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<strong>NMBAQC</strong> workshop 2010<br />

<strong>CUMACEA</strong><br />

Identification guide to British cumaceans<br />

Salma Shalla<br />

July 2011<br />

Atlantistylis borealis Shalla and Bishop 2005


Table of Content<br />

Introduction 3<br />

Morphology 6<br />

Sexual dimorphism 13<br />

Biology 14<br />

Classification 16<br />

Key to the families 16<br />

Family DIASTYLIDAE 19<br />

Family: LAMPROPIDAE 27<br />

Family PSEUDOCUMATlDAE 32<br />

Family BODOTRIIDAE 35<br />

Family NANNASTAClDAE 39<br />

Family LEUCONIDAE 45<br />

References 46<br />

2


Introduction<br />

Cumacea are a very distinctive order of the Class Malacostraca, in the Superorder Peracarida,<br />

distributed from the intertidal zone to the deep sea. Oniscus scorpioides (later re-named<br />

Diastylis scorpioides) was the first cumacean to be described, in 1780. At that time, many<br />

scientists thought that cumaceans were a larval stage of decapods. In 1846, they were<br />

recognized as a separate order by Krøyer. Twenty-five years later about fifty different species<br />

had been described. Since then, the number of known species has dramatically increased, and<br />

is still on the rise. In 1976 Jones listed 885 described species, in 1997 Watling and McCann<br />

listed over 1200 species and currently there are more than 1,593 described species (Anderson,<br />

2010). Although that increase was largely due to the species described from the deep-sea and<br />

Antarctic expeditions, nevertheless many new species were also discovered in shallow<br />

waters, especially in tropical areas.<br />

Cumaceans are small infaunal crustaceans, generally 1-10mm in length, although some<br />

species can reach up to 3.5cm. Typical cumaceans are easily recognizable by the combination<br />

of the following features (Figure 1): an inflated carapace, consisting of the head fused to at<br />

least the first three thoracic somites; anterior margin of the carapace usually extending in<br />

front of the head as a pseudorostrum; second antenna lacking an exopod; pereon composed of<br />

five free thoracic somites; pleon slender and composed of six somites ending in a forked tail.<br />

The telson may be present, reduced, or incorporated into the last abdominal somite.<br />

All the British species are marine, scarcely penetrating into estuaries. Elsewhere, the group is<br />

mainly marine, with only a few adapted to brackish and fresh waters. Relatively few live<br />

between tidemarks while many species are to be found in shallow water offshore, especially<br />

in the tropics, but they are also common in the deep sea, with a few recorded from below<br />

7000 m.<br />

The British cumaceans included by Jones (1976) were those recorded from the continental<br />

shelf within 20 miles of the British coasts and down to 200m depth, and comprised 41<br />

species. An additional 32 species which occur on the upper slopes down to 500m to the west<br />

of the British Isles are included here. The aim of this work is to update Jones (1976), so most<br />

of the keys are adapted from Jones. A few of the figures were also adapted from Sars (1900).<br />

History of cumacean taxonomy in the Northeast Atlantic<br />

Exploration in the deep North East Atlantic started with the “Lightning” cruise in the summer<br />

of 1868 and the “Porcupine” cruises in the summers of 1869 and 1870, led by Wyville<br />

Thomson, a Professor at Edinburgh. The cruises took place in the areas lying to the north and<br />

west of Britain and the Iberian Peninsula. The Rev. A. M. Norman (1879) described 30<br />

species of cumaceans from the „Lightning‟, „Porcupine‟ and „Valorous‟ expeditions, of<br />

which 9 were new species, and 2 new genera. On the basis of these cruises, and with the cooperation<br />

of the Royal Navy, the pioneering voyage of H.M.S. “Challenger” took place<br />

between 1872 and 1876 partly in the Atlantic but also in the Pacific and Southern Ocean.<br />

This was co-organised and led by Wyville Thomson. Fifteen new species of cumaceans were<br />

obtained during the Challenger voyage (Sars, 1887).<br />

Calman (1905) published a description of the cumaceans of the West Coast of Ireland taken<br />

by the “Helga” down to 400 fathoms during the years 1899-1904. Deeper stations produced<br />

3


icher yields (18 and 19 species). In total 48 species were identified, nine being regarded as<br />

new species, of which three represented new genera and one a new family.<br />

The “Ingolf”, “Thor” and other Danish expeditions around the end of the nineteenth century<br />

sampled the seabed in the north Atlantic around the Faeroes, Iceland and Greenland,<br />

recording 66 species, 24 of which were new to science (Hansen, 1920).<br />

The Voyages of Prince Albert I of Monaco took place between 1886 and 1912 in the Bay of<br />

Biscay, off Portugal and near the Azores. Forty species of cumaceans were identified, of<br />

which seven were new (Fage, 1929). The end of this era was completed some decades later<br />

with the Danish round-the-world voyage of the “Galathea” expedition 1950-1952, when the<br />

Bay of Biscay was one of the stations. Thirty species of cumaceans were identified of which<br />

26 species were new and one was referred to a new genus (Jones, 1969). From this time<br />

onwards it was widely accepted that the deep sea was much more diverse than previously<br />

thought.<br />

More recently, to the south west of Ireland, the Institute of Oceanographic Sciences Deacon<br />

Laboratory (IOSDL) carried out an intensive survey of the benthic biology of the Porcupine<br />

Seabight between 1977 and 1986. However, the most extensive deep-sea time series<br />

collection was made by the Scottish Association of Marine Sciences (SAMS) starting 1973 in<br />

the Rockall Trough. Also a shorter time series was taken from the Anton Dohrn SeaMount<br />

since 1980. Primarily from SAMS‟s collection and others, Norman Jones described a large<br />

number of new species, mainly in the family Nannastacidae. A survey of the area around the<br />

Faroe Islands (BIOFAR programme) was carried out between 1987 and 1990 (see Norrevang<br />

et al 1994 for station data). The result of that produced 62 cumacean species, of which 42 are<br />

additional species to the listed 28 species by Hansen (1920) and Stephensen (1929). One of<br />

the 42 additional species is a new leuconid described by Watling and Gerken (1999). That<br />

increases the Faroe cumacean list to 70 species by Gerken and Watling (1999). The Benthic<br />

Invertebrates of Icelandic waters (BIOICE) Programme commenced in 1992 and sampling<br />

continued until 1997. Its main objective was to revise the systematics of the marine<br />

invertebrates of the area. More than 1000 samples were collected around Iceland from 20m<br />

down to 2400m depth. All the cumacean information relating to the BIOICE programme<br />

remains unpublished.<br />

In 1996, 20,000 square kilometres of seabed lying to the west of the Shetland which was<br />

licensed for oil and gas exploration before or during 1996, was mapped and sampled by<br />

Atlantic Frontier Environmental Network (AFEN). In 1998, a further 10,000 square<br />

kilometres north and west of Scotland that covered the 17 th U.K. oil licensing round was<br />

surveyed (see Bett 1997 and 1999 for the survey area and sampling details). From both<br />

surveys 102 cumacean species were identified, 27 of which are of taxonomic importanceeither<br />

new species, or species which were poorly described and causing taxonomic problems<br />

(Shalla, 2001). Out of the 27 problematic species, four new species of the genus Leucon were<br />

described by Shalla and Bishop (2004); one new species of Atlantistylis was described,<br />

constituting the first record of the genus from the North Atlantic (Shalla and Bishop, 2005)<br />

and two new species of the genus Hemilamprops and Mesolamprops were described by<br />

Shalla and Bishop (2007).<br />

At the end of year 2000, the UK Department of Trade and Industry (DTI) conducted a survey<br />

in the area northwest of Scotland in the Faeroe-Shetland channel (2000 “White Zone”<br />

survey). Fifty-two cumacean taxa were recorded from depths of 141-1697m, 11 of which<br />

were of taxonomic importance-either new species, or species which were poorly described<br />

4


and causing taxonomic problems. In 2002, the DTI carried out another survey in the area<br />

north and west of Orkney and Shetland; no reports from this survey have been published.<br />

In 2003, a survey of the north east of the Porcupine Bank, together with other Irish and North<br />

Sea material, lead to the erection of the new genus Monopseudocuma by Mccarthy et al<br />

(2006).<br />

5


Morphology<br />

The cumacean body is divided into three distinct parts, the inflated carapace, the pereon<br />

(thorax) and the pleon (abdomen) (Fig. 1).<br />

The carapace consists of the cephalon (head) fused to at least the first three thoracic somites.<br />

The carapace extends both laterally and ventrally to enclose the branchial cavity. Each side of<br />

the carapace is produced anteriorly to form the pseudorostral lobes. Both lobes project<br />

forward and meet (but are not fused) to form the pseudorostrum (Fig. 1). Projecting<br />

anteriorly through the pseudorostrum is the siphon, which is the anterior part of the branchial<br />

apparatus of the first maxilliped. The siphon is usually short but can be very characteristically<br />

long in some species. In the genus Eudorella and Eudorellopsis, the pseudorostral lobes are<br />

reflexed upward with no obvious pseudorostrum but still with a siphon (Fig. 2). Below the<br />

pseudorostrum the anterior border is excavated to form the antennal notch or sinus which is<br />

defined below by the anterolateral corner, although in many species both are lacking. Behind<br />

the pseudorostrum on the dorsum is the bell-shaped frontal lobe from which the small ocular<br />

lobe projects forward, usually carrying the unpaired eyes (Fig. 1). The shape of the frontal<br />

lobe can vary from triangular to rounded, and the ocular lobe can vary in shape from small or<br />

absent to very long (Fig.2). The eye lobe normally persists (even in eyeless species), except<br />

when the eyes remain separated into two groups as in Nannastacus (Fig. 2). Behind and to the<br />

sides of the frontal lobes are the branchial regions (Fig. 1).<br />

The thorax (pereon) consists of five free thoracic somites (Fig. 1). Individually each is called<br />

a pereonite. Sometimes the first and exceptionally the second and third may be fused with<br />

the carapace, while the third and fourth may be fused together dorsally.<br />

The abdomen (pleon) consists of six cylindrical somites “pleonites” (Fig. 1). The fifth is the<br />

longest, while the sixth is prolonged by a freely articulated telson in five of the families. In<br />

the remaining three, the telson is fused with the sixth pleonite and is not distinct. The freely<br />

articulated telson varies in length and shape. The anus, protected by a pair of valves, opens<br />

on the lower face of the telson or at the end of the sixth pleonite.<br />

The body is covered with a chitinous epidermis, sometimes strongly calcified. Its surface is<br />

often sculptured with grooves, ridges, spines, tubercles or teeth, or has a fine reticulated,<br />

pitted or scaled appearance. It is sometimes covered with sand grains.<br />

The first antenna (antennule) is short (Fig. 3A). It has a peduncle of three articles and two<br />

flagella. The main outer flagellum is usually longer than the accessory inner flagellum, with<br />

two to six articles. The accessory flagellum often has only one article. One or two modified<br />

sensory setae ,“aesthetascs”, may be present at the end of the main flagellum. In males of<br />

some species (e.g. Leptostylis) a brush of sensory filaments is present on the basal articles of<br />

the main flagellum (Fig 3B). Between the bases of the first antenna on the underside of the<br />

front end of the carapace are two small chitinous plates (the epistome), to which the first<br />

antennae are attached (Fig. 3A).<br />

6


Carapace<br />

Pereon<br />

Pleon<br />

A B<br />

Pseudorostrum<br />

Eyelobe<br />

Frontal lobe<br />

Branchial area<br />

Pereonite 1<br />

Pereonite 5<br />

Pereopod 5<br />

Pleonite 1<br />

Pleonite 6<br />

Uropod peduncle<br />

Telson<br />

Exopod<br />

Endopod<br />

Antenna 1<br />

Antenna 2<br />

Pereopod 1<br />

Exopod<br />

Pereopod 5<br />

Pleopod 1<br />

Flagellum of Antenna 2<br />

Figur 1: Diastylis rathkei: A, female dorsal view; B, adult male lateral view (from Sars,<br />

1900).<br />

7


Figure 2: Examples of carapace shape and ornamentation.<br />

8


The second antenna (antenna) is generally rudimentary in females, with from one to five<br />

articles (Fig. 3A). In males, the shape and length changes at different developmental stages.<br />

In mature males (Fig. 3C), it is well developed with a peduncle of five articles and a manysegmented<br />

flagellum. The outer sides of the fourth and fifth articles of the peduncle are<br />

normally thickly covered with setae. The second antenna is bent backwards between the third<br />

and fourth articles of the peduncle, and the flagellum is closely applied to the sides of the<br />

body. It usually reaches to the end of the body but is sometimes shorter. In some species, the<br />

second antenna is used as an organ for clasping the female. The second antenna is attached to<br />

the upper lip (labrum) (Fig. 3A).<br />

The mouth parts consists the upper lip (labrum), paired mandibles, paired first and second<br />

maxillae and lower lip (labium) (Fig. 3A,D-H). The upper and lower lips are of no taxonomic<br />

significance.<br />

The mandibles (Fig. 3D&E) are simple and always without a palp. Each is normally<br />

boat-shaped, pointed at either end, with a strong molar process and toothed incisor process<br />

and in between them a row of setae and a distal lacinia mobilis (Fig.3D). In some families the<br />

mandibule is reduced to a truncate shape (Fig.3E). In Nannastacidae the mandible is<br />

modified to aid the predatory feeding habit, with a styliform molar process.<br />

The first maxillae (maxillulae) (Fig. 3F) lie ventral to the labium (Fig. 3H) and the second<br />

maxillae (Fig. 3G) ventral to the first pair. Both consist of a flattened protopodite bearing<br />

endites and the first pair have a backwardly directed palp, usually with one or two filaments,<br />

which vibrate in the respiratory chamber and cause a current of water to pass the gills.<br />

In cumaceans, the first three thoracic somites are covered by the carapace but only the first<br />

one is thought to be fused to the head. The first three pairs of thoracic appendages are<br />

maxillipeds and the last five are pereopods. Each has a short coxa fused with the sternite,<br />

followed by the basis, usually the longest article, which may carry an exopodite on its<br />

proximal part, normally with a peduncle and setiferous flagellum but sometimes a rudiment.<br />

Exopods are never present on the first and second maxilliped nor on the fifth pereopods, and<br />

the number on the other appendages varies with the species. In the female oostegites are<br />

borne on the coxae of the third maxilliped and the first three pairs of pereopods, lamellar in<br />

form and interlocking to form the brood chamber. The remaining articles are the ischium,<br />

merus, carpus, propodus and dactyl.<br />

The first maxillipeds are short, robust and highly developed (Fig.3I). The coxa carries a large<br />

epipod (the branchial apparatus). The anterior part of the epipod extends forward and,<br />

together with its counterpart, forms the siphon. Its posterior part forms finger-like branchial<br />

lobules (gills), often more numerous in the male than in the female. Water leaves the<br />

branchial chamber through the siphon.<br />

The second maxillipeds (Fig. 3J) usually have a long basis. The propodus and dactyl are bent<br />

in towards one another and may be modified, with large flattened spines. In the female there<br />

is a flattened plate attached to the hind end of the coxa, the rudimentary oostegite, which has<br />

a fringe of long slender setae directed backwards into the brood chamber.<br />

The third maxillipeds (Fig. 3K) are large, and partly cover the first and second pairs and the<br />

mouthparts. They are sometimes similar in structure to the first pereopods but usually differ<br />

considerably. Long feathered setae are present on the basis, acting as a screen to filter the<br />

9


water entering the branchial chamber.<br />

B<br />

K<br />

E<br />

A<br />

G<br />

D<br />

H<br />

epistome<br />

Labrum<br />

C<br />

F<br />

siphon<br />

I<br />

Branchial<br />

apparatus<br />

Figure 3: Detailed view of the head appendages, anterior thoracic appendages and pleopods. A, Antenna<br />

1 and 2 of female diastylid; B, antenna 2 of male Leptostylis; C, antenna 2 of male diastylid; D,<br />

mandible of diastylid; E, mandible of Campylaspis; F-G, maxilla 1 and 2 of Diastylid; H, Labium of<br />

diastylid; I-K; maxillipeds 1-3 of diastylid (all from Sars, 1900).<br />

J<br />

10<br />

coxa<br />

oostegites


The first two pairs of pereopods (Fig. 4) are directed forwards, the first normally reaching<br />

beyond the tip of the pseudorostrum. The second are usually shorter than the first and their<br />

ischia are often fused with the bases. The last three pairs of pereopods are directed<br />

backwards (Fig.4) and are used for burrowing.<br />

Pleopods (Fig. 4) have been reported in the female of only a single species. In the male there<br />

may be one, two, three or five pairs, placed at the hind ends of the pleon sternites, or they<br />

may be absent. The peduncle consists of a short coxa and a long basis, with two short rami,<br />

the outer usually with two articles and the inner with one article, normally carrying long<br />

plumose setae in the adult. The inner ramus is rudimentary in some species.<br />

The uropods are the last pair of abdominal appendages. Each consists of a uniarticulate<br />

peduncle and two rami, the outer ramus (exopod) with two articles and the inner ramus<br />

(endopod) with one to three articles (Fig. 4). Spines and setae are usually present on the<br />

edges of the peduncle and rami. The uropods are used as cleaning organs.<br />

11


carpus<br />

exopod<br />

propodus<br />

merus<br />

dactyl<br />

ischium<br />

basis<br />

Pereopod 1 Pereopod 2 Pereopod 3<br />

Pereopod 4<br />

Pleopod 1<br />

Pleopods<br />

Pleopod 2<br />

Exopod<br />

Uropod<br />

Figure 4: Thoracic and abdominal appendages: pleopods (top to bottom) of Pseudocumatidae,<br />

Diastylidae and Bodotriidae (from Sars, 1900); pereopods and uropods of Leuconidae (from<br />

Shalla and Bishop, 2003).<br />

Endopod<br />

12


Sexual dimorphism<br />

There is always some sexual dimorphism in adults, more pronounced in some families than<br />

in others, but almost always including: better development of the second antenna in the<br />

male; the presence of oostegites (brood pouch) in the female; the presence of pleopods<br />

(except in Nannastacidae) in the male; the presence of more or better-developed exopods on<br />

the thoracic appendages in the male, and sometimes differences in shape and ornamentation<br />

of the carapace, as well as differences in the development of some sense organs.<br />

General distinction between sexes in adult cumaceans:<br />

Female Male<br />

Antenna 2 Rudimentary Well developed, with long<br />

flagellum<br />

Oostegites or brood pouch Present Absent<br />

Pleopods No Yes (except in Nannastacidae)<br />

Exopods on thoracic<br />

appendages<br />

A2 of ♂ Leucon jonesi<br />

(from Bishop 1981)<br />

Less developed More developed (except<br />

Bodotriidae)<br />

13


Biology<br />

Cumaceans brood their eggs after fertilization in the brood pouch in which the embryos are<br />

retained and moult into a stage closely resembling the adult (manca) but lacking the last pair<br />

of pereopods, the secondary sexual characters, and some of the spines and setae. Several<br />

moults follow the manca before the adult stage is reached. The number of moults, the life<br />

cycle and the life span of a cumacean varies with species and environmental conditions (for<br />

detailed studies, see Forsman (1938), Kruger (1940), Zimmer (1941), Corey (1969, 1976,<br />

1983), Bishop (1982), Duncan (1984), Persson (1989), Roccatagliata (1991), Bishop and<br />

Shalla (1994) and Corbera et al. (2000).<br />

The key adopts the terminology of Bishop (1982) who summarised the developmental stages<br />

from a newly hatched manca to an adult in both sexes in the deep-sea species Leucon jonesi<br />

as follows:<br />

Manca: without pereopod 5 and without pleopods.<br />

Juvenile: with five pairs of pereopods and without external sexual differentiation.<br />

Immature males: pleopods not developed externally but visible through the cuticle, antenna 2<br />

in development but unsegmented.<br />

Preadult males: pleopods with peduncle and rami but no setae, antenna 2 in development and<br />

partially segmented with a differentiated peduncle<br />

Adult males: pleopods fully developed and antenna 2 flagellum fully segmented and longer<br />

than body length<br />

Immature females: without oostegites; without pleopods and antenna 2 rudimentary, as in the<br />

following stages.<br />

Preadult females: oostegites in development but not yet forming a brooding pouch.<br />

Brooding females: fully developed oostegites forming a marsupium containing individuals in<br />

different developmental stages.<br />

The majority of species occur in soft deposits and a number in sand where they may show a<br />

preference for a certain range of grain size (Foxon, 1936; Pike and Le Sueur, 1958), possibly<br />

related to the mesh size of their branchial filtering apparatus or the size of grain which they<br />

can manipulate during feeding. They normally burrow into the deposit by means of the last<br />

four pairs of pereopods, remaining at rest usually with only the front part of the carapace<br />

emerging and sometimes also the uropods, but they make fairly frequent swimming<br />

excursions and the adult males of many coastal species may swim up into the plankton,<br />

especially at night, and may rise almost to the surface to form swarms. They may be joined<br />

by smaller numbers of females but it is likely that females do not rise as far from the bottom<br />

nor remain above it for as long a time. The function of this behaviour is presumably to enable<br />

mating to take place, or at least for the males to find the females (from Jones, 1976).<br />

Cumaceans feed on micro-organisms and organic matter from the bottom deposit. Mud-living<br />

species filter small particles and those inhabiting sand clean their food off sand grains. In the<br />

latter case objects carrying food are picked up by the first pereopods and passed on to the<br />

third maxilliped. The food is then cleaned off by the first and second maxillipeds and passed<br />

on to the first and second maxillae and mandibles (Dixon, 1944). The food-manipulating<br />

appendages and mouthparts are suitably armed with spines and bristles. In the filter feeders<br />

14


the first and second maxillae together form a pump by means of which water is drawn<br />

through their setules which filter off particles of food (Dennell, 1934). In Campylaspis and<br />

some related genera the mandibles and second maxilliped are modified as piercing organs<br />

and they probably feed on foraminiferans and perhaps small crustaceans (from Jones, 1976).<br />

15


Family Bodotriidae<br />

Subfamily Vaunthompsoniinae<br />

Subfamily Bodotriinae<br />

Family Leuconidae<br />

Family Nannastacidae<br />

Family Pseudocumatidae<br />

Family Lampropidae<br />

Family Diastylidae<br />

Family Gynodiastylidae*<br />

Family Ceratocumatidae†<br />

Classification<br />

Although Cumaceans are a very recognizable group, their family distinctions are not as<br />

obvious. Various taxonomists have divided the order into 4-26 families, of which eight are<br />

currently accepted. The family definitions overlap to such an extent that the most difficult<br />

task in identification of a species is placement into a family (Day, 1980). A large part of the<br />

problem is due to the significant sexual dimorphism in cumaceans. Thus it is helpful to<br />

identify the sex and the stage of maturity of the specimen before using the key (see section on<br />

biology); but this is not always obvious and can mainly be learnt by experience.<br />

*Species are present predominantly in the southern hemisphere<br />

†British records from mainly deep-water<br />

Useful web links:<br />

http://www.marinespecies.org/cumacea<br />

http://peracarida.usm.edu/CumaceaTaxa.pdf<br />

Key to the Families (Fig. 5)/subfamilies*<br />

(*Applicable to the British species only)<br />

1- With freely articulated telson 2<br />

Without freely articulated telson 5<br />

2- Telson small or large, with 0 or 2 terminal spines 3<br />

Telson mostly well developed, always with 3 or more terminal spines Lampropidae<br />

3-Telson small, without apical spines. Endopod of uropod with only one article 4<br />

Telson larger, with 2 apical spines. Endopod of uropod with 2 or 3 articles<br />

Diastylidae<br />

16


4- ♂ with 3, 4 or 5 pairs of pleopods, pleopods with two rami<br />

Ceratocumatidae†<br />

♂ with 1 or 2 pairs of rudimentary pleopods, pleopods with one ramus (Figure 4)<br />

Pseudocumatidae<br />

5- In both sexes the last four pairs of pereopods either without exopods or with very small<br />

rudimentary exopods. ♂ with 5 pairs of pleopods<br />

Bodotriidae- Bodotriinae<br />

In the female at most the last three pairs of pereopods without exopods, in the male only the<br />

last pair. ♂ with 0, 2 or 5 pairs of pleopods 6<br />

6- Uropodal endopod uniarticulate. In ♀ the last three pairs of pereopods without exopods, in<br />

♂ only the last pair. ♂ without pleopods Nannastacidae<br />

Uropodal endopod with two articles. In ♀ the last two pairs of pereopods without exopods, in<br />

♂ the last pair. ♂ with 5 or 2 pairs of pleopods 7<br />

7- Eyes distinct. ♂ with 5 pairs of pleopods. In ♀ exopods of pereopods 1 well developed, of<br />

pereopods 2-3 well developed or rudimentary. Mandible of normal shape<br />

Bodotriidae-Vaunthompsoniae<br />

Eyes wanting. ♂ with 2 pairs of pleopods. In ♀ exopods of pereopods 1-3 well developed.<br />

Mandibles truncate at base Leuconidae<br />

*Key modified from Jones (1976) and Watling and McCann (1997)<br />

†British records are mainly deep-water, not discussed in the following key.<br />

17


Figure 5: Representative cumaceans from the families included in the present key.<br />

Diastylidae ♀<br />

Lampropidae ♀<br />

Pseudocumatidae<br />

Bodotriidae ♀<br />

Bodotriidae ♂<br />

Nannastacidae ♀<br />

Leuconidae ♀<br />

18


Family DIASTYLIDAE<br />

Diagnosis (as emended by Day, 1980 and Watling and McCann, 1997): Mandibles normally<br />

boat-shaped but widened at base in Diastyloides. First maxilliped branchial filament with<br />

numerous leaflets. Exopods present on maxilliped 3 (except in Paradiastylis) and in females<br />

on pereopods 1 and 2, absent or rudimentary on pereopods 3 and 4, in males present on<br />

pereopods 1-4. Telson variable, usually large, often with a long post-anal part, or short and<br />

poorly armed; bearing one pair of terminal spines or none. Uropods usually long and slender,<br />

endopod with 3 articles. Male antenna 2 with long flagellum reaching at least to the posterior<br />

end of thorax. Males with two pairs of pleopods (except in Atlantistylis which has no<br />

pleopods); no outer process on inner ramus.<br />

Remarks: The Diastylidae is one of the oldest and biggest cumacean families, and is well<br />

represented in the temperate North Atlantic. The family contains around 285 species in 20<br />

genera, with very clear sexual dimorphism. Identification of male specimens is very difficult<br />

especially if the specimen is not at the adult stage, the present key still need to improve on it.<br />

Jones (1976) listed 12 species in 4 genera from depths down to 200m, while the present key<br />

extends the depth range into the lower slopes down to 500m and contains 20 species in 5<br />

genera.<br />

Diastyloides biplicatus (G.O. Sars, 1865)<br />

Diastyloides serratus (G.O. Sars, 1865)<br />

Diastylis boecki Zimmer, 1930<br />

Diastylis bradyi Norman, 1879<br />

Diastylis cornuta (Boeck, 1864)<br />

Diastylis echinata Bate, 1865<br />

Diastylis glabra Zimmer, 1900<br />

Diastylis goodsiri (Bell, 1855)<br />

Diastylis laevis Norman, 1869<br />

Diastylis lucifera (Krøyer, 1837)<br />

Diastylis rathkei (Krøyer, 1841)<br />

Diastylis rugosa Sars, 1865<br />

Diastylis tumida (Liljeborg, 1855)<br />

Brachydiastylis resima (Krøyer, 1846)<br />

Leptostylis ampullacea (Liljeborg, 1855)<br />

Leptostylis longimana (Sars, 1865)<br />

Leptostylis macrura Sars, 1870<br />

Leptostylis villosa Sars, 1869<br />

Makrokylindrus (Adiastylis) josephinae (Sars, 1871)<br />

Makrokylindrus (Adiastylis) longipes (Sars, 1871)<br />

19


Key to the British Species of Diastylidae*<br />

1- Mandibles broad at base, basis of pereopod 2 usually abruptly wider than ischium with<br />

one or two large strong teeth at lower distal corner (Diastyloides) 2<br />

Mandibles narrow at base, basis of pereopod 2 narrow distally or abruptly wider than ischium<br />

but without one or two large strong teeth at lower distal corner 3<br />

2- Carapace with diagonal folds, telson (excluding spines) more than half as long as the<br />

peduncle of the uropod Diastyloides biplicata<br />

Carapace without diagonal folds, telson (excluding spines) about half as long as the peduncle<br />

of the uropod Diastyloides serrata<br />

3-Telson short, shorter than the last abdominal somite (Leptostylis) 4<br />

Telson long, longer than the last abdominal somite 7<br />

4- Lower front edges of carapace with flat-topped teeth Leptostylis villosa<br />

20


Lower front edges of carapace with triangular teeth 5<br />

5- Pereopod 1 with the basis little longer than the carpus<br />

Leptostylis longimana<br />

Pereopod 1 with the basis much longer than the carpus 6<br />

6- Pereopod 1with the basis as long as the carpus, propodus and dactylus combined, peduncle<br />

of the uropod not much longer than its endopod<br />

Leptostylis ampullacea<br />

Pereopod 1with the basis much shorter than the carpus, propodus and dactylus combined,<br />

peduncle of the uropod much longer than its endopod<br />

Leptostylis macrura<br />

Pereopod 1of :<br />

L. macrura L. ampullace L. longimana<br />

Uropod of:<br />

7- Post-anal part of telson shorter than pre-anal part<br />

(Makrokylindrus) 8<br />

Post-anal part of telson longer than or equal to pre-anal part 9<br />

8- Post-anal part of the telson with at most one pair of lateral spines<br />

Makrokylindrus josephinae<br />

Post-anal part of the telson with at least three pairs of lateral spines<br />

Makrokylindrus longipes<br />

L. macrura L. ampullacea<br />

21


9- Pseudorostrum of ♀ strongly upturned, of ♂ slightly upturned, pereopods 2 and 3 widely<br />

separated in ♀and to a lesser extent in ♂ Brachydiastylis resima<br />

Pseudorostrum not strongly upturned, pereopods 2 and 3 not widely separated<br />

(Diastylis) 10<br />

10- Pleopods absent or ,when present, without feathered setae (♀♀ and immature ♂♂)<br />

11<br />

Pleopods present and with long feathered setae (adult ♂♂) 20<br />

11- Carapace with 3-5 vertical folds extends to the posterior half of the carapace and two<br />

pairs of strong dorsal spines<br />

Diastylis rugosa<br />

Carapace without distinct folds or with 1-2 indistinct folds in the anterior part of the carapace<br />

12<br />

12- Carapace with a number of large teeth 13<br />

Only small teeth or none on the carapace 14<br />

13- Read three alternatives<br />

One very prominent pair of teeth and c14 smaller ones on the carapace Diastylis cornuta<br />

One prominent pair of teeth and fewer c6 smaller ones on the carapace Diastylis boecki<br />

Prominent pair of teeth absent. Carapace divided into polygonal areas by numerous teeth,<br />

spines and spinules Diastylis echinata<br />

14- Carapace thickly covered with hairs Diastylis goodsiri<br />

Carapace not thickly covered with hairs 15<br />

D. cornuta D. boecki D. echinata<br />

22


15. The pre-anal part of the telson about as long as the post-anal part Diastylis tumida<br />

The pre-anal part of the telson distinctly shorter than the post-anal part 16<br />

16- Telson with no more than 4 lateral spines on either side Diastylis lucifera<br />

♀<br />

Telson with at least 6 lateral spines on either side 17<br />

17- Two long rows of small teeth on the frontal lobe 18<br />

Where there are teeth on the frontal lobe they are arranged transversely 19<br />

18- Carapace less than one and half as long as broad Diastylis glabra<br />

Carapace more than one and half times as long as broad Diastylis rathkei<br />

propod<br />

us<br />

Diastylis glabra<br />

Diastylis rathkei<br />

19- Prolongation of the hind end of pereonite 5 ending in a short blunt point. Propodus of<br />

pereopod1 about twice as long as dactylus Diastylis laevis<br />

23


Prolongation of hind end of pereonite 5 ending in a long sharp point. Propodus of pereopod 1<br />

much less than twice as long as dactylus Diastylis bradyi<br />

Diastylis laevis<br />

propodus<br />

dactylus<br />

20- No lateral ridges on the carapace 21<br />

Lateral ridges, which may be only faintly defined, present on carapace 22<br />

Diastylis bradyi<br />

21- First article of uropod endopod shorter than the other two articles together<br />

Diastylis lucifera<br />

First article of uropod endopod longer than the other two articles together<br />

Diastyloides serrata<br />

22- No vertical or diagonal ridges nor pseudorostral ridges on the carapace<br />

23<br />

Vertical or diagonal ridges present on carapace, or where absent at least pseudorostral lines<br />

present 28<br />

23- Subrostral lines present or the lateral ridges extend to the subrostral margin of the<br />

carapace 24<br />

Subrostral lines absent, the lateral ridges not reaching to the subrostral margin of the carapace<br />

27<br />

24- No protuberances on the carapace 25<br />

Protuberances present on the carapace in the position where large teeth are present in the<br />

females 26<br />

25- Telson with not more than 4 lateral spines, 3 teeth on the frontal lobe<br />

Diastylis lucifera<br />

24


Telson with more than 4 lateral spines, no teeth on the frontal lobe<br />

Diastylis tumida<br />

26- 2 pairs of protuberances on the carapace Diastylis cornuta<br />

Only one pair of protuberances on the carapace Diastylis boecki<br />

27- Carapace thickly covered with hairs, length over 25mm Diastylis goodsiri<br />

Carapace not thickly covered with hairs, length about 8mm Diastylis lucifera<br />

28- The pseudorostral lines form an angle broadly open towards the underside, propodus of<br />

pereopod 1 about twice as long as dactylus Diastylis laevis<br />

The pseudorostral lines run otherwise or absent 29<br />

29- Pereonites 2-4 without depressions in the mid-line 30<br />

Pereonites 2-4 depressed in the mid-line so that two distinct but low keels are formed<br />

32<br />

30- No pseudorostral lines 31<br />

Pseudorostral lines present Diastylis bradyi<br />

31- 2-3 vertical lines on the carapace Diastylis rugosa<br />

Only 1 oblique line on the carapace Diastylis boecki<br />

32- Pseudorostrum short Diastylis rathkei<br />

Pseudorostrum long Diastylis glabra<br />

*Key modified from Jones 1957 and 1976<br />

25


Makrokylindrus longipes<br />

Immature ♂<br />

Makrokylindrus josephinae ♀<br />

Diastylis echinata<br />

Brooding ♀<br />

Diastylis cornuta<br />

Brooding ♀<br />

26


Family: LAMPROPIDAE<br />

Diagnosis (Day, 1978, Watling and McCann, 1997): Antenna 1 with well developed<br />

flagellum. Mandibles boat-shaped. Maxilla 1 palp absent or bearing 1 or 2 setae. Exopods<br />

present on maxillipeds 3 and pereopod 1 in both sexes; in female present or rudimentary on<br />

pereopods 2-4 or absent from all three; in male exopods present on pereopods 2-4. Pleopods<br />

absent or 1-3 pairs in male, with outer process on inner ramus. Telson moderate to large, well<br />

developed post-anally, with 3-8 terminal and subterminal spines.<br />

Remarks: The family Lampropidae is clearly distinguished by the presence of at least 3<br />

terminal spines on the telson. The family contains around 100 species in 14 genera and is<br />

well represented in the North Atlantic. Of these, Chalarostylis Norman, 1879, Lamprops<br />

Sars, 1863, Hemilamprops Sars, 1883, Mesolamprops Given, 1964, Paralamprops Sars,<br />

1887, and Platysympus Stebbing, 1912 are discussed here. The genus Chalarostylis is defined<br />

by a short telson, a robust first pereopod with a group of long setae on the dactylus, and 3<br />

pairs of pleopods in males. The genera Lamprops, Hemilamprops and Mesolamprops are<br />

very similar; males are readily separated by the absence of pleopods in Lamprops versus the<br />

presence of three and two pairs of pleopods in Hemilamprops and Mesolamprops<br />

respectively. Ascribing females to genus without reference to the corresponding males is<br />

more problematic but, as noted by Haye & Gerken (2005), Hemilamprops and Mesolamprops<br />

both have the basis of the first pereopod much shorter than the rest of the appendage, whereas<br />

in Lamprops it is equal in length to the rest of the appendage. The genus Platysympus is well<br />

defined by the strongly flattened carapace with a pronounced marginal carina, the absence of<br />

exopods on pereopods 2 to 4 in females and the presence of 3 pairs of pleopods in males. The<br />

genus Paralamprops is defined by a broad depressed carapace, with a marginal carina,<br />

exopods on pereopods 3 to 4 rudimentary or absent in females and the presence of 3 pairs of<br />

pleopods in males. Jones (1976) listed 2 species in 2 genera from depths down to 200m,<br />

while the present key extends the depth range to the lower slopes down to 500m and contains<br />

13 species in 6 genera.<br />

Chalarostylis elegans Norman, 1879<br />

Hemilamprops assimilis Sars, 1883<br />

Hemilamprops cristatus (G.O. Sars, 1870)<br />

Hemilamprops normani Bonnier, 1896<br />

Hemilamprops pellucidus Zimmer, 1908<br />

Hemilamprops pterini Shalla and Bishop, 2007 Atlantic<br />

Hemilamprops roseus (Norman, 1863)<br />

Hemilamprops uniplicatus (G.O. Sars, 1872)<br />

Mesolamprops denticulatus, Ledoyer, 1983<br />

Mesolamprops hartleyi Shalla and Bishop, 2007<br />

Paralamprops orbicularis (Calman, 1905)<br />

Platysympus typicus (Sars, 1870)<br />

Lamprops fasciata Sars, 1863<br />

27


Key to the British Species of Lampropidae<br />

1- Telson small, sub equal in length to pleonite 6 and less than half the length of the peduncle<br />

of the uropod Chalarostylis elegans<br />

From Gerken and McCarthy (2007)<br />

Telson long, longer than pleonite 6 and more than half the length of the peduncle of the<br />

uropod 2<br />

2- Basis of pereopod 4 longer than entire length of pereopod 5. Males with 3 pairs of<br />

pleopods Paralamprops orbicularis<br />

Basis of pereopod 4 subequal to, or shorter than, entire length of pereopod 5. Males with 0, 2<br />

or 3 pairs of pleopods 3<br />

3- Carapace strongly flattened with a pronounced marginal carina . Males with 3 pairs of<br />

pleopods Platysympus typicus<br />

Carapace not as above. Males with 0, 2 or 3 pairs of pleopods 4<br />

4- Read three alternatives<br />

Male without pleopods, antennal notch small but distinct, eyes present. Basis of pereopod 1<br />

approximately equal in length to the rest of the appendage. Telson with 5 apical spines, the<br />

median being the longest. Carapace with 3 oblique folds Lamprops fasciata<br />

Male with 2 pairs of pleopods, antennal notch present or absent, eyes present or absent. Basis<br />

of pereopod 1 shorter than the rest of the appendage, carapace without oblique folds<br />

(Mesolamprops) 5<br />

Male with 3 pairs of pleopods, antennal notch usually absent, eyes present or absent. Basis of<br />

pereopod 1 distinctly shorter than the rest of the appendage, carapace with or without oblique<br />

folds (Hemilamprops ) 6<br />

28


5- Dorsal crest denticulate in the anterior half of carapace including eye lobe. Telson slightly<br />

over half but less than three-quarters the length of uropodal peduncle<br />

Mesolamprops denticulatus<br />

Dorsal crest, including the eye lobe smooth, telson slightly over three-quarters the length of<br />

uropodal peduncle Mesolamprops hartleyi<br />

6- Carapace with no lateral folds, telson with three or more apical stout setae 7<br />

Carapace with lateral folds, telson with three apical stout setae 10<br />

29


7- Dorsal crest smooth 8<br />

Dorsal crest serrated 9<br />

8- Telson with eight apical and two pairs of lateral stout setae<br />

Hemilamprops roseus<br />

Telson with six apical and one pair of lateral stout setae<br />

Hemilamprops assimilis<br />

9- Telson with five apical and six pairs of lateral stout setae<br />

Hemilamprops normani<br />

Telson with three apical and seven pairs of lateral stout setae<br />

Hemilamprops pellucidus<br />

Telson with three apical and two to three pairs of lateral stout setae<br />

Hemilamprops cristatus<br />

10- Carapace with prominent lateral fold on each side; branchial region not inflated. Telson<br />

subequal in length to the peduncle of the uropod, with three apical and five to seven pairs of<br />

lateral stout setae Hemilamprops uniplicatus<br />

30


Carapace with relatively inconspicuous lateral folds; branchial region inflated on each side.<br />

Telson about three quarters the length of the peduncle of the uropod, with three apical and<br />

two to three pairs lateral stout setae 11<br />

11- Dorsal crest including eye lobe, antero-lateral corner of carapace and basal part of telson<br />

serrated Hemilamprops cristatus<br />

Hemilamprops cristatus ♀<br />

Dorsal crest including eye lobe, antero-lateral corner of carapace and basal part of telson<br />

smooth Hemilamprops pterini<br />

31


Family PSEUDOCUMATlDAE<br />

Diagnosis (as emended from Jones, 1976 and McCarthy et al, 2006): Telson present but<br />

small. Males with one pair of pleopods or two, the second pair somewhat rudimentary.<br />

Pleopods monoramous. Males with exopods well developed on maxilliped 3 and pereopods<br />

1-4. Females with exopods well developed on maxilliped 3 and pereopods 1-2, and<br />

rudimentary on pereopods 3-4. The uropodal endopod with a single article.<br />

Remarks: Pseudocumatidae is a small family with only three genera present in British<br />

shallow waters. The new genus Monopseudocuma was initiated by McCarthy et al (2006) to<br />

accommodate Pseudocuma gilsoni which differed from the rest of the species in the genus<br />

Pseudocuma by the following combination: 1 pair of pleopods in the male; pereopod 1<br />

normal, fully developed exopods on pereopods 1-4 in males and 1-2 in females, rudimentary<br />

exopods on pereopods 3-4 in females. Antenna 2 in female uni-articulate. Uropods long and<br />

slender. Jones (1976) listed 4 species in 2 genera from depths down to 200m, while the<br />

present key extends the depth range to the lower slopes down to 500m but has no additions to<br />

Jones‟ Pseudocumatid list.<br />

Petalosarsia declivis (Sars, 1865)<br />

Monopseudocuma gilsoni (Gilson, 1906)<br />

Pseudocuma longicornis (Bate, 1858)<br />

Pseudocuma similis Sars, 1900<br />

32


Key to the British Species of Pseudocumatidae<br />

1- The first pereopod shortened and peculiarly formed with expanded flattened carpus<br />

Petalosarsia declivis<br />

♀<br />

♂<br />

The first pereopod not shortened, of normal shape 2<br />

2- With 1 or 2 pairs of pleopods, exopods well developed on maxilliped 3 and pereopods 1-4<br />

(males) 3<br />

Without pleopods, exopods well developed on maxilliped 3 and pereopods 1 and 2 but<br />

rudimentary on pereopods 3 and 4 (females) 5<br />

3- With 1 pair of pleopods. Antenna 2 flagellum extends to the end of pereon, or just beyond.<br />

Uropod peduncle with only one inner spine: endopod with 5-7 stout setulose setae and very<br />

fine setae in between Monopseudocuma gilsoni ♂<br />

With 2 pairs of pleopods. Antenna 2 flagellum long, extends to at least pleonite 5<br />

4<br />

4- Anterolateral angle of the carapace armed with 3 teeth. Uropod peduncle with 1 simple<br />

seta Pseudocuma similis ♂<br />

33


Anterolateral angle without teeth. Uropod peduncle with 4 plumose setae<br />

Pseudocuma longicornis ♂<br />

5- Anterolateral angle of the carapace armed with 3 teeth. Telson broader than long, truncate<br />

Pseudocuma similis♀<br />

Anterolateral angle without teeth. Telson nearly semi- circular 6<br />

6- Uropod peduncle shorter than endopod. Endopod with 5-10 stout setulose setae<br />

Pseudocuma longicornis♀<br />

Uropod peduncle equal to endopod. Endopod with 3-4 stout setulose setae<br />

Monopseudocuma gilsoni♀<br />

34


Family BODOTRIIDAE<br />

Diagnosis (emended from Jones, 1976, Watling 1977 and Day, 1978): No articulate telson.<br />

Pleopods (males only) with a prolonged process on the outer edge of the inner ramus (Fig. 4),<br />

usually 5 pairs, occasionally 2 or 3 pairs. Exopods present on the third maxilliped and at least<br />

pereopod 1, but may occur on pereopods in the following combinations: in the male, 4 or 1,<br />

occasionally 2 or 3 fully developed pairs, 1 and 2 rudimentary, 2 and 2 rudimentary, or 3 and<br />

1 rudimentary; in the female, 3 or 1, occasionally 2 fully developed pairs, 3 and 1<br />

rudimentary, 2 and 2 rudimentary, 2 and 1 rudimentary, or 1 and 2 rudimentary. The<br />

mandibles are not broadened at the base. The number of free thoracic somites is often<br />

reduced.<br />

Remarks: The family Bodotriidae is the second most diverse after the family Nannastacidae,<br />

reaching its highest diversity in the South West Pacific. The family was divided into two<br />

subfamilies, Bodotriinae and Vaunthompsoniinae by Hall (1944) according to the number of<br />

exopods on the pereopods. A third subfamily, Mancocumatinae, was established by Watling<br />

(1977) on the same bases. The family now contains around 360 species in 33 genera. Jones<br />

(1976) listed 11 species in 5 genera from depths down to 200m; the present key extends the<br />

depth range to the lower slopes down to 500m and contains 13 species in 7 genera. The two<br />

additional species are Bathycuma brevirostre (Norman, 1879) recorded from the middle and<br />

lower slopes in the AFEN survey but at 205m west of Ireland by Calman (1905) and<br />

Cyclaspis longicaudata Sars, 1865 recorded from the shelf to lower slopes on both sides of<br />

the Atlantic.<br />

Subfamily: Vaunthompsoniinae<br />

Cumopsis goodsiri (v. Beneden, 1861)<br />

Cumopsis longipes (Dohrn, 1869)<br />

Cumopsis fagei Bacescu, 1965<br />

Vaunthompsonia cristata Bate, 1858<br />

Bathycuma brevirostre (Norman, 1879)<br />

Subfamily: Bodotriinae<br />

Bodotria arenosa Goodsir, 1843<br />

Bodotria pulchella (Sars, 1878)<br />

Bodotria scorpioides (Montagu, 1804)<br />

Bodotria armoricana Le Loeuff & Intes, 1977*<br />

Eocuma dollfusi Calman, 1907<br />

Iphinoe tenella Sars, 1878<br />

Iphinoe trispinosa (Goodsir, 1943)<br />

Iphinoe serrata Norman, 1967<br />

Cyclaspis longicaudata Sars, 1865<br />

* Species not in the key but to be considered.<br />

35


Key to the British Species of Bodotriidae<br />

1- Well developed or rudimentary exopods on pereopods 1-3 or 1-4 in the male and 1-3 in the<br />

female Vaunthompsoniinae 2<br />

Well developed exopods on pereopod 1 only, in both sexes<br />

Bodotriinae 6<br />

2-Telsonic somite not produced posteriorly, exopods on pereopods 2 and 3 rudimentary<br />

3<br />

Telsonic somite produced posteriorly between the uropods, exopods on pereopods 2 and 3<br />

well developed 5<br />

3- Carapace with two pairs of lateral ridges Cumopsis goodsiri<br />

Carapace without lateral ridges. 4<br />

4- Distal article of uropodal endopod with one thin apical spine and several lateral spines;<br />

first article of uropodal exopod shorter than the second Cumopsis longipes<br />

Distal article of uropodal endopod with only one stout spine at the end; first article of<br />

uropodal exopod longer than the second Cumopsis fagei<br />

5- Maxilliped 3 with distal end of basis little or not at all produced and with ischium much<br />

wider than long. Pseudorostral lobes not reaching beyond ocular lobe, eyes present<br />

Vaunthompsonia cristata<br />

Maxilliped 3 with distal end of basis strongly produced and with ischium as long as wide.<br />

Pseudorostral lobes short but reaching beyond ocular lobe, eyes absent<br />

Bathycuma brevirostre<br />

V. cristata<br />

B. brevirostre<br />

36


6- Pereon with 5 free somites, pereonite 1 visible from above<br />

(Iphinoe) 7<br />

Pereon with at most 4 free somites, pereonite 1 hidden from above<br />

9<br />

7- Pleonite 6 with 2 or 4 terminal setae. Adult males with a sternal process on pereonite 2<br />

8<br />

Pleonite 6 with 6 terminal setae. Adult males without sternal process<br />

Iphinoe tenella<br />

8- Carapace with 2-6 teeth dorsally in the female, unarmed in the male. Two terminal setae<br />

on pleonite 6. Antenna 1 with a single aesthetasc<br />

Iphinoe trispinosa<br />

Carapace with 8-20 teeth dorsally in either sex. Four terminal setae on pleonite 6. Antenna 1<br />

with two aesthetascs Iphinoe serrata<br />

9- Pereopod 2 with 7 articles, uropodal endopod uniarticulate with the terminal spine fused to<br />

the article Cyclaspis longicaudata<br />

37


Pereopod 2 with 6 articles (basis and ischium fused), uropodal endopod with 1 or 2 articles<br />

and the terminal spine fused or not fused to the article 10<br />

10- Carapace without lateral horns. Peduncle of the uropods much longer than the rami,<br />

uropodal endopod with 1 or 2 articles and the terminal spine not fused to the article<br />

(Bodotria) 11<br />

Carapace with lateral horns. Peduncle of uropods much shorter than the rami, uropodal<br />

endopod uniarticulate with the terminal spine fused to the article<br />

Eocuma dollfusi<br />

11- Carapace with two carinae on either side Bodotria pulchella<br />

♀<br />

Carapace with a single carina on either side 12<br />

12- Uropodal endopod with two articles Bodotria scorpioides<br />

Uropodal endopod with one article Bodotria arenosa<br />

Bodotria scorpioides Bodotria arenosa<br />

♂<br />

38


Family NANNASTAClDAE<br />

Diagnosis (emended from Jones, 1963 and Watling and McCann, 1997): No articulated<br />

telson. No pleopods. Exopods in male usually present on maxilliped 3 and pereopods 1-4,<br />

rarely on pereopods 1-2 or 1-3; exopods in female usually present on maxilliped 3 and<br />

pereopods 1-2, occasionally absent from maxilliped 3 while present on pereopods1and 2,<br />

rarely absent from all appendages. Mandible normal in shape or with the base widened and<br />

the molar process styliform. Uropod endopod uniarticulate.<br />

Remarks: The family Nannastacidae contains around 352 species in 19 genera and is well<br />

presented in the deep North Atlantic. Jones (1984) described one new genus and 41 new<br />

species and listed overall 99 species from depths exceeding 200m throughout the Atlantic.<br />

Jones (1976) listed 8 species in 3 genera from depths down to 200m. The present key extends<br />

the depth range to the lower slopes down to 500m and contains 17 species in 5 genera.<br />

Procampylaspis armata Bonnier, 1896<br />

Procampylaspis macronyx Hansen, 1920*<br />

Procampylaspis bituberculata Hansen, 1920*<br />

Campylaspis alba Hansen, 1920<br />

Campylaspis affinis Sars, 1870<br />

Campylaspis costata (Sars, 1865)<br />

Campylaspis glabra Sars, 1878<br />

Campylaspis globosa Hansen, 1920<br />

Campylaspis horrida Sars, 1870<br />

Campylaspis intermedia Hansen, 1920<br />

Campylaspis legendrei Fage, 1951<br />

Campylaspis macrophthalma G O Sars, 1879*<br />

Campylaspis rostrata Calman, 1905<br />

Campylaspis rubicunda (Liljeborg, 1855)<br />

Campylaspis sulcata G O Sars, 1870<br />

Campylaspis verrucosa G O Sars, 1865<br />

Campylaspis undata Sars, 1864<br />

Nannastacus brevicaudatus Calman, 1905<br />

Nannastacus longirostris G O Sars, 1879*<br />

Nannastacus unguiculatus (Bate, 1859)<br />

Cumella (Cumella) pygmaea G.O. Sars, 1865<br />

Cumella decipiens Jones, 1984*<br />

Cumella tarda Hansen, 1920*<br />

Cumella sp.aff polita? Jones, 1984*<br />

Styloptocuma gracillimum (Calman, 1905)<br />

Styloptocuma angustata Jones, 1984*<br />

* Species not in the key but to be considered.<br />

39


Key to the British species of Nannastacidae<br />

1- Molar process of mandible styliform, pointed. Anterolateral angle of carapace absent or<br />

only slightly prominent 2<br />

Molar process of mandibles thick and truncate. Carapace of female with well developed<br />

anterolateral angle. 15<br />

2- Maxilla 2 with one lobe; maxilliped 2 with the dactyl shaped like a rake; pereopod 1 with<br />

ischium long; eyelobe extending almost to the end of the pseudorostrum<br />

Procampylaspis armata+<br />

Maxilla 2 rudimentary, without lobes; maxilliped 2 dactyl normal; pereopod 1 with ischium<br />

not specially long (Campylaspis) 3<br />

3- Carapace smooth without tubercles, spines, ridges or lateral grooves 4<br />

Carapace with tubercles, spines or ridges or at least a shallow lateral groove<br />

6<br />

4- Eyelobe small and rudimentary Campylaspis alba<br />

Eyelobe of normal size 5<br />

40


5- Dactyl of pereopod 2 longer than the carpus and propodus together<br />

Campylaspis rubicunda<br />

Dactyl of pereopod 2 not longer than carpus and propodus together<br />

Campylaspis glabra<br />

C. rubicunda C. glabra<br />

6- Carapace with lateral furrow or ridges, but without tubercles, spines or other large<br />

prominences 7<br />

Carapace may or may not have lateral furrow, but with at least a pair of tubercles or many<br />

spines or large prominences, which may or may not form lateral ridges<br />

10<br />

7- Two oblique ridges arising anteriorly and extending for greater part of the length of the<br />

carapace on either side 8<br />

Three or more oblique ridges extending similarly on the carapace, posterior ridge branched<br />

Campylaspis costata<br />

8- Dactyl of pereopod 2 shorter than the carpus Campylaspis undata<br />

Dactyl of pereopod 2 longer than the carpus 9<br />

C. undata<br />

C. sulcata<br />

C. legendrei<br />

9- Dactyl of pereopod 2 with terminal seta. Carapace with longitudinal depressions each<br />

divided into two unequal parts Campylaspis legendrei<br />

Dactyl of pereopod 2 with no terminal seta. Carapace with depressions undivided<br />

Campylaspis sulcata<br />

41


10- Carapace with a few low dorsal protubercles<br />

Campylaspis affinis<br />

Carapace with many conspicuous tubercles 11<br />

11-Carapace with tuberculate ridges or with some large tubercles situated in distinct rows<br />

along the sides 12<br />

Carapace without tuberculate ridges, but low tubercles may be situated along the sides in<br />

rows 14<br />

12- Dactyl of pereopod 2 much longer than the carpus and propodus together. Pseudorostrum<br />

relatively long and prominent. Carapace with few large tubercles<br />

Campylaspis rostrata<br />

Dactyl of pereopod 2 at most as long as the carpus and propodus together. Pseudorostrum<br />

relatively short and not prominent. Carapace with many large tubercles<br />

13<br />

13- Side of carapace with three lateral ridges, the two uppermost bearing large rounded<br />

tubercles Campylaspis intermedia<br />

Side of carapace with two lateral ridges, formed of conical tubercles<br />

Campylaspis horrida<br />

14- Merus of maxilliped 3 as long as carpus and propodus together, tubercles on the carapace<br />

low and rounded, no depressed area on the side of the carapace<br />

Campylaspis verrucosa<br />

42


Merus of maxilliped 3 much shorter than carpus and propodus together, tubercles on the<br />

carapace low and rounded, depressed area on the side of the carpus present<br />

Campylaspis globosa<br />

15- Two ocular groups more or less separated (Nannastacus) 16<br />

A single median ocular group 17<br />

16- Uropods about two-thirds as long as the last two pleonites together<br />

Nannastacus brevicaudatus<br />

Uropods longer than the last two pleonites together<br />

Nannastacus unguiculatus<br />

17- Eye lobe with lenses, narrow, short, not reaching end of pseudorostral lobes<br />

Cumella pygmaea#<br />

Eye lobe without lenses, narrow, generally elongate, reaching to end of pseudorostral lobes or<br />

beyond Styloptocuma gracillimum*<br />

43


Other species not in the key but to be considered:<br />

+Procampylaspis macronyx Hansen, 1920<br />

+Procampylaspis bituberculata Hansen, 1920<br />

#Cumella decipiens Jones, 1984<br />

#Cumella tarda Hansen, 1920<br />

#Cumella sp.aff polita? Jones, 1984<br />

* Styloptocuma angustata Jones, 1984<br />

44


Family LEUCONIDAE<br />

Diagnosis (from Jones, 1976): No free telson. Pleopods without an external process on the<br />

inner ramus, 2 pairs, rarely 1 or 0, in male. In the male, exopods present on the third<br />

maxillipeds and on the first four pairs of pereopods, rarely the first two pairs; in the female<br />

exopods present on the first three pairs, rarely the first two pairs. The mandibles are<br />

broadened at the base. The endopod of the uropod with two articles or rarely uniarticulate.<br />

All the pereon somites are visible from above.<br />

Remarks: The Leuconidae is one of the earliest-recognised cumacean families, established by<br />

Sars (1878). Leuconidae is a very diverse family, containing 121 species in 16 genera. Jones<br />

(1976) listed 4 species in 3 genera from depth down to 200m. The present key extends the<br />

depth range to the lower slopes down to 500m with the addition of only one extra species.<br />

The additional species is Leucon (Crymoleucon) noerrevangi Watling and Gerken (1999),<br />

which was recorded from the west of Shetland at 434m deep. The family has a deep-sea<br />

affinity worldwide. The genus Eudorella needs revision.<br />

.<br />

Leucon (Crymoleucon) noerrevangi Watling and Gerken (1999)<br />

Leucon (Leucon) nasica (Krøyer, 1841)<br />

Leucon acutirostris G O Sars, 1864*<br />

Eudorellopsis deformis (Krøyer, 1846)<br />

Eudorella emarginata (Krøyer, 1846)<br />

Eudorella truncatula (Bate, 1856)<br />

Eudorella species A (Bate, 1856)*<br />

* Species not in the key but to be considered.<br />

45


Key to the British Species of Leuconidae<br />

1- Efferent orifice anterior or anterodorsal 2<br />

Efferent orifice distinctly dorsal, pseudodorsal lobes directed dorsally<br />

3<br />

2- Antenna 1 accessory flagellum extending at least to midlength of main flagellum first<br />

article; frontal lobe with lateral group of spines; pseudorostral lobe with group of spines<br />

Leucon (Crymoleucon) noerrevangi<br />

Antenna 1 accessory flagellum clearly less than half the length of main flagellum first article;<br />

frontal lobe with one spine at most; pseudorostral lobe without group of spines<br />

3- Antenna 1 geniculate between articles 1 and 2. Uropod endopod shorter than exopod<br />

Eudorellopsis deformis<br />

Antenna 1 geniculate between articles 2 and 3. Uropod endopod longer than exopod<br />

4<br />

4- Tooth below sinus at front of carapace small, not projecting beyond upper teeth (applies<br />

best to ♀♀)<br />

Eudorella truncatula<br />

♀<br />

From Watling and Gerken (1999)<br />

Tooth below sinus large, projecting beyond the upper teeth (applies best to ♀♀)<br />

Eudorella emarginaia<br />

♀<br />

♂<br />

46

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