CRABS (BRACHYURA) OF THE GULF OF MEXICO

a catalogue & 

bibliography to the 

CRABS (BRACHYURA) 

OF THE 

GULF OF MEXICO 

LAWRENCE W. POWERS 

ISSN/0082-3349 

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60- Mai/2AJh\_

pnMTTJTRrTT'TnMQ TM MaPTTOR .

Editor: DONALD E. WOHLSCHLAG 

Editorial Assistant: RUTH GRUNDY 

EDITOR S NOTE 

We are offering, "A Catalogue and Bibliography to the Crabs (Brachyura) of the Gulf of 

Mexico," By Lawrence W. Powers as a supplement to Volume 20 of our Contributions in Marine 

Science. We hope that this type of monographic work will stimulate further syntheses. 

Additional copies of this Catalogue and Bibliography may be obtained by writing; 

The Librarian 

Port Aransas Marine Laboratory 

The Universit3r of Texas Marine Science Institute 

Port Aransas, Texas 78373 

Price: $8.00 per copy

ACKNOWLEDGMENTS 

I wish to express my appreciation for those who reviewed the preliminary 

draft of this manuscript. The following offered valuable suggestions for its improvement: 

Lawrence G. Abele, Fenner A. Chace, Jr., Darryl L. Felder, John S. 

Garth, and Austin B. Williams. Samuel R. Haley and Warren Pulich reviewed 

the revised introduction. I assume full responsibility for remaining errors and 

omissions. 

I would like to dedicate this effort to Mrs. Ruth Grundy, librarian of the 

Marine Science Laboratory at Port Aransas, for her initial encouragement of 

this project, continued enthusiasm during compilation, patience with the many 

missed deadlines, and her labors to help see it through to completion. I also 

appreciate the help and support of others at the Marine Science Laboratory, 

especially Donald Wohlschlag, William Brogden, Carl Oppenheimer, Sarah 

Lowrie, Phyllis Picarazzi, and Mary Ellen Tomberlein. Dinah Bowman prepared 

the cover illustration.

TABLE OF CONTENTS 

Page 

Acknowledgments 3 

Table of Contents - 5 

INTRODUCTION 7 

CLASSIFICATION 8 

BIOGEOGRAPHY 10 

DISCUSSION _ 13 

SPECIES ENTRIES - 19 

BRACHYURA 19 

Section DROMIACEA 19 

Superfamily DROMIOIDEA 19 

Family DROMIIDAE 19 

Family PROSOPIDAE 21 

Subfamily HOMOLODROMIINAE 21 

Superfamily HOMOLOIDEA 21 

Family HOMOLIDAE 21 

Family LATREILLIDAE 23 

Section OXYSTOMATA 23 

Superfamily RANINOIDEA 23 

Family RANINIDAE 23 

Superfamily DORIPPOIDEA 26 

Family DORIPPIDAE 26 

Superfamily CALAPPOIDEA 29 

Family CALAPPIDAE 29 

Subfamily CALAPPINAE 29 

Subfamily MATUTINAL 32 

Family LEUCOSIIDAE 35 

Subfamily EBALIINAE 35 

Subfamily LEUCOSIINAE 36 

Subfamily PHILYRINAE 38 

Section HAPALOCARCINIDEA 40 

Superfamily HAPALOCARCINOIDEA 40 

Family HAPALOCARCINIDAE 40 

Section OXYRHYNCHA 41 

Family MAJIDAE 41 

Subfamily ACANTHONYCHINAE 41 

Subfamily INACHINAE 42 

Subfamily MAJINAE 49 

Subfamily MITHRACINAE 50 

Subfamily OPHTHALMIINAE 60 

Subfamily PISINAE 62 

Family PARTHENOPIDAE 67 

Subfamily PARTHENOPINAE 67

Section CANCRIDEA 71 

Superfamily CANCROIDEA 71 

Family ATELECYCLIDAE 71 

Subfamily ATELECYCLINAE 71 

Family CANCRINAE 71 

Subfamily CANCRINAE 71 

Section BRACHYRHYNCHA 72 

Superfamily PORTUNOIDEA 72 

Family PORTUNIDAE 72 

Subfamily POLYBIINAE 72 

Subfamily PORTUNINAE 74 

Superfamily XANTHOIDEA 86 

Family POTAMIDAE 86 

Family XANTHIDAE 87 

Family GERYONIDAE 110 

Family GONEPLACIDAE 111 

Family PALICIDAE 117 

Family PINNOTHERIDAE 119 

Subfamily PINNOTHERINAE 119 

Subfamily PINNOTHERELIINAE 125 

Family GRAPSIDAE 128 

Subfamily GRAPSINAE 128 

Subfamily PLAGUSIINAE 131 

Subfamily SESARMINAE 132 

Subfamily VARUNINAE 136 

Family GECARCINIDAE 138 

Superfamily OCYPODOIDEA 140 

Family OCYPODIDAE 140 

Subfamily OCYPODINAE 140 

BIBLIOGRAPHY 150 

INDEX 184

INTRODUCTION 

The purpose of this catalogue and bibliography is to provide a convenient 

reference source for information on the brachyuran (true) crabs of the Gulf of 

Mexico. Although it is intended primarily for zoologists and others familiar with 

taxonomic resources, the catalogue includes some informal features in order to 

increase its usefulness to students and persons without systematic experience. 

The catalogue is a compilation of recent literature and the taxonomic organization 

of the species entries represents the author's interpretation of revisions by 

carcinological authorities. As a reference source, it is intended to be a compromise 

between the systematic works with formal synonymies and descriptions 

of species and various bibliographies that could be generated by the use of computerized 

key word entries. 

The major part of the catalogue consists of a checklist of species from the 

Gulf of Mexico (referred throughout the text as the Gulf) that was compiled 

from the published literature. Unpublished materials, such as theses, manuscripts, 

and uncirculated reports, were included sparingly. Doctoral dissertations 

were cited in cases where the material was not published elsewhere or when the 

topic was considered essential for inclusion in this work. It was not possible to 

examine and include every known reference and some sources were inadvertently 

missed; omissions and errors should be brought to the attention of the 

author. A complete checklist of the Brachyura of the Gulf has not been published 

since Rathbun's multi-volume work, "The Crabs of America" (1918, 1925, 1930, 

1937). Since that time, numerous local and regional surveys have yielded new 

findings on the distribution and biologj^ of many species, new species have been 

described, and revisions in the sj^stematic organization of several groups have 

appeared. For most non-specialists, much of this documentary literature is inaccessible 

and the taxonomic revisions niay be difficult to evaluate. The present 

work should serve as an introduction to the literature for a species and it should 

also indicate the amount of data and type of research available for a species or 

group of crabs. 

References to the literature are divided into two sections. Those of value in 

identification and characterization of a species are listed under the name and 

source of original description. These citations include synonj-Tns and other 

invalid names; they occasionally emphasize a name or part of a description that 

is not sj^nonymous (e.g., Williams, 1974a, p. 731, figs. 4, 18a (not 18b), 20c, 

etc.). Other references are listed under "Remarks": they provide information 

on the natural historj^ ecology, development, physiology, behavior, pathology, 

or commercial fishery of a species. For some commercially-important species, 

the available literature is large and widely scattered. The citations for these 

species are listed with a minimum of descriptive explanation and the number of 

omitted references will be proportionately higher than for those species with 

only a few known references. 

The species entries consist of the current valid name, the published original 

reference of the species description, common names (if any), taxonomic and

8 Lawrence W. Powers 

descriptive references, geographical and bathymetric distributions, habitats, and 

bibliography with annotations and comments. Species are arranged alphabetically 

viîthin genera and subgenera, as are genera within subfamilies. 

Geographical ranges are based on published sources. If the identification of a 

species or locality is doubtful, the localitj^ is preceded by a question mark. Place 

names are listed from north to south and east to west; eastern Atlantic localities 

follow western Atlantic; eastern Pacific and Indo-Pacific localities are cited last. 

The presence of a species is not assumed in the absence of collection records from 

large gaps in otherwise continuous distributions. For example, many species are 

listed for the Carolinas and the east coast of Florida, without indicating presence 

in Georgian waters. 

Bathymetric ranges are presented in meters and in fathoms, in order to facilitate 

comparisons wth new data as they appear in the literature. Depths are 

omitted for semiterrestrial and terrestrial crabs. 

Habitat descriptions include substrate types, terrain features (land crabs), associations 

with other organisms (commensals, parasites), type of water (marine, 

brackish), and general biotope (estuarine, marsh, rocky intertidal, offshore 

benthic, etc.). Terrestrial habitats are usually described in greater detail than 

aquatic habitats. 

Remarks include literature citations on all aspects of the biolog}^ of the species. 

In many cases, the available literatvire may consist only of one or a few published 

descriptions or citations of the species. Regional lists include faunal surveys 

in the Gulf of Mexico area. These lists may include data on the habitats, 

seasonal occurrence, and biology of the crabs, or they may list only the species 

names. Species identifications were usually verified by a specialist; most of the 

lists were published. Other remarks include discussions of taxonomic revisions, 

identification problems, comparisons with other related species, records of ovigerous 

females, and ecological notes. 

CLASSIFICATION 

A review of attempts to classify decapod crustaceans, including arrangements 

of brachyuran families, was presented by Glaessner (1969) in Treatise on 

Invertebrate Paleontology. The scheme adopted for the Treatise, a compromise 

of paleontological and zoological considerations, is used in the present catalogue. 

Table I presents an outline of the extant families of Brachyura as recognized by 

Glaessner (1969) and modified for the present work. 

The family Hapalocarcinidae is not included in Glaessner (1969) and the 

Palicidae is listed within the Brachyrhyncha as Superfamily Uncertain. Fenner 

Chace, in a review of a preliminary'- manuscript of this catalogue, suggested the 

placement of the hapalocarcinids as a separate section, although Verrill (1908, 

p. 426) proposed the inclusion of these crabs as "a peculiar superfamily group," 

apparently among the Oxystomata. Chace also suggested the inclusion of the 

Palicidae among the Xanthoidea as a matter of convenience. The freshwater 

family of crabs, Potamidae, has been revised by Bott (1955b) and separated into 

two families, the Pseudothelphusidae and the Trichodactylidae. The Prosopidae

I Crabs 

SECTION DROMIACEA de Haan, 1833 

TABLE I 

Classification of Brachyuran Families 

INFRAORDER BRACHYURA 

Superfamily DROMIOIDEA de Haan, 1833 

Family PROSOPIDAE von Meyer, 1860 

Family DYNOMENIDAE Ortmann, 1892 

Family DROMIIDAE de Haan, 1833 

Superfamily HOMOLOIDEA White, 1847 

Family HOMOLIDAE White, 1847 

SECTION OXYSTOMATA H. Milne Edwards, 1834 

SuperfamUy DORIPPOIDEA de Haan, 1841 

Family DORIPPIDAE de Haan, 1841 

Superfamily CALAPPOIDEA de Haan, 1833 

Family CALAPPIDAE de Haan, 1833 

Family LEUCOSIIDAE Samouelle, 1819 

Superfamily RANINOIDEA de Haan, 1841 

Family RANINIDAE de Haan, 1841 

SECTION HAPALOCARCINIDEA Caiman, 1900 

Superfamily HAPALOCARCINOIDEA Caiman, 1900 

Family HAPALOCARCINIDAE Caiman, 1900 

SECTION OXYRHYNCHA Latreille, 1803 

Family MAJIDAE Samouelle, 1819 

Family PARTHENOPIDAE Macleay, 1838 

SECTION BRACHYRHYNCHA Borradaile, 1907 

Superfamily PORTUNOIDEA Rafinesque, 1815 

Family PORTUNIDAE Rafinesque, 1815 

Superfamily XANTHOIDEA Dana, 1851 

Family XANTHIDAE Dana, 1851 

Family GERYONIDAE Colosi, 1924 

Family GONEPLACIDAE Macleay, 1838 

Family POT AMID AE Ortmann, 1896 

Family PINNOTHERIDAE de Haan, 1833 

Family GRAPSIDAE Macleay, 1838 

Family GECARCINIDAE Macleay, 1838 

Family PALICIDAE Bouvier, 1898 

Superfamily OCYPODOIDEA Rafinesque, 1815 

Family OCYPODIDAE Rafinesque, 1815 

Family RETROPLUMIDAE Gill, 1894 

SECTION CANCRIDEA Latreille, 1803 

Superfamily CANCROIDEA Latreille, 1803 

Family CANCRIDAE Latreille, 1803 

Family CORYSTIDAE Samouelle, 1819 

Family ATELECYCLIDAE Ortmann, 1893 

of the Gulf of Mexico 9


includes the subfamily Homolodromiinae Alcock, 1899, usually treated as a 

family in other systematic classifications. The Latreillidae was recognized as a 

separate family by Rathbun (1937), whereas Balss (1957) included the genus 

Latreillia Roux in the Homolidae. Glaessner (1969) tentatively adopted the 

arrangement of Balss, but left the question of placement open; the Latreillidae 

are presented as a separate family in the present work. The families Dynomenidae 

and Retroplumidae do not contain species represented in the Gulf of 

Mexico. 

The arrangement of species and genera within their respective higher taxa 

may differ according to the current stability of nomenclature within the group. 

Within the Xanthidae, subfamilies are not listed because the arrangement of 

genera is incomplete, pending further revisions by Guinot and others. The genus 

Uca of the Ocypodidae has received considerable attention from carcinologists, 

notably Bott (1973) and Crane (1975). Both of these workers have proposed 

subgenera to accommodate findings of morphological, biogeographical, and behavioral 

similarities among species groups. Because the changes are too recent 

to permit evaluation by other specialists, the species of Uca are listed in the 

present text alphabetically; controversial forms, proposed as full species by some 

workers and as subspecies by others, are treated as species for the present. It is 

much easier to combine • separate sets of references and records than to attempt 

to separate them at a later date. Generally, this plan has been followed throughout 

the systematic presentation; when a taxon can be treated in two or more 

ways, the simplest arrangement was selected. 

A discussion of brachyuran evolution is beyond the scope of this catalogue and 

the reader is referred to papers by Bourne (1922), Gordon (1963), Stevcic 

(1971a, 1971b) and the monograph. Treatise on Invertebrate Paleontology, 

which includes the review by Glaessner (1969). This latter reference also compares 

and discusses the various classification schemes that have been proposed 

to account for presumed affinities of living and fossil specimens. 

BIOGEOGRAPHY 

The species catalogue recognizes 352 species of 158 genera, belonging to 22 

families. Enumerations of taxa are somewhat arbitrary, depending on the amount 

of splitting or lumping of various groups. In addition to the 352 species recorded 

from the Gulf of Mexico, two subspecies and two varietal forms are listed as 

distinct entries, three species of uncertain Gulf distribution are included (recorded 

from Cuban waters, coast luispecified), and one species from outside of 

the Gulf is listed because of nomenclatural relationships with a Gulf species. 

Forty-one of the 352 species are presently regarded as endemic to the Gulf; the 

distribution of species and genera by family are listed in Table II. 

The greatest number of endemic species occurred among the Pinnotheridae, 

a family composed mainly of crabs commensal with other organisms. Specificity 

for hosts and other restrictions imposed by the biology of commensal associates 

may account for the high ratio of endemism in this family. The largest families, 

in terms of total numbers of species, contain relatively few species confined to the

Family 

TABLE II 

Endemic and Total Species and Genera of Brachyura 

Droniiidae 

Prosopidae 

Homolidae 

Latreillidae 

Raninidae 

Dorippidae 

Calappidae 

Leucosiidae 

Hapalocarcinidae 

Majidae 

Parthenopidae 

Afelecyclidae 

Cancridae 

Portunidae 

Xanthidae 

Gei-yonidae 

Goneplacidae 

Palicidae 

Pinnotheridae 

Grapsidae 

Gecarcinidae 

Ocypodidae 

Total 

Endemic 

species 

0 

0 

1 

0 

1 

6 

0 

0 

0 

4 

0 

0 

0 

2 

3 

0 

5 

1 

12 

1 

0 

5 

41 

Crabs of the Gulf of Mexico 11 

Species 

5 

2 

4 

1 

7 

16 

13 

15 

1 

77 

13 

1 

1 

27 

69 

1 

19 

9 

31 

21 

4 

15 

352 

Total 

Genera 

Gulf. The Majidae, Xanthidae, Portunidae, and Grapsidae have endemic percentages 

of less than 10% of total Gulf species, whereas the Dorippidae, Goneplacidae, 

and Ocypodidae contain endemic ratios of one-fourth or greater. 

The geographical distribution of Gulf species within other areas is compared 

in Table III. The degree of affinity with the Caribbean fauna is indicated by the 

large percentage of Gulf crabs also occurring in the Greater Antilles (63.1%), 

Lesser Antilles (55.7%), and north coast of South America (34.7%). Within 

these regions, however, there exist considerable differences in collection effort 

and subsequent taxonomic study. Thus, the number of apparent species recorded 

from a particular island or section of continental coastline may be a better measure 

of the number of marine science institutes present than actual species diversity 

or richness. 

Eastern Florida, the Bahamas, and Bermuda contain successively'' fewer Gulf 

species, whereas the combined Virginian and Carolinian Provinces of the Atlantic 

coast contain 44.3% of the total Gulf crabs. The number of Gulf species present 

in the eastern Atlantic comprises only 7.7% of the Gulf total and one-third of 

these belong to the ubiquitous Grapsidae. Only 16 Gulf species (4.5%) are also 

recorded from the eastern Pacific; seven species have been collected from other 

parts of the Indo-Pacific region. 

3 

2 

3 

1 

4 

7 

5 

8 

1 

33 

8 

1 

1 

8 

32 

1 

15 

1 

7 

12 

2 

3 

158


Family 

Dromiidae 

Prosopidae 

Homolidae 

Latreillidae 

Raninidae 

Dorippidae 

Calappidae 

Leucosiidae 


Majidae 

Parthenopidae 

Atelecyclidae 

Cancridae 

Portunidae 

Xanthidae 

Geryonidae 

Goneplacidae 

Palicidae 

Pinnotheridae 

Grapsidae 

Gecarcinidae 

Ocypodidae 

Total 

Total 

species 

5 

2 

4 

1 

7 

16 

13 

15 

1 

77 

13 

1 

1 

27 

69 

1 

19 

9 

31 

21 

4 

15 

352 

TABLE III 

Distribution of Gulf Crabs in Other Areas 

GA 

4 

2 

3 

1 

5 

12 

8 

10 

0 

55 

8 

0 

0 

20 

46 

1 

8 

2 

10 

15 

4 

8 

222 

Caribbean 

LA SA 

3 

2 

2 

0 

1 

5 

8 

10 

1 

53 

8 

1 

0 

17 

49 

0 

5 

4 

3 

13 

4 

7 

196 

3 

0 

0 

0 

1 

0 

4 

5 

0 

26 

4 

0 

0 

16 

34 

0 

1 

2 

2 

13 

4 

7 

122 

wc 

0 

0 

1 

0 

2 

0 

4 

0 

0 

21 

0 

0 

0 

13 

20 

0 

0 

0 

1 

8 

4 

6 

80 

EF 

2 

1 

1 

0 

1 

2 

5 

5 

0 

28 

5 

0 

1 

16 

20 

1 

2 

3 

5 

6 

3 

8 

115 

Western Atlantic 

BA BE NC 

2 

0 

1 

0 

1 

0 

3 

3 

0 

25 

4 

0 

0 

11 

31 

0 

2 

1 

0 

11 

3 

6 

104 

2 

0 

0 

0 

0 

0 

4 

0 

1 

8 

1 

0 

1 

13 

15 

0 

2 

0 

0 

9 

2 

1 

59 

4 

0 

2 

1 

3 

5 

10 

9 

0 

37 

9 

0 

1 

16 

27 

1 

5 

4 

13 

6 

0 

3 

156 

Other 

EA EP 

GA =: Greater Antilles; LA = Lesser Antilles; SA = north coast of South America; WC = western Caribbean, 

including east coast of Central America; EF = east coast of Florida; BA = Bahamas; BE = Bermuda; NC = U.S. 

Atlantic coast, from New England to Georgia; EA = eastern Atlantic, fiom Europe to South Africa and mid-South 

Atlantic islands; EP = eastern Pacific, from California to Chile, including Galapagos. 

The distribution of species within the Gulf of Mexico is presented in Table IV, 

with the Gulf regions diagxammed in Figure 1. Again, some areas have been 

sampled often (e.g., west and northwest coasts of Florida) and other areas have 

received only sporadic attention (east coast of Mexico). More species occur on 

the limestone-based continental shelf of the eastern Gulf than on the sand and 

mud substrates of the western Gulf. The Gulf regions in Figure 1 correspond to 

the geophysical features described by Antoine (1972) as Gulf provinces. The 

dominant sediment types are shown in Figure 2 and reef patches are indicated 

in Figure 3. Most Gulf species are probably West Indian in origin or affinity 

(see discussion below), where the shallow water habitats are similar to Gulf 

regions 1, 2, and 6. The north coast of Cuba (region 7) also contains a relatively 

high species abundance, probably associated with the greater degxee of habitat 

diversity and wide depth range in that part of the Gulf. A detailed analysis of 

species distribution patterns, by families, will not be attempted in the present 

contribution. Table IV summarizes the species recorded to date from each of 

0 

0 

2 

1 

1 

1 

2 

0 

0 

1 

0 

0 

0 

6 

4 

0 

0 

0 

0 

9 

0 

0 

27 

0 

0 

0 

0 

2 

1 

2 

0 

0 

1 

0 

0 

0 

1 

2 

0 

0 

0 

0 

6 

1 

0 

16

Family 

Dromiidae 

Prosopidae 

Homolidae 

Latreillidae 

Raninidae 

Dorippidae 

Calappidae 

Leucosiidae 


Majidae 

Parthenopidae 

Atelecyclidae 

Cancridae 

Portunidae 

Xanthidae 

Geryonidae 

Goneplacidae 

Palicidae 

Pinnotheridae 

Grapsidae 

Gecarcinidae 

Ocypodidae 

Total 

TABLE IV 

Species Distribution Within the Gulf of Mexico 

Total 

species 

5 

2 

4 

1 

7 

16 

13 

15 

1 

77 

13 

1 

1 

27 

69 

1 

19 

9 

31 

21 

4 

15 

352 

1 

5 

1 

2 

1 

5 

8 

11 

13 

1 

61 

10 

0 

1 

23 

61 

1 

12 

5 

16 

16 

2 

6 

261 


2 

2 

0 

0 

0 

1 

2 

5 

8 

0 

38 

6 

1 

0 

11 

28 

1 

5 

5 

12 

3 

0 

7 

145 

Gulf Region (F. Igure 1) 

3 4 5 

these regions. An analysis of breeding tiraies and collection sites of ovigerous 

females, combined with data on circulation patterns of currents, larval maturation 

periods, and tolerances to environmental parameters (temperature, salinity, 

oxygen content, etc.), will be necessary to establish a comprehensive biogeography 

of Gulf crabs. 

DISCUSSION 

The distribution of decapod crustaceans in the western Atlantic has been the 

subject of several discussions in recent years. Hedgpeth (1953) summarized 

much of what was knowia about the geology and zoogeography of the Gulf of 

Mexico. He emphasized the role of Pleistocene sea level changes and their effects 

on continuous distributions of marine organisms in the Carolinian Province, a 

region encompassing shores and adjacent waters from Texas to Cape Hatteras. 

After the Florida peninsula emerged, isolation of the Gulf and Atlantic populations 

occurred for many species not able to circumvent the tropical waters of 

South Florida. The distributions of Ovalipes and Uca were cited by Abele (1970) 

as examples. For this reason the range citations in the present contribution 

attempt to distinguish eastern, southern, and western Florida records. 

3 

0 

0 

0 

2 

2 

5 

3 

0 

20 

4 

1 

0 

13 

26 

1 

8 

2 

10 

8 

2 

8 

118 

0 

0 

1 

0 

2 

1 

3 

1 

0 

7 

0 

1 

0 

6 

4 

1 

2 

1 

1 

6 

3 

7 

47 

0 

0 

0 

0 

2 

1 

3 

1 

0 

1 

0 

1 

0 

6 

0 

0 

0 

0 

0 

1 

3 

3 

22 

6 

2 

0 

0 

0 

0 

1 

4 

1 

0 

15 

3 

0 

0 

11 

7 

0 

8 

3 

2 

3 

3 

6 

69 

7 

2 

2 

3 

1 

4 

11 

5 

3 

0 

31 

4 

0 

0 

16 

35 

1 

8 

4 

6 

11 

3 

7 

161


FIG. 1. Regions of the Gulf of Mexico, based on geological assemblages, depth, and geophysical 

characteristics. Afler Antoine (1972); numbers refer to Table IV and do not correspond 

with province numbers used by Antoine in his Figure 1-1. 

Williams (1965) computed an Atlantic-Gulf disjunct ratio of about 10%, 

based on studies of Carolina decapods. Only 3 (1.4%) of the species presented 

in his Table 1 were considered endemic to the Carolinas, further evidence of 

the ubiquitous nature of the northern fauna. Abele (1970) emphasized the northern 

and southern affinities of the northeastern Gulf decapods, which he believed 

was better characterized as a heterogenous assemblage rather than a typical 

fauna of a province he considered to be ill-defined. In his analysis, 91 species 

(36.5%) of decapods were similar to the typical Carolinian fauna, but many 

species previously thought to have a disjunct distribution have been more 

recently reported from southern Florida (Tabb and Manning, 1961; Rouse, 

1970). Abele listed 33 species (13.4% of the northeast decapod total) as endemic 

and 41 species (16.4%) as tropical in origin, confined to the Caribbean and Gulf 

by higher temperature requirements. The remaining 84 species in his study 

were the ubiquitous fauna that ranged from the Carolinas to the tropics. 

Studies of terrestrial and freshwater crabs can provide further insights into 

possible sources of origin. Chace and Hobbs (1969) listed 57 species of crabs from 

the Caribbean area, none of which had apparent origins in North America alone 

(exclusive of Mexico). They concluded that the majority of endemics in the


FIG. 2. Areas of the Gulf of Mexico based on dominant sediment type. After Lynch (1934). 

Coral reefs and patches are shown in Figure 3. 

A ^ sand, includes other narrow stippled areas adjacent to continental coastline. 

B := sand-mud. 

C = mud. 

D = blue mud. 

E = calcareous mud. 

F := limestone with thin veneer of detrital sediments. 

G = Globigerina ooze. 

H = pteropod ooze. 

Greater Antilles had originated in Central America or southern Mexico and a 

few had arrived from South America. The directions of prevalent surface currents 

(Figure 4) indicate the one-way flow of potential larval populations out of 

the Caribbean into the Gulf. Although the patterns of loop currents within the 

Gulf change seasonally, successive recruitment of Caribbean fauna to various 

parts of the Gulf are possible throughout the year. Various sources (Williams, 

1965 summarizes many of the records) indicate that many Caribbean crab 

species are ovigerous throughout the year; thus Gulf recruitment to a particular 

shore segment would be a function of current pattern and speed, seasonal temperature, 

and length of larval life. Laboratory studies of larval development 

(summarized by Garth, 1965b) indicate that warmer temperatures prolong 

maturation time. Larval populations thus carried into the Gulf could be diverted 

into one of several directions. The westward entering currents sweep the shores


FIG. 3. Location of major coral reefs and other features of the Gulf of Mexico, derived from 

several sources, including Lynch (1954). Dashed line represents the 100 fathom (600 feet) 

isobath. 

1—Florida Keys 

2—Dry Tortugas 

3—Apalachicola Bay and Cape San Bias 

4—Mississippi River Delta 

5—East and West Flower Garden Banks, off Texas 

6—Seven and One-Half Fathom Reef, off Padre Island 

7—Pink Shrimp Grounds in Campeche Bay 

8—Alacran Reef on Campeche Bank 

of Yucatan, eastern Mexico, and eventually, converge with other currents off 

Texas. Central entering currents flow northwest, into region 3 (Figure 1). The 

most complex patterns occur as the eastward currents diverge and converge 

in seasonal patterns of loops. Much of the resultant flow sweeps through the 

Straits of Florida to emerge into the Atlantic as the warm, northward moving 

Gulf Stream. Other loops circulate up the west coast of Florida, reaching as far 

as the Mississippi delta in the winter (Figure 4B). Although these large scale 

current diagrams do not indicate the complexity of water movements in the Gulf 

region, nor do they allow one to evaluate the spatial and temporal variations in 

local currents that might be used for the migration of larval crabs, Figure 4 (after 

Leipper, 1954) does indicate the successive relationship between Caribbean, Gulf, 

and western Atlantic faunal provinces, respectively. Undoubtably, many western

FIG. 4. Surface currents of the Gulf of Mexico (after Leipper, 1954). 

4A. Current patterns in June. 

4B. Current patterns in December. 

Crabs of the Gulf of Mexico 17


Atlantic faunal affinities with West Indian elements occur as a result of direct 

recruitment, bypassing the Gulf of Mexico. 

Specific genera that have been recentlj' investigated and summarized with 

respect to zoogeography include Sesarma (Abele, 1973), Ovalipes (Williams, 

1976), CalUnectes (Williams, 1974a), and Uca (Crane, 1975). Abele (1973) 

divided the six Florida species of Sesarma into those with Carolinian and West 

Indian affinities. Although Hedgpeth (1953) had emphasized salinity and temperature 

tolerances as reasons for the absence of Sesarma cinereum and S. reticulatum 

from southern Florida, Abele (1973) believes that competitive exclusion 

by S. ricordi may account for the absence of the first and possibly the second 

disjunct species. Interspecific competition is an unknown factor in the distribution 

of virtually everj'' brachjûran species, especially the aquatic crabs. An 

exception is the genus Uca, where intensive studies by several authorities over 

the past half century have produced the largest amount of biological data on anj' 

one crab genus. Crane (1975) summarized the zoogeographical and evolutionary 

information on Uca distribution. She remarked on the relatively depauperate 

Atlantic fauna of this genus, compared to the rich and diverse Pacific assemblage, 

also known for many other groups of invertebrates. The differences are attributed 

to cooling during the Oligocene to Pleistocene, which was far more severe in the 

Atlantic than in the Pacific, resulting in the extinction of many sensitive tropical 

species. The isolation of the Caribbean at the end of the Pliocene, due to the 

emergence of Panama, resulted in the Atlantic assemblage, each species of which 

has a Pacific analogue. Only U. subcylindrica, a poorly-known Gulf endemic, 

has no eastern Pacific counterpart. Other Gulf species of Uca, five of which are 

endemic, may be in the process of rapid speciation. The most recentlj^-described 

species, Uca panacea (Novak and Salmon, 1974), may have diverged from U. 

pugilator behaviorally and ecologically with only minimal morphological change. 

Many authorities are reluctant to consider this form a separate species without 

further evidence of isolation or distinction. The same applies to U. virens and 

U. longisignalis (see entries in systematic section), two forms similar to U. 

pugnax of the Atlantic coast. An analysis of courtship displays and habitat requirements 

reveals the divergence of the species not detected from preserved 

specimens alone. This evidence is substantiated by preliminary comparisons of 

isozyme patterns for Atlantic and Gulf populations of U. panacea, U. pugilator, 

U. pugnax, U. virens, and U. longisignalis (Selander, Johnson and Avise, 1971; 

Selander, pers. comm.). Unfortunately, this glimpse of speciation in progress is 

a rare exception to the usual tj^pe of information available. Most accounts of 

zoogeography depend on collection records derived from preserved specimens. 

Misidentifications, erroneous locality labels, and gaps in collection efforts are 

familiar difficulties. 

This contribution is offered as a preliminary step toward compiling the necessary 

information for a synthesis on the distribution and evolution of crabs in the 

Gulf region. It should facilitate the collection of data on particular groups of 

crabs and the comparison of specific lines of evidence relative to ecological, 

behavioral, physiological, or developmental aspects. It can also indicate the rela-


tive lack of information on the majority of species that have been collected and 

formally described, but in which observations on living animals are minimal 

or lacking. Hopefully, it will serve to stimulate increased efforts at erasing these 

defficiencies in our knowledge of some of the most fascinating animals man can 

hope to meet. 

SPECIES ENTRIES 

Infraorder BRACHYURA Latreille, 1803 

SECTION DROMIACEA de Haan, 1833 

Superfamily DROMIOIDEA de Haan, 1833 

Family DROMIIDAE de Haan, 1833 

Dromia Weber, 1795 

Dromia erythropus (George Edwards, 1771) (Cat. Anim. Catesby's Nat. Hist. 

Carolinas, with Linnaean Names) 

Rathbun, 1933, p. 107, fig. 105; Rathbun, 1937, p. 31, text-fig. 11, pi. 6, figs. 1, 2, 

pi. 8, figs. 1, 2; Feldei-, 1973a, p. 44, pi. 6, fig. 2. 

Range: Bermuda; Bahamas; Florida Keys and Dry Tortugas; off Louisiana 

and Texas; north coast of Cuba; Jamaica; Haiti; Puerto Rico; St. Thomas to 

Barbados; Netherlands Antilles; Pemambuco to Sao Paulo, Brazil. 

Depth: shallow water to 360 m (to 197 fm), most common at depths of less 

than46 m (25 fm). 

Habitat: on hard substrates (coral, shell, near rocks); dorsal carapace is always 

covered with sponges or compound ascidians. 

Remarks: Felder (1973) lists collection localities of Seven and One-Half 

Fathom Reef off Texas and a sublittoral prominence about 90 miles south of 

Pecan Island, in the South Atlantic, but a recent (1977) personal communication 

from Dr. Chace indicates that the St. Helena specimens are still not positively 

identified; however, they are not D. erythropus. Coelho and Ramos (1972) 

list this species from Brazil. Hazlett (1971) has examined the antennule chemosensitivity 

of this species. 

Droniidia Stimpson, 1858 

Dromidia antillensis Stimpson, 1858 (Ann. Lye. Nat. Hist. New York 7: 71) 

Hay & Shore, 1918, p. 417, pi. 31, fig. 5; Rathbun, 1933, p. 108, fig. 106; Rathbun, 

1937, p. 33, text-fig. 12, pi. 7, figs. 1-3; Chace, 1940, p. 6; Williams, 1965, p. 143, 

fig. 118; Felder, 1973a, p. 44, pi. 6, figs. 1,3. 

Range: Bermuda; North Carolina; Bahamas; southeast Florida; Florida Keys, 

Straits, and Dry Tortugas; west coast of Florida to Texas; northwest and north 

coasts of Yucatan; off west and south coasts of Cuba; Jamaica; Haiti; Puerto 

Rico; Virgin Islands; Grenada; off Bahia to Espirito Santo, Brazil.

20 LMvrence W. Powers 

Depth: shore to 331 m (to 170 fm). 

Habitat: from hard bottoms (shell, rock, or coral); usually carries a sponge 

or compound ascidian over the dorsal carapace. Uncommon to rare in many 

Gulf areas, but Hildebrand (1955) reported this crab as common on the pink 

shrimp grounds of Campeche, Mexico, in 6 to 16 fm (11 to 29 m) of water. 

Remarks: Rathbun (1937) reported ovigerous females in winter, spring, 

and summer from Florida and the West Indies, and she also listed specimens 

with infestations of bopyrid parasites. Williams (1965) noted crabs that carried 

zoanthoid polyps; specimens from Alligator Harbor in northwest Florida carried 

the ascidian, Eudistoma capsulatum (Wass, 1955). Regional lists include 

Florida (Dragovich and Kelly, 1964; Abele, 1970; Menzel, 1971), Mississippi 

(Franks et al, 1972), Texas (Hildebrand, 1955; Parker, 1959; Leary, 1967), 

and Mexico (Hildebrand, 1955). Felder (1973a) indicates that this species is 

common on Seven and One-Half Fathom Reef, off Texas; Chace (1956) listed 

this species from several Gulf stations of the R/V Oregon. Listed from French 

Guiana by Guinot-Dumortier (1959) and from Brazil by Coelho and Ramos 

(1972). Larval development under laboratory conditions was studied by Rice 

&Provenzano (1966). 

Hypoconcha Guevin, 1854 

Hypoconcha arcuala Stimpson, 1858 (Proc. Acad. Nat. Sci. Philadelphia 1858: 

226) 

Hay & Shore, 1918, p. 418, pi. 31, fig. 3 (not 2); Rathbun, 1937, p. 47, pi. 11, 

figs. 1-4; Williams, 1965, p. 144, fig. 119. 

Range: North Carolina to southern Florida; Dry Tortugas; west coast of Florida 

; St. Thomas, Virgin Islands; Surinam to ELspirito Santo, Brazil. 

Depth: 2 to 40 m (1 to22fm). 

Habitat: sand and shell substrates. Williams (1965) notes that this species is 

always found with a lamellibranch mollusc shell, usually a clam, which it carries 

over its back by its claws and fourth and fifth walking legs. 

Remarks: Listed from Florida by Wass (1955), Dragovich and Kelly (1964), 

Abele (1970), and Menzel (1971). Kircher (1970) studied larval development 

under laboratory conditions. 

Hypoconcha sabulosa (Herbst, 1799) (Vers. Natur. ICrabben u. Krebse, vol. 3, 

p. 57) 

Hay & Shore, 1918, p. 418, pi. 31, fig. 2 (not 3); Rathbun, 1937, p. 44, pi. 8, figs. 

3-4, pi. 9, figs. 1-5; Williams, 1965, p. 145, fig. 120; Felder, 1973a, p. 44, pi. 6. fig. 5. 

Range: off North Carolina; Florida Keys and Dry Tortugas; off Texas; Jamaica; 

Guianas; Moranhao to Bahia, Brazil. 

Depth: 1 to 90 m (to 49 fm). 

Habitat: from sand, shell, and coral bottoms; Williams (1965) states that the 

habits of this crab are similar to those of H. arcuata, but that it is a much rarer 

species.


Remarks: Listed from Florida by Hulings (1961) and Abele (1970) and from 

Brazil by Coelho and Ramos (1972). 

Uypoconcha spinosissima Rathbun, 1933 (Proc. Biol. Soc. Washington 46: 185) 

Rathbun, 1937, p. 46, text-fig. 14, pi. 10, figs. 1-2; Felder, 1973a, p. 4+, pi. 6. fig. 4. 

Range: off North Carolina; Dry Tortugas; west coast of Florida; ? Texas; off 

north coast of Yucatan; Jamaica. 

Depth: 26 to 110 m (14to60fm). 

Habitat: broken coral or shell, sand bottoms. 

Remarks: Although listed by Leary (1967) for Texas, I know of no published 

collection records to verify its presence in the northwestern Gulf. Felder (1973a) 

includes this species, based on the listing by Leary. 

Family PROSOPIDAE von Meyer, 1860 

Subfamily HOMOLODROMIINAE Alcock, 1899 

(Glaessner, 1969, corrected the family name Prosoponidae of von Meyer, 

1860 and listed three subfamilies: Prosopinae, Pithonotinae, and Homolodromiinae. 

Only the latter subfamily is extant and many authors treat 

this group as a family.) 

Dicranodromia A. Milne Edwards, 1880 

Dicranodromia ovala A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 32) 

Rathbun, 1937, p. 60, text-fig. 15, pi. 13, figs. 3-4; Chace, 1940, p. 7; Pequegnat, 

1970, p. 173. 

Range: east and west coasts of Florida; Florida Keys and Straits; off north 

coast of Cuba; northwest Caribbean Sea; Guadeloupe; Barbados. 

Depth: 128 to 895 m (70 to490 fm). 

Habitat: no data available. 

Remarks: Rathbun (1937) reported ovigerous females from Florida in June 

and Chace (1940) reported ovigerous females from Cuba in early May. 

Homolodromia A. Milne Edwards, 1880 

Homolodromia paradoxa A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 

33) 

Rathbun, 1937, p. 58, pi. 13, figs. 1-2, pi. 14, figs. 1-4; Chace, 1940, p. 7. 

Range: north coast of Cuba; off Nevis, Leeward Islands 

Depth: 651 to 896 m (356 to 490 fm), possibly to 1106 m (605 fm). 


Superfamily HOMOLOIDEA White, 1848 

Family HOMOLIDAE White, 1847


Hoinola Leach, 1815 

Honiola barbata (Fabricius, 1793) (Entomol. system., vol. 2, p. 460) 

As Thelxiope barbata—Rathbun, 1937, p. 63, text-fig. 16, pi. 15, figs. 1-2; Chace, 

1940, p. 8; Barnard, 1950; p. 338, fig. 65d-e. 

As Homola barbala—Williams, 1965, p. 146, fig. 121. 

Range: Massachusetts to southern Florida; Florida Keys and Dry Tortugas; 

north coast of Cuba; off east coast of Yucatan (Caribbean); in eastern Atlantic, 

off Naples, Portugal, Azores, Madeira Islands; off South Africa. 

Depth: 55 to 682 m (30 to 373 fm). 

Habitat: sand, shell, and coral substrates; occasionally on mud bottoms. 

Remarks: The genus names Thelxiope and Homola have been interchanged 

by various authors up to 1958, when the latter name was adopted for genus, 

family, and superfamily designation. Gordon (1950) described reproductive 

structures and the evolution of this genus among the Dromiacea. Rice (1964) 

and Rice and Provenzano (1970) studied larval development. Ovigerous females 

occur in June—July off North Carolina and Florida (Williams, 1965) and in 

October off Delaware (Rathbun, 1937). Hartnoll (1970, 1971) noted swimming 

behavior. 

Homola vigil A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 33) 

As Thelxiope wg-JI—Rathbun, 1937, p. 66, pi. 16, figs. 1-3; Chace, 1940, p. 9. 

Range: off Georgia; north and south coasts of Cuba; Guadeloupe; Martinique. 

Depth: 309to804m (169to440fm). 

Habitat: sand, broken shell, and coral bottoms. 

Remarks: Chace (1940) reported ovigerous females from Cuba in April and 

May. 

Homologenus A. Milne Edwards, 1888 

Hotnologenus roslralus (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 

34) 

Rathbun, 1937, p. 70, text-fig. 17, pi. 17, figs. 1-3; Chace, 1940, p. 9; Pequegnat, 

1970, p. 174, fig. 6-1. 

Range: Bahamas; east coast of Mexico in southwest Gulf of Mexico; north and 

south coasts of Cuba; Virgin Islands; near Aves Island (Lesser Antilles); Azores; 

off Morocco. 

Depth: 600 to 1600m (330 to875 fm). 

Habitat: fine sand, mud, and ooze substrates. 

Remarks: Ovigerous females were reported from Cuba in March and May 

(Chace, 1940) and from the Windward Passage in April (Rathbun, 1937). 

Hypsophrys Wood-Mason, 1891 

Hypsophrys noar Williams, 1974 (Proc. Biol. Soc. Washington 87: 485) 

Williams, 1974b, p. 485, figs. 1-12. 

Range: southwest of Dry Tortugas, in Florida Straits.


Depth: 732 m (400 fm). 

Remarks: Williams (1974b) compares this species, known only from the type 

specimen, with the other two species of Hypsophrys. The male holotype carried 

a number of small barnacles, Poecilasma inaequilaterale, on the abdominal setae 

and left cheliped. 

Family LATREILLIIDAE Alcock, 1899 

(This family, including the genera Latreillia Roux and Latreillopsis 

Henderson, was recognized by Ralhbun (1937), but these genera were 

included in the Homolidae by Balss (1957). Glaessner (1969) leaves the 

position of these genera undetermined.) 

Latreillia Roux, 1830 

Latreillia elegans P. Roux, 1830 (Crust. Mediterranea et de son littoral, pi. 22, 

1828(1830)) 

Hay & Shore, 1918, p. 419, pi. 31, fig. 4; Rathbun, 1937, p. 73, text-fig. 18, pi. 20, 

pi. 21, figs. 1-8; Chace, 1940, p, 10; Williams, McCIoskey & Gray, 1968, p. 42, fig. 1. 

Range: Massachusetts to North Carolina; off South Florida; Florida Keys; 

north coast of Cuba; eastern North Atlantic Ocean; Mediterranean Sea; off 

Natal, South Africa. 

Depth: 46 to 366 m (25 to 200 fm). 

Habitat: sand, shell, and coral bottoms; from soft mud; off sponges. 

Remarks: Rathbun (1937) reported ovigerous females from Massachusetts in 

August and from Florida in February. 

SECTION OXYSTOMATA H. Milne Edwards, 1834 

Superfamily RANINOIDEA de Haan, 1841 

Family RANINIDAE de Haan, 1841 

(This family is treated as a "subtribe" Gymnopleura of the "tribe" 

Brachjaira by Bourne (1922). Balss (1957) and Glaessner (1969) include 

this group as family and superfamily within the Oxystomata. The systematic 

position of the raninids depends, in part, on the relative degree 

of specialization and primitiveness assgned to the morphological characters 

of these aberrant crabs.) 

Lyreidus de Haan, 1841 

Lyreidus hairdii Smith, 1881 (Proc. U.S. Nat. Mus. 3: 420) 

Rathbun, 1937, p. 23, pi. 5, figs. 5-6; Chace, 1940, p. 6; Pequegnat, 1970, p. 180. 

Range: Massachusetts; Dry Tortugas; west coast of Florida; off Louisiana, 

Texas, and Mexico; north coast of Cuba; north of Puerto Rico. 

Depth: 119 to 823 m (65 to450 fm). 

Habitat: soft mud substrates.


Remarks: Chace (1940) reported a specimen with rhizocephalan parasites 

from off the north coast of Cuba. Listed from several stations of the R/V Oregon 

by Chace (1956) in the Gulf of Mexico. Pequegnat (1970) considered this species 

to be the most common raninid in the Gulf and he provides some data on 

densities at different depths along the continental slope. 

Ranilia H. Milne Edwards, 1837 

Ranilia conslricla (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 35) 

Rathbun, 1937, p. 20, pi. 4, fig. 5, pi. 5, figs. 1, 2; Gomes Correa, 1970, p. 2; 

Pequegnat, 1970, p. 180. 

Range: Florida Straits; off north coast of Cuba and southeast Gulf of Mexico; 

Rio de Janeiro, Brazil; eastern Atlantic, Ascension Island; Senegal to Congo. 

Depth: off shallow reef (Rathbun, 1937); 183 to 336 m (100 to 200 fm) in 

Gulf of Mexico (Pequegnat, 1970); original type is from 86 m (47 fm), off 

Florida. 

Habitat: coral reefs; hard bottoms in deep water. 

Remarks: Listed from Brazil by Coelho and Ramos (1972). Hartnoll (1971) 

cites an observation by Darwin on a species of Ranilia in the southern Atlantic 

in which swimming was noted, but the extent to which raninids are able to swim 

is not known. 

Ranilia muricala H. Milne Edwards, 1837 (Hist. nat. Crust., vol. 2, p. 196) 

Hay & Shore, 1918, p. 420, pi. 31, fig. 1; Rathbun, 1937, p. 18, pi. 3, figs. 3-6, pi. 4, 

figs. 1-4; WiUiaras, 1965, p. 142, fig. 117. 

Range: North Carolina; Bahamas; Florida Straits; southern to northwestern 

Florida; Swan Island (Caribbean). 

Depth: 9 to 102m (5 to56fm). 

Habitat: offshore, on sandy and broken shell substrates. 

Remarks: Listed from Florida by Wass (1955), Abele (1970) and Menzel 

(1971). Rathbun (1937) listed ovigerous females from North Carolina in September. 

This species has been recovered from fish stomachs in North Carolina 

offshore waters (Williams, 1965). 

Raninoides H. Milne Edwards, 1837 

Raninoides lainarcki A. Milne Edwards & Bouvier, 1923 (Mem. Mus. Comp. 

Zool. 47: 299) 

Ratlibun, 1937, p. 13, text-fig. 8, pi. 1, figs. 3, 4; Chace, 1940, p. 5. 

Range: north of Cuba; north of Puerto Rico; off Colon, Panama (Caribbean). 

Depth: 46 to 366 m (25 to 200 fm). 


Remarks: Chace (1940) notes an error in plate 2 of Rathbun, 1937: figure 3 

is a chela of R. lamarcki and not R. fossor, as the label indicates. The error was 

due to a label interchange on figures a and b of Milne Edwards and Bouvier's 

(1923) original drawing. Manning (1975) has subsequent^ indicated that R.


fossor is a synon3^m of an Indo-West Pacific species, Notosceles chimmonis 

Bourne. 

Raninoides loevis (Latreille, 1825) (Encycl. meth., Hist, nat., vol. 10, p. 268) 

Rathbun, 1937, p. 8, text-fig 3, pi. 1, figs 1, 2; Guinot-Dumortier, 1959, p. 246 

fig. 2a-c; Gomes Correa, 1970, p. 9. 

Range: Florida Keys and Dry Tortugas; southwest coast of Florida; Campeche 

Ba}^, off Tabasco, Mexico; Barbados; Colombia (Caribbean); Guianas to Bahia, 

Brazil; Pacific coasts of Panama and Colombia. 

Depth: 18tol96m (lOtolOZfm). 

Habitat: bottom types include ooze, mud, shelly mud, coral, and broken shell. 

Remarks: Listed from the R/V Oregon collections in the Gulf of Mexico by 

Chace (1956). Guinot-Dumortier (1959) and Knight (1968) compared this 

species with R. benedicti Rathbun. Listed from Brazil by Coelho and Ramos 

(1972). 

Raninoides louisianensis Rathbun, 1933 (Proc. Biol. Soc. Washington 46: 186) 

Common Name: Frog Crab 

Rathbun, 1937, p. 12, text-figs. 6, 7, pi. 1, figs. 5, 6; Pequegnat, 1970, p. 181; Felder, 

1973a, p. 38, pi. 4, fig. 6. 

Range: Gulf of Mexico, from the Mississippi Delta to Campeche Banks. 

Depth: 55 to 400 m (30 to 220 fm). Collection records of the R/V Alaminos 

showed an extension of this species on the middle and outer continental shelf 

to only 115 fm and Pequegnat (1970) believes that the R/V Oregon records at 

200 and 220 fm may be due to trawl contamination from earlier, shallower 

stations. 

Habitat: muddy and fine sand-mud bottoms. 

Remarks: Listed from Texas by Leary (1967) and from the Gulf of Mexico by 

Chace (1956). Ovigerous females were taken in February, June, July, and 

October by the R/V Alaminos (Pequegnat, 1970). Franks et al. (1972) reported 

salinity and temperature ranges of collections off Mississippi. 

Symelhis Weber, 1795 

Symethis variolosa (Fabricius, 1793) (Entomol. System, emend, et aucta, vol. 

2, p. 476) 

Rathbun, 1937, p. 26, text-fig. 10, pi. 5, figs. 7, 8; Gomes Correa, 1970, p. 10. 

Range: North Carolina; southeast Florida; Florida Kej^s and Dry Tortugas; 

north of Puerto Rico; Fernando do Noronha to Bahia, Brazil; Pacific coast of 

Panama. 

Depth: 18 to 110 m (10 to 60fm). 

Habitat: sand, mud bottoms; on calcareous algae; under stones. 

Remarks: Cerame-Vivas and Gray (1966) extended the previously known 

range of this species to North Carolina. Listed from Bra2il by Coelho and Ramos 

(1972) and by Fausto Filho (1974).


Superfamily DORIPPOIDEA deHaan, 1841 

Family DORIPPIDAE de Hann, 1841 

Clythrocerus A. Milne Edwards & Bouvier, 1899 

Clythrocerus nitidus (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 24) 

Rathbun, 1937, p. 109, text-figs. 26,27, pi. 33, figs. 1-2. 

Range: South Carolina; southeast and northwest Florida; Florida Keys; 

Grenada. 

Depth: 12 to 479 m (6.5 to 262 fm). 

Habitat: rock, coral, sand, shell, and gravel bottoms. 

Remarks: Listed by Wass (1955) at 25 fm off Cape San Bias, Florida. Rathbun 

(1937) reported ovigerous females from Florida in February, March, and late 

June. 

Clythrocerus slirnpsoni Rathbun, 1937 (Bull. U.S. Nat. Mus. 166: 121) 

Rathbun, 1937, pi. 121, text-fig. 32, pi. 34, figs. 5, 6. 

Range: west coast of Florida. 

Depth: 183 m (100 fm). 

Remarks: Known only from the single female type specimen. 

Corycodus A. Milne Edwards, 1880 

Corycodus hullatus A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 23) 

Rathbun, 1937, p. 103, pi. 29, figs. 1-4, pi. 30, fig. 1, pi. 31, fig. 1. 

Range: off north coast of Cuba. 

Depth: 320 to457m (175 to250fm). 

Cyclodorippe A. Milne Edwards, 1880 

Cyclodorippe anlennaria A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 

25) 

Rathbun, 1937, p. 104, text-fig. 24, pi. 32, figs. 1, 2; Pequegnat, 1970, p. 177. 

Range: west coast of Cuba; north coast of Yucatan (Gulf); north coast of Cuba; 

Puerto Rico; Lesser Antilles, from Dominica to Grenada. 

Depth: 91 to 653 m (50 to 357 fm). 

Habitat: primarily hard bottoms (coral, sand, and shell). 

Cyclodorippe bouvieri Rathbun, 1934 (Smithsonian Misc. Coll. 91: 1) 

Rathbun, 1937, p. 106, pi. 32, figs. 3-4, pi. 81, figs. 1-2. 

Range: north coast of Cuba; northeast of Puerto Rico. 

Depth: 274 to 549 m (150 to 300 fm). 

Remarks: Rathbun (1937) lists an ovigerous female from Puerto Rico, taken 

in March at 150 fm.

CycZorforippe orn«fa Chace, 1940 (Torreia 3: 19) 

Range: Florida Straits; Florida Keys; Puerto Rico. 

k 

Depth: 128 to 549 m (70 to 300 fm). 

Habitat: sand and rocky bottoms. 


Chace, 1940, p. 19, figs. 7, 8. 

Range: off north coast of Cuba. 

Depth: 375 to 439 m (205 to 240 fm). 

Remarks: Chace (1940) reported an ovigerous female from off Cuba, taken in 

May at 240 fm. 

Cymonomus A. Milne Edwards, 1880 

Cymonomus caecus Chace, 1940 (Torreia 3: 12) 

Chace, 1940, p. 12, figs. 1-2. 

Range: north coast of Cuba. 

Depth: 841m (460fm). 

Cymonomus cubensis Chace, 1940 (Torreia 3: 16) 

Chace, 1940, p. 16, figs. 5-6. 


Depth: 475 to 1006 m (260 to 550 fm). 

Remarks: Chace (1940) reported ovigerous females from off Cuba in March 

and May. He feels that this species is a possible link between the genera 

Cymonomus and Cymopolus and it may be eventually elevated to generic status 

when studied further. 

Cymonomus quadratus A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 26) 

Rathbun, 1933, p. 106, fig. 104; Rathbun, 1937, p. 98, le.xt-fig. 23, pi. 30, fig. 3, 

pi. 31, fig. 3. 

Range: northwest of Dry Tortugas; west, north and south coasts of Cuba; 

Puerto Rico; Lesser Antilles, from St. Croix to Grenada. 

Depth: 185to929m(101to508fm). 

Habitat: soft bottoms, fine sand, mud, and ooze. 

Cymonomus roslralus Chace, 1940 (Torreia 3: 14) 

Chace, 1940, p. 14, figs. 3, 4. 


Depth: 658m (360fm). 

Cymopolus A. Milne Edwards, 1880 

(This genus should not be confused with Cymopolia Roux, a synonym 

of Paliciis Phillipi, of the family Palicidae) 

Cymopolus af;assizi A. Milne Edwards & Bouvier, 1899 (Bull. Mus. Hist. Nat., 

Paris 5: 385) 

Rathbun, 1937, p. lOO, pi. 30, fig, 2, pi. 31, fig. 2.

28 Lawrence W, Powers 

Remarks: Rathbun (1937) lists ovigerous females from off Florida in February 

and March. 

£«/»MsaRoux, 1828 

Ethusa tnascarone americana A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 

8: 30) 

Rathbun, 1933, p. 105, fig. 102; Rathbun, 1937, p. 78, pi. 22, fig. 2, pi. 23, fig. 2; 

Williams, McCloskey & Gray, 1968, p. 43, fig. 2. 

Range: North Carolina; Florida Keys and Dry Tortugas: west coast of Florida; 

Puerto Rico; Virgin Islands; Maranhao to Bahia, Brazil; Gulf of California 

and the Pacific coast of Panama. 

Depth: shallow water to 82 m (to 45 fm). 

Habitat: rock, coral, coarse shell, and sand substrates; from surfaces of algae, 

bryozoans, and seaweeds. 

Remarks: Abele (1970) remarks on the need for revision of this species and 

its subspecies, noting the variation in growth of critical characters, also cited 

by Finnegan (1931) and Garth (1966). Wilhams, McCloskey and Gray (1968) 

observed a specimen in the laboratory, taken from a reef off North Carolina, 

that clasped objects over its carapace, in the manner of dromiid crabs and similar 

to that reported for E. lata in the Pacific (Garth, 1946). Listed from Brazil 

by Rodrigues da Costa (1968a) and Coelho and Ramos (1972). 

Elhusa microphthalma Smith, 1881 (Proc. U.S. Nat. Mus. 3: 418) 

Rathbun, 1937, p. 82, pi. 22, fig. 3, pi. 23, fig. 3; Chace, 1940, p. 10; Pequegnat, 

1970, p. 175. 

Range: Massachusetts to North Carolina; Dry Tortugas; off west coast of 

Florida; ofi Mississippi; off east coast of Mexico and Tabasco (Gulf); north and 

south coasts of Cuba; northeast Caribbean Sea. 


Habitat: soft bottoms, fine sand, mud, and mud with shell. 

Remarks: Listed from the Gulf of Mexico by Chace (1956) and from Texas 

by Leary (1967), but not by Felder (1973a) who didn't include deep sea species. 

Rathbun (1937) reported ovigerous females from off Dry Tortugas in July. 

Ethusa tenuipes Rathbun, 1897 (Proc. Biol. Soc. Washington 11: 110) 

Rathbun, 1937, p. 87, pi. 24, fig. 3, pi. 25, fig. 3; Chace, 1940, p. 11; Williams, 

McCloskey & Gray, 1968, p. 44. 

Range: off North Carolina; east coast of Florida; Florida Keys and Dry Tortugas; 

off Alabama; north and south coasts of Cuba. 

Depth: 46 to 402 m (25 to 220 fm). 

Habitat: sand and coral bottoms. 

Remarks: Rathbun (1937) reported ovigerous females from North Carolina 

in July, from off Cuba in April, and from off Dry Tortugas in July.


Ethusa truncata A. Milne Edwards & Bouvier, 1899 (Bull. Mus. Hist. Nat., 

Paris 5: 384) 

Rathbun, 1937, p. 85, pi. 28, figs. 1-2. 

Range: off west coast of Florida; off Mississippi Delta and Louisiana; northwest 

of Trinidad. 

Depth: 133to219m (73toll9fm). 

Ethusina Smith, 1884 

Ethusina abyssicola Smith, 1884 (Rept. U.S. Comm. Fish Fisher. 1882, p. 349 

(1884)) 

Rathbun, 1937, p. 91, text-fig. 21, pi. 26, fig. 1, pi. 27, fig. 1; Pequegnat, 1970, 

p. 175, fig. 6-2. 

Range: Massachusetts to North Carolina; in deep waters of northwest, northeast, 

and southwest quadrants of Gulf of Mexico; off Cape Frio, Brazil; off west 

coast of Spain. 

Depth: 860 to4061 m (470 to2220 fm). 

Habitat: very soft bottoms, muds and oozes. 

Remarks: Pequegnat (1970) presents e\'idence for two modes of bathymetric 

distribution, one at 860 to 1399 m (470 to 765 fm) with specimens resembling 

typical E. abyssicola, and a deeper mode at 2551 to 4061 m (1395 to 2220 fm) 

with specimens that approach E. faxonii in size and shape of exorbital teeth. 

Further studies may reveal two distinct species. Pequegnat (1970) also reports 

that an ovigerous female with a few advanced embryos was collected at 765 fm 

in mid-November. 

Superfamily CALAPPOIDEA deHaan, 1833 

Family CALAPPIDAE de Haan, 1833 

Subfamily CALAPPINAE de Haan, 1833 

Acanthocarpus Stimpson, 1871 

Acanthocarpus alexandri Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 153) 

Rathbun, 1937, p. 221, pi. 69, figs. 1-2; Chace, 1940, p. 26; Williams, 1965, p. 156, 

fig. 137; Pequegnat, 1970, p. 177, fig. 6-3. 

Range: Massachusetts; North Carolina to south Florida; Florida Keys and 

Dry Tortugas; west and northwest Florida; Mississippi; Texas; east coast of 

Mexico; north coast of Cuba; Puerto Rico to the Grenadines; off Brazil. 

Depth: 68 to 476 m (37 to 260 fm). 

Habitat; primarily soft bottoms, fine sand, mud and ooze. 

Remarks: Chase (1940) and Pequegnat (1970) note that the carapace of this 

k 

• spe 

species is broader than long, contrary to the description of Rathbun (1937). 

Chace (1956) listed this species from the Gulf of Mexico collections of the R/V


Oregon and Pequegnat (1970) reported this crab as the most abundant deep 

water calappid from the Alaminos collections. Rathbun (1937) listed ovigerous 

females from off Florida in June and Pequegnat (1970) listed the same from 

the deep Gulf in early August to mid-November. Listed from Brazil by Coelho 

and Ramos (1972). 

Acanthocarpus bispinosus A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 

19) 


Range: off west and northwest coasts of Florida; Dry Tortugas; Grenadines, 

Windward Islands. 

Depth: 201 to360m (110tol97fm). 

Habitat: mud-shell, coral, and clay-mud bottoms. 

Remarks: The depth and location records off Florida for the R/V Oregon stations 

(Chace, 1956) may be confused. Stations 1007 and 1010 are positioned well 

within the 100 fm isobath, yet the depths for these stations are listed at 180 and 

225 fm, respectively. 

Calappa Weber, 1795 

Calappa angusta A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 18) 

Hay & Shore, 1918, p. 421, pi. 31, fig. 7; Rathbun, 1937, p. 210, pi. 64, figs. 1-6; 

Williams, 1965, p. 154, fig. 134; Pequegnat, 1970, p. 177. 

Range: North Carolina; southeast Florida; Florida Keys and Dry Tortugas; 

west coast of Florida and mid-eastern Gulf of Mexico; off east coast of Mexico; 

off North coast of Yucatan; St. Thomas to Grenada; Barbados. 

Depth: 13 to274m (7 to 150fm). 

Habitat: coral, sand, broken shell, and gravel substrates. 

Remarks: Rathbun (1937) reported ovigerous females from southern Florida 

in Aferch. Chace (1956) recorded this species from the eastern Gulf of Mexico 

and Williams (1965) stated that this crab is more abundant in the Gulf Stream 

than in adjacent inshore waters. Shoup (1968) described shell opening behavior 

by this species. 

Calappa flaniniea (Herbst, 1794) (Versuch Naturgesch. Krabben u. Krebse, vol. 

2, p. 161) 

Common Names: Flame-streaked Box Crab; Shame-faced Crab 

Hay & Shore, 1918, p. 421, pi. 31, fig. 8; Rathbun, 1933, p .103; Rathbun, 1937, 

p. 198 (part), pi. 59, figs. 1-2, pi. 60, fig. 1; Reed, 1941, p. 44; Holthuis, 1958, 

p. 148, figs. 28-35; Williams, 1965, p. 152, figs. 130-131; Felder, 1973a, p. 43, 

pi. 5, fig. 11. 

Range: Massachusetts to south Florida (see Remarks); Bermuda; Bahamas; 

Florida Straits and Keys; Dry Tortugas; west and northwest Florida; all inshore 

Gulf of Mexico areas from Florida to Yucatan, Mexico. 

Depth: shore to 73 m (to 40 fm), rarely to 229 m (125 fm). 

Habitat: hard bottoms, primarily sand.


Remarks: The breeding range for this crab extends only to North Carolina, 

but larvae as far north as New England may occasionally survive a mild winter 

to provide temporary range extensions (Hay and Shore, 1918; Holthuis, 1958). 

Some of the larval stages are figured by Lebour (1944). This species was revised 

by Holthuis (1958), who found at least two species that had been previously 

combined under this name. One of these, C. ocellata, also occurs in the Gulf of 

Mexico, thus previous records, particularly from off west Florida, may refer to 

either or both species. Listed from the R/V Oregon collections in the Gulf bj^ 

Chace (1956). Cheliped modifications associated with shell opening of molluscs 

were described bjr Shoup (1968). 

Calappa gallus (Herbst, 1803) (Versuch Naturgesch. Krabben u. Krebse, vol. 3, 

pt. 3, pp. 18 and 46) 

Common Name: Yellow Box Crab 

Rathbun, 1933, p. 103; Rathbun, 1937, p. 214, pi. 65, figs. 2-3: Barnard, 1950, p. 350, 

fig. 66e-i; Sakai, 1965, p. 55, pi. 21, fig. 3; Fausto Filho, 1967, p. 48, fig. 4, pi. VI, 

figs. 7-8; Sakai, 1976b, p. 131, pi. 39, fig. 2. 

Range: Bermuda; Bahamas; Florida Keys and Dry Tortugas; northwest Cuba; 

Jamaica; Puerto Rico; St. Croix to Barbados; off Campeche snapper banks (Gulf 

of Mexico); Panama (Carib.) to Venezuela; Netherlands Antilles; Ceara to 

Bahia, Brazil; St. Helena Island (So. Atlantic); off western Africa, from Senegal 

to Angola; South Africa; Red Sea; Reunion and Seychelles, in Persian Gulf; off 

India and Maldives; Philippinss; Formosa; Japan; Marshall Inlands; Samoa; 

Hawaiian Islands. 

Depth: low tide mark to 218 m (to 119 fm). 

Habitat: hard substrates; on reef flats; coral, sand, shell and rock bottoms. 

Remarks: Barnard (1950) and Sakai (1965) provide some earlier references 

lor the African and Asian areas, respectivelj^ Coelho and Ramos (1972) and 

Fausto Filho (1974) list Brazilian records for this species. 

Calappa ocellata Holthuis, 1958 (Stud. Fauna Curagao 8: 158) 

Rathbun, 1901, p. 84 (part), pi. 2; Verrill, 1908, p. 420 (part), pi. 25, fig. 1; 

Rathbun, 1937, p. 198 (part), not pi. 59 or 60; Holthuis, 1958, p. 158, figs. 36-40; 

Williams, 1965, p. 153, figs. 132-133. 

As C. oceJata—Fausto Filho, 1967, p. 42, fig. I, pi. 1, figs. 1-2. 

Range: Bermuda; off North Carolina and in Beaufort Harbor (rare); Florida 

Keys and Dry Tortugas; Jamaica; Hispaniola; Puerto Rico; Virgin Islands; 

Caribbean coasts of Panama and Colombia; Netherlands Antilles; Rocas to Pernambuco, 

Brazil. 


Remarks: Plates 59 and 60 of Rathbun (1937) are of C. flammea, not C. 

ocellata. Other records of C. flammea may also include specimens of this species, 

especially reports prior to the revision bj^ Holthuis (1958). Shoup (1968) described 

shell opening behavior by this species and other Calappa. Coelho and 

Ramos (1972) list this crab from Brazil.


Calappa springeriVia\hhun, 1931. 

This species proved to be identical to C. sulcata Rathbun, 1898 when examined 

by Holthuis (1958) and so this name is a junior synonym. Rathbun's earlier 

description was based on a juvenile of the species and her later description, as 

C. springeri, was based on an adult form.. Many of the earlier Gulf of Mexico 

faunal surveys list C. springeri and these can all be referred to Calappa sulcata. 

Calappa sulcata Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 4: 289) 

Common Names: Yellow Box Crab; Shame-faced Crab; Parrot Crab. 

As C. sprmgeri—Rathbun, 1937, p. 205, pi. 60, fig. 1, pi. 61, figs. 1-2. 

As C. sa/c«l«—Rathbun, 1933, p. 103, fig. 99; Rathbun, 1937, p. 211, pi. 64, figs. 

7-8, pi. 65, fig. 1; Holthuis, 1958, p. 179, figs. 51-54; Williams, 1965, p. 155, figs. 

135-136; Fausto Filho, 1967, p. 46, fig. 3, pi. II, figs. 5-6; Felder, 1973a, p. 42, 

pi. 5, fig. 10. 

Range: North Carolina; Chesapeake Bight; Dry Tortugas; Alabama to south 

Texas; Tabasco, Mexico; Puerto Rico; Venezuela to Surinam; Amapa to Sergipe, 

Brazil. 


Habitat: sand and sand-mud bottoms. 

Remarks: Listed from the Gulf of Mexico (some under C. springeri) by Chace 

(1956) and Fotheringham and Brunenmeister (1975), from Mississippi (Franks 

et al, 1972), and off Texas (Gunter, 1950; Hildebrand, 1954; Leary, 1967). 

Hildebrand (1954) reported ovigerous females from Texas in May and that 

claws of this crab were occasionally served at restaurants in Port Aransas. A 

report of mollusc shell-opening by this and other Calappa species (Shoup, 1968) 

provides one of the few behavioral studies. Listed from Chesapeake Bight by 

Musick and McEachran (1972) and from Brazil by Coelho and Ramos (1972). 

Cycloes de Haan, 1837 

Cycloes bairdii Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 237) 

Rathbun, 1933, p. 101, fig. 98; Rathbun, 1937, p. 225, pi. 69, figs. 3-4; Garth, 1946, 

p. 362, pi. 62, figs. 7-8; Williams, McCloskey & Gray, 1968, p. 49, fig. 6. 

V 

Range: Bermuda; Bahamas; North Carolina; southeast Florida; Florida Keys 

and Dry Tortugas; Florida Straits; west coast of Florida; Cuba; Puerto Rico; St. 

Thomas to Barbados; Old Providence Island (Carib.); in the Pacific, from west 

coast of Mexico to Ecuador; Rocas to Espirito Santo, Brazil. 

Depth: 3 to 229 m (1.5 to 125 fm). 

Habitat: sand, rock, coral, and shell bottoms; buries in sand. 

Remarks: Guinot-Dumortier & Dumortier (1961) described a stridulatory 

apparatus in this species and within the genus. Juvenile forms of this crab from 

Brazil were described by Rodrigues da Costa (1968b) and this species is also listed 

from Brazil by Coelho and Ramos (1972) and by Fausto Filho (1974). 

Subfamily MATUTINAE Macleay, 1838 

(Several genera of this subfamily, including Hepatus and Osachila, were


examined by Guinot (1966) and aligned with Aethra of the Parthenopidae, 

along with Actaeomorpha of the Leucosiidae. She proposed a new 

subfamily, Aethrinae, to contain these genera, pending further studies. 

The status of these changes is still in doubt, as is the status of the genus 

Matuta Weber, which is not represented in the Gulf of Mexico. Until 

such studies are available, Hepatus and Osachila are included within the 

Matutinae and they are listed with the other calappids, while recognizing 

their probable relationship to the Parthenopidae.) 

Hepalus Latreille, 1802 

Hepalus ephelilicus (Johansson, in Linnaeus, 1763) (Amoemtates academicae, 

etc., vol. 6, p. 414) 

Common Names: Calico Crab; Leopard Crab; Dolly Varden Crab 

Hay & Shore, 1918, p. 422, pi. 37, fig. 1; Rathbun, 1937, p. 238, pi. 70, figs. 3-4, 

pi. 71, figs. 1-4; Williams, 196S, p. 158, fig. 140; Felder, 1973a, p. 43, pi. 5, fig. 14. 

Range: Chesapeake Bay to south Florida; Florida Keys and Dry Tortugas; 

west coast of Florida to south Texas; Campeche Banks, off Yucatan; Cuba; 

Jamaica; Dominican Republic. 

Depth: near shore to 46 m (to 25 fm). 

Habitat: sand, sand-shell, and mud-sand substrates; found inside passes, channels, 

and harbors, but more common in shallow, open marine waters. Buries in 

substrate, probablj^ nocturnal. Frequently collected with sea anemones attached 

to dorsal carapace. 

Remarks: Calliactis tricolor is the most common anemone found on this crab, 

usually a single anemone located in the middle of the anterior margin, where 

the exhalent current of the crab induces a current over the anemone's basal disc 

(Carlgren and Hedgpeth, 1952). Larval development of this crab was studied by 

Costlow and Bookhout (1962). Gvaj (1957) measured the total gill area. Considerable 

variation exists in the dorsal carapace color patterns: some are spotted 

and others are marked with horizontal bands, including continuous gradations 

between these forms. Ovigerous females are not often collected, but have been 

reported off Texas in July (Hildebrand, 1954). Regional lists include Florida 

(Wass, 1955; Dragovich and Kelly, 1964; Abele, 1970; Menzel, 1971), Mississippi 

(Richmond, 1962; Franks et ah, 1972), Louisiana (Behre, 1950; Hoese and 

Valentine, 1972), Texas (Gunter, 1950; Hedgpeth, 1953; Hildebrand, 1954; 

Parker, 1959; Breuer, 1962; Leary, 1967), Campeche (Hildebrand, 1955), and 

offshore waters of the Gulf (Chace, 1956). Fotheringham and Brunenmeister 

(1975) summarize some of the natural history of this crab. Guinot (1966) 

reviews the taxonomic status of this and related genera. 

Hepalus princeps (Herbst, 1794). 

This name was determined to be a junior synonym of H. pudibundus (Herbst, 

1785) in a revision by Holthuis (1959, p. 167). Rathbun (1937) and earlier 

regional surveys use the junior name and these are referred to H. pudibundus.


Hepatus pudibundus (Herbst, 1785) (Versuch Naturgesch. Krabben u. Krebse, 

vol. l,p. 199) 

As H. princeps—Rathbun, 1933, p. 104, fig. 101; Rathbun, 1937, p. 235, pi. 70, 

figs. 1-2. 

As H. pudibundus—Holthviis, 1959, p. 167, figs. 36-37, 38a-b; Williams, 1965, 

p. 157, figs. 138-139; Guinot, 1966, p. 755, figs. 9, 18, 30; Fausto Filho, 1967, p. 50, 

fig. 5, pi. II, figs. 9-10; Felder, 1973a, p. 43, pi. 5, fig. 13. 

Bange: off Georgia; off Louisiana and Texas; north and south coasts of Cuba; 

Jamaica; Haiti; Puerto Bico; St. Thomas to Guadeloupe; Panama (Carib.); 

Surinam; Bahia to Santa Catarina, Brazil; off Guinea, western Africa; Cape of 

Good Hope, South Africa. 


Habitat: very shallow waters with sand-mud and shell-mud bottoms. Often 

carries anemones and barnacles on carapace. 

Bemarks: The systematic status of this species and the genus have been 

reviewed by Holthuis (1959) and by Guinot (1966). Holthuis reported ovigerous 

females from Surinam in April. Behre (1950) collected this crab at Grand Isle, 

Louisiana, only once or twice, from among oyster beds. Leary (1967) lists this 

species from Texas and Felder (1973a) confirms this with a record of his own. 

Nomura and Fausto Filho (1966) reported biometric data on this crab from 

Brazil and additional Brazilian records were provided by Bodrigues da Costa 

(1968a) and Coelho and Bamos (1972). 

Osachila Stimpson, 1871 

(See remarks under Matutinae with regard to this genus and Hepatus.) 

Osachila antillensis Bathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 4: 

290) 

Rathbun, 1933, p. 104, fig. 100; Rathbun, 1937, p. 251, pi. 77, fig. 2. 

Bange: north coast of Cuba; St. Croix, Virgin Islands; Montserrat; Dominica; 

Barbados; Grenada. 

Depth: 123 to 304 m (67 to 164 fm). 

Habitat: sand, shell, coral, and rock substrates; on sand-mud bottoms. 

Osachila semilevis Bathbun, 1916 (Proc. U.S. Nat. Mus. 50: 652) 

Hay & Shore, 1918, p. 422, pi. 31, fig. 9; Rathbun, 1937, p. 251, pi. 77, fig. 1; 

Williams, 1965; p. 159, fig. 142. 

Bange: North Carolina; Dry Tortugas; northwest Florida. 

Depth: 2 to 91 m(lto50fm). 

Habitat: sand, shell, and rocky bottoms. 

Bemarks: Williams (1965) reports that this species has been collected north 

of Florida only twice, both tim.es in the Beaufort, North Carolina area. He also 

reports ovigerous females from Florida in July. Listed from Florida by Wass 

(1955) andAbele(1970),butnotbyMenzel (1971). 

Osachila tuberosa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 154) 

Hay & Shore, 1918, p. 423, pi. 31, fig. 10; Rathbun, 1937, p. 250, pi. 77, fig. 3;


Williams, 1965, p. 159, fig. 141; Guinot, 1966, p. 748, figs. 3, 15, 17, 27, 31, 34, 35; 


Range: North Carolina (rare); Florida Kej^s and Dry Tortugas; south and 

west coasts of Florida; eastern portions of the Gulf of Mexico. 

Depth: 66 to 183 m (36 to 100 fm), but more common at the shallower end 

of its bathymetric range. 

Remarks: Listed from the Gulf of Mexico by Chace (1956) for the R/V 

Oreeon collections. 

Family LEUCOSIIDAE Samouelle, 1819 

Subfamily EBALIINAE Stimpson, 1871 

EftoZia Leach, 1817 

Ebalia cariosa (Stimpson, 1860) (Ann. Lye. Nat. Hist. New York 7: 238) 

Rathbun, 1937, p. 125, pi. 35, figs. 6-7; Williams, 1965, p. 147, fig. 122. 


west and northwest coasts of Florida; Jamaica; northeastern South America to 

Sao Paulo, Brazil. 


Habitat: from coral, rock, sand, and broken shell substi-ates. 

Remarks: Florida listings include Wass (1955), Tabb & Manning (1961), 

Rouse (1970), Abele (1970), Lyons e^ «/. (1971), and Menzel (1971). Williams 

(1965) reports ovigerous females from North Carolina throughout the summer 

and he notes the death-feigning behavior of this crab when it is captured. The 

crab resembles pebbles and shell pieces with which it is associated. Listed from 


Ebalia stimpsoni A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 22) 

Rathbun, 1933, p. 100, fig. 96; Rathbun, 1937, p. 124, text-fig. 33, pi. 35, figs. 1-3, 

pi. 37, figs. 1-3; Williams, McCloskey & Gray, 1968, p. 46. 

Range: off North Carolina; Bahamas; Florida Keys and Dry Tortugas; west 

coast of Florida; Puerto Rico; Barbados; Maranhao to Bahia, Brazil. 

Depth: 7 to 183 m (4to 100fm). 

Habitat: sand, shell, and coral bottoms. 

Remarks: Listed from Florida by Hulings (1961) and Abele (1970). Williams, 

McCloskey and Gray (1968) reported an ovigerous female from off North Carolina 

in May. Listed from Brazil by Rodrigues da Costa (1968a) and by Coelho 


Lilhadia Bell, 1855 

Lithadia cadaverosa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 159) 


Range: Bahamas; northwest coast of Florida and northeast portion of Gulf.


Depth: 46 to 62 m (25 to 34 fm). 

Habitat: sand, broken shell, and gravel substrates. 

Speloeophorus A. Milne Edwards, 1865 

Speloeophorus elevalus Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 

4: 290) 


Range: Florida Keys; Jamaica; off Cape St. Roque and from Maranhao to 

Alagoas, Brazil. 

Depth: 1 to 83 m (1 to 45 fm). 

Habitat: broken shell substrate. 

Remarks: Listed from Brazil by Coelho (1971a) and Coelho and Ramos 

(1972). 

Speloeophorus nodosus (Bell, 1855) (Trans. Linn. Soc. London 21: 307) 

Hay & Shore, 1918, p. 425, pi. 32, fig. 5; Rathbun, 1933, p. 99, fig. 95; Rathbun, 

1937, p. 142, pi. 40, figs. 1-5; Williams, 1965, p. 148, figs. 123-124. 

Range: North Carolina; Florida Keys and Dry Tortugas; Jamaica; Puerto 

Rico; St. Thomas, Virgin Islands; Guadeloupe. 

Depth: 3 to 18 m (1.5 to 10 fm). 

Habitat: coral reefs and coral-sand bottoms. 

Remarks: Williams (1965) states that this species is rare in the northern 

part of its range and he cites a report hj Pearse and Williams (1951) of its 

occurrence on a reef off North Carolina. This crab readily feigns death when 

captured. 

Speloeophorus ponlifer (Stimpson, 1871) (Ann. Lye. Nat. Hist. New York 10: 

115) 

Hay & Shore, 1918, p. 425, pi. 32, fig. 5; Rathbun, 1933, p. 100; Rathbun, 1937, 

p. 144, pi. 39, figs. 1-3; Williams, 1965, p. 149, figs. 125-126. 

Range: off North Carolina; Florida Keys and Dry Tortugas; west coast of 

Florida; Cuba; Puerto Rico; Barbados. 


Habitat: coral, sand, and shell bottoms. 

llhlias Stimpson, 1871 

Vhlias Umbalus Stimpson, 1871 (Ann. Lye. Nat. Hist. New York 10: 118) 


Range: west of Key West, Florida; north coast of Cuba; Jamaica; Haiti; St. 

Thomas, Virgin Islands. 

Depth: 4 to 64 m (2 to 35 fm). 

Habitat: shell and grassy substrates. 

Subfamily LEUCOSIINAE Samouelle, 1819

Callidactylus Stimpson, 1871 


Callidactylus asper Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 158) 

Rathbun, 1937, p. 193, pi. 58, figs. 1-3; Williams, McCloskey & Gray, 1968, p. 48, 

fig. 5. 

Range: off North Carolina; southeast Florida; Florida Keys and Dry Tortugas; 

off southwest Florida; east coast of Haiti; Maranhao to Alagoas, Brazil. 


Habitat: sand substrates; oHLithothamnium reef. 

Remarks: Listed from Brazil by Coelho (1971a, 1971b) and Coelho and Ramos 

(1972). See remarks for Iliacantha liodactylus. 

Iliacantha Stimpson, 1871 

Iliacantha intermedia Miers, 1886 (Voyage of H.M.S. Challenger, Brachyura, 

vol. 17, p. 302. 


Williams, 1965, p. 151, fig. 129. 

Range: North Carolina; Florida Keys and Dry Tortugas; northwest coast of 

Florida; St. Thomas, Virgin Islands; off Venezuela; from Maranhao to Bahia, 

Brazil. 

Depth: 10to329m(5.5tol80fm). 

Habitat: sand, gravel, coral, and broken shell bottoms. 

Remarks: Williams (1965) comments on the close resemblance between 

juveniles of this species and those of Persephona mediterranea. Listed from 

Brazil by Coelho (1971a) and Coelho and Ramos (1972). 

Iliacantha liodactylus Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 

4: 291) 

Rathbun, 1937, p. 186, text-fig. 41, pi. 55, figs. 1-2; Felder, 1973a, p. 39, pi. 5, fig. 2. 

Range: west coast of Florida; ? Haiti; Puerto Rico; St. John, Virgin Islands; 

Trinidad; Alagoas to Bahia, Brazil. 

Depth: 9 to 130 m (5 to 71 fm). 

Habitat: mud substrates. 

Remarks: Recorded by Chace (1956) from the R/V Oregon cruise in the Gulf 

of Mexico. The locality records cited for this species and for Callidactylus asper 

by Rathbun (1937) are not in agreement with the latitudes and longitudes listed. 

Both species are listed from the east coast of Haiti, as recorded by the Johnson- 

Smithsonian Expedition of 1933, but the coordinates as listed would place C. 

asper east of Puerto Rico and place /. liodactylus on the east coast of the Dominican. 

Republic. Leary (1967) lists /. liodactylus from Texas and Felder (1973a) 

repeats Leary's listing, but I know of no published collection records to verify 

the presence of this species in the northwestern Gulf. Listed from Brazil by 

Coelho (1971b) and Coelho and Ramos (1972).


Iliacanlha sparsa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 156) 

Rathburi, 1937, p. 190, pi. 56, figs. 1-2. 

Range: northwest of the Dry Tortugas; off north and southeast coasts of Puerto 

Rico; Barbados; Maranhao to Bahia, Brazil. 

Depth: 23 to 73 m (13 to 40 fm). 

Habitat: gravel, shell, and coral bottoms; on calcareous algae. 

Remarks: Onl}^ male specimens were listed by Rathbun (1937), one of which 

was heavily infested with parasites. Listed from Brazil by Coelho (1971a) and 

Coelho and Ramos (1972). 

Iliacanlha subglobosa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 155) 


Williams, 1965, p. 150, fig. 128; Pequegnat, 1970, p. 179. 


northwest Florida; north coast of Cuba; Jamaica; Lesser Antilles, from Montserrat 

to Barbados; from Amapa to Alagoas, Brazil. 

Depth: 27 to 393m (15to215fm). 

Habitat: coral, sand, and rock bottoms. 

Remarks: Listed by Chace (1956) from the Gulf of Mexico. Williams (1965) 

reported ovigerous females from the Gulf in June. Brazilian records include 

Coelho (1971a, 1971b) and Coelho and Ramos (1972). 

Subfamily PHILYRINAE Rathbun, 1937 

Myropsis Stimpson, 1871 

Myropsis quinquespinosa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 157) 

Rathbun, 1937, p. 164, pi. 46, figs. 1-3; Chace, 1940, p. 24; Williams, McCloskey & 

Graj', 1968, p. 46, fig. 4; Pequegnat, 1970, p. 179; Felder, 1973a, p. 39, pi. 5, fig. 5. 

Range: Massachusetts; North Carolina; Bahamas; southeast Florida; Florida 

Keys and Dry Tortugas; west coast of Florida; all portions of Gulf of Mexico; 

off Alabama, Texas, and Campeche Bank iji Mexico; north and south coasts 

of Cuba; Jamaica; Puerto Rico; Lesser Antilles, from Martinique to Grenada; 

Barbados; Venezuela. 

Depth: 91 to 329 m (50 to 185 fm), rare report to 1045 m (572 fm). 

Habitat: commonly on mud bottoms, also on sand and shell substrates. 

Remarks: Rathbun (1937) reported ovigerous females from Florida in May 

and July and Williams, McCloskey and Gray (1968) reported an ovigerous 

female from off North Carolina in July, where they believe this species is restricted 

to a depth range of 120 to 160 m. Chace (1956) recorded this crab from 

50 fm off Sabine, Texas and it is listed from Texas by Leary (1967). Pequegnat 

(1970) notes that the depth record of 572 fm may be excessive.

Persephona Iêach, 1817 

Persephona aquilonaris Rathbun, 1933. 


This name was given to the western Atlantic species, including the Gulf of 

Mexico form, orginally to indicate a subspecies of P. punctata. A revision of the 

genus by Guinot-Dumortier (1959) indicated that the two forms were distinct 

species and she extended the range through the Caribbean to South America, 

where it co-exists with P. punctata. Rathbun (1937) had restricted P. punctata 

aquilonaris to North America. At the same time as her revision, Guinot- 

Dumortier (1959) indicated that the species illustrated by Herbst, P. mediterranean 

may be the correct name of P. aquilonaris. Rathbun (1937, p. 153) listed 

P. mediterranea as a synonym of P. punctata, but she indicated that the type 

locality (Mediterranean Sea) was probably incorrect. Abele (1970) reviews the 

synonymy and notes that L. B. Holthuis verified the identity and synonymy of 

P. mediterranea and P. aquilonaris. Abele (1970) also notes that the figure in 

Rathbun (1933, p. 99, fig. 94) for P. punctata from Puerto Rico is a figure of P. 

mediterranea. 

Persephona crinila Rathbun, 1931 (J. Washington Acad. Sci. 21: 128) 

Rathbun, 1937, p. 163, pi. 43, figs. 2-3, pi. 4-4', figs. 1-3; Felder, 1973a, p. 39, pi. 5, 

fig. 3. 

Range: northwest Florida to Texas; Trinidad; Ilha Sao Sebastiao, Brazil. 

Depth: 5.5 to 91 m (3 to 50 fm). 

Habitat: mud and mud-sand bottoms; more common in open marine waters 

than is P. mediterranea. 

Remarks: Hildebrand (1954) collected an ovigerous female from off Texas in 

June. Wass (1955) stated that this species is rarer than P. aquilonaris (^ P. 

mediterranea) off northwest Florida and Menzel (1971) listed this crab as 

uncommon on sand bottoms in the same area. Franks et al. (1972) extended the 

known depth range to 50 fm, from collections off Mississippi. Listed from the 

Gulf collections of the R/V Oregon by Chace (1956). 

Persephona mediterranea (Herbst, 1794) (Versuch Naturgesch. Krabben u. 

Krebse,voI.2,p. 150) 

Common Name: Purse Crab 

As P. punclala—Hay & Shore, 1918, p. 423, pi. 32, fig. 9 (part); Rathbun, 1933, 

p. 99 (part), fig. 94. 

As P. punctata aquilonaris—Rathbun, 1937, p. ISA; pi. 42, figs. 6-7; Williams, 

1965, p. 150, fig. 127. 

As P. aguaonaris—Guinot-Dumortier, 1959, p. 429, figs. 7, 9; Felder, 1973a, p. 42, 

pi. 5, fig. 4. 

As P. mediterranea—Abele, 1970, p. 62. 

Range: New Jersey to south Florida; Dry Tortugas; west coast of Florida to 

south Texas; off Campeche, Mexico; Guadeloupe, Lesser Antilles; French 

Guiana; Santa Catarina, Brazil. 

Depth: 4 to 55 m (2 to 30 fm).


Habitat: sand, shell, and coral bottoms; inshore waters, in passes; in very 

shallow water just below low tide mark; from mud-shell substrates in northwestern 

Gulf of Mexico. 

Remarks: For explanation of nomenclatural revisions, see P. aquilonaris. 

Williams (1965) reported abundant colonies of this crab, with ovigerous females 

present throughout the spring and summer in North Carolina waters. Regional 

listings include the Gulf (Chace, 1956; Fotheringham and Brunenmeister, 1975), 

Florida (Wass, 1955; Tabb and Manning, 1961; Dragovich and Kelly, 1964), 

Louisiana (Behre, 1950), and Texas (Hedgpeth, 1953; Hildebrand, 1954; Leary, 

1967). 

Persephona punclala aquilonaris Rathbun, 1933. 

Revised by Guinot-Dumortier (1959) to a separate species and later determined 

to be a junior synonym of P. mediterrnnea (Herbst). See P. aquilonaris 

for nomenclatural discussion. 

SECTION HAPALOCARCINIDEA Verrill, 1908 

Superfamily HAPALOCARCINOIDEA (Verrill, 1908) 

Family HAPALOCARCINIDAE Caiman, 1900 

(The taxonomic status of these unique crabs is uncertain. Verrill (1908, 

p. 426) proposed the designation "Hapalocarcinidea" for "a peculiar 

superfamily," apparently including them among the Oxystomata. Glaessner 

(1969) does not include this group in his classification schemes. 

Chace (personal comm.) has suggested the retention of "Hapalocarcinidea" 

for a section name and the appropriate ending for the superfamily 

name, both of which are credited to Verrill. The status and 

relationships of the hapalocarcinids will undoubtably change with further 

study.) 

Cryplochirus Heller, 1861 

Cryplochirus corallicola (Verrill, 1908) (Trans. Connecticus Acad. Arts Sci. 

13:427) 

Rathbun, 1937, p. 262 (part), text-fig. 47, pi. 78, figs. 5-7. 

Range: Florida Straits; Dry Tortugas; Bermuda; Dominica; Maranhao to 

Bahia, Brazil. 

Depth: 0 to 75 m (0 to 41 fm). 

Habitat: in ovoid cavities in the upper surfaces of corals, such as Meandra 

areolata and Meandrina sp. 

Remarks: Serene (1966) reviews the taxonomy and geographical distribution 

of hapalocarcinids. Rathbun (1937) commented on the relationship between the 

crabs and their living coral habitat: "the opening of the den is usually semicircular 

or lunate, commonly oblique to surface of coral. The downturned, rough,


and dirt-covered front of the crab serves as an operculum, closing the aperture. 

Full grown crabs are probably unable to leave their dens." The western African 

record listed by Rathbun (1937) was referred to Troglocarcinus balssihy Monod 

(1956). Listed from Brazil (as Troglocarcinus corallicola) by Coelho (1971a) 

and Coelho and Ramos (1972). Fausto Filho (1974) provides some habitat notes 

and extended the known depth to 75 m, based on Brazilian specimens collected 

by Coelho. 

SECTION OXYRHYNCHA Latreille, 1803 

Family MAJIDAE Samouelle, 1819 

Subfamily ACANTHONYCHINAE Stimpson, 1870 

Acanlhonyx Latreille, 1825 

Acanthonyx petiverii H. Milne Edwards, 1834 (Hist. Nat. Crust., vol. 1, p. 343) 

Rathbun, 1925, p. 142, text-fig. 52, pi, 44, pi. 222, figs. 1-6; Rathbun, 1933, p. 13, 

fig. 11; Garth, 1938, p. 225, pi. O, fig. 3, pi. 25, fig. 2; Felder, 1973a, p. 53, fig. 14. 

Range: Bahamas; southeast and northwest Florida; Texas; Cuba; .Jamaica; 

Puerto Rico; Virgin Islands; Netherlands Antilles; Panama (Carib.) to Rio de 

Janeiro, Brazil; along the Pacific coast, from Baja California to Caldera, Chile; 

Galapagos Islands. 

Depth: shore to 29 m (16 fm). 

Habitat: in tide pools of rockj'^, surf-beaten shores; algal-covered surfaces and 

on seaweeds; sandy shores; coral flats. 

Remarks: Leary (1967) lists this crab from Texas and Felder (1973a) found 

this crab on rock jetties in southern Texas. Chace (1966) figured four types of 

carapace variation in this species. Abele (1970) also collected this crab, a single 

female, from seaweed on jetties in northwest Florida. Listed from Brazil by 

Coelho (1971a, 1971c) and Coelho and Ramos (1972). 

Epialtus H. Milne Edwards, 1834 

Epiallus bituberculalus H. Milne Edwards, 1834 (Hist Nat. Crust., vol. 1, p. 

345) 

Rathbun, 1925, p. 148, text-figs. 53a, 54, pi. 45, figs. 3-4; Rathbun, 1933, p. 14, 

fig. 15; Garth, 1958, p. 228. 

Range: east coast of Florida; Key West, Florida; Puerto Rico; Panama (Carib.) 

to Colombia; Ceara to Pernambuco, Brazil. 

Depth: shallow waters. 

Habitat: like others of this genus, on hard surfaces and in tide pools, feeding 

among seaweed and other plant growth. 

Remarks: Although this species has been listed from Chile and from southern 

California, Garth (1958) recognizes an exclusively Atlantic range. Listed from 

Brazil by Coelho (1971c) and Coelho and Ramos (1972).


Epiahus dilalalus A. Milne Edwards, 1878 (Crust. Reg. Mex., p. 140) 

Rathbun, 1925, p. 153, text-fig. 53j, pi. 45, fig. 2; Rathbun, 1933, p. 15, fig. 14; 

Williams, 1965, p. 249, figs. 228, 233D. 

Range: North Carolina; Bahamas; south Florida; Dry Tortugas; Isia Mujeres, 

Yucatan (Carib.); Puerto Rico; St. Thomas, Virgin Islands. 

Depth: 5 to 22 m (2.5 to 12 fm). 

Habitat: sand, coral, and broken shell substrates. 

Remarks: Yang (1968) describes the development of this crab to the first 

adult stage under laboratory conditions. The taxonomic status of the elongated 

form is still unclear and is described separate^ below. 

Epiahus dilalalus forma elongala Rathbun, 1923 (Proc. Biol. Soc. Washington 

36: 72) 

Rathbun, 1925, p. 154, fig. 53k, pi. 48. 

Range: Florida Keys; south, west, and northwest coasts of Florida. 

Depth: 2tol4m(l to7.5fm). 

Habitat: on sandy bottoms with grass; in patches of Sargassum. 

Remarks: Abele (1970) notes the considerable variation in this species and 

genus and he expresses reservations about the distinct status of this form. Florida 

listings include Wass (1955), Abele (1970), Menzel (1971), and Lyons et al, 

(1971). Wass (1955) collected this crab from Sargassum, off Ochlockonee Bay, 

Florida. Rathbun (1925) reported ovigerous females from Florida in January 

and February; other records include ovigerous females from Florida in March 

and July. 

Epiahus longiroslris Stimpson, 1860 (Ann. Lye. Nat. Llist. New York 7: 199) 

Rathbun, 1925, p. 151, text-figs. 53g, 56. 

Range: Key West and west coast of Florida; Cuba; Jamaica; St. Thomas, 

Virgin Islands; northeast Brazil. 

Depth: shallow water, 3 to 5 m (2 to 3 fm) off Cuba; 19 to 54 m (10 to 30 fm) 

off Brazil. 

Remarks: Listed from Brazil by Coelho (1971c). 

Mocosoa Stimpson, 1871 

Mocosoa crebripunclala Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 128) 

Rathbun, 1925, p. 159, text-fig. 59, pi. 49, figs. 3-4. 

Range: Florida Straits; off Cape San Bias, northwest Florida; Maranhao to 

Espirito Santo, Brazil. 

Depth: 27 to 131m (15 to 72fm). 

Remarks: Listed from Brazil by Coelho (1971a) and by Coelho and Ramos 

(1972). 

Subfamily INACHINAE Macleay, 1838

Aepinus Rathbun, 1897 

Crabsof the Gulf of Mexico 43 

Aepinus seplemspinosus (A. Milne Edwards, 1879) (Crust. Reg. Mex., p. 185) 

Rathbun, 1925, p. 92, text-figs. 28^29, pi. 32, figs. 3-4, pi. 219, figs. 1-3; Rathbun, 

1933,p. 10,fig. 7. 

Range: Bahamas; Florida Straits; Dry Tortugas; west and northwest coasts 

of Florida; Puerto Rico; St. Thomas, Virgin Islands; Para to Bahia, Brazil. 

Depth: 13 to 85 m (7 to 47 fm). 

Habitat: on hard surfaces, mainly coral; from calcareous algae and rock. 

Remarks: Listed from off northwest Florida by Wass (1955). Coelho (1971c) 

extended the known depth range and commented on the ecology of this crab in 

Brazil. Also listed from Brazil by Coelho and Ramos (1972) and Fausto Filho 

(1974). 

Anasinius A. Milne Edwards, 1880 

Anasimus lalus Rathbun, 1894 (Proc. U.S. Nat. Mus. 17: 58) 

Rathbun, 1925, p. 65, pi. 214; Guinot-Dumortier, 1960, p. 177, fig. 18a-b; Williams, 

1965, p. 240, figs. 217, 223F; Felder, 1973a, p. 49, pi. 7, fig. 5. 

Range: North Carolina to south Florida; Florida Keys; northwest Florida to 

off coast of Tabasco, Mexico; west of Trinidad; Guianas; Amapa, Brazil. 

Depth: 48 to 161m (26 to 88 fm). 

Habitat: mud, sand, broken shell, and coral bottoms. 

Remarks: Rathbun (1925) compares differences in the morphology of adults 

and juveniles. Reports of ovigerous females were summarized by Williams 

(1965); females apparently carry eggs throughout the year in the Gulf of Mexico. 

Holthuis (1959) describes this species from off Surinam, where it is ovigerous 

from April to August. Sandifer and van Engel (1972) studied larval development 

under laboratory conditions. Regional lists include Florida (Wass, 1955), 

Mississippi (Franks et al, 1972), Texas (Hildebrand, 1954; Leary, 1967), 

Mexico (Hildebrand, 1954), and the Gulf of Mexico collections of the R/V 

Oregon (Chace, 1956). Listed from Brazil by Coelho (1971c) and Coelho and 

Ramos (1972). 

^nomaZol/iir Miers, 1879 

Anomalolhir frontalis (A. Milne Edwards, 1879) (Cioist. Reg. Mox., p. 189) 

Rathbun, 1925, p. 25, pi. 8, fig. 1, pi. 9, fig. 1, pi. 207; Chace, 1940, p. 56. 

Range: north coast of Cuba; Montserrat to Barbados. 

Depth: 133 to 421 m (73 to 230 fm). 

Habitat: sand, coral, and broken shell bottoms. 

Remarks: Rathbun (1925) compares this species with A. furcillatus. 

Anomalolhir furcillatus (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 125) 

Rathbun, 1925, p. 24, text-fig. 6, pi. 8, fig. 2, pi. 9, fig. 2, pi. 206; Rathbun, 1933, 

p. 6, fig. 2; Chace, 1940, p. 55; Williams, 1965, p. 236, figs. 212,223A. 

Range: North Carolina; Florida Straits and Keys; Dry Tortugas; off northwest


Florida; north coast of Cuba; north coast of Yucatan; Jamaica; St. Croix, Virgin 

Island to Grenada. 

Depth: 55 to 686 m (30 to 375 fm), more commonly at depths of greater than 

183 m (100 fm). 

Habitat: mud, sand, shell, stone, and coral bottoms. 

Remarks: Rathbun (1925) listed ovigerous females from the northeast qviadrant 

of the Gulf in February and March. 

Arachnopsis Stimpson, 1871 

Arachnopsis filipes Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 121) , 

Rathbun, 1925, p. 89, text figs. 26-27, pi. 32, figs. 1-2, pi. 219, figs. 4-5; "Williams, 

McClosfcey & Gray, 1968, p. 58. 

Range: off North Carolina; Florida Straits; Dry Tortugas; west coast of 

Florida; off Dominica; Barbados; Ceara to Rio Grande do Norte, Brazil. 

Depth: 27 to 238 m (15 to 130 fm). 

Habit: sand, shell, and coral bottoms. 

Remarks: Listed by Wass (1955) from northwest Florida and b)^ Chace (1956) 

from R/V Oregon collections in the Gulf of Mexico. Recorded from Brazil by 

Coelho (1971a, 1971b) and Coelho and Ramos (1972). 

Balrachonolus Stimpson, 1871 

Balrachonotus fragosus Stimpson, 1871 (Bvdl. Mus. Comp. Zool. 2; 122) 

Ratlibun, 1925, p. 123, text-fig. 48, pi. 39, figs. 1-4; Rathbun, 1933, p. 13, fig. 10; 

Williams, 1965, p. 238, figs. 214, 223C. 

Range: North Carolina; south, west and northwest coasts of Florida; Dry 

Tortugas; north coast of Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin Islands; 

Curasao, Netherlands Antilles; Rio de Janeiro, Brazil. 

Depth: shore to 137 m (to75fm). 

Habitat: sand, shell and coral bottoms; rarely on mud. 

Remarks: Listed from northwest Florida by Wass (1955), Hulings (1961), 

and Abele (1970). Williams (1965) reported ovigerous females from Dry Tortugas 

in June. 

Collodes Stimpson, 1860 

Collodes armalus Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 4: 252) 

Rathbun, 1925, p. 122, pi. 217, fig. 6. 

Range: off Havana, Cuba. 

Remarks; known only from the type specimen, collected in 1893. 

Collodes leplocheles Rathbun, 1894 (Proc. U.S. Nat. Mus. 17: 53) 

Rathbun, 1925, p. llr, text-fig. 42, pi. 38, figs. 5-6; Pequegnat, 1970, p. 181; Felder, 

1973a,p. 49,pl. 7,fig.3. 

Range: all quadrants of Gulf of Mexico except southeast, off coasts of Florida, 

Alabama, and Texas; off Vera Cruz, Mexico.


Depth: 124to384m (68to210fm). 

Habitat: gray mud, broken shell substrates. 

Remarks: Chace (1956) reported the first record of this species from off Texas, 

taken at 150 fm, and Leary (1967) also lists this crab from Texas. Pequegnat 

(1970) reported two ovigerous females, taken in August from 111 fm. 

Collodes trispinosus Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 120) 

Rathbun, 1925, p. 107, text-fig. 32, pi. 36, figs. 5-6; Williams, 1965, p. 239, figs. 

215, 223D. 

Range: North Carolina to south Florida; Florida Straits and Keys; Dry Tortugas; 

west and northwest coasts of Florida. 

Depth: 7tol50m(4to82fm). 

Habitat: sand, broken shell, and gravel substrates. 

Remarks: Listed from northwest Florida by Wass (1955). Williams (1965) 

lists ovigerous females from North Carolina in October and from Florida in July. 

Euprognatha Stimpson, 1871 

Euprognatha gracilipes A. Milne Edwards, 1878 (Crust. Reg. Mex., p. 184) 

Rathbun, 1925, p. 101, pi. 34, figs. 3-4; Rathbun, 1933, p. 11. 

Range: Florida Keys; north coast of Yucatan; off north coast of Cuba; Puerto 

Rico; St. Croix, Virgin Islands; Barbados; Amapa to Sao Paulo, Brazil. 

Depth: 72 to 368 m (39 to 201 fm). 

Habitat: sand, shell, coral, and rocky bottoms. 

Remarks: Rathbun (1925) comments on variation within this species and the 

possibility of two different subspecies within the described population. Listed 

from Brazil by Coelho (1971c) and Coelho and Ramos (1972). 

Euprognatha rastellifera acuta A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 

' 348) 

Rathbun, 1925, p. 96, pi. 34, figs. 1-2; Rathbun, 1933, p. 11, fig. 8; Chace, 1940, p. 57. 

Range: Massachusetts; North and South Carolina; Florida Keys; north coast 

of Cuba; Puerto Rico; St. Kitts; Martinique. 

Depth: 102 to 708m (56 to 387 fm). 

Habitat: sand, coral, and shell substrates. 

Remarks: There is considerable geographical overlap among the three forms 

of this species, the typical form, E. r. acuta, and E. r. marthae. The subspecies 

acuta is distributed over the entire species range, although it is more common in 

the southern portion, whereas marthae is listed by Rathbun (1925) as restricted 

to north of southern Florida, along the Atlantic coast. Williams (1965) notes 

the need for clarification of these forms and their distributions. 

Inachoides H. Milne Edwards & Lucas, 1843 

Inachoides forceps A. Milne Edwards, 1879 (Crust. Reg. Mex., p. 199) 

As/, laevis—Rathbun, 1925, p. 61 (part), text-fig. 17, not pi. 22, figs. 3-6; Rathbun, 

1933,p. 8,fig. 4. 

As l. forceps—Garth, 1938, p. 101; Williams, McCloskey & Gi-ay, 1968, p. 60, 

fig. 14.


Range: North Carolina; west and northwest coasts of Florida; Jamaica; Puerto 

Rico; Guadeloupe; St. Thomas, Virgin Islands; Guianas to Santa Catarina, Brazil, 


Habitat: sand, gravel, and coral substrates. 

Remarks: Garth (1958) separated the Pacific and Atlantic populations, previously 

combined as /. laevis, into two distinct species, retaining laevis for the 

original Pacific form designated by Stimpson. Williams, McCloske)'- and Gray 

(1968) cite pi. 22 of Rathbun (1925) as a synonymy, but this is of a specimen 

from the Pacific coast of Panama and it should be referred to /. laevis. Listed 

from northwest Florida by Wass (1955) and Menzel (1971) and from Brazil 

byCoelho (1971c) and Coelho and Ramos (1972). 

Inachoides laevis Stimpson, 1860. 

Restricted to Pacific records by Garth (1958); all Atlantic records are referred 

to /. forceps A. Milne Edwards. 

Metoporhaphis Stimpson, 1860 

Metoporhaphis calcarata (Saj'-, 1818) (J. Acad. Nat. Sci. Philadelphia 1: 455) 

Hay & Shore, 1918, p. 454, pi. 37, fig. 5; Rathbun, 1925, p. 21, text-fig. 5, pis. 6-7; 

Williams, 1965, p. 243, figs. 221, 223J; Felder, 1973a, p. 48, pi. 7, fig. 2. 

Range: North Carolina to south Florida; Florida Keys; west coast of Florida 

to south Texas; Rio de Janeiro, Brazil. 


Habitat: primarily hard surfaces, common on sand; rock, coral and grassy 

areas; among hydroids at North Carolina (Williams, 1965) and oyster beds at 

Grand Isle, Louisiana (Behre, 1950). 

Remarks: Regional lists include Florida (Wass, 1955; Tabb and Manning, 

1961; Dragovich and Kelly, 1964; Abele, 1970; Lyons et al., 1971; Menzel, 

1971), Mississippi (Richmond, 1968), Louisiana (Behre, 1950), and Texas 

(Breuer, 1962). Ovigerous females are known from South Carolina in August 

(Williams, 1965) and from Florida in March and August (Wass, 1955). Abele 

(1970) noted sexual dimorphism in the shape of the chelae. Wass (1955) commented 

on the ability of this crab to remain suspended in water by the rhythmic 

waving of the elongate legs, which are covered with fine setae on the distal 

portion. Listed from Brazil by Coelho and Ramos (1972). 

Podochela Stimpson, 1860 

Podochela curvirostrts (A. Milne Edwards, 1879) (Crust. Reg. Mex., p. 196) 

Rathbun, 1925, p. 50, pis. 19, 210; Chace, 1940, p. 56. 

Range: Florida Straits; north coast of Cuba; Caribbean coast of Yucatan; 

Montserrat; Barbados; Grenadines. 


Habitat: coral, sand, and broken shell bottoms.


Remarks: Rathbun (1925) reported an ovigerous female from off Cuba in 

May. 

Podochela gracilipes Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 126) 

Hay & Shore, 1918, p. 454, pi. 37, fig. 6; Rathbun, 1925, p. 47, text-fig. 12, pi. 17; 

Williams, 1963, p. 243, figs. 220,2231. 

Range: North and South Carolina; Florida Keys and Straits; Drj^ Tortugas; 

west coast of Florida to off Alabama; north coast of Yucatan; Barbados; Colombia 

(Carib.); Guianas to Santa Catarina, Brazil. 

Depth: 6 to 220 m (3 to 120 fm). 

Habitat: sand, gravel, broken shell, rocky, and coral bottoms. 

Remarks: Ovigerous females are known from North Carolina in December 

(Williams, 1965) and from Florida in March (Rathbun, 1925. Listed from shallow 

waters in Brazil by Coelho and Ramos (1972). 

Podochela lamelligera (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 126) 


Range: southeast Florida; off Key West, Florida; off northwest Florida. 

Depth: 38 to 110 m (21 to 60 fm). 

Habitat: sand, coral, and rocky substrates. 

Remarks: Rathbun (1925) reports an ovigerous female from off Cape Florida 

in March. Listed by Chace (1956) from the Gulf of Mexico collections of the 

R/V Oregon. 

Podochela macrodera Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 196) 

Rathbun, 1925, p. 44, text-fig. 11, pi. 16; Rathbun, 1933, p. 8. 

Range: Bahamas; Florida Keys; west coast of Florida; off Caribbean coast of 

Yucatan; Cuba; Puerto Rico; Virgin Islands; Guadalupe; Curasao, Netherlands 

Antilles; Brazil. 

Depth: shallow water to 53 m (to 29 fm), rare to 91 m (50 fm). 

Habitat: coral, sand, and gravel bottoms; from sponges; among rocks. 

Remarks: Rathbun (1925) listed ovigerous females from off Florida in February 

and from Cuba in June. Coelho (1971c) listed this crab from depths of 

20 to 53 m off Brazil. 

Podochela riisei Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 196) 


figs. 1-2, pi. 208, fig. 2; Rathbun, 1933, p. 7, fig. 3; Chace, 1940, p. 56; Williams, 

1965, p. 241, figs. 218, 223G. 

Range: North Carolina; Bermuda; Bahamas; Florida Keys and Dry Tortugas; 

west and northwest coasts of Florida; Campeche, off Yucatan; Cuba; Jamaica; 

Puerto Rico; St. Thomas, Virgin Islands; Rio de Janeiro and south of Pernambuco, 

Brazil. 


Habitat: coral, sand, shell, rock, and gravel substrates; on alga beds, grasses 

on sand bottoms; among Sargassum (Wass, 1955).


Remarks; Records of ovigerous females include the west coast of Florida in 

March, April, June and September (Lyons et al., 1971) and from November 

through February (Rathbun, 1925). These crabs are often covered with a number 

of other organisms, including actinians, sponges, and rhizocephalan barnacles 

(Rathbun, 1925), and bryozoans, ascidians, and a red algae, Calathamnion 

byssoideum, as reported by Wass (1955). Regional lists include Florida (Wass, 

1955; Tabb and Manning, 1961; Dragovich and Kelly, 1964; Abele, 1970; Lyons 

et al, 1971; Menzel, 1971), Campeche (Hildebrand, 1955), and the Gulf of 

Mexico (Chace, 1956). Coelho (1971c) listed it from depths of 24 to 90 m, off 

Brazil. 

Podochela sidneyi Rathbun, 1924 (Proc. U.S. Nat. Mus. 64: 1) 

Rathbun, 1925, p. 39, pis. 12-13; Williams, 1965, p. 242, figs. 219, 223H; Felder, 

I973a,p.49„pl. 7,fig. 4. 

Range: North Carolina; Dry Tortugas; west coast of Florida to central Texas 

coast; off north coast of Yucatan; northwest coast of Cuba. 

Depth: shallow water to 187 m (tol02fm). 

Habitat: mud, broken shell, sand, coral, and rock bottoms; on alga-covered 

surfaces. 

Remarks: Hildebrand (1954) collected specimens off Louisiana and Texas with 

small Styela (ascidians) attached to the legs, and some were covered with a 

dense hydroid growth. Listed from Florida by Wass (1955), Abele (1970), and 

Menzel (1971) and from Texas by Hildebrand (1954) and Leary (1967). 

Pyrojmaifl Stimpson, 1871 

Pyromaia arachna Rathbun, 1924 (Proc. U.S. Nat. Mus. 64: 1) 

Rathbun, 1925, p. 131, pis. 42-43; Pequegnat, 1970, p. 182. 

Range: off Soutli Carolina; off west coast of Florida to off east coast of Mexico; 

throughout all quadrants of the Gulf of Mexico. 

Depth: 183to384m (100to210fm). 

Habitat: off mud, mud-sand, and mud-shell bottoms. 

Remarks: Rathbun (1925) reports ovigerous females off Florida in March. 

Listed from the Gulf collections of the R/V Oregon by Chace (1956) and from 

Texas by Leary (1967). 

Pyromaia cuspidala Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 110) 

Hay & Shore, 1918, p. 455, pi. 38, fig. 4; Rathbun, 1925, p. 129, text-fig. 49, pi. 4); 

Chace, 1940, p. 57; Williams, 1965, p. 240, figs. 216, 223E,- Pequegnat, 1970, p. 181. 

Range: North Carolina; south Florida; Florida Keys and Straits; Dry Tortugas; 

west coast of Florida; off Caribbean coast of Yucatan; north and south coasts 

of Cuba. 

Depth: 27 to 549 m (15 to 300 fm). 

Habitat: mud, sand, rock, coral, and pebble substrates. 

Remarks: Chace (1940) noted that this crab autotomizes its legs readily. Williams 

(1965) reports ovigerous females from off Florida in February and July.

Slenor/iync/iMs Lamarck, 1818 


Slenorhynchus selicornis (Herbst, 1788) (Versuch Naturgesch. Krabben u. 

Krebse,vol. l,p. 229) 

Common Names: Arrow Crab; Arana del Mar 

Hay & Shore, 1918, p. 435, pi. 37, fig. 8; Rathbun, 192S, p. 13, te.xt-fig. 3, pis. 2-3; 

Rathbun, 1933, p. 6, fig. 1; Chace, 1940, p. 5S; Williams, 1965, p. 244, figs. 222, 

223K; Felder, 1973a, p. 48, pi. 7, fig. 1; Pequegnat & Ray, 1974, p .237, figs. 11-12; 

Yang, 1976, p. 157. 

Range: North Carolina to south Florida; Bermuda; Bahamas; Florida Keys 

and Straits; Dry Tortugas; west coast of Florida to south Texas; north and east 

coasts of Yucatan; north and south coasts of Cuba; Jamaica; Puerto Rico; St. 

Thomas, Virgin Islands to Dominica; Netherlands Antilles; Colombia; Maranhao 

to Santa Catarina, Brazil. 

Depth: near surface to 1489 m (814 fm). 

Habitat: rock, coral, sand, sand-shell, and pebble bottoms; from sponges; off 

wharf pilings and rock jetties. 

Remarks: The eastern Atlantic records cited by Rathbun (1925) are referred 

to S. lanceolatus (Brulle) by Yang (1967) and Barr (1975). Yang (1976) provides 

evidence that 5. selicornis contains two species, one in shallow water, the 

other a deep water form. Williams (1965) states that this crab is more commonly 

collected at depths of 100 fm or less. Various authors use a variation of 

the genus name (e.g., Stenorynchus, in Williams, 1965). Yang (1967) describes 

larval stages. Ovigerous females occur off Texas in May and June (Hildebrand, 

1954) and throughout the spring and summer over most of its range. Regional 

lists include Florida (Wass, 1955; Menzel, 1971), Mississippi (Franks et ah, 

1972), Texas (Hildebrand, 1954; Leary, 1967), and the mid-ocean Gulf (Chace, 

1956). Hartnoll (1965a) provided some notes on the biology and growth of 

Jamaican populations and Barr (1971, 1975) studied its biology in the Virgin 

Islands. Its occurrence in Brazil was noted by Coelho (1971c) and Coelho and 

Ramos (1972). Agonistic behavior of this crab is discussed by Schone (1968) and 

antennule chemosensitivity was tested by Hazlett (1971). Herrnkind, Stanton 

and Conklin (1976) described commensal relationships with an anemone in 

Florida. 

Subfamily MAJINAE Samouelle, 1819 

Temnonolus A. Milne Edwards, 1875 

Temnonotus granulosus A. Milne Edwards, 1875 (Crust. Reg. Mex., p. 83) 

As T. simplex—"Rathhun, 1925, p. 342, pi, 249, figs. 10-12, 

As T. granulosus—Rathhun, 1925, p. 341, pi. 249, figs. 7-9; Chace, 1940, p. 65, 

fig. 22. 

Range: north coast of Cuba; Barbados. 

Depth: 183 to 478 m (100 to 260 fm). 

Habitat: coral, broken shell bottoms.


Remarks: Rathbun (1925) commented on the likelihood that the female type 

of T. granulosus was conspecific with the males of T. simplex^ the only known 

specimens of these two species. Chace (1940) synonymized the two names on 

the basis of a male T. gi-anulosus taken off Cuba. He also noted that distinctions 

in individual specimens are attributable to age variation as well as sexual dimorphism. 

Temnonolus simplex A. Milne Edwards, 1875 

Synonj^m of T. granulosus A. Milne Edwards, as revised by Chace (1940). 

Subfamily MITHRACINAEBalss, 1929 (sensu Garth, 1958). 

(This subfamily consists of Mithracinae Balss, 1929 plus Macrocoelominae 

Balss, 1929, derived from the Majinae Periceroida of Alcock, 1895. 

Glaessner treats the genera of this group under the subfamily Majinae 

Samouelle, 1819.) 

Coelocerus A. Milne Edwards, 1875 

Coelocerus spinosus A. Milne Edwards, 1875 (Crust Reg. Mex., p. 85) 

Rathbun, 1923, p. +46, text-fig. 130, pi. 263, pi. 264, figs. 1-2; Felder, 1973a, p. 49, 

pl. 7, fig. 6. 

Range: west coast of Florida to off Alabama. 

Depth: 24 to 64 m (13 to 35 fm). 

Habitat: rock, coral, and sand bottoms. 

Remarks: Garth (1958) states that this genus may be a link between Libinia 

and Neodoclea of the Pisinae and Stenocionops and Macrocoeloma of the Mithracinae. 

Hemus A. Milne Edwards, 1875 

Hemus crislulipes A. Milne Edwards, 1875 (Crust. Reg. Mex., p. 88) 

Rathbun, 1925, p. 345, text-fig. 110, pl. 124, fig. 1, pl. 248, figs. 9-15; Rathbun, 

1933, p. 21, fig. 23. 

Range: northwest coast of Florida; north coast of Yucatan; Puerto Rico; 

Curacao, Netherlands Antilles, Maranhao to Pernambuco, Brazil. 

Depth 15 to 69 m (8 to 38 fm). 

Habitat: sand, rock, and coral bottoms; in horn sponges and in the coral 

Poriies porites. 

Remarks: Listed by Wass (1955) and Menzel (1971) from northwest Florida. 

Brazilian listings include Coelho (1971c), Coelho and Ramos (1972), and Fausto 

Filho (1974). 

MacrocoeZoma Miers, 1879 

Macrocoeloma catnplocerum (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 112) 

Hay & Shore, 1918, p. 437, pl. 38, fig. 12; Rathbun, 1923, p .469, pl. 174, fig. 4, pl. 

270, fig. 2; Williams, 1965, p. 264, figs. 244, 245K.


Range: North Carolina; southeast Florida; Florida Keys; south Florida to 

northwest Florida. 

Depth: 4 to 35 m (2 to 19 fm). 

Habitat: rock, sand, coral, and broken shell substrates; often from grassy areas. 

Remarks: Rath bun (1925) reported specimens covered with various sponges, 

hydroids, bryozoans, ascidians, and infestations of rhizocephalid barnacles. Ovigerous 

females were listed from North Carolina in August and from Florida in 

January to March. Wass (1955) and Menzel (1971) list this species from northwest 

Florida and Lyons et al. (1971) state that it is ovigerous in April, June, 

and October at Crystal River, Florida. 

Macrocoelonia diplacanlhutn (Stimpson, 1860) (Ann. Lye. Nat. Hist. New 

York 7: 183) 

Rathbun, 1925, p. 478, pi. 169, fig. I, pi. 269, figs. 1-3; Rathbun, 1933, p. 36. 

Range: Bahamas; Key West, Florida; Cuba; Jamaica; Puerto Rico; Virgin 

Islands; Guadeloupe; Curagao, Netherlands Antilles; Old Providence Island 

(Carib.). 

Depth: 5 to 24 m (3 to 13 fm). 

Habitat: off shallow reefs; sandy substrates. 

Remarks: Rathbun (1925) listed specimens infected with rhizocephalid barnacles 

and others that were encrusted with algae. 

Macrocoelonia eulheca (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 112) 

Rathbun, 1925, p. 484, text-fig. 137, pi. 170, fig. I, pi. 171, fig. I; Rathbun, 1933, 

p. 37. 

Range: Bahamas; southeast Florida; Florida Keys; north coast of Cuba; St. 

Croix, Virgin Islands; Barbados; Caribbean coast of Panama; Maranhao to Bahia, 

Brazil. 

Depth: 30 to 214m (16toll7fm). 

Habitat: rock, broken shell, and coral substrates. 

Remarks: Chace (1956) listed this crab from the R/V Oregon collections in the 

Gulf of Mexico. Listings from Brazil include Coelho (1971a, 1971c) and Coelho 


Macrocoelonia intermedium Rathbun, 1901 (Bull. U.S. Fish. Comm. 20: 75) 

Rathbun, 1925, p. 486, text-fig. 138, pi. 170, fig. 2, pi. 171, fig. 2. 

Range: north coast of Cuba; Dominica; Caribbean coast of Panama. 

Depth: 62 to 298 m (34 to 163 fm). 

Habitat: coral and broken shell bottoms. 

Macrocoeloma laevigatum (Stimpson, 1860) (Ann. Lye. Nat. Hist. New York 

7:181) 

Rathbun, 1925, p. 483, text-fig. 136, pi. 169, figs. 2-3; Rathbun, 1933, p. 36. 

Range: Florida Keys; north coast of Cuba; Jamaica; St. Thomas, Virgin 

Islands; Guadeloupe; Piaui to Alagoas, Brazil. 

Depth: shore to 31 m (to 17 fm).


Habitat: rock and sand bottoms, often weedy. 

Remarks: Listed from Brazil by Coelho (IQZla, 1971c) and Coelho and Ramos 

(1972). 

Macrocoeloma septemspinosum (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 

113) 

Rathbun, 1925, p. 477, pi. 173, figs. 2^3. 

Range: South Carolina; Bahamas; Florida Keys; northeast quadrant of Gulf; 

Ceara to Rio Grande do Norte, Brazil. 

Depth: shallow water to 145 m (to 79 fm), rarely to 212 m (116 fm). 

Habitat: sand, broken shell, and coral substrates; on calcareous algae. 

Remarks: Listed by Chace (1956) from the Gulf of Mexico. Brazilian records 

include Coelho (1971a, 1971c) and Coelho and Ramos (1972). 

Macrocoeloma suhparallelum. (Stimpson, 1860) (Ann. Lye. Nat. Hist. New 

York 7: 182) 

Rathbun, 1925, p. 480, pi. 172; Rathbun, 1933, p. 36. 

Range: north coast of Cuba; Jamaica; Haiti; Puerto Rico; St. Thomas, Virgin 

Islands; Guadeloupe; Barbados; Old Providence Island (Carib.); Rio Grande do 

Norte to Pernambuco, Brazil. 


Habitat; on coral reefs; in tide pools; on bottoms of sand, grasses and weeds. 

Remarks: Brazilian records include notes on its occurrence in the littoral zone 

(Coelho, 1971c) and listings by Coelho and Ramos (1972) and Fausto Filho 

(1974). 

Macrocoeloma Irispinosum trispinosum. (Latreille, 1825) (Encyc. meth.. Hist. 

Nat, vol 10, p. 142) 

Common Names: Grass Crab; Sponge Crab; Decorator Crab 

Hay & Shore, 1918, p. 457, pi. 38, fig. 11; Rathbun, 1925, p. 466, te.vt-fig. 132, 

pi. 166, fig. 1, pi. 167; Rathbun, 1933, p. 35, fig. 31; Williams, 1965, p. 263, fig. 243; 

Felder, 1973a, p. 53, pi. 7, fig. 9. 

Range: North Carolina; Bermuda; south Florida to northwest Florida; off 

Louisiana and Texas; Gulf and Caribbean coasts of Yucatan, Mexico; Cuba; 

Jamaica; Puerto Rico; St. Thomas to St. Lucia; Curasao, Netherlands Antilles; 

Piaui to Bahia, Brazil. 


Habitat: sand, rock and shell bottoms; among submerged mangrove roots; 

from wharf pilings; from floating masses of Sargassum. 

Remarks: Rathbun (1925) listed three varieties of this species, two of which 

are listed here as subspecies (the typical form and M. t. nodipes), and a third, 

considered an intermediate form. Behre (1950) listed an unspecified form of 

Macrocoeloma from Louisiana and Menzel (1971) and Lyons et al. (1971) 

record this crab as uncommon in Florida. Ovigerous female records were summarized 

by Williams (1965). Hartnoll (1965a) described the biology and growth 

of this crab in Jamaica. Many specimens are encrusted with sponges, which are


attached to the hairs of the carapace and legs, providing the basis for two of its 

common names. Records from Brazil include Coelho (1971a, 1971c), Coelho and 

Ramos (1972) and Fausto Filho (1974). 

Macrocoeloina trispinosum nodipes (Desbonne, 1867) (Crust, de la Guadeloupe, 

p. 15) 

Rathbun, 1925, p. 468, pi. 166, fig. 2, pi. 168, fig. 2; Rathbun, 1933, p. 36; Williams, 

1965, p. 264. 

Range: North Carolina; Bermuda; south to northwest coasts of Florida; 

Florida Keys and Dry Tortugas; Cuba; Puerto Rico; Antigua; Brazil. 

Depth: shore to 48 m (to26fm). 

Habitat: sand, shell, rock, and coral bottoms; grassy areas. 

Remarks: Rathbun (1925) lists several specimens that were covered with 

sponges and one crab that was encrusted with worm tubes. She reported ovigerous 

females from Florida in December and from Cuba in June. Listed from 

Florida by Wass (1955), Abele (1970), and Menzel (1971), and from the Gulf 

collections of the R/V Oregon by Chace (1956). Abele (1970) treats this form 

as a separate species and notes the need for revision of this genus. 

Macrocoeloina trispinosum variety Rathbun, 1925 (Bull. U.S. Nat. Mus. 129: 

468) 

Rathbun, 1925, p. 468, pi. 168, fig, 1; Rathbun, 1933, p. 36. 

Range: North Carolina; Bahamas; southeast and west coasts of Florida; Florida 

Keys and Dry Tortugas; Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin 

Islands; Curagao, Netherlands Antilles. 


Habitat: sand, shell, rock, and coral bottoms. 

Remarks: Williams (1965) followed the practice of Rathbun (1925) in treating 

this variety as an unnamed but distinct form, an intermediate linking the 

typical subspecies and M. t. nodipes. If the latter is raised to the status of a full 

species (as in Abele, 1970), then this variety will require reevaluation as a subspecies 

or species. 

Microphrys H. Milne Edwards, 1851 

Microphrys anlillensis Rathbun, 1920 (Proc. Washington Acad. Sci. 33: 24) 

As M. platysoma—Hay & Shore, 1918, p. 459, pi. 38, fig. 9. 

As M. ontiHensis—Rathbun, 1925, p. 498, text-fig. 141, pi. 176, figs. 3, 4; Rathbun, 

1933, p. 38; Williams, 1965, p. 260, figs. 240,245G. 

Range: North Carolina; Bimini; west coast of Florida; north coast of Cuba; 

Jamaica; Puerto Rico; Brazil. 

Depth: 4 to 38 m (2 to 21 fm). 

Habitat: mud, coral, sand, shell, and weed bottoms. 

Remarks: Ovigerous females were reported from North Carohna in September 

(Rathbun, 1925) and from Florida in June and from Bimini in November (Williams, 

1965). Listed from Brazil by Coelho (1971c) and Coelho and Ramos 

(1972).


Microphrys hicornutus (Latreille, 1825) (Encyc. meth., Hist. Nat. Insectes, 

vol. 10, p. 141) 

Common Name: Dirty Decorator Crab 

Hay & Shore, 1918, p. 459, pi. 38, fig. 10; Rathbun, 1925, p. 489, text-fig. 139, 

pi. 175; Rathbun, 1933, p. 37, fig. 32; Williams, 1965, p. 259, figs. 239, 245F. 

Range: North Carolina to south Florida; Bermuda; Bahamas; Florida Keys 

and Dry Tortugas; west and northwest coasts of Florida; north coast of Cuba; 

Jamaica; Puerto Rico; St. Thomas to Barbados; Old Providence Island in the 

Caribbean Sea; Caribbean coast of Panama to Venezuela; Curasao; Trinidad; 

Island of Santa Anna to Santa Catarina, Brazil. 

Depth: shallow water to 30 m (to 16.5 fm). 

Habitat: common on coral reefs; on a variety of substrates, including rock, 

shell, sand, and mud; among grass, mangrove roots, and on sponges; often covered 

with anemones, algae, sponges, etc. 

Remarks: Rathbun (1925) notes that Latreille's original type locality, "Nouvelle 

HoUande," is an error. Ovigerous females occur from March to August in 

the Caribbean and from November to January in the West Indies, Brazil, and 

Venezuela (Williams, 1965). Wass (1955) and Menzel (1971) list this species 

as rare in northwest Florida. Listed from Brazil by Coelho (1971a), Coelho and 

Ramos (1972), and Fausto Filho (1974). Hartnoll (1965a) commented on the 

biology and growth of this crab in Jamaica and Coelho (1971c) provided ecological 

notes on Brazilian specimens. Zoeal stages have been described by Lebour 

(1944) and by Hartnoll (1964b). Hazlett (1972a, 1972b) and Hazlett and Estabrook 

(1974) analyzed agonistic behavior and Hazlett (1971) tested the antennular 

chemosensitivity of this species. Williams (1965) lists records of copepods 

and a tapeworm from this crab. 

Microphrys plalysoma (Rathbun, 1901), not (Stimpson, 1860). 

Specimens from Puerto Rico were designated by Rathbun (1920) as M. 

antillensis, to which all of the Atlantic specimens are referred, separating them 

from the Pacific form of Stimpson. 

Mithrax Desmarest, 1823 

(Although Latreille, 1817 had been traditionally recognized as the author 

of this genus. Garth (1958) questioned the validity of Latreille's citation 

and attributes the first valid citation to Desmarest.) 

Subgenus Mithraculus White, 1847 

Mithrax (Mithraculus) cinctimanus (Stimpson, 1860) (Amer. J. Sci. 29: 132) 


Range: Bahamas; southeast Florida; Florida Keys and Dry Tortugas; Jamaica; 

Puerto Rico; St. Thomas, Virgin Islands; St. Martin; Antigua; Curasao. 

Depth: shallow water. 

Habitat: on coral reefs; rocky bottoms; inside sponges.


Milhrax (Milhraculus) coryphe (Herbst, 1801) (Natur. ICrabben u. Krebse, 

vol. 3, pt. 2, p. 8) 

Rathbun, 192S, p. 426, pi. 153; Rathbun, 1933, p. 31. 

Range: Bahamas; southeast Florida; Florida Keys; north coast of Cuba; 

Jamaica; Puerto Rico; St. Thomas, Virgin Islands to Barbados; Caribbean coast 

of Panama to Colombia; Curasao; Trinidad; Ceara and Fernando de Noronha to 



Habitat: in cavities of corals, rocks, and sponges; on sand, broken shell, grass, 

and mud bottoms. 

Remarks: Listed from Brazil by Coelho and Ramos (1972). 

Milhrax (Milhraculus) forceps (A. Milne Edwards, 1875) (Crust. Reg. Mex., 

p. 109) 

Hay & Shore, 1918, p. 457, pi. 38, fig. 1; Rathbun, 1925, p. 431, pi. 156; Rathbun, 

1933, p. 32; Chace, 1940, p. 67; "Williams, 1965, p. 258, figs. 238, 245E; Pequegnat 

& Ray, 1974, p. 236, figs. 1-4. 

Range: Bermuda; North Carolina to south Florida; Florida Keys and Dry 

Tortugas; west and northwest coasts of Florida; West Flower Garden Bank, off 

Texas; south coast of Cuba; Puerto Rico; St. Thomas, Virgin Islands; Barbados; 

Old Providence Island (Carib.); Netherlands Antilles; Venezuela; Trinidad; 

Ceara to Rio de Janeiro, Brazil. 

Depth: intertidal to 90 m (to 49 fm). 

Habitat: under stones and dead coral; in crevices along rocky shores and reefs; 

in sponges; on sand, shell, coral, rock, and grass bottoms. 

Remarks: Ovigerous females have been reported from Florida in November to 

February, from the Gulf of Mexico in February, and from the southern Caribbean 

in April, mid-summer, September, and November (Williams, 1965). Listed 

from northwest Florida by Wass (1955), Abele (1970), and Menzel (1971). 

Chace (1956) recorded this crab from the Gulf of Mexico collections of the R/V 

Oregon and Pequegnat and Ray (1974) state that this species is one of the most 

common brachyurans on the West Flower Garden reefs, often taken on sponges. 

Williams (1965) reports this crab from the sponge, Stematumenia strobilinia, off 

North Carolina. Lebour (1944) described some of the larval stages. Threat 

behavior was described and illustrated by Schone (1968). Listed from Brazil by 

Coelho (1971a, 1971c), Coelho and Ramos (1972) and FaustoFilho (1974). 

Milhrax (Milhraculus) ruber (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 

118) 


Range: north coast of Cuba; Puerto Rico; St. Thomas, Virgin Islands to Barbados; 

Curasao. 

Depth: shallow water to 46 m (to 25 fm), rare to 154m (84fm). 

Habitat: sand, shell, grass, coral, and mud substrates; off coral reefs; in 

sponges.


Mithrax (Milhraculus) sculptus (Lamarck, 1818) (Hist. Anim. sans Vert., Aôl. 

5, p. 242) 

Rathbun, 1923, p. 422, text-figs. 123-126, pi. 132; Rathbun, 1933, p. 31. 

Range: Bahamas; southeast Florida; Florida Keys and Dry Tortugas; north 

coast of Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin Islands; Antigua; Barbados; 

Swan Island and Old Providence Island (Carib.); Belize; Curagao; Rio 

Grande do Norte to Bahia, Brazil. 


Habitat: abundant on coral reefs; under stones at low tide; on sand, shell, grass, 

and mud bottoms. 

Remarks: Hartnoll (1965a) provides data on the biology and growth of this 

crab in Jamaica. Listed from Brazil by Coelho and Ramos (1972) and by Fausto 

Filho (1974). 

Subgenus Mithrax Desmarest, 1823 

Mithrax (Mithrax) acuticornis Stimpson, 1871 (Bull. MILS. Comp. Zool. 2: 116) 

Rathbun, 1925, p. 388, pi. 136, figs. 1-2, pi. 257, fig. 1; Rathbun, 1933, p. 29, fig. 28; 


Range: southeast coast of Florida; Florida Keys and Straits; Dry Tortugas; 

west and northwest coasts of Florida; off Texas; north and east coasts of Yucatan, 

Mexico; Puerto Rico; Santa Cruz to Grenadines in the Lesser Antilles; Amapa 

to Espirito Santo, Brazil. 

Depth: Iltol03m(6to56fm). 

Habitat: sand, mud, broken shell, rock, and coral substrates. 

Remarks: Rathbun (1925) compared this species with the young of M. 

cornutus and M. spinosissimus, with which it can be confused. Felder (1973a) 

provided the first northwestern Gulf of Mexico records, based on specimens collected 

off Galveston and Port Mansfield, Texas. Listed from Brazil by Coelho 

(1971a, 1971c) and Coelho and Ramos (1972). 

Mithrax (Mithrax) cornutus Saussure, 1857 (Rev. Mag. Zool., ser. 2, 9: 501) 

Common Names: Coral Crab; Red Spider Crab 

Rathbun, 1923, p. 386, pi. 137, figs. 3-4, pi. 236. 

Range: Bermuda; Florida Straits; north coast of Cuba; between Jamaica and 

Haiti; Dominica; Martinique; off Bahia, Brazil. 

Depth: shallow water to 1077 m (589 fm). 

Habitat: coral, sand, and broken shell bottoms. 

Remarks: Listed by Chace (1956) from the Gulf of Mexico. 

Mithrax (Mithrax) hispidus (Herbst, 1790) (Natur. Krabben u. Krebse, vol. 1, 

p. 245 (not p. 247)). 

Common Name: Coral Crab 

Rathbun, 1925, p. 406, text-fig. 124, pis. 143-146, pi. 147, fig. 3; Rathbun, 1933, 

p. 30; Williams, 1963, p. 236, figs. 236, 243C; Pequegnat & Ray, 1974, p. 236, 

figs. 5-10.

Crabs of the Gulf of Mexico 5 7 

Bange: Delaware Bay to south Florida; Bermuda; Bahamas; Florida Keys and 

Dry Tortugas; West Flower Garden Bank, off Texas; Jamaica; Curagao; Para 

to Sao Paulo, Brazil. 


Habitat: sand, shell, and stone bottoms; on coral reefs; inside sponges; occasionally 

on the sea grass, Halodule. 

Remarks: Plentiful on the coral reefs of the West Flower Garden Bank 

(Pequegnat and 'Raj, 1974). Listed from Brazil by Coelho (1971a, 1971c) and 

by Coelho and Ramos (1972). 

Mllhrax (Mithrax) holderi Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 117) 

Rathbun, 1925, p. 392, pi. 138, figs. 1-2, pi. 257, fig. 2; Rathbun, 1933, p. 29. 

Range: Florida Keys and Dry Tortugas; north and south coasts of Cuba; 

Jamaica; Puerto Rico; Virgin Islands. 


Habitat: coral bottoms. 

Mithrax (Mithrax) pilosus Rathbun, 1892 (Proc. U.S. Nat. Mus. 15: 262) 

Rathbun, 1925, p. 394, pi. 138, fig. 3, pi. 258; Rathbun, 1933, p. 29. 

Range: Bahamas; Florida Keys and Dry Tortugas; Vera Cruz, Mexico; Cuba; 

Puerto Rico; St. Thomas, Virgin Islands to Barbados; Caribbean coast of Panama; 

Venezuela. 

Depth: data not available. 

Habitat: rare on stony bottoms; off reefs. 

Mithrax (Mithrax) pleuracanthus Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 

116) 

As M. rfepressus—Hay & Shore, 1918, p. 458, pi. 38, fig. 2. 

As M. pleuracanlhus—ilay & Shore, 1918, p. 458, pi. 38, fig. 3; Rathbun, 1925, 

p. 411, pi. 150; Rathbun, 1933, p. 31; Williams, 1965, p. 257, figs. 237, 245D. 

Range: Bermuda; North and South Carolina; southeast Florida; Florida Kej^s 

and Dry Tortugas; Bahamas; west and northwest coasts of Florida; north coast 

of Yucatan; north coast of Cuba; Puerto Rico; St. Thomas, Virgin Islands; St. 

Martin; Old ProAddence Island (Carib.); Curasao; Venezuela. 


Habitat: common on rocky, gravel, and broken shell substrates; occasionally 

on sand and mud bottoms; in the sponge Stemalumenia strobilinia at Tortugas 

(Pearse, 1934). Often encrusted with bryozoans, serpulid worms, etc. (after 

Williams, 1965). 

Remarks: Listed as common in northwest Florida by Wass (1955), Abele 

(1970), and Menzel (1971). Williams (1965) notes that this species is often 

associated with Mithrax forceps on the banks off North Carolina in April, from 

St. Thomas in July, and from Venezuela in September.


Milhrax (Milhrax) spinosissiinus (Lamarck, 1818) (Hist. Nat. Anim. sans 

Vert., vol. 5, p. 241) 

Common Names: Cangrejo de la Santa Virgen; Cabouca; Lazy Crab 

Rathbun, 1925, p. 383, pi. 135; Rafhbun, 1933, p. 29; Chace, 1940, p. 67; Williams, 

1965, p. 254, figs. 234, 245A. 

Range: North or South (?) Carolina; Florida Keys and Dry Tortugas; north 

and south coasts of Cuba; Jamaica; Haiti; Virgin Islands; Guadeloupe. Type 

locality of "Ile-de-France" is erroneous. 


Habitat: among rocks and on coral sand bottoms; crab is often encrusted with 

stalked barnacles, bryozoans, serpulid worms, red foraminiferans, etc. 

Remarks: Ovigerous females were reported from Cuba in May and June by 

Rathbun (1925). Hazlett and Rittschof (1975) reported on spatial patterns of 

activity. Brownell, Provenzano and Martinez (1977) reported on attempts to 

commercially culture this crab in Venezuela. 

Milhrax (Mithrax) verrucosus H. Milne Edwards, 1832 (Mag. Zool., vol. 2, 

class 7, pi. 4) 

Rathbun, 1925, p. 400, pi. 144; Ratlibun, 1933, p. 30; Williams, 1965, p. 255, figs. 

235, 245B. 

Range: South Carolina; Bahamas; southeast Florida; Florida Keys and Dry 

Tortugas; north coast of Cuba; Swan Islands (Carib.) Jamaica; Hispaniola; 

Puerto Rico; St. Thomas, Virgin Islands; Guadeloupe; Curasao; Martinique; 

Rocas and Fernando do Noronha, Brazil. 

Depth: shallow wâter, near shore. 

Habitat: among rocks; hides in crevices and holes; often found near madrepores; 

has nocturnal habits. 

Remarks: Found in Porf^g^ ;3o/-fte5 at Curasao. Pearse (1932b) determined the 

freezing point of hemolymph from this crab at Dry Tortugas. Listed from Brazil 

by Coelho (1971a, 1971c), Coelho and Ramos (1972) and by Fausto Filho 

(1974). 

Stenocionops Desmarest, 1823 

Slenocionops furcala furcula (Oliver, 1791) (Encyc. meth., Hist. Nat., Insects, 

vol. 6, p. 174) 

Common Names: Decorator Crab; Macca Crab 

Rathbun, 1925, p. 449, text-fig, 131, pis, 160-161; Rathbun, 1933, p. 33, fig. 30; 

Guinot-Dumortier, 1960, p. 180, fig. 21a-b. 

Range: Georgia; Florida (location unspecified); ? Gulf of Mexico; Jamaica; 

Puerto Rico; St. Thomas, Virgin Islands; Dominica; Barbados; French Guiana; 

Paraiba to Rio de Janeiro, Brazil; South Africa. 

Depth: shallow water to 64 m (to 35 f m). 

Habitat: mud, sand, coral, rock, and shell bottoms; on wharf pilings. 

Remarks: The inclusion of this form in the Gulf fauna is doubtful. Although 

Leary (1967) and Felder (1973a) report this form from Texas, both reports are


based on a listing by Hildebrand (1954) in which the subspecies identification 

may have been in error. Felder (1973a) indicates that more recent collections 

in the northwestern Gulf are of crabs similar to S. furcata coelata, and other 

records confirm the presence of this latter form in the region. Neither of the 

two specimens cited bj^ Rathbun (1925, p. 542) are determined for the Gulf. 

Guinot-Dumortier (1960) remarked that the male pleopod of this species is very 

similar to the pleopods of three Pacific species figured by Garth (1958). Listed 

fromBrazil by Coelho and Ramos (1972). 

Stenocionops furcata coelata (A. Milne Edwards, 1878) (Bull. Soc. Philom., 

ser. 7,2:224) 

Hay & Shore, 1918, p. 460, pi. 39, fig. 3; Kathbun, 1925, p. +50, pi. 164; Kathbun, 

1933, p. 34; Williams, 1965, p. 261, figs. 241, 245H; Felder, 1973a, p. 53, pi. 7, 

fig. 12. 

Range: North Carolina to south Florida; Florida Keys and Dry Tortugas; west 

and northwest coasts of Florida; Alabama to Texas; north and east coasts of 

Yucatan; north coast of Cuba; Puerto Rico; St. Lucia; Barbados. 


Habitat: sand, shell, and coral bottoms, usually of coarse material; on shelly 

reefs off North Carolina. 

Remarks: Early Gulf records distinguishing this subspecies from the typical 

form may be confused (Felder, 1973a). A single specimen taken off St. Joseph 

Island, Texas (Hildebrand, 1954) is probably coelata; it was covered with algae, 

hydroids, bryozoans, three small Ostrea, and three large Calliactis tricolor. 

Listed from the Gulf of Mexico (Chace, 1956) and from northwest Florida 

(Wass, 1955; Hulings, 1961; Abele, 1970). Ovigerous females occur in Florida 

from March to August (Williams, 1965). 

Stenocionops spinimana (Rathbun, 1892) (Proc. U.S. Nat. Mus. 15: 240) 

Hay & Shore, 1918, p. 460, pi. 39, fig. 2; Rathbun, 1925, p. 457, pi. 267; Williams, 

1965, p. 262, figs. 242, 2451; Pequegnat, 1970, p. 182. 

Range: North and South Carolina; Florida Keys; west coast of Florida to off 

Mississippi. 

Depth: 37 to 227 m (20tol24fm). 

Habitat: mud, sand, shell, coral, and rock bottoms. 

Remarks: Williams (1965) noted the age-related variability in the morphology 

of this and related species. Rathbun (1925) listed ovigerous females from 

South Carolina in December. Listed by Chace (1956) from the Gulf of Mexico 

and by Franks et al. (1972) off Mississippi at 37 to 91 m depth. 

Stenocionops spinosissima (Saussure, 1857) (Rev. Mag. Zool., ser. 2, 9: 501) 

Rathbun, 1925, p. 455, pi. 165, fig. 2, pi. 264, figs. 3^., pi. 265; Chace, 1940, p. 67; 

Williams, McCloskey & Gray, 1968, p. 62; Pequegnat, 1970, p. 182; Felder, 1973a, 

p.52,pl. 7,fig. 11. 

Range: North Carolina; south and southwest Florida; off Texas and east coast 

of Mexico; north coast of Cuba; Haiti; Guadeloupe; Dominica; Rio de Janeiro 

and Fernando de Noronha, Brazil.


Depth: 46 to 480 m (25 to 260 fm); center of depth distribution in Gulf is at 

110tol83m(60tol00fm) (Pequegnat, 1970). 

Habitat: mud and sand bottoms. 

Remarks: Hildebrand (1954) found this crab to be common off the Texas 

coast at depths of 25 to 37 fm (46 to 68 m) and he reported ovigerous females 

in February and April. Listed from Texas by Leary (1967) and from the Gulf 

of Mexico by Chace (1956). Recorded from Brazil by Coelho and Ramos (1972) 

andbyFaustoFilho (1974). 

Teleophrys Stimpson; 1860 

Teleophrys ornatus Rathbun, 1901 (Bull. U.S. Fish. Comm. 20: 65) 

Rathbuii, 1925, p. 444, text-fig. 129, pi. 139, figs. 3-4, pi. 262, figs. 8-9; Rathbun, 

1933, p. 33, fig. 29. 

Range: off northeast Yucatan (Gulf); Puerto Rico; St. Croix; Fernando de 

Noronha, Brazil. 

Depth: 7 to 44 m (4 to 24 fm). 

Habitat: rock and coral bottoms. 

Remarks: Listed from Brazil by Coelho and Ramos (1972) and Fausto Filho 

(1974). 

r/ioeBell, 1836 

Thoe puella Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 178) 

Rathbun, 1925, p. 348, text-figs. 111-112, pi. 125, figs. 1-2; Rathbuij, 1933, p. 21, 

fig. 24. 

Range: Florida Keys and Dry Tortugas; Jamaica; Puerto Rico; St. Thomas, 

Virgin Islands; Guadeloupe; Curasao. 

Depth: Shallow water. 

Habitat: in and on coral reefs. 

Subfamily OPHTHALMIINAE Balss, 1929 

Picroceroirfes Miers, 1886 

Pieroceraides luhularis Miers, 1886 {Challenger Rept., Zool. 17: 77) 

Rathbun, 1925, p. 354, text-fig. 115, pi. 126, pi. 254, figs. 2-5. 

Range: Bahamas; southeast Florida; north and south coasts of Cuba; between 

Jamiaica and Haiti: St. Thomas, Virgin Islands; Maranhao to Espirito Santo, 

Brazil. 


Habitat: coral and broken shell bottonas. 

Remarks: Listed from Brazil by Coelho (1971c), Coelho and Ramos (1972), 

and Fausto Filho (1974).

Pif/io Bell, 1835 


Pitho aculeata (Gibbes, 1850) (Proc. Amer. Assoc. Adv. Sci. 3: 171) 

Rathbun, 1925, p. 357, text-fig. 116c, pi. 127, pi. 251, fig. 1; Ratlibun, 1933, p. 23, 

fig. 26. 

Range: Bahamas; Florida Keys and Dry Tortugas; west coast of Florida; north 

coast of Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin Islands; Guadeloupe; 

Old Providence Island (Carib.); Netherlands Antilles. 

Depth: Shallow water. 

Habitat: sand, shell, coral, grass, and mud bottoms; among algae in lagoons; 

on Sargassum; on banks at low tide. 

Pitho anisodon (von Martens, 1872) (Arch. f. Naturg. 38: 83) 

Rathbun, 1925, p. 368, text-figs. 116b, 117d, 118, pi. 131, pi. 251, fig. 2; Rathbun, 

1933, p. 24. 

Range: Bahamas; south, west, and northwest coasts of Florida; Florida Keys; 

north coast of Cuba; Jamaica; Puerto Rico; Guadeloupe; Curagao, Netherlands 

Antilles. 


Habitat: rock, sand, mud, coral, and grass bottoms. 

Remarks: Reported from Florida by Wass (1955), Tabb and Manning (1961), 

Dragovich and Kelly (1964), Abele (1970), and Menzel (1971). Tabb and 

Manning (1961) noted the presence of ovigerous females in Florida Bay when 

salinities were fully marine and in Tampa Bay in March. Abele (1970) remarked 

on the variability of lateral and orbital spines with size of crab. 

Pitho laevigata (A. Milne Edwards, 1875) (Crust. Reg. Mex., p. 116) 

Rathbun, 1925, p. 372, pi. 132, figs. 3-4, pi. 133, fig. 3, pi. 250, figs. 11-13. 

Range: west and northwest coasts of Florida; Antilles, loc. unspec; Colombia; 

Trinidad. 

Depth: shallow water, hsted from 10.5 m (about 6 fm). 

Habitat: coral, rock, and grass bottoms. 

Remarks: Listed from northwest Florida by Wass (1955). Rathbun (1925) 

described a variety of this crab based on a female collected from the west coast 

of Florida. 

Pitho Iherininieri (Schramm, 1867) (Crust. Guadeloupe, Desbonne & Schramm, 

p. 20) 

Hay & Shore, 1918, p. 459, pi. 38, fig. 8; Rathbun, 1925, p. 362, text-fig. 116a, 117b, 

pi. 128, figs. 1-2, pi. 129, figs. 1-2, pi. 252, fig. 2; Rathbun, 1933, p. 24; Williams, 

1965, p. 246, figs. 224,233A. 

Range: North and South Carolina; Bahamas; Florida Keys; west and northwest 

coasts of Florida; Vera Cruz, Mexico; north coast of Cuba; Jamaica; Puerto 

Rico; St. Thomas, Virgin Islands to Martinique; Old Providence Island (Carib.); 

Curagao; Cape St. Roque to Sao Paulo, Brazil. 


Habitat: coral, sand, shell, rock, and grass bottoms; rarety on mud.


Remarks: Listed from northwest Florida b)^ Wass (1955); recorded from 

Brazil by Coelho (1971c), Coelho and Ramos (1972) and Fausto Filho (1974). 

Ovigerous females have been collected from the Bahamas and Florida in May to 

November and from Brazil in December (Williams, 1965). 

Pilho mirabilis (Herbst, 1794) (Naturg. Krabben u. Krebse, vol. 2, p. 152) 

Rathbun, 1925, p. 366, text-figs. 116d, 117c, pi. 128, fig. 3, pi. 129, fig. 3, pi, 253, 

fig. 1; Rathbun, 1933, p. 24. 

Range: Bahamas; Florida Keys; Puerto Rico; Guadeloupe. 


Habitat: rock and coral bottoms. 

Remarks: Only part of the original species description by Herbst is valid 

(Rathbun, 1925). 

Tyc/i€ Bell, 1835 

Tyche emarginata White, 1847 (Ann. Mag. Nat. Hist. 20: 206) 

Hay & Shore, 1918, p. 461, pi. 39, fig. 4; Rathbun, 1925, p. 508, pi. 272, pi. 273, 

figs. 7-12,- Garth, 1946, p. 406, text-fig. 1; Williams, 1965, p. 247, figs. 225, 226, 

233B. 

Range: North Carolina; Bahamas; Florida Keys and Dry Tortugas; west and 

northwest coasts of Florida; St. Thomas, Virgin Islands; Guadeloupe; Cape St. 

Roque, Brazil. 


Habitat: shell and rock bottoms. 

Remarks: Garth (1946) compared this species with its Pacific analogue, T. 

lamellifrons. Listed from Florida by Wass (1955) and Lyons et al. (1971) and 

from the Gulf collections of the R/V Oregon by Chace (1956). 

Subfamily PISINAE Dana, 1852 

Chorinns Latreille, 1825 

Chorinus heros (Herbst, 1790) (Naturg. Krabben u. Krebse, vol. 1. pi. 18, fig. 

2) 

Rathbun, 1925, p. 305, text-fig. 101, pi. 107, pi. 246, figs. 3-5; Rathbun, 1933, p. 20, 

fig. 21. 

Range: Bermuda; Florida Keys and Dry Tortugas; Cuba; Caribbean coast of 

Yucatan, Mexico; Jamaica; Hispaniola; Puerto Rico; St. Croix; Barbados; Ceara 

to Bahia, Brazil. 


Habitat: sand, shell, rock, and coral bottoms. 

Remarks: Listed from the Gulf of Mexico by Chace (1956) and from Brazil 

by Coelho (1971c) and Coelho and Ramos (1972).

Holoplites Rathbun, 1894 


Holoplites armala (A. Milne Edwards, 1880) (Crust. Reg. Mex., p. 348) 

Rathbun, 1925, p. 307, text-fig. 102, pi. 108, pi. 245, figs. 6-8; Chace, 1940, p. 64; 

BuIIis & Thompson, 1965, p. 12. 

Range: north coast of Cuba; St. Vincent; Grenadines; Barbados; Para, Brazil. 

Depth: 161 to 798 m (88 to 387 fm). 

Habitat: coral, shell, and rock bottoms. 


Lifoinja Leach, 1815 

Libinia dubia H. Milne Edwards, 1834 (Hist. Nat. Crust., vol. 1, p. 300) 

Hay & Shore, 1918, p. 456, pi. 38, fig. 5; Rathbun, 1925, p. 313, text-figs. 105-106, 

pis. 114-115, 122, fig. 1; Williams, 1965, p. 252, figs. 232, 233G; Felder, 1973a, 

p. 52, pi. 7, fig. 8. 

Range: Cape Cod, Massachusetts to south Florida; Bahamas; Florida Keys; 

west Florida to south Texas; Cuba; off Gabon, western Africa. 


Habitat: primarily on mud and mud-sand substrates; also on sand, gravel, and 

shell bottoms; often near shore and occasionally in tide pools. Juveniles are usually 

covered with hydroids, ascidians, sponges, etc., but older adults are almost 

always clean. 

Remarks: Gulf reports of this species may be confused with those for L. 

emarginata, especially records of juveniles for which identification is difficult 

and further complicated by the variety of attached organisms. Regional lists 

include Florida (Wass, 1955; Tabb and Manning, 1961; Dragovich and Kelly, 

1964; Lyons et al., 1971; Menzel, 1971), Mississippi (Richmond, 1962; Christmas 

and Langley, 1973), Louisiana (Behre, 1950; Hoese and Valentine, 1972), 

and Texas (Leary, 1967). Fotheringham and Brunenmeister (1975) describe 

some aspects of natural history for this crab in the Gulf. 

Tabb and Manning (1961) found that larger adults were prevalent in the 

sponge-algae areas of Florida Bay, whereas smaller individuals were more common 

in the Thalassia beds. Wass (1955) states that this species is more common 

than L. emarginata in the shallower harbor and bay waters, but that the reverse 

is true in the more marine outer waters of northwest Florida. This crab occurred 

in salinity ranges of 20.4 to 39 ppt in Texas bays after a drought (Hoese, 1960) 

and down to 12 ppt at Crystal River, Florida (Lyons et al., 1971). In Tampa 

Bay, juveniles are common on Gracilaria beds and this species is the most common 

spider crab of the area (Dragovich and Kelly, 1964). Ovigerous females 

have been reported in Florida from January to July (Dragovich and Kelly, 1964; 

Lyons e/a/., 1971). 

Several authors have commented on the presence of young L. dubia in or on the 

cabbagehead jellyfish, Stomolophus meleagris. Corrington (1927) and Gutsell 

(1928) record this relationship in Carolina waters and Williams (1965) states 

that small crabs have been found in the genital pits and subumbrellar space.


Jachowski (1963) noted a similar relationship with Aurelia aurita. Wass (1955) 

noted that one female crab was covered with 93 barnacles {Balanus) and Pearse 

(1952) reported copepods in the gills and on the carapace of this crab in Texas 

waters. 

Development of this species in the laboratory was described by Sandifer and 

van Engel (1971). Ayres (1938) compared relationships between habitat and 

oxygen consumption and Gray (1957) correlated habitat and gill surface area 

in this species. Other physiological studies include tolerance to desiccation and 

salinity changes (Pearse, 1929) and the physiological activitj^ and neurosecretions 

of the pericardial organs (Berlind and Cooke, 1970). 

Libinia eniarginala Leach, 1815 (Zool. Misc., 2: 130) 

Common Name: Common Spider Crab 

Hay & Shore, 1918, p. 456, pi. 38, fig. 6; Rathbun, 1925, p. 311, text-figs. 103-104', 

pis. 110-113; Williams, 1965, p. 232, figs. 231, 233H; Felder, 1973a, p. 52, pi. 7, 

fig. 7. 

Range: Nova Scotia to south Florida; Florida Keys; west coast of Florida to 

Mexico. Pacific coast records are probably erroneous (Garth, 1958). 

Depth: shore to 49m (to27fm),rarely to 124 m (68 fm). 

Habitat: on all types of substrates, but most common on mud and mud-sand 

in shallow waters. 

Remarks: Gulf records of this species and L. dubia may be confused due to 

similarity of the juvenile stages. Wass (1955) compares the key morphological 

features of the two species. Listed from Florida (Wass, 1955; Menzel, 1971), 

Mississippi (Richmond, 1962; Franks ei al, 1972), Louisiana (Behre, 1950; 

Hoese and Valentine, 1972), Texas (Gunter, 1950; Hildebrand, 1954; Parker, 

1959; Leary, 1967), Mexico (Hildebrand, 1954), and the Gulf of Mexico (Chace, 

1956). 

Hildebrand (1954) states that this species is the most common large spider 

crab in the westez-n Gulf of Mexico, reversing the numerical dominance relationship 

with L. dubia that is found in the eastern Gulf. Like L. dubia, young L. 

cmarginala are often associated with the cabbagehead jellyfish, Siomolophus 

meleagris. Ovigerous females are present in the western Gulf in February and 

are common in July. Hoese and Valentine (1972) collected a crab from the 

Chandeleur Islands that was covered with the bryozoan, Bugula. Musick and 

McEachran (1972) reported a depth range of 18 to 51 m for this crab in Chesapeake 

Bight. Aldrich (1976) reported predaticn by this crab on the starfish, 

Asterias. Forward (1977) described shadow responses of the larval stages. 

Other studies include: measurement of gill surface area in relation to habitat 

(Gray, 1957), biochemical adaptations to temperature variation (Vernberg and 

Vernberg, 1968), osmoregulation (Gilles, 1970), physiology of molting (Skinner 

and Graham, 1972), sterol synthesis in larval stages (Whitney, 1969), vitellogenesis 

(Hinsch and Cone, 1969), sperm structure (Hinsch, 1973), growth biometrics 

(Aldrich, 1974), reproductive physiology (Hinsch, 1970), neural fine 

structure (Skobe and Nunnemacher, 1970), antennule chemosensitivity (Haz-


lett, 1971), pericardial organ neurosecretion (Berlind and Cook, 1970), and 

behavior related to copulation and reproduction (Hinsch, 1968). 

Lihinia erinacea (A. Milne Edwards, 1879) (Crust. Reg. Mex., p. 202) 

Rathbun, 1925, p. 321, pi. 109. 

Range: Florida Keys; southeast to northwest Florida; north coast of Cuba. 

Depth: 4 to 68 m (2 to 37 fm). 

Habitat: sand-mud and rock bottoms; in patches of moss. 

Remarks: The status of this species is confused due to a paucity of mature 

specimens. Rathbun (1925) remarked on the close resemblance of this specie.^ 

to L. dubia, but she also listed differences between the two forms. Tabb and 

Manning (1961) also questioned the status of this crab as a distinct species and 

Abele (1970) compared the types of the two species and concluded that further 

examination of more mature specimens will be required. Abele (1970) also 

noted that the type specimen of L. erinacea is in the Museum of Comparative 

Zoology (Harvard) and not in the Paris Museum, as indicated bj'^ Rathbun 

(1925). Listed as uncommon at Apalachee Bay, Florida hj Menzel (1971). 

Libinia rhoinhoidea Streets, 1870 (Proc. Acad. Nat. Sci. Philadelphia, p. 106) 

Rathbun, 1925, p. 323, pis. 116-117, pi. 24.5, figs. 1-3. 

Range: west and north coasts of Cuba; off Merida, Yucatan (Gulf coast). 

Depth and Habitat: no data available. 

Remarks: Rathbun (1925) believed that the tj^pe locality of "East Indies" 

was probably an error. She also compared this species with L. dubia and noted 

variation in size of the spines. 

mhilia A. Milne Edwards, 1878 

^'ibiIia anlilocapra (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 110) 

Rathbun, 192S, p. 290, text-fig. 97, pis. 102-103, 239; Williams, 1965, p. 251, figs. 

230, 233F. 

Range: North Carolina; southeast Florida; off Mobile, Alabama; Gulf of 

Campeche, Mexico; St. Vincent; Barbados. 

Depth: 71 to256m (39to 140fm). 

Habitat: mud, sand, broken shell, rock, and coral bottoms. 

Remarks: Ovigerous females were collected at St. Vincent in February and at 

Barbados in March (Rathbun, 1925). Recorded by Chace (1956) from the Gulf 

of Mexico, some distance west of Dry Tortugas. 

PeZi« Bell, 1836 

Pelia miillca (Gibbes, 1850) (Proc. Amer. Assoc. Adv. Sci. 3: 171) 


figs. 2-3; Ratiibun, 1933, p. 18, fig. 19; Williams, 1965, p. 250, figs. 229, 233E; 


Range: Massachusetts to south Florida; Florida Keys; west coast of Florida to


south Texas; north coast of Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin 

Islands. 


Habitat: shell or rock rubble; mud, sand, gravel, and coral substrates; among 

hydroids, ascidians, and sponges on wharf pilings; from Chaetopterus tubes. 

Remarks: Ovigerous females have been collected from Florida in February to 

July, through the summer in the Carolinas, and from Massachusetts in July. 

Regional lists include Florida (Wass, 1955; Tabb and Manning, 1961; Menzel, 

1971), Louisiana (Behre, 1950; Hoese and Valentine, 1972), and Texas (Felder, 

1973a). Tabb and Manning (1961) reported that all of the crabs collected in 

Florida Bay were covered with algae, sponges, or hydroids; Wass (1955) noted 

that this crab is common in clumps of the ascidian, Styela. Gray (1961) reported 

the crab in tubes of the annelid, Chaetopterus, in North Carolina. Hartnoll 

(1965a) studied growth and other aspects of biology of this species in Jamaica. 

Rochinia A. Milne Edwards, 1875 

Rochinia crassa (A. Milne Edwards, 1879) (Crust. Reg. Mex., p. 203) 

Rathbun, 1925, p. 210, text-figs. 83-84, pis. 68-69, 226; Chace, 1940, p. 62; 

Williams, McCloskey & Gray, 1968, p. 60; Pequegnat, 1970, p. 183. 

Range: Massachusetts to south Florida; Florida Straits; off Alabama and 

Texas; east coast of Mexico; north coast of Cuba. 


Habitat: mud, sand, and coral oozes. 

Remarks: Rathbun (1925) reported ovigerous females from off South Carolina 

in December. Musick and McEachran (1972) listed this crab from 194 m depth 

in Chesapeake Bight. Listed by Chace (1956) from the R/V Oregon collections 

in the Gulf of Mexico. 

Rochinia hystrix (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 124) 

Rathbun, 1925, p. 214, pis. 70-71; Rathbun, 1933, p. 17, fig. 17; Chace, 1940, p. 62. 

Range: Florida Keys and Dry Tortugas; north coast of Cuba; off northwest 

Florida and Mississippi; Puerto Rico; Martinique to Barbados. 

Depth: 150 to 708 m (82to387fm). 

Habitat: sand, coral, and rocky substrates. 

Remarks: Rathbun (1925) listed ovigerous females from off Key West in 

February. Chace (1940) commented on sample numbers from off Cuba and 

Chace (1956) listed this species from the Gulf collections of the R/V Oregon. 

Rochinia tanneri (Smith, 1883) (Proc. U.S. Nat. Mus. 6: 4) 

Rathbun, 1925, p. 216, pi. 227, fig. 1; WilHams, McCloskey & Gray, 1968, p. 60, 

fig. 15. 

Range: Massachusetts to North Carolina; southeast Florida; off Key West and 

Florida Straits. 

Depth: 128 to 708 m (70 to 387 fm). 

Habitat: sand and shell bottoms.


Rochinia umhonala (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 115) 

Rathbun, 1925, p. 222, text-fig. 85, pi. 72, pi. 73, fig. 1; Chace, 1940, p. 63; 

Williams, McCloskey & Gray, 1968, fig. 16; Pequegnat, 1970, p. 183. 

Range: North Carolina to south Florida; Florida Straits, off Key West; off 

Mississippi; St. Vincent, Windward Islands. 

Depth: 161 to 900 m {88 to 492 fm). 

Habitat: sand, gravel, coral, shell, and foraminiferan substrates. 

Remarks: Listed by Chace (1956) from the Gulf of Mexico. 

Family PARTHENOPIDAE Macleay, 1838 

Subfamily PARTHENOPINAE Macleay, 1838 

Cryptopodia H. Milne Edwards, 1834 

Crypiopodia concava Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 137) 

Rathbun, 1925, p. 553, text-fig. 151, pi. 202, figs. 3-4, pi, 282, figs. 6-11; Rathbun, 

1933, p. 42, fig. 37; Williams, McCloskey & Gray, 1968, p. 64. 

Range: off North Carolina; Bahamas; Florida Ke5^s and Dry Tortugas; west 

coast of Florida; St. Thomas, Virgin Islands; Ceara to Bahia, Brazil. 

Depth: 7 to 62 m (4 to 34 fm). 

Habitat: mud, sand, shell, and coral bottoms. 


Heterocrypta Stimpsou, 1871 

Heterocrypta granulata (Gibbes, 1850) (Proc. Amer. Assoc. Adv. Sci. 3: 173) 

Common Name: Pentagon Crab 


figs. 1-2, pi. 282, figs. 1-3; Rathbun, 1933, p. 43, fig. 38; Williams, 1965, p. 270, 

figs. 251, 252E; Felder, 1973a, p. 45, pi. 6, fig. 6. 

Range: Massachusetts to Georgia; Florida Keys and Straits; west coast of 

Florida to south Texas; Jamaica; Puerto Rico; St. Thomas, Virgin Islands; Ceara 

to Bahia, Brazil. 

Depth: 4tol37m(2to75fm). 

Habitat: mud, sand, gravel, shell, rock, and coral bottoms; this crab is difficult 

to detect on pebble and shell substrates, where its form and coloration provide 

excellent camouflage. 

Remarks: Williams (1965) reports ovigerous females throughout the summer 

off North Carolina. Regional lists include Florida (Wass, 1955; Tabb and Manning, 

1961; Dragovich and Kelly, 1964; Rouse, 1970; Menzel, 1971), Louisiana 

(Behre, 1950) and Texas (Hedgpeth, 1953; Parker, 1959; Leary, 1967). Listed 

from Brazil by Coelho and Ramos (1972).


Leiolamhrus A. Milne Edwards, 1878 

Leiolambrus nitidus Rathbun, 1901 (Bull. U.S. Fish. Comm. 20: 80) 

Rathbun, 1925, p. 545, pi. 199, pi. 281, fig. 1; Rathbun, 1933, p. 41, fig. 35; Guinot- 

Dumortier, 1960, p. 182, figs. 23a-b, 26; Felder, 1973a, p. 45, pi. 6, fig. 7. 

Range: Gulf of Mexico, from off Alabama to south Texas; Jamaica; Puerto 

Rico; French Guiana. 

Depth: 7 to 73 m (4 to 40 fm). 

Habitat: mud, mud-sand, and mud-shell bottoms. 

Remarks: Regionallists include Alabama (Chace, 1956), Louisiana (Dawson, 

1966), and Texas (Hildebrand, 1954; Leary, 1967). Hildebrand (1954) reported 

ovigerous females from off Texas in June. Listed from French Guiana by Guinot- 

Dumortier (1960). 

Mesorhoea Stimpson, 1871 

Mesorhoea sexspinosa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 136) 

Rathbun, 1925, p. 547, text-fig. 150, pi. 200; Rathbun, 1933, p. 42, fig. 36; Williams, 

McCloskey & Gray, 1968, p. 64, fig. 17. 

Range: North Carolina; Florida Keys and Dry Tortugas; northwest coast of 

Florida; Puerto Rico; Virgin Islands. 

Depth: 7 to 49 m (4 to 27 fm). 

Habitat: sand and shell substrates. 

Remarks: Rathbun (1925) reported an ovigerous female from off northwest 

Florida in January. 

Parlhenope Weber, 1795 

Parlhenope (Parlhenope) agonus (Stimpson, 1871) (Bull. Mus. Comp. Zool. 

2: 131) 

As P. ogoiMi—Hay & Shore, 1918, p. 462, pi. 39, fig. 5; Williams, 1965, p. 266, 

figs. 246, 252A; Pequegnat, 1970, p. 183. 

As P. agonus—Rathbun, 1925, p. 313, text-fig. US, pis. 178-179, pi. 273, figs. 1-3; 

Rathbun, 1933, p. 39. 

Range: North Carolina; Florida Straits; Dry Tortugas; west coast of Florida; 

Puerto Rico; Trinidad; off Guianas. 

Depth: 46 to 391 m (25 to 214 fm). 

Habitat: sand, broken shell, gravel, coral, and mud bottoms. 

Remarks: Ovigerous females were reported from northwest Florida in March 

(Rathbun, 1925) and from the Guianas in September (Williams, 1965). Chace 

(1956) lists this crab from the eastern Gulf of Mexico. 

Subgenus Plalylanihrus Stimpson, 1871 

Parlhenope (Platylanibrus) fralerculus (Stimpson, 1871) (Bull. Mus. Comp. 

Zool. 2: 130) 

Rathbun, 1925, p. 323, pis. 186-187, pi. 190, fig. 2; Williams, 1965, p. 269, figs. 

249, 252D.


Range: North Carohna; southeast Florida; Florida Keys and Straits; Dry 

Tortugas; west and northwest coasts of Florida; northeast coast of Yucatan; 

Barbados; miouth of Amazon River, Brazil; Surinami. 

Depth: 7 to 201m (4tollOfm). 

Habitat: sand, shell, gravel, rock, and coral bottoms. 

Remarks: Williams (1965) listed ovigerous females from south Florida in 

Majr and from northwestern Florida in August. Listed by Chace (1956) from 

the northwestern Gulf of Mexico and by Holthuis (1959) from Surinam. 

Parllienope (Platylambrus) pourtalesii (Stimpson, 1871) (Bull. Mus. Comp. 

Zool. 2: 129) 

Hay & Shore, 1918, p. 462, pi. 39, fig. 6; Rathbun, 1925, p. 521, pis. 182-183, 276; 

Rathbun, 1933, p. 39, fig. 33; Chace, 1940, p. 53; Williams, 1965, p. 268, figs. 248, 

252C; Pequegnat, 1970, p. 183; Felder, 1973a, p. 48, pi. 6, fig. 9. 

Range: New Jersey to south Florida; Florida Keys and Straits; Dry Tortugas; 

north coast of Yucatan; off north coast of Cuba; Grenada. 

Depth: 18to348m(10tol90fm). 

Habitat: primariljr mud and sand-mud bottoms; also on sand, shell, and gravel 

substrates. Pequegnat (1970) states that Gulf populations are probably centered 

at the mdddle of the continental shelf. 

Remarks: Ovigerous females are kno-wn from North Carolina in December 

(Williams, 1965) and from the southeast Gulf in July (Pequegnat, 1970). 

Listed by Chace (1956) from off Yucatan. Leary (1967) includes this species 

on a Texas list and this is repeated by Felder (1973a), but I know of no actual 

records to confirm collection of this crab in the northwestern Gulf. 

Parllienope (Platylambrus) serrala (H. Milne Edwards, 1834) (Hist. Nat. 

Crust., vol. 1, p. 357) 

As Platylambrus serralus—Hay & Shore, 1918, p. 463, pi. 39, fig. 7. 

As Parthenope serr«t«—Rathbun, 1925, p. 516, pis. 180-181, pi. 275, figs. 7-10; 

Rathbun, 1933, p. 39; Williams, 1965, p. 267, figs. 247, 252B; Turkay, 1968, p. 251; 

Felder, 1973a, p. •l-S, pi. 6, fig. 8. 

Range: North Carolina; Bermuda; Bahamas; southeast and south Florida; 

Florida Keys and Dry Tortugas; west Florida to off Texas; Bay of Campeche, 

off Yucatan; north coast of Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin 

Islands; Curagao; Surinam; Bahia, Brazil. 


Habitat: mainly on mud and mud-sand bottoms; also from sand, shell, gravel 

and coral substrates. 

Remarks: Gore (1977) reviews this species and concludes that two distinct 

species have been confused under this name. Ovigerous females are listed from 

North Carolina in June, from Florida in summer, from Cuba in October, and 

from Surinam in May to June (Williams, 1965). Hildebrand (1955) reported 

this crab present on the pink shrimp grounds at Campeche. Wass (1955) and 

Menzel (1971) indicated that this crab was rare off northwestern Florida; Chace 

(1956) listed a single record from off Mississippi for the collections of the R/V 

Oregon. Listed from Texas by Leary (1967), but Felder (1973a) cites only a


single collection record from the northwestern Gulf, a specimen taken from a 

snapper stomach off south Texas. Listed from Surinam by Holthuis (1959). 

Solenolainbrus Stimpson, 1871 

Solenolainbrus deceinspinosus Rathbun, 1894 (Proc. U.S. Nat. Mus. 17: 84) 


Range: off northwest Florida; Puerto Rico. 

Depth: 82 to 110 m (45 to 60 fm). 

Habitat: sand and sand-mud bottoms. 

Solenolainbrus tenellus Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 134) 

Hay & Shore, 1918, p. 463, pi. 39, fig. 8; Rathbun, 1925, p. 541, pi. 194, figs. 3-4, 

pi. 279, figs. 5-9; Rathbun, 1933, p. 41; Williams, 1965, p. 270, fig. 250. 

Range: North Carolina; southeast Florida; Bahamas; Florida Keys; west coast 

of Florida; Barbados. 

Depth: 55to210m (30to 115fm). 

Habitat: sand, broken coral, and rocky bottoms. 

Remarks: Ovigerous females are known from Florida in May-June and from 

Barbados in May (Rathbun, 1925). 

Solenolainbrus typicus Stimpson, 1871 (Bull. Mus. Comp. 2k)ol. 2: 133) 

Rathbun, 1925, p. 537, text-fig. 148, pis. 192-193, pi. 279, figs. 1-4; Rathbun, 1933, 

p. 40, fig. 34; Chace, 1940, p. 53; Williams, McCloskey & Gray, 1968, p. 63; 


Range: North Carolina; Bahamas; southeast Florida; Florida Keys and Straits; 

Dry Tortugas; off Texas; north and south coasts of Cuba. 

Depth: 91 to 618 m (50 to338 fm). 

Habitat: sand, broken shell, and coral substrates. 

Remarks: Location of the R/V Alaminos record is confused. Table 6-9 of 

Pequegnat (1970, p. 201) lists this species from the southwestern Gulf and a 

synopsis of distribution (p. 184) includes a range from the Bahamas to northwest 

Florida, but the actual collecting station of record (64-A-10-13) is located 

off south Texas. 

Thryrolambrus Viathhun, 1894 

Thyrolambrus astroides Rathbun, 1894 (Proc. U.S. Nat. Mus. 17: 83) 

Rathbun, 1925, p. 532, text-fig. 147, pj. 196, pi. 280, figs. 5-6. 

Range: off north coast of Cuba; Mauritius and Andaman Sea, in the In do- 

Pacific region. 

Depth: 123 to 366 m (67 to 200 fm) off Cuba; 66 m (36 fm) in Andaman Sea. 

Habitat: coral bottoms.

Tutankhamen Rathbun, 1925 


Tutankhamen cristatipes (A. Milne Edwards, 1880) (Crust. Reg. Mex., p. 352) 


Range: Pourtales Plateau, Florida Straits; St. Vincent, Windward Islands. 

Depth: 227 to366 m( 124 to200 fm). 

SECTION CANCRIDEA Latreille, 1803 

Superfamily CANCROIDEA Latreille, 1803 

Family ATELECYCLIDAE Ortmann, 1893 

Subfamily ATELECYCLINAE Ortmann, 1893 

Trachycarcinus Faxon, 1893 

Trachycarcinus spinulifer Rathbun, 1898. 

Determined by Pequegnat (1970) to be a synonym of Trichopeltarion nobile 

A. Milne Edwards, to which all previous records are referred. 

Trichopeltarion A. Milne Edwards, 1880 

Trichopeltarion nobile A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 20) 

As Trachycarcinus spinulifer—Rathbun, 1930, p. 166, text-figs. 26-27, pis. 70-71. 

As Trichopeltarion no6i7e—Rathbun, 1930, p. 168, pi. 73; Pequegnat, 1970, p. 184, 

figs. 6-4, 6-5. 

Range: off northwest Florida and Mississippi; east coast of Mexico; Bay of 

Campeche; off St. Lucia. 

Depth: 274to752m (150to411fm). 

Habitat: mud bottoms. 

Remarks: Pequegnat (1970) synonymized the two species on the basis of 

fresh material collected by the R/V Alaminos in the Gulf of Mexico. The largest 

of 9 females collected was ovigerous, from 195 fm depth, in September. Listed by 

Chace (1956), as T. spinulifer, from off Mississippi. 

Family CANCRIDAE Latreille, 1803. 

Subfamily CANCRINAE Latreille, 1803 

Cancer Linnaeus, 1758 

Cancer borealis Stimpson, 1859 (Ann. Lye. Nat. Hist. New York 7: 50) 

Common Names: Jonah Crab; Northern Crab 

Hay & Shore, 1918, p. 434, pi. 35, fig. 2; Ratlibun, 1930, p. 182, text-figs. 30-31; 

Chace, 1940, p. 38; Williams, 1965, p. 175, fig. 156. 

Range: Nova Scotia to east coast of Florida; Bermuda; south coast of Florida; 

south of Dry Tortugas, in Florida Straits.



Habitat: immature and sub-adult stages occur intertidally among rocks and 

in shallower harbors and bays; larger specimens are found over a wide range of 

depths on sand, shell, gravel, mud, and ooze substrates. 

Remarks: Ovigerous females were listed from Florida in June (William^s, 

1965). This species has been collected in the western portions of the Florida 

Straits and can be considered to inhabit the Gulf of Mexico. Cancer irroratus 

Say, 1817 has been collected in the Florida Straits slightly west of 80° W, but is 

more typically restricted to the Atlantic coast and is not included in the Gulf 

fauna at the present time (based on pers. comm. of F. A. Chace). 

SECTION BRACHYRHYNCHA Borradaile, 1907 

(This group of crabs includes over 3000 species, world-wide. Older systematic 

literature traditionall}'^ divided the Brachyrhyncha into two subgroups, 

based on the general shape of the carapace: the Cyclometopa 

(round-fronted) or "cancroid" crabs and the Catametopa (square-fronted) 

or "grapsoid" crabs. The first group, as treated by Rathbun (1930), consisted 

of the families Portunidae, Xanthidae, Potamidae, Atelecyclidae, 

and Cancridae. These last two families are treated by some authors and 

in the present list within a separate section, the Cancridea, following the 

format of Glaessner (1969). Sakai (1965) groups the Goneplacidae with 

the other cancroid families in his key to the Brachyrhyncha, whereas 

Rathbun (1918) treated the Goneplacidae as a grapsoid family. This 

family has long been recognized as containing genera that link certain 

genera of the Xanthidae and Grapsidae, thus blurring many of the sharp 

distinctions between graspoid and cancroid characteristics. The position 

of the Palicidae is treated as "uncertain" by Glaessner (1969), but it 

is included here with the Xanthoidea as a matter of convenience, suggested 

by Chace (pers. comm.)). 

Superfamily PORTUNOIDEA Rafinesque, 1815. 

Family PORTUNIDAE Rafinesque, 1815 

Subfamily POLYBIINAE Ortmann, 1893 

(Ovalipes is listed as a genus of the Macropipinae in Glaessner (1969), 

but an addendum, p. R627, indicates the synonymy with Polybiinae.) 

Balhynecles Stimpson, 1871 

Balhynecies superba (Costa, 1853) (Fauna Regno Napoli, Addiz. Decapodi 

Brachyuri, p. 19) 

Milne Edwards & Bouvier, 1923, p. 311; Rathbun, 1930, p. 28, pis. 9-10; Chace, 

1940, p. 30; Monod, 1956, p. 183, figs. 210-212; Williams, McCIoskey & Gray, 1968, 

p. 50; Christiansen, 1969, p. 70, fig. 28, map 22.


Range: Massachusetts to south Florida; Florida Keys and Straits; west of Dry 

Tortugas and mid-eastern Gulf of Mexico; off Alabama and Mississippi; northeast 

of Yucatan; north and south coasts of Cuba; in the eastern Atlantic—Norway; 

north of Scotland; west coast of France; Cape Verde Islands; Mediterranean 

and Black Seas. 

Depth: 100 to 1435 m (55 to 769 fm). 

Habitat: on sand, gravel, shell and coral bottoms; rocky areas. 

Remarks: Most of the Gulf of Mexico records are from Chace (1956). Musick 

and McEachran (1972) collected this crab at depths of 159 to 274 m in Chesapeake 

Bight. Roberts (1969) studied larval development and epizoites of this 

species were described by Lewis (1976). 

Benlhochascon Alcock & Anderson, 1899 

Benthochascon schmiui Rathbun, 1931 (J. Washington Acad. Sci. 21: 125) 

Rathbun, 1931a, p. 125, pis. 1-2; Pequegnat, 1970, p. 187, fig. 6-6. 

Range: off Dry Tortugas; deep waters off Mississippi to Texas; off southern 

Gulf coast of Mexico; recently found off New England. 

Depth: 201 to 510 m (110 to 279 fm). 

Habitat: mud, silt-clay substrates. 

Remarks: This species was first reported from the Dry Tortugas by Rathbun 

(1931a), based on a specimen collected by W. L. Schmitt. Schmitt (1931) provided 

some note on this crab. Reported by Chace (1956) from the Gulf collections 

of the R/V Oregon. Pequegnat (1970) collected numerous specimens from 

various parts of the Gulf of Mexico and Wigley and Messersmith (1976) collected 

a single male from 252 m depth off southern New England. 

Ovalipes Rathbun, 1898 

(This genus has undergone a number of recent revisions, resulting in 

considerable confusion to non-specialists and complicating the comparison 

of field records in the non-systematic literature. Prior to revision by 

Williams (1962), Ovalipes ocellatus was considered to be represented 

in the Atlantic by the typical form and in the Gulf of Mexico by a subspecies, 

O. o. guadulpensis. Many of the earlier surveys and checklists 

reported O. ocellatus from the Gulf, but without specifying subspecies 

or authority. Felder (1973a) listed both forms for the northwestern Gulf, 

but with reservations, noting the nomenclatural confusion in the records. 

Tiirkay (1971) had discovered, in the meantime, that the original type 

of Saussure's guadulpensis belonged to the genus Macropipus and not 

Ovalipes and that it was named for a locality in the Azores rather than 

in the Caribbean. The next available name for the western Atlantic form 

thus became Ovalipes ocellatus floridanus, as used by Hay and Shore 

(1918) and recognized by Tiirkay (1971). Two forms of this crab were 

recognized by Williams (1962, 1965) and by Stephenson and Rees 

(1968), Form a from the Carolinian Province off the southeastern United 

States and Form b from the Gulf of Mexico. Williams (1976) finally


presented evidence that the two forms of O. o. floridanus should be 

regarded as separate species, but only one, O. floridanus, inhabits the 

Gulf of Mexico. See Williams (1976) for a nomenclatural history of these 

species.) 

Ovalipes floridanus Hay & Shore, 1918 (Bull. U.S. Bur. Fish. 35: 427) 

Common Names: Lady Crab; Sand Crab 

As O. ocellatus floridanus—Hay & Shore, 1918, p. 427, pi. 32, fig. 8; Turka3s 

1971, p. 139, fig. 3. 

As O. ocellatus guadulpensis—Rathbun, 1930, p. 23, pi. 4 (pai-t, the Pensacola, 

Alabama, and Gulf of Mexico specimens). 

As O. guadulpensis (Form 6;—Stephenson & Rees, 1968, p. 243, pis. 37D, 40F, 

41E, 42K, fig. IK. 

As O. guadulpensis—Williams, 1965, p. 161 (part, Gulf specimens only); Felder, 

1973a, p. 54, pl. 8, fig. 2. 

As O. /Zoi-iV/anus—Williams, 1976, p. 206, fig. la-d. 

Range: southwest Florida to south Texas. 

Depth: near surface and shallow water to 31 m (to 17 fm). 

Habitat: mainly sandy substrates; also on coral and broken shell bottoms. In 

coastal lagoons, channels, and ba}^s in southern and central Texas. 

Remarks: Williams (1976) describes and compares the two species of Ovalipes 

previously considered to be O. guadulpensis. The Carolinian form was named 

O. stephensoni, Form a of Stephenson and Rees (1968). The type locality for 

O. floridanus is Pensacola, Florida. Williams (1976) lists ovigerous females in 

February from Florida. Abele (1970) noted that juveniles were common on 

shallow grass flats in northwest Florida, while adults were more often found 

offshore on sandy bottoms. Regional lists include, as O. guadulpensis or 0. 

ocellatus guadulpensis, Florida (Wass, 1955, Abele, 1970; Menzel, 1971), Mississippi 

(Richmond, 1962; Franks et al., 1972; Christmas and Langley, 1973), 

Louisiana (Belire, 1950; Hildebrand, 1954), Texas (Gunter, 1950; Leacy, 1967), 

and the northwestern Gulf (Fotheringham and Brunenmeister, 1975). Caine 

(1974) provided a detailed description of feeding behavior, ecology, and burrowing 

activities. Abele (1970) commented on behavior of the animal in sand 

substrates. Notes on the related species are provided in Williams (1965). 

Ovalipes guadulpensis and Ovalipes ocellatus guadulpensis (Saussure, 1858). 

Both names are invalid and all Gulf of Mexico reports should be refeiTed to 

Ovalipes floridanus Hay and Shore, 1918. See Williams (1976) for a review of 

the nomenclatural history of this species. 

Subfamily PORTUNINAE Rafinesque, 1815 

Arenaeus Dana, 1851 

Arenaeus cribrarius (Lamarck, 1818) (His. Nat. Anim. sans Vert., vol 5, p. 

259) 

Common Name: Speckled Crab 

Hay & Shore, 1918, p. 434, pl. 34, fig. 3; Rathbun, 1930, p. 134, pl. 58, figs. 2-3,

Crabs of ihe Gulf of Mexico 75 

pis. 39-60; Rathbun, 1933, p. 50; Williams, 1963, p. 173, fig. 133; Felder, 1973a, 

p. 55, pi. 8, fig. 4. 

Range: Massachusetts to North Carolina; Bermuda; east coast of Florida; 

Florida Keys and Tirj Tortugas; west coast of Florida to Tabasco, Mexico; Jamaica; 

Puerto Rico; Dominica to St. Lucia; Belize to Colombia; Curagao; Ceara 



Habitat: in surf zone of sandy beaches; an adroit swimmer, it is seldom 

washed ashore and it is rarely found in estuaries and back lagoons. 

Remarks: Williams (1965) lists ovigerous females from Florida in August. 

Regional lists include Florida (Wass, 1955; Abele, 1970; Menzel, 1971), Mississippi 

(Richmond, 1962), off the Mississippi Delta (Chace, 1956), Louisiana 

(Behre, 1950; Hoese and Valentine, 1972), Texas (Gunter, 1950, Hildebrand, 

1954), and the northwestern Gulf of Mexico (Fotheringham and Brunenmeister, 

1975). Listed from Brazil by Coelho and Ramos (1972). 

Callinecles Stimpson, 1860 

(This commercially important genus has bL;en revised and reviewed 

by Williams (1974a), including details of reproductive morphology, 

notes on larAâl development, fossil records, a discussion of zoogeographic 

affinities, and keys for the identification of the 14 species known worldwide. 

In the Gulf region, as well as elsewhere, confusion has existed as 

to the identity of the lesser or smaller blue crab species, cited locally as 

Callinectes danae and C. ornatus. A prior revision by Williams (1966) 

established a new species, C. similis, for the Gulf populations and many 

of the east coast specimens, limiting the former two species to the southeastern 

Gulf and beyond. The present list includes eight species in the 

Gulf of Mexico, five of which occur only in the eastern or southern margins 

of the region. Only C. ralhhunae is endemic to the Gulf; C. sapidus 

and C. similis range widely along the U.S. Gulf and Atlantic coasts and 

beyond (see Figures 24-27 in Williams, 1974a).) 

Callinectes bocourti A. Milne Edwards, 1879 (Crust. Reg. Mex., p. 226) 

Rathbun, 1930, p. 128, text-figs. 15g, 16e, 17h, 18f, pi. 55; Rathbun, 1933, p. 49; 

Holthuis, 1959, p. 201, text-fig. 47, pi. 3, fig. 2; Chace & Hobbs, 1969, p. 127, 

text-figs. 35, 37a; Williams, 1974a, p. 767, figs. 12,18j, 20m, 22j, 27. 

Range: southeast Florida; Mississippi (rare); Jamaica; Hispaniola; Puerto 

Rico; Dominica; Virgin Islands to Barbados; Trinidad; Netherlands Antilles; 

Belize to Panama; Colombia to Santa Catarina, Brazil. 

Depth: shallow water, near shore and inshore waters. 

Habitat: in shallow, brackish waters of estuaries and river mouths. Tolerates 

salinities as low as 5 ppt; females usually move to saltier waters after mating. 

Williams (1974a) notes that this species is often associated with C. sapidus, but 

appears to be more tolerant than the latter to stagnant and polluted waters. 

Collected from a mud bottom in 3 m of water in Biloxi Bay, Mississippi (Perry, 

1973). See Chace and Hobbs (1969) for extensive habitat notes in Dominica.


Remarks: The first reports of this species in North America were from Florida 

(Provenzano, 1961), from Mississippi, the only Gulf records (Perry, 1973), and 

a second Florida finding (Gore and Grizzle, 1974). Norse (1972) noted habitat 

preferences in Jamaica. Listed from Brazil by Coelho (1971a) and Coelho and 

Ramos (1972). 

Callinectes danae Smith, 1869 (Trans. Connecticut Acad. Sci. 2: 7) 

Rathbun, 1930, p. 118 (part)), text-figs. 15d, 16d, 17b, 18d, pi. SI; Rathbun, 1933, 

p. 49; Chace, 1940, p. 33; Chace & Hobbs, 1969, p. 130, fig. 37b; Holthuis, 1959, 

p. 201; Williams, 1966, p. 86, figs. 2A-D, 4C-D; Jones, 1968, p. 187; Williams, 

1974a, p. 746, figs. 7,18e, 20e-f, 22e, 24. 

Range: Bermuda; southeast Florida; Dry Tortugas; north coast of Cuba; 

Caribbean coast of Yucatan; Jamaica; Hispaniola; Puerto Rico; St. Croix to 

Barbados; Trinidad; Netherlands Antilles; Belize to Panama; Colombia to Santa 

Catarina, Brazil. 


Habitat: from nearly fresh to full sea water, possibty in hypersaline lagoons; 

Occurs among mangroves and in mudd}'- estuaries in Brazil; Off beaches and in 

open water. 

Remarks: Williams (1974a) believes that earlier records of this species from 

Chile are erroneous. Records of this species along the U.S. Gulf coast should be 

referred to C. similis, as per the revision by Williams (1966, 1974a). Records 

from east Florida indicated this crab's presence on the ocean side of islands in 

Biscayne Bay and its absence from the Florida Keys (Park, 1969). Norse (1972) 

noted habitat preferences in Jamaica. Listed from Brazil b}^ Coelho and Ramos 

(1972). Morrison and Morrison (1952) studied hemolymph coagulation in 

Bermuda specimens. 

Callinectes exasperatus (Gerstaecker 1856) (Arch. f. Naturg. 22: 129) 

Rathbun, 1930, p. 130, text-figs. I5f, 16f, 17e, 18e, pi. 56; Conti-eras, 1930, p. 236, 

fig. 7; Rathbun, 1933, p. 49; Chace, 1940, p. 4-1.; Chace & Hobbs, 1969, p. 131, fig. 

37c; Felder, 1973a, p. 58, pi. 8, fig. 8; Williams, 1974a, p. 757, figs. 9, 18g, 20i, 

22g, 26. 

Range: Bermuda; Bahamas; southeast Florida; Florida Keys and Dr};- Tortugas; 

Vera Cûz to Yucatan, Mexico; north and south coasts of Cuba; Jamaica; 

Hispaniola; Puerto Rico; St. Croix to Grenadines; Trinidad; Netherlands 

Antilles; Caribbean coast of Yucatan to Panama; Venezuela and Isla de Margarita; 

Rio Grande do Norte to Santa Catarina, Brazil. 

Depth: shore to 7.5 m (to 4 fm). 

Habitat: estuarine and marine waters, possibl}^ fresh water; around river 

mouths and in mangrove areas. 

Remarks: Williams (1974a) comments on a lack of specimens from the 

Guianas and northern Brazil and that a locality record from Chile is erroneous. 

Leary (1967) listed this species from Texas on the basis of a single specimen 

from near Port Aransas, tentatively identified by Gordon Gunter, but not available 

for confirmation by Williams. Leary (1967) stated that the frontal teeth 

pattern was very similar to C. danae (= C. similis), thus the Gulf record for this


species is questionable. Felder (1973a) includes this species in his key for the 

northwestern Gulf on the basis of the same specimen. Beports of this crab in 

southern Florida are provided by Futch (1965) and Park (1969). Norse (1972) 

noted habitat preferences in Jamaica. Listed from Brazil by Coelho and Bamos 

(1972). 

Callinecles marginalus (A. Milne Edwards, 1861) (Arch. Mus. Hist. Nat., Paris 

10: 318) 

Rathbun, 1930, p. 123, figs. 15e, 16cl, IZd, 18c, pi. 53; Contreras, 1930, p. 235, fig. 6; 

Rathbun, 1933, p. 49; Chace & Hobbs, 1969, p. 131, fig. 37d; Felder, 1973a, p. 59, 

pi. 8, fig. 9; Williams, 1974a, p. 722, figs. 3, 18b (not 18a), 20a, 22b, 27. 

Bange: North Carolina; Bermuda; Bahamas; southeast Florida; Florida Keys 

and Dry Tortugas; southwest Florida; Louisiana (rare); Vera Cruz to Yucatan, 

Mexico; north and south coasts of Cuba; Jamaica; Hispaniola; Puerto Bico; St. 

Croix to Grenadines; Tobago and Trinidad; Netherlands Antilles; Yucatan to 

Panama; Colombia to Sao Paulo, Brazil; in eastern Atlantic—Senegal to Angola; 

Mauritania; Cape Verde Islands. 

Depth: intertidal to 15 m, rarely to 25 m (to 14 fm). 

Habitat: shallow water on sand and mud fiats; edges of mangrove swamps; 

grass flats, oyster bars, and rocky pools; often in brackish water, rarely in open 

marine water. 

Bemarks: The records from North Carolina and Louisiana are extralimital 

occurrences. Behre (1950) listed this crab as a common component of the Sargassum 

community in Louisiana waters, but Felder (1973a) commented that 

this species is by no means common and that Behre probably confused it with 

Portunus sayi. Williams (1974a) reports spawning records throughout the range 

of this crab, from December to July. Norse (1972) noted habitat preferences in 

Jamaica. Listed from Brazil by Coelho and Bamos (1972) and from West Africa 

by Forest and Guinot (1966). 

Callinecles ornalus Ordway, 1863 (J. Boston Soc. Nat. Hist. 7: 571) 

Rathbun, 1930, p. 114 (part), text-figs. 15b, 16a, 17a, 18b, pi. 50; Contreras, 1930, 

p. 232 (part), fig. 4; Rathbun, 1933, p. 48, fig. 40; Chace, 1940, p. 33; Chace & 

Hobbs, 1969, p. 132, fig. 37e; Holthuis, 1959, p. 200; Williams, 1965, p. 172 (part); 

Wilfiams, 1966, p. 84, figs. lA-B, 4A-B; Williams, 1974a, p. 739, figs. 6, I8d, 20d, 

22d, 25. 

Bange: Bermuda; Bahamas; North Carolina to south Florida; Florida Keys and 

Dry Tortugas; west Florida to Tampa Bay; Gulf coast of Yucatan; north and 

south coasts of Cuba; Jamaica; Hispaniola; Puerto Bico; St. Croix to Barbados; 

Trinidad; Netherlands Antilles; Caribbean coast of Yucatan to Belize; Colombia 



Habitat: sand, mud, shell substrates; off sponges; near river mouths and bays; 

in fresh waters, but more common in waters of moderate salinities (Williams, 

1974a). 

Bemarks: As with many portunids, juveniles of this species may be difficult 

to distinguish from others, especially those of C. danae and C. similis. Becords


of C. ornatus in New Jersey, Louisiana, and Texas (Rathbun, 1930; Leary, 1967) 

should be referred to C. .similis, based on the revision and restriction of this species 

by Williams (1966). Brues (1927) discussed the ecology of this crab and Norse 

(1972) noted habitat preferences for the species in Jamaica. Listed from Brazil 

byCoelho (1971a) and Coelho and Ramos (1972). 

CalUnectes ralhbunae Contreras, 1930 (An. Inst. Biol. Univ. Nac. Auton., Mex. 

1: 238) 

Contreras, 1930, p. 238, figs. 9-10; Felder, 1973a, p. 38, pi. 8, fig. 10; Williams, 

1974a, p. 772, figs. 13, 19a, 20n, 22k, 27. 

Range: eastern coast of Mexico, from mouth of Rio Grande to southern Vera 

Cruz. 

Depth: verj^ shallow waters. 

Habitat: estuarine waters of ditches, lagoons, and river mouths; in shallow 

coastal bays and a broad range of salinities. 

Remarks: This species is apparently localized to the central Mexican coast, 

with occasional specimens as far north as the Rio Grande. Listed by Leary 

(1967) for Texas, based on a single male specimen found at the mouth of the 

Rio Grande River (H. H. Hildebrand, collector). 

CalUnectes sapidus Rathbun, 1896 (Proc. U.S. Nat. Mus. 18: 352) 

Common Names: Blue Crab; Common Edible Crab 

Ha}-- & Shore, 1918, p. 432, pi. 35, fig. 1; Rathbun, 1930, p. 99, text-figs. ISa, 16c, 

17c, 18a, 19, pi. 47; Holthuis, 1961, p. SO, pi. 1, fig. 2, pi. 2, fig, 2; Williams, 196S, 

p. 168, fig. 151; Chace & Hohhs, 1969, p. 133, figs. 36, 37f; Christiansen, 1969, 

p. 72, fig. 29; Felder, 1973a, p. 55, pi. 8, fig. 7; Williams, 1974a, p. 778, figs. 1, 16, 

17, 19d,21,23b-c, 26. 

As C. sapidus acutidens—Rathbun, 1930, p. Ill, text-fig. I5c, pi. 48; Contreras, 

1930, p. 228, fig. 1; Rathbun, 1933, p. 48. 

Range: Bermuda; Bahamas; Nova Scotia to south Florida; Florida Keys and 

Dry Tortugas; south Florida to Yucatan, along entire Gulf coast of the United 

States and Mexico; north and south coasts of Cuba; Jamaica; Haiti; Puerto Rico; 

Virgin Islands; Dominica; Trinidad and Tobago; Netherlands Antilles; Yucatan 

to Guatamala; Nicaragua to Panama; Venezuela; central Brazil to northern 

Argentina; in the eastern Atlantic—Denmark; Netherlands and adjacent North 

Sea; southwest France; Golfo di Genova; northern Adriatic Sea; Aegean Sea; 

western part of Black Sea; eastern Mediterranean Sea; Japan (Sakai, 1976a). 

Williams (1974a) reviews the distribution of this species and cites pertinent 

literature on the ship transport of crab larvae and their introduction into the 

Old World during recent times. 

Depth: shore (intertidal) to 90 m (49 fm), more commonly to 35 m (19 fm). 

Habitat: occurs in a wide range of salinities, from freshwater to hypersaline; 

along coasts in shallow water off ocean beaches (mainly females), in bays, 

estuaries, lagoons, ponds, ditches; well upstream in larger rivers (mainly males). 

Copeland and Bechtel (1974) listed an optimum salinity range of 0 to 27 ppt 

and optimum temperature range of 10 to 35°C, but extremes of 117 ppt (Hildebrand, 

1957) and 45°C. (personal observations) under natural conditions are


tolerated for short periods. Mahood et al., (1970) showed that temperature and 

salinity tolerances were interdependent. Blue crabs are also able to tolerate low 

oxygen conditions and they are occasionally found in very polluted and anoxic 

waters. Along the Texas coast, blue crabs in shallow ponds on sand flats are 

exposed to temperatures in excess of 40°C. during midday. These crabs have 

been observed to leave the water for Salicornia patches around the pools, where 

they rely on aerial respiration for 2 to 4 hours in the cooler (30 to 35°C.), 

humidity-saturated environment (personal observations). Blue crabs can travel 

some distance overland at night and during wet periods, again relying on aerial 

respiration. 

Remarks: The vast literature, much of it non-technical or of a commercial 

nature, precludes anything approaching a complete bibliography on this species. 

The original type for this species was obtained from the eastern coast of the 

United States, a variant of the form that is more typical throughout its range. 

The "typical" form, most often encountered from Florida southward, was considered 

a subspecies by Rathbun (1930) and many others, C. sapidus acutidens, 

so-named because of the surface features and pronounced spines, teeth, and 

prominent ridges. Because the type-based form from farther north was the basis 

for comparison, confusion existed for some years over the designation of a type 

to replace the original, which had been lost. Williams (1974a) discusses these 

variations and the nomenclatural historj^ of this crab, agreeing with Chace and 

Hobbs (1969) that a variety of extreme forms exist and that they could be considered 

separate species if they were considered in isolation from each other. 

However, these forms are intergraded and form a continuum, without morphological, 

bathymetrical, or geographical discontinuity, thus all the forms of Callinectes 

sapidus are considered, at present, to represent a single species in the 

process of local speciation which is still morphologically incomplete. 

Recent literature compilations on this species were provided by Cronin et al. 

(1957) and by Tagatz and Hall (1971). Gulf regional Hsts include Florida 

(Wass, 1955; Tabb and Manning, 1961; Dragovich and Kelly, 1964; Rouse, 

1970; Menzel, 1971; Lyons et al, 1971), Mississippi (Richmond, 1962; Franks 

et al., 1972; Christmas and Langley, 1973), Louisiana (Behre, 1950; Darnell, 

1959; Hoese and Valentine, 1972), Texas (Gunter, 1950; Lledgpeth, 1950; Hoese, 

1960; Copeland, 1965; Leary, 1967; More, 1969; Copeland and Bechtel, 1974), 

Mexico (Contreras, 1930; Hildebrand, 1957), and north coast of Cuba (Chace, 

1940). 

Hay (1905) and Churchill (1919) provided comprehensive life history studies. 

Other information on ecology includes: habitat relationships in Texas (Hedgpeth, 

1953; Simmons, 1957; Breuer, 1962; Fotheringham and Brunenmeister, 1975; 

Trent, Pullen and Proctor, 1976), megalops ecology in Maryland (Cargo, 1960), 

habitat notes in Mississippi (Franks et al., 1972; Christmas and Langley, 1973), 

effects of environmental variables on juveniles (Holland, Aldrich and Strawn, 

1971), larval ecology in Virginia (Sandifer, 1973), seasonal population changes 

in Chesapeake Bight (Musick and McEachran, 1972), field observations of freshwater 

populations (Gunter, 1938), temperature and thermal tolerance (Tagatz, 

1969a), and habitats in Jamaica (Norse, 1972).


Behavioral studies include: agonistic displays (Jachowski, 1974), larval 

shadow responses (Forward, 1977), analyses of swimming behavior (Spirito, 

1972), predation on oyster spats (Lunz, 1947), quahogs (Carriker, 1951; Haven 

and Andrews, 1957), and gastropods (Hamilton, 1976), predator avoidance 

(Gunter, 1954), sex recognition (Teytaud, 1971), and climbing behavior on 

vegetation (Abbott, 1967). 

Developmental studies include: lar\âl rearing in the laboratory (Costlow 

and Bookhout, 1959, 1960a; Bust and Carlson, 1960; Davis, 1965), feeding of 

larvae (Sulkin and Epifanio, 1975), variability in morphology of larvae (Tj'ler 

and Cargo, 1963; Costlow, 1965), hormonal control of development (Costlow, 

1963), zoeal growth and survival (Sandoz and Rogers, 1944), effects of pesticides 

on larval development (Bookhout and Costlow, 1976), salinity-temperature effects 

on larvae (Costlow, 1967; Rosenberg and Costlow, 1976), growth of juveniles 

(Gray and Newcombe, 1938; Tagatz, 1969b; Holland, Aldrich and Strawn, 

1971), and development of cheliped laterality (Hamilton, Nishimoto and Halusky,1976). 

Physiological studies include: effects of salinity on growth at terminal molt 

(Haefner and Shuster, 1964), ionic and osmotic regulation ( Gifford, 1962a; Tan 

and van Engel, 1966; Mantel, 1967; Copeland and Fitzjarrell, 1968; Ballard and 

Abbott, 1969; Tagatz, 1971; Gerard and Gilles, 1972; Mangum and Amende, 

1972; Lynch et al., 1973; Towles et al., 1976; Mangum et al., 1976; Mangum 

and Towle, 1977), respiration and respiratory pigments (Ayres, 1938; Gray, 

1957; Horn and Kerr, 1963, 1969; Bonaventura et al., 1974; Engel and Eggert, 

1974; Mangum and Weiland, 1975; Weiland and Mangum, 1975; Lewis and 

Haefner, 1976; Laird and Haefner, 1976), hemolymph volume (Gleeson and 

Zubkoff, 1977), responses of megalops to pressure (Naylor and Isaac, 1973), 

gross anatomy and fine structure (Cochran, 1935; Cronin, 1947; Pyle and 

Cronin, 1950; Jahromi and Govind, 1976), ultrastructure of sperm (G. C. Brown, 

1966), partial albinism (Sims and Joyce, 1966), neuroendocrinology (Costlow, 

1963; Payen et al., 1971; Andrews, Copeland and Fingerman, 1971; Skinner and 

Graham, 1972; Andrews, 1973; Ludolph, Paganalli and Mote, 1973), biochemical 

adaptations (Vernberg and Vernberg, 1968; Robert and Gray, 1972), neurobiology 

(Mendelson, 1963, 1966; Skobe and Nunnemacher, 1970; Hazlett, 1971; 

White and Spirito, 1973; Mtiynard and Dando, 1974; Reingold, 1975; Steinacker, 

1975), fluoride metabolism (Moore, 1971), effects of DDT (Sheridan, 1975), 

homolymph coagulation (Morrison and Morrison, 1952), and temperature effects 

on growth and metabolism (Tagatz, 1969a; Leffler, 1972). 

Studies on parasitology and pathology include: parasitic barnacles (Himnes, 

1941a; Reinhard, 1950a, 1950b; Adkins, 1972a), external barnacles (Williams 

and Porter, 1964), effects of nemerteans on reproduction (Hopkins, 1947), 

infections of nemerteans in gills (Pearse, 1949), amoebic infections (Sprague and 

Beckett, 1966; Sawyer, 1969; Sprague, Beckett and Sawyer, 1969; Pauley, Newman 

and Gould, 1975), effects of amoeba infections on hemocjê values (Sawyer, 

Cox and Higginbottom, 1970), gas-bubble disease (Johnson, 1976), microsporid 

infections (Sprague, 1965, 1966), bacterial infections (Krantz, Colwell and


Lovelace, 1969; Cook and Lofton, 1973), fungal infections (Couch, 1942; Newcombe 

and Rogers, 1947; Rogers-Talbert, 1948; Bland eZ al., 1976) andtrematode 

infections (Overstz'eet and Perry, 1972). 

Commercial fishery and mariculture reports include: Texas fishery (Daugherty, 

1952a; More, 1969), Louisiana fishery (Jaworski, 1970; Adkins, 1972b), 

Mississippi fishery (Perret, 1967; Perry, 1975), Chesapeake Bay fishery (Truitt, 

1939; van Engel, 1958; Miller, Sulkin and Lippson, 1975), methods for handling 

soft crabs (Haefner and Garten, 1974), migration in bays and estauries (Fiedler, 

1930), and mariculture (Rust and Carlson, 1960; Sulkin and Epifanio, 1975). 

Tlie preceding does not include unpublished reports of state Fish and Game 

Commissions, which contain statistics on commercial landings and are further 

sim:imarized by publications of the U.S. Fish and Wildlife Service. 

Callinectes sirnilis Williams, 1966 (Tulane Stud. Zool. 13: 87) 

Common Name: Lesser Blue Crab 

As C danae—Raihhun, 1930, p. 118 (part). 

As C. onialus—Hay & Shore, 1918, p. 433, pi. 34, fig. 2; Conlreras, 1930, p. 231 

(part), ? fig. 4; Ratlibun, 1930, p. 114 (part); Williams, 1965, p. 172, fig. 152. 

As C. si"mi7i.s—Williams, 1966, p. 87, figs. 3, 4E^F; Felder, 1973a, p. 58, pi. 8, fig. 1; 

Williams, 1974a, p. 731, figs. 4, 18a (not 18b), 20c, 22a, 24. 

Range: Delaware Bay to southern Florida; Florida Keys; northwest Florida 

to Campeche, Yucatan, including Gulf coast of United States and Mexico. 


Habitat: in ocean waters, near shore on sand and mud bottoms, often in association 

with populations of C. sapidus; in bays and estuaries, rarely below 

salinities of 15 ppt (Hoese, 1960 lists a range of 4.7 to 45 ppt), most common 

at 25 to 37 ppt; temperature ranges of 13 to 29°C. in the Gulf, slightly higher 

in Texas bays. 

Remarks: With the exception of southwest Florida, all Gulf coast records of 

C. danae and C. ornatus for the United States and Mexico should be referred to 

C. sirnilis. Regional lists, including records under danae and ornatus, include 

Florida (Wass, 1955; Tagatz, 1967; Abele, 1970), Mississippi (Richmond, 1962 

Franks et al., 1972; Christmas and Langley, 1973), Louisiana (Behre, 1950 

Dawson, 1966), Texas (Gunter, 1950; Daugherty, 1952b; Hildebrand, 1954 

Simmons, 1957; Hoese, 1960; Breuer, 1962; Leary, 1967; Hoese ei al, 1968), 

and Campeche (Hildebrand, 1955). Nocturnal swimming at the surface was 

noted by Franks e:« «Z. (1972). 

Cronius Stimpson, 1860 

Cronius ruber (Lamarck, 1818) (Hist. Nat. Anim. sans Vert., vol. 5, p. 260) 

Rathbun, 1930, p. 139, pis. 62-63; Rathbun, 1933, p. 51; Gartli, 1965a, p. 15; 

Williams, 1965, p. 174, fig. 154; Felder, 1973a, p. 55, pi. 8, fig. 3. 

Range: South Carolina to south Florida; Dry Tortugas; off Texas; off Campeche, 

Mexico; Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin Islands to 

Dominica; Caribbean coast of Panama; Amapa to Santa Catarina, Brazil; in


eastern Atlantic—from Cape Verde Island and Senegal to Angola; in eastern 

Pacific—from Lower California (Mexico) to Peru; Galapagos Islands; Clipperton 

Island. 


Habitat: sandy bottoms; from areas of reefs, rocks, and shell rubble. 

Remarks: Listed from Texas by Leary (1967) and confirmed by Felder 

(1973a). Garth and Stephenson (1966) commented on Pacific distribution and 

African collections were examined by Forest and Guinot (1966). Listed from 

Brazil to a depth of 105 m bj' Coelho and Ramos (1972). 

Cronius tumidulus (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 149) 

Rathbun, 1930, p. 142, pi. 64; Rathbmi, 1933, p. 51, fig. 43. 

Range: Bermuda; Bahamas; Florida Keys and Dry Tortugas; west coast of 

Florida; north and south coasts of Cuba; Jamaica; Puerto Rico; Virgin Islands; 

Netherlands Antilles; Old Providence Island (Carib.); Ceara to Bahia, Brazil. 

Depth: 5 to 73 m (3 to 40 fm). 

Habitat: coral, sand, and rock bottoms; grass, seaweed, and Sargassum. 

Remarks: Listed from Brazil bj^ Coelho and Ramos, 1972. 

Lupella Rathbun, 1897 

Lupella forceps (Fabricius, 1793) (Entom. Syst. omend. auct., vol. 2, p. 449) 

Rathbun, 1930, p. 133, pi, 57; Rathbun, 1933, p. 50, fig. 41. 

Range: north coast of Cuba; Jamaica; Haiti; Puerto Rico; St. Thomas, Virgin 

Islands; Martinique. 

Depth: 13 to 15 m (7 to 8.5 fm). 


Portunus Weber, 1795 

(This genus has traditionally been divided into subgenera, based on 

morphological critera. Modem studies of Portunus (Stephenson and Rees, 

1967; Stephenson, Williams, and Lance, 1968) have raised serious doubts 

about the subgeneric relationships and they suggest that further study 

will be necessary to define the complex phylogenetic affinities of this 

group, including the closely related genera, Arenaeus and Callinectes. 

As in Williams (1965), subgeneric catagories are deleted here.) 

Portunus anceps (Saussure, 1858) (Mem. Soc. Phys. Hist. Nat. Geneve 14: 

434) 


1933, p. 46; Williams, 1965, p. 163, fig. 145. 

Range: North Carolina; Bennuda; Bahamas; Florida Keys; north and south 

coasts of Cuba; Caribbean coast of Yucatan; Jamaica; Puerto Rico; St. Thomas, 

Virgin Islands to Guadeloupe; Panama to Bahia, Brazil. 

Depth: surface to 103 m (to 56 fm).


Habitat: mainly on sandy bottoms, often with weeds or grass; also on mud, 

shell, and stone substrates; on coral reefs; buries in sand; in shallow waters, 

brackish ponds. 


in October and from Cuba in June. Listed from Brazil by Coelho (1971a) and 

by Coelho and Ramos (1972). 

Portunus binoculus Holthuis, 1969 (Bull. Mar. Sci. 19: 409) 

Holthuis, 1969, p. 409, fig. 1. 

Range: Bahamas; Florida Straits; north coast of Cuba; east of Yucatan, in 

Caribbean Sea; off Caribbean coasts of Panama and Colombia. 

Depth: 74 to 291 m (40 to 159 fm). possibly to a range of 63 to 467 m (34 to 

255 fm). 

Remarks: Holthuis (1969) notes that some of the specimens of P. spinicarpus 

in Rathbun (1930) may be P. binoculus and these may include a record from 

Tortugas at 37 m (20 fm). 

Portunus depressifrons (Stimpson, 1859) (Ann. Lye. Nat. Hist. New York 7: 

58) 


1933, p. 47; Williams, 1965, p. 166, fig. 149. 

Range: North Carolina; Bermuda; south Florida; Florida Keys and Dry Tortugas; 

Bahamas; Gulf of Campeche, off Yucatan; north coast of Cuba; Culebra; 

St. Thomas, Virgin Islands; Aruba, Netherlands Antilles; Old Providence Island 

(Carib.). 

Depth: surface to 29 m (tol6fm). 

Habitat: shallow water coves and inlets with sandy bottoms; on coral, shell, 

and grass-covered sand substrates. 

Remarks: Listed from northwest Florida by Wass (1955), Abele (1970), and 

Menzel (1971). Williams (1965) notes the lack of recent specimens from the 

Florida Keys. Ovigerous females have been reported from Florida in June and 

August, the Caribbean in August (Rathbtxn, 1930), and from Campeche in 

August (Williams, 1965). Chace (1956) recorded this species off the north coast 

of Yucatan. Abele (1970) provided detailed notes on the color of a live female 

and he listed it as common in St. Andrews Bay, Florida. 

Portunus floridanus Rathbun, 1930 (Bull. U.S. Nat. Mus. 152: 82) 

Rathbun, 1930, p. 82, pi. 40. 

Range: Key West, Florida. 

Depth: 82m (45 fm). 

Habitat: coral reefs. 

Portunus gibbesii (Stimpson, 1859) (Ann. Lye. Nat. Hist. New York 7: 57) 

Hay & Shore, 1918, p. 428, pi. 33, fig. 1; Rathbun, 1930, p. 49, pis. 16-17; Williams, 

1965, p. 164, fig. 146; Felder, 1973a, p. 60, pi. 8, fig. 16. 

Range: Massachusetts to south Florida; Florida Keys and Straits; Dry Tor-


tugas; west coast of Florida to south Texas; Campeche, Mexico; Venezuela; 

Surinam. 

Depth: surface to 88 m (to 48 fm), rarely deeper. 

Habitat: mud, sand, and broken shell bottoms; usually in deeper off-shore 

waters of Gulf coast and in deeper parts of near-shore marine passes, inlets, and 

bays. 

Remarks: Regional lists include Florida (Wass, 1955; Tabb and Manning, 

1961; Rouse, 1970; Abele, 1970; Menzel, 1971), Mississippi (Richmond, 1968 

Franks et al., 1972; Christmas and Langley, 1973), Louisiana (Dawson, 1966 

Hoese and Valentine, 1972), Texas (Gunter, 1950; Hildebrand, 1954; Parker, 

1959; Leary, 1967), and from the Gulf (Chace, 1956; Fotheringham and 

Brunenmeister, 1975). Hildebrand (1954) reported this crab abundant at 6 to 

10 fm at Campeche. Tabb and Manning (1961) found this crab feeding on concentrations 

of cyprinodont fishes in Coot Bay, Florida. Musick and McEachran 

(1972) listed it from Chesapeake Bight at depths of 18 to 49 m. Felder (1973a) 

notes that this species is often found in association with P. spinimanus. Rathbun 

(1930) reported ovigerous females from Florida in April; Williams (1965) 

cites other ovigerous females from North Carolina to Surinam, from February 

to June. Gray (1957) measured the gill area of this crab and compared it with 

habitat preferences. 

Porlunus ordwayi (Stimpson, 1860) (Ann. Lye. Nat. Hist. New York 7: 224) 


1933, p. 46; Chace, 1940, p. 31; Williams, 1965, p. 166, fig. 148. 

Range: Massachusetts; North Carolina; Bermuda; Bahamas; southeast Florida; 

Florida Keys and Dry Tortugas; west and northwest coasts of Florida; north 

and south coasts of Cuba; Gulf and Caribbean coasts of Yucatan; Jamaica; Puerto 

Rico; St. Thomas, Virgin Islands to Dominica; Old Providence Island (Carib.); 

Para and Fernando de Noronha to Bahia, Brazil. 

Depth: surface to 106 m (58 fm), rarely deeper. 

Habitat: sand, gravel, broken shell, and coral substrates. 

Remarks: Rathbun (1930) listed an ovigerous female from Florida in March. 

Chace (1956) recorded this species from off northwest Florida. Listed from 

Brazil by Coelho (1971a), Coelho and Ramos (1972), and Fausto Filho (1974). 

Porlunus sayi (Gibbes, 1850) (Proc. Amer. Assoc. Adv. Sci., 3rd Meeting, 7: 

178) 

Common Name: Sargassum Crab 

Hay & Shore, 1918, p. 428, pi. 33, fig. 2; Rathbun, 1930, p. 37, text-figs. 6-7, pi. 14; 

Rathbun, 1933, p. 46, fig. 39; Chace, 1940, p. 31; Williams, 1965, p. 163, fig. 144; 


Range: Nova Scotia to south Florida; Bermuda; Bahamas; Florida Keys and 

Dry Tortugas; west coast of Florida to south Texas; north and south coasts of 

Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin Islands; Trinidad; Guiana; 

Brazil; in eastern Atlantic Ocean; off Kerguelon Island in the Indian Ocean. 

Depth: pelagic, at surface.


Habitat: normally among Sargassuni, floating on surface; also on other flotsam; 

occasionally swimming freely. 

Remarks: Larval stages from Bermuda were described by Lebour (1944) and 

Coventry (1944) described Caribbean collections. Williams (1965) listed ovigerous 

females in the northeast Gulf and in the West Indies from February to 

August and from off Massachusetts in September. Regional lists include Florida 

(Abele, 1970), Mississippi (Franks et al, 1972), Louisiana (Behre, 1950), Texas 

(Leary, 1967), and the Gulf (Chace, 1956; Fotheringham and Brunenmeister, 

1975). Autotomy and regeneration of chelae was studied by Zeleny (1908); 

Hartnoll (1971) described swimming behavior. 

Porlunus sehae (H. Milne Edwards, 1834) (Hist. Nat. Crust., vol. 1, p. 455) 

Rathbun, 1930, p. 79, pis. 34-35; Rathbun, 1933, p. 46. 

Range: Bermuda; Florida Keys and Straits; Dry Tortugas; south coast of 

Cuba; Jamaica; Puerto Rico; St. Tliomas, Virgin Islands; Dominica; Netherlands 

Antilles. 

Depth: 4tol8m (2tolOfm). 

Habitat: sand, rocky, and grass-covered sandy bottoms. 

Porlunus spinicarpus (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 148) 

Hay & Shore, 1918, p. 429, pi. 33, fig. 3; Rathbun, 1930, p. 92„ pi. 45; Rathbun, 

1933, p. 47; Chace, 1940, p. 32; Williams, 1965, p. 167, fig. 150; Holthuis, 1969, 

p. 415, fig. 1; Felder, 1973a, p. 60, pi. 8, figs, 13-14. 

Range: North and South Carolina; southeast Horida; Florida Ke5rs and Dry 

Tortugas; west coast of Florida to south Texas; north and south coasts of Cuba; 

Puerto Rico; Trinidad; Caribbean coast of Colombia; Guianas to Santa Catarina, 

Brazil. 

Depth: 9 to 550 m (5 to 300 fm). 

Habitat: on sand, gravel, coral, broken shell, and mud substrates. 

Remarks: Larval development was described by Bookhout and Costlow (1974). 

Williams (1965) listed ovigerous females from Texas in November. Rathbun 

(1930) noted that despite large numbers of this crab in the Florida Keys, fish 

stomachs do not contain this species, which she attributes to the Crab's extendable 

carpal spines. Pearse (1932a) reported the presence of the barnacle, Dichelastis 

sinvala^ on crabs from North Carolina. Holthuis (1969) expanded the 

previously known depth range to 550 m and he reported ovigerous females from 

the Caribbean from January to September, with some females carrying sacculinid 

barnacles. Musick and McEachran (1972) collected this crab from a depth of 

101 m in Chesapeake Bight. Regional lists included Florida (Hulings, 1961; 

Abele, 1970), Mississippi (Richmond, 1962; 1962; FrankseZo/., 1972),Louisiana 

(Dawson, 1966), and Texas (Hildebrand, 1954; Chace, 1956; Lear)^ 1967). 

Listed from Brazil by Rodrigues da Costa (1968a), Coelho (1971a) and Coelho 


Porlunus spinimanus Latreille, 1819 (Nouv. Diet. Hist. Nat., ed. 2, vol. 28, p. 

47) 

Hay & Shore, 1918, p. 429, pi. 33, fig. 4; Rathbun, 1930, p. 62, text-fig. 10, pis.


26-28; Ratlibun, 1933, p. 46; Chace, 1940, p. 31; Williams, 1965, p. 165, fig. 147; 


Range: New Jersey to south Florida; Bermuda; Bahamas; Florida Keys and 

Dry Tortugas; west Florida to south Texas; Campeche, off Mexico; south coast 

of Cuba; Jamaica; Puerto Rico; Hispaniola; Trinidad; Aruba, Netherlands 

Antilles; Bahia to Santa Catarina, Brazil. 

Depth: surface to 91 m (50 fm). 

Habitat: waters of inlets, canals, and harbors; on sand, gravel, broken shell, 

and mud bottoms; occasionally on Sargassum. 

Remarks: Lebour (1950) raised and described larvae collected at Bermuda. 

Holthuis (1959) listed ovigerous females from Surinam in May, August, and 

September; Williams (1965) summarized other records of ovigerous females. 

Regional lists include Florida (Wass, 1955; Tabb and Manning, 1961; Abele, 

1970; Menzel, 1971), Mississippi (Richmond, 1962), Louisiana (Behre, 1950), 

Texas (Hildebrand, 1954; Parker, 1959; Lear>s 1967), and Campeche (Hildebrand, 

1954). This crab was abundant at Campeche at depths of 6 to 10 fm, often 

in association with P. gibbesii. Reported from the Gulf by Chace (1956) and 

described generally by Fotheringham and Brunenmeister (1975). Musick and 

McEachran (1972) found this crab at 49 m depth in Chesapeake Bight. Gray 

(1957) measured gill area. Lobo de Mesquita (1972) reported biometrical data 

on Brazilian specimens; listed from Brazil by Coelho and Ramos (1972). 

Porlunus veiUralis (A. Milne Edwards, 1879) (Crust. Reg. Mex., p. 215) 

Rathbun, 1930, p. 43, pi. 13, figs. 1-2; Rathbun, 1933, p. 46; Chace, 1940, p. 31; 


Range: Georgia to east coast of Florida; Dry Tortugas; ? Texas; north and 

west coasts of Cuba; Jamaica; Puerto Rico; St. Thomas, "Virgin Islands; Barbados; 

Rio Grande do Norte to Rio de Janeiro, Brazil. 

Depth: shallow water, near shore, to 25 m (14 fm). 

Habitat: sandy beaches; tide pools; on surface of open waters. 

Remarks: Felder (1973a) questioned repoi-ts by Parker (1959) and Trott (unpublished) 

of this crab's occurrence in Texas; these records have not been verified 

by later collections. Rathbun (1930) reported ovigerous females from Dry 

Tortugas in August. Listed from Brazil hy Coelho and Ramos (1972). 

Porlunus vocans (A. Milne Edwards, 1878) (Bull. Soc. Philom. Paris, ser. 7, 

vol. 2, p. 225) 

Rathbun, 1930, p. 60, text-figs. 8-9, pi. 25. 

Range; north coast of Cuba; between Jamaica and Haiti; Ascension Island, 

in the South Atlantic Ocean. 

Depth: 37 to 309 m (20 to 169 fm). 


Superfamily XANTHOIDEA Dana, 1851 

Family POTAMIDAE Ortmann, 1896

The Freshwater Crabs 


The familial name, based on the genus Potamon Savigny, 1816, was corrected 

from the original Potamonidae of Ortmann by an ICZN decision (Opinion 712, 

p. 342, in 1964). A revision of the freshwater crabs by Bott (1955b) split this 

family into two: the Pseudothelphusidae and the Trichodactylidae. Various 

other schemes for classification have been proposed (Pretzmann, 1965; Bott, 

1968; Smalley, 1970). Gulf region species are confined to northern Cuba and 

southern Mexico, but insufficient material and information is available for most 

species to yield accurate ranges or to present a satisfactory organization of taxonomic 

relationships. Because these crabs are inhabitants of rivers, lakes, and 

caves, species ranges tend to be restricted to small areas; several are known only 

from a single type specimen. Species occurring in the center of a land mass, 

such as Potamocarcinus (Typhlopseudothelphusa) mocinoi in caves near Comitan 

(Chiapas) Mexico (Rioja, 1952) could be assigned to a coastal region based 

on watershed drainage patterns. The Cuban species listed by Chace and Hobbs 

(1969) are not localized to either the Gulf or the Caribbean side of the island 

in the records cited. For these reasons, these crabs are not included in the present 

work. Further references, including some older systematic papers, can be found 

in the papers cited above. There exists a clear need for more extensive work on 

this group of brachyurans. 

Family XANTHIDAE Dana, 1851 

This is a large family of crabs (about 1000 species and more than 130 genera) 

that has traditionally posed a number of taxonomic problems. Many of the 

species are small in size and appear morphologicallj^ similar. Individual variability 

and the large number of closely related species has often made definitive 

identification difficult, so that earlier collection records must be used cautiously. 

Rathbun (1930) did not subdivide her account of the xanthids into subfamilies. 

Guinot (1971) offers a number of systematic revisions and she comments 

at some length on affinities, but she also avoids listing the 51 genera that she 

treats under subfamilies. The same procedure is followed here, by arrangement 

of the 33 Gulf genera in alphabetical order, without regard for proposed affinity 

within the family. The xanthids are currently being revised by various workers 

around the world, so that a better organized and more accurate representation 

of this family should be forthcoming. 

Aclaea de Haan, 1833 

Aclaea acantha (H. Milne Edwards, 1834) (Hist. Nat. Crust., vol. 1, p. 379) 

Rathbun, 1930, p. 261, pi. 105, fig. 5, pi. 106, figs. 1-2; Rathbun, 1933, p. 57. 

Range: Bahamas; Florida Keys and Dry Tortugas; northwest coast of Cuba; 

Jamaica; Haiti; Puerto Rico; Guadeloupe; St. Bartholomew; Fernando de Noronha, 

Brazil. 

Depth: surface to 22 m (tol2fm).


Habitat: sand, shell, coral bottoms; from coral reefs; off mud and grassy bottoms. 

Remarks: Habitat and color of specimens from Brazil were described by 

FaustoFilho (1974). 

Actaea bifrons Rathbun, 1898 (Bull. Lab. Nat. Hist. Stats Univ. Iowa 4: 262) 

Rathbun, 1930, p. 255, text-fig. 41, pi. 104, figs. 3-6; Rathbun, 1933, p. 56, fig. 48. 

Range: Key West, Florida; Puerto Rico; Virgin Islands; St. Bartholomew; 

Barbados; Curasao; Colon, Panama. 


Habitat: in coral, Porites fareata; on shoal banks, coral bottoms, sponge areas. 

Aciaea /Wmeri Rathbun, 1894 (Proc. U.S. Nat. Mus. \7: 85) 


Range: Bahamas; east coast of Florida; Florida Keys; north coast of Cuba; 

Plaiti; Virgin Islands; Curagao. 


Habitat: from sponges and among coral {Porites furcata). 

Aciaea rufopunctala nodosa Stimpson, 1860. 

Transferred to a new genus, Paractaea, by Guinot (1969b). See Paractaea 

rufopunctata nodosa (Stimpson, 1860). 

Actaea setigera (H. Milne Edwards, 1834). 

Transferred to a new genus, Platyactaea, by Guinot (1967b). See Platyactaea 

setigera (H. Milne Edwards, 1834). 

Carpilius Leach, 1823 

Carpilius corallinus (Herbst, 1783) (Natur. Krabben u. Krebse, vol. 1, p. 133) 

Common Names: Queen Crab; Coral Crab 

Rathbun, 1930, p. 240, pis. 97-99; Rathbun, 1933, p. 53; Chace, 1940, p. 33; Guinot, 

1968b, p. 157, fig. 9; Guinot, 1968c, p. 321, figs. 10-11; Pequegnat & Ray, 1974, 

p. 237, figs. 13-15. 

Range: Bermuda; Bahamas; West Flower Garden Bank, off Texas; north coast 

of Cuba; Jamaica; Puerto Rico; Virgin Islands; Guadeloupe; Dominica; Curasao; 

Old Providence Island (Carib.); Pemambuco and Ceara, Brazil. 

Depth: 2 to 46 m (1 to 25 fm). 

Habitat: on coral reefs; sandj^, coral, and stone substrates. 

Remarks: This is the largest crab found in the Gulf of Mexico and Caribbean 

area and it is used for food in the West Indies. It reaches carapace widths of more 

than 15 cm. Garth (1965a) compared this species with C. convexus from the 

Pacific. Pequegnat and Raj^ (1974) reported some observations on mating and 

other behavior seen on a coral reef off the Texas coast. Saraiva da Costa (1968) 

described the biology and fishery of this species at Ceara, Brazil. Listed from 

Brazil by FaustoFilho (1968).

Carpoporus Stimpson, 1871 


Carpoporus papulosus Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 139) 

Rathbun, 1930, p. 269, pi. 110, figs. 3-6, pi. Ill; Williams, 1965, p. 186, figs. 168, 

183B; Guinot, 1967b, p. 551, figs. 18-19, 22. 

Range: North Carolina; Florida Keys and Dry Tortugas; west and northwest 

coasts of Florida; Alabama; north of Yucatan, Mexico. 

Depth: 33 to 110 m (18to60fm). 

Habitat: sand, broken shell, and coral bottoms. 

Remarks: Wass (1955) listed this species from northwest Florida. 

Calaleptodius Guinot, 1968 

Calaleplodius floridanus (Gibbes, 1850) (Proc. Amer. Assoc. Adv. Sci. 3: 175) 

As Leplodius /iorirfanus—Rathbun, 1930, p. 297, pi. 137, figs. 1-2, pi. 138, fig. 1; 

Rathbun, 1933, p. 57. 

As Calaleplodius floridanus—Gmnot, 1968a, p. 706, figs. 20, 23, 29. 

Range: Bermuda; Bahamas; Florida Keys and Drjr Tortugas; northwest coast 

of Florida; north coast of Cuba; .Jamaica; Puerto Rico; Virgin Islands; Antigua; 

Barbados; Curasao; Panama to Colombia (Caribbean coasts; Abolhos Islands to 



Habitat: on coral and stone reefs; in Sargassum; in living sponges; from sand, 

shell, grassy, mud bottoms; intertidal pools in rocky areas; under rocks. 

Remarks: Menzel (1971) listed this crab as rare on oyster reefs at Apalachee 

Bay, Florida. Listed from Florida by Abele (1970). Sulkin (1973) described 

larval depth regulation and Epifanio (1972) studied the effects of dieldrin on 

larval development. Hazlett (1976) described agonistic behavior in this crab. 

Listed from Brazil bj^ Fausto Filho (1974), he provided notes on color of crab 

and its habitat. 

CWorof/ieZ/a Rathbun, 1897 

Chlorodiella longimana (H. Milne Edwards, 1834) (Hist. Nat. Crust., vol. 1, 

p. 401) 

Rathbun, 1930, p. 462, pi. 186; Rathbuii, 1933, p. 68, fig. 58. 

Range: Bahamas; east coast of Florida; Florida Keys and Dry Tortugas; 

Jamaica; Puerto Rico; Virgin Islands; Martinique; Barbados; Curagao. 

Depth: 5 to 154 m (3 to 84 I'm). 

Habitat: coral reefs; rock and stone substrates; in sponges. 

Remarks: Rathbun (1930) also listed this species from West Africa. 

Domecia Eydoux & Souleyet, 18'1'2 

Domecia acanthophora acanthophora (Desbonne & Schramm, 1867) (Crust. 

Guadeloupe, p. 35) 

As D. /lispida—Rathbun, 1930, p. 554, pi. 227; Pequegnat & Ray, 1974, p. 237, 

figs. 16-17.


As D. acanthopliora acanlhophora—Guinot, 1964, p. 271, figs. 4-5, 7-8, 15; 

Williams, McCloskey & Graj^, 1968, p. 52. 

Range: North and South Carolina; east coast of Florida; Florida Keys and 

Dry Tortugas; West Flower Garden Bank, off Texas; Cuba; Jamaica; Puerto 

Rico; Barbados; Curasao; Alagoas to Pernambuco, Brazil. 


Habitat: on coral reefs; among sponges; rocky and coral bottoms. 

Remarks: The Pacific specimens of D. hispida listed by Rathbun (1930) 

remain with the species indicated; the Atlantic specimens were referred by 

Guinot (1964) to D. acanthophora. Patton (1967) studied the ecology of this 

species on coral reefs (Acropora) off Puerto Rico. A single male was taken from 

Oculina off North Carolina by Williams, McCloskey and Gray (1968). 

Domecia hispidalLydoux & Souleyet, 1842. 

Atlantic specimens were referred to D. acanthophora (Desbonne & Schramm, 

1867) by Guinot (1964). 

Eri/j/iia Latreiile, 1817 

Eriphia gonagra (Fabricius, 1781) (Species Insectorum, p. 505) 

Common Name: Calico Crab 

Hay & Shore, 1918, p. 439, pi. 35, fig. 6; Rathbun, 1930, p. 545, text-fig. 83, pi. 222; 

Rathbun, 1933, p. 76, fig. 64; Williams, 1965, p. 182, figs. 164A-C, 165; Felder, 

1973a,p. 64,pl. 9, fig. 5. 

Range: North Carolina; Bermuda; Bahamas; Florida Keys and Drj^ Tortugas; 

south Texas; north coast of Cuba; Jamaica; Puerto Rico; Virgin Islands; Antigua; 

Barbados; Trinidad; Aruba, Netherlands Antilles; Caribbean coasts of Panama 

and Colombia; Parahyba, Brazil to Argentina. 

Depth: intertidal to shallow water subtidal. 

Habitat: coral and stone reefs; under rocks and in crevices of intertidal pools; 

on rock jetties; brackish ponds; in seaweed and sponges. 

Remarks: 0vigorous females are known from south Florida and the West 

Indies during March to September (Williams, 1965). Furtado-Ogawa (1972) 

noted habitats in Brazil. 

Etisus H. Milne Edwards, 1834 

Etisus maculatus (Stimpson, 1860) (Ann. Lye. Nat. Hist. New York 7: 210) 

As Ptiymodius maculatus—Rathbun, 1930, p. 295, pi. 136; Rathbun, 1933, p. 57, 

fig. 49. 

As Etisus maculatus—Guinot, 1969b, p. 234. 

Range: Florida Keys and Dry Tortugas; Bahamas; north coast of Cuba; Puerto 

Rico; Virgin Islands. 

Depth: low tide mark and shallow water. 

Habitat: coral reefs; among rocks. 

Remarks: In transferring this species to Etisus, Guinot (1969b) remarks that


it becomes the sole representative of this genus in the Atlantic, other members 

of the genus being Indo-Pacific. 

Eucratodes A. Milne Edwards, 1880 

Eucratodes agassisii A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 347) 

Rathbun, 1930, p. 471, pi. 190; Rathbun, 1933, p. 68, fig. 59; Guinot, 1969a, p. 722, 

figs. 145-146; Pequegnat, 1970, p. 188. 

Range: off Mississippi; Yucatan Channel (Caribbean); Puerto Rico. 

Depth: 156 to 315 m (85 to 172 fm). 

Habitat: mud, sand, and shell bottoms. 

Remarks: Pequegnat (1970) collected an ovigerous female from 100 fm off 

Mississippi in early December. 

Eurypanopeus A. Milne Edwards, 1880 

Eurypanopeus abbreviatus abbreviatus (Stimpson, 1860) (Ann. Lye. Nat. Hist. 

New York 7: 211) 

Rathbun, 1930, p. 405, text-fig. 63, pi. 172, figs. 1-2; Rathbun, 1933, p. 64, fig. 55; 

Williams, 1965, p. 194, figs. 178, 183K; Felder, 1973a, p. 68, pi. 9, fig. 13. 

Range: South Carolina; Bahamas; Florida Keys; ? Louisiana; Texas; Jamaica; 

Haiti; Puerto Rico; Virgin Islands; Antigua; Barbados; Curasao; Trinidad; 

Colombia to Venezuela; Parahyba to Santa Catarina, Brazil. 

Depth: intertidal to shallow subtidal. 

Habitat: under rocks; on stone and coral reefs; under sponges and bryozoans; 

on oyster beds. 

Remarks: Behre (1950) listed a specimen of E. crenatus from Grand Isle, 

Louisiana. Felder (1973a) notes that TL. crenatus is known primarily from the 

Pacific coast of South America and that Behre's record may be a mistaken identification 

of E. abbreviatus. Williams (1965) listed ovigerous females from the 

West Indies during April to November and in southern Brazil from August to 

November. Furtado-Ogawa (1972) commented on ecology of this species in 

Brazil. 

Eurypanopeus abbreviatus ater Rathbun, 1930 (Bull. U.S. Nat. Mus. 152: 407) 


Range: Vera Cruz, Mexico. 

Remarks: Known only from the single male type specimen; no other data 

available. 

Eurypanopeus depressus (Smith, 1869) (Proc. Boston Soc. Nat. Hist. 12: 283) 


figs. 3-4; Williams, 1965, p. 195, figs. 179, 183L; Felder, 1973a, p. 67, pi. 9, fig. 17. 

Range: Massachusetts to south Florida; Bermuda; west coast of Florida to 

Texas; St. Martin, Leeward Islands. 

Depth: intertidal to 48 m (to 26 fm).


Habitat: very common on oyster bars; on muddj^ and stony shores, usually 

intertidal; on wharves and submerged pilings; in eel grass; brackish waters, to 

4.5 ppt salinity. 

Remarks: Developmental studies include descriptions of larval stages (Costlow 

and Bookhout, 1961b) and data on larval ecology in Chesapeake Bight (Sandifer, 

1973). Life history information was provided by Ryan (1956). Hyman (1925) 

figured zoeal stages. Lunz (1937) observed the association of this crab with 

oysters; McDermott (1960) noted the threat to oyster spats in New Jersey. 

Abele (1970) noted that the presence of the red spot on the third maxillipeds 

was not consistent among specimens, but is found in crabs only associated with 

03rsters. Records of ovigerous females are summarized in Williams (1965). 

Regional lists include Florida (Wass, 1955; Tabb and Manning, 1961; Dragovich 

and Kelly, 1964; Abele, 1970; Lyons et al, 1971; Menzel, 1971), Mississippi 

(Christmas and Langlej^ 1973), Louisiana (Behre, 1950), Texas (Leary, 1967), 

and the northwestern Gulf of Mexico (Fotheringham and Brunenmeister, 1975). 

Ayres (1938) studied resiDiration in relation to habitat preferences. 

Eurypanopeus dissimilis (Benedict & Rathbun, 1891) (Proc. U.S. Nat. Mus. 

14: 366) 


Range: west coast of Florida; north coast of Cuba; Jamaica; Nicaragua; Trinidad; 

Brazil. 

Habitat: Listed from a salt water lagoon in Nicaragua and from harbors in 

Florida and Jamaica, no depths given. 

Eurypanopeus turgidus (Rathbun, 1930). 

This is a manuscript name used bj^ Abele (1970) for Panopeus turgidus 

Rathbun, 1930 and listed by Menzel (1971, p. 80) for northwest Florida. Dr. 

Fenner A. Chace, Jr. examined the identification of Abele's specimens and compared 

them with species of Eurypanopeus. Until official revision of the two 

genera is complete, the present work will continue to recognize this species as a 

member of the genus Panopeus, while lecognizing its probable affinity with 

Eurypanopeus. See Panopeus turgidus. 

Eurylium Stimpson, 1859 

Eurylium limosuin (Saj^, 1818) (J. Acad. Nat. Sci., Philadelphia 1: 446) 

Hay & Shore, 1918, p. 438, pi. 35, fig. 7; Rathbun, 1930, p. 423, pi, 176, figs. 1-2; 

Rathbun, 1933, p. 65, fig, 56; Chace, 1940, p. 34; Williams, 1965, p. 199, figs. 182, 

1830; Chace & Hobbs, 1969, p. 153, figs. ^6, 46b; Felder, 1973a, p. 65, pi. 9, fig. 4. 

Range: Bermuda; Bahamas; New York to south Florida; Florida Ke3^s and 

Tirj Tortugas; west and northwest coasts of Florida; Louisiana; north and south 

coasts of Cuba; Jamaica; Hispaniola; Puerto Rico; Virgin Islands; Islas Los 

Roques; Curasao; Belize; Caribbean coast of Panama; Maranhao to Sao Paulo, 

Brazil. 

Depth: intertidal to shallow subtidal, near shore.


Habitat: muddy shores, especially among mangroves; burrows along tidal 

stream banks, burrows partially filled with water; under stones at high tide 

mark. 

Remarks: Williams (1965) notes that modern records limit the northern 

extent of the range to South Carolina. Regional lists include Florida (Wass, 

1955; Tabb and Manning, 1961; Abele, 1970; Menzel, 1971; Subrahmanyam 

etai, 1976), and Louisiana (Behre, 1950; Hoese and Valentine, 1972). Manning 

(1961) compared growth of this crab with that in Menippe mercenaria and 

Panopeus herbstii. Warner (1969) provided ecological data on this species in 

Jamaica and Ryan (1956) described the life history of populations observed in 

Chesapeake Bay. Teal (1959) studied respiration of this crab under different 

field conditions. 

Glyploxanthosus A. Milne Edwards, 1879 

Glyploxanthosus erosus (Stimpson, 1859) (Ann. Lye. Nat. Hist. New York 

7:51) 

Rathbun, 1930, p. 263, pi. 107; Williams, 1965, p. 185, figs. 167, 183A; Guinot, 

1967b, p. 551, fig. 30; Felder, 1973a, p. 60, pi. 9, fig. 9. 

Range: Bahamas; North Carolina; east coast of Florida; Florida Keys and Dry 

Tortugas; west and northwest coasts of Florida; Louisiana and Texas (uncommon) 

; Campeche Banks and off Yucatan, Mexico; Guadeloupe. 


Habitat: sand, broken shell, coral, and rock bottoms; from rocks and algal 

mats {Halimeda) in shallow water; sponges and coral reefs of deeper water. 

Remarks: Ovigerous females are known from off northwest Florida in January 

(Williams, 1965). Listed from Florida by Wass (1955) and Chace (1956), from 

Louisiana by Behre (1950), and from Texas by Leary (1967). 

Heteraclaea Lockington, 1877 

Heleraclaea ceralopus (Stimpson, 1860) (Ann. Lye. Nat. Hist. New York 7: 

215) 

Rathbun, 1930, p. 530, pi. 212, figs. 5-8; pi. 213; Guinot, 1968a, p. 721, figs. 50, 56. 

Range: Bahamas; east coast of Florida; Florida Keys and Dry Tortugas; north 

coast of Cuba; Curagao; Trinidad; Barbados. 


Habitat; on coral reefs; in sponges and coral. 

Hexapanopeus Rathbun,1898 

Hexapanopeus anguslifrons (Benedict & Rathbun, 1891) (Proc. U.S. Nat. Mus. 

14: 373) 


figs. 1-2; Williams, 1965, p. 188, figs. 170, 183D; Felder, 1973a, pi. 9, fig. 24. 

Range: Massachusetts to South Carolina; Bahamas; west and northwest coasts 

of Florida; Mississippi to Texas; Jamaica. 

^


Depth: near shore subtidal to 139 m (to 76 fm). 

Habitat: most commonly on muddy bottoms; occasionally on sand, shell, and 

gravel substrates; on oyster beds. 

Remarks: Developmental studies include data on growth conditions for zoeal 

stages (Chamberlain, 1961), descriptions of larval stages as reared in the laboratory 

(Costlow and Bookhout, 1966a), and notes on larval ecology in Chesapeake 

Bight (Sandifer, 1973). Williams (1965) summarized records of ovigerous 

females. Ryan (1956) described the general life history of this crab in Chesapeake 

Bay. Hazlett (1976) studied agonistic behaviors. Regional lists include 

Florida (Wass, 1955; Dragovich and Kelly, 1964; Abele, 1970; Menzel, 1971; 

L3'ons et al, 1971), Louisiana (Behre, 1950), and Texas (Hedgpeth, 1953; 

Leary, 1967). 

Hexapanopeus hemphillii (Benedict & Rathbun, 1891) (Proc. U.S. Nat. Mus. 

14: 374) 

Rathbun, 1930, p. 400, pi. 171, figs. 1-2, 6; Rathbun, 1933, p. 63. 

Range: Florida Keys and Dry Tortugas; west coast of Florida; north coast of 

Cuba; Puerto Rico; St. Thomas, Virgin Islands. 


Habitat: coral, sand, and gravel bottoms; off turtle grass flats. 

Hexapanopeus lobipes (A. Milne Edwards, 1880) (Crust. Reg. Mex., p. 331) 

As Lophopanopeus lobipes—Rathbun, 1930, p. 329, text-fig. 50, pi. 155, figs. 3-5. 

As Hexapanopeus lobipes—Menzies, 1948, p. 23. 

Range: Bahamas; off Key West, in Florida Straits; northwest of Dry Tortugas. 

Depth: 68 m (37 fm); also recorded from shallow water. 

Habitat: sand bottoms; coral banks. 

Remarks: Menzies (1948) noted that this species did not fit the generic description 

of Lophopanopeus and he transferred it to Hexapanopeus, the American 

genus it most closely resembled. 

Hexapanopeus paulensis Rathbun, 1930 (Bull. U.S. Nat. Mus. 152: 395) 

Rathbun, 1930, p. 395, pi. 170, figs. 5-6; Williams, 1965, p. 189, figs. 171, 183E; 


Range: South Carolina; northwest Florida; Texas; Sao Paulo, Brazil. 

Depth: inter tidal to 5 m (3 fm). 

Habitat: sand, broken shell, and rock substrates; off rock jetties; among 

sponges, ascidians, and bryozoans. 

Remarks: Originally listed from South Carolina by Lunz (1937). Additional 

records, from northwest Florida, were provided by Abele (1970), including 

ovigerous females taken in July. 

Hexapanopeus quinquedenlalus Rathbun, 1901 (Bull. U.S. Fish Comm. for 

1900,2: 31) 

Rathbun, 1930, p. 402, text-fig. 62. 

Range: northwest Florida; Puerto Rico. 

Depth: 15 to22m (8.5 to 12fm).


Habitat: coral sand, sand-grass, rocky, and sticky mud bottoms. 

Remarks: Rathbun (1930) noted the resemblance between this species and 

H. sinaloensis, a rare species from the west coast of Mexico. Abele (1970) provided 

the first Gulf records and noted a close resemblance to H. paulensis. Abele 

(1970) states that Fenner Chace, Jr., of the U.S.N.M. compared the Floridian 

specimens with the Puerto Rico holotj-pe and noted the differences between 

them. The paucity of specimens of these three similar forms only permits a 

tentative designation of this crab; until further material and revision is available, 

this list follows the nomenclature of Abele (1970, p. 78). 

Leptodius A. Milne Edwards, 1863 

Leptodius agassizzi A. Milne Edwards, 1880. 

Transferred to a new genus by Guinot (1968a), see Pseudomedaeus agassizii. 

Leptodius floridanus (Gibbes, 1850). 

Transferred to a new genus by Guinot (1968a), see Cataleptodius floridanus. 

Leptodius parvulus (Fabricius, 1793) (Entom. Syst. Auct. et emend., vol. 2, p. 

451) novo comb. Rathbun, 1930 (Bull. U.S. Nat. Mus. 152:305) V ^^'^^'''"'•^ a- 

Rathbun, 1930, p. 303, pi. 141, figs. 1-3; Rathbun, 1933, p. 38, fig. 30. ' 

Range: Bermuda; Bahamas; Florida Keys; Jamaica; Haiti; Puerto Rico; Barbados; 

Curagao; Fernando deNoronha, Brazil. 

Habitat: shallow water, near shore; under rocks; in tide pools. 

Remarks: Fausto Filho (1974) provided notes on color and habitat of Brazilian 

specimens. 

Lobopilumnus A. Milne Edwards, 1880 

Lobopilumnus agas.'sisii (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 142) 


Williams, 1965, p. 181, figs. 157G, 163. 

Range: Bermuda; North Carolina; Florida Keys and Dry Tortugas; west and 

northwest coasts of Florida; north of Yucatan; north coast of Cuba; Trinidad. 


Habitat: sand, gravel, rock, coral, and broken shell substrates; under stones 

and dead corals; in sponges. 

Remarks: Four environmental forms of this highly variable crab were recognized 

by Rathbun (1930): L. a. typica, L. a. bermudensis, L. a. pulchella, and 

L. a. trinidadensis. The distributions of the various forms overlap to some extent. 

Verrill (1908) described this species in Bermiida. Reported from the northeast 

Gulf by Chace (1956) and listed from northwest Florida by Abele (1970). 

Ovigerous females are known from Florida and Cuba during February to July 

(Williams, 1965). Pearse (1934) noted the presence of this crab inside the 

loggerhead sponge, Spheciospongia vesparia.


Lophopanopeus Rathhun, 1898 

Lophopanopeus distinclus Rathbun, 1898. 

Transferred to the genus Micropanope by Menzies (1948), see Micropanope 

distincta (Rathbun, 1898). 

Lophopanopeus lobipes (A. Milne Edwards, 1880). 

Transferred to the genus Hexapanopeus by Menzies (1948), see Hexapanopeus 

lobipes (A. Milne Edwards, 1880). 

MeZyfoia Stimpson, 1871 

Melyhia ihalamlla Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 144) 

Rathbun, \930, p. 562, pi. 230. 

Range: Dry Tortugas; north coast of Cuba; Jamaica; Haiti; Culebra and 

Vieques Islands; Barbados; Curagao; Colon, Panama; Abrolhos Islands, Brazil. 

Depth: low tide mark to 368 m (to 201 fm), most common to 82 m (45 fm). 

Habitat: coral, sand, rock, and broken shell bottoms. 

Remarks: The deepest recorded specimens (192 and 201 fm) are from off 

Havana, Cuba. 

Menippe de Haan, 1833 

Menippe mercenaria (Say, 1818) (J. Acad. Nat. Sci. Philadelphia 1: 448) 

Common Name: Stone Crab 

Hay & Shore, 1918, p. 439, pi. 35, fig. 8; Rathbun, 1930, p. 472, text-fig. 78, pis. 

191-193; Williams, 1965, p. 183, figs. 164D-E, 166; Felder, 1973a, p. 64, pi. 9, figs. 

2^3. 

Range: North Carolina to south Florida; Bahamas; Florida Keys; southwest 

Florida to south Texas; off Yucatan Gulf coast; north coast of Cuba; Jamaica. 

Depth: surface and intertidal to 51 m (to 28 fm). 

Habitat: in estuaries and bays of near-marine salinity; from sand, shell, clay, 

and mud substrates; in deeper waters on offshore reefs; in turtle grass (Thalassia) 

beds off northwest Florida; post-larval crabs are common in deeper channels and 

bays, under shell fragments; older juveniles and adults are among rocks, under 

stones, and on and among oyster bars. 

Remarks: Except for the blue crab (Callinectes sapidus), this species is the 

most important crab commercially harvested on the Gulf coast, primarily in 

Florida. Williams (1965) summarizes much of the literature and accounts of 

the natural history of stone crabs in Texas are provided by Powell and Gunter 

(1968) and by Futch (1966) for Florida populations. Fotheringham and Brunenmeister 

(1975) offer general comments on the stone crab in the northwestern 

Gulf. 

Developmental studies include data on fertilization (Binford, 1913), zoeal 

stage descriptions (Porter, 1960), studies of effects of temperature and salinity 

on larvae (Ong and Costlow, 1970), effects of Mirex on larvae (Bookhout et al., 

1972), and studies of larval energy budgets (Mootz and Epifanio, 1974). Studies


of behavior include those on mating (Flartnoll, 1969; Savage, 1971b), stridulation 

(Guinot-Dumortier and Dumortier, 1960), and shadow responses of larvae 

(Forward, 1977). Physiological research includes data on hormonal control of 

reproduction (Cheung, 1967, 1969), respiration in relation to habitat (A3a'es, 

1938), gill area measurements (Gray, 1957), respiration and metabolism 

(Leffler, 1973), respiration and osmoregulation (Karandieva and Lee, 1967), 

and digestion and energjr balance (Suchenia and Claro Madruga, 1967). 

Information on growth, molting, and fisheries includes reports by Manning 

(1961), Savage and McMahan (1968), Savage (1971a), and Savage, Sullivan 

and Kalman (1974, 1975). Cheung (1973, 1976) studied regeneration of claws 

in relation to molting. Menzel and Hopkins (1956) described predation of stone 

crabs on oysters in Louisiana. Iversen and Beardsley (1976) studied shell disease 

in south Florida populations. Humes (1941a) noted the presence of a parasitic 

barnacle. 

Regional lists include Florida (Wass, 1955; Tabb and Manning, 1961; 

Dragovich and Kelly, 1964; Abele, 1970; Menzel, 1971; Lyons et al., 1971), 

Mississippi (Richmond, 1962; Franks et al., 1972; Christmas and Langley, 1973), 

Louisiana (Behre, 1950; Hoese and Valentine, 1972), Texas (Gunter, 1950; 

Whitten, Rosene and Hedgpeth, 1950; Hedgpeth, 1953; Hildebrand, 1954; Simmons, 

1957; Parker, 1959; Hoese, I960; Leary, 1967), and the northeastern 

Gulf (Chace, 1956). 

Menippe nodifrons Stimpson, 1859 (Ann. Lye. Nat. Hist. New York 7: 53) 

Rathbun, 1930, p. 479, pi. 198, fig. 3, pi. 199. 

Range: east coast of Florida; ? Louisiana; north and south coasts of Cuba; 

Jamaica; Virgin Islands; Trinidad; Caribbean coasts of Panama and Colombia; 

Paraiba to Sao Francisco do Sul, Brazil; Gabon, West Africa. 

Depth: shallow water, near shore. 

Habitat: tide pools; under rocks; near dock pilings; on sponges and among 

brji-Qzoans. 

Remarks: Rathbun's (1930) record of this crab from Louisiana has been questioned 

by Felder (1973a) because extensive collecting along the northwestern 

Gulf coast has produced no further specimens. The only other Gulf record would 

be that of a single male collected near Havana, Cuba (Rathbun, 1930). 

Micropanope Stimpson, 1871 

(As restricted by Guinot (1967a, 1968b), this genus would include only 

the type species, M. sculptipes Stimpson, and one other, leaving a number 

of species formerly in Micropanope without an apparent generic name. 

Following a suggestion of Garth (pers. comm.), this list retains these 

orphaned species, not included in other genera by Guinot, as members of 

Micropanope, pending further clarification of their taxonomic status.) 

Micropanope barbadensis (Rathbun, 1921) (Bull. Lab. Nat. Hist. State Univ. 

Iowa 9: 73), novo comb. Rathbun, 1930 (Bull. U.S. Nat. Mus. 152: 446) 

iO Rathbun, 1930, p. -M-G, text-fig. 72.


Range: Dry Tortugas; Barbados. 


Habitat: froni coral heads. 

Micropanope distincla (Rathbun, 1898) (Bull. Lab. Nat. Hist. State Univ. Iowa 

4: 272) 

As Lophopanopeus dislinctus—Rathbun, 1930, p. 331, pi. 155, figs. 1-2. 

As Micropanope dislincta—Menzies, 1948, p. 24. 

Range: North Carolina; Florida Straits; Dry Tortugas; northwest Florida; 

Barbados. 

Depth: 48tol85m(26tol01fm). 


Remarks: Menzies (1948) transferred this species to Micropanope and he 

considered it to be close to M. sculptipes Stimpson, the type species of the genus. 

Cerame-Vivas and Gray (1966) extended the known range (listed as Lophopanopeus) 

to North Carolina. 

Micropanope lobifrons A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 327) 


Range: south Florida, in Gulf Stream; Dry Tortugas; off northwest Florida; 

off north coast of Cuba; Puerto Rico; Virgin Islands; Santa Cruz Island (Carib.); 

Grenada; Barbados; Colon, Panama. 

Depth: 37 to 311 m (20 to 170 fm). 

Habitat: sand, coral, rock, and broken shell substrates. 

Remarks: Guinot (1968b) retained this species in Micropanope along with 

the type species, while revising the genus. 

Micropanope nuUingi (Rathbun, 1898) (Bull. Lab. Nat. Hist. State Univ. Iowa 

4: 271), novo comb. Rathbun, 1930 (Bull. U.S. Nat. Mus. 152: 450) 

Ratlibun, 1930, p. 450, text-fig. 74; Rathbun, 1933, p. 67, fig. 57; Williams, 1963, 

p. 194, figs. 177,183J; Felder, 1973a, p. 66, pi. 9, fig. 22. 

Range: North Carolina; east coast of Florida; Florida Keys and Dry Tortugas; 

west and northwest coasts of Florida; Texas; north coast of Yucatan; north coast 

of Cuba; Jamaica; Puerto Rico; Virgin Islands; Old Providence Island (Carib.); 

Rio Grande do Norte, Brazil. 


Habitat: rock, sand, coral, and broken-shell substrates; from boulder-covered 

beaches; from clumps of Porites and Halimede. 

Remarks: Felder (1973a) notes that this species may be eventually placed in 

a different genus on the basis of differences in male first pleopods. Williams 

(1965) listed ovigerous females from Florida in July. 

Micropanope pusilla A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 327) 


Range: Dry Tortugas; northwest of Key West; west and northwest coasts of 

Florida; Alabama; north coast of Cuba; Jamaica; Puerto Rico; Virgin Islands.


Depth: 31 to 311 m (17 to 170 fm). 


Remarks: Listed from Florida by Wass (1955), Abele (1970), and Menzel 

(1971). Rathbun (1930) reported ovigerous females from the Gulf of Mexico 

during February-March and June-July. Abele (1970) noted that this species does 

not fit any of the genera reviewed or erected by Guinot (1968a). 

Micropanope sculplipes Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 140) 

Rathbun, 1930, p. 428, pi. 178, figs. 1-3; Rathbun, 1933, p. 66; Williams, 1965, 

p. 193, fig. 175; Felder, 1973a, p. 66, pi. 9, fig. 15. 

Range: North and South Carolina; Florida Keys and Drjr Tortugas; west and 

northwest coasts of Florida; Texas; St. Croix; Grenada; Barbados. 

Depth: 9 to 311 m (5 to 170 fm). 


Remarks: Listed from Florida by Wass (1955). Cerame-Vivas and Gray 

(1966) extended the known range of this crab to North Carolina. 

Micropanope spinipes A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 326) 

Rathbun, 1930, p. 443, text-fig. 71, pi. 181, figs. 1-2; Pequegnat & Ray, 1974, p. 238, 

figs. 18-22. 

Range: Bermuda; Bahamas; Florida Keys; West Flower Garden Bank, off 

Texas; Curasao; Alagoas and off the Abrolhos Islands, Brazil. 


Habitat: sand and coral bottoms; off coral reefs; in sponges. 

Micropanope truncalifrons Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. 

Iowa 4: 274) 


Range: off north coast of Cuba; off Caribbean coast of Yucatan. 

Depth: 238 to 355 m (130 to 194 fm). 

Habitat: coral sand bottoms. 

Remarks: This species may be eventually transferred to the genus Nanocassiope. 

Guinot (1971, p. 1076) noted a resemblance between this species and A^. 

melanodactylus (A. Milne Eldwards), the type species of the genus recently 

erected by Guinot (1967a). 

Micropanope urinalor (A. Milne Edwards, 1881) (Crust. Reg. Mex., p. 289) 

Rathbun, 1930, p. 451, pi. 182, figs. 3,^, pi. 183, figs. 1-3; Chace, 1940, p. 34; 

Williams, McCloskey & Gray, 1968, p. 51, fig. 7. 

Range: off North Carolina; Florida Keys; north and south coasts of Cuba; St. 

Croix. 

Depth: 146 to457m (80 to250 fm). 


Micropanope xanlhiformis (A. Milne Edwards, 1880). 

Transferred to a new genus, Nanoplax, by Guinot (1967a). See Nanoplax 

xanthiformis.


IS'anoplax Guinot, 1967 

Nanoplax xanthtformis (A. Milne Edwards, 1880) (Crust. Reg. Mex., p. 353) 

As MicTopanope xanlhiformis—Rathbun, 1930, p. 442, pi. 180, figs. 7-8; Rathbun, 

1933, p. 67:, Williams, 1965, p. 193, figs. 176,1831. 

As Nanoplax xanlhiformis—Guinot, 1967a, p. 362, fig. 16. 

Range: North Carolina to off south Florida; Florida Keys and Dry Tortugas; 

northwest coast of Florida; north coast of Cuba; off Caribbean coast of Yucatan; 

Puerto Rico; Dominica; Barbados; Grenada; Curagao; off Cape Frio, Brazil. 

Depth: 9 to 333 m (5 to 182 fm). 

Habitat: sand, broken shell, coral, and mud substrates. 

Remarks: Guinot (1967a) discussed the affinities of this species and genus 

with certain genera of Goneplacidae. Listed from northwest Florida (as Micropanope) 

by Wass (1955) and Hulings (1961). Ovigerous females are known 

from Florida in June and August and from North Carolina in October (Williams, 

1965). 

Neopanope A. Milne Edwards, 1880 

Neopanope packardii (Kingsley, 1879) (Proc. Boston Soc. Nat. Hist. 20: 152) 

Rathbun, 1930, p. 380, text-fig. 59, pi. 168, figs. 5-6; Abele, 1972a, p. 269, figs. 

1B,3A. 

Range: southeast and south Florida; Bahamas; Florida Keys and Dry Tortugas; 

west and northwest coasts of Florida; Louisiana; north coast of Cuba. 

Depth: lowtide mark to 74m (to 135 fm). 

Habitat: sand, gravel, rock, and coral substrates; occasionally in mud; grass 

beds, algal beds, marshes; on Styela. 

Remarks: A systematic review and morphological key for this genus was presented 

by Abele (1972a). Abele (1971) provided scanning electron micrographs 

of the gonopods of this crab and other Neopanope. Larval development was 

described by Costlow and Bookhout (1967); larvae of various Neopanope species 

were compared by McMahan (1967). Records of ovigerous crabs were listed by 

Rathbun (1930). Listed from Florida by Tabb and Manning (1961), Dragovich 

and Kelly (1964), Abele (1970), Rouse (1970), Menzel (1971), and Lyons 

et al. (1971), and from Louisiana by Hoese and Valentine (1972). 

Neopanope texana (Stimpson, 1859) (Ann. Lye. Nat. Hist. New York 7: 55) 

As ;V. lexana Icvano—Rathbun, 1930, p. 367, text-fig. 57, pi. 168, figs. 1-2. 

As TS. texana—Abele, 1972a, p. 266, figs, lA, 2A, 2E, 3B, 3C; Felder, 1973a, p. 68, 

pi. 9, fig. 19; not Williams, 1965, p. 190 (= N. sayi). 

Range: west coast of Florida (south as far as Charlotte County) to south 

Texas. 


Habitat: Thalassia grass flats; mud, sand, rock, and gravel substrates; among 

barnacles, clumps of ascidians. 

Remarks: Abele (1972a) reviewed the status of the two related species. A''. 

texana and N. sayi. The latter form is restricted in distribution to the east coast

7'^ 


of the United States, whereas A^. texana occurs only along the Gulf coast. Reports 

of A^. texana in southern Florida (Tabb and Manning, 1961) are attributed by 

Abele to A^. packardii. Williams (1965) listed this species from North Carolina, 

but Abele (1972a) states that the figures are of A^. sayi. Florida listings include 

Wass (1955), Dragovich and Kelly (1964), Abele (1970), Lyons et al. (1971), 

and Menzel (1971). Other listings (some as A', texana sayi (= A', sayi), but all 

are referred to A^. texana) include the northwestern Gulf (Fotheringham and 

Brunenmeister, 1975) and Texas (Simmons, 1957; Parker, 1959; Hoese, 1960; 

Breuer, 1962; Hoese and Jones, 1963; Keith and Hulings, 1965; Leary, 1967). 

McMahan (1967) described lanâe reared in the laboratory. Landers (1954) 

noted predation by this species on clams. Ryan (1956) gave accounts of life 

history for A^. sayi, which should be very similar to that of N. texana. 

Panopeus H. Milne Edwards, 1834 

Panopeus americanus Saussure, 1857 Rev. Mag. ZooL, ser. 2, vol. 9: 502) 


Range: Bahamas; Florida Keys; west coast of Florida; north coast of Cuba; 

Jamaica; Dominican Republic; Puerto Rico; St. Thomas, Virgin Islands; Guadeloupe; 

Trinidad; Caribbean coast of Colombia; Rio Parahyba do Norte to Santa 

Catarina, Brazil. 

Depth: inter tidal to 22 m (to 12 fm). 

Habitat: under stones, on mud flats; on mangroves; sand, shell, and mud bottoms; 

grass flats; under sponges and bryozoans. 

Rem.arks: Listed from south Florida by Tabb and Manning (1961). 

Panopeus bermudensis Benedict & Rathbun, 1891 (Proc. U.S. Nat. Mus. 14: 

376) 

Rathbun, 1930, p. 360, text-fig. 56, pi. 165; Rathbun, 1933, p. 62; Garth, 1961, 

p. 149; Felder, ig73a, p. 69, pi. 9, fig. 20. 

Range: Bermuda; Bahamas; west coast of Florida; PTexas; north coast of 

Cuba; Jamaica; Puerto Rico; St. Thomas, Virgin Islands; Trinidad; Old Providence 

Island (Carib.); Colombia to Santa Catarina, Brazil; in Pacific—from 

Magdalena Bay, Mexico to Peru. 


Habitat: oyster beds; rocky tide pools; under rocks, sponges, bryozoans, debris, 

and among fouling organisms; from bays and near-marine waters; on mangrove 

roots; from sand bottoms. 

Remarks: Felder (1973a) listed this species from Texas on the basis of a tentative 

identification. Garth (1961) noted the variability of this species over its 

entire geographical range and pointed to the possibility of future segregation of 

this species into distinct Pacific and Atlantic forms. 

Panopeus harttii Smith, 1869 (Proc. Boston Soc. Nat. Hist. 12: 280) 


Range: Florida Keys and Dry Tortugas; Isla de Pinos, Cuba (Carib.); Puerto


Rico; St. Thomas, Virgin Islands; Antigua; Barbados; Pernambuco to Sao Paulo, 

Brazil. 

Depth: low tide mark and shallow waters. 

Habitat: on rocks and coral reefs. 

Remarks: Color and habitat in Brazil were described by Fausto Filho (1974). 

Panopeus herbstii H. Milne Edwards, 1834 (Hist. Nat. Crust., vol. 1, p. 403) 

Common Name: Common Mud Crab 


pis. 156-157; Chace, 1940, p. 34; Ryan, 1956, p. 147, text-figs. 4B, 5B, 9A, pi. IC; 

Edmondson, 1962, p. 277; Williams, 1965, p. 196, figs. 180, 183M; Chace & Hobbs, 

1969, p. 154, figs. 46c, 47; Felder, 1973a, p. 69, pi. 9, fig. 21. 

Range: Bermuda; Bahamas; Massachusetts to south Florida; Florida Keys 

and Dry Tortugas; west coast of Florida to Veracruz, Mexico; north and south 

coasts of Cuba; Jamaica; Haiti; Puerto Rico; Virgin Islands; Antigua to Barbados; 

Ti'inidad; Islas Los Roques; Netherlands Antilles; Belize; Caribbean coast 

of Panama to Venezuela; Rio Parahyba do Norte, Brazil to Uruguay; in Pacific— 

Hawaiian Islands. 

Depth: intertidal to 22m (12 fm). 

Habitat: muddy bottoms of bays and estuaries; shell, rock, and stone bottoms; 

oyster beds; among mangroves and in banks of tidal streams; burrows in mud 

banks and under stones or shells; on coral and rock reefs; brackish tide pools; 

sandy beaches with rocks. 

Remarks: Larval development has been described by Costlow and Bookhout 

(1961a) and Costlow, Bookhout and Monroe (1962). Sulkin (1973) reported on 

larval depth regulation and Sandifer (1973) gathered data on larval ecology in 

Chesapeake Bight. Schwartz and Cargo (1960) recorded this crab in Virginia 

and Maryland. Life history studies include those of Ryan (1956) in Chesapeake 

Bay and Warner (1969) in Jamaica. Rathbun (1930) listed three forms, in addition 

to the typical form, of this species and provided distribution records for 

each. Furtado-Ogawa (1972) noted individual variations and habitat differences 

for this species in Brazil. Williams (1965) provided a summary of the many 

studies on this crab. Relationships with oysters and role as a molluscan predator 

are included in reports by McDermott and Flower (1953), Ryan (1956), Menzel 

and Nichy (1958) and McDermott (1960). Perkins (1975) describes the fine 

structure of a haplosporid parasite of this crab and Humes (1941a) noted the 

presence of a parasitic barnacle. Forward (1977) studied shadow responses of 

the larv'âe. Physiological studies include reports on antennule chemosensitivity 

(Hazlett, 1971), coagulation (Morrison and Morrison, 1952), thoracic neurosection 

(Maynard, 1961a, 1961b; Maynard and Maynard, 1962), gill area 

(Gray, 1957), respiration in relation to habitat (Ayres, 1938; Teal, 1959), 

amino acid metabolism (Boone and Claybrook, 1977), and respiration and 

metabolism (Leffler, 1973). Regional lists include Florida (Wass, 1955; Dragovich 

and Kelly, 1964; Abele, 1970; Menzel, 1971; Subrahmanyam et al, 1976), 

Mississippi (Richmond, 1968; Christmas and Langley, 1973), Louisiana (Behre, 

1950; Hoese and Valentine, 1972), Texas (Leary, 1967), and the northwestern 

Gulf of Mexico (Fotheringham and Brunenmeister, 1975).

Crabs of the Gulf oj Mexico 103 

Panopeus occidentalis Saussure, 1857 (Rev. Mag. Zool., ser. 2, vol. 9: 502) 

Rathbun, 1930, p. 348, text-fig. 55, pi. 161; Rathbun, 1933, p. 61; Williams, 1965, 

p. 198, figs. 181,183N. 

Range: Bermuda; Bahamas; North Carolina to southeast Florida; Florida 

Keys and Dry Tortugas; west coast of Florida; north coast of Cuba; Jamaica; 

Hispaniola; Puerto Rico; Virgin Islands; Guadeloupe; Old Providence Island 

(Carib.); Curasao; Trinidad; Colon, Panama to Santa Catarina, Brazil. 

Depth: intertidalto 18 m (to 10 fm). 

Habitat: sand, shell, rock, and gravel bottoms; among ascidians, sponges, and 

seaweed; on mangrove roots; under rocks; on pilings and piers. 

Remarks: Although both of the environmental forms listed by Rathbun (1930) 

were recorded from Louisiana by Behre (1950), Felder (1973a) doubts the 

validity of the Grand Isle records and believes that they may represent P. herbstii. 

Williams (1965) provided a good summary of data on this crab. Listed from 

Florida by Tabb and Manning (1961) and by Lyons et al. (1971). De Oliveira 

(1940) described specimens from Brazil and provided notes on life history; Furtado-Ogawa 

(1972) noted individual variations and habitat differences in Brazil. 

Panopeus rugosus A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 314) 

Rathbun, 1930, p. 353, pis. 162-163. 

Range: Florida Keys and Dry Tortugas; west and northwest coasts of Florida; 

north coast of Cuba; Haiti; Virgin Islands; Puerto Rico; Honduras to Nicaragua; 

^ Curagao; Bahia to Santa Catarina, Brazil. 

^ • Depth: low tide mark to 51 m (to 28 fm). 

'] Habitat: sand, shell, rock, and coral bottoms; coral reefs; on pilings. 

I Remarks: listed from northwest Florida by Wass (1955) 

* 

\^ Panopeus turgidus Rathbun, 1930 (Bull. U.S. Nat. Mus. 152: 364) 

- _ Rathbun, 1930, p.364,pl. 166;Felder, 1973a,p. 68,pl.9, fig. 18. 

Range: northwest coast of Florida to Texas. 

[. Depth: near shore, shallow waters. 

K Habitat: bay and near-marine waters; found on or among rocks, shells, debris, 

p- and vegetation. 

f. Remarks: Listed from northwest Florida by Wass (1955) and from Louisiana 

j^ by Behre (1950). Abele (1970) collected a specimen from northwest Florida 

I and Fenner Chace, Jr. of the USNM confirmed its identity by comparison with 

the type, suggesting it was similar to the genus Eurypanopeus. Abele (1970) 

listed it in his thesis as Eurypanopeus turgidus and this was repeated in the list by 

Menzel (1971). As indicated under this latter name, this present listing will 

continue to use Panopeus turgidus until a definitive study or revision is available. 

Paractaea Guinot, 1969 

Paractaea rujopunctala nodosa (Simpson, 1860) (Ann. Lye. Nat. Hist. New 

York 7: 203) 

As Aclaea rufopunctata nodosa—Ralhbun, 1930, p. 257, pi. 105, figs. 1-2;


Rathbun, 1933, p. 56; Holthuis & Gottlieb, 1956, p. 287; Williams, McCloskey & 

Gray, 1968, p. 51. 

As Paractaea rufopunctata forma nodosa—Guinot, 1969b, p. 252, fig. 25. 

Range: North Carolina; Bahamas; east coast of Florida; Florida Keys and Dry 

Tortugas; north coast of Cuba; Jamaica; Haiti; Puerto Rico; Virgin Islands; 

Barbados; Curasao; Cape Frio, Brazil; Ascension Island, South Atlantic. 

Depth: 5 to 212 m (3 to 115 fm). 

Habitat: coral, broken shell, stone, and sand bottoms; in sponges. 

Remarks: This genus was erected by Guinot (1969b) for several species of 

Actaea. She compared the differences and similarities of this form and forma 

africana and other forms of the rufopunctata complex, deciding that further 

research was needed to clarify the systematic relationships of the genus. 

ParaZiomer« Rathbun, 1930 

Paraliomera dispar (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2; 140) 

Rathbun, 1930, p. 244, lext-fig. 38, pi. 101, figs. 4-5; Rathbun, 1933, p. 54. 

Range: Bermuda; Florida Keys and Dry Tortugas; north coast of Cuba; Jamaica; 

Haiti; Puerto Rico; Antigua; Barbados; Curagao; Caribbean coast of 

Colombia. 


Habitat: sand, shell, grassy, and mud bottoms; coral reefs and rocky areas. 

Paraliomera longimana (A. Milne Edwards, 1865) (Nouv. Arch. Mus. Hist. 

Nat, Paris 1: 221). 

Rathbun, 1930, p. 243, pi. 101, figs. 1-3; Rathbun, 1933, p. 53, fig. 46. 

Range: Florida Keys and Dry Tortugas; Veracruz, Mexico; Puerto Rico; 

Virgin Islands; Barbados; Curasao. 


Habitat: coral reefs and rocky and grassy substrates. 

Phymodius A. Milne Edwards, 1863 

Phymodius maculalus (Stimpson, 1860). 

This species was transferred to Etisus by Guinot (1969b), thus leaving the 

genus Phymodius unrepresented in the Gulf of Mexico. 

Pilumnoides H. Milne Edwards & Lucas, 1843 

Pilumnoides nudifrons (Stimpson, 1871) (Bull. Mus. Comp. Zool. 2: 143) 


Range: Florida Straits and Keys; Barbados. 

Deptli: 128 to 556 m (70 to 304 fm). 

Habitat: sand and rock substrates.

Piluninus L,each, 1815 


Piluinnus caribaeus Desbonne & Schramm, 1867 (Crust. Guadeloupe, p. 32) 


Range: Bahamas; Florida Keys; north coast of Cuba; Jamaica; Puerto Rico; 

Vieques and Culebra; Virgin Islands; Guadeloupe; Curagao; Bahia to Sao Paulo, 

Brazil. 


Habitat: mud, sand, shell, grassy, and coral bottoms. 

Piluinnus dasypodus Kingsley, 1879 (Proc. Boston Soc. Nat. Hist, 20: 155) 


p. 178, figs. 157C, 159; Felder, 1973a, p. 61, pi. 9, fig. 7. 

Range: North and South Carolina; Florida Keys and Dry Tortugas; south to 

northwest Florida; Mississippi; Texas; north and south coasts of Cuba; Jamaica; 

Puerto Rico; Culebra; Virgin Islands; Martinique; Curasao; Pernambuco to 

Santa Catarina, Brazil. 

Depth: 1 to29m (0.5 to 16 fm). 

Habitat: sand, shell, rocky, and coral bottoms; from mangrove roots; with 

fouling materials on pilings, buoys and jetties; off sponges. 

Remarks: Sandifer (1974) studied larval development. Williams (1965) reviewed 

records of ovigerous females. Ecological notes were provided by Lunz 

(1937), Pearse (1934), and Pearse and Williams (1951). Listed from Florida 

(Wass, 1955; Abele, 1970; Menzel, 1971; Lyons et ah, 1971), Mississippi 

(Christmas and Langley, 1973) and from Texas (Leary, 1967). 

Pilumnus diomedeae Rathbun, 1894 (Proc. U.S. Nat. Mus. 17: 85) 


Range: north coast of Cuba; Caribbean coast of Yucatan. 

Depth: 238 to 337 m (130 to 184 fm). 

Habitat: coral and sand bottoms. 

Pilumnus floridanus Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 141) 


p. 179, figs. 157D, 160; Felder, 1973a, p. 61, pi. 9, fig. 8; Pequegnat & Ray, 1974, 

p. 238, figs. 23-24. 

Range: North Carolina; Bahamas; Florida Keys and Dry Tortugas; west and 

northwest coasts of Florida; off Texas; north of Yucatan; Honduras; Jamaica; 

Puerto Rico; Culebra; Virgin Islands; Venezuela. 


Habitat: sand, shell, gravel, rock, and coral substrates; mud and grassy bottoms; 

inside sponges. 

Remarks: Listed from the Gulf by Chace (1956) and from Florida by Wass 

(1955) and Lyons et al. (1971). Williams (1965) reported ovigerous females 

from Florida in March to August and from North Carolina in February. Pearse 

and Williams (1951) collected this crab from sponges in North Carolina waters.


Pilumnus gemmalus Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 214) 

Rathbun, 1&30, p. 513, pi. 207, figs. 1-3; Rathbun, 1933, p. 72. 

Range: Dry Tortugas; Culebra; Virgin Islands; Curagao. 


Habitat: shallow water lagoons; coral and rock bottoms; in corals; on seavvoods. 

Pilumnus holosericus Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 

4: 268) 

Rathbun, 1930, p. 519, text-fig. 81, pi. 207, figs. 8-9; Rathbun, 1933, p. 73, fig. 61. 

Range: Bahamas; Dry Tortugas; Puerto Rico; Virgin Islands; Trinidad; 

Curasao. 


Habitat: near shore, under stones; coral reefs; rocky areas. 

PiZMmnMsZwclews Stimpson, 1871 (Bull. Mus. Comp.Zool. 2: 142) 


Williams, 1965, p. 180, figs. 157E, 161. 

Range: North and South Carolina; Florida Keys and Dry Tortugas; west 

coast of Florida; north coast of Cuba. 

Depth: low tide mark to 15 m (8 fm). 

Habitat: sand, shell, rock, coral, and mud substrates; among sponges and seaweed; 

under stones; on buoys and pilings; in beds of Thalassia. 

Remarks: Williams (1965) listed records of ovigerous females; Limz (1937) 

provided notes on ecology of South Carolina populations, listed from Florida by 

Wass (1955), Tabb and Manning (1961) and Lyons gZ aZ. (1971). Rouse (1970) 

found this crab to be the most common pilumnid collected in Florida Bay. 

Pilumnus longleyiRathhun, 1930 (Bull. U.S. Nat. Mus. 152: 502) 


Range: Bahamas; Florida Keys and Dry Tortugas. 


Habitat: rocky and coral bottoms. 

Remarks: Rathbun (1930) noted that this species is easily confused with P. 

caribaeus and P. sayi; she compared the three species morphologically and described 

the young of P. longleyi. 

Pilumnus marshi Rathbun, 1901 (Bull. U.S. Fish. Comm. for 1900. vol. 20, pt. 

2, p. 41) 

Rathbun, 1930, p. 499, text-fig. 80; Rathbun, 1933, p. 72. 

Range: Dry Tortugas; Virgin Islands. 


Habitat: coral bottoms. 

Pilumnus pannosus Rathbun, 1896 (Proc. U.S. Nat. Mus. 19: 142) 


p. 181, figs. 157F, 162; Felder, 1973a, p. 64, pi. 9, fig. 12. 

Range: North Carolina; Bahamas; southeast Florida; Florida Keys; west and 

northwest coasts of Florida; Texas; Puerto Rico; Jamaica; Virgin Islands.


Depth: 1 to 16 m (to9fm). 

Habitat: sand, shell, rock, and coral substrates; on jetties and reefs with fouling 

material; with sponges and corals. 

Remarks: Williams (1965) listed records of ovigerous females. Listed from 

Florida by Wass (1955) and from Texas by Leary (1967). Pearse and Williams 

(1951) collected this crab from submerged reefs off North Carolina. 

Pilumnus sayi Rathbun, 1897 (Ann. Inst. Jamaica 1: 15) 

Common Name: Hairy Crab 

Hay & Shore, 1918, p. 440, pi. 35, fig. 4; Kathbun, 1930, p. 484, pi. 200, figs. 1-2, 

pi. 201, figs. 4-7; Kathbun, 1933, p. 71; Williams, 1965, p. 177, figs. 157A-B, 158; 


Range: North Carolina to Georgia; Bahamas; east coast of Florida; Dry Tortugas; 

west and northwest coasts of Florida; Louisiana and Texas; Jamaica; 

Puerto Rico; Guadeloupe; Curasao. 


Habitat: sand, shell, rock, coral, and gravel bottoms; on offshore reefs; among 

fouling material on jetties, pilings, and buoys. 

Remarks: Listed from Florida by Wass (1955), Dragovich and Kelly (1964), 

Abele (1970), Menzel (1971) and Lyons et al. (1971), the latter group remarking 

on the considerable variation in morphology of this species. Lunz (1939) 

reported collecting P. marshi off a shoal in North Carolina, but Williams (1965) 

believed this to be an aberrant P. sayi. Pearse (1934) reported this crab from inside 

a sponge. Collected by Chace (1956) in the northeastern Gulf. Chamberlain 

(1961) studied the physiological ecology of larval and megalops stages. 

Pilumnus spinosissimûs Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. Iowa 

4: 265) 

Kathbun, 1930, p. 494, text-fig. 79, pi. 200, figs. 7-8. 

Range: Florida Keys and Dry Tortugas. 

Depth: 5 to 11 m (3 to 6 fm). 

Habitat: from rocks, shell, and coral substrates. 

Platyactaea Guinot, 1967 

Platyactaea setigera (H. Milne Edwards, 1834) (Hist. Nat. Crust., vol. 1, p. 390) 

As Actaea setigera—Kathbun, 1930, p. 251, pi. 103; Kathbun, 1933, p. 55. 

As Platyactaea setigera—Guinot, 1967b, p. 561, fig. 36. 

Range: Bermuda; Bahamas; Florida Keys and Dry Tortugas; north coast of 

Cuba; Jamaica; Puerto Rico; Virgin Islands; Antigua; Barbados; Trinidad; 

Curasao; Caribbean coast of Colombia. 

Depth: intertidal and shallow water. 

Habitat: coral reefs; among and under rocks. 

Remarks: Guinot (1967b) placed this species in a new genus and discussed the 

systeniatic affinities of related species.


Platypodia spectahilis (Herbst, 1794). 

Platypodia Bell, 1835 

This species was transferred to a new genus, Platypodiella, by Guinot (1967b). 

leaving this genus without a representative in the Gulf of Mexico. 

Platypodiella Guinot, 1967 

Platypodiella spectahilis (Herbst. 1794) (Natur. Krabben u. Krebse, vol. 2, p. 

153) 

As Platypodia spectabUis—Rathhun, 1930, p. 247, text-fig. 39, pi. 102, fig. 4; 

Ralhbun, 1933, p. 54, fig. 47. 

As Platypodiella spectabilis—Guinot, 1967b, p. 562; Felder, 1973a, p. 65, pi. 9, 

fig. 10. 

Range: Bermuda; Bahamas; Florida Keys; Texas; Veracruz, Mexico, Jamaica; 

Puerto Rico; Virgin Islands; Guadeloupe; Martinique; Barbados; Curagao; Fernando 

de Noronha, Brazil. 

Depth: 4 to 13 m (2 to 7.5 fm). 

Habitat: coral reefs; under stones. 

Remarks: This species is illustrated by Forest & Guinot (1966, fig. 7). Listed 

from Brazil and described by Fausto Filho (1974). Guinot (1967b) transferred 

this species from Platypodia to the newly-erected genus. 

Pseudomedaeus Guinot, 1967 

Pseudoniedaeus agassizii (A. Milne Edwards, 1880) (Crust. Reg. Mex., p. 270) 

As Leptodius agassizii—Hay & Shore, 1918, p. 441, pi. 34, fig. 6; Rathbun, 1930, 

p. 307, pi. 141, figs. 4-5; Williams, 1965, p. 192, figs. 174,183H. 

As Pseudomedaeus agassizii—Guinot, 1968a, p. 726, fig. 25; Felder, 1973a, p. 67, 

pi. 9, fig. 11. 

Range: North Carolina; Florida Straits and Keys; Dry Tortugas; west and 

northwest coasts of Florida; Louisiana and Texas; Virgin Islands. 

Depth: 7 to 82 m (4 to 45 fm). 

Habitat: sand, shell, rock, and coral bottoms; from sponges. 

Remarks: Costlow and Bookhout (1968a) described larval development. Williams 

(1965) noted morphological variability over the range of this species and 

reported ovigerous females from April to November. Listed from Florida by 

Wass (1955) andAbele (1970). Abele (1970) commented on variation in anteriolateral 

teeth and tuberculation of the chelae. 

Rhithropanopeus Rathbun, 1898 

Rhithropanopeus harrisii (Gould, 1841) (Rept. Invert. Massachusetts, p. 326) 


figs. 7-8; Williams, 1965, p. 187, figs. 169, 183C; Christiansen, 1969, p. 81, fig. 33, 

map 27; Felder, 1973a, p. 67, pi. 9, fig. 14. 

Range: New Brunswick to south Florida; west and northwest coasts of Florida;


Mississippi to Veracruz, Mexico; introduced to west coast of United States and to 

various parts of Europe in recent times. 

Depth: intertidal to 37 m (to 20 fm), most common to 9 m (5 fm). 

Habitat: freshwater and estuarine areas; upper bays with salinities usually 

less than 20 ppt; on 03rster reefs; on living and dead vegetation; under rocks and 

in old cans and other debris. 

Remarks: Ryan (1956) provided an account of the life history of this crab, 

based on studies in Chesapeake Bay. Williams (1965) reviewed much of the 

pertinent literature on this species. Studies of larval forms include those of Connolly 

(1925), Hood (1962), Chamberlain (1962), Costlow (1966), Costlow, 

Bookhout and Monroe (1966), Bookhout et al (1972), Ott and Forward (1976), 

andSandifer (1973). 

The European populations were considered a separate subspecies, R. h. trldentaius 

Maitland, by Buitendijk and Holthuis (1949), but this was questioned by 

Wolff (1954). Christiansen (1969) reviewed the distributional data for this 

species in Europe and treated the different populations as conspecific, with no 

distinct subspecies recognized. Reports of this crab on the U.S. Pacific coast include 

Jones (1940), Felice (1958), and Ricketts and Calvin (1968). Regional 

lists include Florida (Wass, 1955; Tabb and Manning, 1961; Abele, 1970; Menzel, 

1971), Mississippi (Christmas andLangley, 1973), Louisiana (Behre, 1950), 

Texas (Hedgpeth, 1953; Leary, 1967), and the northwestern Gulf (Fotheringham 

and Brunenmeister, 1975). 

Physiological studies include data on osmoregulation (Jones, 1941; Verway, 

1957; Kalber and Costlow, 1966; R. I. Smith, 1967), effects of pesticides on larval 

development (Bookhout and Costlow, 1976), metabolism and larval development 

(Rosenberg and Costlow, 1976), effects of juvenile hormone on larvae (Christiansen, 

Costlow and Monroe, 1977a, b), sterol synthesis in larvae (Whitney, 1969), 

androgen glands (Payen, Costlow and Charniaux-Cotton, 1971), and eyestalk 

hormones (Skorkowski, 1972). 

Tetraxanlhus Rathbun, 1898 

Tetraxanthus bidenlalus (A. Milne Edwards, 1880) (Crust. Reg. Mex., p. 353) 

As T. rugosus Rathbun—Rathbun, 1930, p. 459, pi. 185. 

As T. bidenlatus—Chace, 1939, p. 52; Chace, 1940, p. 36. 

Range: Florida Keys; north and south coasts of Cuba; Grenada. 

Depth: 168to293m (92to 160fm). 


Remarks: The species described and illustrated by Rathbun (1930) as T. bldentatus 

was actually a new species, which Chace (1939) named T. rathbunae 

(= T. rathbuni). The species Rathbun (1930) listed as T. rugosus n. sp. is T. 

bldentatus. 

Tetraxanlhus rathbunae Chace, 1939 (Mem. Soc. Cubana Hist. Nat. 13: 52) 

As T. bldentatus (A. Milne Edwards)—Rathbun, 1930, p. 458, pi. 184. 

As T. rathbunae—Chace, 1939, p. 52; Chace, 1940, p. 37. 

As T. rathbuni—Pequenat, 1970, p. 195.

110 iMwrence W. Powers 

Range: North Carolina; southeast Florida; Florida Keys; west coast of Florida; 

north and south coasts of Cuba; off Mississippi; Campeche, off Yucatan; off 

Grenda; off Cape Frio, Brazil. 

Depth: 108 to 476 m (59 to 260 fm). 

Habitat: mud, coral, shell, sand, and rock bottoms. 

Remarks: This is the species described by Rathbun (1930) as T. bidentatus 

(A. Milne Edwards), but it is not Xanthodes bidentatus A. Milne Edwards, the 

original designation of the species Tetraxanthus bidentatus. Chace (1956, p. 19) 

listed this species from the Gulf of Mexico. 

Tetraxanthus rugosus Rathbun, 1930. 

This is a junior synonym of Xanthodes bidentatus {= Tetraxanthus bidentatus), 

therefore Chace (1939, p. 52) substituted T. rathbunae for Rathbun's T. 

bidentatus and T. rugosus is synonymized with T. bidentatus (A. Milne 

Edwards). 

Xantho Leach, 1814 

Xantho denticulata White, 1847 (List Crustacea British Mus., p. 17) 

As Xanlhodius denticulatus—Rathbun, 1930, p. 314, pi. 314, pi. 145, fig. 1, pi. 146. 

As Xantho denticulata—Monod, 1956, p. 280, figs. 335-339; Forest c& Guinot, 

1961, p. 60, fig. 51. 

Range: Bermuda; Bahamas; Florida Keys and Dry Tortugas; northwest 

Florida; Jamaica; Puerto Rico; Virgin Islands; Antigua; Barbados; Colon, 

Panama; Curagao; Trinidad; Pemambuco to Abrolhos Islands, Brazil, Gulf of 

Guinea, west coast of Africa. 


Habitat: tide pools; coral reefs; under rocks. 

Remarks: Guinot (1968a, p. 711) commented on the relationships between 

species of this genus and other related genera, but she did not discuss X. denticulata, 

other than to raise the question of differences between specimens from 

west Africa and South America. Monod (1956) compared this crab with species 

of other genera, especially Cycloxanthops. Listed from northwest Florida by 

Abele (1970). 

Xanlhodius Stimpson, 1859 

Xanlhodius denticulatus (White, 1847). 

Monod (1956) listed synonymies. This species was transferred to the genus 

Xanthodius by Rathbun (1930, p. 314); other authors have continued to treat 

it as a species of Xantho. Not Xantho denticulata Stimpson, 1860, a west coast 

species (= Xanthodius stimpsoni (A. Milne Edwards, 1879) proposed by Rathbun, 

1930). 

Family GERYONIDAE Colosi, 1924 

(The systematic status of this family and of the Goneplacidae is still 

subject to revision by Guinot and others. Balss (1957) placed this genus


between the Xanthidae and Carcinoplacinae as a link to the Goneplacidae. 

Bouvier (1940) included Geryon in the Xanthidae, but Rathbun (1937), 

Sakai (1939) and Barnard (1950) treated it as a genus of the Goneplacidae. 

In the present list, only the type genus, Geryon, is listed in the 

family; Bathyplax is retained in the Goneplacidae until further studies 

are available (refer to Guinot, 1969a, 1971).) 

Geryon Kreyer, 183 7 

Geryon quinquedens Smith, 1879 (Trans. Connecticut Acad. Arts Sci. 5: 35) 

Common Name: Deep Sea Red Crab 

Rathbun, 1937, p. 271, not pis. 85-86; Chace, 1940, p. 38; Pequegnat, 1970, p. 189, 

fig. 6-7; Wigley, Theroux & Murray, 1975, p. 1. 

Range: Nova Scotia to South Carolina; east coast of Florida; Florida Straits; 

off Key West; Dry Tortugas; off Alabama and Texas; off northeast Mexico; 

north coast of Cuba; off Cape Frio, Brazil. 

Depth: 40 to 2153 m (22 to 1178 fm), most common at 320 to 914 m (175 to 

500 fm). 

Habitat: primarily on mud and ooze substrates, occasionally on sand or shellmud 

bottoms. 

Remarks: Chace (1940) compared this species with G. affinis, which is very 

similar in appearance. Plates 85 and 86 in Rathbun (1937) are of G. affinis. 

This latter species has also been caught off the coast of Florida, but it is more 

common in southern areas of the Atlantic and in the Indian Ocean. Le Loeuff 

et al. (1974) recently reported G. quinquedens from the Ivory Coast of Africa. 

When alive, there are color differences between these two species (Schroeder, 

1959). Leone (1951) used serological techniques to investigate the taxonomic 

status of this crab. Musick and McEachran (1972) reported it from depths of 

168 m in Chesapeake Bight. Accounts of the red crab fishery include those of 

Schroeder (1959), McRae (1961), Meade and Gray (1973), Holmsen and Mc 

Allister (1974), Ganz and Herrmann (1975), and Wigley, Theroux and Murray 

(1975). The latter study contains details of substrate and temperature data, 

a review of the life history of this crab, and in situ bottom photographs of the 

animal. Gray (1969) also provided an account of the biology of this species. 

Haefner and Musick (1974) reported its occurrence in Norfolk Canyon. Pequegnat 

(1970) indicated a center of depth range at about 914 m (500 fm) 

within the Gulf of Mexico, but all of the crabs taken by the R/V Alaminos at 

depths greater than 1170 m (640 fm) were juveniles. Haefner (1977) investigated 

reproduction in females. 

Family GONEPLACIDAE Macleay, 1838 (sensuBalss, 1957) 

(Guinot (1969a) has proposed several revisions in the systematic relationships 

of this family, most of them based on presumed affinities with 

genera of the Xanthidae, from which the Goneplacidae may have been 

derived. In the present list, the Gulf genera are listed alphabetically, as


they were for the Xanthidae, without regard to subfamilj^ alignment. The 

genus Geryon is treated under a separate family, Geryonidae.) 

Bathyplax A. Milne Edwards, 1880 

Balhyplax lyphla A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 16) 

Rathbun, 1918, p. 19, (ext-fig. 4, pi. 2; Rathbun, 1933, p. 77, fig. 67; Chace, 1940, 

p. 43; Williams, McCloskey & Gray, 1968, p. 52, fig. 8; Pequegnat, 1970, p. 192, 

figs. 6-9,6-10. 

Range: off North Carolina; west coast of Florida; Mississippi; Texas and 

Mexico; west coast of Cuba; St. Croix; St. Lucia; off Recife, Brazil. 

Depth: 402 to 878 m (220 to 480 fm); at 1106 m (605 fm) off Cuba. Pequegnat 

(1970) determined the highest densities of crabs at 512 m (280 fm) in the 

Gulf of Mexico. 

Habitat; mainly from muddy substrates; also from coral bottoms. Pequegnat 

(1970) reported that blackened specimens were commonly collected, apparently 

due to contact with natural oil seepage. 

Remarks: Recorded from the Gulf of Chace (1956). Pequegnat (1970) considers 

this crab to be the most common deep-water species in the Gulf of Mexico; 

ovigerous females were collected in August, November, and December. Guinot 

(1971) listed this species under the Xanthidae and under the Goneplacidae 

because of the uncertainty of its systematic affinities. 

Chacellus Guinot, 1969 

Chacellus filiformis Guinot, 1969 (Bull. Mus. Nation. Hist. Nat. 41: 722) 

Guinot, 1969, p. 722, figs. 135-136, pi. V, fig. 4. 

Range: between Bahamas and east coast of Florida; off northwest Florida. 

Depth: 183to223m(100tol22fm). 

Chasmocarcinus Rathbun, 1898 

Chasmocarcinus cylindricus Rathbun, 1901 (Bull. U.S. Fish. Comm. for 1900, 

vol.2: 10) 

Rathbun, 1918, p. 59, text-figs. 28-29; Rathbun, 1933, p. 80, fig. 73; Chace, 1940, 

p. 49; Pequegnat, 1970, p. 195. 

Range: Mississippi; off Louisiana; Campeche, off Yucatan; north and south 

coasts of Cuba; Jamaica; Puerto Rico. 

Depth: 13 to 1906m (7 to 1075 fm). 

Habitat: mud bottoms; sand, rock, and coral substrates. 

Chasmocarcinus mississippiensis Rathbun, 1931 (Proc. Biol. Soc. Washington 

44: 71) 

Rathbun, 1931b, p. 71; Felder, 1973a, p. 70, pi. 10, fig. 2. 

Range: off coasts of Mississippi, Louisiana, and Texas. 

Depth: 4 to 91 m (2to50fm). 

Habitat; sand and mud bottoms.


Remarks: Dawson (1966) reported this species from off Grand Isle, Louisiana; 

Franks et al. (1972) obtained a single specimen from off Mississippi at 50 

fm. Felder (1973a) reported specimens from Padre Island, Texas. 

Chasmocarcinus ohliquus Rathbun, 1898 (Bull. Lab. Nat. Hist. State Univ. 

Iowa 4: 286) 

Rathbun, 1918, p. 58, text-fig. 27, pi. 14, figs. 1-2; Chace, 1940, p. 48. 

Range: southeast of Bahamas; north and south coasts of Cuba. 

Depth: 177 to 503 m (97 to 275 fm). 

Habitat: mud and ooze substrates. 

Eucratopsis Smith, 1869 

Eucratopsis crassimanus (Dana, 1852) (Proc. Acad. Nat. Sci. Philadelphia, for 

1851, vol. 5:248) 

Rathbun, 1918, p. 52, text-fig. 22, pi. 12, fig. 3, pi. 159, figs. 1-2; Guinot, 1969a, 

p. 258, figs. 6, 10, 25. 

Range: Florida Keys; south and west coasts of Florida; Yucatan; Jamaica; 

Bahia to Rio de Janeiro, Brazil. 

Depth: shallow water to 14 m (to 7.5 fm). 

Habitat: sand, coral, and broken shell substrates. 

Remarks: Tabb and Manning (1961) collected ovigerous females in October 

from Oyster Bay in south Florida. 

Euphrosynoplax Guinot, 1969 

Euphrosynoplax clausa Guinot, 1969 (Bull. Mus. Nation. Hist. Nat. 41: 720) 

Guinot, 1969a, p. 720, figs. 127, 139, pi. IV, fig. 3; Pequegnat, 1970, p. 194. 

Range: Dry Tortugas; off Alabama and Mississippi; Campeche, Yucatan. 

Depth: 91 to210m (50to 115fm). 

Euryplax Stimpson, 1859 

Euryplax nitida Stimpson, 1859 (Ann. Lye. Nat. Hist. New York 7: 60) 

Ratlibun, 1918, p. 34, pi. 7; Rathbun, 1933, p. 78, fig. 69; Williams, 1965, p. 202, 

fig. 185; Guinot, 1969a, p. 512, figs. 39, 41, 47, 56-57, pi. II, fig. 1; Felder, 1973a, 

p. 70, pi. 10, fig. 1. 

Range: off North Carolina; Bermuda; Florida Keys; south and west coasts of 

Florida to Texas; Puerto Rico; St. Thomas, Virgin Islands. 

Depth: shallow water to 90 m (to49fm). 

Habitat: sand, shell, rock, and coral substrates; sandy grass flats. 

Remarks: Menzel (1971) listed this crab as rare at Apalachee Bay and Abele 

(1970) collected only a single specimen in his study of the northeastern nearshore 

Gulf. Range reports for the western Gulf of Mexico are scant and questionable. 

Rathbun (1918) listed one specimen from New Orleans and Williams 

(1965) indicated a range extending to Texas. Felder (1973a) cites these reports 

but adds no new records.


Frevillea A. Milne Edwards, 1880 

Frevillea barbala A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 8: 15) 

As Goneplax barhata—Rathbun, 1918, p. 26, pi. 4, figs. 1, 3, pi. 5. 

As Frevillea barbala—Guinot, 1969a, p. 513, pi. II, fig. 2. 

Range: west coast of Florida; Yucatan (Gulf); north coast of Cuba; off 

Grenada. 

Depth: 55 to 168 m (30 to 92 fm). 

Habitat: sand, broken coral bottoms. 

Frevillea hirsuta (Borradaile, 1916) (Brit. Antarctic Exped., 1910, Zool., vol. 3, 

no. 2, p. 99) 

As Goneplax ftirsuta—Rathbun, 1918, p. 28, text-fig. 7; Williams, 1965, p. 201, 

fig. 184. 

As Frevillea hirsuta—Guinot, 1969a, p. 513, te.Yt-figs. 33, 40, 58-59, pi. II. fig. 3. 

Range: off North Carolina; off west and northwest Florida and Alabama; north 

of Yucatan; off Rio de Janeiro, Brazil. 

Depth: 73 to 146 m (40 to 80 fm). 

Remarks: Collected by the R/V Oregon from the Gulf of Mexico (Chace, 

1956). 

Frevillea tridenlalaA. Milne Edwards, 1880. 

Transferred to the genus Trapezioplax by Guinot (1969a). Trapezioplax 

tridentata. 

Glyploplax Smith, 1870 

Glyploplax smithii A. Milne Edwards, 1880 (Crust. Reg. Mex., p. 336) 

Rathbun, 1918, p. 51, pi. 13, figs. 3-4; pi. 158, figs. 7-10; Milne Edwards & Bouvier, 

1923, p. 328, pi. 5, fig. 5; Williams, McCIoskey & Gray, 1968, p. 55, fig. 11; Guinot, 

1969a, p. 259. 

Range: Bermuda; off North Carolina; west coast of Florida; Cape Catoche, 

Yucatan (Gulf coast). 

Depth: 24 to 55 m (13 to 30 fm). 

Habitat: sand, coral, gravel and rock bottoms. 

Remarks: Guinot (1969a) believed that this species should be excluded from 

the genus, based on differences from the type species, G. pugnax, a Pacific form 

from Central America. 

Goneplax Leach, 1814 

(All three of the recognized Gulf species of this genus have been transferred 

to other genera by Guinot (1969a). Goneplax harhata and G. hirsuta 

are referred to Frevillea and G. tridentata is referred to Trapezioplax.) 

Neopilumnoplax Serene, 1969 

Neopilumnoplax americana (Rathbun, 1898) (Bull. Lab. Nat. Hist. State Univ. 

Iowa 4: 283) 

As Pilumnoplax americana—Rathbun, 1918, p. 21, text-figs. 5-6; Williams,

McCloskey & Gray, 1968, p. 52, fig. 9. 


As Pieopilumnoplax americana—Guinot, 1969a, p. 689, figs. 83-84. 

Range: off North Carolina and Georgia; Florida Keys and Straits; north coast 

of Cuba; Guadeloupe; Espirito Santo, Brazil; Arabian Sea. 

Depth: 128 to 805 m (70 to 440 fm). 

Habitat: sand, shell, coral, and rocky substrates. 

Remarks: Chace (1940) recovered this crab from the stomach of a smooth 

dogfish {Muslelus canis) from off Havana, Cuba. Listed from Brazil by Rodrigues 

da Costa (1968a) and Coelho and Ramos (1972). 

Panoplax Stimpson, 1871 

Panoplax depressa Stimpson, 1871 (Bull. Mus. Comp. Zool. 2: 151) 

Rathbun, 1918, p. 47, text-fig. 21, pi. 12, figs. 1-2; Rathbun, 1933, p. 80, fig. 72; 

Guinot, 1969a, p. 264, figs. 3, 12, 28, ? 29; Bright & Pequegnat, 1974, p. 33. 

Range: Dry Tortugas; west coast of Florida; West Flower Garden Bank, off 

Texas; north coast of Cuba; north coast of Yucatan; Puerto Rico. 


Habitat: sand, coral, and broken shell bottoms. 

Remarks: Listed by Chace (1956) from off the west coast of Florida and by 

Bright and Pequegnat (1974) from silty-sand bottoms at West Flower Garden 

coral reef, at 330 foot depth. 

Pilumnoplax Stimpson, 1858 

Pilumnoplax a/nericawa Rathbun, 1898. 

Transferred to the genus Neopilumnoplax by Guinot (1969a). See Neopilumnoplax 

americana. 

Pilumnoplax data (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 18) 

As Eucratoplax elata—A. Milne Edwards, 1880, p. 18 (original type description, 

female holotype in Paris Museum, type locality is West Florida, 13 fms.). Not 

Rathbun, 1898, p. 281. 

As Pilumnoplax c/nla—Guinot, 1969a, p. 688. Not Rathbun, 1918, p. 23. 

Range: West Florida, tjqje locality. 

Depth: 24m (13 fm). 

Remarks: Only the original description of A. Milne Edwards (1880) is valid 

for this species. All of the other material described by Rathbun (1918, p. 23) has 

been referred by Guinot (1969a, p. 688, 716-717) to other genra. The male 

specimen description of Rathbun (1918, p. 23) was referred to Robertsella 

mystica (Guinot, 1969a, p. 716) and the females and juveniles were referred to 

Thalassoplax angusta (Guinot, 1969a, p. 717). This leaves the original type specimen 

to represent this poorly known species in the Gulf of Mexico. 

Pilutnnoplax nilida Chace, 1940 (Torreia3; 44) 

Chace, 1940, p. 44, figs. 17-18; Guinot, 1969a, p. 689. 


Depth: 348to476m( 190to260fm).

116 hawrence W. Powers 

Remarks: Guinot (1969a) retained this species in the genus, but she commented 

on the obscurity of its relationships. 

Prionoplax allanlica Kendall, 1891. 

Prionoplax H. Milne Edwards, 1852 

Referred to Frevillea tridentata by Guinot (1969a), who later decided that 

F. tridentata was different enough from other Frevillea to establish a new genus, 

Trapezioplax, for this species. See Trapezioplax tridentata. 

Rober(sella Guinot, 1969 

Roherlsella mystica Guinot, 1969 (Bull. Mus. Nation. Hist. Nat. 41: 716) 

As Pilumnoplax data in Rathbun (not A. MiJne Edwards)—Rathbun, 1918, p. 23 

(part, male description only), pi. 3 (part, male only). 

As Robertsella mystica—Guinot, 1969a, p. 716, figs. 132-133, pi. V, fig. 4. 

Range: off southeast coast of Florida, Florida Straits. 

Depth: 353 m (193 fm). 

Habitat: sand bottom. 

Remarks: Although this species has not been recorded from the Gulf of 

Mexico, it is included here because of the confusing nomenclatural history of 

Pilumnoplax elata. At present, this new species and genus includes only the 

mature male specimen described by Rathbun (1918, p. 23 and part of pi. 3). 

Speocarcinus Stimpson, 1859 

Speocarcinus carolineiisis Stimpson, 1859. 

This species was reported from the Gulf of Mexico, prior to revision by Guinot 

(1969a, p. 710), who referred the Gulf specimens to S. lobatus. The Carolinean 

specimens described in Williams (1965) are S. carolinensis. 

Speocarcinus lobatus Guinot, 1969 (Bull. Mus. Nation, Hist. Nat. 41: 710) 

As S. caroliiiensis—Rathbun, 1918, p. 39 (part, specimen from Dry Tortugas 

only). 

As S. /o6aJ«s—Guinot, 1969a, p. 710, text-figs. 124-125, pi. IV, fig. 2; Felder 

1973a,p. 70,pl.l0, fig. 3. 

Range: Dry Tortugas; off Louisiana and Texas. 


Habitat: probably inhabits burrows of polychaetes and crustaceans as does 

S. carolinensis. 

Remarks: Dawson (1966) reported S. carolinensis from off Grand Isle, Louisiana 

and Felder (1973a) reported that specimens taken from that same area at 

later dates were S. lobatus. 

Telraplax Rathbun, 1901 

Telraplax quadridenlata (Rathbun, 1898) ((Bull. Lab. Nat. Hist. State Univ. 

Iowa 4: 287) 

Rathbun, 1918, p. 32, text-figs. 9-10, pi. 6, figs. 3-4; Rathbun, 1933, p. 78, fig. 68; 

Guinot, 1969a, p. 256, figs. 1,14, 26.

Range: north coast of Cuba; Puerto Rico, Curasao. 

Depth: 8 to 22 m (4.5 to 12 fm). 


Thalassoplax Guinot, 1969 


Thalassoplax angusla Guinot, 1969 (Bull. Mus. Nation. Hist. Nat. 41: 717) 

As Pilumnoplax elala in Rathbun (not A. Milne Edwards)—Rathbun, 1918, p. 23 

(part, female and juveniles, includes female on pi. 3). 

As Thalassoplax angusla—Guinot, 1969a, p. 717, figs. 131-132, pi. IV, fig. 2; 


Range: east coast of Florida; off northwest Florida, Alabama and Mississippi; 

off east coast of Mexico; off Campeche, Yucatan. 

Depth: 183 to 752m (100 to411 fm). 

Habitat: mud, sand, broken shell bottoms. 

Remarks: Pequegnat (1970) added several new records for the Gulf of Mexico 

and also commented on differences in morphological descriptions between the 

Alaminos specimens and that provided \)y Guinot (1969a, p. 717). The legends 

on Guinot's plates were transposed; the correct citations appear above and in 

Pequegnat (1970, p. 192). As previously discussed for Robertsella rnystica, 

Rathbun's (1918, p. 23) description of Pilumnoplax elata does not conform to the 

type description of this species provided by A. Milne Edwards; thus all of the 

material included in Rathbun was transferred to new species and genera by 

Guinot (1969). Thalassoplax angusta includes the females and juveniles described 

by Rathbun, but not the mature male nor the original female holotjqDe 

of P. elala. 

Trapesioplax Guinot, 1969 

Trapesioplax Iridenlala (A. Milne Edwards, 1880) (Bui. Mus. Comp. Zool. 8: 

16) 

As Goneplax Iridenlala—Rathbun, 1918, p. 29. 

As Prionoplax allanlica—Rathbun, 1918, p. 30, text-fig. 8, pi. 6, figs. 1-2. 

As Trapezioplax Iridenlala—Guinot, 1969a, p. 713, figs. 128-129,142. 

Range: Florida Keys and Dry Tortugas; west coast of Florida; Barbados. 

Depth: 13 to42 m (7to23fm). 

Habitat: sand, coral, shell, and mud bottoms. 

Remarks: Guinot (1969a) established this new genus based on a separation 

of Frevillea tridentata from the other species of Frevillea. Trapezioplax tridentata 

includes the original type, Goneplax tridentata, and Prionoplax atlantica, listed 

as separate species by Rathbun (1918). 

Family PALICIDAE Bouvier, 1898 (= CYMOPOLIDAE Faxon, 1895) 

PaKcMs Philippi, 1838 

Palicus affinis A. Milne Edwards & Bouvier, 1899 (Bull. Mus. Hist. Nat. Paris 

5: 122)

118 'Lawrence W. Powers 

As CymopoUa affinis—TKathhuTi, 1918, p. 196, text-fig. 121, pi. 46, pi. 47, fig. 3; 

Rathbun, 1933, p. 85. 

Range: southeast and west coasts of Florida; Dry Tortugas; Virgin Islands; 

Barbados; Guianas to Espirito Santo, Brazil. 

Depth: 33to214m (18toll7fm). 

Habitat: sand, shell, and coral substrates. 

Remarks: Listed from Brazil by Rodrigues da Costa (1968a), Coelho (1971c), 

and Coelho and Ramos (1972). 

Palicus alternaliis Rathbun, 1897 (Proc. Biol. Soc. Washington 11: 95) 

As CymopoUa nflernaln—Ratlibun, 1918, p. 188, text-fig. 117, pis. 42-43. 

As Palicus alternatus—Williams, 1965, p. 215, fig. 200. 

Range: North Carolina; Florida Keys; west and northwest coasts of Florida. 

Depth: 7 to 110 m (4 to 60 fm). 

Habitat: sand, gravel, broken shell, coral, and sand-mud bottoms. 

Remarks: Ovigerous females are known from Florida during January to 

August and from North Carolina in October (Williams, 1965). 

Palicus cursor (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 29) 

As CymopoUa cursor—Rathbun, 1918, p. 215, text-figs. 130-131, pi. 52, figs. 1-2; 

Chace, 1940, p. 50. 

Range: North Carolina; Florida Keys; northwest coast of Florida; north coast 

of Cuba; St. Christopher; Dominica; Barbados. 

Depth: 206to530m( 107to290fm). 

Habitat: sand ooze, sand-mud, sand, and broken shell bottoms. 


in October, from Florida in March, and from the Antilles in January-February. 

Palicus denlalus (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 28) 

As CymopoUa dentata—Rathbun, 1918, p. 202, text-fig. 124. 

As PaUcus denlalus—Pequegnat, 1970, p. 197. 

Range: Florida Keys; west coast of Florida; off Alabama; off Barbados. 

Depth: 27 to 139 m (15 to 76 fm). 


Palicus faxoni Rathbun, 1897 (Proc. Biol. Soc. Washington 11: 96) 

As CymopoUa faxoni—'Rathhuit, 1918, p. 194, text-fig. 120, pi. 45,2-3. 

As PaUcus /a-voni—Williams, 1965, p. 216, fig. 201. 

Range: North Carolina; east coast of Florida; northeast of Yucatan (Gulf); ? 

off Cape Frio, Brazil. 

Depth: 59 to 93 m (32 to 51 fm). 

Habitat: sand substrates. 

Palicus gracilipes (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 29) 

As CymopoUa graciUpes—Ratlibun, 1918, p. 221, text-fig. 133, pi. 52, figs. 3-4; 

Chace, 1940, p. 51. 

Range: Bahamas; north of Yucatan; north coast of Cuba.

Depth: 112 to 545 m (61 to 298 fm). 

Habitat: sand and shell bottoms. 

Palicus gracilis (Smith, 1883) (Proc. U.S. Nat. Mus. 6: 20) 


As Cvmopolia gracitfs—Rathbun, 1918, p. 218, text-fig. 132, pi. 50, pi. 51, fig. 1; 

Chace, 1940, p. 50. 

As Palicus gracilis—Pequegnat, 1970, p. 195, fig. 6-11. 

Range: off Massachusetts; east coast of Florida; northwest Florida; Louisiana 

to central east coast of Mexico; north coast of Cuba; Curasao. 

Depth: 183 to 686m (100 to 375 fm). 

Habitat: fine sand and mud substrates. 

Remarks: Chace (1956) reported this species in the Gulf of Mexico and 

Pequegnat (1970) reported ovigerous females from the same area in August and 

November. He further noted that this crab may be able to swim, but is probably 

not pelagic. 

Palicus obesus (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 27) 

As Cymopolia obesa—Ralhbun, 1918, p. 205, text-fig. 125, pi. 49. 

As Palicus obesus—Pequegnat, 1970, p. 197. 

Range: off northwest Florida and Mississippi; Campeche, Mexico. 

Depth: 24 to 220 m (13 to 120 fm). 

Remarks: Collected by the R/V Oregon from the northeastern Gulf of Mexico 

(Chace, 1956). 

Palicus sica (A. Milne Edwards, 1880) (Bull. Mus. Comp. Zool. 8: 29) 

As Cymopolia sico—Rathbuii, 1918, p. 208, text-fig. 127, pi. 40, figs. 3-4; Rathbun, 

1933, p. 85, fig. 78; Chace, 1940, p. 49. 

As Palicus sicus—Milne Edwards & Bouvier, 1902, p. 56, pi. 10, figs. 7-11, pi. 11, 

fig. 9; Pequegnat, 1970, p. 198. 

Range: Florida Keys and Straits; west coast of Florida; north coast of Cuba; 

Puerto Rico; Virgin Islands; Barbados; Grenada. 

Depth: 27 to 348 m (15 to 190 fm). 

Habitat: sand, mud, shell, and coral bottoms. 

Remarks: Pequegnat (1970) reported ovigerous females from the Gulf of 

Mexico in mid-July. Rathbun (1918) described the colors of freshly-preserved 

specimens. Chace (pers. comm.) notes that the specific name sica is used as a 

noun in opposition (L. = curved dagger) and thus should not be changed due 

to the transfer of the species to the masculine genus Palicus. 

Family PINNOTHERIDAE de Haan, 1833 

Subfamily PINNOTHERINAE de Haan, 1833 

Dissodactylus Smith, 1870 

Dissodactylus alcocki Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 124) 

Rathbun, 1918, p. 124, text-figs. 70-71, pi. 28, figs. 3-4; Schmitt, McCain & 

Davidson, 1973, p. 16.


Range: off delta of Mississippi River. 

Depth: 64m(35fm). 

Habitat: sand-mud bottom. 

Remarks: This species is known only from a female type and a damaged male 

pai'atype. 

Dissodaclylus horradailei Rathbun. 1918 (Bull. U.S. Nat. Mus. 97: 121) 

Rathbun, 1918, p. 121, text-fig. 68, pi. 27, figs. 5-8; Schmitt, McCain & Davidson, 

1973, p. 16. 

Range: off southeast and southwest coasts of Florida; Jamaica. 

Depth: 49 to 55 m (27 to 30 fm). 

Habitat: fine white sand. 

Dissodaclylus caZmani Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 125) 

Rathbun, 1918, p. 125, text-figs. 72-73, pi. 28, figs. 5-6; Schmitt, McCain & 

Davidson, 1973, p. 16. 

Range: east coast of Florida; northwest coast of Cuba. 


Habitat: coral, sand, gravel, and rock bottoms; near shore. 

Dissodaclylus crinilichelis Moreira, 1901 (Arch. Mus. Nac. Rio de Janeiro 11: 

37) 

As D. encopei—Rathbun, 1918, p. 119, text-fig. 67, pi. 27, figs. 1-4; Williams, 

McQoskey & Gray, 1968, p. 56, fig. 12. 

As D. crinilichelis—Rathbun, 1933, p. 83, fig. 76; Schmitt, McCain & Davidson, 

1973, p. 17. 

Range: off North Carolina; northwest coast of Florida; Jamaica; Puerto Rico; 

Belize; Caribbean coast of Colombia; Paraiba to Rio Grande do Sul, Brazil. 


Habitat: fine white sand, coral, and broken shell bottoms; on Halodule (sea 

grass); with the echinoids Encope marginata, E. michelini (sand dollars) and 

Clypeaster subdepressus (sea biscuit). 

Remarks: L. H. Hyman (1955) commented on host relationships with 

echinoids (as D. encopei). Listed from Florida by Wass (1955), Abele (1970), 

and Menzel (1971); listed from Brazil by Coelho and Ramos (1972) and Rodrigues 

da Costa (1971). 

Dissodaclylus encopei Rathbun, 1901. 

A junior synonym of D. crinitichelislsAoTeiTa, 1901. 

Dissodaclylus juvenilis Bouvier, 1917 (Bull. Mus. Nat. Hist. Natur. Paris 23: 

397) 

Milne Edwards & Bouvier, 1923, p. 349, text-figs. 11-12, pi. 9, figs. 3-4; Schmitt, 

McCain & Davidson, 1973, p. 17. 

Range: north of Yucatan, Mexico. 

Dissodaclylus mellilae (Rathbun, 1900) (Amer. Natural. 34: 590) 

Hay & Shore, 1918, p. 444, pi. 36, fig. 1; Rathbun, 1918, p. 117, text-fig. 66, pi. 28,

Crabs of t he Giilf of Mexico 121 

figs. 7-8; Williams, 1965, p. 209, fig. 192; Williams, McCloskey & Gray, 1968, 

p. 37; Rogers, 1968, p. 318; Schmitt, McCain & Davidson, 1973, p. 18. 

Range: Massachusetts to South Carolina; northwest coast of Florida; Texas. 

Depth: 9 to 52 m (5 to 28 fm). 

Habitat: sand and broken shell bottoms; areas of scattered sponges and coral 

heads; with the echinoids Mellita quinquesperforata., Encope micheUni, Echinarachinius 

parma, and Clypeaster subdepressus. 

Remarks: Larval stages have been described by 0. W. Hyman (1924), Lebour 

(1928), Aikawa (1937) and Costlow and Bookhout (1966b). Host relationships 

were described by Johnson (1952), L. H Hyman (1955), Gray (1961), Gray, 

McCloskey and Weihe (1968), and MacGinitie and MacGinitie (1968, p. 314). 

Regional lists include Florida (Wass, 1955; Abele, 1970; Menzel, 19.71). Not 

listed by Felder (1973a) for Texas, but see Rogers (1968) for a report on this 

species at Galveston. 

Dissodiiclylus primilivus Bouvisr, 1917 (Bull. Mus. Nat. Hist. Natur. Paris 23: 

394) 

Milne Edwards & Bouvier, 1923, p. 346, text-fig. 8, pi. 8, figs. 3-4, pi. 9, fig. 1; 

Schmitt, McCain & Davidson, 1973, p. 20. 

Range: west of Tortugas, Florida. 

Remarks: The above location is the orAj known record for this species. 

Dissodactylus stehbingi Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 123) 


1973, p. 20. 

Range: Virginia; west and northwest coasts of Florida. 

Depth: 9to 10m (17fm). 

Habitat: on sea biscuits, Clypeaster subdepressus, in an area of scattered 

sponges and coral heads (northwest Florida). 

Remarks: Wass (1955) and Menzel (1971) provide some ecological notes on 

this crab. 

Fafeifl Dana, 1851 

Fabia byssomiae (Say, 1818) (J. Acad. Nat. Sci. Philadelphia 1: 451) 

Rathbun, 1918, p. 103, text-fig. 36, pi. 24, figs. 6, 8; Schmitt, McCain & Davidson, 

1973, p. 22. 

Range: west coast of Florida; northwest coast of Cuba. 


Habitat: in bivalve molluscs, Hiatella arctica; located on beds of Alcyonium 

and between individuals of aggregating ascidians. 

Fabia leHinfte Cobb, 1973 (Crustaceana25: 70) 

Cobb, 1973, p. 70, figs. 1-2. 

Range: off northwest Florida to Alabama. 

Depth: 5 to 18 m (3tol0fm). 

Habitat: commensal in bivalves, Tellina magna Spengler (females in mantle 

cavit}^, males in excurrent siphon); from sandy bottoms.


Orthotheres Sakai, 1969 

Orthotheres serrei (Rathbun, 1909) (Bull. Mus. Hist. Nat. Paris 2: 69) 

As Pinnolheres serrei—Rathbun, 1918, p. 84, text-fig. 41, pi. 19, figs. 1-7; Rathbun, 

1933, p. 82. 

As Orlholheres serrei—Sakai, 1969, p. 275; Schmitt, McCain & Davidson, 1973, 

p. 27. 

Range: northwest Cuba; Jamaica; Puerto Rico. 

Habitat: at surface; on reef flats; in mantle cavity of Strombus. 

Orthotheres strombi (Rathbun, 1905) (Proc. Acad. Nat. Sci. Philadelphia 1905: 

371) 

As Pinnotheres strombi—Rathbun, 1918, p. 90, text-fig. 45, pi. 20, figs. 1-2. 

As Orthotheres sJromfti—Sakai, 1969, p. 275; Schmitt, McCain & Davidson, 1973, 

p. 27. 

Range: west and northwest coasts of Florida. 

Habitat: commensal in the gastropods Strombus pugilis, S. alatus, and Pleuroploca. 

Remarks: Listed from Florida by Wass (1955), Abele (1970), and Menzel 

(1971). 

Parapinnixa Holmes, 1894 

Parapinnixa bouvieri Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 111) 

Rathbun, 1918, p. Ill, text-fig. 60, pi. 25, figs. 4-10; Rathbun, 1933, p. 83, fig. 75; 

Williams, 1963, p. 208, fig. 191; Schmitt, McCain & Davidson, 1973, p. 31. 

Range: South Carolina; northeast of Yucatan (Gulf); Puerto Rico. 

Depth: 5 to 73 m (3 to 40 fm). 

Habitat: coral and sand bottoms; among ventral spines of a sea urchin. 

Remarks: Williams (1965) lists ovigerous females from Florida and notes the 

association of this crab wdth a sea urchin. 

Parapinnixa hendersoni Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 109) 


1973, p. 32. 

Range: west coast of Florida; northwest Cuba; Curagao; Maranhao to Bahia, 

Brazil. 

Depth: 38 to 55 m (21 to 30 fm). 

Habitat: free-swimming, pelagic (Cuba, in Rathbun, 1918); on sand and 

broken coral bottom in Florida. 

Remarks: Recorded from Brazil by Righi (1967) and Coelho and Ramos 

(1972). 

Pinnaxodes HeWer, 1865 

Pinnaxodes floridensis Wells & Wells, 1961 (Bull. Mar. Sci. Gulf Carib. 11: 

267) 

Weils & Wells, 1961, p. 267, figs. 1-2; Schmitt, McCain & Davidson, 1973, p. 34. 

Range: west and northwest coasts of Florida.


Ilabitat: commensal in the cloaca and respiratory tree of the holothurian, 

Theelothuria princeps (Selenlza), which buries in sand; juvenile crabs are found 

in the anterior digestive tract of the host. 

Remarks: Wells and Wells (1961) provided data on the natural history, 

ecology, and morphology of this crab and Pearce (1966) reviewed the biology 

and host relationships. Abele (1970) listed the angel wing mollusc, Cyrtopleura 

cosiata, as a host for a sexual intermediate form of this crab. Listed from northwest 

Florida by Menzel (1971). 

Pinnotheres Bosc, 1801-1802 

Pinnotheres geddesi Miers, 1880 (J. Linn. Soc. London, Zool. 15: 86) 

Rathbun, 1918, p. 70, text-fig. 32, pi. 16, figs. 1-4; Rathbun, 1933, p. 82; Schmitt, 


Range: Veracruz, Mexico; eastern Cuba (Atlantic); Puerto Rico; ? Jamaica. 

Habitat: commensal in mangrove oysters (? Crassostrea rhizophorae) and 

Ostrea. 

Pinnotheres guerini H. Milne Edwards, 1853 (Ann. Sci. Nat. Zool. Paris 20: 

219) 

Rathbun, 1918, p. 101, text-fig, 52; Rathbun, 1933, p. 83; Schmitt, McCain & Davidson, 

1973, p. 48. 

Range: Cuba (location not specified); Puerto Rico. 

Habitat: reported from oysters. 

Remarks: The location of the type specimen in Cuba is unspecified, thus this 

species may not be present in the Gulf of Mexico. 

Pinnotheres hemphilli Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 99) 

Rathbun, 1918, p. 99, text-fig. 51, pi. 23; Schmitt, McCain & Davidson, 1973, p. 48. 

Range: Cedar Keys, Florida. 

Habitat: intertidal. 

Remarks: Known only from a single type specimen. 

Pinnotheres hirtimanus H. Milne Edwards, 1853 (Ann. Sci. Nat. Zool. Paris 

20: 219) 

Rathbun, 1918, p. 101; Schmitt, McCain & Davidson, 1973, p. 48. 

Range: Cuba, location unspecified. 

Remarks: Known only from the single type specimen. 

Pinnotheres maculatus Say, 1818 (J. Acad. Nat. Sci. Philadelphia 1: 450) 

Common Names: Mussel Crab; Pea Crab 


pi. 17, figs. 3-6; Rathbun, 1933, p. 82, fig. 74; Williams, 1965, p. 206, fig. 190; 

Felder, 1973a, p. 74, pi. 10, figs. 10-11; Schmitt, McCain & Davidson, 1973, p. 53. 

Range: Massachusetts to south Florida; west coast of Florida to Texas; northwest 

Cuba; Jamaica; Puerto Rico; Virgin Islands; Uruguay and Argentina. 

Depth: surface to 50 m (to 27 fm).


Habitat: commensal in a variety of bivalve molluscs; young of both sexes and 

often adult males are free-swimming; most common in the mantle cavities of 

mussels, Mytilus edulis; in tubes of the polychaetes Arenicola and Chaetopterus 

(C. pergamentaceus and C. variopedatus); from mud, sand, shell and gravel 

substrates. Other molluscan hosts include: Atrina rigida, A. seminuda, A. serrata, 

Anomia simplex, Argopecten gibba, A. irradians, Cyrtopleura costata. Modiolus 

modiolus, M. tulipa, Mya arenaria, and Placopecten magellanica. 

Remarks: This species has a large literature, catalogued by Schmitt, McCain 

and Davidson (1973). Larval stages were described by 0. W. Hyman (1924), 

Aikawa (1937) and Costlow and Bookhout (1966b). Life history data is provided 

in MacGinitie and MacGinitie (1968) and by Christensen and McDermott 

(1959). Caine (1975) studied feeding behavior and physiology and Kruczynski 

(1975) measured food intake and digestion. Pearce (1964) described reproductive 

aspects. Larval shadow responses were studied by Forward (1977) and behavior 

in relation to hosts was described by SasUy and Menzel (1962) and by 

Eidemiller (1969). The effects of this crab on the growth and biology of its 

scallop hosts were studied by Kruczynski (1971, 1972). Sandifer (1973) commented 

on larval ecology in Virginia and Fotheringham and Brunenmeister 

(1975) described this crab as it occurs in the Gulf of Mexico. Regional lists include 

Florida (Wass, 1955; Tabb and Manning, 1961; Abele, 1970; Menzel, 

1971; Godcharles and .laap, 1973) and Texas (Leary, 1967). Listed from Brazil 

by Rodrigues da Costa (1971) and Coelho and Ramos (1972). 

Pinnotheres inoseri Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 94) 


1973, p. 58. 


Depth: 1.5to5.5m (lto3fm). 

Habitat: commensal in sea squirts (tunicates) and from the brachial cavity of 

an ascidian, Polycarpa obtecta; off rocky bottoms with grass and thin layers of 

sand and mud. 

Remarks: Rathbun (1918) did not list sea squirts as commensals, only as 

present in the dredges in which the crabs were found. Pearce (1966) provided 

information on life histoiy and Hartnoll (1964a) described a larval stage. Listed 

from Florida by Godcharles and Jaap (1973), including data from the collection 

locality. 

Pinnotheres ostreum Say, 1817 (J. Acad. Nat. Sci. Philadelphia 1: 67) 

Common Names: Oyster Crab; Common Pea Crab 


figs. 3-6; Williams, 1965, p. 203, figs.. 187-189; Felder, 1973a, p. 75, pi. 10, figs. 

12-14; Schmitt, McCain & Davidson, 1973, p. 61. 

Range: Massachusetts to south Florida; Texas; northwest Cuba; Guadeloupe; 

Pernambuco to Santa Catarina, Brazil. 

Habitat: parasitic in oysters and present in other bivalve molluscs, including: 

Crassostrea virginica, C. rhizophorae, Anomia simplex, Mytilus edulis, and


Pecten spp.; occasionally in polychaete {Chaetopterus) tubes; only the first crab 

("invasive") stage is free-swimming; found primarily in shallow bays and other 

suitable oyster habitats. 

Remarks: The large literature on this species was catalogued by Schmitt, 

McCain and Davidson (1973). Earlier biologists thought that this crab was a 

commensal of oysters, but its parasitic nature was definitely established, as summarized 

by Stauber (1945), Flower and McDermott (1953) and Haven (1958). 

Information on larval stages can be found in O. W. Hyman (1924), Lebour 

(1928), Aikawa (1937), Costlow and Bookhout (1966b) and in Sandoz and 

Hopkins (1947). Natural history of this species is reviewed by Christensen and 

McDermott (1959) and by MacGinitie and MacGinitie (1968); Williams 

(1965) summarized much of the current literature. Beach (1969) studied the life 

history of this crab in North Carolina. Hartnoll (1971) noted modifications for 

swimming activity. Listed from Texas by Hedgpeth (1953), Breuer (1962) and 

Leary (1967). 

Pinnotheres serreiVvaXhhun, 1909. 

Transfen-ed to a new genus, Ortholheres, by Sakai (1969). Refer to Orthotheres 

serrei. 

Pinnotheres shoemakeri Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 95) 

Rathbun, 1918, p. 95, text-fig. 48, pi. 22, figs. 1-4; Rathbun, 1933, p. 83; Schmitt, 


Range: west coast of Florida; St. Thomas, Virgin Islands. 

Pinnotheres stromhi Rathbun, 1905. 

Transferred to a new genus, Orthotheres, by Sakai (1969). Refer toOrthothere^ 

strombi. 

Subfamily PINNOTHERELIINAE Alcock, 1900 

Pinmxo White, 1846 

Pinnixa chacei Wass, 1955 (Quart. J. Flor. Acad. Sci. 18: 160) 

Wass, 1955, p. 160, figs. 5-9; Felder, 1973a, p. 71, pi. 10, fig. 5; Schmitt, McCain & 

Davidson, 1973, p. 104. 

Range: northwest Florida; Louisiana and Texas. 

Habitat: intertidal, commensal with burrowing shrimp, Callinassa islagrande, 

living in upper part of burrow; on sandy bottoms. 

Remarks: Listed from Florida by Wass (1955) and Menzel (1971), from 

Louisiana by Behre (1950) as Pinnixa sp., and from Texas (Leary, 1967). 

Pinnixa chaetopterana Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 

235) 

Hay & Shore, 1918, p. 44.5, pi. 36,rig. 4; Rathbun, 1918, p. 151, text-figs. 93-94, pi. 

33, figs, 3-6; Williams, 1965, p. 210, fig. 194; Felder, 1973a, p. 74, pi. 10, fig. 8; 

Srhmitt, McCain & Davidson, 1973, p. 104,


Range: Massachusetts to Florida; northwest Florida to Texas; Pernambuco 

to Rio Grande do Sul, Brazil. 


Habitat: mud, shell, and gravel bottoms; there are two forms of this crab 

along the northern Gulf coast: the larger is a commensal with the polychaetes 

Amphitrite ornata and Chaetopterus variopedatus, living inside the tubes of the 

hosts; the smaller form occupies the upper portion of burrows of Callinassa 

jamaicense louisianensis. 

Remarks: Larval stages were described by 0. W. Hyman (1924), Lebour 

(1928), and Aikawa (1937). Sandifer (1973) noted aspects of larval ecology 

in Virginia. Williams (1965) summarized current literature on this species and 

MacGinitie and MacGinitie (1968) provided a general account of its life history. 

Johnson (1952) described a "host factor" for this crab. Behavioral studies include 

Pearse (1913) and Gra}^ (1961), including notes on symbiotic relationships. 

Craig (1974) measured temperature tolerances and oxygen consumption. Listed 

from Florida by Wass (1955), Menzel (1971), and Godcharles and Jaap (1973) 

and from Mississippi by Richmond (1962) and Christmas and Langley (1973). 

Listed from Brazil by Righi (1967), Rodrigues da Costa (1971) and Coelho and 

Ramos (1972). 

Pinnixa cristata VâXhhun, 1900 (Amer. Natural. 34: 589) 

Hay & Shore, 1918, p. 446, pi. 36, fig. 5; Rathbun, 1918, p. 134, text-fig, 78, pi. 29, 

figs. 8-9; Williams, 1965, p. 210, fig. 193; Felder, 1973a, p. 74, pi. 10, fig. 6; Schmitt, 


Range: North and South Carolina; Louisiana and Texas. 

Habitat: intertidal beaches; shallow sand and sand-mud substrates of brackish 

to marine waters; usually commensal with callinassid shrimps and other 

burrowers. 

Remarks: Hedgpeth (1950) described these crab from salt flats that border 

the bays and intercoastal waterways of Texas. MacGinitie and MacGinitie 

(1968) included information on the ecology of this species. Listed from Louisiana 

by Behre (1950). 

Pinnixa cylindrica (Say, 1818) (J. Acad. Nat. Sci. Philadelphia 1: 452) 


figs. 5, 8; Milne Edwards & Bouvier, 1923, p. 343; Williams, 1965, p. 213, fig. 197; 

Schmitt, McCain & Davidson, 1973, p. 106. 

Range: Massachusetts to South Carolina; west and northwest coasts of Florida. 

Depth: shallow water to 37 m (to20fm). 

Habitat: commensal with Arenicola cristata (lugworm) in the non-tubular 

burrows; young crabs occur near the intertidal zone of slimy shores. 

Remarks: McDermott (1962) provided a general account of this species, 

which is also summarized by Williams (1965). Sandifer (1973) commented on 

larval ecology in Virginia. Listed by Menzel (1971) from northwest Florida. 

Pinnixa floridana Rathbun, 1918 (Bull, U.S. Nat. Mus. 97: 138) 

Rathbun, 1918, p. 138, te.\t-fig. 82, pi. 30, figs. 4-7; Williams, McCloskey & Gray, 

1968, p. 57, fig. 13; Schmitt, McCain & Davidson, 1973, p. 110.


Range: North Carolina; west and northwest coasts of Florida. 

Habitat: shallow water, possibly in tubes of the polychaete, Diopatra cuprea; 

collected from a compound ascidian growing on a soft coral; from under rocks 

in 10 feet of water. 

Remarks: Rathbun (1918) commented on morphological variation between 

the sexes of this species. Listed from Florida by Wass (1955) andMenzel (1971). 

Pinnixa leplosynaptae Yfass, 1968 (Tulane Stud. Zool. 14: 137) 

Wass, 1968, p. 137, figs. 1-6; Schmitt, McCain & Davidson, 1973, p. 112. 


Habitat: found on the surface of the holothurian Leptosynapta crassipatina. 

Remarks: Listed from northwest Florida by Menzel (1971). 

Pinnix(dlunzi G\a55e\\ 1937 (Charleston Mus. Leaflet 9: 3) 

Glassell, 1937, p. 3, figs. 1-8; Williams, 1965, p. 214, figs. 198-199; Felder, 1973a, 

p. 71, pi. 10, fig. 4; Schmitt, McCain & Davidson, 1973, p. 114. 

Range: Virginia to Georgia; Mississippi to Texas. 


Habitat: on beaches, under drift material; in burrows of echiurans (Thalassema 

hartmani) and possibly other burrowers. 

Remarks: Boesch (1971) listed this crab from an echiuran burrow in Virginia. 

Felder (1973b) reported a specimen taken from a red snapper stomach 

from a reef off Texas. 

Pinnixa pearsei Wass, 1955 (Quart. J. Flor. Acad. Sci. 18: 164) 

Wass, 1955, p. 164, figs. 10-13; Schmitt, McCain & Davidson, 1973, p. 116. 

Range: northwest coast of Florida. 

Habitat: commensal in tubes of the polychaete, Diopatra, from sand-mud 

beaches. 

Remarks: Listed from northwest Florida by Abele (1970) andMenzel (1971). 

Although Menzel (1971) listed this crab as a commensal of an undetermined 

annelid, Abele (1970) stated that the crab does not seem to be restricted to commensal 

relationships where it was common in the sand-mud intertidal zone of 

Alligator Harbor's south shore. 

Pinnixa relinens Rathbun, 1918 (Bull. U.S. Nat. Mus. 97: 139) 

Ratlihun, 1918, p. 139, text-figs. 83-84, pi. 41, figs. 1-2; Williams, 1965, p. 212, 

fig. 196; Felder, 1973a, p. 74, pi. 10, fig. 7; Schmitt, McCain & Davidson, 1973, p. 

118. 

Range: Chesapeake Bay; west coast of Florida; Texas. 


Habitat: mud bottoms of estuarine and marine waters; from burrows of the 

callinassid shrimp, Upogebia affinis. 

Remarks: Rathbun (1918) believed this crab to be allied with P. floridana. 

Listed from Florida by Wass (1955) and Menzel (1971). Williams (1965) included 

this species in the Carolina fauna, even though it hadn't yet been collected 

from that area. Based on the wide range of locales but paucity of specimens, 

it is probably a rare species.

128 Lawrence W .Powers 

Pinnixa sayana Stimpson, 1860 (Ann. Lye. Nat. Hist. New York 7: 236) 


figs. 2-4; Williams, 1965, p. 212, fig. 195; Felder, 1973a, p. 74, pi. 10, fig. 9; Schmitt, 


Range: Massachusetts to North Carolina; west coast of Florida; Louisiana; 

Amapa to Sao Pavilo, Brazil. 

Depth: surface to 75 m (to 41 fm). 

Habitat: free-swimming; on sandy beaches in drift material; from mud bottoms; 

in tubes of lugworm, Arenicola cristata. 

Remarks: Larval descriptions include 0. W. Hyman (1924), Lebour (1928), 

and Aikawa (1929, 1937). Regional lists include Louisiana (Behre, 1950). 

Schmitt, McCain and Davidson (1973) note that the host record of Arenicola 

may be due to synonymy of P. sayana with P. cylindrica by Hay and Shore 

(1918) and so may be in error. Listed from Brazil by Righi (1967), Rodrigues 

da Costa (1968a), Coelho (1971a) and Coelhoand Ramos (1972). 

Family GRAPSIDAE, Macleay, 1838 

Subfamily GRAPSINAE Macleay, 1838 

Geograpsus Stimpson, 1858 

Geograpsus lividus (H. Milne Edwards, 1837) (Hist. Nat. Crust., vol. 2. p. 85) 

Rathbim, 1918, p. 234, pi. 55; Rathbun, 1933, p. 87, fig. 80; Garth, 1965a, p. 26; 

Forest & Guinot, 1966, p. 91; Chace & Hobbs, 1969, p. 157, figs. 48, 52n-c. 

Range: Bermuda; Florida Keys, north and south coasts of Cuba; Jamaica; 

Puerto Rico; Virgin Islands to Barbados; Netherlands Antilles to Trinidad; Old 

Providence Island (Carib.); Caribbean coast of Colombia to Sao Paulo, Brazil; 

eastern Atlantic, from Senegal to Angola; Cape Verde Islands; eastern Pacific, 

from soutl.em part of Baja California to northern Chile; Clipperton Island; Galapagos 

Islands; Hawaiian Islands. 

Habitat: supralittoral, near the splash zone of rocky areas and stone beaches; 

from middle to upper intertidal, beneath stones. 

Remarks: Hartnoll (1965b) provided ecological notes on populations in Jamaica. 

Chace and Hobbs (1969) commented on ecology of this crab in Dominica. 

Listed from Braziil by Coelho and Ramos (1972) and Fausto Filho (1974). 

Goniopsis de Haan, 1833 

Goniopsis cruenlala (Latreille, 1803) (Hist. Nat. Crust., vol. 6, p. 70) 

Common Names: Mangrove Crab; Tree Crab 

Rathbun, 1918, p. 237, text-fig. 136, pi. 57; Rathbun, 1933, p. 87, fig. 81; Chace, 

19


lands to Barbados; Netherlands Antilles; Belize; Old Providence Island (Carib.); 

Surinam to Rio de Janeiro, Brazil; eastern Atlantic, from Senegal to northern 

Angola. 

Plabitat: mangrove swamps, along roots and on trunks of trees; on wet muddy 

marine shores, along inlets and estuaries; intertidal to supratidal. 

Remarks: Leary (1967) listed this species from Texas, based on a collection 

by Hildebrand in 1958, but its occurrence along the Gulf coast is scattered and 

rare. Ecological studies include field work in Jamaica (Hartnoll, 1965b; Warner, 

1969) and Dominica (Chace and Hobbs, 1969). Behavioral data is provided by 

Schone and Schone (1963), Schone (1968), and Warner (1970). Physiological 

studies include data on thoracic neurosecretion (Maynard, 1961a, 1961b; Mayard 

and Maynard, 1962) and coagulation (Morrison and Morrison, 1952). 

Listed from Brazil by Coelho and Ramos (1972). 

Grapsws Lamarck, 1801 

Grapsus grapsus (Linnaeus, 1758) (Syst. Nat., ed. 10, vol. 1, p. 630) 

Common Names: Rock Crab; Cliff Crab; Sally Lightfoot 

Rathbun, 1918, p. 227, text-fig. 135, pis. 53-54; Rathbun, 1933, p. 86, fig. 79; 

Garth, 1965, p. 25; Forest & Guinot, 1966, p. 90; Chace & Hobbs, 1969, p. 163, figs. 

50, 52g-i; Felder, 1973a, p. 78, pi. 11, fig. 15. 

Range: Bermuda; Bahamas; southeast and south Florida; Texas; north and 

south coasts of Cuba; Jamaica; Puerto Rico; Hispaniola; Virgin Islands to Barbados; 

Netherlands Antilles to Trinidad; Old Providence Island and Swan Island 

(Carib.); Colombia to northern Brazil; eastern Atlantic, from Portugal to 

Angola; Cape Verde Islands and Azores; St. Helena Island; Ascension Island; 

eastern Pacific, from central Baja California to central Chile; Galapagos Islands; 

Clipperton Island. 

Habitat: intertidal and supratidal zones of rocky areas, stone beaches, and 

sea walls; wdthin reach of splash from surf and wave action; in crevices and 

cracks of rock cliffs near water's edge. 

Remarks: Reports of this crab in the Gulf of Mexico are confined to the north 

coast of Cuba and from the rock jetties of Texas, where they are rare. Listed 

from Texas by Leary (1967), based on collections by Hildebrand. Also recorded 

from Brazil by Coelho and Ramos (1972) and Fausto Filho (1974). Ecological 

studies include those of Hartnoll (1965b) in Jamaica, Chace and Hobbs (1969) 

in Dominica, and Johnson (1965) on the relation of behavior to development 

and ecology. Social behavior was studied by Wright (1966, 1968), Schone and 

Eibl-Eibesfeldt (1965), Kramer (1967), Schone (1968), and Eibl-Eibesfeldt 

(film, 1963). Hartnoll (1971) noted the ability of this crab to swim. Physiological 

and anatomical studies include work on gill anatomy (Chen, 1933), coagulation 

(Morrison and Morrison, 1952), thoracic neurosecretion (Maynard, 

1961a, 1961b; Maynard and Maynard, 1962), and neural fine structure (Skobe 

and Nunnemacher, 1970).


Pachygrapsus Randall, 1840 

Pachygrapsus gracilis (Saussure, 1858) (Mem. Soc. Phys. Hist. Nat. Geneve 

15: 443) 

Common Name: Wharf Crab 

Rathbun, 1918, p. 249, pi. 60, fig. 3, pi. 61, fig. 1; Rathbun, 1933, p. 89; Holthuis, 

1959, p. 239, pi. 10, fig. 3; Forest & Guinot, 1966, p. 92; Chace & HOVDS, 1969, p. 

167, figs. 51, 52); Felder, 1973a, p. 79, pi, 11, figs. 3-4,11. 

Range: Bermuda; Bahamas; south Florida; Texas; north and south coasts 

of Cuba; Jamaica; Puerto Rico; Virgin Islands; Caribbean coast of Columbia; 

Pemambuco to Bahia, Brazil; eastern Atlantic, from Senegal to Zaire. 

Habitat: mangrove roots; along river banks near the sea; on pilings, wharves, 

stone jetties; rocky areas, just above the water level; intertidal to near supratidal. 

Remarks: Extensive notes on the natural history of this crab in Jamaica were 

provided by HartnoU (1965b) and by Warner (1969). Felder (1973a) listed 

some collection localities in Texas, but it is absent from collection lists of west 

Florida, Mississippi, and Louisiana. Listed from Brazil by Coelho and Ramos 

(1972). 

Pachygrapsus transversus (Gibbes, 1850) (Proc. Amer. Assoc. Adv. Sci. 3: 

181. 

Common Name: Mottled Shore Crab 

Hay & Shore, 1918, p. 447, pi, 36, fig, 9; Rathbun, 1918, p. 244, pi. 61; Rathbun, 

1933, p. 88, fig. 82; Williams, 1965, p. 217, fig. 202; Forest & Guinot, 1966, p. 91; 

Chace & Hobbs, 1969, p. 169, fig, 52k; Felder, 1973a, p. 79, pi. 11, figs. 5, 10. 

Range: North Carolina; Bermuda; Bahamas; east coast of Florida; Florida 

Keys and Dry Tortugas; Louisiana to east coast of Mexico; north coast of Cuba; 

Jamaica; Hispaniola; Puerto Rico; St. Thomas, Virgin Islands to Barbados; 

Trinidad; Netherlands Antilles; Bahia, Brazil to Uruguay; eastern Atlantic, 

from Mediterranean Sea to Angola; eastern Pacific, from central California to 

Peru; Galapagos Islands; Easter Island. 

Habitat: mainly in rocky areas near the tideline; beneath stones and on 

wharves and pilings; occasionally found among mangrove roots and on sandy 

beaches. 

Remarks: Leobur (1944) figured some of the larval stages. Ecological studies 

include Hartnoll (1965b) in Jamaica and Verrill (1908) in Bermuda. Listed 

from Louisiana (Behre, 1950) and Texas (Whitten, Rosene and Hedgpeth, 

1950; Leary, 1967). Pearse (1932a) reported a protozoan from the gill cavity 

of this crab. Hazlett (1971) studied antennule chemosensivity and Alves (1974) 

tested salinity tolerances. Listed from Brazil by Coelho and Ramos (1972) and 

FaustoFilho (1974). 

PZancs Bowdich, 1825 

Planes cyaneus Dana, 1852 (Proc. Acad. Nat. Sci. Philadelphia 5: 250) 

As P. minutus—Barnard, 1950, p. 120.


As P. cyaneus—Chace, 1951, pp. G5-103, figs, lb, 2b, 2e, 2h, 2in-o, 3i-ii; Chace 

1966, p. 646; Sakai, 1965, p. 197, pi. 93, figs. 3-4; Felder, 1973a, p. 78, pi. 11, fig. 1. 

Range: rare occurrence in Texas; south Atlantic, at St. Helena Island and 

off west coast of Africa; throughout eastern Pacific and westward to the northwestern 

Pacific and Indian Ocean. 

Habitat: pelagic, on floating objects, debris, and gulfweed; open ocean, but 

occasionally washed up with flotsam onto beaches. 

Remarks: Chace (1951) provided a definitive taxonomic review, but this crab 

was considered at that time to be confined to the Pacific Ocean. In 1966, Chace 

reported it from St. Helena Island in the south Atlantic and noted that Barnard's 

(1950) citations of P. minuius from South Africa may have been partly or entirely 

records of P. cyaneus. Crosnier (1967) reported this crab from West Africa. 

Shirley (1974) found two specimens washed up on the beaches at Padre 

Island, Texas, in driftwood. This species should be considered extralimital for 

the Gulf of Mexico, although re-examination of Planes in collections may reveal 

additional specimens of this species. 

Subfamily PLAGUSIINAE Dana, 1851 

Percnon Gistel, 1848 

Percnon gibbesi (H. Milne Edwards, 1853) (Ann. Sci. Nat. ser. 3, Zool. 20: 

146 and 180) 

Common Name: Spray Crab 

Rathbun, 1918, p. 337, pi. 105; Rathbun, 1933, p. 93; fig. 88; Schmitt, 1939, p. 24; 

Garth, 1965, p. 34; Williams, 1965, p. 224. 

Range: North Carolina, Bermuda; Bahamas, south Florida; Florida Keys; 

north coast of Cuba.; Jamaica; Puerto Rico; Colon, Panama; Brazil; eastern Atlantic, 

from the Azores to South Africa; eastern Pacific, from Baja California to 

Chile; Galapagos Islands; Clipperton Island. 

Habitat: low tide zone of rocky areas; surf zone, on rock and pebble bottoms, 

commensal with Diadema in Puerto Rico. 

Remarks: Rathbun (1918) questioned the inclusion of the Pacific and Atlantic 

populations of this crab into one species, but subsequent authors have treated 

them as identical. Verrill (1908) commented on this crab in Bermuda and Hartnoil 

(1965b) described its biology and ecology in Jamaica. Garth (1946) described 

it from the Galapagos Islands. Schmitt (1939) provided the key characters 

for the genus. The habitat and color of Brazilian specimens was provided 

byFaustoFilho (1974). 

PZagMsia La treille, 1806 

Plagusiadepressa (Fabricius, 1775) (Syst. entom., 1775), p. 406 

Common Name: Cliff Crab 

Rathbun, 1918, p. 332, text-fig. 154, pi. 101; Rathbun, 1933, p. 93, fig. 87; Monod,


1956, p. 455, figs. 614-617; Williams, 1665, p. 223, fig. 207; Forest & Guinot, 1966, 

p. 93; Chace & HobLs, 1969, p. 192, figs. 62r-t, 63; Feldtr, 1973a, p. 75, pi. 11, fig. 13. 

Range: North and South Carolina; Bermuda; Florida Keys and Dry Tortugas; 

Texas; Cuba; Jamaica; Hispaniola; Puerto Rico; Virgin Islands; Dominica to 

Barbados; Trinidad; Netherlands Antilles; Ceara to Pemambuco, Brazil; eastern 

Atlantic, from the Azores and Madeira and Senegal to Angola. 

Habitat: in fissures and crevices of rocks; in tide pools; on jetties; intertidal. 

Remarks: Chace and Hobbs (1969) provided notes on color patterns of living 

crabs. Haratnoll (1965b) studied the ecology of this crab in Jamaica and notes 

on swimming behavior were given in Hartnoll (1971). Physiological studies 

include Morrison and Morrison (1952) on coagulation and data on thoracic 

neurosecretion (Marynard, 1961a, 1961b; Maynard and Maynard, 1962). Listed 

from Brazil by Coelho (1971a), Coelho and Ramos (1972), and Fausto Filho 

(1974). 

Subfamily SESARMINAE Dana, 1852 

Aratus H. Milne Edwards, 1853 

Aratus pisonii (H. Milne Edwards, 1837) (Hist. Nat. Crust., vol. 2: 76) 

Common Names: Mangrove Crab; Tree Crab 

Rathbun, 1918, p. 323, pi. 96; Rathbun, 1933, p. 92, fig. 83: Chace & Hobbs, 1969, 

p. 172, figs. 54, 58a. 

Range; Bahamas; southeast to southwest Florida; norlh and south coasts of 

Cuba; New Providence Island (Atlantic); Jamaica; Puerto Rico; Virgin Islands 

to Guadeloupe; Netherlands Antilles; Belize; Rio Parahyba do Norte to Sao 

Paulo, Brazil; Nicaragua to Peru, in eastern Pacific. 

Habitat: along shores of estuaries and near fresh, brackish or marine waters; 

on rocks, piles, and wharves; commonly in mangroves, on which this crab can 

easily climb. 

Remarks: Warner (1968) described larval development. Hartnoll (1965b) 

provided extensive notes on the biolog}'- of this crab in Jamaica, including ecology, 

growth, feeding, behavior, and reproduction. Warner (1967, 1969, 1970) 

also studied this species in Jamaica. Hartnoll (1971) briefly commented on 

swimming activity. Listed from south Florida by Tabb and Manning (1961) 

and from Brazil by Coelho and Ramos (1972). 

Cyclograpsus H. Milne Edwards, 1837 

Cyclograpsus integer H. Milne Edwards, 1837 (Hist. Nat. Crust., vol. 2: 79) 

Common Name: Marsh Crab 

Rathbun, 1918, p. 326, pi. 97, figs. 1-2; Rathbun, 1933, p. 92, fig. 86; Monod, 1956, 

p. 451, figs. 609-612; Chace & Hobbs, 1969, p. 173, figs. 55, 58b-cl; Felder, 1973a, 

p. 75, pi, 11, figs. 12, 14. 

Range: Bermuda; Bahamas; south Florida; Florida Keys; Texas; Cuba; Jamaica; 

Hispaniola; Puerto Rico; St. Croix; Dominica; Islas Los Roques and


Caribbean coast of Colombia; Ceara to Pernambuco, Brazil; eastern Atlantic, 

from Senegal to Zaire. 

Habitat: burrows in marshy marine areas; among rocky and stony areas of 

the intertidal zone and up to the high tide line. 

Remarks: Felder (1973a) provided the only other specific Gulf record in addition 

to the previous record for the Florida Keys. HartnoU (1965b) commented 

on the ecology of this crab in Jamaica. Listed from Brazil by Coelho and Ramos 

(1972) and Fausto Filho (1974). 

Sesarjna Say, 1817 

Subgenus Holomelopus H. Milne Edwards, 1853 

Sesarma (Holomelopus) americanum Saussure, 1858 (Mem. Soc. Hist. Nat. 

Geneve 14: 441) 

As 5. tampicense—Bathbun, 1918, p. 307, text-fig. 151, pi. 88, 

AsS. americanum—Chace & Hobbs, 1969, p. 178, figs. 62a-f. 

Range: Tampico, Me.xico; St. Thomas, Virgin Islands. 

Habitat: soft mud, along river banks. 

Remarks: Chace and Hobbs (1969) determined that S. tampicense Rathbun 

was a junior synonym of 5. americanum Saussure. Behre (1950) tentatively 

listed the species from Louisiana (as S. tampicense), but noted that Chace had 

examined the specimens and preferred not to record the species as indicated. 

Abele (1972b) mentions the similarities betv\'een several of the western Atlantic 

members of the genus; he notes the distinct differences between S. angustipes 

Dana and S. americanum. 

Sesarma (Holomelopus) anguslipes Dana, 1852. 

This species was restricted by Abele (1972b) to the specimens from Brazil 

and Trinidad; refer to his paper for a discussion of synonymy. Material listed 

under this name by Rathbun (1918, p. 331) was synonymized with S. roberti 

by Chace and Hobbs (1969, p. 184). 

Sesarma (Holomelopus) benedicli Rathbun, 1897 (Proc. Biol. Soc. Washington 

11: 90) 

Bathbun, 1918, p. 316, pi. 93; Holthuis, 1959, p. 248, fig. 62; Abele, 1973, p. 379, 

figs. lA, IG. 

Range: Key West, Florida; Guyana and Surinam; Bra2;il. 

Habitat; under wood and stones on banks of brackish and almost freshwater 

streams. 

Remarks: The female specimen from Key West (MCZ 6236) listed by Rathbun 

(1918) is the only Gulf of Mexico record; all others are from South America. 

Ecological notes were provided by Holthuis (1959). 

Sesarma (Holomelopus) cinereum-(Bosc, 1802) (Hist. Nat. Crust, vol. 1, 

an X, p. 204) 

Common Names: Square-backed Fiddler; Wharf Crab; Wood Crab; Friendly 

Crab


As S. cinei-ea—Ha,y & Short, 1918, p. 449, pi. 36, fig. 11. 

As S. cinereum—Rathbun, 1918, p. 300, text-fig, 149, pi. 83; Williams, 1965, p. 

222, fig. 206; Felder, 1973a, p. 78, pi. 11, fig. 6; Abele, 1973, p. 377, figs. IB, IH. 

Range: Maryland to southeast Florida; southwest Florida to Vera Cruz, 

Mexico. 

Habitat: on wharves, pilings, and other wooden objects; stone jetties and rocky 

areas; in Spartina marshes and along the edges of mangrove swamps; burrows 

from the high tide mark to well inland in mud and sand substrates; frequently 

found on boats and ships. 

Remarks: Abele (1973) states that previous records of S. cinereum from the 

West Indies and elsewhere in the Caribbean were based on juvenile specimens 

of S. ricordi and S. americanum. Regional lists include Florida (Wass, 1955; 

Menzel, 1971; Subrahmanyam et al., 1976), Mississippi (Richmond, 1968), 

Louisiana (Behre, 1950; Hoese and Valentine, 1972) and Texas (Hedgpeth, 1953; 

Leary, 1967). Hedgpeth (1953) presented a map, showing the ranges of Sesarma 

in the northern Gulf of Mexico. Williams (1965) listed ovigerous females from 

North Carolina in May to November and from the mouth of the Potomac River 

in January. Abele (1973) collected ovigerous females from Florida in June and 

from Texas in Julj^ Sandifer (1973) commented on larval abundance in 

Virginia. 

Larval development was studied by Costlow, Bookhout and Monroe (1960) 

and Costlow and Bookhout (1960b). Ecological notes were provided by Williams 

(1965) and by Fotheringham and Brunenmeister (1975). Physiological studies 

include observations on gill area (Gray, 1957), oxygen consumption (Teal, 

1959), and tolerance to dilute salt water (Pearse, 1929). Observations on the 

behavior of this crab in captivity were made by Oler (1941) and by Duncker 

(1934). 

Sesarina (Holometopus) miersii Rathbun, 1897 (Proc. Biol. Soc. Washington 

11: 91) 

Rathbun, 1918, p. 303 (part), pi. 84; Chace & Hobbs, 1969, p. 180, figs. 59, 62g-i; 

Abele, 1972b, p. 166, figs. IB, IC, 2B, 2C; Abele, 1973, p. 380, fig. II. 

Range: Bahamas; Key West, Florida; south coast of Cuba; Swan Island 

(Carib.); Dominica. "^ 

Habitat: marshy tidal flats. 

Remarks: Abele (1972b) reviewed the status of this crab and the confusion 

in nomenclature that existed from Rathbun's (1897) description of Mier's original 

material. Only the original specimens from the Bahamas became the type 

material for this species. Specimens from Brazil in Rathbun (1918) are now 

designated S. angustipes and the specimen from Uruguay is Metasesarma 

ruhripes. Hartnoll (1965b) found no evidence of S. miersii in Jamaica and later 

authors agree that the observations of Andrews cited in Rathbun (1918, p. 304) 

refer to S. roberti (see Abele, 1973). The only Gulf of Mexico record is that 

from Key West (Abele, 1973).


Sesartna (Holomelopus) ricordi H. Milne Edwards, 1853 (Ann. Sci. Nat., ser. 

3,Zool. 20: 183) 

Common Name: Beach Crab 

Rathbun, 1918, p. 309 (part), pi. 89: Rathbun, 1933, p. 91; Holthuis, 1959, p. 246, 

pi. 11, fig. 3; Chace and Hobbs, 1969, p. 183, fig. 62k; Abele, 1973, p. 378, fig. IJ. 

Range: Bermuda; Bahamas; southeast Florida; Florida Keys; west coast of 

Florida; north coast of Yucatan; Cuba; Jamaica; Hispaniola; Puerto Rico; Virgin 

Islands to Trinidad; Curasao; Old Providence Island (Carib.); Yucatan to Surinam. 

Habitat: from intertidal zone to about 50 meters inland; in low-lying pine 

woods; edges of mangrove swamps; burrows in grassy areas above sandy beaches; 

under driftwood and among rocks, along shorelines. 

Remarks: Previous records of this species from Mississippi have been identified 

as S. cinereum by Chace (in Hedgpeth, 1953) and the material from Brazil 

was determined to be 5". ang«rfz/jg5 by Abele (1972b). Abele (1973) listed ovigerous 

females from Florida in May, June and August and from Panama in January. 

Larval development was studied by Diaz and Ewald (1968). The ecology 

and other aspects of biology of this crab in Jamaica were reported by HartnoU 

(1965b), Warner (1969), and Standing (1972). 

Sesarina (Holomelopus) roberli H. Milne Edwards, 1853 (Ann. Sci. Nat. ser. 

3,Zool. 20: 182) 

Common Name: Brackish-water Crab 

As S. angustipes—Rathbun, 1918, p. 311, pi. 90. 

As S. roberti—Hathhun, 1918, p. 312, pi. 91; Rathbun, 1933, p. 91; Monod, 1956, 

p. 443, figs. 602-604; Chace & Hobbs, 1969, p. 184, figs. 60, 621^n. 

Range: Veracruz, Mexico to Nicaragua; north and south coasts of Cuba; 

Jamaica; Hispaniola; Puerto Rico; Virgin Islands to Trinidad; Venezuela. 

Habitat: streams, rivers, and bays, including a wide range of freshwater and 

brackish environments; from marine shorelines to upland elevations of 1000 

feet; burrows in steep muddy banks; among mangroves; on rocks in streams; 

among stony areas at bases of cliffs. 

Remarks: Monod (1956) questioned the occurrence of S. roberti on Goree 

Island off West Africa, the indicated type-locality for this species. HartnoU 

(1965b) reported on the biology of this crab in Jamaica (under the name S. 

angustipes, which he used as a senior synonym of S. roberti). Chace & Hobbs 

(1969) maintained the distinction between these two species, at least until a 

South American specimen with Caribbean characteristics is discovered. Abele 

(1972b) reviewed these reports and also regarded the two forms as separate 

species. 

Sesartna (Holomelopus) ta/n/iicense Rathbun, 1914. 

This name was determined by Chace & Hobbs (1969) to be a junior synonym 

of S. americanum.


Subgenus Sesarma Say, 1817 

Sesarma (Sesarma) curacaoense de Man, 1892 (Notes Leyden Mus. 14: 257) 

Common Name: Mangrove Crab 

Rathbun, 1918, p. 293, text-fig. 147, pi. 78, figs. 1-2, pi, 160, fig. 3; Rathbun, 1933, 

p. 90; Holthuis, 1959, p. 242; Chace & Hobbs, 1969, p, 188, figs, 61, 62p; Abele, 

1973,p. 380, figs. IC, IF. 

Range: Key West, Florida; south and southwest Florida; north coast of Cuba; 

Jamaica; Puerto Rico; Curasao; Bahia, Brazil. 

Habitat: muddy banks of rivers and ditches, including brackish water; in 

mangrove swamps, under rocks and litter; intertidal zone, in clumps of oysters 

and among rocks. 

Remarks: Abele (1973) commented on sexual maturity and size ranges, but 

he did not report ovigerous females in the Florida material he studied. Tabb and 

Manning (1961) listed this crab from mangroves at Whitewater Bay in south 

Florida. Ecology of this species in Jamaica was studied by Hartnoll (1965b) and 

Warner (1969). Listed from Brazil by Coelho and Ramos (1972). 

Sesarma (Sesarma) reliculaluni (Say, 1817) (J. Acad. Nat. Sci. Philadelphia 

1: 73) 

Common Name: Marsh Crab 

AsS. reticulata—Hay & Shore, 1918, p. 448, pi. 36, fig, 12. 

As S. reticulatum—Rathbun, 1918, p. 290, pi. 77; Williams, 1965, p. 221, fig. 205; 

Felder, 1973a, p. 78, pi. 11, fig. 7; Abele, 1973, p, 380, fig. ID, IE. 

Range: Massachusetts to east coast of Florida; west coast of Florida to central 

Texas. 

Habitat: Spartina salt marshes; burrows in soft muds and sand-mud; under 

rocks and litter of intertidal streams and near-marine to brackish waters. 

Remarks: Lai-val descriptions were provided by 0. W. Hyman'(1924) and 

Costlow and Bookhout (1962). Sandifer (1973) reported on larval ecology in 

Virginia. Crichton (1960) gave a general account of life history as noted in 

Delaware marshes. Regional lists include Florida (Wass, 1955; Menzel, 1971; 

Subrahmanyam et al, 1976), Louisiana (Behre, 1950), and Texas (Hedgpeth, 

1953; Leary, 1967). The specimen listed by Tabb and Manning (1961) for this 

species was determined by Abele (1973) to be S. curacaoense. Humes (1941b) 

described a parasitic copepod in the gill chambers of this crab. Physiological 

studies include work on gill area (Gray, 1957), oxygen consumption (Teal, 

1959), rhythmic activity (Palmer, 1967), antennule chemosensitivity (Hazlett, 

1971), and melanophore hormones (Fingerman, Nagabhushanam and Philpott, 

1961). 

Subfamily VARUNINAE H. Milne Edwards, 1852 

EuchirograpsusH. Milne Edwards, 1853 

Euchirograpsus amêricanus A. Milne Edwards, 1880 (Bull. Mus. Comp. Zool. 

8: 18) 

Hay & Shore, 1918, p. 448, pl. 36, fig, 7; Rathbun, 1918, p. 282, text-fig. 144, pi.


74; Chace, 1940, p. 52; Williams, 1965, p. 220, fig. 204; Turkay, 1975, p. 114, figs. 

6^7, 16b, 20, 24. 

Range: North and South Carolina; south Florida; north and south coasts of 

Cuba; St. Lucia; Barbados; Colombia to Venezuela. 

Depth: 31 to 508 m (17 to 278 fm). 

Habitat: rocky, coral, and sand substrates. 

Remarks: Listed from off the Carolines by Cerame-Vivas, Williams and Gray 

(1963) and Cerame-Vivas and Gray (1966). Williams (1965) listed ovigerous 

females from Florida in March to September. Recorded off the Atlantic coast 

of Cuba by Chace (1956). The Pacific specimens listed by Garth (1946) were 

referred to a new species, E. pacificus, and the specimen (USNM 17672) listed 

by Rathbun (1918) from off Yucatan was referred to a new species, E. antillensis, 

byTiirkay (1975). 

Euchirograpsus antillensis Tiirkay, 1975 (Senckenbergiana Biol. 56: 112) 

As E. aniericanus—Rathbun, 1918, p. 283 (part, Yucatan specimen only). 

As E. antiffensis—TiJrkay, 1975, p. 112, figs 4-5,16a, 19, 2S. 

Range: off Havana, Cuba; Arrowsmith Banks, between Cuba and Yucatan; 

south of Florida Keys; Bahamas. 

Depth: 192to430m(105lo235fm). 

Remarks: Tiirkay (1975) compared this new species with E. americanus and 

the other species of the genus, which now number a total of six. 

Plalychirograpsus de Man, 1896 

Plalychirograpsus speclahilis de Man, 1896 (Zool. Anz. 19: 292) 

Common Names: River Crab; Saber Crab 

As P. lypicujs—Rathbun, 1918, p. 278, text-figs. 141-143, pi. 73; Bolivar y Pieltain, 

1945, p. 267-270, figs. 1-5. 

As P. specln6i7f.«—Buitendijk, 1950, p. 280, fig. lb; Monod, 1956, p. 426, text-figs. 

584-588. 

Range: Gulf coast of Mexico; west coast of Florida. 

Habitat: burrows in clay banks, just above the water line, along rivers; shallow 

rocky areas of rivers; known from altitudes of greater than 100 feet, up to 

140 miles from the sea. 

Remarks: This species was described on the basis of only a few specimens, all 

from Ttibasco, Mexico. Marchand (1946) discovered a large population of these 

crabs along the Hillsborough River in west Florida, which empties into the Gulf 

near Tampa. These crabs were found to have originated in Mexico from where 

they were transported on logs and in lumber for Tampa, beginning about 1915. 

Marchand (1946) provided notes on ecology, behavior, and feeding habits of the 

west Florida populations, under the name P. typicus. Buitendijk (1950) determined 

this latter name to be a junior synonym of P. specLabilis. 

Plalychirograpsus typicus Rathbun, 1914. 

Determined b}^ Buitendijk (1950) to be a junior synon3'^m of P. spectabilis 

deMan, 1896.


Family GECARCINIDAE Macleay, 1838 

Cardisoma Latreille, 1825 

Cardiosoma guanhunii Latreille, 1825 (Encycl. Meth., Hist. Nat., Entom., vol. 

10, p. 685) 

Common Names: Great Land Crab; White Land Crab; Mulatto Land Crab; 

Juej'-; Tourlourou; Guanhumi; Guaiamu 

Rathbun, 1918, p. 341, text-fig. 155, pis. 106-107; Rathbun, 1933, p. 94, fig. 89; 

Hright, 1966, p. 191, fig. 4-1; Forest & Guinot, 1966, p. 94; Chace & Hobbs, 1969, 

p. 195, figs. 64, 67a-c; Tiirkay, 1970, p. 345; Rright & Hogue, 1972; p. 16; Felder, 

1973a, p. 79, pi. 12, figs. 1,4. 

Range: Bermuda; Bahamas; southeast Florida; Florida Keys; Louisiana and 

south Texas; eastern Mexico to Colombia; north and south coasts of Cuba; Jamaica; 

Puerto Rico; St. Thomas, Virgin Islands to Barbados; Trinidad; Netherlands 

Antilles; Colombia to Sao Paulo, Brazil. 

Habitat: low-lying coastal areas, especially mangrove swamps; open fields; 

along rivers, streams, drainage canals and ditches; under buildings; saline soils 

with high water tables; primarily nocturnal, but diurnal in heavily shaded 

areas and on days when the sky is heavily overcast. 

Remarks: This is the largest land crab in the Gulf of Mexico region. It is 

commercially harvested as a food item on some islands of the West Indies, especially 

Puerto Rico. In other areas, such as southern Florida, this crab is considered 

an agricultural pest because of the damage caused to fields by the large, 

extensive burrows and also due to the fondness of the crabs for young, growing 

shoots. 

Regional lists include Louisiana (Behre, 1949, 1950) and Texas (Leary, 

1967; Felder, 1973a). Listed from Cuba by Chace (1940) and from Brazil by 

Coelho and Ramos (1972). Accounts of natural history were provided by Gifford 

(1962b), Feliciano (1962), and Folheringham and Brunenmeister (1975). 

Henning (1975a, 1975b) studied the biology of this crab in northern Columbia, 

including extensive observations on behavior and ecology. Wright (1968) described 

agonistic behavior, especially chela displays during social encounters. 

Herreid (1963) investigated feeding behavior and Herreid and Gifford (1963) 

reported on the burrow as a habitat and on ionic regulation by the crab. 

Developmental stages were described by Moreira (1913) and Costlow and 

Bookhout (1968b). Costlow and Bookhout (1968c) studied the effects of various 

environmental factors on development. Physiological studies include work on 

calcium metablism (Gifford and Johnson, 1962), growth and morphometries 

(Herreid, 1967), various terrestrial adaptations (Bliss, 1963, 1968), aerial respiration 

(Cameron, 1975), respiratory pigments (Redmond, 1962), and uric acid 

metabolism (Gifford, 1968), osmoregulation by larval stages (Kalber and Costlow, 

1968), neurobiology of autotomy and leg elevation (Moffett, 1975), pericardial 

organ neurosecretion (Cooke and Goldstone, 1970; Berlind and Cooke, 

1970; Berlind, Cooke and Goldstone, 1970) neural control of walking (Barnes, 

Spirito and Evoy, 1972; Spirito, Evoy and Barnes, 1972; Evoy and Fourtner,


1973; Fourtner and Evoy, 1973; Moffett, 1975) and biochemistry (Quinn and 

Lane, 1966,1967). 

Studies of economic and commercial impact include those of de Oliviera 

(1946) on the fishery and ecology of this crab in Brazil and Feliciano (1962) on 

the fishery in Puerto Rico. Humes (1958) described a copepod from the gill 

chambers of this crab. 

Gecarcinus Leach, 1814 

Gecarcinus lateralis (Freminville, 1835) (Ann. Sci. Nat., ser. 2, Zool. 3: 224) 

Common Names: Black Land Crab; Common Land Crab 

Rathbun, 1918, p. 3SS, text-fig. 161, pis. 119-120; Rathbun, 1933, p. 9S, fig. 91; 

Chace & Holthuis, 1948, p. 26; Chace & Hobbs, 1969, p. 198, figs. 6S, 67e-g; TiJrkay, 

1970, p. 337, figs. 2a-c; Bright & Hogue, 1972, p. 21; Felder, 1973a, p. 82, pi. 

12, figs. 2-3; Tiirkay, 1973, p. 974, fig. 2. 

Range: Bermuda; Bahamas; southeast Florida; Florida Keys; south Texas to 

north coast of Yucatan; north and south coasts of Cuba; Jamaica; Hispaniola; 

Puerto Rico; St. Thomas, Virgin Islands to Barbados; Netherlands Antilles; 

Honduras to Costa Rica; Caribbean coast of Columbia to Surinam. 

Habitat: burrows in dry, sandy areas; in back dunes and on dune ridges; up 

to 1000 foot elevation in Dominica (Chace and Hobbs, 1969); in wooded areas 

of dune ridges and back dunes of eastern Florida, under logs and leaf litter; primarily 

nocturnal, but diel in heavily wooded habitats. 

Remarks: The status of this species and of G. quadratus have been unclear 

for some time. Tiirkay (1970) listed G. quadratus as a subspecies of G. lateralis, 

but an examination of types in the Paris Museum led Tijrkay (1973) to conclude 

that the two are synonymous. Most other workers have listed them as distinct 

and separate species, yet recognizing the complex distribution pattern of 

G. quadratus on both sides of Central America. Some specimens of G. lateralis 

have also been reported from the Pacific coast. Specimens collected in Texas 

(Ray, 1967; Britton, 1976; personal collections) are of G. lateralis. 

Regional lists include Texas (Ray, 1967; Britton, 1976; Felder, 1973a; Fotheringham 

and Brunenmeister, 1975) and Mexico (Cabrera, 1965, zoeaonly). Listed 

from Costa Rica by Bright (1966). Reports on ecology and natural history of this 

species include Bhss and Sprague (1958b), Weitzman (1963), Bliss (1968), 

Chace and Hobbs (1969), and Klaassen (1975). This crab has been used for a 

variety of physiological studies, especially for research on molting and regeneration 

(Hodge, 1956a, 1956b; Bliss, 1960a, 1960b, 1966; Bliss and Boyer, 1964; 

Bliss etal., 1972; Skinner, 1965, 1966; Skinner and Graham, 1972; Mason, 1970; 

Holland and Skinner, 1976; Yamaoka and Skinner, 1976). Other physiological 

reports include work on osmoregulation and water balance (Bliss, 1963; Bliss, 

Wang and Martinez, 1966; Mantel, 1968; Copeland, 1968), aerial respiration 

(Cameron, 1975), lipid metabolism (O'Connor and Gilbert, 1968), coagulation 

(Morrison and Morrison, 1952; Stutman and DoUiver, 1968), neuroendocrinology 

(Hodge and Chapman, 1958; Bliss and Sprague, 1958a; Maynard, 1961a, 

1961b; Maynard and Maynard, 1962; Weitzman, 1969; Mantel et al., 1975),


oxygen transport in hemolymph (Redmond, 1968), saline composition for lab 

experiments (Skinner, Marsh and Cook, 1965), neural fine structure (Skobe and 

Nunnemacher, 1970), and sensitivity to substrate vibrations (Klaassen, 1973). 

Gecarcinus quadralus Saussure, 1853 (Rev. Mag. Zool., ser. 2, vol. 5, p. 360) 

Common Names: Red Land Crab; Whitespot Crab 

Rathbun, 1918, p. 358, text-fig. 162, pis. 121-122; Garth, 1948, p. 58; Bright, 1966, 

p. 190, fig. 4G; Tiirkay, 1970, p. 338, fig. 4; Bright & Hogue, 1972, p. 20; Turkay, 

1973, p. 974. 

Range: Veracruz, Mexico to Turbo, Columbia (Carib.); St. Croix; Jamaica; 

Barbados; in eastern Pacific, from Acapulco, Mexico to La Libertad, Ecuador. 

Habitat: well above the high tide mark of sandy beaches; mangrove and 

other heavily-vegetated areas of marine shores; under debris and other litter. 

Remarks: As indicated for G. lateralis, this species may be regarded as distinct, 

as a subspecies of G. lateralis, or as completely synonymous with the latter. 

Tiirkay (1970) published a biogeographical distribution map of the two forms 

which shows considerable overlap in the ranges of the two species. The two forms 

are listed here as separate species, but those workers accepting Tiirkay's (1973) 

synonj^my of the two forms can combine the references and locality records. 

Gecarcinus ruricola (Linnaeus, 1758) (Syst. Nat., ed. 10, vol. 1, p. 626) 

Common names: Black Crab; Mountain Crab; Blue Land Crab; Red Tourlourou 

Rathbun, 1918, p. 352, text-fig. 160, pis. 117-118; Chace & Holthuis, 1948, p. 26; 

Chace & Hobbs, 1969, p. 200, figs. 66, 67h-j; Turkay, 1970, p. 336, fig. la-f; Bright 

& Hogue, 1972, p. 20. 

Range: Bahamas; southeast Florida; north and south coasts of Cuba; Cayman 

Islands; Jamaica; Navassa Island (Carib.); Hispaniola; Puerto Rico; St. Croix 

to Barbados; Curasao; Old Providence and Swan Islands (Carib.). 

Habitat: closer to the intertidal zone than other species of this genus; on low 

and marshy grotmd and on lower slopes of island mountains, up to elevations of 

500 m; in wooded dune areas of southeast Florida (rare, personal observation). 

Remarks: Descriptions of ecology and behavior are found in Rathbun (1918) 

and Chace and Hobbs (1969); a summary is provided by Bright and Hogue 

(1972). Listed from the south coast of Cuba hj Chace (1940). This species 

occurs among dense populations of G. lateralis in southeast Florida, but is rather 

rare. 

Superfamily OCYPODOIDEA Rafinesque, 1815 

Family OCYPODIDAE Rafinesque, 1815 

Subfamily OCYPODINAE Rafinesque, 1815 

Ocypode albicans Bosc, 1801-1802. 

Ocypode Weber, 1795 

Junior synonym for Ocypode quadrata, used by Rathbun (1918) and others 

prior to revision by Holthuis (1959).

Ocypoda arenaria Sa}^ 1817. 


Junior synonym and invalid generic name for Ocypode quadrata. used by 

Cowles (1908) and some other early studies. 

Ocypode quadrata (Fabricius, 1787) (Mantissa insect . . ., vol. 1, p. 315) 

Common Names: Ghost Crab; Sand Crab; Racing Crab 

As O. albicans—Rathhxm, 1918, p. 367, pis. 127-128; Rathbun, 1933, p. 96, fig. 92. 

As O. quadrata—Hohhxiis, 1959, p. 259; Williams, 1965, p. 225, fig. 208; Chace & 

Hobbs, 1969, p. 204, figs. 68-69; Feldcr, 1973a, p. 82, pi. 12, figs. 5, 8. 

Range: Bermuda; Bahamas; Rhode Island to south Florida; Florida Keys and 

Dry Tort.ugas; west coast of Florida, around entire Gulf coast to Yucatan; north 

and south coasts of Cuba, through West Indies to Barbados; from Yucatan, along 

east coast of Central America and the north coast of South America to Estado de 

Santa Catarina, Brazil; most Caribbean Islands, including Netherlands Antilles, 

Old Providence Island, etc. Megalops have been collected as far north as 

Massachusetts. 

Habitat: on sandy beaches, from high v\'ater line to back dunes areas; younger 

crabs burrow closer to water line and among beach vegetation; along waveexposed 

shores, protected harbor beaches, ba3rs, intracoastal canals, and lagoons; 

juveniles are mainly diel and older adults are primarily nocturnal, depending 

on degree of disturbance by man and various environmental factors. Adults 

usually burrow well back from the waterline, but often feed at the driftline. 

Remarks: Regional lists include Florida (Wass, 1955; Menzel, 1971), Mississippi 

(Richmond, 1962), Louisiana (Behre, 1950; Hoese and Valentine, 1972), 

and Texas (Whitten, Rosene and Hedgpeth, 1950; Hedgpeth, 1953; Leary, 

1967; Fotheringham and Brunenmeister, 1975). Bright and Hogue (1972) include 

this species in their world-list of land crabs; listed from Brazil by Coelho 

(1971a), Coelhoand Ramos (1972), and Fausto Filho (1974). 

Diaz and Costlow (1972) described and illustrated larval stages raised under 

laboratory conditions. Haley (1969) provided data on growth and morphometries 

of Texas populations; reproductive C3^cling, female morphometries, and 

population d3^namics were covered in Haley (1967, 1972). Hughes (1973) 

described mating behavior in the laboratory and compared the mating functions 

of burrows in several ghost crab species. Population densities and interactions 

v\dth man were studied by Teerling (1970). Accounts of general natural history 

were provided by Cowles (1908) for populations in the Tortugas (as Ocypoda 

arenaria) and by Milne and Milne (1946) for New Jersey. Williams (1965) 

summarized many of the recent studies. Chace and Hobbs (1969) described 

color phases of Dominican populations. Burrow construction and ecology in 

Texas was reported by Hill and Hunter (1973) and predatory behavior on mole 

crabs was noted by Fales (1976). Schone (1968) investigated agonistic displays 

and these were also presented in a film (Schone and Eibl-Eibesfeldt, 1965). 

Physiological studies include work on oxygen consumption (Pearse, 1929; 

Ayres, 1938; Vemberg, 1956; Gray, 1957), water relations and the role of the 

pericardial sac (Blass, 1963, 1968), ionic regulation and respiration (Flemister


and Flemister, 1951; Flemister, 1958), ionic and osmotic regulation (Gifford, 

1962a), gill and "kidney" histophysiology (Flemister, 1959), biochemistry of 

terrestrial adaptations (Vernberg and Vernberg, 1968), thoracic neurosecretion 

(Maynard, 1961a, 1961b; Maynard and Maynard, 1962), and visual perception 

(Schone and Schone, 1961). Studies of acoustic perception and related behavior 

include Horch and Salmon (1969), Horch (1971), and Salmon and Horch 

(1972). 

t/ca Leach, 1814 

(This genus of intertidal ocypodids, along with a few other Australo- 

Asian genera, are commonly known as fiddler crabs. A number of subgenera 

have been proposed for this large and diverse genus. Bott (1973) 

split this genus into 10 genera and Crane (1975) also created a number 

of subgenera. Although Crane's (1975) monograph is a comprehensive 

and monumental work, taxonomic precedence must be given to Bott 

(1973) with regard to most of these proposed changes. For the present, 

and in agreement with a review by von Hagen (1976), this compilation 

will avoid the use of subgenera and will continue the use of IJca, with 

the species arranged alphabetically.) 

t/ca fcwrgers/Holthuis, 1967 (Zool. Meded. Leiden 42: 52) 

As U. inordax—Rathbun, 1918, p. 391 (part), not text-fig. 166, nor pi. 134, figs. 

3^; Maccagno, 1928, p. 46 (part); de Oliviera, 1939a, p. 138; Holthuis, 1939, p. 

265. 

As U. feurgrersi'—Holthuis, 1967, p. 32; Chace & Hcbbs, 1969, p. 207, figs. 70, 

71a-d; Gibbs, 1974, p. 84; Crane, 1973, p, 168, figs. 26F, 31H, 34G, 66F, 100, pi. 

24E-H, map 12. 

Range: Bahamas; east coast of Florida; northeast (Gulf) coast of Yucatan; 

north and south coasts of Cuba; Jamaica; Hispaniola; Puerto Rico; St. Thomas, 

Virgin Islands to Trinidad; Curasao; east coast of Yucatan to Guatamala; Caribbean 

coast of Panama; Venezuela to Rio de Janeiro, Brazil. 

Habitat: sheltered mud flats; sloping mud banks and mud-sand areas at mouths 

of streams; along shores of lagoons and estuaries; often near mangroves; intertidal; 

above high tide mark in mangrove thickets of Florida, associated with V. 

rapax. 

Remarks: Earlier references confused this species with V. mordax, to which it 

is similar in morphology, ecology, and geographical range. This species was also 

recorded as V. affinis by Holthuis (1959), when he distinguished it from V. 

mordax in Surinam. Records of this crab from west Africa are questioned by 

Crane (1975). Crane (1957) included data on waving displays as part of her 

description of V. mordax. Adaptation to intertidal zone habitats was reported by 

von Hagen (1970b) and Salmon (1967) obtained sound recordings of legwagging 

(as V. mordax). Gibbs (1974) investigated the ecology of this crab on 

Barbuda and Gibbs and Bryan (1972) studied cation composition of the exoskeleton.


Vca leplodaclyla Rathbun, 1898 (Ann. New York, Acad. Sci. 11: 227) 

Rathbun, 1918, p. 420, pi. 156; Maccagno, 1928, p. 41 (part), not text-fig. 25; 

Rathbun, 1933, p. 98; de Oliviera, 1939a, p. 126, pi. 5, text-figs. 25-28, pi. 6, fig. 

29, pi. 8, fig. 47, pi. 13, figs. 61-62; Chace & Hobbs, 1969, p. 212, figs, 71g-h; von 

Hagen, 1970a, p. 227; Crane, 1975, p. 304, text-figs. 37M, 56F, 60N-O, 69K-L, 101, 

map 17, pi. 41A-D. 

Range: Bahamas; west coast of Florida (not recently); east coast of Yucatan; 

north coast of Cuba; Jamaica; Puerto Rico; St. Croix; Curasao; Venezuela to 

Santa Catarina, Brazil. 

Habitat: relatively sandy tidal flats of marine waters; sometimes at supratidal 

levels, where burrows are covered only by spring tides; occasionally in mud or 

claj' substrates, in partial shade of mangrove trees. 

Remarks: Behavioral studies include Matthews (1930), Crane (1957), and 

Gerlach (1958b). Ecological data were provided by Matthews (1930), de Oliviera 

(1939a, 1939b, 1939c), and Crane (1957). Bott (1973) lists this species under 

his genus Leptuca; Crane (1975) lists it under her subgenus Celuca. Listed from 

Brazil by Coelho (1971a) and Coelho and Ramos (1972). 

Vca longisignalis Salmon & Atsaides, 1968 (Proc. Biol. Soc. Washington 81: 

279) 

As U. pugnax rapax—Rathbun, 1918, p. 397 (part), not. pi. 140. 

As U. longisignalis—Salmon & Atsaides, 1968b, p. 279, text-figs. 1-4, 6, 7. 

As U. rapax longisignalis—Crane, 1975, p. 190, map 14. 

Range: northwest Florida to south Texas. 

Habitat: similar to that of U. rapax; often on exposed salt flats and algal beds; 

among marsh vegetation; substrates of mud, mud-sand, and sand-mud. 

Remarks: Felder (1973a) comments on the distribution of U. pugnax, U. 

virens and U. longisignalis and retains the name of U. pugnax for all forms in 

the northwestern Gulf. Crane (1975) listed this form as a subspecies of U. rapax, 

which until recently was a subspecies of U. pugnax. In a review of Crane's 

(1975) monograph, von Hagen (1976) commented that an examination of Salmon's 

holotypes left "no doubt that U. rapax longisignalis is a synonym of U. 

minax (Le Conte)." The present list treats this crab as a separate species, mainly 

on the basis of Salmon and Atsaides (1968b) evidence of behavioral separation 

and on the lack of agreement among other authorities as to which subspecies or 

species it is most closely allied with. Subrahmanyam et al. (1976) listed this 

crab from northwest Florida and Powers (1975) noted its occurrence in Texas. 

Included in the key to northwestern Gulf Uca by Fotheringham and Brunenmeister 

(1975). Abele (1970) provided habitat notes on populations at Alligator 

Harbor, Florida. 

Vca minax (Le Conte, 1855) (Proc. Acad. Nat. Sci. Philadelphia 7: 403) 

Common Names: Red-Jointed Fiddler Crab; Brackish Water Fiddle Crab 

Hay & Shore, 1918, p. 451, pi. 37, fig. 3; Rathbun, 1918, p. 389, pi. 137; Maccagno. 

1928, p. 48, text-fig. 31; Crane, 1943a, p. 220, text-fig. lb; Williams, 1965, p. 227, 

figs. 209A, 210B; Felder, 1973a, p. 85, pi. 12, fig. 12; Crane, 1975, p. 176, figs. 

67D, 8IK, 100, pi. 25E-H, map 12.


Range: Massachusetts to northeast Florida; northwest Florida to Louisiana, 

possibly Texas. 

Habitat: brackish to fresh waters of estuaries, bays and streams; drainage 

ditches and canals; usually located some distance from marine waters, but often 

subject to some tidal influence, particularly along the Atlantic coast; burrows 

in mud banks and among marsh vegetation, often supratidal; occasionally at 

edges of fields or woodlands. 

Remarks: Crane (1975) questioned the presence of this species from Texas, 

stating that records listed by Rathbun (1918) from Texas were found to be 

U. rapax longisignalis and U. pugnax virens; von Hagen (1976) believes that 

U. {rapax) longisignalis is a synonym of U. minas. If the latter is true, then 

the range of U. minax would definitely include Texas, but the form described 

by Salmon and Atsaides (1968b) as U. longisignalis does not inhabit freshwater 

and brackish areas. Listed from northwest Klorida by Wass (1955), Abele 

(1970), and Menzel (1971), from Louisiana by Behre (1950) and Felder 

(1973a), and from Texas by Leary (1967), but the latter is probably based on 

Rathbun (1918). 

Gray (1942) and Miller (1965) provided descriptions of natural history and 

Williams (1965) summarized much of the current literature. Hyman (1920, 

1922) described post-larval development and behavior, including spawning. 

Ecological studies include Teal (1958) in Georgia, Miller and Maurer (1973) 

on distribution in relation to salinity. Whiting (1972) and Whiting and Moshiri 

(1974) on distribution in relation to substrate, and Kerwin (1971) on distribution 

in relation to marsh vegetation. Salmon (1967) studied distribution in 

Florida. Miller (1961) compared feeding adaptations in this and other Uca 

species. Physiological studies include work on gill area (Gray, 1957), osmotic 

and ionic regulation (Cole, 1971), the relationship between respiration and 

habitat (Teal, 1959), tolerance to desiccation (Pearse, 1929), acclimation to 

temperature (Vernberg, 1959), tidal rhythms of color change (Fingerman, 

Lowe and Mobberly, 1958), and radiation sensitivity (Engel, 1973). Nimmo 

et al. (1971) studied PCB absorbtion from sediments. Behavioral studies include 

descriptions of waving displays (Crane, 1943a, 1957; Salmon, 1965), sound 

production (Salmon, 1965), and vibration reception (Salmon and Horch, 1973). 

Vcamordax (Smith, 1870). 

Although recorded from the Gulf of Mexico by Rathbun (1918, p. 391), these 

and earlier records had confused this species with U. vocalor, U. rapax, and TJ. 

burgersi. Crane (1975, p. 173) restricts TJ. mordax to the continental coast, from 

Guatamala to Brazil, plus the island of Trinidad. 

Uca panacea Novak & Salmon, 1974 (Proc. Biol. Soc. Washington 87: 313) 

Common Name: Sand Fiddler Crab 

As v. pugf/ator—Rathbun, 1918, p. 400 (part), not pi. 141 and pi. 169, fig. 2; 

Felder, 1973a, p. 83; Crane, 1975, p. 223 (part), not pi. 29E-H, part map 16, other 

figs, indet. 

As v. panacea—Novak & Salmon, 1974, p. 316, figs. 1-8.


Range: northwest Florida to south Texas. 

Habitat: sandy areas of marshes and tidal flats; often supratidal, intertidal 

in some areas; well inland on Texas barrier islands; similar to habitats of Z7. 

pugilator when intertidal. 

Remarks: This species is morphologically similar to Z7. pugilator, but it has 

been separated from the latter on the basis of biochemical (Selander, Johnson 

and Aviso, 1971) and behavioral studies (Novak and Salmon, 1974). Color variants 

of "Z7. pugilator''' from Florida were noted b}"- Rao & Fingerman (1968), 

a species-specific character present only in living specimens. It is likely that 

many of the studies reported as "Z7. pugilator" when collected from the central 

and western Gulf coasts, actually utilized U. panacea. Uca pugilator has been 

collected as far west as central Texas (Carl Thurman, pers. comm.; pers. obseiT. 

of author), indicating greater geographical overlap between the two species 

than reported by Novak and Salmon (1974). Other authors (Felder, 1973a; 

Crane, 1975) regarded the two forms as one species. The southern limits of Z7. 

panacea have not been defined yet, but may extend into northeastern Mexico. 

Hedgpeth (1950) commented on the ecology of this crab on salt flats in Texas 

and Powers (1973) provided data on burrow densities. Ecological and behavioral 

data on Texas barrier island populations were presented by Powers (1975) and 

Powers and Cole (1976). Studies prior to 1974, using the name U. pugilator, 

may include either or both species; regional lists and references are listed under 

TJ. pugilator, but many of these studies are undoubtably of U. panacea. 

Vca pugilator (Bosc, 1802) (Hist. Nat. Crust., vol. 1, an X, p. 197) 

Common Name: Sand Fiddler Crab 

Hay & Shore, 1918, p. 452, pi. 37, fig. 2; Rathbun, 1918, p. 400 (part), pi. 141, pi. 

160, fig. 2; Maccagno, 1928, p. 44, text-fig. 28; Crane, 1943a, p. 220; Williams, 

1965, p, 232, figs. 209C, 210C-D, 211; Crane, 1975, p. 223 (part), text-figs. 37K, 

69F, 101,pI.29E-H,map 16 (part). 

Range: Bahamas; Massachusetts to south Florida; Florida Keys; west and 

northwest coasts of Florida; Mississippi to Texas; ? Santo Domingo; ? Old Providence 

Island (Carib.) 

Habitat: sandy and sand-mud substrates; intertidal to supratidal marshes; 

burrows on open sand flats or among thick clumps of grasses and other vegetation. 

Remarks: This species is listed by Bott (1973) in the genus Planuca and by 

Crane (1975) in the subgenus Celuca. The status of the Caribbean specimens 

needs to be re-examined with respect to U. pugilator and U. panacea. Because 

of the widespread use of this animal in experimental studies, the taxonomic relationships 

and variability of morphological, behavioral, and physiological features 

need to be defined and established for both of these closety-related forms. 

A partial listing of the large literature on U. pugilator, including some of TJ. 

panacea, follows. 

Accounts of natural history include Pease (1914) in Massachusetts, Schwartz 

and Safir (1915) in New York, and studies by 0. W. Hyman (1920, 1922) and 

Dembowski (1925, 1926). Developmental studies were reported by Hyman


(1920), Gray (1942) on transient prezoea, and by Hernkind (1968b). Miller 

(1968) investigated asymmetry during growth and Vemberg and Costlow (1966) 

studied handedness. Ecological studies include work on habitat preferences in 

Georgia (Teal, 1958), feeding efficiency (Miller, 1961), habitats in the Bahamas 

(Coventry, 1944), habitats in Massachusetts (Knopf, 1966), distribution in relation 

to thermal tolerance (Miller and Vernberg, 1968), thermal relations of 

crab and microhabitat (Smith and Miller, 1973), and capture-recapture methods 

(Hockett and Kritzler, 1972). 

Sand Fiddlers have been the subjects of many behavioral studies: displays and 

courtships (Pease, 1914; Dembowski, 1925, 1926; Crane, 1943a, 1957), threat 

displays (Schone, 1968; Aspey, 1971), sound production and visual signals (Burkenroad, 

1947; Salmon and Stout, 1962; Salmon, 1965, 1967; Salmon and Atsaides, 

1969; Salmon and Horch, 1972), burrowing activity (Teal, 1958; Coward, 

Gerhardt and Crockett, 1970), visual orientation (Herrnkind, 1968a, 1968c, 

1972), feeding (Miller, 1961), locomotion (Baird and Burleson, 1970), and 

larval shadow responses (Forward, 1977). 

Physiological studies include work on molting (Abramowitz and Abramowitz, 

1940; Guyselman, 1953; Stewart and Green, 1969; Skinner and Graham, 

1972; Fingerman and Fingerman, 1976; Weis, 1976a), regeneration (Weis, 

1976b, 1976c, 1977a, 1977b; Weis and Mantel, 1976), color changes and chromatophores 

(Carlson, 1935, 1936; Brown and Sandeen, 1948; Brown and Webb, 

1948; Brown, 1950; Guyselman, 1953; Webb, Bennett and Brown, 1954; Fingerman 

and Yamamoto, 1967; Barnwell, 1968a; Rao and Fingerman, 1968; Fingerman, 

Rao and Ring, 1969; Coohill and Fingerman, 1975), metabolism (W. B. 

Vemberg and Vemberg, 1972), rhythmical activity and physiolog}^ (Brown 

et al., 1955; Fingerman, 1956, 1957; Fingerman, Lowe and Mobberly, 1958; 

Barnwell, 1966, 1968b), biochemistry (Eisen et cd., 1973), sensitivity to anemone 

toxin (Blanquet, 1968), reproduction and endocrinology (Darby, 1935; 

Brown and Jones, 1949; Sandeen, 1950; Fingerman and Fitzpatrick, 1956; Fingerman 

and Couch, 1967; Rao, Fingerman and Bartell, 1967; Rao and Fingerman, 

1969, 1970; Fingerman, 1970, 1973; Bartell, Rao and Fingerman, 1971; 

Fielder, Rao and Fingerman, 1971), thermoregulation and temperature adaptations 

(Edwards, 1950; Orr, 1955; Demeusy, 1957; Wilkins and Fingerman, 

1965; Vemberg, DeCoursey and Padgett, 1973), osmoregulation (Pease, 1929; 

Teal, 1958; Green et al., 1959; Evans, Cooper and Bogan, 1976), toxicology 

(Nimmo et al., 1971; DeCoursey and Vernberg, 1972; O'Hara, 1973), respiration 

(Gray, 1957; Teal, 1959; Wilkins and Fingerman, 1965; Smith and Miller, 

1973; Silverthorn, 1975a, 1975b), sensory perception (Salmon and Atsaides, 

1969; Horch and Salmon, 1969; Salmon, 1971; Langdon, 1971; Avent, 1974; 

Hyatt, 1974, 1975; Salmon, Horch and Hyatt, 1977), neu: obiologj' (Nunnemacher, 

1965; Andrews, 1973), radiation sensitivity (Engel, 1973), and infection 

by bacteria (Spindler-Barth, 1976). 

Regional lists include Florida (Wass, 1955; Tabb and Manning, 1961; Menzel, 

1971), Louisiana (Behre, 1950; Hoese and Valentine, 1972), and Texas (Hedgpeth, 

1950, 1953; Whitten, Rosene and Hedgpeth, 1950; Simmons, 1957; Leary, 

1967), but the Louisiana and Texas records probably refer to Uca panacea.

Ilea pugnax (Smith, 1870). 


This is another species with a history of frequent nomenclatural changes. 

Most older references have included this crab as a Gulf species, but Salmon and 

Atsaides (1968b) referred the Gulf populations of U. pugnax to new species, 

U. virens and V. longisignalis. Crane (1975) maintained U. virerLS as a subspecies 

of U. pugnax, but she placed U. longisignalis as a subspecies of V. rapax. 

Until a revision by Tashian and Vernberg (1958), Z7. rapax was considered a 

subspecies of U. pugnax; thus all four forms are closety related. However, von 

Hagen (1976) synonymizes Z7. virens with U. rapax and considers U. longisignalis 

to be synonymous with U. minaxl The present list treats each form separately, 

maintaining each species presented by Salmon and Atsaides (1968b) 

and excluding U. pugnax as a Gulf species. 

Vca rapax (Smith, 1870) (Trans. Connecticut Acad. Arts Sci. 2: 134) 

As V. pugnax rnpn;c—Rathbun, 1918, p. 397 (part), pi. 140; Maccagno, 1928, p. 

45, text-fig. 29; Rathbun, 1933, p. 97; de Oliviera, 1939a, p. 134. 

As V. rapax—Tashian & Vf3rnberg, 1958, Holthuis, 1959, p. 266, text-figs. 64d-f, 

65, pi. 14, figs. 4-6, pi. 15, fig. 3; Chace & Hobbs, 1969, p. 214, figs. 73a-b; von 

Hagen, 1970a, p. 226; Crane, 1973, p. 190, figs. 52C-DD, 54F, 67C, 86, 91E-F, 

100, pis. 27A-D, 45C-F, map 14. 

Range: Bahamas; east coast of Florida; Florida Keys; southwest coast of Florida; 

northeast coast of Mexico to northeast Yucatan; north and south coasts of 

Cuba; Jamaica; Hispaniola; Puerto Rico; St. Thomas, Virgin Islands to Trinidad 

and Tobago; Netherlands Antilles; east coast of Yucatan to Guatamala; Caribbean 

coast of Panama to Santa Catarina, Brazil. 

Habitat: mud, sand-mud, and mud-sand flats; edges of mangroves; along 

rivers and streams on flats and banks. 

Remarks: This species may also occur infrequently along the northwestern 

Gulf coast, but Crane (1975) attributes records of this crab to U. rapax longisignalis. 

Felder (1973a) listed V. rapax from the same area, but past records may 

be erroneous with regard to the several similar species involved. Listed from 


Behavioral studies include observations on waving displays (Crane, 1943a, 

1957), combat between males (Crane, 1957, 1967), visual and acoustical signalling 

(Salmon and Atsaides, 1968a), kinaesthetic orientation (von Iâgen, 

1967), orientation to burrows (von Hagen, 1970b), and feeding (Miller, 1965). 

Warner (1969) studied the natural historj^ of this crab in Jamaica and Holthuis 

(1959) provided ecological notes and populations in Surinam. Smith and Miller 

(1973) measured thermal adaptations. Barnwell (1963) observed motor activity 

and the rhythmicity of color changes in populations in Brazil. Handedness and 

its relationship to development was analyzed by Vernberg and Costlow (1966). 

Adaptations to particular tidal levels were observed by von Hagen (1970b). Salmon 

(1971) measured vibration receptivity and van Delft (1968) studied daily 

rhythms of color changes.


Vca speciosa (Ives, 1891) (Proc. Nat. Acad. Sci. Philadelphia 1891: 179) 

Rathburi, 1918, p. 408, pi. 143; Chace & Hobbs, 1969, p. 213, figs. 73c-d; von Hagen, 

1970a, p. 227; Crane, 1975, p. 236, text-figs. 68G, 101, map 15, pi. 31A-D. 

Range: southeast Florida; Florida Keys; west and northwest coasts of Florida; 

northeast Yucatan and northwest Cuba. 

Habitat: wet, muddy substrates; mid to high intertidal zone; commonly foimd 

in mangroves. 

Remarks: Specimens from Curagao reported by Rathbun (1918) were referred 

to U. cumulanta by Chace and Hobbs (1969); Crane (1975) referred the Jamaican 

specimen of Chace and Hobbs (1969) to V. cumulanta. Crane (1957) 

provided some preliminary data on courtship displays and Salmon (1967) analyzed 

waving patterns of the crabs. Miller (1965) studied the distribution and 

ecology of this species. Listed from Florida by Wass (1955), Tabb and Manning 

(1961) andSubrahmanyamgfaZ. (1976). 

Vca spmicar/ja Rathbun, 1900 (Amer. Natural. 34: 586) 

As U. spinicnrpa—Rathbun, 1918, p. 411, pi. 148; Felder, 1973a, p. 83, pi. 12, 

fig, 11. 

As U. speciosa spinicarpa—Crane, 1973, p. 239, figs. 68K, 101, pi. 31E-H, map 13. 

Range: Alabama to northeastern coast of Mexico. 

Habitat: muddy banks of coastal freshwater ponds and streams; muddy, 

brackish beaches of the Gulf; grassy mud flats off bays (after Felder, 1973a). 

Remarks: This crab was considered a separate, but allied species to U. speciosa 

by Rathbun (1918) and subsequent workers. Crane (1975) placed it as a subspecies 

of the latter. Felder (1973a) listed several personal collections in Louisiana 

and Mississippi. Listed from Texas by Learj'- (1967) and Fotheringham and 

Brunenmeister (1975) comment on its presence in the northwestern Gulf, providing 

a key for comparison with other Uca species. Bott (1973) placed this 

species and U. speciosa in his genus Leptuca; Crane's (1975) subgenus designation 

is Celuca. 

Vca suhcylindrica (Stimpson, 1859) (Ann. Lye. Nat. Hist. New York 7: 63) 

Rathbun, 1918, p. 419, pi. 155, pi. 160, fig. 3: Felder, 1973a, p. 83, pi. 12, fig. 10; 

Crane, 1975, p. 209, figs. 67, 100, pi. E-H,map 11. 

Range: Texas to northeastern coast of Mexico. 

Habitat: banks of freshwater streams; brackish water areas; on mud flats 

and algal beds, often some distance upstream from mouths of rivers and creeks. 

Remarks: This species is uncommon and has a restricted range. Listed by 

Fotheringham and Brunenmeister (1975) for the northwestern Gulf. Very little 

is known about this crab's ecology, behavior, or other biological aspects. 

Vca thayeri Rathbun, 1900 (Proc. Washington Acad. Sci. 2: 134) 

Rathbun, 1918, p. 406, text-fig. 169, pi. 144; Rathbun, 1933, p. 98; Holthuis, 1959, 

p. 275, text-figs. 68b-c, pi. 16; Chace & Hobbs, 1969, p. 216, text-figs. 73e-f; 

von Hagen, 1970a, p. 226; Crane, 1973, p. 112, figs. 46K, 56E, 60H-I, 73A-B, 811, 

821, 99, map 11, pi. 17. 

Range: east and southwest coasts of Florida; north and south coasts of Cuba;


Jamaica; Hispaniola; Puerto Rico; Guadeloupe; Trinidad; Tobago; Guatamala 

and Panama (Caribbean coasts) Venezuela to Sao Paulo, Brazil. 

Habitat: deep mud an banks of streams and estuaries, among mangrove 

swamps; burrows are often shaded by vegetation. 

Remarks: Ecological studies include Gerlach (1958a) in Brazil Warner (1969) 

in Jamaica, and Salmon (1967) in Florida. Crane (1957) described daily behavioral 

displays and Barnwell (1963) reported on daily and tidal rhythms of 

activity. Bott (1973) placed this species in his genus Planuca; Crane (1975) 

designated the subgenus Boboruca. listed from Brazil by Coelho and Ramos 

(1972). 

Vca virens Salmon & Atsaides, 1968 (Proc. Biol. Soc. Washington 81: 281) 

As v. pKgna.i;—Felder, 1973a, p. M. 

As V. DiVens—Salmon & Atsaides, 1968b, p. 281, figs. 2-3, 5-7. 

As V pugnax virens—Crane, 1975, p. 203, map 10. 

As U. rapax—von Hagen, 1976, p. 224. 

Range: Mississippi to Coatzacoalcos (central Gulf coast), Mexico. 

Habitat: muddy sand, sand-mud, and mud substrates of salt marshes; algal 

flats, close to bays, estuaries and inlets; often among marsh vegetation. 

Remarks: Studies that have recorded "Uca pugnax" from the Gulf coast may 

represent records of V. virens or V. longisignalis; the occasional presence of U. 

rapax along the Texas and Mexico coasts is also possible. Salmon and Atsaides 

(1968b) reported analyses of waving and acoustic signalling. Powers (1975) 

and Powers and Cole (1976) provided some data on habitats of this crab in Texas. 

See Uca pugnax and Uca longisignalis for a discussion of nomenclatural changes 

in the pugnax-rapax species group. 

Vca vocator (Herbst, 1804) (Versuch. Natur. Kxabben u. ICrebse, vol. 1. pi. 59, 

fig- 1) 

As U. morrfa.v—Rathbun, 1918, p. 391 (part), pi. 134, figs. 3-4. 

As U. murifecenla—Crane, 1943b, p. 38, text-figs. Id-f, pi, 1, figs. 1-2. 

As U. rocalor—Holthuis, 1959, p. 269, text-figs. 66-67, pi. 14, fig. 1, pi. 15, fig. 1; 

Chace & Hobbs, 1969, p. 217, figs. 73g-j, 74; von Hagen, 1970a, p. 225; Crane, 1975, 

p. 27, figs. 66D, 100, pi. 23E-G, pi. 24A-D, map 13. 

Range: Tampico, Mexico; Belize to Guyana; Puerto Rico; Santo Domingo; 

Guadeloupe; Dominica; Trinidad and Tobago; Paraiba to Pemambuco, Brazil; 

? Santa Catarina, Brazil. 

Habitat: grassy marshes; mud flats; flat banks of streams and rivers; in darnp 

mud among mangroves. 

Remarks: This species was presented by Rathbun (1918) with U. mordax. 

Ecological data includes studies by Crane (1943b) in Venezuela, by Holthuis 

(1959) in Surinam, by Chace and Hobbs (1969) in Dominica, and by von Hagen 

(1970a, 1970c), who also commented on sound production and other aspects 

of behavior. The only record for this species from the Gulf of Mexico is that 

listed by Crane (1975) for Tampico, Mexico. Listed from Brazil by Coelho and 

Ramos (1972) for Paraiba and Pemambuco, but Crane (1975) questions the 

record from Santa Catarina.


Ucides Rathbun, 1897 

Vcides cordalus (Linnaeus, 1763) (Amoen. Acad., vol. 6, p. 414) 

Common Names: Pagurus; Kaburi 

Rathbun, 1918, p. 347, text-fig. 158, pis. 110-113, pi. 159, figs. 3-4; Rathbun, 1933, 

p. 95, fig. 90; Bott, 1955, p. 66; Bright, 1966, p. 191; Chace & Hobbs, 1969, p. 219, 

figs. 75-76; Tiirkay, 1970, p. 351, fig. lOa-d; Bright & Hogue, 1972, p. 14. 

Range: Bahamas; southeast Florida; northeast Mexico to Panama; north and 

south coasts of Cuba; Jamaica; Hispaniola; Puerto Rico; St. Thomas, Virgin 

Islands to Grenada; Colombia to Santa Catarina, Brazil. 

Habitat: swampy ground, among mangrove roots; in areas of standing brackish 

water; on mud flats, among Uca and Cardlsoma burrows. 

Remarks: Chace and Hobbs (1969) transferred this genus to the Ocypodidae 

from the Gecarcinidae and Tiirkay (1970) accepted this transfer. Although the 

genus did not fit conveniently into any of the existing subfamilies, Chace and 

Hobbs (1969) felt that it was more closely allied with the Ocypodinae than with 

others. Turkay (1970) placed the Pacific species, U. occidentalism under U. 

cordatus as a subspecies. Manning and Provenzano (1961) comment on Ucides 

in Florida. Ecological and behavioral notes were provide by Chace and Hobbs 

(1969) and Bright and Hogue (1972). Warner (1969) discussed the ecology of 

this crab in Jamaica and de Oliviera (1946) studied its biology in Brazil. Listed 

from Brazil by Coelho and Ramos (1972). De Souza and Caland (1968) reported 

on bacterial infections in this species. Ogawa et al. (1973a, 1973b) 

described commercial processing of this crab for food in Brazil. Alves (1975) 

studied reproductive biology of Brazilian populations. 

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Acanthocarpus 29 

A. alexandri 29 

A. bispinosus 30 

Acanthouychinae 41 

Acanlhonyx 41 

A. petiuerii 41 

Actaea 87, 104 

A. acantha 87 

A. bifrons 88 

A. palmeri 88 

/I. rufopunctata nodosa 88, 

103 

/I. êi;ge/a 88, 107 

Aciaeomorpha 33 

Aepinus 43 

/I. septemspinosus 43 

Aethra 33 

Aelhrinae 33 

Anasimus 43 

/I. /aûi 43 

Anomalothir 43 

/I. frontalis A'i 

A. furcillalus 43 

Arachnopsis 44 

/I. filipes 44 

arafia del mar 49 

Aratus 132 

/I. pisonii 132 

Arenaeus 74, 82 

/I. cribrarius 74 

arrow crab 49 

assemblages, faunal 13-14 

ATELECYCLIDAE 9, 11-13, 

71,72 

Atelecyclinae 71 

Bathynectes 72 

S. superba 72 

Bathyplax 111, 112 

S. /:r/'A/n 112 

Batrachonotus 44 

S. fragosus 44 

beach crab 135 

Benthochascon 73 

S. schmilti 73 

black crab 140 

black land crab 139 

blue crab 78-81, 96 

blue land crab 140 

Boboruca 149 

INDEX 

box crab 30 

BRACHYRHYNCHA 8, 9, 72 

brackish-water crab 135 

brackish-water fiddler crab 143 

Cabouca 58 

Calappa 30 

C. angusta 30 

C./Zammea30, 31 

C gallus 31 

C ocellata 31 

C. springeri 32 

C. sulcata 32 

CALAPPIDAE 9,11-13,29 

Calappinae 29 

CALAPPOIDEA 9, 29 

Calico crab 33, 90 

Callidactylus 37 

C. a^per 37 

Callinectes 18, 75, 82 

C. bocourti 75 

C. da/zae 75, 76, 77, 81 

C. exasperatus 76 

C. marginalus 77 

C. ornazui 75, 77-78, 81 

C.rathbunae 75, 78 

C.sapirfus 75, 78-81,96 

C. sapidus acutidens 78, 79 

C. similis 75, 76, 77, 78, 81 

Cancer 71 

C. borgalis 71 

C.irroratus 72 

CANCRIDAE 9,11-13, 71, 72 

CANCRIDEA 9, 71, 72 

Cancrinae 71 

cancroid crabs 72 

CANCROIULA 9, 71 

cangrejo de la Santa Virgen 58 

Carcinoplacinae 111 

Cardisoma 138, 150 

C. guanhumi 138 

Carolinean Province 11, 13-14, 

73 

Carpilius 88 

C. convexus 88 

C. corallinus 88 

Carpoporus 89 

C. papulosus 89 

Cataleptodius 89 

C. floridanus 89, 95 

Catametopa 72 

Celuca 143, 145,148 

Chacellus 112 

C. fillformis 112 

Chasmocarcinus 112 

C. cylindricus 112 

C. mississippiensis 112 

C. obliquus 113 

Chlorodiella 89 

C longimana 89 

Chorinus 62 

C /zeroi 62 

cliff crab 129, 131 

Clythrocerus 26 

C. /zz'iz'dui 26 

C. stimpsoni 26 

Coelocerus 50 

C spinosus 50 

Collodes 44 

C armalus 44 

C. leplocheles 44 

C trispinosus 45 

common edible crab 78 

common land crab 139 

common mud crab 102 

common pea crab 124 

corrunon spider crab 64 

coral crab 56, 88 

Corycodus 26 

C. bullatus 26 

Corj'Stidae 9 

Cronius 81 

C. ruber 81 

C. tumidulus 82 

Cryptochhus 40 

C corallicola 40 

Cryptopodia 67 

C concava 67 

Cyclodorippe 26 

C antennaria 26 

C bouvieri 26 

C ornata 27 

Cycloes 32 

C. fcraVdii 32 

Cyclograpsus 132 

C. i/zieger 132 

Cyclometopa 72 

Cycloxanlhops 110 

Cymonomus 27 

C caecus 27


C. cubensis 27 

C. quadratus 9,7 

C. rosiratus 27 

CymopoUa 27 

C.affinisn?, 

C. alternata 118 

C. cursor 118 

C. denlata 118 

C. faxoni 118 

C. gracilipes 118 

C. gracilis 119 

C. obesa 119 

C.sicaW^ 

CYMOPOLIDAE 117 

Cymopolus 91 

C. agassizi 27 

decorator crab 52, 58 

deep sea red crab 111 

Dicranodromia 21 

D. ovala 21 

dirt}'' decorator crab 54 

disjunct distribution 14, 18 

Dissodaclylus 119 

D.alcocki 119 

D. borradailei 120 

D. calmani 120 

D. crinitichelis 120 

D.encopei 120 

D. juvenilis 120 

D.melliiae 120 

D. primitivus 121 

D. stebbingi 121 

Dolly Varden crab 33 

Dornecia 89 

D. acanthophora acanlhophora 

89-90 

D, AwpiWa 89-90 

D0RIPPIDAE9, 11-13, 26 

DORIPPOIDEA 9, 26 

Dromia 19 

D. erylhropus 19 

DR0MIACEA9,19,22 

Dromidia 19 

D. antillensis 19 

DR0MIIDAE9,11-13, 19 

DROMIOIDEA 9, 19 

Dj'nomenidae 9, 10 

Ebalia 35 

£. cariosa 35 

£. stimpsoni 35 

Ebaliinae 35 

edible crab 78 

endemic species 11, 14 

Epiallus 41 

JB. biiuberculatus 41 

£. dilaiatus 42 

£. dilaiatus forma elongata 

42 

£. longirostris 42 

Eriphia 90 

£. gonagra 90 

Ethusa 28 

£. Zaza 28 

£. rnascarone americana 28 

£. rnicrophlhahna 28 

£. teruiipes 28 

£. truncata 29 

Ethusina 29 

£. abyssicola 29 

£. favonii 29 

£Ziiui 90, 104 

£. maculatus 90 

Euchirograpsus 136 

£. americanus 136, 137 

£. antillensis 137 

£. pacificus 137 

Eucratodes 91 

£. agassizii 91 

Eucralopsis 113 

£. crassimanus 113 

Euphrosynoplax 113 

£. clausa 113 

Euprognatha 45 

£. gracilipes 45 

£- raslellijera acuta 45 

£. raslellijera niarlhae 45 

Eurypanopeus 91, 103 

£. abbreviatus abbreviatus 91 

£. abbreviatus ater 91 

£. crenatus 91 

£• depressus 91 

£. dissimilis 92 

E.turgidus 92, 103 

Euryplax 113 

£. ni'Zi'Az 113 

Eurytium 92 

£. limosum 92 

£aWa 121 

£. byssorniae 121 

F. teWnae 121 

fiddler crab (see Uca) 

flame-streaked box crab 30 

freshwater crab 87 

Frevillea 114 

£. barbata 114 

£. hirsuta 114 

F. irWe«toall4, 116, 117 

friendly crab 133 

frog crab 25 

GECARCINIDAE 9, 11-13, 138 

Gecarcinus 139 

G. lateralis 139, 140 

G.(/ua(Zra/ui 139,140 

G. ruricola 140 

Geograpsus 128 

G. /iyiWui 128 

Geryon 111, 112 

G. ajji'railll 

G.quinquedens 111 

GERYONIDAE 9,11-13,110, 

112 

ghost crab 141 

Glyptoplax 114 

G. pugnax 114 

G.smithii 114 

Glyploxanthus 93 

G. erosus 93 

GONEPLACIDAE 9,11-13, 72, 

100, 110, 111 

Goneplax 114 

G. ta?-tetoll4 

G. hirsuta 114 

G. tridentata 117 

Goniopsis 128 

G. cruentata 128 

GRAPSIDAE 9, 72, 128 

Grapsinae 128 

grapsoid crabs 72 

Grapsus 129 

G. grapsus 129 

grass crab 52 

great land crab 138 

guaiamu 138 

guanhumi 138 

Gymnopleura 23 

hairy crab 107 

HAPALOCARCINIDAE 8, 9, 

11-13,40 

HAPALOCARCINIDEA 9, 40 

HAPALOCARCINOIDEA 9, 

40 

Hemus 50 

//. crislulipes 50 

Hepatus 32, 33 

//. epheliticus 33

H. princeps 33, 34 

H. pudibundus 33, 34 

Heieraclaea 93 

H. ceratopus 93 

Heterocrypla 67 

//. granulala 67 

Hexapanopeus 93, 96 

//. anguslijrons 93 

//. hemphillii 94 

//. fo^>;>ei 94, 96 

//. paulensis 94, 95 

//. quinquedentatus 94 

//. sinaloensis 95 

Holonietopus 133 

Holopliles 63 

//. armata 63 

Homola 22 

//. barbala 22 

//. f fg77 22 

HOMOLIDAE 9,10,11-13, 

21,23 

Homolodromia 21 

//. paradoxa 21 

Homolodroiniinae 10, 21 

Honiologenus 22 

//. rostratus 22 

HOMOLOIDEA 9, 21 

Hypoconcha 20 

//. arcuata 20 

//. sabulosa 20 

//. spinosissima 21 

Hypsophrys 22-23 

//. noar 22 

Iliacantha 37 

/. intermedia 37 

I. liodactylus 37 

/. sparsa 38 

/. subglobosa 38 

Inachinae 42 

Inachoides 45 

/. jorcepts 45, 46 

/. laevis 45, 46 

isozyme patterns 18 

Jonah crab 71 

juey 138 

kaburi 150 

lady crab 74 

land crab 138,139, 140, 150 

Lalreillia 10, 23 

L. elegans 23 

LATREILLIDAE 10, 11-13, 

23 

Latreillopsis 23 

lazy crab 58 

Leiolambrus 68 

L. nilidiis 68 

LEUCOSIIDAE 9,11-13, 33, 

35 

Leucosiinae 36 

leopard crab 33 

Leplodius 95 

L. agassizzi 95, 108 

L. floridanus 89, 95 

Z,. parvulus 95 

Lepiuca 143, 148 

lesser blue crab 81 

libinia 50, 63 

L. J«Z)M 63, 64, 65 

L. emarginala 63, 64 

L. erinacea 65 

L. rhomboidea 65 

Lithadia 35 

L. cadaverosa 35 

Lobopilumnus 95 

Z^. agassizii 95 

loop currents 15 

Lophopanopeus 94, 96 

L. distinctus 96, 98 

L. /o^>;pei 94, 96 

Lupella 82 

i. forceps 82 

Lyreidus 23 

i. bairdii 23 

macca crab 58 

Macrocoeloma 50 

M. campLocerum 50 

M. diplacanihum 51 

M, eutheca 51 

M. intermedium 51 

yW. laevigatum 51 

M. seplemspinosiim 52 

Af. subparallelum 52 

M. trispinosum trispinosum 

52 

M.trispinosum nodipes 52, 53 

M. trispinosum variety 

Rathbun 53 

Macrocoelomjnae 50 

Macropipinae 72 

Macropipus 73 

MAJIDAE 9, 11-13,41 


Majinae 49, 50 

mangrove crab 128, 132, 136 

marsh crab 132, 136 

Malula 33 

Matutinae 32, 33 

Melybia 96 

M. thalamila 96 

Menippe 96 

M. rnercenaria 93, 96-97 

M. nodifrons 97 

Mesorhoea 68 

iVf. sexspinosa 68 

Melasesarma rubripes 134 

Mgloporhaphis 46 

M. calcarata 46 

Micropanope 96, 97 

Af. barbadensis 97 

M. dislincta 96, 98 

7W. lobifrons 98 

M. nullingi 98 

M. p««7/a 98 

M. sculptipes 97, 98, 99 

iW. spinipes 99 

M. truncatifrons 99 

M. urinaior 99 

M. xanlhiformis 99, 100 

Microphrys 53 

iV/. anlillensis 53 

iV/. bicornulus 54 

M. plalysoma 53, 54 

Mithracinae 50 

Mithraculus 54 

Mithrax 54, 56 

iVf. (Mithraculus) cinctimanus 

54 

M. (Mithraculus) coryphe 55 

M. (Mithraculus) forceps 

55,57 

M. (Mithraculus) ruber 55 

7W. (Mithraculus) sculplus 

56 

M. (Mithrax) acuticornis 56 

M. (Mithrax) cornutus 56 

7W. (Mithrax) depressus 57 

7W. (Mithrax) hispidus 56 

M. (Mithrax) holderi 57 

M. (Mithrax) pilosus 57 

M. (Mithrax) pleuracanthus 

57 

M. (Mithrax) spinosissimus 

56,58 

M. (Mithrax) verrucosus 58 

Mocosoa 42


M. crebripunciaia 42 

mottled shore crab 130 

mountain crab 140 

mud crab 102 

mulatto land crab 138 

mussel crab 123 

Myropsis 38 

M. quinquespinosa 38 

Nanocassiope 99 

N. melanodaclylus 99 

Nanoplax 99, 100 

A', xanthijormis 99, 100 

Neodoclea 50 

Neopanope 100 

N. packardii 100, 101 

N. sayi 100, 101 

N. texana 100 

N. texana sayi 101 

Neopilumnoplax 114, 115 

N. americana 114, 115 

Nibilia 65 

N. antilocapra 65 

northein crab 71 

Notosceles chimrnonis 25 

Ocypode 140 

O, albicans 140 

O. argnaria 141 

O. quadrala 140, 141 

OCYPODIDAE 9,10, 11-13, 

140 

OCYPODOIDEA 9, 140 

Ocypodinae 140 

Ophthalmiinae 60 

Orlhotheres 122,125 

O. ierrei 122, 125 

O. strombi 122, 125 

Osachila 32, 33, 34 

O. anlillensis 34 

O. semilevis 34 

O. tuberosa 34 

Oualipes 13, 18, 72, 73 

O. floridanus 74 

O. guadulpensis 73, 74 

O. ocellatus 73 

O. slephensoni 74 

OXYRHYNCHA 9, 41 

OXYSTOMATA 8, 9, 23 

oyster crab 124 

Pachygrapsus 130 

P. gracilis 130 

P. Iransversus 130 

pagurus150 

PALICIDAE 8, 9,11-13, 27, 

72, 117 

Palicus27, 117 

P.affinism 

P. alternatus 118 

P. cursor 118 

P.dentalus 118 

P. /ajronz 118 

P. gracilipes 118 

P. gracilis 119 

P. ofeiz/i 119 

P. szca 119 

Panopeus 101 

P. arnericanus 101 

P. berniudensis 101 

P. fozrZri; 101 

P. herbstii 93, 102 

P. occidentalis 103 

P. rugosus 103 

P. «urg;rfui92, 103 

Panoplax 115 

P. depressa 115 

Paraclaea 88, 103 

P. rufopunciala nodosa 88, 

103 

Paraliontera 104 

P. dispar 104 

P. longirnana 104 

Parapinnixa 122 

P. bouvieri 122 

P. hendersoni 122 

parrot crab 32 

Parlhenope 68 

P. (Parlhenope) agonus 68 

(Plalylambrus) fraterculus 

68 

P. (Plalylambrus) pourtalesii 

69 

P. (Plalylambrus) serrala 69 

PARTHENOPIDAE 9, 11-13, 

33,67 

Parthenopinae 67 

pea crab 123,124 

Pe/w 65 

P. mutica 65 

pentagon crab 67 

Percnon 131 

P. g/i-feii 131 

Periceroida 50 

Persephona 39 

P. aquilonaris 39 

P. crinila 39 

P. mediterranea 37, 39 

P. punctata 39 

P. punctata aquilonaris 39, 40 

Philyrinae 38 

Phyjodius 90, 104 

P. rnaculalus 90, 104 

Picroceroides 60 

P. tubularis 60 

Pilumnoides 104 

P. nudifrons 104 

Pilumnoplax 115 

P. americana 114,115 

P. etoall5, 116,117 

P. raVzWa 115 

Pilumnus 105 

P. caribaeus 105 

P. dasypodus 105 

P. diomedeae 105 

P. floridanus 105 

P. gemmalus 106 

P. holosericus 106 

P. /acZe!isl06 

P. longleyi 106 

P. mariW 106, 107 

P. pannosus 106 

P. ray; 106,107 

P. spinosissimus 107 

Pinnaxodes 122 

P. floridensis 122 

Pinnixa 125 

P. chacei 125 

P. chaetoplerana 125 

P. cristata 126 

P. cylindrica 126, 128 

P.floridana 126,127 

P. leplosynaptae 127 

P. /wnz/ 127 

P. pearsei 127 

P. retinens 127 

P. sayana 128 

Pinnothereliinae 125 

Pinnotheres 123 

P. geddesi 123 

P. guerini 123 

P. hemphilli 123 

P. hirlimanus 123 

P. maculalus 123 

P. moseri 124 

P. ostreum 124 

P. ierre; 122, 125 

P. shoemakeri 125 

P. strombi 122, 125

PINNOTHERIDAE 9, 10,11- 

13, 119 

Pinnotherinae 119 

Pisinae 50, 62 

Pilho 61 

P. aculeata 61 

P. anisodon 61 

P. laevigata 61 

P. Iherminieri 61 

P. mirabilis 62 

Plagusiinae 131 

Plagusia 131 

P. depressa 131 

Planes 130 

P. cyaneus 130-131 

P. minutus 130,131 

Planuca 145, 149 

Plalyactaea 88, 107 

P. ie«gera88, 107 

Platychirograpsus 137 

P, spgciabilis 137 

P. typicas 137 

Plaiylamhrus 68 

Platypodia 108 

P. spectabilis 108 

Platypodiella 108 

P. spectabilis 108 

Podochela 46 

P. curvirostris 46 

P. gracilipes 47 

P. lamelligera 47 

P. macrodera 47 

P. n'i.?ei 47 

P. sidneyi 48 

Polybiinae 72 

PORTUNIDAE 9, 11-13, 72 

Portuninae 74 

PORTUNOIDEA 9, 72 

Portunus 82 

P. anceps 82 

P. binoculus 83 

P. depressijrons 83 

P. floridanus 83 

P. gibbesii 83, 86 

P. ordivayi 84 

P. iar; 77, 84 

P.ietae 85 

P. spinicarpus 83, 85 

P. spinimanus 84, 85 

P. ventralis 86 

P. vocans 86 

POTAMIDAE 8, 11-13, 72, 86 

PoLamocarcirms 87 

Potamon 87 

Potamonidae 87 

Prinoplax 116 

P.atlantica 116, 117 

PROSOPIDAE 8, 9,11-13, 21 

Pseudomedaeus 108 

P. agassizii 93, 108 

Pseudothelphusidae 8, 87 

purse crab 39 

Pyrornaia 48 

P. arachna 48 

P. cuspidata 48 

queen crab 88 

racing crab 144 

Ranilia 24 

/». constricta 24 

/». rnuricata 24 

RANINIDAE9, 11-13, 23 

RANINOIDEA 9, 23 

Raninoides 24 

/». bengdicti 25 

/». fossor 24 

/?. lamarcki 24 

/». loevis 25 

7i. louisianensis 25 

red crab, deep-sea 111 

red-jointed fiddler crab 143 

red land crab 140 

red spider crab 56 

red tourlourou 140 

Retroplumidae 9, 10 

Rhithropanopeus 108 

/?. /zarT-wz; 108 

/». harrissi Iridentatus 109 

river crab 137 

Robertsella 116 

i?. myszfcall5, 116, 117 

Rochinia 66 

/?. crassa 66 

fi. hyslrix 66 

/». lanneri 66 

i?. umbonata 67 

rock crab 129 

saber crab 137 

sally lightfoot 129 

sand crab 74, 141 

sand fiddler crab 144, 145 

sargassum crab 84 

Sesarma 18, 133 

S. (Holometopus) arnericanum 

133, 134, 135 


S. (Holometopus) angustipes 

133, 134, 135 

S. (Holometopus) benedicti 

133 

S. (Holometopus) miersii 134 

S. (Holometopus) cinereum 

18,133-134, 135 

S. (Holometopus) miersii 134 

S. (Holometopus) ricordi 18, 

134, 135 

S. (Holometopus) roberti 

133, 134, 135 

6'. (Holometopus) tampicense 

133, 135 

5. (Sssarma) curacaoense 136 

S. (Sesarma) reticulatum 

18,136 

Sesarminae 132 

sliame-faced crab 30, 32 

shore crab 130 

Solenolambrus 70 

S. decemspinosus 70 

S. tenellus 70 

S. typicus 70 

speckled crab 74 

Speloeophorus 36 

S. elevatus 36 

S. nodosus 36 

S. poniifer 36 

Speocarcimus 116 

S. carolinensis 116 

S. Zofea/i/i 116 

spider crab (see MAJIDAE) 

sponge crab 52 

spray crab 131 

square-backed fiddler crab 133 

Stenocionops 50, 58 

S. furcata coelata 59 

S. furcata furcata 58 

S. spinimana 59 

S. spinosissima 59 

Stgnorhynchus 49 

S. lanceolatus 49 

S. seticornis 49 

stone crab 96 

swimming crab (see 

PORTUNIDAE) 

Symethis 23 

S. variolosa 25 

Teleophrys 60 

r. ornatus 60 

Temnonotus 49


T. granulosus 49-50 

T. simplex 49, 60 

Tetraplax 116 

T. quadridentata 116 

Tetraxanthus 109 

T.hidentaius\m,\\Q 

T.rugosus 109, 110 

T. rathbunae 109 

Thalassoplex 117 

T.arigusta lis, 117 

Thelxiope 22 

T. barbala 22 

r. vigil 22 

T/zoe 60 

T. puella 60 

Thyrolarnbrus 70 

T. astroides 70 

tourlourou 138 

Trnchycarcinus 71 

T. spinulifer 71 

TrapezzopfezlH, 116, 117 

T.trideniata\l4;li6,ti7 

tree crab 128, 132 

Trichodactylidae 8, 87 

Trichopeltarion 71 

r. /2oWte 71 

Troglocarcinus 41 

T. iafof 41 

T. corallicola AA 

Tutankhamen 71 

T. cristaiipes 71 

Tyc/it? 62 

T. emarginata 62 

T. lamellifrons 62 

Typhlopseudolhelphusa 87 

Z7cal0, 13, 18,142, 150 

f/, ff/jTzn/i 142 

C/. burger si 1'1'2, 144 

C/. cumulanla 148 

C/. leplodaclyla 143 

f/. longisignalis 18, 143, 147, 

149 

C/, mz«az 143,144, 147 

[/. rnordax 142, 144, 149 

C/. murifecenla 149 

[/. panacea 18, 144, 145 

C/. pugilator 18, 144, 145- 

146 

[/. joufe'«ax 18, 143, 147,149 

[/. pugnax repax 143, 144, 

147 

C/. pugnax virens 144, 1'1'9 

Z7. rapax 142,143, 147,149 

V. rapax longisignalis 143, 

141s 147 

U. speciosa 148 

[/. speciosa spinicarpa 148 

i7. spinicarpa 148 

C/. subcylindrica 18, 148 

C/. thayeri 148 

C/. yzVeni' 18,143, 147, 149 

U. vocator 144, 149 

U aides 130 

Z7. cordatus 150 

[/. occidenlalis 150 

C//zfoi 36 

[/. limbaius 36 

Varuninae 136 

Virginian Province 11 

wharf carb 130,133 

white land crab 138 

whitespot crab 140 

wood crab 133 

XANTHIDAE 9, 10,11-13, 

72,87,111,112 

Xantho 110 

X. denliculala 110 

Xanthodes bidentatus 110 

Xanthodius 110 

X. denticulatus 110 

JC. slimpsoni 110 

XANTHOIDEA 8, 9, 72,86 

yellow box crab 31, 32

CRABS (BRACHYURA) OF THE GULF OF MEXICO

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