277 

mastersii, Metapenaeus . . . . . . . . . . . . 910, 912 

Mastigoteuthid squids . . . . . . . . . . . . . . . . 799 

MASTIGOTEUTHIDAE . . . . . . . . . . . . 798-799 

Mastigoteuthis . . . . . . . . . . . . . . . . . . . . 799 

Mastigoteuthis cordiformis . . . . . . . . . . . . . 799 

Matuta banksii . . . . . . . . . . . . . . . . . . . 1095 

Matuta crebripunctata . . . . . . . . . . . . . . . 1095 

Matuta lesuerii . . . . . . . . . . . . . . . . . . . 1095 

Matuta lunaris . . . . . . . . . . . . . . . . . . . 1095 

Matuta peronii . . . . . . . . . . . . . . . . . . . 1095 

Matuta planipes . . . . . . . . . . . . . . . . . . 1094 

Matuta victor . . . . . . . . . . . . . . . . . . . 1095 

MATUTINAE . . . . . . . . . . . . . 1050, 1091, 1115 

mauritiana, Actinopyga . . . . . . . . . . . 1167-1168 

mauritiana, Justitia . . . . . . . . . . . . . . . . 1023 

mauritianus, Portunus . . . . . . . . . . . . . . . 1124 

maxillipedo, Parapenaeopsis . . . . . . . . . 942, 944 

maximus, Cetorhinus . . . . . . . . . . . . . . . 1274 

Meat crabs . . . . . . . . . . . . . . . . . . . . . 1126 

mederi, Episesarma . . . . . . . . . . . . . 1143, 1145 

megalops, Squalus . . . . . . . . . . . . . . . . 1230 

melanobranchius, Parmaturus . . . . . . . . . . 1291 

melanopterus, Carcharhinus . . . . . . . . 1333, 1343 

melantho, Solenocera . . . . . . . . . . . 884, 886-887 

melanurus, Squalus . . . . . . . . . . . . . . . . 1230 

membranaceus, Octopus . . . . . . . . . . . . . . 819 

Menippe granulosav . . . . . . . . . . . . . . . . 1107 

Menippe rumphii . . . . . . . . . . . . . . 1103, 1107 

MENIPPIDAE . . . . . . . . . . . . . . . . 1050, 1103 

menisorrah, Carcharhinus . . . . . 1328, 1334, 1346 

merguiensis, Penaeus . . . . . 890, 916, 921, 923, 926 

mestus, Sepia . . . . . . . . . . . . . . . . . . . . 760 

Metanephrops . . . . . . . . . 975, 977, 982, 985, 992 

Metanephrops andamanicus . . . . . . . . . . . . 992 

Metanephrops australiensis . . . . . . . . . . . . 992 

Metanephrops neptunus . . . . . . . . . . . . . . 993 

Metanephrops sibogae . . . . . . . . . . . . . . . 989 

Metanephrops sinensis . . . . . . . . . . . . . . . 993 

Metanephrops thomsoni . . . . . . . . . . . . . . 990 

Metanephrops velutinus . . . . . . . . . . . . . . 994 

Metanephros spp. . . . . . . . . . . . . . . . . . . 974 

Metapenaeopsis . . . . . . . . . . . . . . . . 889-890 

Metapenaeopsis barbata . . . . . . . . . . . . . . 907 

Metapenaeopsis barbeensis . . . . . . . . . . . . . 908


Metapenaeopsis lamellata . . . . . . . . . . . . . 930 

Metapenaeopsis mogiensis . . . . . . . . . . . . . 930 

Metapenaeopsis novaeguineae . . . . . . . . 908, 931 

Metapenaeopsis palmensis . . . . . . . . . . . . . 908 

Metapenaeopsis rosea . . . . . . . . . . . . . . . . 931 

Metapenaeopsis stridulans . . . . . . . . . . 908, 932 

Metapenaeopsis toloensis . . . . . . . . . . . . . . 932 

Metapenaeopsis wellsi . . . . . . . . . . . . . . . 933 

Metapenaeus . . . . . . . . . . . . . . . 853, 890, 910 

Metapenaeus affinis . . . . . . . . . . . . . . . . . 933 

Metapenaeus anchistus . . . . . . . . . . . . 909, 938 

Metapenaeus benettae . . . . . . . . . . . . . . . 934 

Metapenaeus brevicornis . . . . . . . . . . . . . . 934 

Metapenaeus burkenroadi . . . . . . . . . . . . . . 912 

Metapenaeus conjunctus . . . . . . . . . . . . . . 935 

Metapenaeus dalli . . . . . . . . . . . . . . . 912, 935 

Metapenaeus demani . . . . . . . . . . . . . . . . 936 

Metapenaeus dobsoni . . . . . . . . . . . . . . . . 936 

Metapenaeus eboracensis . . . . . . . . . . . . . . 937 

Metapenaeus elegans . . . . . . . . . . . . . 937, 940 

Metapenaeus endeavouri . . . . . . . . . 909-910, 938 

Metapenaeus ensis . 910, 912, 933, 935, 937-938, 940 

Metapenaeus ensis baramensis . . . . . . . . . . . 910 

Metapenaeus incisipes . . . . . . . . . . . . . . . . 910 

Metapenaeus insolitus . . . . . . . . . . . . . . . 938 

Metapenaeus intermedius . . . . . . . . . . 909, 911 

Metapenaeus lysianassa . . . . . . . . . . . . . . 939 

Metapenaeus macleayi . . . . . . . . . . . . . . . 939 

Metapenaeus mastersii . . . . . . . . . . . . 910, 912 

Metapenaeus monoceros . . . . . . . . . . . . . . 910 

Metapenaeus moyebi . . . . . . . . . . . . . . . . 912 

Metapenaeus papuensis . . . . . . . . . . . . . . . 940 

Metapenaeus philippinensis . . . . . . . . . . . . . 910 

Metapenaeus suluensis . . . . . . . . . . . . . . . 940 

Metapenaeus tenuipes . . . . . . . . . . . . . . . 941 

Metasepia pfefferi . . . . . . . . . . . . . . . . . . 757 

microcheirus, Sepia (Sepiella) . . . . . . . . . . . 756 

microdon, Pseudotriakis . . . . . . . . . . . . . 1296 

microstoma, Hemigaleus . . . . . . . 1308-1309-1310 

microstoma, Negogaleus . . . . . . . . . . . . . . 1309 

Middle shrimp . . . . . . . . . . . . . . . . . . . . 911 

Milandre belette . . . . . . . . . . . . . . . . . . 1311 

Milandre chicor . . . . . . . . . . . . . . . . . . . 1310 

Milandre faucille . . . . . . . . . . . . . . . . . . 1309 

Milandre harpon . . . . . . . . . . . . . . . . . . 1308 

milberti, Carcharhinus . . . . . . . . . . . . . . 1345 

miliaris, Actinopyga . . . . . . . . . . . . . 1169-1171 

Milk shark . . . . . . . . . . . . . . . . . . . . . 1354 

MIMILAMBRIDAE . . . . . . . . . . . . . . . . . 1050 

Mino nylon shrimp . . . . . . . . . . . . . . . . . . 970 

mira, Sepia . . . . . . . . . . . . . . . . . . . . . . 760 

mirabile, Macrobrachium . . . . . . . . . . . . . 968 

miranda, Histioteuthis . . . . . . . . . . . . . . . . 787 

Mitre squid . . . . . . . . . . . . . . . . . . . . . . 774 

mitsukurii, Squalus . . . . . . . . . . . . . . . . 1230 

Miyakea . . . . . . . . . . . . . . . . . . . . . . . 842 

Miyakea nepa . . . . . . . . . . . . . . . . . 842, 847 

Mogi velvet shrimp . . . . . . . . . . . . . . . . . . 930 

mogiensis, Metapenaeopsis . . . . . . . . . . . . . 930 

mokarran, Sphyrna . . . . . . . . . . 1364-1365-1366 

molleri, Etmopterus . . . . . . . . . . . . . . . . 1226 

MOLPADIDA . . . . . . . . . . . . . . . . . 1159-1160 

Moluccas brush shrimp . . . . . . . . . . . . . . . 960 

moluccensis, Atyopsis . . . . . . . . . . . . . . . . 960 

moluccensis, Centrophorus . . . . . . . . . . . . 1223 

Monkey river prawn . . . . . . . . . . . . . . . . . 967 

monoceros, Metapenaeus . . . . . . . . . . . . . . 910 

monodon manillensis, Penaeus . . . . . . . . . . . 925 

monodon, Penaeus . . . . . . . . . . . . 890, 921-922 

Moon crabs . . . . . . . . . . . . . . 1057, 1091, 1115 

Moroteuthis . . . . . . . . . . . . . . . . . . . . . 784 

morsei, Eupryma . . . . . . . . . . . . . . . . . . 714 

Mosaic crab . . . . . . . . . . . . . . . . . . . . 1049 

Mosaic drop-arm octopus . . . . . . . . . . . . . . 822 

mototi, Octopus . . . . . . . . . . . . . . . . . . . 825 

Mottled Sally-light-foot . . . . . . . . . . . . . . . 1142 

Moyebi shrimp . . . . . . . . . . . . . . . . . . . . 912 

moyebi, Metapenaeus . . . . . . . . . . . . . . . . 912 

mozambica, Sepia . . . . . . . . . . . . . . . . . . 744 

Mud crabs . . . . . . . . . . . 1057, 1098, 1103, 1110 

Mud lobster . . . . . . . . . . . . . . . . . . . . . 1143 

Mud lobsters . . . . . . . . . . . . . . . . . 976, 1048 

Mud shrimps . . . . . . . . . . . . . . . 855, 976, 1048 

Mud spiny lobster . . . . . . . . . . . . . . . . . 1020 

mülleri, Physodon . . . . . . . . . . . . . . . . . 1357 

Musical furry lobster . . . . . . . . . . . . . . . . 1004 

Musola austral . . . . . . . . . . . . . . . . . . . 1304 

Musola celestrial . . . . . . . . . . . . . . . . . . 1304 

Musola gris . . . . . . . . . . . . . . . . . . . . . 1304 

Musolón aleta larga . . . . . . . . . . . . . . . . 1296 

Mustelus . . . . . . . . . . . . . . . . . . . . . . 1297 

Mustelus antarcticus . . . . . . . . . . . . . . . 1304 

Mustelus griseus . . . . . . . . . . . . . . . . . 1304 

Mustelus manazo . . . . . . . . . . . . . . . . . 1304 

Myomenippe fornasinii . . . . . . . . . . . 1107, 1109 

Myomenippe granulosa . . . . . . . . . . . . . . 1107 

Myomenippe hardwickii . . . . . . . . . . 1103, 1107 

Myopsid squids . . . . . . . . . . . . . . . . . 691, 694 

MYOPSIDA . . . . . . . . . . . . . . . . . . . . . . 688 

Myopsids . . . . . . . . . . . . . . . . . . . . . . . 688 

Mysid shrimps . . . . . . . . . . . . . . . . . . . . 855 

MYSIDACEA . . . . . . . . . . . . . . . . . . . . . 855 

MYXINIDAE . . . . . . . . . . . . . . . . . . . . 1192 

N 

nakamurai, Hexanchus . . . . . . . . . . . . . . 1210 

NANNOSQUILLIDAE . . . . . . . . . . . . . . . . 835 

Natal Sally-light-foot . . . . . . . . . . . . . . . . 1142 

natalis, Gecarcoidea . . . . . . . . . . . . . . . . 1150 

natator, Charybdis . . . . . . . . . . . . . 1120-1121 

natator, Charybdis . . . . . . . . . . . . . . . . . 1121 

Nautile bouton . . . . . . . . . . . . . . . . . . . . 711 

Nautile flammé . . . . . . . . . . . . . . . . . . . . 711 

NAUTILIDAE . . . . . . . . . . . . . . . . . . 688, 709 

Nautilo común . . . . . . . . . . . . . . . . . . . . 711 

Nautilo ombligo . . . . . . . . . . . . . . . . . . . . 711 

Nautilus . . . . . . . . . . . . . . . . . 690, 698, 709 

Nautilus belauensis . . . . . . . . . . . . . . . . . 711 

Nautilus macromphalus . . . . . . . . . . . . . . 711 

Nautilus pompilius . . . . . . . . . . . . . . . . . 711


Nautilus repertus . . . . . . . . . . . . . . . . . . . 711 

Nautilus scrobiculatus . . . . . . . . . . . . . . . . 711 

Nautilus stenomphalus . . . . . . . . . . . . . . . 711 

Nautiluses . . . . . . . . . . . . . . . . . . . . . . 688 

Nebrius concolor . . . . . . . . . . . . . . . . . 1260 

Nebrius doldi . . . . . . . . . . . . . . . . . . . 1260 

Nebrius ferrugineus . . . . . . . . . . . . . . . . 1260 

Needle cuttlefish . . . . . . . . . . . . . . . . . . . 736 

Needle shrimp . . . . . . . . . . . . . . . . . . . . 948 

Negaprion . . . . . . . . . . . . . . . . . . . . . 1312 

Negaprion acutidens . . . . . . . . . . . . 1350, 1352 

neglectus, Saron . . . . . . . . . . . . . . . . . . . 962 

Negogaleus longicaudatus . . . . . . . . . . . . 1311 

Negogaleus microstoma . . . . . . . . . . . . . . 1309 

Negogaleus tengi . . . . . . . . . . . . . . . . . . 1311 

nehereus, Harpodon . . . . . . . . . . . . . . . . 1357 

Nematopalaemon tenuipes . . . . . . . . . . . . . 968 

Neon flying squid . . . . . . . . . . . . . . . . . . . 793 

nepa, Miyakea . . . . . . . . . . . . . . . . . 842, 847 

NEPHROPIDAE . . . . . . 977, 982, 996, 1001, 1007 

Nephrops sibogae . . . . . . . . . . . . . . . . . . 989 

Nephrops thomsoni . . . . . . . . . . . . . . . . . 990 

Nephropsis . . . . . . . . . . . . . . . . . . . 977, 986 

Nephropsis stewarti . . . . . . . . . . . . . . . . . 991 

Neptune lobster . . . . . . . . . . . . . . . . . . . 993 

neptunus, Metanephrops . . . . . . . . . . . . . . 993 

Neritic squids . . . . . . . . . . . . . . . . . . . . . 688 

Nervous shark . . . . . . . . . . . . . . . . . . . 1333 

New Caledonia blackfish . . . . . . . . . . . . . . 1171 

New Caledonia catshark . . . . . . . . . . . . . . 1288 

New Guinea river shark . . . . . . . . . . . . . . 1360 

niaukang, Centrophorus . . . . . . . . . . . . . 1224 

nipponensis, Sepiolina . . . . . . . . . . . . . . . 718 

nipponicus, Nototodarus . . . . . . . . . . . . . . 792 

Nipponololigo beka . . . . . . . . . . . . . . . . . 772 

Nipponololigo spp. . . . . . . . . . . . . . . . . . 780 

Nipponololigo sumatrensis . . . . . . . . . . . . . 773 

Nipponololigo uyii . . . . . . . . . . . . . . . 772-773 

nobilis, Holothuria . . . . . . . . . . . . . . . . 1181 

nobilis, Holothuria (Microthele) . . . . . . . . 1183 

noctiluca, Loliolus . . . . . . . . . . . . . . . . . 780 

nocturnus, Octopus . . . . . . . . . . . . . . 801, 816 

noronhai, Odontaspis . . . . . . . . . . . . . . . 1267 

Northern rough shrimp . . . . . . . . . . . . . . . . 949 

Northern velvet shrimp . . . . . . . . . . . . . . . . 931 

Northern wobbegong . . . . . . . . . . . . . . . . 1248 

Northwest lobster . . . . . . . . . . . . . . . . . . 992 

Nototodarus gouldi . . . . . . . . . . . . . . . . . 792 

Nototodarus hawaiiensis . . . . . . 788, 792, 795-796 

Nototodarus nipponicus . . . . . . . . . . . . . . . 792 

Nototodarus philippinensis . . . . . . . . . . 792, 795 

novaeguineae, Metapenaeopsis . . . . . . . 908, 931 

novemdentatus, Ibacus . . . . . . . . . . . . . . 1035 

nuchalis, Thysanoteuthis . . . . . . . . . . . . . . 797 

Nurse sharks . . . . . . . . . . . . . . . . . 1197, 1260 

O 

obesum, Cardisoma . . . . . . . . . . . . . . . . 1149 

obesus, Triaenodon . . . . . . . . . . . . . 1325, 1358 

obscurus, Carcharhinus . . . . . . . . . . 1326, 1344 

obtusus, Triaenodon . . . . . . . . . . . . . . . . 1329 

occidentalis, Enoplometopus . . . . . . . . . . . 1000 

occidua, Solitosepia . . . . . . . . . . . . . . . . . 747 

Oceanic paddler crab . . . . . . . . . . . . . . . 1144 

Oceanic squids . . . . . . . . . . . . . . . . . . . . 688 

Oceanic whitetip shark . . . . . . . . . . . . . . . 1341 

oceanica, Scylla serrata var. . . . . . . . . . . . 1126 

ocellata, Sepiella . . . . . . . . . . . . . . . . . . 762 

ocellatum, Hemiscyllium . . . . . . . . . . . . . 1258 

OCTOPODA . . . . . . . . . . . . . . . . . . 689, 801 

OCTOPODIDAE . . . . . . . . . . . . . . . . 689, 800 

Octopods . . . . . . . . . . . . . . . . . . . . . . . 694 

OCTOPOTEUTHIDAE . . . . . . . . . . . . . 782, 797 

Octopoteuthis . . . . . . . . . . . . . . . . . . . . 797 

Octopus . . . . . . . . . . . . . . . . . . . . . . . 800 

Octopus abaculus . . . . . . . . . . . . . . . 800, 822 

Octopus aculeatus . . . . . . . . . . . . . . . 800, 822 

Octopus aegina . . . . . . . . . . . . . 811, 815, 819 

Octopus alpheus . . . . . . . . . . . . . . . . 801, 823 

Octopus arakawai . . . . . . . . . . . . . . . . . . 817 

Octopus aspilosomatis . . . . . . . . . . . . 801, 823 

Octopus australis . . . . . . . . . . . . . . . . . . 824 

Octopus bimaculatus . . . . . . . . . . . . . . . . 812 

Octopus bocki . . . . . . . . . . . . . . . . . . . . 801 

Octopus cf. luteus . . . . . . . . . . . . . . . . . . 814 

Octopus cyanea . . . . . . . . . . . 696, 812, 817-818 

Octopus cyaneus . . . . . . . . . . . . . . . . . . . 818 

Octopus defillipi . . . . . . . . . . . . . . . . . . . 801 

Octopus dierythraeus . . . . . . . . . . . . . 801, 824 

Octopus dollfusi . . . . . . . . . . . . . . . . . . . 811 

Octopus exannulatus . . . . . . . . . . . . . . . . 825 

Octopus graptus . . . . . . . . . . . . . . . . 801, 813 

Octopus hardwickei . . . . . . . . . . . . . . . . . 811 

Octopus horridus . . . . . . . . . . . . . . . 800-801 

Octopus kagoshimensis . . . . . . . . . . . . . . . 815 

Octopus luteus . . . . . . . . . . . . . . . . . 801, 814 

Octopus macropus . . . . 689, 800-801, 810, 814, 816 

Octopus magnocellatus . . . . . . . . . . . . . . . 812 

Octopus marginatus . . . . . . . . . . . . . . . . . 815 

Octopus marmoratus . . . . . . . . . . . . . . . . 812 

Octopus membranaceus . . . . . . . . . . . . . . . 819 

Octopus mototi . . . . . . . . . . . . . . . . . . . 825 

Octopus nocturnus . . . . . . . . . . . . . . 801, 816 

Octopus ornatus . . . . . . . . . . . . . . . . 801, 817 

Octopus polyzenia . . . . . . . . . . . . . . . . . . 826 

Octopus sp. A . . . . . . . . . . . . . . . . . 811, 819 

Octopus striolatus . . . . . . . . . . . . . . . . . . 815 

Octopus tetricus . . . . . . . . . . . . . . . . . . . 818 

Octopus vulgaris . . . . . . . . . . . . . . . . 689, 800 

Octopus wolfi . . . . . . . . . . . . . . . . . . . . 801 

Octopuses . . . . . . . . . . . . . . . . . . . 688-689 

Ocypode . . . . . . . . . . . . . . . . . . . . . . 1152 

Ocypode ceratophthalma . . . . . . . . . . 1152-1153 

Ocypode cordimanus . . . . . . . . . . . . . . . 1153 

OCYPODIDAE . . . . . 1050, 1060, 1138, 1147, 1152 

Ocypodids . . . . . . . . . . . . . . . . . . . . . 1048 

Ocythoe tuberculata . . . . . . . . . . . . . . . . . 802 

OCYTHOIDAE . . . . . . . . . . . . . . . . . . . . 802 

ODONTASPIDIDAE . . . . . . . . . 1196, 1264, 1314 

Odontaspis ferox . . . . . . . . . . . . . . . . . 1267


Odontaspis herbsti . . . . . . . . . . . . . . . . . 1267 

Odontaspis kamoharai . . . . . . . . . . . . . . . 1268 

Odontaspis noronhai . . . . . . . . . . . . . . . 1267 

Odontaspis taurus . . . . . . . . . . . . . . . . . 1266 

Odontaspix ferox . . . . . . . . . . . . . . . . . . 1266 

Odontodactylid mantis shrimps . . . . . . . . 829, 832 

ODONTODACTYLIDAE . . . . . . . . . . . . 829, 832 

Odontodactylus . . . . . . . . . . . . . . . . . . . 829 

Odontodactylus scyllarus . . . . . . . . . . . 829, 834 

Oegopsid squids . . . . . . . . . . . . . . . . . . . 694 

OEGOPSIDA . . . . . . . . . . . . . . . . . . 688-689 

Oegopsids . . . . . . . . . . . . . . . . . . . 691, 694 

officinalis, Sepia . . . . . . . . . . . . . . . . . . . 725 

Old woman octopus . . . . . . . . . . . . . . . . . 810 

oligolinx, Rhizoprionodon . . . . . . . . . . 1354-1356 

olivacea, Scylla . . . . . . . . . . . . . . . 1126-1127 

omani, Sepia . . . . . . . . . . . . . . . . . . . . . 762 

Ommastrephes bartramii . . . . . . . . . . . . . . 793 

Ommastrephes caroli stenodactyla . . . . . . . . . 793 

Ommastrephes sloani pacificus . . . . . . . . . . . 795 

Ommastrephid squid . . . . . . . . . . . . . . . . . 788 

OMMASTREPHIDAE . . . . . . . . . . 689, 784, 788 

Ommastrephids . . . . . . . . . . . . . 694, 788, 797 

Onefin catshark . . . . . . . . . . . . . . . . . . 1291 

ONYCHOTEUTHIDAE . . . . . . . . . . . . . 784, 789 

Onychoteuthids . . . . . . . . . . . . . . . . . 784, 797 

Onychoteuthis . . . . . . . . . . . . . . . . . . . . 784 

Onykia . . . . . . . . . . . . . . . . . . . . . . . . 784 

OPHIDIIFORMES . . . . . . . . . . . . . . . . . 1189 

Ophiuroids . . . . . . . . . . . . . . . . . . 1158-1159 

opilio, Chionoecetes . . . . . . . . . . . . . . . . 1055 

opipara, Acanthacaris . . . . . . . . . . . . . . . . 988 

opipara, Glyptosepia . . . . . . . . . . . . . . . . 746 

opipara, Sepia . . . . . . . . . . . . . . . . . . . . 746 

OPISTHOTEUTHIDAE . . . . . . . . . . . . . . . . 801 

Orange mud crab . . . . . . . . . . . . . . . . . . 1127 

Orange shrimp . . . . . . . . . . . . . . . . . . . . 929 

oratoria, Oratosquilla . . . . . . . . . . . . . 843, 847 

Oratosquilla . . . . . . . . . . . . . . . . . . 842-843 

Oratosquilla oratoria . . . . . . . . . . . . . 843, 847 

Oratosquillina . . . . . . . . . . . . . . . . . 842-843 

Oratosquillina gravieri . . . . . . . . . . . . 843, 848 

Oratosquillina interrupta . . . . . . . . . . . . . . 843 

Oratosquillina perpensa . . . . . . . . . . . 843, 848 

Oratosquillina quinquedentata . . . . . . . . . . . 849 

Oratosquillina solicitans . . . . . . . . . . . . . . 849 

ORECTOLOBIDAE . . . . . . . . . . . . . . . . 1245 

Orectolobus japonicus . . . . . . . . . . . . . . 1247 

Orectolobus maculatus . . . . . . . . . . . . . . 1247 

Orectolobus ornatus . . . . . . . . . . . . . . . . 1248 

Orectolobus wardi . . . . . . . . . . . . . . . . . 1248 

Oriental spear lobster . . . . . . . . . . . . . . . 1024 

orientalis, Dromia . . . . . . . . . . . . . . . . . 1088 

orientalis, Thenus . . . . . . . . . . . . . . . . . 1040 

ORITHYIIDAE . . . . . . . . . . . . . . . . . . . 1050 

ornata, Sepiella . . . . . . . . . . . . . . . . . . . 762 

Ornate spiny lobster . . . . . . . . . . . . . . . . 1018 

Ornate wobbegong . . . . . . . . . . . . . . . . . 1248 

ornatum, Scyllium . . . . . . . . . . . . . . . . . 1256 

ornatus, Octopus . . . . . . . . . . . . . . . 801, 817 

ornatus, Orectolobus . . . . . . . . . . . . . . . 1248 

ornatus, Panulirus . . . . . . . . . . . . . . . . 1018 

oshimai, Loligo . . . . . . . . . . . . . . . . . . . 775 

oualaniensis, Sthenoteuthis . 689, 694, 788, 793-794 

oualaniensis, Symplectoteuthis . . . . . . . . . . . 794 

Ovalbone cuttlefish . . . . . . . . . . . . . . . . . . 741 

Ovalipes . . . . . . . . . . . . . . . . . . . . . . 1122 

Ovalipes australiensis . . . . . . . . . . . . . . . 1122 

Ovalipes punctatus . . . . . . . . . . . . . . . . 1122 

oxyrinchus, Isurus . . . . . . . . . . . . . 1277-1278 

OZIIDAE . . . . . . . . . . . . . . . . . . . 1050, 1103 

Ozius guttatus . . . . . . . . . . . . . . . . . . . 1109 

Ozius tuberculosus 

P 

. . . . . . . . . . . . . . . . 1109 

Pacific golden crab . . . . . . . . . . . . . . . . . 1134 

pacifica, Histioteuthis celetaria . . . . . . . . . . . 787 

pacifica, Rossia . . . . . . . . . . . . . . . . . . . 716 

pacificus pacificus, Todarodes . . . . . . . . . . . 795 

pacificus pusillus, Todarodes . . . . . . . . . . . . 795 

pacificus ssp., Todarodes . . . . . . . . . . . 788, 796 

pacificus, Alloposus . . . . . . . . . . . . . . . . . 802 

pacificus, Ommastrephes sloani . . . . . . . . . . . 795 

pacificus, Todarodes . . . . . . . . . . . . . . . . . 795 

pacificus, Todarodes pacificus . . . . . . . . . . . . 795 

Paddler crabs . . . . . . . . . . . . . . . . . 1060, 1138 

pagenstecheri, Sepia . . . . . . . . . . . . . . . . . 750 

pageorum, Acanthosepion . . . . . . . . . . . . . . 753 

PAGURIDAE . . . . . . . . . . . . . . . . . 1048, 1154 

Painted harlequin shrimp . . . . . . . . . . . . . . . 963 

Painted spiny lobster . . . . . . . . . . . . . . . . 1021 

Palaemon concinnus . . . . . . . . . . . . . . . . 969 

PALAEMONIDAE . . . . . . . . . . . . . . . 958, 964 

palari, Eledone . . . . . . . . . . . . . . . . . . . 801 

palasorra, Scoliodon . . . . . . . . . 1354-1355, 1357 

palauensis, Actinopyga . . . . . . . . 1169-1170-1172 

palawanense, Episesarma . . . . . . . . . 1143, 1145 

Pale catshark . . . . . . . . . . . . . . . . . . . . 1286 

Palibythus . . . . . . . . . . . . . . . . . . . . . 1001 

Palibythus magnificus . . . . . . . . . . . 1004, 1028 

Palinurellus . . . . . . . . . . . . . . . . . . . . 1001 

Palinurellus wieneckii . . . . . . . . . . . . . . 1004 

PALINURIDAE . . . . . 977, 979, 984, 997, 1002, 1005 

Palinustus . . . . . . . . . . . . . . . . . . 1006, 1011 

Palinustus unicornutus . . . . . . . . . . . . . . 1025 

Palinustus waguensis . . . . . . . . . . . . . . . 1025 

Palm theif . . . . . . . . . . . . . . . . . . . . . . 1155 

palmata, Sepia . . . . . . . . . . . . . . . . . . . . 738 

palmensis, Metapenaeopsis . . . . . . . . . . . . . 908 

PANDALIDAE . . . . . . . . . . . . . . . . . 855, 969 

Pandalus . . . . . . . . . . . . . . . . . . . . . . . 957 

Panning’s blackfish . . . . . . . . . . . . . . . . . 1170 

Panulirus . . . 975, 977, 1005-1006, 1008, 1012, 1021 

Panulirus albiflagellum . . . . . . . . . . . 1015, 1017 

Panulirus burgeri . . . . . . . . . . . . . . . . . 1016 

Panulirus dasypus . . . . . . . . . . . . . . . . . 1016 

Panulirus fasciatus . . . . . . . . . . . . . 1020-1021 

Panulirus homarus . . . . . . . . . . . . . . . . 1016 

Panulirus japonicus . . . . . . . . . . . . . . . . 1017 

Panulirus longipes . . . . . . . . . . . . . 1015, 1017


Panulirus longipes femoristriga . . . . . . . 1015, 1017 

Panulirus longipes longipes . . . . . . . . . . . . 1017 

Panulirus ornatus . . . . . . . . . . . . . . . . . 1018 

Panulirus pascuensis . . . . . . . . . . . . . . . 1026 

Panulirus penicillatus . . . . . . . . . . . . 1019, 1041 

Panulirus polyphagus . . . . . . . . . . . . 1020-1021 

Panulirus stimpsoni . . . . . . . . . . . . . . . . 1026 

Panulirus versicolor . . . . . . . . . . . . . . . . 1021 

Papua shrimp . . . . . . . . . . . . . . . . . . . . . 940 

Papuan cuttlefish . . . . . . . . . . . . . . . . . . . 747 

Papuan epaulette shark . . . . . . . . . . . . . . 1258 

papuensis, Metapenaeus . . . . . . . . . . . . . . 940 

papuensis, Sepia . . . . . . . . . . . . . . . . . . . 747 

Paragaleus tengi . . . . . . . . . . . . . . . . . . 1311 

Paralithodes camtschaticus . . . . . . . . . 1048, 1055 

paramamosain, Scylla . . . . . . . . . . . 1127-1128 

Parapenaeopsis . . . . . . . . . . . . . . . . 853, 890 

Parapenaeopsis arafurica . . . . . . . . . . . . . 941 

Parapenaeopsis cornuta . . . . . . . . . . . 942, 944 

Parapenaeopsis coromandelica . . . . . . . . . . 942 

Parapenaeopsis cultrirostris . . . . . . . . . . . . . 913 

Parapenaeopsis gracillima . . . . . . . . . . . . . 943 

Parapenaeopsis hardwickii . . . . . . . . . . . . . 913 

Parapenaeopsis hungerfordi . . . . . . . . . . . . 943 

Parapenaeopsis maxillipedo . . . . . . . . . 942, 944 

Parapenaeopsis sculptilis . . . . . . . . . . . 913, 944 

Parapenaeopsis tenella . . . . . . . . . . . . . . . 945 

Parapenaeopsis uncta . . . . . . . . . . . . . . . . 945 

Parapenaeopsis venusta . . . . . . . . . . . . . . . 946 

Parapenaeus . . . . . . . . . . . . . . . . . . 889-890 

Parapenaeus fissuroides . . . . . . . . . . . . . . 946 

Parapenaeus fissurus . . . . . . . . . . . . . . . . 946 

Parapenaeus longipes . . . . . . . . . . . . . . . . 947 

PARASCYLLIIDAE . . . . . . . . . . . . . . . . . 1241 

Parascyllium collare . . . . . . . . . . . . . . . 1242 

PARATHELPHUSIDAE . . . . . . . 1048, 1050, 1147 

Parmaturus . . . . . . . . . . . . . . . . . . . . 1280 

Parmaturus melanobranchius . . . . . . . . . . 1291 

Paromola japonica . . . . . . . . . . . . . 1083-1084 

Paromola macrochira . . . . . . . . . . . . 1083-1084 

Parribacus . . . . . . . . . . . . . . . 977, 1028, 1031 

Parribacus antarcticus . . . . . . . . . . . . . . 1037 

Parribacus caledonicus . . . . . . . . . . . . . . 1041 

Parribacus holthuisi . . . . . . . . . . . . . . . 1042 

Parribacus scarlatinus . . . . . . . . . . . . . . 1042 

Parribacus ursus major . . . . . . . . . . . . . . 1037 

PARTHENOPIDAE . . . . . . . . . . . . . . . . . 1050 

parva, Sepiola . . . . . . . . . . . . . . . . . . . . 717 

parvispina, Heterocarpus . . . . . . . . . . . 969-970 

parysatis, Amplisepia . . . . . . . . . . . . . . . . 738 

pascuensis, Panulirus . . . . . . . . . . . . . . . 1026 

Pastel odontodactylid mantis shrimp . . . . . . . . . 834 

paucidentatus, Penaeus semisulcatus . . . . . . . . 925 

paucus, Isurus . . . . . . . . . . . . . . . . 1277-1278 

Pearsonothuria graeffei . . . . . . . . . . . . . . 1184 

Peau bleue . . . . . . . . . . . . . . . . . . . . . 1353 

pectinata, Solenocera . . . . . . . . . . . . . 887-888 

pectinulata, Solenocera . . . . . . . . . . . . 887-888 

Pejegato aleton . . . . . . . . . . . . . . . . . . . 1286 

Pejegato cabezón . . . . . . . . . . . . . . . . . 1286 

Pejegato de agallas negras . . . . . . . . . . . . 1291 

Pejegato de Borneo . . . . . . . . . . . . . . . . 1287 

Pejegato esponjoso . . . . . . . . . . . . . . . . 1287 

Pejegato jaspeado . . . . . . . . . . . . . . . . . 1288 

Pejegato mallero . . . . . . . . . . . . . . . . . . 1289 

Pejegato paliducho . . . . . . . . . . . . . . . . . 1286 

Pejegato pintado . . . . . . . . . . . . . . . . . . 1291 

Pejegato velero . . . . . . . . . . . . . . . . . . . 1291 

Pelagic squids . . . . . . . . . . . . . . . . . . . . 688 

Pelagic thresher . . . . . . . . . . . . . . . . . . 1271 

pelagicus, Alopias . . . . . . . . . . . . . . 1271-1273 

pelagicus, Portunus . . . . . . 1115, 1123-1124-1125 

Pelocarcinus cailloti . . . . . . . . . . . . . . . . 1150 

Pelocarcinus marchei . . . . . . . . . . . . . . . 1150 

Penaeid shrimps . . . . . . . . . . . . . . . . 855, 889 

PENAEIDAE . . . . . . . . . . 855, 869, 876, 889, 952 

PENAEIDEA . . . . . . . . . . . . . . . . . . . . . 858 

Penaeids . . . . . . . . . . . . . . . . . . . . . . . 889 

Penaeoid shrimps . . . . . . . . . . . . 856, 866, 889 

PENAEOIDEA . . . . . . 855-856, 858, 866, 956, 958 

Penaeoids . . . . . . . . . . . . . . . . . . . . . . 855 

Penaeopsis . . . . . . . . . . . . . . . . . . . 889-890 

Penaeopsis eduardoi . . . . . . . . . . . . . . 947, 948 

Penaeopsis rectacuta . . . . . . . . . . . . . 947-948 

Penaeus . . . . . . . . . . . . . . . 853, 889-890, 934 

Penaeus ashiaka . . . . . . . . . . . . . . . . . . . 925 

Penaeus bubulus . . . . . . . . . . . . . . . . . . . 922 

Penaeus caesius . . . . . . . . . . . . . . . . . . . 919 

Penaeus canaliculatus . . . . . . . . . . . . . . . 914 

Penaeus carinatus . . . . . . . . . . . . . . . . . . 922 

Penaeus esculentus . . . . . . . . . . . . . . 915, 925 

Penaeus indicus . . . . . . . . . . . 916, 921, 923, 926 

Penaeus japonicus . . . . . . . . . . . . . . 914, 917 

Penaeus jejunus . . . . . . . . . . . . . . . . . . . 919 

Penaeus latisulcatus . . . . . . . . . . . 914, 918-919 

Penaeus latisulcatus hathor . . . . . . . . . . . . . 918 

Penaeus longistylus . . . . . . . . . . . . . . 918-919 

Penaeus marginatus . . . . . . . . . . . . . . . . 920 

Penaeus merguiensis . . . . . 890, 916, 921, 923, 926 

Penaeus monodon . . . . . . . . . . . . . 890, 921-922 

Penaeus monodon manillensis . . . . . . . . . . . 925 

Penaeus penicillatus . . . . . . . . 916, 921, 923, 926 

Penaeus plebejus . . . . . . . . . . . . . . . . . . 924 

Penaeus semisulcatusv . . . . . . . . . . . . 915, 925 

Penaeus semisulcatus exsulcatus . . . . . . . . . . 922 

Penaeus semisulcatus paucidentatus . . . . . . . . 925 

Penaeus silasi . . . . . . . . . . . . 916, 921, 923, 926 

Penaeus sp. . . . . . . . . . . . . . . . 914, 917, 922 

Penaeus teraoi . . . . . . . . . . . . . . . . . . . . 920 

Pencil squids . . . . . . . . . . . . . . . . . . . . . 764 

penicillatus, Panulirus . . . . . . . . . . . 1019, 1041 

penicillatus, Penaeus . . . . . . . . 916, 921, 923, 926 

Pentanchus . . . . . . . . . . . . . . . . . . . . . 1279 

Pentanchus profundicolus . . . . . . . . . . . . 1291 

peregrina, Solitosepia rozella . . . . . . . . . . . . 752 

Periscope crabs . . . . . . . . . . . . . . . . . . 1048 

Periscope shrimp . . . . . . . . . . . . . . . . . . . 929 

perlo, Heptranchias . . . . . . . . . . . . . . . . 1210 

Perna spp. . . . . . . . . . . . . . . . . . . . . . 1107 

peronii, Ibacus . . . . . . . . . . . . . . . . . . . 1035


peronii, Matuta . . . . . . . . . . . . . . . . . . 1095 

perpensa, Oratosquillina . . . . . . . . . . . 843, 848 

Petit taupe . . . . . . . . . . . . . . . . . . . . . 1278 

Pez toro . . . . . . . . . . . . . . . . . . . . . . . 1266 

Pfeffer’s flamboyant cuttlefish . . . . . . . . . . . . 757 

pfefferi, Metasepia . . . . . . . . . . . . . . . . . 757 

Pharaoh cuttlefish . . . . . . . . . . . . . . . . . . 748 

pharaonis, Sepia . . . . . . . . . . . . . . . 744, 748 

philargius, Calappa . . . . . . . . . 1091, 1094, 1097 

Philippine night octopus . . . . . . . . . . . . . . . 816 

Philippine sawshark . . . . . . . . . . . . . . . . 1234 

philippinensis, Metapenaeus . . . . . . . . . . . . 910 

philippinensis, Nototodarus . . . . . . . . . . 792, 795 

Philippines rough shrimp . . . . . . . . . . . . . . . 951 

Phoberus brevirostris . . . . . . . . . . . . . . . . 988 

Phoberus caecus sublevis . . . . . . . . . . . . . . 988 

Phoberus tenuimanus . . . . . . . . . . . . . . . . 988 

Photololigo . . . . . . . . . . . . . . . . 689, 691, 765 

Photololigo chinensis . . . . . 765-766, 774-775, 777 

Photololigo duvaucelii . . . . . . . . . . . . 774-775 

Photololigo edulis . . . . . . . . . . 765-766, 774, 776 

Photololigo singhalensis . . . . . . . . . . . . . . 777 

Photololigo sp. 1 . . . . . . . . . . . . . . . . 766, 776 



Photololigo spp. . . . . . . . . . . . . . . . . . . . 780 

Physodon mülleri . . . . . . . . . . . . . . . . . 1357 

picta, Hymenocera . . . . . . . . . . . . . . . . . 963 

pictor, Lophozozymus . . . . . . . . . . . . 1049, 1098 

Pigeye shark . . . . . . . . . . . . . . . . . . . . 1329 

PILUMNIDAE . . . . . . . . . . . . . . . . 1058, 1112 

Pilumnids . . . . . . . . . . . . . . . . . . . . . . 1112 

Pincer lobsters . . . . . . . . . . . . . . . . . . . . 977 

Pink lanternshark . . . . . . . . . . . . . . . . . . 1227 

Pink velvet shrimp . . . . . . . . . . . . . . . . . . 931 

Pink-fingered vinegar crab . . . . . . . . . . . . . 1145 

Pinkfish . . . . . . . . . . . . . . . . . . . . . . . 1177 

Pintarroja australiana . . . . . . . . . . . . . . . 1289 

Pintarroja coral . . . . . . . . . . . . . . . . . . . 1288 

Pintarroja enana . . . . . . . . . . . . . . . . . . 1290 

Pintarroja rabonegro . . . . . . . . . . . . . . . . 1290 

Pintarroja salamanquesa . . . . . . . . . . . . . . 1289 

PIRIMELIDAE . . . . . . . . . . . . . . . . . . . 1050 

plagiosum, Chiloscyllium . . . . . . . 1253-1256-1257 

Plagusia tuberculata . . . . . . . . . . . . . . . 1146 

Plain-body night octopus . . . . . . . . . . . . . . . 823 

Plain-spot ocellate octopus . . . . . . . . . . . . . 825 

plangon adhaesa, Solitosepia . . . . . . . . . . . . 749 

plangon, Sepia . . . . . . . . . . . . . . . . . . . . 749 

planipes, Matuta . . . . . . . . . . . . . . . . . 1094 

Platypodia granulosa . . . . . . . . . . . . . . . 1049 

PLATYXANTHIDAE . . . . . . . . . . . . . . . . 1050 

playfairi, Hypoprion . . . . . . . . . . . . . . . . 1343 

plebejus, Penaeus . . . . . . . . . . . . . . . . . . 924 

Plesiopenaeus edwardsianus . . . . . . . . . . . . 874 

pleurotaenia, Carcharhinus . . . . . . . . . . . . 1327 

plumbeus, Carcharhinus . . . . . . . . . . . . . 1345 

Podophthalmus . . . . . . . . . . . . . . . . . . 1123 

Podophthalmus vigil . . . . . . . . . . . . . . . 1123 

Podopilumnus fittoni . . . . . . . . . . . . . . . . 1113 

Poison ocellate octopus . . . . . . . . . . . . . . . 825 

Poisonous crabs . . . . . . . . . . . . . . . . . . 1049 

POLYCHELIDAE . . . . 977-978, 983, 996, 1002, 1007 

Polynesian furrow lobster . . . . . . . . . . . . . 1023 

Polynesian golden crab . . . . . . . . . . . . . . 1135 

polyphagus, Panulirus . . . . . . . . . . . 1020-1021 

polyzenia, Octopus . . . . . . . . . . . . . . . . . 826 

pompilius, Nautilus . . . . . . . . . . . . . . . . . 711 

Ponderisepia eclogaria . . . . . . . . . . . . . . . 744 

Pondicherry shark . . . . . . . . . . . . . . . . . 1338 

Porbeagles . . . . . . . . . . . . . . . . . . . . . 1274 

porosus, Carcharhinus . . . . . . . . . . . . . . 1359 

Port Jackson shark . . . . . . . . . . . . . . . . . 1240 

Porter crabs . . . . . . . . . . . . . . . . . 1050, 1083 

PORTUNIDAE . . . . . . . . . 1050, 1059, 1091, 1115 

Portunids . . . . . . . . . . . . . . . . . . . . . 1115 

Portunus . . . . . . . . . . . . . . . 1120, 1124,-1125 

Portunus mauritianus . . . . . . . . . . . . . . . 1124 

Portunus pelagicus . . . . . . 1115, 1123-1124-1125 

Portunus sanguinolentus . . . . . . . . . . 1115, 1125 

Portunus trituberculatus . . . . . . . 1115, 1124-1125 

portusjacksoni, Heterodontus . . . . . . . . . . 1240 

Pota cárdena . . . . . . . . . . . . . . . . . . . . . 794 

Pota costera . . . . . . . . . . . . . . . . . . . . . 796 

Pota hawaiana . . . . . . . . . . . . . . . . . . . . 792 

Pota japonesa . . . . . . . . . . . . . . . . . . . . 795 

Pota saltadora . . . . . . . . . . . . . . . . . . . . 793 

POTAMIDAE . . . . . . . . . . . . . 1048, 1050, 1147 

potamiscus, Leptocarpus . . . . . . . . . . . . . . 966 

POTAMOTRYGONIDAE . . . . . . . . . . . . . . 1196 

Poulpe des sables . . . . . . . . . . . . . . . . . . 815 

Poulpe nain . . . . . . . . . . . . . . . . . . . . . . 811 

Poulpe vieille femme . . . . . . . . . . . . . . . . . 810 

poupini, Chaceon . . . . . . . . . . . . . . 1132, 1135 

POUPINIIDAE . . . . . . . . . . . . . . . . . . . 1083 

Poupiniids . . . . . . . . . . . . . . . . . . . . . 1083 

Prickly deep-sea lobster . . . . . . . . . . . . . . . 988 

Prickly redfish . . . . . . . . . . . . . . . . . . . 1189 

Prickly shark . . . . . . . . . . . . . . . . . . . . 1212 

Prionace . . . . . . . . . . . . . . . . . . . . . . 1312 

Prionace glauca . . . . . . . . . . . . . . . . . . 1353 

prionota, Sepia . . . . . . . . . . . . . . . . . . . 747 

PRISTIDAE . . . . . . . . . . . . . . . . . . . . . 1233 

PRISTIOPHORIDAE . . . . . . . . . 1211, 1213, 1233 

Pristiophorus cirratus . . . . . . . . . . . . . . . 1234 

Pristiophorus japonicus . . . . . . . . . . . . . . 1234 

Pristiophorus sp. . . . . . . . . . . . . . . . . . 1234 

Pristiophorus sp. B . . . . . . . . . . . . . . . . 1234 

profundicolus, Pentanchus . . . . . . . . . . . . 1291 

profundorum, Deania . . . . . . . . . . . . . . . 1225 

profundus, Alopias . . . . . . . . . . . . . . . . . 1272 

prominentis, Solenocera . . . . . . . . . . . . . . . 884 

Pronghorn decorator crab . . . . . . . . . . . . . 1137 

Pronghorn spiny lobster . . . . . . . . . . . . . . 1019 

PROSCYLLIIDAE . . . . . . . 1196, 1264, 1293, 1295, 

1298, 1306, 1313 

Proscyllium habereri . . . . . . . . . . . . 1293, 1295 

Protozygaena taylori . . . . . . . . . . . . . . . 1356 

Pseudocarcharias kamoharai . . . . . . . . . . 1268 

PSEUDOCARCHARIIDAE . . . . . . 1196, 1264, 1268


Pseudocarcinus gigas . . . . . . . . . . . . . . . 1055 

PSEUDOTRIAKIDAE . . . . . . . . . . . . 1196, 1296 

Pseudotriakis microdon . . . . . . . . . . . . . . 1296 

Pterygioteuthis . . . . . . . . . . . . . . . . . . . . 781 

pubescens, Ibacus . . . . . . . . . . . . . . 1035-1036 

pubescens, Ibacus ciliatus . . . . . . . . . . . . 1036 

Puerulus . . . . . . . . . . . . . . . . 977, 1006, 1012 

Puerulus angulatus . . . . . . . . . . . . . . . . 1027 

Puerulus velutinus . . . . . . . . . . . . . . . . 1027 

Pugnose caridina . . . . . . . . . . . . . . . . . . . 961 

Pulpo marmóreo . . . . . . . . . . . . . . . . . . . 811 

Pulpo perforado . . . . . . . . . . . . . . . . . . . 810 

Pulpo reticulado . . . . . . . . . . . . . . . . . . . 815 

punctatum, Chiloscyllium . . 1253-1254, 1256-1257 

punctatus, Ovalipes . . . . . . . . . . . . . . . . 1122 

Purple land crab . . . . . . . . . . . . . . . . . . 1150 

Purple mud crab . . . . . . . . . . . . . . . . . . 1128 

Purpleback flying squid . . . . . . . . . . . . . . . 794 

pusillus, Todarodes pacificus . . . . . . . . . . . . 795 

pygmaeus, Idiosepius . . . . . . . . . . . . . . . . 721 

Pygmy cuttlefish . . . . . . . . . . . . . . . . . . . 721 

Pygmy cuttlefishes . . . . . . . . . . . . . . . . . . 688 

Pygmy ribbontail catshark . . . . . . . . . . . . . 1295 

Pygmy shark . . . . . . . . . . . . . . . . . . . . 1228 

Pyroteuthis . . . . . . . . . . . . . . . . . . . . . . 781 

Q 

quadrispinosa, Deania . . . . . . . . . . . . . . 1225 

Queen crab . . . . . . . . . . . . . . . . . . . . . 1055 

Queensland river shark . . . . . . . . . . . . . . 1359 

Quelvacho . . . . . . . . . . . . . . . . . . . . . 1223 

Quelvacho chino . . . . . . . . . . . . . . . . . . 1224 

Quelvacho de aleta corta . . . . . . . . . . . . . 1223 

Quelvacho negro . . . . . . . . . . . . . . . . . . 1224 

quinquedentata, Oratosquillina . . . . . . . . . . 849 

R 

radamae, Carcharhinus . . . . . . . . . . . . . . 1326 

radcliffei, Eridacnis . . . . . . . . . . . . . . . . 1295 

Rainbow shrimp . . . . . . . . . . . . . . . . . . . 944 

Ram’s horn squid . . . . . . . . . . . . . . . . . . . 722 

rancureli, Squalus . . . . . . . . . . . . . . . . . 1231 

Ranina dentata . . . . . . . . . . . . . . . . . . . 1090 

Ranina ranina . . . . . . . . . . . . . . . . 1089-1090 

ranina, Ranina . . . . . . . . . . . . . . . 1089-1090 

RANINIDAE . . . . . . . . . . . . . 1056, 1089, 1115 

Raoulius cultrifer . . . . . . . . . . . . . . . . . . 834 

raphidea, Harpiosquilla . . . . . . . . . . . . . . 841 

rappiana, Sepia . . . . . . . . . . . . . . . . . . . 744 

Rathbun’s vinegar crab . . . . . . . . . . . . . . . 1145 

rayneri, Galeocerdo . . . . . . . . . . . . . . . . 1349 

Rays . . . . . . . . . . . . . . . . . . . . . . . . 1236 

Razor mud shrimp . . . . . . . . . . . . . . . . . . 884 

Reaper cuttlefish . . . . . . . . . . . . . . . . . . . 760 

rectacuta, Penaeopsis . . . . . . . . . . . . . 947-948 

recurvirostra, Sepia . . . . . . . . . . . . . . . . . 750 

Red egg crab . . . . . . . . . . . . . . . . . . . . 1049 

Red reef crab . . . . . . . . . . . . . . . . . . . . 1111 

Red reef lobster. . . . . . . . . . . . . . . . . . . 1000 

Red shrimp . . . . . . . . . . . . . . . . . . . . . . 957 

Red-banded lobster . . . . . . . . . . . . . . . . . 990 

Red-eyed crabs . . . . . . . . . . . . . . . . . . 1049 

Red-spot king prawn . . . . . . . . . . . . . . . . . 919 

Red-spot night octopus . . . . . . . . . . . . . . . 824 

Red-spotted mitten lobster . . . . . . . . . . . . . 1042 

Redtail prawn . . . . . . . . . . . . . . . . . . . . . 923 

Reef box crab . . . . . . . . . . . . . . . . . . . 1097 

Reef crabs . . . . . . . . 1048-1049, 1058, 1098, 1110 

Reef lobsters . . . . . . . . . . . . . . . . . . 978, 995 

Reef odontodactylid mantis shrimp . . . . . . . . . 834 

reesi, Loligo . . . . . . . . . . . . . . . . . . . . . 766 

regalis, Arctides . . . . . . . . . . . . . . . . . . 1041 

remotus, Carcharhinus . . . . . . . . . . . . . . 1331 

Renard . . . . . . . . . . . . . . . . . . . . . . . 1273 

Renard à gros yeux . . . . . . . . . . . . . . . . 1272 

Renard pélagique . . . . . . . . . . . . . . . . . 1271 

repertus, Nautilus . . . . . . . . . . . . . . . . . . 711 

Réquiem babosse . . . . . . . . . . . . . . . . . 1326 

Réquiem de sable . . . . . . . . . . . . . . . . . 1344 

Réquiem macuire . . . . . . . . . . . . . . . . . 1340 

Réquiem océanique . . . . . . . . . . . . . . . . 1341 

Réquiem plombe . . . . . . . . . . . . . . . . . . 1345 

Requiem sharks . . . . . . . . 1196-1197, 1312-1313 

Réquiem soie . . . . . . . . . . . . . . . . . . . . 1335 

Réquiem taureau . . . . . . . . . . . . . . . . . . 1339 

Requiem tisserand . . . . . . . . . . . . . . . . . 1332 

Requin à joues blanches . . . . . . . . . . . . . . 1334 

Requin à longue dorsale . . . . . . . . . . . . . . 1296 

Requin à museau pointu . . . . . . . . . . . . . . 1354 

Requin à nez rude . . . . . . . . . . . . . . . . . 1342 

Requin à tache noire . . . . . . . . . . . . . . . . 1346 

Requin aiguille gris . . . . . . . . . . . . . . . . . 1355 

Requin aiguille réchine . . . . . . . . . . . . . . . 1356 

Requin aveugle des roches . . . . . . . . . . . . 1244 

Requin aveugle gris-bleu . . . . . . . . . . . . . . 1244 

Requin babosse . . . . . . . . . . . . . . . . . . 1326 

Requin baleine . . . . . . . . . . . . . . . . . . . 1263 

Requin baleinier . . . . . . . . . . . . . . . . . . 1336 

Requin balestrine . . . . . . . . . . . . . . . . . 1329 

Requin baliai . . . . . . . . . . . . . . . . . . . . 1338 

Requin bordé . . . . . . . . . . . . . . . . . . . . 1340 

Requin bouledogue . . . . . . . . . . . . . . . . 1339 

Requin carpette à collarette . . . . . . . . . . . . 1242 

Requin carpette à moustache . . . . . . . . . . . 1242 

Requin chat golloum . . . . . . . . . . . . . . . . 1295 

Requin chat gracile . . . . . . . . . . . . . . . . . 1295 

Requin chat pygmé . . . . . . . . . . . . . . . . 1295 

Requin citron faucille . . . . . . . . . . . . . . . . 1352 

Requin corail . . . . . . . . . . . . . . . . . . . . 1358 

Requin crocodile . . . . . . . . . . . . . . . . . . 1268 

Requin cuivre . . . . . . . . . . . . . . . . . . . . 1331 

Requin dagsit . . . . . . . . . . . . . . . . . . . 1328 

Requin de Galapagos . . . . . . . . . . . . . . . 1337 

Requin dormeur à crête . . . . . . . . . . . . . . 1240 

Requin dormeur taureau . . . . . . . . . . . . . . 1240 

Requin dormeur zèbre . . . . . . . . . . . . . . . 1240 

Requin epée . . . . . . . . . . . . . . . . . . . . 1357 

Requin féroce . . . . . . . . . . . . . . . . . . . 1267 

Requin gracile . . . . . . . . . . . . . . . . . . . 1327 

Requin grandes ailes . . . . . . . . . . . . . . . . 1350


Requin gris . . . . . . . . . . . . . . . . . . . . . 1345 

Requin grise . . . . . . . . . . . . . . . . . . . . 1210 

Requin marteau halicorne . . . . . . . . . . . . . 1364 

Requin marteau planeur . . . . . . . . . . . . . . 1363 

Requin nerveux . . . . . . . . . . . . . . . . . . 1333 

Requin océanique . . . . . . . . . . . . . . . . . 1341 

Requin perlon . . . . . . . . . . . . . . . . . . . 1210 

Requin pointe blanche . . . . . . . . . . . . . . . 1325 

Requin pointes noires . . . . . . . . . . . . . . . 1343 

Requin sable tacheté . . . . . . . . . . . . . . . . 1266 

Requin sagrin . . . . . . . . . . . . . . . . . . . 1351 

Requin sombre . . . . . . . . . . . . . . . . . . . 1344 

Requin soyeux . . . . . . . . . . . . . . . . . . . 1335 

Requin tachete . . . . . . . . . . . . . . . . . . . 1347 

Requin taureau . . . . . . . . . . . . . . . . . . . 1266 

Requin tigre . . . . . . . . . . . . . . . . . . . . 1349 

Requin tigre commun . . . . . . . . . . . . . . . 1349 

Requin tisserand . . . . . . . . . . . . . . . . . . 1332 

Requin zèbre . . . . . . . . . . . . . . . . . . . . 1262 

Requin-chabot à taches blanches . . . . . . . . . 1256 

Requin-chabot bambou . . . . . . . . . . . . . . 1257 

Requin-chabot élégant . . . . . . . . . . . . . . . 1255 

Requin-chabot épaulette . . . . . . . . . . . . . . 1258 

Requin-chabot gris . . . . . . . . . . . . . . . . . 1253 

Requin-chabot grivelé . . . . . . . . . . . . . . . 1258 

Requin-chabot marqueterie . . . . . . . . . . . . 1259 

Requin-chabot moine . . . . . . . . . . . . . . . 1259 

Requin-chabot ocellé . . . . . . . . . . . . . . . . 1258 

Requin-hâ . . . . . . . . . . . . . . . . . . . . . 1302 

Requin-hâ aile blanche . . . . . . . . . . . . . . . 1302 

Requin-hâ elegant . . . . . . . . . . . . . . . . . 1303 

Requin-hâ long nez . . . . . . . . . . . . . . . . 1303 

Requin-hâ voile . . . . . . . . . . . . . . . . . . . 1302 

Requin-marteau commun . . . . . . . . . . . . . 1366 

Requin-nourrice fauve . . . . . . . . . . . . . . . 1260 

Requin-tapis barbu . . . . . . . . . . . . . . . . . 1247 

Requin-tapis moustache . . . . . . . . . . . . . . 1247 

Requin-tapis paste . . . . . . . . . . . . . . . . . 1248 

Requin-tapis savetier . . . . . . . . . . . . . . . . 1248 

Requin-tapis tacheté . . . . . . . . . . . . . . . . 1247 

Requin-tigre houareau . . . . . . . . . . . . . . . 1330 

Reticulated swellshark . . . . . . . . . . . . . . . 1289 

RETROPLUMIDAE . . . . . . . . . . . . . . . . . 1048 

rex, Decorisepia . . . . . . . . . . . . . . . . . . . 751 

rex, Sepia . . . . . . . . . . . . . . . . . . . 751, 759 

Rhincodon typus . . . . . . . . . . . . . . . . . 1263 

RHINCODONTIDAE . . . . . . 1196, 1263, 1274, 1314 

Rhiniodon typus . . . . . . . . . . . . . . . . . . 1263 

Rhizoprionodon . . . . . . . . . . . . . . . . . . 1312 

Rhizoprionodon acutus . . . . . . . . . . 1354—1356 

Rhizoprionodon oligolinx . . . . . . . 1354-1355-1356 

Rhizoprionodon taylori . . . . . . . . . . . 1354-1356 

rhoda, Arctosepia . . . . . . . . . . . . . . . . . . 739 

Rhomboid crabs . . . . . . . . . . . . . . . 1058, 1114 

Rhomboid squids . . . . . . . . . . . . . . . . . . . 797 

rhomboidalis, Loligo . . . . . . . . . . . . . . . . 773 

rhombus, Thysanoteuthis . . . . . . . . . . . . . . 797 

Rhynchocinetes durbanensis . . . . . . . . . . . . 971 

Rhynchocinetes uritai . . . . . . . . . . . . . . . . 971 

RHYNCHOCINETIDAE . . . . . . . . . . . . 958, 971 

Ridgeback shrimp . . . . . . . . . . . . . . . . . . 882 

Ridged swimming crab . . . . . . . . . . . . . . . 1121 

Robber harpiosquillid mantis shirmp . . . . . . . . . 841 

Rock shrimps . . . . . . . . . . . . . . . . . . . . . 952 

Roe crabs . . . . . . . . . . . . . . . . . . . . . 1126 

rosea, Metapenaeopsis . . . . . . . . . . . . . . . 931 

Rosecone cuttlefish . . . . . . . . . . . . . . . . . 752 

rosenbergii dacqueti, Macrobrachium . . . . . . . 964 

rosenbergii, Macrobrachium . 855, 957, 964, 966-967 

Roshna prawn . . . . . . . . . . . . . . . . . . . . 965 

Rossia . . . . . . . . . . . . . . . . . . . . . . . . 715 

Rossia australis . . . . . . . . . . . . . . . . . . . 715 

Rossia bipapillata . . . . . . . . . . . . . . . . . . 716 

Rossia pacifica . . . . . . . . . . . . . . . . . . . . 716 

rostrata, Sepia . . . . . . . . . . . . . . . . . . . . 740 

rostridentata, Aristaeomorpha . . . . . . . . . . . 872 

rotundum, Cardisoma . . . . . . . . . . . 1149, 1151 

Rough redeyed crab . . . . . . . . . . . . . . . . 1106 

Rough river prawn . . . . . . . . . . . . . . . . . . 967 

Roussette à taches brunes . . . . . . . . . . . . 1292 

Roussette nuageuse . . . . . . . . . . . . . . . . 1292 

rouxii, Sepia . . . . . . . . . . . . . . . . . . . . 2748 

Royal Spanish lobster . . . . . . . . . . . . . . . 1041 

rozella peregrina, Solitosepia . . . . . . . . . . . . 752 

rozella, Sepia . . . . . . . . . . . . . . . . . . . . 752 

rozella, Solitosepia . . . . . . . . . . . . . . . . . 752 

Rugose land crab . . . . . . . . . . . . . . . . . 1151 

rugosus, Scyllarus . . . . . . . . . . . . . . . . . 1043 

rumphii, Dromia . . . . . . . . . . . . . . . . . . 1087 

rumphii, Menippe . . . . . . . . . . . . . . 1103, 1107 

S 

Saa liu . . . . . . . . . . . . . . . . . . . . . . . . 815 

Sagre lucifer . . . . . . . . . . . . . . . . . . . . 1226 

Sagre porte-feu à queue courte . . . . . . . . . . 1226 

Sailback houndshark . . . . . . . . . . . . . . . . 1302 

Salicoque balafrée . . . . . . . . . . . . . . . . . . 882 

Salicoque canif . . . . . . . . . . . . . . . . . . . . 881 

Salicoque chinoise de vase . . . . . . . . . . . . . 887 

Salicoque des vases côtières . . . . . . . . . . . . 883 

Salicoque peigne . . . . . . . . . . . . . . . . . . . 887 

Sally-light-foots . . . . . . . . . . . . . . . . 1060, 1138 

Saltarelle nez-camus . . . . . . . . . . . . . . . . . 961 

Saltarelle soldat . . . . . . . . . . . . . . . . . . . 960 

Sand bird octopus . . . . . . . . . . . . . . . . . . 815 

Sand crab . . . . . . . . . . . . . . . . . . . . . 1122 

Sand devils . . . . . . . . . . . . . . . . . . . . . 1235 

Sand fish . . . . . . . . . . . . . . . . . . . . . . 1179 

Sand tiger sharks . . . . . . . . . . . . . . . . . 1264 

Sandbar shark . . . . . . . . . . . . . . . . . . . 1345 

Sandtiger shark . . . . . . . . . . . . . . . . . . 1266 

sanguinolentus, Portunus . . . . . . . . . 1115, 1125 

Sargassum spp. . . . . . . . . . . . . . . . . . . 1144 

Saron marmoratus . . . . . . . . . . . . . . . . . . 962 

Saron neglectus . . . . . . . . . . . . . . . . . . . 962 

sauteri, Galeus . . . . . . . . . . . . . . . . . . . 1290 

Saw sharks . . . . . . . . . . . . . . . . . . . . . 1233 

Sawedged spooner . . . . . . . . . . . . . . . . . 1101 

Sawfishes . . . . . . . . . . . . . . . . . . . . . 1233 

scabra var. versicolor, Holothuria (Metriatyla) 1179- 1180


scabra, Holothuria . . . . . . . . . . 1173-1174, 1180 

scabra, Holothuria (Metriatyla) . . . . . . 1179-1180 

Scaeurgus . . . . . . . . . . . . . . . . . . . . . . 801 

Scalloped hammerhead . . . . . . . . . . . . . . 1364 

Scalloped spiny lobster . . . . . . . . . . . . . . . 1016 

scarlatinus, Parribacus . . . . . . . . . . . . . . 1042 

Scarlet shrimp . . . . . . . . . . . . . . . . . . . . 874 

Schizophrys . . . . . . . . . . . . . . . . . . . . 1136 

Schizophrys aspera . . . . . . . . . . . . . . . . 1137 

Schizophrys dama . . . . . . . . . . . . . . . . . 1137 

schultzi, Galeus . . . . . . . . . . . . . . . . . . 1290 

scintillans, Watasenia . . . . . . . . . . . . . . . . 781 

Scoliodon . . . . . . . . . . . . . . . . . . . . . 1312 

Scoliodon acutus . . . . . . . . . . . . . . . 1351, 1354 

Scoliodon ceylonensis . . . . . . . . . . . . . . . 1351 

Scoliodon laticaudus . . . . . . . . 1351, 1354-1357 

Scoliodon palasorra . . . . . . . . . 1354-1355, 1357 

Scoliodon sorrakowa . . . . . . . . . . . . . . . 1357 

Scoliodon walbeehmi . . . . . . . . . . . . . . . 1354 

scorpio, Cloridopsis . . . . . . . . . . . . . . 842, 846 

Scribbled night octopus . . . . . . . . . . . . . . . 813 

scrobiculatus, Nautilus . . . . . . . . . . . . . . . 711 

sculptilis, Parapenaeopsis . . . . . . . . . . 913, 944 

Sculptured mitten lobster . . . . . . . . . . . . . . 1037 

SCYLIORHINIDAE . . . . . . . 1241, 1279, 1293, 1313 

Scyliorhinus garmani . . . . . . . . . . . . . . . 1292 

Scyliorhinus torazame . . . . . . . . . . . . . . 1292 

Scylla . . . . . . . . . . . . . . 1120, 1124, 1126-1127 

Scylla olivacea . . . . . . . . . . . . . . . . 1126-1127 

Scylla paramamosain . . . . . . . . . . . . 1127-1128 

Scylla serrata . . . . . . . . . . . . 1115, 1126-1128 

Scylla serrata var. oceanica . . . . . . . . . . . . 1126 

Scylla tranquebarica . . . . . . . . . . . . 1127-1128 

SCYLLARIDAE 977, 979, 984, 997, 1002, 1008, 1028 

Scyllarides . . . . . . . . . . . . . . . 977, 1028, 1032 

Scyllarides haanii . . . . . . . . . . . . . . . . . 1038 

Scyllarides squammosus . . . . . . . . . . . . . 1039 

Scyllarus . . . . . . . . . . . . . . . . . . . 1028, 1030 

Scyllarus bertholdii . . . . . . . . . . . . . . . . 1043 

Scyllarus cultrifer . . . . . . . . . . . . . . . . . 1028 

Scyllarus rugosus . . . . . . . . . . . . . . . . . 1043 

scyllarus, Odontodactylus . . . . . . . . . . 829, 834 

Scyllium ornatum . . . . . . . . . . . . . . . . . 1256 

Scymnodon . . . . . . . . . . . . . . . . . . . . . 1213 

Scymnodon squamulosus . . . . . . . . . . . . . 1228 

Sea cucumbers . . . . . . . . . . . . 1158, 1163, 1180 

Sea stars . . . . . . . . . . . . . . . . . . . . . . 1158 

Sea urchins . . . . . . . . . . . . . . . . . . . . . 1158 

sealei, Carcharhinus . . . . . . . . . . . . 1334, 1346 

sebana, Eriphia . . . . . . . . . . . . . . . 1106, 1108 

sedili, Trachypenaeus . . . . . . . . . . . . . 928, 950 

Seiche aiguille . . . . . . . . . . . . . . . . . . . . 736 

Seiche andreana . . . . . . . . . . . . . . . . . . . 737 

Seiche baisers . . . . . . . . . . . . . . . . . . . . 745 

Seiche dorée . . . . . . . . . . . . . . . . . . . . . 742 

Seiche géante . . . . . . . . . . . . . . . . . . . . 738 

Seiche gracile . . . . . . . . . . . . . . . . . . . . 739 

Seiche grandes mains . . . . . . . . . . . . . . . . 744 

Seiche hameçon . . . . . . . . . . . . . . . . . . . 750 

Seiche kobi . . . . . . . . . . . . . . . . . . . . . . 743 

Seiche madokai . . . . . . . . . . . . . . . . . . . 759 

Seiche moisson . . . . . . . . . . . . . . . . . . . 760 

Seiche petites mains . . . . . . . . . . . . . . . . . 740 

Seiche pharaon . . . . . . . . . . . . . . . . . . . 748 

Selenka’s sea cucumber . . . . . . . . . . . . . . 1187 

semisulcatus exsulcatus, Penaeus . . . . . . . . . . 922 

semisulcatus paucidentatus, Penaeus . . . . . . . . 925 

semisulcatus, Penaeus . . . . . . . . . . . . 915, 925 

Sentinel crab . . . . . . . . . . . . . . . . . . . . 1123 

Sepia (Sepiella) microcheirus . . . . . . . . . . . . 756 

Sepia aculeata . . . . . . . . . . . . . . . . . . . . 736 

Sepia acuminata . . . . . . . . . . . . . . . . . . . 759 

Sepia affinis . . . . . . . . . . . . . . . . . . . . . 756 

Sepia andreana . . . . . . . . . . . . . . . . 737, 743 

Sepia andreanoides . . . . . . . . . . . . . . . . . 743 

Sepia apama . . . . . . . . . . . . . . . . . . . . . 738 

Sepia bandensis . . . . . . . . . . . . . . . . . . . 757 

Sepia bartletti . . . . . . . . . . . . . . . . . . . . 758 

Sepia braggi . . . . . . . . . . . . . . . . . . . . . 739 

Sepia brevimana . . . . . . . . . . . . 740, 754, 761 

Sepia con punta . . . . . . . . . . . . . . . . . . . 736 

Sepia cottoni . . . . . . . . . . . . . . . . . . . . . 758 

Sepia cultrata . . . . . . . . . . . . . . . . . . . . 759 

Sepia dorada . . . . . . . . . . . . . . . . . . . . . 742 

Sepia elliptica . . . . . . . . . . . . . . . . . 741-742 

Sepia esculenta . . . . . . . . . . . . . . 740-742, 748 

Sepia faraónica . . . . . . . . . . . . . . . . . . . . 748 

Sepia formosana . . . . . . . . . . . . . . . . . . . 748 

Sepia galei . . . . . . . . . . . . . . . . . . . . . . 747 

Sepia ganchuda . . . . . . . . . . . . . . . . . . . 750 

Sepia gigante . . . . . . . . . . . . . . . . . . . . 738 

Sepia grácil . . . . . . . . . . . . . . . . . . . . . . 739 

Sepia hedleyi . . . . . . . . . . . . . . . . . . . . . 751 

Sepia hercules . . . . . . . . . . . . . . . . . . . . 744 

Sepia indica . . . . . . . . . . . . . . . . . . . . . 736 

Sepia inerme . . . . . . . . . . . . . . . . . . . . . 756 

Sepia kobí . . . . . . . . . . . . . . . . . . . . . . 743 

Sepia kobiensis . . . . . . . . . . . . . . . . 737, 743 

Sepia kobiensis var. albatrossi . . . . . . . . . . . 743 

Sepia kobiensis var. andreanoides . . . . . . . . . 743 

Sepia kobiensis var. beppauna . . . . . . . . . . . 743 

Sepia kobiensis var. crassa . . . . . . . . . . . . . 743 

Sepia kobiensis var. toyamensis . . . . . . . . . . . 743 

Sepia labiada . . . . . . . . . . . . . . . . . . . . . 745 

Sepia latimanus . . . . . . . . . . . . . . . . . . . 744 

Sepia lycidas . . . . . . . . . . . . . . . . . . . . . 745 

Sepia madokai . . . . . . . . . . . . . . . . . 751, 759 

Sepia mazicorta . . . . . . . . . . . . . . . . . . . 740 

Sepia mazuda . . . . . . . . . . . . . . . . . . . . 744 

Sepia mestus . . . . . . . . . . . . . . . . . . . . . 760 

Sepia mira . . . . . . . . . . . . . . . . . . . . . . 760 

Sepia mozambica . . . . . . . . . . . . . . . . . . 744 

Sepia officinalis . . . . . . . . . . . . . . . . . . . 725 

Sepia omani . . . . . . . . . . . . . . . . . . . . . 762 

Sepia opipara . . . . . . . . . . . . . . . . . . . . 746 

Sepia pagenstecheri . . . . . . . . . . . . . . . . . 750 

Sepia palmata . . . . . . . . . . . . . . . . . . . . 738 

Sepia papuensis . . . . . . . . . . . . . . . . . . . 747 

Sepia pharaonis . . . . . . . . . . . . . . . . 744, 748 

Sepia plangon . . . . . . . . . . . . . . . . . . . . 749


Sepia prionota . . . . . . . . . . . . . . . . . . . . 747 

Sepia rappiana . . . . . . . . . . . . . . . . . . . . 744 

Sepia recurvirostra . . . . . . . . . . . . . . . . . 750 

Sepia rex . . . . . . . . . . . . . . . . . . . . 751, 759 

Sepia rostrata . . . . . . . . . . . . . . . . . . . . 740 

Sepia rouxii . . . . . . . . . . . . . . . . . . . . . 748 

Sepia rozella . . . . . . . . . . . . . . . . . . . . . 752 

Sepia segadora . . . . . . . . . . . . . . . . . . . 760 

Sepia singaporensis . . . . . . . . . . . . . . . . . 750 

Sepia smithi . . . . . . . . . . . . . . . . . . 753, 755 

Sepia stellifera . . . . . . . . . . . . . . . 740-741, 754 

Sepia subaculeata . . . . . . . . . . . . . . . . . . 745 

Sepia sulcata . . . . . . . . . . . . . . . . . . . . . 761 

Sepia tigris . . . . . . . . . . . . . . . . . . . . . . 748 

Sepia vietnamica . . . . . . . . . . . . . . . . . . 761 

Sepia vossi . . . . . . . . . . . . . . . . . . . . . . 762 

Sepia whitleyana . . . . . . . . . . . . . . . 753, 755 

SEPIADARIIDAE . . . . . . . . . . . . . . . . 712, 719 

Sepiadariids . . . . . . . . . . . . . . . . . . . . . 696 

Sepiadarium kochii . . . . . . . . . . . . . . 694, 720 

Sepiadarium malayense . . . . . . . . . . . . . . . 720 

Sepiella inermis . . . . . . . . . . . . . . . . . . . 756 

Sepiella mandroni . . . . . . . . . . . . . . . . . . 756 

Sepiella ocellata . . . . . . . . . . . . . . . . . . . 762 

Sepiella ornata . . . . . . . . . . . . . . . . . . . . 762 

Sepiella weberi . . . . . . . . . . . . . . . . . . . 763 

SEPIIDAE . . . . . . . . . . . . . . . . 689, 723, 765 

Sepioids . . . . . . . . . . . . . . . . . . . . 694, 696 

Sepiola birostrata . . . . . . . . . . . . . . . 716-717 

Sepiola mariposa . . . . . . . . . . . . . . . . . . . 716 

Sepiola parva . . . . . . . . . . . . . . . . . . . . 717 

Sepiola trirostrata . . . . . . . . . . . . . . . 716-717 

Sépiole gros yeux . . . . . . . . . . . . . . . . . . 718 

Sépiole papillon . . . . . . . . . . . . . . . . . . . 716 

SEPIOLIDAE . . . . . . . . . . . . . . . . . . 712, 719 

Sepiolids . . . . . . . . . . . . . . . . . . . . . . . 696 

Sepiolina . . . . . . . . . . . . . . . . . . . . . . . 718 

Sepiolina nipponensis . . . . . . . . . . . . . . . . 718 

Sepioloidea lineolata . . . . . . . . . . . . . . . . 720 

Sepioteuthis . . . . . . . . . . . . . . . 725, 765, 797 

Sepioteuthis arctipinnis . . . . . . . . . . . . . . . 778 

Sepioteuthis lessoniana . . . . 688-689, 765, 778, 797 

SERGESTIDAE . . . . . 858, 860, 869, 876, 891, 953 

Sergestoid shrimps . . . . . . . . . . . . . 855-856, 858 

SERGESTOIDEA . . . . . 855-856, 858, 866, 956, 958 

serrata var. oceanica, Scylla . . . . . . . . . . . 1126 

serrata, Scyllav . . . . . . . . . . . 1115, 1126-1128 

serrulatus, Acetes . . . . . . . . . . . . . . . . . . 864 

SESARMINAE . . . . . . . . . . . . . . . . . . . 1048 

Sevengill sharks . . . . . . . . . . . . . . . . . . 1208 

Sharpnose sevengill shark . . . . . . . . . . . . . 1210 

Sharpnose sharks . . . . . . . . . . . . . . . . . 1312 

Short-spined nylon shrimp . . . . . . . . . . . . . . 970 

Shortclub cuttlefish . . . . . . . . . . . . . . . . . . 740 

Shortfin mako . . . . . . . . . . . . . . . . . . . 1277 

Shortleg river prawn . . . . . . . . . . . . . . . . . 968 

Shortnose spurdog . . . . . . . . . . . . . . . . . 1230 

Shortspine spurdog . . . . . . . . . . . . . . . . 1230 

Shorttail lanternshark . . . . . . . . . . . . . . . 1226 

Siboga lobster . . . . . . . . . . . . . . . . . . . . 989 

sibogae australiensis, Haliporoides . . . . . . . . . 881 

sibogae madagascariensis, Haliporoides . . . . . . 881 

sibogae, Acetes . . . . . . . . . . . . . . . . . . . 864 

sibogae, Apristurus . . . . . . . . . . . . . . . . 1286 

sibogae, Doryteuthis . . . . . . . . . . . . . . 776-777 

sibogae, Haliporoides . . . . . . . . . . . . . . . . 881 

sibogae, Heterocarpus . . . . . . . . . . . . 969-970 

sibogae, Hymenopenaeus . . . . . . . . . . . . . . 881 

sibogae, Loligo . . . . . . . . . . . . . . . . . . . 777 

sibogae, Metanephrops . . . . . . . . . . . . . . . 989 

sibogae, Nephrops . . . . . . . . . . . . . . . . . . 989 

Sicklefin lemon shark . . . . . . . . . . . . . . . 1352 

Sicklefin weasel shark . . . . . . . . . . . . . . . 1309 

Sicyonella . . . . . . . . . . . . . . . . . . . . . . 858 

Sicyonia cristata . . . . . . . . . . . . . . . . . . . 954 

Sicyonia lancifera . . . . . . . . . . . . . . . 952, 954 

SICYONIIDAE . . . . . . . . . 855, 869, 876, 890, 952 

signatus, Atergatopsis . . . . . . . . . . . . . . . 1102 

silasi, Penaeus . . . . . . . . . . . . 916, 921, 923, 926 

Silky shark . . . . . . . . . . . . . . . . . . 1335, 1341 

Sillago . . . . . . . . . . . . . . . . . . . . . . . 1333 

Silvertip shark . . . . . . . . . . . . . . . . . . . 1325 

similis, Bohadschia . . . . . . . . . . . . . 1173-1174 

sinensis, Eriocheir . . . . . . . . . . . . . . . . . 1055 

sinensis, Metanephrops . . . . . . . . . . . . . . . 993 

sinensis, Solenocera . . . . . . . . . . . . . . . . . 883 

Singapore rough shrimp . . . . . . . . . . . . . . . 950 

Singapore vinegar crab . . . . . . . . . . . . . . 1146 

singaporense, Episesarma . . . . . . . . . 1143, 1146 

singaporensis, Sepia . . . . . . . . . . . . . . . . . 750 

singhalensis, Doryteuthis . . . . . . . . . . . . . . 774 

singhalensis, Photololigo . . . . . . . . . . . . . . 777 

Sixgill sharks . . . . . . . . . . . . . . . . . . . . 1208 

Slender bambooshark . . . . . . . . . . . . . . . 1255 

Slender cuttlefish . . . . . . . . . . . . . . . . . . . 739 

Slender sawtail catshark . . . . . . . . . . . . . . 1290 

Slender smooth-hound . . . . . . . . . . . . . . . 1295 

Slendertail lanternshark . . . . . . . . . . . . . . 1226 

Slipper lobster . . . . . . . . . . . . . . . . . . . 1038 

Slipper lobsters . . . . . . . . . . . . . . 977, 979, 1028 

Sliteye shark . . . . . . . . . . . . . . . . . . . . 1351 

sloani pacificus, Ommastrephes . . . . . . . . . . . 795 

Small furrow lobster . . . . . . . . . . . . . . . . 1022 

Small-spot night octopus . . . . . . . . . . . . . . . 814 

Smalleye pigmy shark . . . . . . . . . . . . . . . 1229 

Smalleyed squillid mantis shrimp . . . . . . . . . . 847 

Smallfin gulper shark . . . . . . . . . . . . . . . . 1223 

Smalltooth sand tiger . . . . . . . . . . . . . . . . 1267 

Smelt-whiting . . . . . . . . . . . . . . . . . . . . 1333 

Smith’s cuttlefish . . . . . . . . . . . . . . . . . . . 753 

smithi, Sepia . . . . . . . . . . . . . . . . . . 753, 755 

smithii, Eriphia . . . . . . . . . . . . . . . 1106, 1108 

Smooth fan lobster . . . . . . . . . . . . . . . . . 1035 

Smooth hammerhead . . . . . . . . . . . . . . . 1366 

Smooth redeyed crab . . . . . . . . . . . . . . . 1108 

Smooth spooner . . . . . . . . . . . . . . . . . . 1100 

Smooth squillid mantis shrimp . . . . . . . . . . . . 846 

Smooth stone crab . . . . . . . . . . . . . . . . . 1109 

Smooth-shelled swimming crab . . . . . . . . . . 1130 

Smoothhounds . . . . . . . . . . . . . . . . . . . 1297


Smoothshell shrimp . . . . . . . . . . . . . . . . . 945 

Snaggletooth shark . . . . . . . . . . . . . . . . 1310 

Snake fish . . . . . . . . . . . . . . . . . . . . . 1175 

Snow crab . . . . . . . . . . . . . . . . . . . . . 1055 

Soldier brush shrimp . . . . . . . . . . . . . . . . . 960 

Soldier crabs . . . . . . . . . . . . . . . . . . . . 1048 

Solenocera . . . . . . . . . . . . . . . . . . . . . . 875 

Solenocera alfonso . . . . . . . . . . . . . . 884-885 

Solenocera alticarinata . . . . . . . . . 882, 885, 887 

Solenocera australiana . . . . . . . . . . . . 884, 886 

Solenocera choprai . . . . . . . . . . . 882, 885, 887 

Solenocera crassicornis . . . . . . . . . . . . . . . 883 

Solenocera halli . . . . . . . . . . . . . . . . 884, 886 

Solenocera indicus . . . . . . . . . . . . . . . . . . 883 

Solenocera koelbeli . . . . . . . . . . . . . . 884, 887 

Solenocera kuboi . . . . . . . . . . . . . . . . . . 883 

Solenocera melantho . . . . . . . . . . . 884, 886-887 

Solenocera pectinata . . . . . . . . . . . . . 887-888 

Solenocera pectinulata . . . . . . . . . . . . 887-888 

Solenocera prominentis . . . . . . . . . . . . . . . 884 

Solenocera sinensis . . . . . . . . . . . . . . . . . 883 

Solenocera sp. . . . . . . . . . . . . . . . . . . . . 882 

Solenocera subnuda . . . . . . . . . . . . . . . . . 883 

Solenocerid shrimps . . . . . . . . . . . . . . . . . 875 

SOLENOCERIDAE . . . . . . 855, 869, 875, 890, 953 

solicitans, Oratosquillina . . . . . . . . . . . . . . 849 

Solitosepia genista . . . . . . . . . . . . . . . . . . 747 

Solitosepia lana . . . . . . . . . . . . . . . . . . . 747 

Solitosepia occidua . . . . . . . . . . . . . . . . . 747 

Solitosepia plangon adhaesa . . . . . . . . . . . . 749 

Solitosepia rozella . . . . . . . . . . . . . . . . . . 752 

Solitosepia rozella peregrinav . . . . . . . . . . . . 752 

Solitosepia submestus . . . . . . . . . . . . . . . . 747 

Solrayo . . . . . . . . . . . . . . . . . . . . . . . 1267 

sordidus, Linuparus . . . . . . . . . . . . . . . . 1024 

sorrah, Carcharhinus . . . . . . . . . . . . . . . 1347 

sorrakowa, Scoliodon . . . . . . . . . . . . . . . 1357 

Southern bobtail squid . . . . . . . . . . . . . . . . 714 

Southern mauxia shrimp . . . . . . . . . . . . . . . 864 

Southern rough shrimp . . . . . . . . . . . . . . . . 927 

Southern velvet shrimp . . . . . . . . . . . . . . . . 908 

Spadenose shark . . . . . . . . . . . . . . . . . . 1357 

Spanner crab . . . . . . . . . . . . . . . . . . . . 1090 

Spanner crabs . . . . . . . . . . . . 1056, 1089, 1115 

Spear shrimp . . . . . . . . . . . . . . . . . . . . . 913 

Speartooth shark . . . . . . . . . . . . . . . . . . 1359 

Speckled carpetshark . . . . . . . . . . . . . . . 1259 

Speckled catshark . . . . . . . . . . . . . . . . . 1291 

Spectacled box crab . . . . . . . . . . . . . . . . 1097 

Sphyrna blochii . . . . . . . . . . . . . . . . . . 1363 

Sphyrna diplana . . . . . . . . . . . . . . . . . . 1364 

Sphyrna lewini . . . . . . . . . . . . . . . 1364-1366 

Sphyrna mokarran . . . . . . . . . . 1364-1365-1366 

Sphyrna tudes . . . . . . . . . . . . . . . . . . . 1365 

Sphyrna zygaena . . . . . . . . . . . . . . 1364-1366 

Sphyrnid sharks . . . . . . . . . . . . . . . . . . 1197 

SPHYRNIDAE . . . . . . . . . 1196-1197, 1314, 1361 

Spider crabs . . . . . . . . . . 1050, 1059, 1083, 1136 

Spider prawn . . . . . . . . . . . . . . . . . . . . . 968 

Spindle crabs . . . . . . . . . . . . . . . . . . . . 1083 

spinea, Actinopyga . . . . . . . . . . . . . 1169-1171 

Spined pygmy shark . . . . . . . . . . . . . . . . 1229 

Spineless cuttlefish . . . . . . . . . . . . . . . . . . 756 

spinimana, Thalamita . . . . . . . . . . . . . . . 1129 

spinipes, Atyopsis . . . . . . . . . . . . . . . . . . 960 

Spinner shark . . . . . . . . . . . . . . . . . . . 1332 

Spiny claw swimming crab . . . . . . . . . . . . . 1129 

Spiny greasyback shrimp . . . . . . . . . . . . . . 909 

Spiny lobster . . . . . . . . . . . . . . . . . . . . . 975 

Spiny lobsters . . . 977, 1005-1006, 1037-1038, 1040 

Spiny spooner . . . . . . . . . . . . . . . . . . . 1102 

Spirula spirula . . . . . . . . . . . . . . . . . . . 722 

spirula, Spirula . . . . . . . . . . . . . . . . . . . 722 

SPIRULIDAE . . . . . . . . . . . . . . . . . . . . . 722 

Splendid lanternshark . . . . . . . . . . . . . . . 1227 

Splendid spooner . . . . . . . . . . . . . . . . . 1101 

splendidus, Etisus . . . . . . . . . . . . . . . . . 1101 

splendidus, Etmopterus . . . . . . . . . . . . . . 1227 

Sponge crabs . . . . . . . . . . . . 1056, 1083, 1085 

Spongehead catshark . . . . . . . . . . . . . . . 1287 

spongiceps, Apristurus . . . . . . . . . . . . . . 1287 

Spongicola venusta . . . . . . . . . . . . . . . . . 955 

Spooner crabs . . . . . . . . . . . . . . . . . . . 1098 

Spotless smooth-hound . . . . . . . . . . . . . . 1304 

Spottail shark . . . . . . . . . . . . . . . . . . . . 1347 

Spotted marbled shrimp . . . . . . . . . . . . . . . 962 

Spotted reef crab . . . . . . . . . . . . . . . . . . 1111 

Spotted squillid mantis shrimp . . . . . . . . . . . . 846 

Spotted wobbegong . . . . . . . . . . . . . . . . 1247 

Spottedbelly rock crab . . . . . . . . . . . . . . . 1109 

Spotty bobtail squid . . . . . . . . . . . . . . . . . 717 

Spotty cuttlefish . . . . . . . . . . . . . . . . . . . 762 

Squale bouclé . . . . . . . . . . . . . . . . . . . 1212 

Squale bouclé du Pacifique . . . . . . . . . . . . 1212 

Squale liche . . . . . . . . . . . . . . . . . . . . 1225 

Squale moustache . . . . . . . . . . . . . . . . . 1224 

Squale nain . . . . . . . . . . . . . . . . . . . . . 1229 

Squale pygmée . . . . . . . . . . . . . . . . . . . 1228 

Squale-chagrin cagaou . . . . . . . . . . . . . . 1223 

Squale-chagrin commun . . . . . . . . . . . . . . 1223 

Squale-chagrin de l’Atlantique . . . . . . . . . . . 1224 

Squale-chagrin quelvacho . . . . . . . . . . . . . 1224 

Squale-grogneur velouté . . . . . . . . . . . . . . 1228 

Squale-savate à long nez . . . . . . . . . . . . . 1225 

Squale-savate lutin . . . . . . . . . . . . . . . . . 1225 

Squalelet féroce . . . . . . . . . . . . . . . . . . 1228 

SQUALIDAE . . . . . . . 1196-1197, 1211, 1213, 1314 

Squaliolus . . . . . . . . . . . . . . . . . . . . . 1213 

Squaliolus aliae . . . . . . . . . . . . . . . . . . 1229 

Squaliolus laticaudus . . . . . . . . . . . . . . . 1229 

Squalus brevirostris . . . . . . . . . . . . . . . . 1230 

Squalus japonicus . . . . . . . . . . . . . . . . . 1229 

Squalus megalops . . . . . . . . . . . . . . . . . 1230 

Squalus melanurus . . . . . . . . . . . . . . . . 1230 

Squalus mitsukurii . . . . . . . . . . . . . . . . 1230 

Squalus rancureli . . . . . . . . . . . . . . . . . 1231 

Squalus sp. A . . . . . . . . . . . . . . . . . . . 1231 

Squalus sp. B . . . . . . . . . . . . . . . . . . . 1231 

Squalus sp. F . . . . . . . . . . . . . . . . . . . 1232 

squammosus, Scyllarides . . . . . . . . . . . . . 1039


squamosus, Centrophorus . . . . . . . . . . . . 1224 

squamulosus, Scymnodon . . . . . . . . . . . . 1228 

Square-shelled crab . . . . . . . . . . . . . . . . 1113 

Squat lobsters . . . . . . . . . . . . . . . . 976, 1048 

Squatina australis . . . . . . . . . . . . . . . . . 1237 

Squatina japonica . . . . . . . . . . . . . . . . . 1237 

Squatina sp. A . . . . . . . . . . . . . . . . . . . 1237 

SQUATINIDAE . . 1196, 1211, 1214, 1235, 1246, 1314 

Squid . . . . . . . . . . . . . . . . . . . . . . . . . 692 

Squids . . . . . . . . . . . . . . . . 688, 694, 696, 698 

Squillid mantis shrimps . . . . . . . . . . . . . 830, 842 

SQUILLIDAE . . . . . . . . . . . . . . . . . . 830, 842 

Squillids . . . . . . . . . . . . . . . . . . . . . . . 842 

Sri Lanka crab . . . . . . . . . . . . . . . . . . . 1127 

Staregaze cuttlefish . . . . . . . . . . . . . . . . . 746 

Starry cuttlefish . . . . . . . . . . . . . . . . . . . . 754 

Starspotted smooth-hound . . . . . . . . . . . . . 1304 

Steenstrup’s bay squid . . . . . . . . . . . . . . . . 780 

Stegostoma fasciatum . . . . . . . . . . . . . . . 1262 

Stegostoma tygrinus . . . . . . . . . . . . . . . . 1262 

Stegostoma varium . . . . . . . . . . . . . . . . . 1262 

STEGOSTOMATIDAE . . 1197, 1250, 1261-1262, 1269 

stellifera, Sepia . . . . . . . . . . . . . . 740-741, 754 

stenodactyla, Ommastrephes caroli . . . . . . . . 793 

stenodactylus, Atypopenaeus . . . . . . . . . . . . 929 

stenomphalus, Nautilus . . . . . . . . . . . . . . . 711 

Stenopodid shrimps . . . . . . . . . . . . . . 856, 955 

STENOPODIDAE . . . . 855, 870, 877, 891, 953, 955 

STENOPODIDEA . . . . 855-856, 859, 866, 955, 958 

Stenopodids . . . . . . . . . . . . . . . . . . . . . 855 

Stenopus . . . . . . . . . . . . . . . . . . . . . . . 955 

Stenopus hispidus . . . . . . . . . . . . . . . . . . 955 

stewarti, Nephropsis . . . . . . . . . . . . . . . . . 991 

Sthenoteuthis oualaniensis . . 694, 689, 788, 793-794 

STICHOPODIDAE . . . . . . . . . . . . . . 1160, 1185 

Stichopus chloronotus . . . . . . . . . . . . . . 1186 

Stichopus godeffroyi . . . . . . . . . . . . . . . . 1187 

Stichopus horrens . . . . . . . . . . . . . . . . . 1187 

Stichopus variegatus . . . . . . . . . . . . . . . 1188 

stimpsoni, Panulirus . . . . . . . . . . . . . . . 1026 

Stingrays . . . . . . . . . . . . . . . . . . . . . . 1196 

STOMATOPODA . . . . . . . . . . . . . . . . . . . 855 

Stone crabs . . . . . . . 1048, 1057, 1098, 1103, 1110 

Stork shrimp . . . . . . . . . . . . . . . . . . . . . 941 

Stout red shrimp . . . . . . . . . . . . . . . . . . . 873 

strahani, Hemiscyllium . . . . . . . . . . . . . . 1259 

Straight-tooth weasel shark . . . . . . . . . . . . 1311 

stridulans, Metapenaeopsis . . . . . . . . . . 908, 932 

strigosus, Grapsus . . . . . . . . . . . . . . . . . 1142 

Striking cuttlefish . . . . . . . . . . . . . . . . . . . 749 

striolatus, Octopus . . . . . . . . . . . . . . . . . . 815 

Striped dumpling squid . . . . . . . . . . . . . . . . 720 

Striped hinge-beak shrimp . . . . . . . . . . . . . . 971 

Striped reef lobster . . . . . . . . . . . . . . . . . . 999 

Stumpy bobtail squid . . . . . . . . . . . . . . . . . 715 

Stumpy spined cuttlefish . . . . . . . . . . . . . . . 757 

styliferus, Exopalaemon . . . . . . . . . . . . . . 965 

subaculeata, Sepia . . . . . . . . . . . . . . . . . . 745 

sublevis, Phoberus caecus . . . . . . . . . . . . . . 988 

submestus, Solitosepia . . . . . . . . . . . . . . . . 747 

subnuda, Solenocera . . . . . . . . . . . . . . . . 883 

sulcata, Sepia . . . . . . . . . . . . . . . . . . . . 761 

Sulu shrimp . . . . . . . . . . . . . . . . . . . . . . 940 

suluensis, Metapenaeus . . . . . . . . . . . . . . . 940 

sumatrensis, Nipponololigo . . . . . . . . . . . . 773 

Sundaic paddler crab . . . . . . . . . . . . . . . 1144 

SUNDATHELPHUSIDAE . . . . . . . . . . 1050, 1147 

superciliosus, Alopias . . . . . . . . . 1271-1272-1273 

Surf redfish . . . . . . . . . . . . . . . . . . . . . 1168 

Swimming crabs . . . . . . . . . . . . . . . 1059, 1115 

Swordtip squid . . . . . . . . . . . . . . . . . . . . 776 

Symplectoteuthis oualaniensis . . . . . . . . . . . 794 

SYNAPTIDAE . . . . . . . . . . . . . . . . . . . 1160 

SYNAXIDAE . . . . . . 977, 979, 983, 996, 1001, 1008 

T 

Taiwan gulper shark . . . . . . . . . . . . . . . . 1224 

Taiwan mauxia shrimp . . . . . . . . . . . . . . . . 863 

Taningia . . . . . . . . . . . . . . . . . . . . . . . 797 

Tapicero barbudo . . . . . . . . . . . . . . . . . . 1247 

Tapicero japonés . . . . . . . . . . . . . . . . . . 1247 

Tapicero manchado . . . . . . . . . . . . . . . . 1247 

Tapicero ornamentado . . . . . . . . . . . . . . . 1248 

Tapicero zapatilla . . . . . . . . . . . . . . . . . . 1248 

tasmanica, Eupryma . . . . . . . . . . . . . . . . 714 

Tasselled wobbegong . . . . . . . . . . . . . . . 1247 

Taupe bleu . . . . . . . . . . . . . . . . . . . . . 1277 

taurus, Carcharias . . . . . . . . . . 1196, 1266-1267 

taurus, Eugomphodus . . . . . . . . . . . . . . . 1266 

taurus, Odontaspis . . . . . . . . . . . . . . . . . 1266 

Tawny nurse shark . . . . . . . . . . . . . . . . . 1260 

taylori, Rhizoprionodon . . . . . . . . . . . 1354-1356 

taylori, Protozygaena . . . . . . . . . . . . . . . 1356 

TELMESSINAE . . . . . . . . . . . . . . . . . . 1050 

temmincki, Carcharhinus . . . . . . . . . . . . . 1350 

temmincki, Eulamia . . . . . . . . . . . . . . . . 1350 

temmincki, Lamiopsis . . . . . . . . . . . . 1350, 1352 

tenella, Parapenaeopsis . . . . . . . . . . . . . . . 945 

tengi, Negogaleus . . . . . . . . . . . . . . . . . 1311 

tengi, Paragaleus . . . . . . . . . . . . . . . . . 1311 

tenuicrustatus, Grapsus . . . . . . . . . . . . . . 1142 

tenuimana, Acanthacaris . . . . . . . . . . . . . . 988 

tenuimanus, Phoberus . . . . . . . . . . . . . . . . 988 

tenuipes, Metapenaeus . . . . . . . . . . . . . . . 941 

tenuipes, Nematopalaemon . . . . . . . . . . . . . 968 

teraoi, Penaeus . . . . . . . . . . . . . . . . . . . 920 

tetricus, Octopus . . . . . . . . . . . . . . . . . . 818 

TEUTHIDA . . . . . . . . . . . . . . . . . . . . . . 688 

Teuthoids . . . . . . . . . . . . . . . . . . . . . . . 694 

Thai vinegar crab . . . . . . . . . . . . . . . . . . 1145 

Thalamita . . . . . . . . . . . . . . . . . . . . . 1129 

Thalamita crenata . . . . . . . . . . . . . . . . . 1129 

Thalamita spinimana . . . . . . . . . . . . . . . 1129 

Thalassina . . . . . . . . . . . . . . . . . . . . . 1143 

Thalassina anomala . . . . . . . . . . . . . . . . 1048 

Thalassina spp. . . . . . . . . . . . . . . . . . . 1048 

THALASSINIDAE . . . . . . . . . . . . . . 976, 1048 

THALASSINIDEA . . . . . . . . . . . . . . . 855, 976 

THAUMASTOCHELIDAE . . 977, 983, 996, 1001, 1007 

Thelenota ananas . . . . . . . . . . . . . . . . . 1189


Thelenota anax . . . . . . . . . . . . . . . . . . 1190 

Thenus . . . . . . . . . . . . . . . . . . . . 977, 1028 

Thenus orientalis . . . . . . . . . . . . . . . . . 1040 

THIIDAE . . . . . . . . . . . . . . . . . . . . . . 1050 

Thin shrimp . . . . . . . . . . . . . . . . . . . . . . 943 

thomsoni, Metanephrops . . . . . . . . . . . . . . 990 

thomsoni, Nephrops . . . . . . . . . . . . . . . . . 990 

Three-spot swimming crab . . . . . . . . . . . . . 1125 

Thresher shark . . . . . . . . . . . . . . . . . . . 1273 

Thresher sharks . . . . . . . . . . . . . . . 1197, 1269 

THYSANOTEUTHIDAE . . . . . . . . . . . . . . . 797 

Thysanoteuthis . . . . . . . . . . . . . . . . . . . . 797 

Thysanoteuthis nuchalis . . . . . . . . . . . . . . . 797 

Thysanoteuthis rhombus . . . . . . . . . . . . . . 797 

Tiburón acebrado . . . . . . . . . . . . . . . . . 1262 

Tiburón aleta negra . . . . . . . . . . . . . . . . 1332 

Tiburón aleton . . . . . . . . . . . . . . . . . . . 1350 

Tiburón alinegro . . . . . . . . . . . . . . . . . . 1346 

Tiburón arenero . . . . . . . . . . . . . . . . . . 1344 

Tiburón azul . . . . . . . . . . . . . . . . . . . . 1353 

Tiburón baboso . . . . . . . . . . . . . . . . . . . 1326 

Tiburón baleta . . . . . . . . . . . . . . . . . . . 1329 

Tiburón ballena . . . . . . . . . . . . . . . . . . . 1263 

Tiburón ballenero . . . . . . . . . . . . . . . . . . 1336 

Tiburón cariblanco . . . . . . . . . . . . . . . . . 1334 

Tiburón ciego de roca . . . . . . . . . . . . . . . 1244 

Tiburón ciego gris . . . . . . . . . . . . . . . . . 1244 

Tiburón cobrizo . . . . . . . . . . . . . . . . . . . 1331 

Tiburón cocodrilo . . . . . . . . . . . . . . . . . . 1268 

Tiburón de arrecifes . . . . . . . . . . . . . . . . 1328 

Tiburón de Borneo . . . . . . . . . . . . . . . . . 1330 

Tiburón de clavos . . . . . . . . . . . . . . . . . 1212 

Tiburón de Galápagos . . . . . . . . . . . . . . . 1337 

Tiburón de Milberto . . . . . . . . . . . . . . . . 1345 

Tiburón de Pondicherry . . . . . . . . . . . . . . 1338 

Tiburón de puntas blancas . . . . . . . . . . . . . 1325 

Tiburón de puntas negras . . . . . . . . . . . . . 1343 

Tiburón grácil . . . . . . . . . . . . . . . . . . . . 1327 

Tiburón jaquetón . . . . . . . . . . . . . . . . . . 1335 

Tiburón macuira . . . . . . . . . . . . . . . . . . 1340 

Tiburón negro espinoso . . . . . . . . . . . . . . 1212 

Tiburón nervioso . . . . . . . . . . . . . . . . . . 1333 

Tiburón oceánico . . . . . . . . . . . . . . . . . . 1341 

Tiburón ojuelo . . . . . . . . . . . . . . . . . . . 1351 

Tiburón rabo manchado . . . . . . . . . . . . . . 1347 

Tiburón sarda . . . . . . . . . . . . . . . . . . . . 1339 

Tiburón segador . . . . . . . . . . . . . . . . . . 1352 

Tiburón trompudo . . . . . . . . . . . . . . . . . 1342 

Tiburón trozo . . . . . . . . . . . . . . . . . . . . 1345 

Tiger dumpling squid . . . . . . . . . . . . . . . . . 720 

Tiger shark . . . . . . . . . . . . . . . . . . 1197, 1349 

tigris, Sepia . . . . . . . . . . . . . . . . . . . . . 748 

tilstoni, Carcharhinus . . . . . . . . . . . . 1340, 1348 

Tintorera . . . . . . . . . . . . . . . . . . . . . . 1349 

tjutjot, Carcharhinus . . . . . . . . . . . . . . . . 1334 

Todarodes pacificus . . . . . . . . . . . . . . 788, 795 

Todarodes pacificus pacificus . . . . . . . . . . . . 795 

Todarodes pacificus pusillus . . . . . . . . . . . . 795 

Todarodes pacificus ssp. . . . . . . . . . . . 788, 796 

Todaropsis eblanae . . . . . . . . . . . . . . . . . 796 

Tollo cigarro . . . . . . . . . . . . . . . . . . . . 1228 

Tollo coludo elegante . . . . . . . . . . . . . . . . 1295 

Tollo coludo grácil . . . . . . . . . . . . . . . . . 1295 

Tollo coludo pigmeo . . . . . . . . . . . . . . . . 1295 

Tollo flecha . . . . . . . . . . . . . . . . . . . . . 1225 

Tollo lucero diablo . . . . . . . . . . . . . . . . . 1226 

Tollo lucero mocho . . . . . . . . . . . . . . . . . 1226 

Tollo mandarín . . . . . . . . . . . . . . . . . . . 1224 

Tollo pigmeo . . . . . . . . . . . . . . . . . . . . 1228 

Tollo pigmeo espinudo . . . . . . . . . . . . . . . 1229 

Tollo trompalarga . . . . . . . . . . . . . . . . . . 1225 

Tolo velvet shrimp . . . . . . . . . . . . . . . . . . 932 

toloensis, Metapenaeopsis . . . . . . . . . . . . . 932 

Tope shark . . . . . . . . . . . . . . . . . . . . . 1302 

Topes . . . . . . . . . . . . . . . . . . . . . . . . 1297 

torazame, Scyliorhinus . . . . . . . . . . . . . . 1292 

Toro bacota . . . . . . . . . . . . . . . . . . . . . 1266 

Torpedo shrimp . . . . . . . . . . . . . . . . . . . . 944 

Toutenon japonais . . . . . . . . . . . . . . . . . . 795 

Toutenon souffleur . . . . . . . . . . . . . . . . . . 796 

toyamensis, Sepia kobiensis var. . . . . . . . . . . 743 

Trachypenaeopsis . . . . . . . . . . . . . . . 889-890 

Trachypenaeus . . . . . . . . . . . . . . . . . . . . 890 

Trachypenaeus anchoralis . . . . . . . . . . . . . 948 

Trachypenaeus asper . . . . . . . . . . . . . 927, 950 

Trachypenaeus curvirostris . . . . . . . . 927-928, 950 

Trachypenaeus fulvus . . . . . . . . . . . . . . . . 928 

Trachypenaeus gonospinifer . . . . . . . . . . . . 949 

Trachypenaeus granulosus . . . . . . . . . . . . . 949 

Trachypenaeus longipes . . . . . . . . . . . 927, 950 

Trachypenaeus malaiana . . . . . . . . . 927-928, 950 

Trachypenaeus sedili . . . . . . . . . . . . . 928, 950 

Trachypenaeus unicus . . . . . . . . . . . . . . . . 928 

Trachypenaeus villaluzi . . . . . . . . . . . . . . . 951 

tranquebarica, Scylla . . . . . . . . . . . . 1127-1128 

treba, Arctosepia . . . . . . . . . . . . . . . . . . . 739 

TREMOCTOPODIDAE . . . . . . . . . . . . . . . . 802 

Tremoctopus violaceusv . . . . . . . . . . . . . . . 802 

Trépang . . . . . . . . . . . . . . . . . . . . 1163-1164 

Trépang à canaux blancs . . . . . . . . . . . . . 1178 

Trépang curry . . . . . . . . . . . . . . . . . . . 1188 

Trépang rose . . . . . . . . . . . . . . . . . . . . 1177 

Trépang vert . . . . . . . . . . . . . . . . . . . . 1186 

Triaenodon . . . . . . . . . . . . . . . . . . . . . 1312 

Triaenodon obesus . . . . . . . . . . . . . 1325, 1358 

Triaenodon obtusus . . . . . . . . . . . . . . . . 1329 

TRIAKIDAE . . . 1196, 1264, 1294, 1297, 1306, 1313 

Triakis . . . . . . . . . . . . . . . . . . . . . . . 1297 

tricuspidatus, Carcharias . . . . . . . . . . . . . 1266 

trigonus, Linuparus . . . . . . . . . . . . . . . . 1024 

trirostrata, Sepiola . . . . . . . . . . . . . . 716-717 

trispeculare, Hemiscyllium . . . . . . . . . . . . 1259 

trispinosus, Gecarcinus . . . . . . . . . . . . . . 1113 

trituberculatus, Portunus . . . . . . . 1115, 1124-1125 

Tropical sawshark . . . . . . . . . . . . . . . . . 1234 

True crabs . . . . . . . . . . . . . . . . . . . . . 1048 

True lobster . . . . . . . . . . . . . . . . . . . . . . 974 

True lobsters . . . . . . . . . . . . . . . . 976-977, 982 

truncata, Charybdis . . . . . . . . . . . . . 1120, 1131 

Tsivakihini paste shrimp . . . . . . . . . . . . . . . 862


tuberculata, Ocythoe . . . . . . . . . . . . . . . . . 802 

tuberculata, Plagusia . . . . . . . . . . . . . . . 1146 

Tuberculated Sally-light-foot . . . . . . . . . . . . 1146 

tuberculosus, Ozius . . . . . . . . . . . . . . . . 1109 

tudes, Sphyrna . . . . . . . . . . . . . . . . . . . 1365 

Two-spot locust lobster . . . . . . . . . . . . . . . 1043 

Twospined arm swimming crab . . . . . . . . . . 1130 

tygrinus, Stegostoma . . . . . . . . . . . . . . . . 1262 

typicus, Carcinoplax longimanus . . . . . . . . . 1114 

typus, Rhincodon . . . . . . . . . . . . . . . . . 1263 

typus, Rhiniodon . . . . . . . . . . . . . . . . . . 1263 

U 

Uca spp. . . . . . . . . . . . . . . . . . . . . . . 1048 

Umbrella squids . . . . . . . . . . . . . . . . . . . 787 

Uncta shrimp . . . . . . . . . . . . . . . . . . . . . 945 

uncta, Parapenaeopsis . . . . . . . . . . . . . . . 945 

Unicorn blunthorn lobster . . . . . . . . . . . . . 1025 

unicornutus, Palinustus . . . . . . . . . . . . . 1025 

unicus, Trachypenaeus . . . . . . . . . . . . . . . . 928 

Upogebia spp. . . . . . . . . . . . . . . . . . . . 1048 

UPOGEBIIDAE . . . . . . . . . . . . . . . . 976, 1048 

uritai, Rhynchocinetes . . . . . . . . . . . . . . . 971 

Uroteuthis . . . . . . . . . . . . . . . . . . . . . . 764 

Uroteuthis bartschi . . . . . . . . . . . . . . . . . 779 

ursulae, Crumenasepia . . . . . . . . . . . . . . . 748 

ursus major, Parribacus . . . . . . . . . . . . . . 1037 

urvillei, Cardisoma . . . . . . . . . . . . . . . . 1149 

utilis, Etisus . . . . . . . . . . . . . . . . . . . . 1101 

uyii, Nipponololigo . . . . . . . . . . . . . . 772-773 

V 

Vampyromorphs . . . . . . . . . . . . . . . . . . . 696 

vanrooyeni, Carcharhinus . . . . . . . . . . . . . 1339 

Variable squillid mantis shrimp . . . . . . . . . . . . 849 

variegatus, Stichopus . . . . . . . . . . . . . . . 1188 

varium, Stegostoma . . . . . . . . . . . . . . . . 1262 

Varuna . . . . . . . . . . . . . . . . . . . . 1138, 1144 

Varuna litterata . . . . . . . . . . . . . . . . . . 1144 

Varuna yui . . . . . . . . . . . . . . . . . . . . . 1144 

velutinus, Metanephrops . . . . . . . . . . . . . . 994 

velutinus, Puerulus . . . . . . . . . . . . . . . . 1027 

Velvet dogfish . . . . . . . . . . . . . . . . . . . 1228 

Velvet lobster . . . . . . . . . . . . . . . . . . . . . 994 

Velvet shrimp . . . . . . . . . . . . . . . . . . . . . 933 

Velvet whip lobster . . . . . . . . . . . . . . . . . 1027 

Venus flower basket shrimp . . . . . . . . . . . . . 955 

Venus shrimp . . . . . . . . . . . . . . . . . . . . . 955 

venusta, Parapenaeopsis . . . . . . . . . . . . . . 946 

venusta, Spongicola . . . . . . . . . . . . . . . . . 955 

vericeli, Justitia . . . . . . . . . . . . . . . . . . 1023 

verreauxi, Amplisepia . . . . . . . . . . . . . . . . 738 

versicolor, Episesarma . . . . . . . . . . . . . . 1143 

versicolor, Holothuria . . . . . . . . . . . . . . . 1180 

versicolor, Holothuria (Metriatyla) . . . . . . . 1179 

versicolor, Holothuria (Metriatyla) scabra var.. 1179-1180 

versicolor, Panulirus . . . . . . . . . . . . . . . 1021 

versuta, Arctosepia . . . . . . . . . . . . . . . . . 739 

verweyi, Apristurus . . . . . . . . . . . . . . . . 1287 

victor, Matuta . . . . . . . . . . . . . . . . . . . 1095 

Viet Nam cuttlefish . . . . . . . . . . . . . . . . . . 761 

Vietnamese crest prawn . . . . . . . . . . . . . . . 965 

Vietnamese squillid mantis shrimp . . . . . . . . . . 848 

vietnamica, Sepia . . . . . . . . . . . . . . . . . . 761 

vietnamicus, Exopalaemon . . . . . . . . . . . . . 965 

vigil, Podophthalmus . . . . . . . . . . . . . . . 1123 

villaluzi, Trachypenaeus . . . . . . . . . . . . . . 951 

Vinegar crabs . . . . . . . . . . . . 1048, 1060, 1138 

violaceus, Tremoctopus . . . . . . . . . . . . . . . 802 

Violet vinegar crab . . . . . . . . . . . . . . . . . 1143 

Violet-spotted reef lobster . . . . . . . . . . . . . . 999 

virilis, Aristeus . . . . . . . . . . . . . . . . . . . 873 

vitiensis, Bohadschia . . . . . . . . . . . . . . . 1174 

VITRELEDONELLIDAE . . . . . . . . . . . . . . . 802 

Voss’ cuttlefish . . . . . . . . . . . . . . . . . . . . 762 

vossi, Loligo . . . . . . . . . . . . . . . . . . 766, 774 

vossi, Sepia . . . . . . . . . . . . . . . . . . . . . 762 

vulgaris, Acetes . . . . . . . . . . . . . . . . . . . 865 

vulgaris, Octopus . . . . . . . . . . . . . . . 689, 800 

vulpinus, Alopias . . . . . . . . . . . . . . 1271-1273 

W 

waddi, Brachaelurus . . . . . . . . . . . . . . . 1244 

waguensis, Palinustus . . . . . . . . . . . . . . 1025 

walbeehmi, Scoliodon . . . . . . . . . . . . . . . 1354 

wardi, Orectolobus . . . . . . . . . . . . . . . . 1248 

Watasenia . . . . . . . . . . . . . . . . . . . . . . 781 

Watasenia scintillans . . . . . . . . . . . . . . . . 781 

Water crabs . . . . . . . . . . . . . . . . . . . . 1126 

Weasel sharks . . . . . . . . . . . . . 1196-1197, 1305 

Web’s cuttlefish . . . . . . . . . . . . . . . . . . . . 763 

weberi, Caridina . . . . . . . . . . . . . . . . . . . 961 

weberi, Heteroteuthis . . . . . . . . . . . . . . . . 715 

weberi, Sepiella . . . . . . . . . . . . . . . . . . . 763 

wellsi, Metapenaeopsis . . . . . . . . . . . . . . . 933 

Western king prawn . . . . . . . . . . . . . . . . . 918 

Western school shrimp . . . . . . . . . . . . . . . . 935 

Whale shark . . . . . . . . . . . . . . . . . . . . 1263 

Whale sharks . . . . . . . . . . . . . . . . . 1196, 1263 

wheeleri, Carcharhinus . . . . . . . . . . . . . . 1328 

Whiskered velvet shrimp . . . . . . . . . . . . . . . 907 

White shark . . . . . . . . . . . . . . . . . . . . . 1197 

White sharks . . . . . . . . . . . . . . . . . . . . 1274 

White teatfish . . . . . . . . . . . . . . . . . . . . 1181 

White threads fish . . . . . . . . . . . . . . . . . 1178 

White whisker spiny lobster . . . . . . . . . . . . 1015 

White-striped octopus . . . . . . . . . . . . . . . . 817 

Whitecheek shark . . . . . . . . . . . . . . . . . 1334 

Whitefin topeshark . . . . . . . . . . . . . . . . . 1302 

Whitespotted bambooshark . . . . . . . . . . . . 1256 

Whitetip reef shark . . . . . . . . . . . . . . . . . 1358 

Whitley’s cuttlefish . . . . . . . . . . . . . . . . . . 755 

whitleyana, Sepia . . . . . . . . . . . . . . . 753, 755 

whitleyanum, Acanthosepion . . . . . . . . . . . . 755 

wieneckii, Palinurellus . . . . . . . . . . . . . . 1004 

Winghead shark . . . . . . . . . . . . . . . . . . 1363 

Witch prawn . . . . . . . . . . . . . . . . . . . . . 914 

Wobbegongs . . . . . . . . . . . . . . . . . . . . 1245 

wolfi, Octopus . . . . . . . . . . . . . . . . . . . . 801 

Wood shrimp . . . . . . . . . . . . . . . . . . . . . 935


woodmasoni, Erugosquilla . . . . . . . . . . 842, 846 

X 

Xanthid stone crabs . . . . . . . . . . . . . . . . 1098 

XANTHIDAE . . . . . . 1048-1049, 1057, 1061, 1098, 

1103, 1110, 1114 

Xanthids . . . . . . . . . . . . . . . . . . . . . . 1112 

xera, Arctosepia braggi . . . . . . . . . . . . . . . 739 

Y 

Yellow moon crab . . . . . . . . . . . . . . . . . . 1096 

Yellow shrimp . . . . . . . . . . . . . . . . . . . . . 934 

yokoyae, Loligo . . . . . . . . . . . . . . . . . . . 773 

York shrimp . . . . . . . . . . . . . . . . . . . . . . 937 

yui, Varuna . . . . . . . . . . . . . . . . . . . . 1144 

Z 

zambezensis, Carcharhinus . . . . . . . . . . . . 1339 

Zebra bullhead shark . . . . . . . . . . . . . . . . 1240 

Zebra shark . . . . . . . . . . . . . . . . . . . . 1262 

Zebra sharks . . . . . . . . . . . . . . . . . 1197, 1262 

zebra, Heterodontus . . . . . . . . . . . . . . . . 1240 

Zorro . . . . . . . . . . . . . . . . . . . . . . . . 1273 

Zorro ojón . . . . . . . . . . . . . . . . . . . . . 1272 

Zorro pelágico . . . . . . . . . . . . . . . . . . . 1271 

Zosimus aeneus . . . . . . . . . . . . . . . 1049, 1098 

zygaena, Sphyrna . . . . . . . . . . . . . . 1364-1366

FAO SPECIES IDENTIFICATION GUIDE FOR FISHERYPURPOSES 

THE LIVING MARINE RESOURCES OF THE 

WESTERNCENTRAL 

PACIFIC 

ISSN 1020-6868 

Volume 2. Cephalopods, crustaceans, holothurians and sharks

FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES 

THE LIVING MARINE RESOURCES OF THE 

WESTERN CENTRAL PACIFIC 

VOLUME 2 

Cephalopods, crustaceans, holothurians and sharks 

edited by 

Kent E. Carpenter 

Department of Biological Sciences 

Old Dominion University 

Norfolk, Virginia, USA 

and 

Volker H. Niem 

Marine Resources Service 

Species Identification and Data Programme 

FAO Fisheries Department 

with the support of the 

South Pacific Forum Fisheries Agency (FFA) 

and the 

Norwegian Agency for International Development (NORAD) 

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS 

Rome, 1998

ii 

The designations employed and the presentation of material in 

this publication do not imply the expression of any opinion 

whatsoever on the part of the Food and Agriculture Organization 

of the United Nations concerning the legal status of any 

country, territory, city or area or of its authorities, or concerning 

the delimitation of its frontiers and boundaries. 

M-40 

ISBN 92-5-104051-6 

All rights reserved. No part of this publication may be reproduced by 

any means without the prior written permission of the copyright owner. 

Applications for such permissions, with a statement of the purpose and 

extent of the reproduction, should be addressed to the Director, 

Publications Division, Food and Agriculture Organization of the United 

Nations, via delle Terme di Caracalla, 00100 Rome, Italy. 

© FAO 1998

Carpenter, K.E.; Niem, V.H. (eds) 

FAO species identification guide for fishery purposes. The living marine resources 

of the Western Central Pacific. Volume 2. Cephalopods, crustaceans, holothurians 

and sharks. 

Rome, FAO. 1998. 687-1396 p. 

SUMMARY 

This multivolume field guide covers the species of interest to fisheries of the major 

marine resource groups exploited in the Western Central Pacific. The area of coverage 

includes FAO Fishing Area 71 and the southwestern portion of Fishing Area 77 

corresponding to the South Pacific Commission mandate area. The marine resource 

groups included are seaweeds, corals, bivalves, gastropods, cephalopods, stomatopods, 

shrimps, lobsters, crabs, holothurians, sharks, batoid fishes, chimaeras, 

bony fishes, estuarine crocodiles, sea turtles, sea snakes, and marine mammals. The 

introductory chapter outlines the environmental, ecological, and biogeographical factors 

influencing the marine biota, and the basic components of the fisheries in the Western 

Central Pacific. Within the field guide, the sections on the resource groups are arranged 

phylogenetically according to higher taxonomic levels such as class, order, and family. 

Each resource group is introduced by general remarks on the group, an illustrated 

section on technical terms and measurements, and a key or guide to orders or families. 

Each family generally has an account summarizing family diagnostic characters, biological 

and fisheries information, notes on similar families occurring in the area, a key 

to species, a checklist of species, and a short list of relevant literature. Families that 

are less important to fisheries include an abbreviated family account and no detailed 

species information. Species in the important families are treated in detail (arranged 

alphabetically by genus and species) and include the species name, frequent synonyms 

and names of similar species, an illustration, FAO common name(s), diagnostic characters, 

biology and fisheries information, notes on geographical distribution, and a 

distribution map. For less important species, abbreviated accounts are used. Generally, 

this includes the species name, FAO common name(s), an illustration, a distribution 

map, and notes on biology, fisheries, and distribution. Each volume concludes with its 

own index of scientific and common names. 

iii

FAO Species Identification Guide for Fishery Purposes Western

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?