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Recent Brachiopoda from the océanographie

expédition SEAMOUNT 2 to the north-eastern

Atlantic in 1993

Alan LOGAN

Centre for Coastal Studies and Aquaculture, University of New Brunswick,

P.O. Box 5050, Saint John, N.B., E2L 4L5 (Canada)

logan@unbsj.ca

Logan A. 1998. — Recent Brachiopoda from the océanographie expédition SEAMOUNT 2

to the north-eastern Atlantic in 1993. Zoosystema 20 (4) : 549-562.

KEYWORDS

Brachiopods,

Recent,

SEAMOUNT 2,

north-eastern Adantic.

ABSTRACT

Eight species of recent brachiopods belonging to the généra Neocrania,

Dyscolia, Abyssothyris , Stenosarina, Eucalathis , Platidia, Pbaneropora and

Dallina hâve been identified from collections from the 1993 SEAMOUNT 2

expédition to Meteor, Hyères, Irving-Cruiser, Plato, Atlands, Tyro and

Antialtair seamounts in the north-eastern Atlantic. The species misidentified

by Jeffreys (1878) as Terebratula vitre a var. sphenoidea [non Philippi, 1844] is

described as Stenosarina davidsoni n.sp. The affmities of the SEAMOUNT 2

brachiopods are with the Mauritanian biogeographic province. Diversity and

number of stations yielding brachiopods increase from south to north in the

cluster of six seamounts (Meteor-Tyro) south of the Azores. Brachiopod

diversity for the seven seamounts as a whole is less than for the Canary

Islands to the east. There is an as yet unexplained absence from the sea¬

mounts of deeper water species belonging to such généra as Pelagodiscus,

Hispanirhynchia, Terebratulina, Gryphus, Megerlia and Macandrevia , which

commonly occur around island archipelagos such as Madeira, the Canaries and

the Cape Verdes, as well as off the Iberian coast and the African mainland.

ZOOSYSTEMA • 1998 • 20(4)

549

Logan A.

MOTS CLÉS

Brachiopoda,

actuel,

SEAMOUNT 2,

Atlantique Nord-Est.

RÉSUMÉ

Brachiopodes actuels récoltés lors de l'expédition océanographique SEA¬

MOUNT 2 dans l’océan Atlantique Nord-Est en 1993. Huit espèces de bra¬

chiopodes actuels appartenant aux genres Neocritnia, Dyscoha, Abyssothyris,

Stenosarina, Eucalathis, P hindi a, Phaneropora et Dallina ont été identifiées à

partir des collections réalisées lors de l’expédition SEAMOUNT 2 de 1993

aux pitons sous-marins Mcteor, Hyèrcs. Irving-Cruiser, Plato, Adantis, Tyro

et Antialtair dans l’océan Atlantique Nord-Est. L espèce incorrectement

identifiée par J effreys (1878) comme Terebratula vitrea var. sphenoidea [non

Philippi, 1844] est décrite comme Stenosarina datndsoni n.sp. Les brachio¬

podes de SEAMOUNT 2 montrent une affinité avec la faune de la province

biogéographique mauritanienne. La diversité et le nombre de stations où ces

brachiopodes ont été observés augmentent du sud au nord dans le groupe des

six pitons (Meteor-Tyrü) au sud des Açores. La diversité en brachiopodes

pour la totalité des sept pitons est inférieure à celle des îles Canaries à Pest.

On observe sut ces pitons, sans pouvoir f expliquer actuellement, une absence

des espèces plus profondes appartenant aux genres Pelagodiscus, Hispani-

rhynchia , Terebratulina, Gryphus, Megerlia et Macandrevia qui se rencontrent

normalement dans les archipels tels que Madère, les Canaries et les îles du

Cap-Vert, ainsi que le long des côtes ibérique et africaine.

INTRODUCTION

The SEAMOUNT 2 océanographie expédition

in 1993 to Meteor, Hyères, Irving-Cruiser, Plato,

Atlantis, Tyro and Antialtair seamounts in the

north-eastern Atlantic, west of Madeira and the

Canary lslands and notth and south of the

Azores, sampled 165 stations (Gofas 1993). Of

these, forty-eight stations (29%) yiclded hrachio-

pods. This proportion is comparable to chose for

brachiopods described front other Atlantic and

Caribbean collections (Cooper 1977; Logan

1983, 1988) and confions once again thac they

are not a comrnon constituent of bottom

samples except in high latitudes (Foster 1974).

The location of ail sampling stations is shown in

Figure 1.

At the time of the study, the fifieen stations sam-

pled off the western side of Grand Canary Island

had not been sorted for brachiopods. Two out of

forty-six stations (4%) on Meteor seamount yiel-

ded a cocal of three species, five out of twenty-

four stations (21%) on Hycres seamount yielded

a total of foui' species, thirteen out of rhirty-five

stations (37%) on Irving-Cruiser seamounts yiel¬

ded four species, eighc out of fourteen stations

(57%) on Plato seamount yielded six species,

fourteen out of twency-one stations (67%) on

Atlantis seamount yielded eighc species, four out

of six stations (67%) on Tyro seamount yielded

five species and two oui of four stations (50%)

on Antialtair seamount yielded five species of bra¬

chiopods.

The following brachiopods hâve been recorded

from the SEAMOUNT 2 Expédition and appear

in the systematic section below:

Neocrania anomala (Muller, 1776);

Dyscolia wyviUei (Davidson, 1878);

Abyssothyris atlantica Cooper, 1977;

Stenosarina davidsoni n.sp.;

Eucalathis ergmtica Fischer et Oehlert, 1890;

Plaudui anmnioides (Scacchi et Philippi, 1844);

Phaneropora incerta (Davidson, 1880);

Dallina septigera (Loven, 1846).

550

ZOOSYSTEMA • 1998 • 20(4)

SEAMOUNT 2 brachiopods from the N.E. Atlantic

SYSTEMATICS

The supra-ordinal classification of Williams et al.

(1996) has been used in this report, and, in

accordance with usage in the forthcoming revised

treatise on brachiopods, the terms "dorsal” and

“ventral” are used tor “brachial” and "pedicle”

when describing the two valves. Since most spe-

cies hâve been fully described elsewhere, only

brief notes, illustrations of salient characteristics

and a citation to a recent reliable description

hâve been given here. Synonymies are therefore

partial except for the newly-named species

Stenosarina davidsoni Logan, which has been

fully redescribed and illustrated. Ail types and

figured specimens are deposited in the collections

ot the Muséum national d’Histoire naturelle

(MNHN) in Paris, France. The exact location,

depth and species identified at each station are

shown in the Appendix.

Fig. 1. — Map of the south-east North Atlantic to show the location of the seven seamounts from which brachiopods were obtained

by the SEAMOUNT 2 expédition.

ZOOSYSTEMA • 1998 • 20 (4)

551

Logan A.

Sub-phylum CRANIIFORMEA

Popov, Bassett, Holmer et Latine, 1993

Class CRANLATA Williams, Carlson,

Brun ton, Holmer et Popov, 1996

Ordet CRAN1 IDA Waagen, 1885

Suborder CRANJ1D1NA Waagen, 1885

Superfamily CranIAŒA Menke, 1828

Family CRANIIDAE Menke, 1828

Genus Neocrania Lee et Brun ton, 1986

Neocrania anomala (Millier, 1776)

(Fig. 2A-C)

Crania anomala - Logan 1979: 27, figs 4, 5; pi. 1,

figs 1-10.

Stations. — Irving-Cruiser: DW 215-216,

DW 237; Plato: DW 240, DW 248; Atlantis:

DW 254-257, DW 265, TS 270, DW 274; Tyro:

275, DW 277; Anrialtair: DW 281, DW 283.

Depth range. — 270-1665 m.

Remarks

This species, one of the commonest brachiopods

in the SEAMOUNT 2 collections, is a typical

member of the brachiopod fauna of the eastern

Atlantic (Fischer & Oehlert 1891; Brunton and

Curry 1979; Anadon 1994), being found as far

south as the Cape Verdc Tslands (Logan 1988)

and ranging grcatly in depth (Cooper 1981; this

report), lt is also common throughout the wes¬

tern Mcditerranean, where it again ranges consi-

derably in depth, although occurring most

commonly in cryptic habitats in shallow water

(Logan 1979; Logan & Noble 1983; Templado

& Luque 1986) h is easily recognizable by its

cone-shaped upper (dorsal) valve, subcircular

outline, concentric growth Iinés and endopunc-

tatc shell, while internally the adductor muscle

scars of both valves arc arranged in pairs, poste-

riorly and anteriorly, on white callus pads, with a

centrally-placed brachial protractor scar in the

upper valve and a prominent médian muscle scar

in the very thin areached (ventral) valve. The

shape of the shell, and particuiarly the attached

valve, is variable, since it is strongly influenced

by the irregularities of the substrate to which it is

cemented (Fig. 2C). This substrate conststs

mainly of coarse sedimentary, voleame or bioclas-

tic material in the SEAMOLJNT 2 collections.

Ail specimens examined showed the typical dor¬

sal valve internai characteristies of N. anomala

rarher rhan N. turbinata (Poli, 1795) which

Brunton (1988) rcdcscribed Iront the eastern

Mediter-ranean.

Order TEREBRATUL1DA Waagen, 1883

Suborder TEREBRATULIDINA Waagen, 1883

Superfamily TEREBRATULACEA Waagen, 1883

Family DyscOLUDAE Fischer et Oehlert, 1891

Genus Dyscolia Fischer et Oehlert, 1890

Dyscolia wyvillei (Davidson, 1878)

(Fig. 2D-F)

Dyscolia wyvillei - Cooper 1977: 56, pl. 15, figs 1-10.

Stations. - - Meteor: DW 173; Hvères: DW 186,

DW 203; Atlantis: DW 264.

Depth range. — 795-1520 m.

Remarks

This very large species occurs in the eastern

Atlantic (Fischer &t Oehlert 1891; Logan 1983;

Saiz Salinas 1989) and the Caribbean (Cooper

1977) where the shells are usuaJly collected in a

very worn State. The loop of the brachial skele-

ton is rarely preserved in Recent specimens and

the soft parts almost never, so the nature of the

lophophore is uncertain, although one of

Fischer & Oehlert’s figures (1891, pl. 6, fig. 3i)

suggesrs rhat it is trocholophous (see Cooper

1983 for further discussion). The specimens

from SEAMOUNT 2 arc ail badly worn and

show no brachial loops but the size and thickness

of the shell, the large foramen and the distinctive

fine radial ornamentation allow a positive identi¬

fication to be made.

Family TerebratULIDÆ Gray, 1840

Genus Abyssothyris Thomson, 1927

Abyssothyris atlantica Cooper, 1977

(Fig. 2G-M)

Abyssothyris atlantica Cooper, 1977: 58, pl. 20,

figs 1-10.

552

ZOOSYSTEMA • 1998 • 20(4)

SEAMOUNT 2 brachiopods from the N.E. Atlantic

Fig. 2. — A-C, Neocrania anomala (Müller); D-F, Dyscolia wyvillei (Davidson); G-M, Abyssothyris atlantica Cooper. A, B, SEMs of

exterior and interior of upper (dorsal) valve, the lattor showing details of the muscle Beats DW 274, 280 m, Atlantis seamount,

MNHN LBIMM-BBA-2781, 2782; C, trngmont ot volconlc rock wlth se verni attacher) lower (ventral) valves. DW 256. 340-345 m,

Atlantis seamount. MNHN L6IMM-0nA-?783. D-F, ventral valve exleiior showlng fine radial ornamentation, ventral volve Interior of

same specimen snowmg worn hinge teeth and dorsal valve interior showing sockels and adductor sons DW 173, Meteor ssnrnount,

920 m, MNHN LBIMM-BRA-2784, 2785; S J, SEM ventral, dorsal, sido and antorior views to show external ornement and slightly

sulcate antenor commissure, DW 279. Tyro 98$mr>unt, 780-805 m, MNHN LBIMM-BRA-2786, 2787. 2788, 2789; K, SEM ot dorsal

valve Interior to show loop of brachial sketeton, same locality and depth as préviens Specimen, L, enlergemanl ot previ0(rs spedi-

men. MNHf'l I RtMM-pRA-2790. M SEM ot dorsal valve rptorlor wlth plectolophous lophophore covermg loop of brachial skeleton,

same locality and depth as previous specimen, MNHN LBIMM-BRA-2791. Scale bars: A, B, C, 1 mm; D-F, 1 cm; G-J, K, L, M, 1 mm.

ZOOSYSTEMA • 1998 • 20(4)

553

Logan A.

Stations. — Meteor: DW 173; Plato: DW 241;

Atlantis: DW 258; Tyro: DW 277-279.

Depth range. — 420-1070 m.

Remarks

This species was lirsr described from over

2500 iti dcptli. off Cape Fear, South CaroJina, by

Cooper (1977) who narned it fot very small

elongate Abyssothyns wich a broadly sulcate antc-

rior margin. Specimens from SEAMOUNT 2

are less sulcate than the types but otherwise simi-

lar enough to the western Atlantic examples to

be identifted with them. The loop of Abyssothyns

is rounded anteriorly with a slight médian angu¬

lation. The plectolophous lophophore is small

and tighdy coiled (Fig. 2M) but laclcs the médian

connecting band ilJustrated by Cooper (1983)

for A. elongata. and said to be typical of the

genus. However, Coopers illustratcd specimen

(op. rit., pl. 16, ftg. 19) came from depths of ovcr

3600 m, whereas A. atlantica Iront SEA¬

MOUNT 2 occurs at much shallower depths.

Members of the genus typically occur in deep

water and the development of the lophophore

may be related to the effects of great depths, such

as hydrostatic pressure and oxygen content

(Zezina 1975). This is the First record of the

genus from the eastern Atlantic.

Genus Stenosarina Cooper, 1977

Stenosarina davidsoni n.sp.

(Fig. 3A-J)

Terebratula vitrea var. sphenoidea [not Philippi,

1844] — Jeffreys 1878: 404, pl. 22, fig. 6?

Liothyris sphenoidea \sensu Jeffreys, 1878] - Davidson

1886: 12, pl. 2, figs 17, 18 only.

Terebratula (Liothyrina) sphenoidea [ sensu Jeffreys,

1878] - Fischer & Ochlert 1891, p. 58, pl. 3,

fig. 8a-m.

Liothyrina sphenoidea | sensu Jetfreys, 1878] —

Blochmann 1908: 619, pl. 37, fig. 11; pl. 38, fig. 19a-

c; pl. 39, fig. 23a-b.

Dallithyris ? aff. D .? sphenoidea [sensu leffreys,

1878]- Cooper 1981: 1 5, pl. 2, figs 36-38.

Stenosarina sphenoidea [sensu Jeffreys, 1878] - Cooper

1983:272, pl. 14, figs 15-19.

Stenosarina sphenoidea [sensu Jeffreys, 1878] — Logan

1988:64.

Mateiuaj EXAMINED. — Atlantis seamount,

DW 258, holotype MNHN LBIMM-BRA-2778,

paratypes MNHN I.BIMM-BRA-2779, 2780.

Stations. — Hyéres: DW 184, DW 186, DW 197,

DW 203; Irving-Cruiser: DW 218-219, DW 225-

227, DW 229. DW 231, DW 237-238; Plato:

DW 241-242, DW 246-248, DW 251; Atlantis;

DW 254-256, DW 258-259, DW 263. DW 265,

DW 271 ; Tyro; DW 277-278; Antialtair: DW 283.

Depth range. — 275-1715 m.

Description

Exterior

Shell white, thin and translucent, rarely excee-

ding 20 mm in length and 15 rnm in width;

elongate ovaJ in outline, widest point just ante¬

rior to mid-valve, with anterior margin gently

rounded. Both valves more or less evenly convex,

with latéral commissure weakly concave towards

ventral side near mid-valve; anterior margin recti-

marginate. Beak suberect, labiate; foramen

moderate in size, permesorhyridid.

Interior

Ventral valve interior with small, narrowly elon¬

gate teeth; short pedicle collar. Dorsal valve with

narrow, parailel-sided loop which occupies about

one third the length of the valve. Cardinal pro-

cess small, Socket ridges curved, flanking a Socket

roofed at its proximal end. Outer hinge plates

triangular and concave, extending anteriorly to

join crural bases. Crura typical for the genus, as

described and illustrated in detail bv Cooper

(1983:271,272).

Remarks

This species, the commonest brachiopod in the

SEAMOUNT 2 collections, has in the past been

assigned to the généra Terebratula, Liothyris,

Liothyrina, Dallithyris? and Gryphus by earlier

aurhors, prior to Stenosarina by Cooper in 1983,

under the spécifie epithet sphenoidea. Davidson

(1886), Fischer & Üehlert (1891), Blochmann

(1908), Dali (1920), Muir-Wood (1959),

Cooper (1981, 1983) and Logan (1988) bave ail

discussed the problems that hâve arisen from

554

ZOOSYSTEMA • 1998 • 20(4)

SEAMOUNT 2 brachiopods from the N.E. Atlantic

Fig. 3. — A-J, Stenosarina davidsoni n.sp.; K-T, Eucalalhis ergastica (Fischer et Oehlert). A-D, ventral, dorsal, side and anterior

views of holotype to show external arnament and rectimarginato anterior commissure, DW 258, Atlantis seamount. -120-460 m,

MNHN LBIMM-BRA-2778, H, F, ventral and dorsal views ot paratypo t, MNHN LBIMM-BBA 2779; G, H, ventral and dorsal views of

paratype 2, -;ame localily and deplh as previous specimen. MNHN LBIMM-BRA-2780; I, J. 9EM émargements ol loop of Prachlal

skeleton of dorsal valve, straight and fllled, to show V-shaped rtolch m transverse band. samo locality and deplh as previous spéci¬

men. MNHN L8IMM-BRA-2792. K. L SEMs of exlerini ol ventral and dorsal valves to show ornsmont. DW 264, Atlantis seamount,

795-830 m, MNI IN LBIMM URA-2793, 2794. M. N. SEMs ol inlerior of ventral and dorsal valve* to Htiow denlilion end brachial ske-

leton, DW 283, Anlialtalf seamount, 1175-1210 m, MNHN LBIMM-BRA-2795. 2796: o r, SEMs of extorior and inlerior of ventral and

dorsal valves ol fuvemle speci-nens to show subiioed omtjTrtenl end mdimentary hrachinl skelelori, DW 274 Atlantis seamount,

280 m, MNHN LBIMM-BRA-2797 to 2802. Scale bars: A-D, E, F, G, H, 1 cm; I, J, K, L, M, N, O-T, 1 mm.

ZOOSYSTEMA • 1998 • 20(4)

555

Logan A.

Jeffreys’nrisidentification of the eastern Atlantic

Recent form wirh Pbilippis (1844) Pliocène spe-

cies Terebratula sphenoidea. Dali (1920), Muir-

Wood (1959) and Cooper (1981, 1983) hâve

pointed out the main différences, which need

not be repeared here, between the Pliocène and

modem form. The above synonymy contains

only chose references wherc good descriptions

and/or illustrations ensure identification with the

modem form.

Several aurhors (Muir-Wood 1959; Cooper

1981) hâve called for the establishment ot a new

species tiame to avoid furcher confusion of

modem and Pliocène forms. Jeffreys (1878) reco-

gnized two vatieties of Terebratula vitrea , for

which he used two names cstablished by Philippi

(1844): min or and sphenoidea. Neirher is an

appropriate désignation for the modem eastern

Atlantic form. A further complication is thar

Jeffreys’figure is fainrly costellate, a condition

that I hâve not seen in typical eastern Atlantic

Stenosarina and which Muir-Wood (1959) feels

is represented erroneously. Enough material has

been found in the présent srudy to warrant a

redescriprion of this species as a new taxon,

which I hetc propose to call Stenosarina davidso-

ni in honour oi the eminent brachiopod

palaeontologist Thomas Davidson.

The species occurs in rhe eastern Atlantic as far

south as the Canary and Cape Verde Islands

(Logan 1988). Cooper (1981) is in error in sta-

ting that it occurs in the Mediterranean.

Family CANCELLOTHYRIDIDAE Thomson, 1926

Subfamily EUCALATHINAE Muir-Wood, 1965

Genus Eucalathis Fischer et Oehlert, 1890

Eucalathis ergastica Fischer et Oehlert, 1890

(Fig. 3K-T)

Eucalathis ergastica - Cooper 1981: 12, pl. 1, figs 1-8.

-Logan 1983: 173.

Stations. — Me tco r; DW 136; Hyères: DW 203;

Plato: DW 249; Atlantis: DW 255, DW 258,

DW 261, DW 264, TS 270, DW 274; Tyro:

DW 278; Antialtair: DW 283.

Depth RANCE. — 280-1800 m.

Remarks

Eucalathis ergastica and its dosely related conge-

ner E. tuheraia (Jeffreys), originally described

front the south-east North Atlantic by Fischer &

Oehlert (1891), are common brachiopods in the

SEAMOUNT 2 and CANCAP collections

(Logan 1983, 1988, this report). Cooper (1981)

and Saiz Salinas (1989) hâve also recorded this

species front the Bay of Biscay and Anadon

(1994) from the coast of Asrurias, norrhern

Spain. The différences between the two species

are ntainly in their size, shape and ornament,

E. tuherata being smaller and more rounded,

with more nunterous beaded radial costella.

Juvéniles ot E. ergastica (Fig. 4E-J) hâve a more

subdued omantent of less elosely-spaeed costellae

than rhe adult forms. The species occurs olf the

Azores and the Canary Islands, the north coast of

Spain and rite Moroccan coast (Cooper 1981).

Suborder TEREBRATELLID1NA

Muir-Wood, 1955

Superfamily Terebratellacea King, 1850

Family PLATIDIIDAE Thomson, 1927

Genus Platidia Costa, 1852

Platidia anomioides (Scacclii et Philippi, 1844)

(Fig. 4A-D)

Platidia anomioides — Cooper 1977: 122, pl. 20,

figs 11-19; pl. 33, figs 15-17. - Logan 1979: 60,

figs 17, 18; pl. 7, figs 1-11.

Stations. — lrving-Cruiser: DW 209, DW 225;

Plato- DW 242; Atlantis: DW 254-258, DW 262-

265, TS 270. DW 274; Antialtair: DW 281.

Depth range. — 275-1160 m.

Remarks

This heavily-spiculate species is relatively com¬

mon throughout the North Atlantic, particularly

on its eastern side (Bruntoti &: Curry 1979;

Cooper 1981; Logan 1983; Saiz Salinas 1989;

Anadon 1994). lt also occurs in the Caribbean

région (Cooper 1977) but is commonest in the

Mediterranean (Logan 1979) where, as in the

556

ZOOSYSTEMA • 1998 • 20(4)

SEAMOUNT 2 brachiopods from the N.E. Adantic

Fig. 4. — A-D, Piatidia anomioides (Scacchi et Philippi); E-G, Phaneropora incerta (Davidson); H-N, Daltina septigera (Loven).

A, B, SEMs of exterior of ventral and dorsal valves, to show convex ventral valve, fiat dorsal valve, and amphithyridid pedicle fora¬

men shared t>y both valves. DW 258 and 256, respeclively, Atlanlis seamount, 420-480 m and 340-345 m, MNHN LBIMM-BRA-

2803, 2804, C, D, SEMs showtng tilted dorsal and side vievvs of mterior of dorsal valve to show spiculés and brachial support for

lophophore, DW 264, Atlaotis seamount, 795-830 m, MNHN LBIMM-BRA-2805, E, SEM ol fnterior ot ventral valve to show hrnge

teeth, DW 261, Atlanlis seamount, 1190-1340 m. MNHN LBIMM-BRA-2806; F, G, SEMs al dorsal and side viows of Interior ol dorsal

valve to show ïockets and btachial support for lophophore. same locality and depth as previous specimen, MNHN L6IMM-BRA-

2807; H-K, ventral, dorsal, side andanterior views to show axternal ornament and sulcate anterior commissure. DW 241. Plato sea¬

mount. 695 m, MNHN LBIMM-BRA-2808, L-N, SEMs ol ventral, dorsal and side dorsal views ol two intertors to show teeth, sockets

and brachial loop, DW 281, Antiallair seamount, 300 m, MNHN LBIMM-BRA-2009, 2ei0. Scale bars; A, B, C, D, 1 mm; E, F, G,

0.5 mm; H-K, 1 cm; L-N, 1 mm.

ZOOSYSTEMA • 1998 • 20(4)

557

Logan A.

Atlantic, it is usually attachée! to bioclastic

débris. Logan (1979) has shown that P. ano-

mioides can be disnnguishcd most reliably from

its closely-related congener P. davidsoni

(Deslongchamps) by slight différences in the

configuration of the brachial skcleton. Removal

of the lophophore of several SEAMOUNT 2

specimens to reveal this supporring structure has

confirmed that they are indeed P anomioides.

Family PHANEROPORIDAE Zezina, 1981

Genus Phaneropora Zezina, 1981

Phaneropora incerta (Davidson, 1880)

(Fig. 4E-G)

Phaneropora incerta - Logan 1983: 177, pl. 1,

figs 11-17.

Stations. — Atlantis: DW 255, DW 261, TS 270.

Depth range. — 335-1340 m.

Remarks

This species was redescribed and illustrated from

Madeira, the Canary and Cape Verde Islands by

Logan (1983, 1988), who discussed the diffé¬

rences between the Atlantic species and the type

species P. galatheae (Muir-Wood, 1965) from the

Pacific. Davidson’s species was questionably assi-

gned to Platidia Costa by Fischer & Oehlert

(1891) and, in fact, the internai skeletal characte-

ristics oi Phaneropora are not unlike rhuse of

Platidia, but externaJly the two généra are quite

different in shape. Furthermore, the former

genus lacks the large amphithyridid foramen sha-

red by both valves, which results in a large

hemispherical notch m each valve of Platidia.

Family Daixinidae Beecher, 1893

Genus Dallina Beecher, 1893

Dallina septigera (Loven, 1846)

(Fig. 4H-N)

Dallina septigera - Cooper 1981: 23, pl. 2, figs 41-43.

Stations. — Hyères: DW 200, DW 203; Irving-

Cruiser: DW 219; Plato: DW 241-242, DW 248;

Atlantis: DW 258, DW 263; Tyro: DW 277-279;

Antialtair: DW 281, DW 283.

Depth range. — 420-1210 m.

Remarks

Dallina septigera is a commun species throughout

the north-eastern Atlantic and has been well des-

cribed and illustrated by previous authors

(Davidson 1886; Fischer 6t Oehlert 1891;

Arkins 1960; d’Flondt 1976; Brunton & Curry

1977, Cooper 1981; Logan 1983, 1988;

Anadon, 1994). It has a distinctive triangular

outline, is widest anteriorly and has a sulcate

anterior commissure. The brachial skeleton

lorms a long loop with a wide hood-like trans¬

verse ribbon (Fig. 4M, N).

DISCUSSION

The earliest expéditions ro make extensive collec¬

tions of brachiopods in the eastern Atlantic

island archipelagos vvere those of die French

research vessels Travailleur in 1882 and Talisman

in 1883 (Fischer Ht Oehlert 1891). Silice lhen six

Dutch CANCAP expéditions to the same

régions between 1976 and 1982 bave significant-

ly added to our knowledge ot brachiopods from

this région (Logan 1983, 1988, 1993), alrhough

there is siill a dearth of information on hrachio-

pods from the Azores. In terms of biogeographic

affinities, the SEAMOUNT 2 brachiopods are

typical of rhe Mauritanian faunal province. The

affinities of the Mauritanian brachiopods to

those front the Lusttanian and Mediterranean

faunal provinces hâve been discussed by Logan

(1993). No species new to science hâve been des-

cribed in this study, alrhough the genus

Abyssothyris has been recorded for the fitst time

Iront the eastern Atlantic, and the lormer species

Stenosarina sphenoidea (Jeffreys) [not Philippi]

has been renamed Stenosarina davidsoni n.sp.

There is a graduai increase in rhe ttumber of sta¬

tions yieldtng brachiopods, and the number of

species identified, from south to north in the

cluster of six seamounts (Meteor-Tyro) south of

tbe Azores. Nevertheless, diversity for the whole

seamount région is low, only eight species of bra-

558

ZOOSYSTEMA • 1998 • 20 (4)

SEAMOUNT 2 brachiopods from the N.E. Atlantic

chiopods being identified in this study, compa-

red to sixteen species in the Canary-Salvage

Islands area (Logan 1993). This relative paucity

is not believed to be an arrifacc of the sampling

and sorting eftlciency of the collectors; rather it

may be, at least partially, explained by rhe

depths, sampling merhods and borrom types of

the stations sampled. Depths range from about

300 m to over 3000 m, widi most over 500 m,

so that predominantly shallow-water species

belonging to the généra Argyrotheca Dali, 1900;

Megathiris d'Orbigny, 1847, Thecidellina

Thomson, 1915; Pajnudim Logan, 1988, and

Lacazella Munier-Chalmas, 1881, which were

commonly obtained by the earlier CANCAP

expéditions at depths of less than 600 m (Logan

1983, 1988, 1993), were not encountered in

SEAMOUNT 2 samples. More diflicult to

explain, however, is rhe absence from the sea-

mounts of deeper warer species belonging to

généra such as Pelagûdiscus , Hispanirhynchia ,

Terebratulina, Gryphtis, Megerlia and Macan-

drevia, which commonly occur around island

archipelagos such as Madctra, the Canaries and

Cape Vcrde Islands, as well as oit rhe Iberian

coast and rhe African maintand (Fischer &

Oehlert 1891; d’Hondt 1976; Logan 1983,

1988; Saiz Salinas 1989; Anadon 1994). Most

SEAMOUNT 2 botrom samples were obtained

by either dredge or epibenthic sied (Gofas pers.

comm,), vv h créas rhe CANCAP samples were

obtained mainly by Van Vccn grab (Logan

1983). It is possible that some small and/or fragi¬

le brachiopod shells might hâve been lost or

damaged by the sampling methods employed by

SEAMOUNT 2.

Acknowledgements

I am indebted to the SEM unir of the University

of New Brunswick for photography and UNBSJ

Research Tecbnical Ofiftcer Willred Morris for

computcrized improvements to the plates, using

Photoshop Research on Recent brachiopods is

supported by the Natural Sciences and Engineer¬

ing Research Council ot Canada Research Grant

OGP 4331 to the author. This is rcscarch publi¬

cation nutnber 9 ol the Centre tor Coastal

Studies and Aquaculture of the LIniversiry ot

New Brunswick.

REFERENCES

Anadon N. 1994. — Braquiôpodos actuales de la pla-

taforrna y talus continental de la costa de Asturias

(Norte de Espana), Boletin de la Real Sociedad

espahola de historia natural , Madrid, Seccion

Biologia 91 (1-4); 65-77.

Atkins D. I960. — A note on Dallina septigera

(Lovtn) (Brachiopoda, Dallinidae). Journal of the

Marine Biological Association of the U.K. 39:91-99.

Blochmann F. 1908. — Zur Systematik und geoqra-

phischen Verbreitung der Brachiopoden. Zeitschrift

fur wissatscbaftHche Zoologie, Leipzig, 90: 596-644.

Brunton C. H. C. 1988. — Some brachiopods Irom

the eastern Mediterrancan Ses. Israël Journal of

Zoology3 5, 1988-1989. 151-169.

Brunton C. H. C. & Curry G. B. 1979. — Brirish

Brachiopods; 1-64, in Kermack D. M. & Barnes

R. S. K (eds). Synapses of the British Raima (N. S.),

17- Academie Press.

Cooper G. A. 1977. — Brachiopods from the

Caribbean Sea and adjacent waters. Studies in

Tropical Oceanography. Rosenstiel School of Marine

and Aimosphcric Science, University of Miami

Press, 14: 1-211.

— 1981. - Brachiopoda from the Gulf of Gascogne,

France (Recent). Smithsauian Contributions to

Paleobwlogy 44: 1-35.

— 1983. — The Terebratulacea (Brachiopoda),

Triassic to Récent: a sllldy of the brachidia (loops).

Smithsonian Contributions to Paleobiology 50:

1-445.

Dali W. H. 1920. — Annotared list of the Recent

Brachiopoda m the collection of the United States

National Muséum, with descriptions of thirty-ihree

new forms. ProceedingS of the United States National

Muséum 57 (2314): 261-377.

Davidson T. 1886. — A monograpli of Recent

Brachiopoda. Transactions of the Ltnnatitn Society of

London (2) 4, Zoology. Pt 1: 1-74.

Fischer P. & Oehlert D-P. 1891. — Brachiopodes.

Expéditions scientifiques du "Travailleur'' et du

"Talisman''pendant les années 1880, 1881, 1882,

1883 : 1-140. G. Masson Éditeur, Paris.

Foster M. W (974. — Recent Anrarctic and

Subantarctic brachiopods. Anturcttc Research Sériés,

American Geophysical Union 1 1: 1-189.

Gofas S. 1993. — Mission océanographique SEA¬

MOUNT 2. Compte rendu et liste des stations.

Rapport non publié : 1-30.

Hondt J.-L. d 1976. — Sur quelques brachiopodes

actuels (océan Atlantique. Méditerranée,

Kerguelen). Bulletin du Muséum national d’Histoire

naturelle (3) 350, Zool. 243 : 1-13.

Jeffreys J. G. 1828. —On the Mollusca ptocured

during the “Ligluning’ and "Porcupinc expédi¬

tions, 1808-1870. ProceedingS of the Znological

Society of London, I 393-416.

Logan Â. 1979. — The Recent Brachiopoda of the

ZOOSYSTEMA • 1998 • 20(4)

559

Logan A.

Mediterranean Sea. Bulletin de l’Institut océanogra¬

phique , Monaco, 72 (1434) : 1-112.

— 1983. — Brachiopoda collccted by CANCAP I-III

expéditions to the south-east North Atlantic.

1976-1978. CANCAP-project, Contribution

No. 34, Zoologische Mcdedelingen Leiden 57 (18):

165-189.

— 1988. — Brachiopoda collected by CANCAP IV

and VI expéditions to the south-east North

Atlantic. 1980-1982. CANCAP-project, Contri¬

bution No. 73, Zoologische Mcdedelingen Leiden 62

(5): 59-74.

— 1993. — Recent brachiopods from the Canarian-

Cape Verdcan région: diversiry, biogeographic affi-

nities, bathymétrie range and life habits.

CANCAP-project, Contribution No. 91, Courier

Forschungsinstitut Settekenberg 159: 229-233.

Logan A. & Noble J. P. A. 1983. — Recent brachio¬

pods front Malta. Central Mediterranean N attirai ut

1 (2): 33-39.

Muir-Wood H. M. 1959. — Report on the

Brachiopoda of the John Murray Expédition, in

The John Murray Expédition 1933-1934, Scientific

Reports 10 (6): 283-317. British Muséum (Natural

History).

Philippi R. A. 1844. — Enumeratio Molluscorum

Sicitiae 2 : 1-303. Halle, Germany.

Saiz Salinas J. 1. 1989. — Verzeichnis der rezenten

Brachiopoden (Brachiopoda) von den iberischen

Kusten tintl den angrenzenden Meeren. Bonner

Zoologisc he Beitraege 40: 141-154.

Templado J. & Luque A. A. 1986. — Braquiopodos

de los fondos de Corallium rubrum IL.) prdximos a

la isla de Alboran (SL de Espana). Baletin del instito

espaiwl de oceanogra/ia 3 (4): 11-114.

Williams A., Carlson S. J., Brunton C. H. C... Holmer

L.. F.. & Popov L. 1996. — A supra-ordinal classifi¬

cation o( the Brachiopoda. Philosophical

Transactions of the Royal Society of London, B 351:

1171-1193.

Zezina O. N. 1975. — Deep-sea brachiopods from

the south-west Pacific and Scotian Sea. Akademia

Nauk SSSR. Trudy Instituta Okeanologii 103:

247-258.

Submitted on 26 September 1997;

accepted on 24 March 1998.

APPENDIX

STATION LIST

Station

Location

Depth

Species

METEOR

DW 136

30°01'N - 28°28’W

305 m

Eucalathis ergastica

DW 173

30°03’N - 28°43'W

920-1070 m

Dyscolia wyvillei.

Abyssothyris attantica

HYÈRES

DW 184

31°24'N - 28°52'W

675-705 m

Stenosartna davidsoni

DW 186

31°26'N - 28°51’W

1520 m

Dyscolia wyvillei ,

Stenosaiina davidsoni

DW 197

31°18’N - 28°33’W

1370-1480 m

Stenosarina davidsoni

DW 200

31°19’N - 28°36’W

1060-1100 m

Dallina sepligera

DW 203

31 °09’N - 28°43’W

845 m

Stenosarina davidsoni,

Dallina sepligera,

Dyscolia wyvillei,

Eucalathis ergastica

560

ZOOSYSTEMA • 1998 • 20(4)

SEAMOUNT 2 brachiopods from the N.E. Adantic

Station

Location

Depth

Species

IRVING-CRUISER

DW 209

31°59'N - 27°56’W

435-460 m

Platidia anomioides

DW 215

31°53’N - 28°02’W

270-275 m

Neocrania anomala

DW 216

31°53'N - 28°02’W

270 m

Neocrania anomala

DW 218

31°52’N - 28°53’W

420-480 m

Stenosarina davidsoni

DW 219

32°00’N - 27°53’W

760 m

Dallina septigera,

Stenosarina davidsoni

DW 225

32°08'N - 28°10’W

1030-1035 m

Platidia anomioides,

Stenosarina davidsoni,

? Terebratulina sp.

DW 226

32°06'N - 28°08’W

580 m

Stenosarina davidsoni

DW 227

32°07’N - 28°08'W

695-730 m

Stenosarina davidsoni

DW 229

32°02'N - 28°24’W

1715 m

Stenosarina davidsoni

DW 231

32“01 ’N - 27°54'W

745-750 m

Stenosarina davidsoni

TS 234

32°00'N - 27°41 ’W

1860-1880 m

? Pelagodiscus sp

DW 237

32 D 16’N - 27“31’W

670-715 m

Neocrania anomala,

Stenosarina davidsoni

DW 238

32'17'N - 27"32’W

890-900 m

Stenosarina davidsoni

PLATO

DW 240

33°12’N - 29°01’W

565-575 m

Neocrania anomala

DW 241

33°11’N - 28°59’W

695 m

Dallina septigera,

Stenosarina davidsoni,

Abyssothyris atlantica

DW 242

33°11’N - 28°56’W

690-710 m

Stenosarina davidsoni,

Platidia anomioides,

Dallina septigera

DW 246

33°13’N - 29°36’W

520-550 m

Stenosarina davidsoni

DW 247

33°13’N - 29°35’W

580-625 m

Stenosarina davidsoni

DW 248

33°13’N - 29°32’W

670-735 m

Dallina septigera,

Stenosarina davidsoni,

Neocrania anomala

DW 249

33°12’N - 29°15'W

1700-1800 m

Eucalathis sp. ci. ergaslica

DW 251

33°13’N - 29°28'W

900-985 m

Stenosarina davidsoni

ATLANTIS

DW 254

34°05’N - 30°13’W

275-280 m

Platidia anomioides,

Neocrania anomala,

Stenosarina davidsoni

DW 255

34°05’N - 30°15’W

335-340 m

? Eucalathis sp.,

Platidia anomioides,

Neocrania anomala,

Stenosarina davidsoni,

Phaneropora inceria

DW 256

34°06’N - 30°16'W

340-345 m

Neocrania anomala,

Platidia anomioides,

Stenosarina davidsoni

DW 257

34°04’N - 30°15’W

330-338 m

Neocrania anomala,

Platidia anomioides

DW 258

33°59’N - 30°12’W

420-460 m

Stenosarina davidsoni,

Platidia anomioides,

Dallina septigera,

? Eucalathis sp.,

Abyssothyris atlantica

DW 259

33°59'N - 30°1 l’W

635-665 m

Stenosarina davidsoni

DW 261

34°22’N - 30°27’W

1190-1340 m

Phaneropora inceria,

Eucalathis ergaslica

DW 262

34°23’N - 30°29’W

1000-1160 m

Platidia anomioides

ZOOSYSTEMA • 1998 • 20(4)

561

Logan A.

Station

Location

Depth

Species

DW 263

34°26’W - 30°32’W

610-655 m

Dallina septigera,

Platidia anomioides,

Stenosarina davidsoni

DW 264

34°24’N - 30°31'W

795-830 m

Dyscolia wyvillei,

Platidia anomioides,

Eucalathis ergastica

DW 265

34°28’N - 30°35’W

540-545 m

Neocrania anomala,

Stenosarina davidsoni,

Platidia anomioides

TS 270

34°04’N - 30°14’W

330 m

Neocrania anomala,

? Eucalathis sp.,

Platidia anomioides,

? Phaneropora sp.

DW 271

33°54’N - 30°09'W

1220 m

Stenosarina davidsoni

DW 274

34°05’N - 30°13’W

280 m

Platidia anomioides,

Neocrania anomala,

Eucalathis ergastica

TYRO

DW 275

34°03’N - 28°18’W

1590-1665 m

Neocrania anomala

DW 277

33°59'N - 28°20’W

945-1000 m

Neocrania anomala,

Abyssolhyris atlantica,

Stenosarina davidsoni,

Dallina septigera

DW 278

33°57’N - 28°22’W

890-925 m

Dallina septigera,

Stenosarina davidsoni,

Eucalathis ergastica,

Abyssothyris atlantica

DW 279

33°55’N - 28°23’W

760-805 m

Dallina septigera,

Abyssothyris atlantica

ANTIALTAIR

DW 281

43°34’N - 22°25’W

900 m

Dallina septigera,

Platidia anomioides.

Neocrania anomala

DW 283

43°34’N - 22°19'W

1175-1210 m

Stenosarina davidsoni,

Dallina septigera,

Neocrania anomala,

Eucalathis ergastica

562

ZOOSYSTEMA • 1998 • 20(4)

The Paracyclops fimbriatus -compIex (Copepoda,

Cyclopoida): a révision

Süphan KARAYTUG

Department of Zoology, The Natural History Muséum,

Cromwell Road, London SW7 5BD (UK)

School of Biological Sciences, Queen Mary and Westfield College,

Mile End Road, London El 4NS (UK)

s.karaytug@nhm.ac.uk

Geoffrey A. BOXSHALL

Department of Zoology, The Natural History Muséum, Cromwell Road, London SW7 5BD (UK)

gab@nhm.ac.uk

Karaytug S. & Boxshall G. A. 1998. — The Paracyclops fimbriatus-compiex (Copepoda,

Cyclopoida): a révision. Zoosystema 20 (4): 563-602.

KEYWORDS

Copepoda,

I‘. fnnhrmrus,

P. imminuta ,

P. chiltoni ,

taxonomy,

Palaearctic.

ABSTRACT

The type .species of Paracyclops , P. fimbriatus (Fischer, 1853), is redescribed

based on material froni rhe vieilli ty of St Pctersburg in Russia, one of the type

localities. The taxonomie status of P. fimbriatus, P. chiltoni ( Thomson, 1882)

and P. imminuta Kiefer, 1929 are stabiliscd and a neotype is designated for

P. fimbriatus. It is revealed thar faillira lo separate P imminuta front the other

two species of the complex was the main factor responsible for the taxonomie

confusion concerning ail rhree. Characters tr.lditionally used by taxonomists,

such as proportional measurements of the caudal rami and the terminal

spines of the third endopodal segment of leg 4 are found to vary considerably

within populations as well as between populations. Since there is extensive

overlap in variability expressed beeween thèse three variable species, it is

therefore concluded that most early records are unrelîablc. Previously

overlooked characters which can help to differentiate benveen them are

introduced, including the ornamentation of the frontal surface of the

coxobasis of antenna, of the posterior surface of leg 1, and of the caudal rami.

ZOOSYSTEMA • 1998 - 20 (4)

563

Karaytug S. & Boxshall G. A.

MOTS CLÉS

Copepuda.

P. fimbriatus,

P imminuta ,

P cbiUom ,

taxonomie,

Paléarctique.

RÉSUMÉ

Le complexe Paracyclops fimbri.itus (Copepuda, Cyclopoida) : une révision.

L’espèce-type de Paracyclops, P. fimbriatus (Fischer, 1853) est rodée ri te à partir

de matériel collecté à Si Pctersbourg en Russie, une localité-type. Le statut

systématique do P (hnbriatus , do P. chiltoni ( Thomson, 1882) et P. imminuta

Kief'er, 1929 est redéfini et un néotype est désigné pour P. jim brin tus. Il

apparair que la difficulté de séparer P. imminuta des deux aurres espèces est la

raison principale de la confusion systématique régnant dans ce complexe

d'espèces. Les caractères utilisés jusqu'à présent, en particulier les mesures

relatives des branches furcales et des épines terminales du troisième segment

de Tendopodite de P4 présentent des variations considérables pour une

population donnée et également entre populations et se recouvrent en partie

chez les trois espèces. Cette variabilité extrême ne permet pas de les séparer

clairement, les données anciennes paraissant parfois incertaines. D’autres

caractères pouvant aider à différencier les trois espèces sont introduits comme

Tornementarktn de la face frontale du coxdbasis de l'antenne, de lu face

postérieure de PI (basipodite et coxopndite), des rames hi ica les et la structure

du réceptacle séminal de la femelle.

INTRODUCTION

The earliesr record assigned to Paracyclops

fimbriatus (Fischer, 1853) by Dussart & Defaye

(1985) is thut of the Dattish naturalise O. H

Millier who described it under the name Cyclops

crassicomis Muller, 1785. Milliers (1785) des¬

cription and illustrations were based on an early

copepodid stage and are grossly inadéquate, so

thaï it is almost impossible to déterminé vvhether

he was dealing with a cyclopoid or harpacticoid.

The type material of Cyclops crassicomis cannot he

traced and prohably is. no longer exranr. If

C. crassicomis is a cyclopoid it Can he best

considered as spccies inccrtae sedis in the family

Cyclopidae. Unfoftunately Sars (1863) identified

some offris spécimens as C. crassicomis although

he expressed some réservations abour cnn-

specificity wirh Milliers material. In spite of this

uncertainty C. crassicomis became established in

the literaturc since other workers (Uljanin 18 7 5;

Brady 1878; Herrick 1882) followed Sars and

ignored oroverlooked Fischer’s (1853) important

paper on the cyclopids from St Petersburg. It was

not until 1892 that Brady (1892) recognized

Fischers (1853) Cyclops fimbriatus and discarded

Milliers C. crassicomis as a valid spccies ol

cyclopid. San; (1913-1918) independently

arrivée! ar the sanie conclusion and regarded it as

very qucstionable rhat Cyclops crassicomis belongs

to rhe genus Cyclops.

I lie engins ol the current taxonomie confusion

surrounding the genus Paracydops in general, and

rhe so-called P fimbriatus complex in particular,

are threefold.

1 irstly, Fischers (1853) original description ot the

type spccies P fimbriatus , Itill and valid by

contemporary mid-nineceenth centurv standards

but inadéquate by modem standards, lias heen

accepred as the standard référencé and conse-

quently has led to prohlems in later taxonomie

work. It is therelore not surprising thaï the

spccies hxs been reported Iront a vvidc range of

freshwater habitats ail over the world.

A second factor chat has contributed significantly

to the taxonomie confusion is the publication of

varions incompletely described species or forms

thar are closely related to rhe type-spectes and can

564

ZOOSYSTEMA ■ 1998 -20(4)

Révision of Paracyclops fimbriatus -complex

be considcrcd as forming part of a P. fimbriatus

species complex: these include P. chiltoni

(Thomson, 1882), P. finitimus Kiefer, 1928.

7? abnobemis Kiefer, 1929 and P. fimbriatus forma

irnminuta Kiefer, 1929.

Finally, in the course of tins study it became

apparent tliat the traditional means ol d i fièrent-

iating species within the genus Paracyclops , such

as the morphologv ol the caudal rami and leg 5,

are insufficient and in sonie cases completely

misleading since virtually no accoLint lias been

taken of intra- or inter-population variation.

Recent progress in copepod sysrematics lias raised

the level of taxonomie resolution of these

freshwater copepods and it has been démons-

trated scvetal tintes that many cyclopoid species,

reported earlier as cosniopolitan in freshwater

habitats, iiave a restricted geographical distri¬

bution (Kiefer 1981; Van de Veldc 1984; Reid

1998). A similar situation has been demonstrated

for the Cladocera (Frey 1980, 1982) and the

Rotifera (Dumont 1983).

Prior to this révision, the geographical records

and major synonyms of P. fimbriatus and

P. chiltoni wete sumniariv.ed by Dussart & Defaye

(1985) and indicated cosniopolitan distributions

for both. So, it became vital to describe typical

P. fimbriatus in detail. Artempts were made to

locate the type material but Fischer’s material is,

in ail probability, no longer extant. Unfor-

tunately. Fischer did not designate a single type

locality. Fischer’s (1853) original paper was

entitled “Contribution to the knowledge ol

Cyclopoids from the area ol St Petersburg” and in

this paper, it was stated that the P. fimbriatus

material was collected from Madeira island, the

vicinity of Baden-Baden (Gerniany), Iwanofskoje

and Peterhof (arotind St Petersburg). There was

tio clear indication on which material h is original

description was based.

An attempt was made to collect Paracyclops

material froin the vicinity of St Petersburg. No

Paracyclops material was found in the Peterhof

pond but abundant Paracyclops material was

collected Irom orher localities in the vicinity ol

St Petersburg. The redescription given bere is

based on a neotype seleeted from this material,

and supplemented by exanvitiation ol numerous

other collections from the Palaearctic région.

Examination of four unsorted samples from the

vicinity of St Petersburg revealed lour Paracyclops

species: P fimbriatus, P chiltoni, P. irnminuta and

P poppei. Paracyclops puppci can easily be sepa-

tated from the other tliree species, but P. fim¬

briatus, P. chiltoni and P. irnminuta are easily

conflised. Traditional means of diffeteniiating

between these taxa, such as the morphology of

the caudal rami, the proportional length of

terminal spincs of leg 4 and the structure ol leg 5

are insufficient and in some cases misleading,

thcreforc most carly records are unrcliable.

The ohject ol this paper is to redescribe

P fimbriatus, P irnminuta and P chiltoni in detail,

to identify their major synonyms and to

introduce previously overiooked characters which

are important in differentiating them.

METHODS

Specimens were dissected and mounted in

lactophenol. Broken glass-fibres were added to

preveut the appendages from being compressed

by the coverslip and to facilitate rotation and

manipulation which allowed viewing from ail

sides. Ail drawings were made with the aid of a

caméra lucida using an Olympus B1I-2

microscope cquipped with Nomarski differenrial

inrcrfcrence contrast and ail measurements made

with an ocular micrometcr. Body lengths were

measured frotn the base ol the rostrum to the

posterior edge of the caudal rami. Body vvidth is

given as the widest part of the ccp halo thorax. In

the spinc and seta formula ol the swimming legs

Roman numcrals and Arabie numerats are used

lor spines and setae, respectively. The rermi-

nology proposcd by Huys & Boxshall (1991) is

adopted. The ternis “frontal” and “caudal”

introduced by Van de Vclde (1984) to dénoté the

anterior and posterior surlace of the antennary

coxobasis are also adopted.

ARBREVIATrONS

NHM The Natural History Muséum, London;

MNIIM Muséum national d’Histoire naturelle,

Paris;

7.M 7oologi.sk Muséum, Oslo;

USNM United States National Muséum,

Smithsonian Institution, Washington.

ZOOSYSTEMA • 1998 • 20 (4)

565

Karaytug S. & Boxshall G. A.

SYSTEMATICS

Paracyclops fimbriatus (Fischer, 1853)

(Figs 1-9)

Original description:

Cyclops fimbriatus Fischer, 1853 [Bull. Soc. Imp. Nat.

Moscou 26, 1; 94, Figs 19-28, 30].

Synonymy:

Cyclops fimbriatus Fischer, 1853: 94, pl. 111, Figs

19-28,30.

Cyclops crassicomis O, F. Millier, 1785 sensu Brady

1878: 118, 1 19, pl. 23, Figs 1-6.

Cyclops soli Kokubo, 1912: 102, pl. Il, Figs 24-26.

Platycyclops fimbriatus (Fischer, 1853) - Sars 1913-

1918: 81,82, pl. L.

Cyclops {Paracyclops) fimbriatus Fischer, 1853 - Pesta

1928: 114-116, Figs 96A-U, 97A-C. - Reuter-

Schiltz &t Hoffmann 1970: 229-231, figs 27-31.

Paracyclops fimbriatus I. typiat (Fischer, 1853) — Kiefer

1929: 50, Àbb„ 53.

Paracyclops abnobensis Kiefer, 1929: 51, Abb. 57-59.

Paracyclops vagus l.indberg, 1939: 45-50, figs la-l.

Type MATERIAL. —A neotype is desienated herein and

deposited in the collection of die NHM, Reg,

No. 1997.1762. lt is a Female collected at

P Dudelgofka, St Petersburg, Russia and dissected on

4 slides.

OTHER MATERIAL. — Russia. River Igora (Neva

systein), about 10-15 km east ol St Pctcrsburg, depth

0.5 ni, t 18 "C, 22.VII.1996, coll. V, Alekseev:

14 9 9. — P. Dudcrgofka, St Pctcrsburg,

27.VII, 1996, depth 0.5 m. t 18 “C, coll. V. Alekseev:

4 9 9. — River Ravan (Ladova System), about

100 km east ol St Pctcrsburg, depth 0.2-0.5 m,

t 17 °C, 22.VI1.1996, coll. V. Alekseev: 2 95. -

St Petersburg, watcv supply syslcm, pond |W$, July

1984, coll V Alekseev 1 d, — Baikal pulp &paper

mill, pond oFcleaning System, in the vicinity ot l.ake

Baikal, 9.V1T.1984, coll. V. Alekseev: 4 9 9,’3 d <3.

Turkcy. Demirkoprü, coll. IJ. Demirhindi: 7 9 9

(NHM 1966.3.24.23).

Sweden. Lakc Maint. Uplaiid: 4 9 9 (Norman

Collection, NHM 1911.11.8.40935-939).

Ircland. Locality unspecified: 24 9 9, 7 d d

(Norman Collection. 1911.11.8.40915-934; selected

Iront more th.ui 200 specirnens in mixed sample,

ntostly P immitnaa). — Rossmore, Co. Monaghan:

1 9 (NHM 1911.11.8,40914).

Palestine. 33 9 9 (NHM 1938.3.9.83-89 1002a).

Norway. 69 9 9, 16 d d (G. O. Sars Collection ZM,

F20478: no locality is given by C. O. Sars on the ori¬

ginal label but on a second label is written “Norv”

which indicatcs rhat the material is probably From

Norvvav. This material contains a mixture of

P fimbriatus and P. imnunuta.)

Mongolia. Lake Orog Nuur, Cioby Desert, 11.VII.

1995, coll. V Alekseev: 1 9, 14 dd.

India. Nagarjuna Universiiv réservoir fed from River

Krishna, a VilI I9-6, coll. Y. Ranga Reddy: 7 2 9,

6 dd. — River Krishna ai Vijayawada,

16 ,, 3I N - 80’39’E , January 1974: 3 9 9.

Finland. Province ol South Hame. tminicipality of

I.ammi, Lake Paajarvi, western arm, organic poze

mixed with minerai parrides at 3-5 m depth,

61'04'N - 25“08’E, 18.XI.1996, coll. ). Sarvala:

25 9 9, 1 6-

Germanv- Oldenburg, 2.VI 11.1996, coll. T. Ishida:

26 9 9, 12 dd.

Japan. Tomakomai, Hokkaido, 4. XL. 1987: 4 9 9,

8 dd. — Samt locality. 12.V. 1987: 6 9 9, 1 d. —

Ryuku, I2.V. 1996, coll. I. Ishida: 1 2, l d . — Lake

Biwa, Honshu, 17-18.111.1986, coll. T. Ishida: 2 9 9

dissected on two slides. — Hokkaido Lbiiversity,

Sappom, Hokkaido, 9.V1II.1987, coll, F ishida:

6 9 2 whole mounts on I si idc, 2 S d whole rnounts

on I slide, 2 2 Ç djsçecred on slide. — Tokachi,

Hokkaido, S.VIII. 1983, coll. F Ishida: 1 9, 1 d

whole mounts.

U/bekistan. Fown Muynak near Aral Sea,

Kashltadavya Région. Chauvak Réservoir, Tashkent

Région, coll. 1. M. Mïrabdullaycv: 6 9 9.

Scotland. Loch Ncss, I9.VI1.I992: 7 9 9, 5 d d.

Distribution. — Examination of numerous

collections ol Paracyclops front the Palaearctic, Nearctic

régions, and From Africa lias revcalcd thaï P fimbriatus

is nor as widely distributed as previously believed

(Karaytug 1998). Based on examined tnarerials,

P. fimbriatus is, at présent, known to lie disrributed in

Europe and Asia extending eastwards to înclude

litrkey, Palestine, China, Japan and India.

P. fimbriatus appears to be widely distributed

throughout the Palaearctic région.

RlDLSCItin ION

Adult female

Body length and width not including caudal

setae given in Table 1. Prosome (Fig. IC) with

céphalothorax narrowing anteriorly and three

Iree pedigerous somites decreasing in width From

anterior to posterior. Third pedigerous somite

with minurely denticulate hyaline Irill along the

posterior margin. Third and fourtli pedigerous

somites with patch ol spinules at each

postérolatéral corner. Urosome (Fig. 2A, B)

566

ZOOSYSTEMA • 1998 - 20(4)

Révision of Paracyclops fimbriatus-œmp\ex

Fig. 1 . — R fimbriatus-, A-C, E, F, neotype, adult 9 ; D. adult 6 ; A, maxillule; B, maxilliped; C, body, dorsal; D, body, dorsal; E, maxilla

F, mandible. Scale bars: A, B, E, F, 50 pm; C, D, 400 pm.

ZOOSYSTEMA • 1998 • 20 (4)

567

Révision of Paracyclops fimbriatus-comp\e\

Fig. 3. — P fimbriatus ; neotype, adult S; A. antennule with arrow indicating spiniform seta at anterodista! corner on third segment

B, antenna, coxobasis and first endopodal segment, frontal; C, antenna, caudal. Scale bars: A, 50 pm; B, C, 25 pm.

ZOOSYSTEMA • 1998 • 20 (4)

Karaytug S. & Boxshall G. A.

consisting of fifth pedigerous somite, génital

double-somite and thrce hcc abdominal somites.

Fifth pedigerous somite with Iringe-like elongate

setules ar postérolatéral angles. Génital double-

somite, second and third abdominal somites

ornamenred with irregular pattern of Fine pirs

dorsally as figured (Fig. 2A). Séminal réceptacle

divided into broad anteiior and posterior lobes

(Fig. 2B). Médian copulatory pore located

ventrally about halfway along lengrh of génital

double-somite. Posterior margin of génital

double-somite and following two abdominal

somites with finely incised hyaline fri11. Anal

somite with ventral spinulaf rovv extending

laterally and dorsally almost to either side of anal

operculum. Anal operculum weakly developed.

smooth; and opening bordered by spinular row

(Fig. 2A).

Caudal rarni with slightly convex inner margin;

very variable in length and shape (Figs 2A, B,

7A-F); armed with six setae (Fig. 2A); seta (1)

absent; antérolatéral seta (II) plumose with

spinules at base on dorsolateral surface;

postérolatéral seta (III) unilaterally plumose with

spinules along dorsal surface, spinular row at base

ventrally extending dorsally; terminal accessory

seta (VI) plumose; outer terminal seta (IV) and

inner terminal seta (V) well-developed and

heterogeneously ornamented (Fig. ICi).

Antennule 8-segmented (Fig. 3A). First segment

with ventral spinuiar tows in proximal half.

Segment 3 with partial suture line and spiniform

seta near anterodistal corner (arrowed in Fig. 3A).

Segmenv 5 wivh characteristic short aesthetasc.

Segment 7 with aesthetasc located distally on

anteroventral margin. Apical segment with

aesthetasc fused basally to adjacent seta. Setal

formula 8, 12, 6, 5, 2 + I aesthetasc, 2,2 +

aesthetasc, 7 + aesthetasc.

Antenna 4-segmented (Fig. 30, comprising

coxobasis and 3-segmented endopod. Coxobasis

with complex ornamentation on caudal (Fig. 30

and frontal (Fig. 3B) surfaces as figured, and

armed with two inner spinulose setae plus very

long outer spinulose seta representing exopod.

First endopodal segment with inner distal

spinulose seta and spinules along outer margin.

Second endopodal segment wirh ni ne setae, of

which six along inner margin and three arranged

around distal inner corner; ornamented with

spinules along outer margin. Third endopodal

segment armed with seven setae around apex;

outer margin ornamented with short spinules

proximally and long spinules distally.

Labrum (Fig. 2F.) torming broad posterior

ourgrowth. Disral margin with strong teeth;

anterior surface ornamented with paired groups

of long spinules.

Mandible (Fig. 1F) consisring of well-developed

coxal gnarhobase and reduced palp. Gnathobasis

blades pointed, nmstly simple, dorsal seta with

spinules along inner rim. Paip represented by

minute segment, bearing three spinulose setae,

two of which very long and one short. Coxa with

spinular row along outer margin between palp

and gnathobàse, and with lacerai group of

spinules near insertion of palp.

Maxillule (Fig. IA) consisting of powerful

praecoXa and reduced 2-segmented palp.

Praecoxal arthrite armed with six setae

articulating at base and five spines fused to

segment; proximalmost articulating spine

spinulose, other spines naked. Proximal segment

of palp representing fused coxa and basis, bearing

one strong spine and two naked setae apically,

plus outer spinulose seta representing exopod.

Distal segment of palp, representing endopod,

armed with three setae, outermost seta spinulose.

Maxilla 5-segmemed (Fig. 1E) comprising

praecoxa, coxa, basis and 2-segmented endopod.

Praecoxa with spinular rows on outer margin and

dorsally. Praecoxal enditc with two spinulose

setae. Coxa with proximal endite represented by

single spinulose seta; distal endite cylindrical,

with strong spinulose spine and naked seta

apically. Basis drawn out into poWerfi.il curved

claw bearing coarse spinules along middle part of

inner margin; accessory armature consisung of

strong spine; with spinular row along convex

margin and naked seta. First endopodal segment

carrving two setae, second with three setae.

Maxilliped 4-segmented (Fig. 1 B) comprising

syncoxa, basis, and 2-segmented endopod.

Syncoxa armed with ihree spinulose setae

representing endifes, tew r long spinules arranged

near base of setae; ornamented with spinular row

near outer distal angle. Basis armed with one

spinulose and one naked sera; ornamented with

570

ZOOSYSTEMA • 1998 -20(4)

Révision of Paracyclops fimbriatus-comp\ex

Fig. 4. — P fimbriatus; A, B, E, G, neotype, adult 9 ; C. non-type adult 9 ; D, adult S ; F. non-type 9 ; A. leg 2, anterior; B, C, intercoxal

sclerite and coxa of leg 2, posterior; D, terminal endopodal segment of leg 1 ; E. F, intercoxal sclerite and coxa of leg 1, posterior; G,

leg 1, anterior view. Scale bar: 100 pm.

ZOOSYSTEMA ■ 1998 - 20(4)

571

Karaytug S. & Boxshall G. A.

two transverse rows of spimiles near outer

margin. First endopodal segment bearing claw-

like seta with spinules at midlength. Second

endopodal segment with three setae, one of

which naked; other spinulose.

Legs 1 to 4 with 3-segmented protopod (Figs 4A,

G, 5A, F). Praecoxa represented by triangular

sclerite at oüter proximal angle; each proropodal

segment with spinular row on outer corner of

margin. Coxa with complex ornamentation on

both anterior and posterior surfaces as tlgured.

Basis with plumose outer seta. Endopodal

segments with long spinules aiong outer margins.

Ail spines on segments of both rami with spinules

at thelr bases. Exopodal segments I and 2 with

short spinules along outer margins. Legs 2 to 4

each with spinular rows on anterior surface of

endopodal segments 1 and 2 and exopodal

segment I and posteriorly on exopodal

segments I and 2. Legs 2 and 3 with posterior

spinular rows on endopodal segment 2.

Leg 1 (Fig. 4G): coxa with inner plumose seta.

Basis with spinulose spine on inner margin

bearing rwo groups of spinular rows at hase, one

of which long and fringe-like; also omamented

with spinular rows ameriorly near base of

endopod. Intercoxal sclerite omamented with

spinular rows anteriorly (Fig. 4G) and posteriorly

(Fig. 4E, F). Exopodal segments 1 and 2 and

endopodal segment 2 with spinular rows

posteriorly. Spine of exopodal segment 1 with

flagellate apex. Seta next to outermost spine of

terminal exopodal segment scmispinulose

(Fig. 4G).

Leg 2 (Fig. 4A): intercoxal sclerite with spinular

rows anteriorly and posreriorly (Fig. 4A-C). Coxa

with inner spinulose spine.

Leg 3 (Fig. 5F): intercoxal sclerite with (Fig. 5G)

or without (Fig. 5F) spinular row anteriorly, with

two spinular rows posreriorly (Fig. 5H). Coxa

with inner spinulose spine.

Leg 4 (Fig. 5A); intercoxal sclerite with patch of

spinules on anterior surface (Fig. 5A) and with

three spinular rows posteriorly (Fig. 5B, G). Inner

coxai spine with group oi setules mainly

originating posteriorly (Fig. 5C). Basis with

spinular row near inner margin posteriorly

(Fig. 5B, G). Endopodal segment 2 without

spinular row posteriorly.

Spine and seta formula as follows:

Coxa

Basis

Exopod

Endopod

Leg 1

0-1

1-1

: 1-1; II!. 5

0-1; 0-1; 1, 1, 4

Leg 2

0-1

1-0

1-1

; 1-1; III, 1, 5

0-1; 0-2; 1. 1, 4

Leg 3

0-1

1-0

1-1

; 1-1; III, 1, 5

0-1; 0-2; 1, 1, 4

Leg 4

0-1

1-0

1-1

; 1-1 ; II, 1, 5

0-1; 0-2;1, 11,2

Leg 5 (Fig. 2C) comprising single free segment,

armed with one long (almost twice as long as

inner spine) multispinulose outer seta, one strong

inner spine with 3-4 spinules around base, and

One plumose seta in middJe. Leg 6 (Fig. 2A)

represented by one plumose seta and one tiny

spinule dorsolatcrally.

Adult male

Body iength 716 pm and width 260 pm

(St Petersburg), the body Iength and width

measurements of the males front other localities

are given in Table 1. Body (Fig. 1D) differing

iront adult temale as follows: urosome

6-segmented (Fig. 8A, C), comprising fîfth

pedigerous, génital and four free abdominal

somites; genitaJ, third, fourth and fifth

urosontites omamented with cuticular pits

dorsally (also présent ventraliy on Itturth

urosomite). Caudal rami shorter than female.

Antennule digeniculate (Fig. 9A, C, D),

indistinctly 15-segmented. Segment 1 armed

with eight setae; one seta (seta A) large and

modified by ornamentation of strong spinules in

proximal and mid sections, tapering to fine point

distally (see inset in Fig. 9D); ornamenred with

row of spinules ventraliy (Fig. 9E). Segment 2

with four setae. Segment 3 with two setae.

Segment 4 with two setae plus aesthetasc.

Segments 5 and six each witlt two setae.

Segments 7 and 8 separated from each other by

extensive arthrodial membrane (Fig. 9D):

segment 7 with rwo setae, segment 8 with two

setae. Segment 9 with two setae plus a short

aesthetasc, fused to segment 8. Segment 10

(= ancestral segment XV) produced on one side

into sheath enclostng segment 11 ventraliy;

armed with two setae, one omamented with long

setules u ni latéral ly, other longer and naked.

Segment 11 bearing curved seta omamented with

572

ZOOSYSTEMA ■ 1998 -20(4)

Révision of Paracydops flmbriatus-complex

Fig. 5. — P fimbriatus ; A, B, F, neolype, adult 2; C. D. E, non-type 2 ; A, leg 4 . anterior; B, C, intercoxal sclerite and coxa of leg 4,

posterior; D, first exopodal segment of leg 4 , posterior (Finland); E, terminal endopodal segment of leg 4 . anterior: F, leg 3, anterior;

G, intercoxal sclerite of leg 3, anterior; H, intercoxal sclerite and coxa of leg 3, posterior. Scale bar: 100 pm.

ZOOSYSTEMA • 1998 - 20(4)

573

Karaytug S. &C Boxshall G. A.

Table 1. — Body length (BL) and width (BW) measurements (in gm) of adult Paracyclops fimbriatus from various localities (N, number

of specimens measured).

LOCALITY

SEX

(mean ± SD)

RANGE

(mean ± SD)

RANGE

St Petersburg (River Igora)

885 ± 92.5

684-1017

309 ± 12.6

289-333

St Petersburg (P. Dudergofka)

916 ±91.5

815-993

370 ± 12.5

358-383

Russia (Vicinity of

862 ± 52

802-894

314 ±7.5

306-321

Lake Baikal)

754 ±16

738-770

254 ±5

249-259

Mongolia

743

296

<î

651 ±19.3

605-672

228 ±15.4

202-249

India

624 ± 22.9

600-662

231 ±6.5

222-240

558 ± 27

527-589

204 ± 3.5

200-207

Turkey

776 ± 67.5

707-854

302 ±15.3

289-326

Finland

893 ± 48.4

808-983

329 ± 9.3

313-351

786

262

Sweden

923 ± 30.4

901-944

340 ± 27.6

320-359

Germany

834 ± 38.2

786-926

336 ± 13.1

318-359

696 ±31.6

641-728

280 ± 4.5

272-285

Japan

862 ±41.6

777-909

308 ± 13.6

282-328

748 ± 42.4

693-810

259 ±4.9

247-268

Ireland

831 ±40.9

767-901

336 ± 16.2

311-363

721 ± 22.3

699-757

276 ±7

268-285

Scotland

916 ±50.9

827-998

337 ±21.5

295-359

816 ±80.3

722-913

287 ± 18.7

264-307

Palestine

743 ± 48.1

709-777

256 ± 2.8

254-258

Uzbekistan

782 ± 32.5

724-846

258 ± 19

234-284

double rovv of strong denticles, plus one naked

seta (Fig. 9A, C). Segment 12 armed with minute

naked seta, plus short, highly chitinized spine.

Segment 13 armed with one short spinulate seta

proximally, four short naked sérac, plus one

modified element attached to segment by short

stalk (Fig. 9A); main part of modified element

lying along surface of segment and ornamented

with longitudinal ridges and small central pore.

Geniculation located between segments 13 and

14. Segments 14 and 15 parily fused (Fig. 9B),

forming cttrved subchela-like section: segment 14

armed with two setae and two modified éléments

each ornamented with longitudinal ridges and

a central pore (as proximal element on seg¬

ment 13). Apical segment tapering distally;

armed with eleven setae and one aesthetasc,

mosdy origtnating on outer (= posteriori surface.

Segmentai fusion pattern as follows: 1-V, Vf-VII,

VIH, IX, X, XI, Xll. XIII, XIV, XV, XVI, XVII,

XVIH-XX, XXi-XXIII, XXIV-XXVW.

Ail other appendages as in female except:

coxobasis of antenna with additional spinular

row near inner spinulose setae (arrowed in

Fig. 8E), innermost seta of coxobasis srrongly

spinulated. One inner seta on endopodal

segment 3 ot leg 1 spinulose (Fig. 4D). Outer seta

of fifth leg pluinose and less developed than in

574

ZOOSYSTEMA • 1998 -20 (4)

Révision of Paracyclops fimbriatus-c omplex

Fig 6 — R limbriatus • non-type, adult 9; A-C, leg 5, ventral (India); D E, leg 5, ventral (St Petersburg); F, leg 5, ventral (Germany);

G, leg 3, intercoxal sclerite, posterior (Lake Baikal); H (anterior), I (posterior), leg 4, intercoxal sclerite (Russia); J, K, leg 4, intercoxal

sclerite, posterior (India). Scale bars: A-F, 25 pm; G-K, 50 pm.

ZOOSYSTEMA • 1998 • 20 (4)

575

Karaytug S. & Boxshall G. A.

Fig. 7. P fimbriatus; A-F, non-type, adult 9 ; G, adult S ; A-E (Europe), F (India), caudal rami, dorsal; G, caudal rami (India) Scale

bar: 50 pm. '

576

ZOOSYSTEMA • 1998 -20(4)

Révision of Paracyclops fimbriatus-conup\tx

Fig. 8. — P. fimbriatus ; adult 6\ A, urosome, dorsal; B, leg 4, intercoxal sclerite, posterior; C, urosome, ventral; D, detail of leg 5 and

leg 6, ventral; E, antenna, coxobasis and first endopodal segment, caudal. Scale bars: A, C, 100 pm; B, E, 25 pm; D, 50 pm.

ZOOSYSTEMA • 1998 -20(4)

577

Karaytug S. & Boxshall G. A.

female (Fig. 8D). Sixth leg (Fig. 8D) armed with

one inner spine (as Long as third urosomal somite

and omamemed with about elevcn spinules at

base) and two outer plumose setae.

Variability

Females

Measurements of body lengths and widths are

given in Table I. One female from St Petersburg

possessed an extra spinular row on the anal

somite ventrally (Fig. 2D). Specimens from

Finland consistently bad two spinular rows on

exopodal segment 1 of ieg 4 on posterior sutface

(Fig. SD). Tbese rows appcarcd inconsistcntly on

specimens from St Petersburg and Turkey. I bis

variability is also i I Inst rat ed in Damian-

Georgescu’s (1963) drawings baxed on material

from Romania. Tbe exopodal spines of the

swimming legs and the inner spine of leg 5

(Fig. 6F) of some specimens from Oldenburg

(Germany) appeared to be stouter tban in other

populations. Despite tbe fact that material Irom

Germany had stouter spines on swimming legs,

as did the Finish material, the two posterior

spinular rows on the First exopodal segment of

leg 4 were not consistently présent in German

material. Additional variability was noted (or the

spinules on anterior surface of ihe intercoxal

sclerite of leg 3 which may be présent or absent in

any population examined (Fig, 5F, G) and some

specimens from the viciniry of Lake BaikaI had

three rows of spinules posreriorly on rhe

intercoxal sclerite (Fig. 6G),

Variability in the arrangement and number of

posterior spinular rows on rhe intercoxal sclerite

of leg 4, as well as in the shape of rhe inner coxal

spine (degree of curvature) on leg 4 was also

noted in some specimens from the victnity of

Lake BaikaI (Fig. 6H, I) and from India (Fig. 6J,

K). The material from India espccially showcd

interesting variability in leg 5 (cf Fig. 6A-C) as

the inner spine was usually longer than in other

material examined and there were spinules

présent at the base of the middle seta in some

specimens (Fig. 6B, C). Variability in the struc¬

ture of the inner spine of leg 5 in specimens from

St Petersburg région is shown in Figure 6D, E,

but no consistent pattern was observed in this

variability.

Males

In material from Japan, a group of spinules

(arrowed in Fig. 8E) on the coxobasis of antenna

can be very small or absent in some specimens.

DlFFLRENTIAL DIAGNOSIS

P. fnnbriatus is distinguished by the following

unique combination of characters: the structure

of leg 5 (Fig. 2C), the spinular ornamentation of

coxobasis of antenna in both sexes (Figs 3B, C,

8E), the structure of the teceptaculum seminis

(Fig. 2B), the absence of the aesthetasc on the

first segment ofthe male antennule (Fig. 9A).

7? fimbriatus and P imminuta are easilv confused

as a rcsult of their close morphological similarity.

Nevcrthelcss, rhe absence of the wcll-dcveloped

spinular row on the frontal surface of the

coxobasis of the antenna in P fimbriatus (Fig. 3C)

can be used unequivocally to separate females of

P fimbriatus from P imminuta. The spinular row

is sexually dimorphic and, in the adulr male of

P fimbriatus (Fig. 8E), the spinules arc Icss well-

dcveloped than in P imminuta (arrowed in

Fig. 1 5D). In addition, the ornamentation ofthe

mid-distal spinular ornamentation on the

posterior surface of leg 1 in P imminuta is

significamly different (arrowed in Fig. 13B) from

that of P. fimbriatus (Fig, 4F). P fimbriatus can

easily be separated from P chiltoru by the length

of outer seta ol leg 5 in the female (Fig. 2C), by

the absence of cuticular dépréssions on the

ventral surface of the caudal rami in the female

and by the structure of the séminal réceptacle

(Fig. 2B),

Remarks

Most early descriptions of this species lack detail

and several subspecies hâve been described on the

basis of characters that are very variable. Most

early records are, therefore, unreliable. Below,

comparions are made only with the important

descriptions that conform to /.’ fimbriatus as

described above. Howevcr, one record of

"P fnnbriatus" from Iran (Lindberg 1941) should

be mentioned hère because the material described

by Lindberg appears to be significantly different

from the typical form described in detail above,

on the basis of ver)' short inner spine of leg 5

(Lindberg 1941: 477, figs a, c). This record does

578

ZOOSYSTEMA • 1998 • 20 (4)

Révision of Paracyclops fimbriatus-complex

Fig. 9. — P. limbriatus ; adult S, antennule; A, anteroventral showing setation; B, terminal segment, posteriori C, ventral showing

segmentation: D, dorsal showing segmentation; E. first segment showing setation, anteroventral. Scale bars: A, B, 25 pm; C-E,

50 pm.

ZOOSYSTEMA • 1998 • ZO (4)

57Q

Karaytug S. & Boxshall G. A.

not conform to P fimbriatus as deftned here and

may rcp resent a ncw species.

After Fischer (1853). a more detailcd description

of P. fimbriatus was given by Brady (1878). In

Brady's drnwings the caudal rami are inserted

very vvide apait. This character is typical of

P. fini b ri ri tus and is never lound in R imminuta.

Later, further improved illustrations were

provided by SchtneiJ (1892) showittg long caudal

rami, the structure of leg 5 and the séminal

réceptacle. These characters, though variable,

help to confirm the identity of his material as

7? fimbriatus. After Schmeil, similar descriptions

were given by Sars (1913-1918) as Platycyclops

fimbriatus and by Pesta (1928) under the name

Cyclops (Paracydops) fimbriatus.

Although Kokubo (1912) presumably overlooked

the third seta of leg 5 or possibly examined an

aberrant specimen, his description of Cydops soit

indicates tliat it is a synonym of R fimbriatus.

Kiefer (1929) separated /.’ abnobensis Irom

P. fimbriatus on the tnisis of shorter (not quite

4 times as long as broad) but widely separated

caudal rami. In addition, in leg 4 the inner apical

spine is twice as long as the third endopodal

segment and the outer seta of leg 5 is relarively

long. These characters do not differ significantly

from those of P fimbriatus. It is now widely

accepted that the length of caudal rami is a very

variable feature within species of Cyclopidae in

general. Individual examination ot specimens in

samples of P fimbriatus from several localities in

Europe, especially those specimens with short

caudal rami (sec Fig. 7A), revealed no consistent

variation in characters other than length of

caudal rami to support the validity of R. abno-

bensis. Conscquentiy, P abnobenis is treated here

as a synonym of P. fimbriatus, followîng Dussart

& Defaye (1985). Variation in the length of

caudal rami was also indicated by Monchenko’s

(1974) description of P fimbriatus in which the

caudal rami are shown as very short and widely

separated.

Lindberg (1939) descri bed P va pu s as a new

species from India but later synonymized it with

P fimbriatus (Lindberg 1958). Study of material

from India that was idcncified as P vagtts by

Ranga Reddy Radhakrishna (1984) revealed

variation in leg 5 {cfi Fig. 6A-C), as the inner

spine was usually longer than in other material

examined and there were spinales présent at the

base of the middle seta in soute specimens

(Fig. 6B, C). Consistent variation, however, was

not observed and no additional characters

supporring the status of P vagin as a distinct

species or subspecies were lound. In accordance

with Lindberg (1958) P vagus is treated here as a

synonym of P fimbriatus.

Paracyclops imminuta Kiefer, 1929

(Figs 10-16)

Original description:

Paracyclops fimbriatus f. imminuta Kiefer, 1929 [Zur

Kcnnrnis einiger Artgruppen der Süsswasser-

Cyclopidcn. Z. wiss. Zool. 133: 1-56, figs 1-61].

Synonymv:

Cydops fimbriatus Fischer var: Kiefer, 1926: 278.

Paracydops fimbriatus f. imminuta Kiefer, 1929: 49,

50. Àbb. 54-56.

Paracyclops fimbriatus Fîschcr. 1853 sensu Gurney

1933': 121 126, figs 1438-1458.

Paracyclops fimbriatus orievtalis Alekseev, 1995:

133-138. figs 1,2.

TYPE LOCAUTY. — Germanv: Mains water System of

Oefitigen (Kiefer 1929).

Material examined. — lt has not been possible to

examine the type material of Paracydops fimbriatus

f. imminuta Kiefer, 1929. It is not listed in the

catalogue of the Kiefer collection (Frauke 1989) and

may not lie extant.

England. Co. Durham: 26 9 9, 10 d d (Norman

Collection, NHM 1911.11.8.40884-903). —

Richmond Purk. Surrev, 5.VI.I967, colJ. A. I Bruce:

9 ? 9, 3 dd (NHM 1967.9.1.242). — Rainton

Meadows, Durham: 34 9 9, Il od (Norman

Collection, NHM 1900.3.29.261). - Somerset: 2 9

(NHM 1953.6.29.1). — Reechwood Paris: 1 9

(NHM 1965.4.1.1.5). — La agites s Cave, Deep

Threshold, pardy dried up pool, pli 6.7, t 21.5 °C,

conductivity 725 ms/cm, coll. M. Moseley: 3 9 9. —

Fltghgate: 3 99 (NHM 1951.11.9.3). — Derby

Power Station: I 9 (NF1M 1963.7.16.3). —

Manchester: 2 9 9, 1 d (NHM 1961.3.10.f). —

Langues.*. Peninsula (NGR SC) 281653) from damp

pebble/gravel on floor, 2"-3” below ground, (material

was collected from a sea cave which is only inundated

by the sea during storms, freshwater enfers as seepage

at other times), 30.VII.1994: 3 9 9,4 d d.

580

ZOOSYSTEMA • 1998 - 20 (4)

Révision of Paracyclops fimbriatus-c omplex

Fig. 10. — P. imminuta-, adult 2; A, antennule; B, body, dorsal; C, maxilliped; D, maxilla. Scale bars: A, C, D, 50 pm; B, 200 pm.

ZOOSYSTEMA • 1998 *20(4)

581

Karaytug S. & Boxshall G. A.

Fig. 11. — P. imminuta ; adult 9; A, urosome, ventral; B, urosome, dorsal; C, leg 5, ventral; D-F, caudal rami, dorsal. Scale bars: A,

B, F, 50 [irr\; C, D, 25 pm; E, 100 \}m.

coi

ZOOSYSTEMA • 1998 *20 (4)

Révision of Paracyclops fimbriatus-comp\ex

Fig. 12. — P. imminuta-, adult 9 ; A, antenna, coxobasis showing variant pattern of spinulation, caudal; B, antenna, caudal; C, antenna,

coxobasis and first endopodal segment, frontal; D, mandible; E, maxillulary palp; F, maxillule. Scale bars: 50 pm.

ZOOSYSTEMA • 1998 -20 (4)

583

Karaytug S. & Boxshall G. A.

Russie. River Jgors (Neva System), about 10-15 km

east ot St Perersbutg, 22. V 11.1096, depth 0.5 m,

t 18 ”C, coll. V. Alckseev: I 9. — P. Dudergofka,

St Petcrsburg. 27.VÜ.I996, dcprlt 0.5 m, t 18 °C. coll.

V. Alekscev: I V. — St Petcrsburg vvater supply System

pond, Au gu s r 1984, coll. V. Alekscev: 2 9 9.

Ircland. Sevcral fbinales and males selccted Iront more

titan 200 spécimens (Norman Collection, NHM

1911.11.8.40915-934).

Norway. 69 9 9, 16 66 (G. O. Sars Collection, ZM

F20478; no localiry is given by G. O. Sars on the

original label but on a second label is written: “Norv",

which indicatcs tinte the inateri.il is probably from

Norway; thi.s material contants a mixture of

/:’ fimbriatus and /.' imminuta.).

Sweden. Pake Malar, Upland: 4 9 9,266 (Norman

Collection, NHM 1911.11.8.40935-939).

Grccce. Delphi: 1 ?, 1 6 (Gurney Collection, NHM

1937.11.16.620-1).

Israël. 3 9 9 on rwo slidcs (Glassman Collection; En

Quntran, 1 ES cop 226301 ); 1 2 on ! siide (Glassman

Collection; En Ziv, IFS cop 200701),

Adores. Pico, 12.X. 1971: 2 9 9 dissected on three

si ides (Th. Monod Collection n” 15190, MNH'N); 2

66 niounrcd on one slide H h. Monod Collection

n u 15188, MNFIN). lerceir.t, 15.X.1971: I 9

Fig. 13. — P. imminuta ; A, B, D, E, adult 9 ; C, adult <?; A, leg 1, anterior; B, intercoxal solerite and coxaof leg 1, posteriori C, terminal

endopodal segment of leg 1; D, intercoxal sclerite and coxa of leg 3, posterior; E, leg 3, anterior. Scale bar: 50 pm.

584

ZOOSYSTEMA • 1998 * 20(4)

Révision of Paracyclops fimbriatus-comp\a

Fig. 14. — P. imminuta ; adult $; A, leg 4, anterior; B, intercoxal sclerite and coxa of leg 4, posterior; C, intercoxal sclerite and coxa

of leg 2, posterior; D, leg 2, anterior. Scale bar: 50 pm.

ZOOSYSTEMA • 1998 * 20 (4)

585

Karaytug S. & Boxshall G. A.

dissected on ont' siide (Tli Monod n° 1 5208, MNHN

Collection).

Gibraltar. Leonora Cave, from pool with rotren rope

in it: 1 ?. 5 6 6 (NHM 1958.8.5.1).

France. Lac Léman, August 1962, coll. B. Dussart:

3 9? (MNHN-Cp897).

Distribution. — On the basis of examined material

only, the distribution of P. bnminuta extends across

Europe to central Russia, and southwards to include

Israël.

Redescription

Adult female

Body lengths and widths are given in Table 2.

Urosome ornamcnted with finer surface pits

dorsally than P. fimbriatus as figured (compare

Figs 11 A, B and 2A). Génital double-somite

hardly increasing in width antcriorly, whereas in

P. fïmbriatus geniral-doubk somite increasing in

width anteriorly. Séminal réceptacles dividcd into

broad anterior and narrower posterior lobes

(Fig. Il A), lobes narrower and more distinctly

separated than chose of P. fimbriatus. Caudal ranci

positioned in parallel configuration, and

frequently very close to each other; in some

specimens slightly converging proximally, parallel

distally (Fig. 1 IA, B, D-F).

Coxobasis of antenna with complex ornament¬

ation on caudal and frontal surfaces as figured

(Fig. 12A-C); well-devcloped spinular row

présent on caudal surface (arrowed in Fig. 12A,

B).

Coxa ot leg I ornamented with long continuous

spinular row across posterior surface (arrowed in

Fig. 13B). Intercoxal sclerite ornamented with

group of spinules on anterior surface (Fig. 13A).

Leg 3 (Fig. I3E) with intercoxal sclerite

ornamented with three spinular rows on

posterior surface (Fig. I3D); distal row of

spinules (arrowed in Fig. 13D) longer than chat

of P. fimbriatus. Leg 4 with inner coxal spine

slightly stouter than that of P. fimbriatus

(Fig. 14A); distal spinular row on posterior

surface of intercoxal sclerite hairdfke and much

longer than that of P fïmbriatus (Fig. 14B); inner

apical spine of endopodal segment 3 as long as

segment, ratio of length of inner apical spine

relative to outer apical spine usually smalier than

in P. ftmbrîatus.

Spine and seta formula as follows:

Coxa

Basis

Exopod

Endopod

Leg 1

0-1

1-1

1-1:1-1:111.5

0-1; 0-1; 1, F 4

Leg 2

0-1

1-0

1-1; 1-1; III, 1, 5

0-1: 0-2: 1, 1, 4

Leg 3

0-1

1-0

1-1; 1-1; III. 1, 5

0-1; 0-2; 1, 1. 4

Leg 4

0-1

1-0

I l ; 1-1 ; II, 1. 5

0-1; 0-2; 1, II. 2

Leg 5 with inner spine serrate-like, usually

stouter than in P. fïmbriatus (Fig. 11C). Leg 6

(Fig. 11 B) represented by one plumose seta and

two tiny spinules located dorsolaterally.

Adult male

Body lengths and widths excluding caudal setae

are given in Table 2. DifFering from adult female

as follows: urosontal somites (Fig. 15C)

ornamented with cuticular pits dorsally as

figured. Caudal rami slightly curved inwards

proximally, usually parallel lor most of length.

Coxobasis ol antenna with spinular row near base

of inner setae (arrowed in Fig. 15D). Innermost

seta of coxobasis more strongly spinulated than in

female (arrowed in Fig. 15D). Antcnnule similar

to that of male P. fïmbriatus cxcept aesrhetnsc on

llrst segment (arrowed in Fig. 16F., F) présent

and setiform. Aesthecasc on ancestral seg¬

ment XIV much longer than in P. fïmbriatus

(arrowed in Fig. 16E). Segmentai fusion pattern

as follows: l-V, VI-YII, VIH. IX, X,XI.XII, XIII,

XIV, XV, XVI, XVI1, XVIII-XX. XXI-XXIII,

XXIV-XXVMI.

Varia liii.iTY

Females

The relative width, length and shape of caudal

rami can varv between specimens from any

population as illustrated in figure 1 IB, D-F. The

arrangement ol the spinular row on the caudal

surface of the antenna (arrowed in Fig. 12A, B)

may also vary slightly. T he length ratio of the

outer apical spine relative to the irincr apical

spine on the distal endopod segment ol leg 4 can

be very variable. The form and length of the

inner spine ofleg 3 can also be variable with some

populations or individuals having a stouter spine

(Fig. I1C) than others.

586

ZOOSYSTEMA • 1998 - 20(4)

Révision of Paracyclops fimbriatus-c omplex

Fig. 15. — R imminuta ; adult 6\ A, urosome, ventral; B, detail of leg 5 and leg 6, ventral; C, urosome, dorsal; D, antenna, coxobasis

and first endopodal segment, caudal. Scale bars; A, C, 100 pm; B. 50 |jm; D, 25 pm.

ZOOSYSTEMA • 1998 *20(4)

587

Karaytug S. & BoxshaJl G. A.

Fig. 16. — P. imminuta-, adult S\ A, body, dorsal; B. antennule showing segmentation, dorsal; C, antennule showing segmentation,

ventral; D, detail of modified seta on tirst segment of the antennule; E, antennule showing setation, anteroventral; F, antennule, first

segment showing setation, anteroventral. Scale bars: A, 400 gm; B, C, 100 gm, D, 25 gm; E, F, 50 gm.

588

ZOOSYSTEMA • 1998 *20(4)

Révision ot Paracyclops fimbriatus-compiex

Table 2. — Body lengtb (BL) and width (BW) measurements (in gm) of adult Paracyclops imminuta from different localities (N, number

of spécimens measured).

LOCALITY

SEX

(mean + SD)

RANGE

(mean ± SD)

RANGE

St Petersburg

846 1 26.2

827-864

31011.4

309-311

England

1014151.7

918-1082

343 1 15.4

305-359

960 1 60.3

872-1079

321 123.6

275-354

Ireland

835 1 32

781-880

318114.2

293-342

805 1 30.6

757-858

292 19.4

282-311

Sweden

10721128.7

981-1163

351 1 23.3

334-367

819

299

Greece

670

247

670

243

Azores (Pico)

791 1 74.2

738-843

298 1 7.1

293-303

Israël

777 182.7

718-835

286 1 1.4

285-287

Gibraltar

910 1 37

849-946

292 1 7.5

280-301

France

921 1143.8

835-1087

315 + 25.4

299-344

DlFFERENTIAL DIAGNOS1S

The close morphological similariry between

P. fimbriatus and P. imminuta makes it easy to

confuse them. howevet, there are two characters

that hâve proven to be extremcly constant both

within and between populations ot P. imminuta.

These characters are the spinular ornamentation

on the caudal surface of the antennary coxobasis

(arrowed in Fig. 12A, B) and the présence of the

mid-distal spinular row on rhe posterior surface

of leg 1 (arrowed in Fig. 13B). The latter

character also distinguisbes P. imminuta trom

other Paracyclops species. In addition, the

following combination of flve characters also

separate.s P imminuta trom P. fimbriatus-. (1) the

spinular row near rhe base of inner setae on the

coxobasis ofantenna which is bener developed in

the male ot P. imminuta (arrowed in Fig. 15D)

than in P fimbriatus (Fig. SE); (2) rhe aesthetasc

on the first segmenr ot male antennule (arrowed

in Fig, 16E, F) which is présent only in

P. imminuta ; (3) the inner coxal spine of leg 4

(Fig. 14A), which is stouter than in P. fimbriatus-,

(4) the form of the distal spinular row on the

posterior surface of the intercoxal sclerite ol

legs 3-4 (Fig. 14A, B), the spinules of which are

hair-like and much longer than in P. fimbriatus-,

(5) the aesthetasc on ancestral segment XIV

(arrowed in Fig. 16F) which is much longer than

that of P. fimbriatus.

Remarks

Kiefer (1929) first separated this form from

P. fimbriatus on the basis of rhe shorter caudal

rami which are about 3-4 tintes longer than

broad. P. fimbriatus forma imminuta sensu

Srantek-Husek ( 1939), Paracyclops fimbriatus var.

imminutus sensu Rylov (1963) and Paracyclops

fimbriatus f. imminutus sensu Stcrba (1935) may

be considered as conspecific with P imminuta on

the basis o( rhe caudal rami which are illustrated

as being close to each other and slightly

converging in the proximal part. The usetulness

of this character is however lirnited sincc it is

known to bc extremcly variable.

Borutzkys (1930) usage ot the caudal rami and

the leg 5 ol male in his identification of

P. fimbriatus. 1. imminuta is very doubtful since

Kiefer did nor refer to male characters in his

original drawings. Perhaps the most complété

description ot P. imminuta was given by Gumey

(1933) under the liante P. fimbriatus. Fortunately,

Gurrtey figured thecoxaoflcg 1 in posterior view

and il can bc seen that the pattern of the spinular

row is very similar to that of P. imminuta

(arrowed in Fig. 13B). Also, rhere is one relatively

large spinulc at the base ot the outer seta of the

antennary coxobasis in the position corres¬

pond ing to the spinular row in P. imminuta

(arrowed in Fig. 12A, B) and probably repre-

senting the end of this spinular row. As remarked

ZOOSYSTEMA • 1998 - 20(4)

589

Karaytug S. & BoxshaJl G. A.

earlier, rhese rwo characrers utiequivocally

separate P. bnmimita from P. fimbriatus and

strongly indicate thar Gurney’s matcria! belonged

to the former species. tn addition, other less

significant characters which serve to separate

P. imminuta from P. fimbriatus can lie observed in

Gurneys drawings, such as the shape of rhe

caudal rami in both sexes, the structure of the

séminal réceptacle and the relatively short inner

spine on the female leg 5 (Fig. 1 IC), ail of which

are more characteristic for P imminuta than

P. fimbriatus.

The validity ol P. fimbriatus orientalis is proble-

matic. This subspecies was incompletely describ-

ed (Alelcseev 1995). Apart from the spinular

ornamentation on the coxobasis of the antenna,

the illustrations of this species lack sufficient

detail. No type material was designated by

Aleksecv (1995) but finir females and three males

collected and identifted as P fimbriatus orientalis

by V. Alekseev were re-examined. None of rhese

specimens matched Alekseev's original drawings

of P. fimbriatus orientalis. In tact the material

represented P. fimbriatus and showed variation in

the shape of the caudal rami and in the spinular

ornamentation on the posterior surface of the

mtercoxal sclerite of leg 4, as also found in

Indian, Mottgolian and Uzbekistani material.

From the original drawings alone it is extremely

difficult to assign P. fimbriatus orientalis to an v of

the species recognised herein. The pattern of

spinulation on the coxobasis of antenna is the

only indication that Alekseev might hâve been

dealing with the species redescribed here as

P. imminuta.

Paracyclops chiltoni (Thomson, 1882)

(Figs 17-23)

Original description:

Cyclops chiltoni Thomson, 1882 [ Tram. Proc. New

Zealand'. Inst. 15: 93-116].

Synonymy:

Cyclops chiltoni Thomson, 1882: 97, pl. IX, figs 11-19.

Paracyclopsfinitimus Kiefer, 1929: 51, Abb. 60, 61.

Paracyclops fimbriatus Fischer, 1853 sensu Karaytug &

Boxshall 1996: 42-66, figs 1-22.

Typk lûCALITY. — New Zealand, Eyreton.

Matp.IUAI KXAMINKD. — h has not been possible to

locale Thomson’-, type material. It is nor stored in the

collection of the National Muséum ol New Zealand.

One vial with forty-seven specimens from New

Zealand is stored in the Smifhsonian Institution,

Washington, 19. G. [ lie redescription is based on this

material.

New Zealand. South Island, Southland, le Anau,

Upakerora River, New Zealand, 45°2TI8S -

167°43'18E, 15.IV.1990, col!. T. Ishida: 33 $9,

8 dd. 6 copepodids (USNM. car. 250692).

Easter Island. Rano Aroi crater (almost completely

overgrown by Pulygonum), small puddle of open water

in littoral zone, 19.IX.1990, col!. Il |. Dumont:

2 9 9,21 dd.

South Afirica. 52 2 9. 44 dd, coll. A. D. Harrison

(NHM 1966.3.22.9).

England. Herüotdshire, percolating filters' ot sewage

works: 50 2 9, 65 dd (NFTM 1968.7.1.1). —

London, Regent’s Parle pond. Numcrous specimens

examined from culrurcd material.

Russia. P Dudergofka. St Petersburg, 27.V1I 1996,

deprh 0.5 m, t 13 °C, coll. V, Alekseev: 4 9 9.

Crozet Island. 28.11.1969. 2 d d dissected on

rwo slides (Ni LM 1970.4.30.3),

Tahiti. 20.1.1981: 2 9 9, 6 dd (MN11N,

unregistered material).

Brazil. I.eaf I itter and small quanttty of moist

superficial soil collected from bank of a rill near to the

emrancc of Gruta da Tapagcm (Tapagem Cave),

24 C “38T2”S 48' J 23'50' , W, Eldorado Paulista, State of

Sâo Paulo, 31.VIII. 1988, coll. Carlos F.. F. Roeha:

18 $ 9. 17 d d obtained from a culture.

Azores. Tetteira, mnunted on three slides (undis-

sected), 2 9, ! d (Th. Monod Collection n° 15210,

MNHN). — Flores, mounted on one slide (undis-

sected), ail Th. Monod Collection: I 9 (n° 15324,

MNHN); 1 9 9, 1 d, mounted on two slides (1 9

dissected) (n a 15292, MNHN); 1 9, parcially dissect¬

ed on one slide (n n 15079, MNHN): l 9, dissected

and mounted on one slide (n n 15087, MNHN); l 9,

mounted on one slide (n° 15138. MNHN); I 9,

dissected and mounted on one slide (n° 15317,

MNHN); I 9. dissected and mounted on one slide

(n° 15335, MNHN); 1 6, mounted on one slide

(n° 15274. MNHN). Pico. 12.X.1971, coll.

Th. Monod: 1 2 , mounted on one slide (Th. Monod

Collection n° 15189, MNHN).

Distribution. — P. chiltoni shows the widest

distribution within the genus Paracyclops and is the only

true cosmopolitan spcctes at présent (Karaytug 1998). It

occurs at extremely isolated sites such as F.astcr Island

and Hawaii in the Pacific Océan, Crozet Island in

Southern Indian Océan, and in New Zealand, However

records of P chiltoni Imm sortie other régions could not

be confirmed because material was unavailable.

590

ZOOSYSTEMA • 1998 - 20(4)

Révision of Paracyclopsfimbriatus-c omplex

Fig. 17. — P. chiltonr, adult 9 ; A, maxillule; B, body, dorsal; C, maxilliped; D. mandible; E, labrum; F, maxilla. Scale bars: A, C-F,

50 pm; B, 200 (jm.

ZOOSYSTEMA • 1998 -20(4)

“591

Karaytug S. & Boxshall G. A.

Fig. 18. — P. chiltoni ; adult 5; A, urosome, dorsal; B, urosome, ventral; C, leg 5, ventral; D, adult S, first segment of antennule

showing setation, anteroventral, with inset showing detail of modified seta, dorsal. Scale bars: A, B, 50 pm; C, D, 25 pm.

592

ZOOSYSTEMA • 1998 *20 (4)

Karaytug S. & BoxshaJI G. A.

Table 3. — Body length (BL) and width (BW) measurements (in pm) of Paracyclops chiltoni from different localities. (N, number of

specimens measured).

LOCALITY

SEX

(mean ± SD)

RANGE

(mean ± SD)

RANGE

New Zealand

627 + 36 1

556-738

251 + 14.9

230-291

581 ± 29.2

531-630

214 ±5.1

207-222

England

713 ±61.2

588-802

289 ±24.6

237-353

(Hertfordshire)

649 ± 52.5

598-751

240 ±7.8

227-254

Easter Island

555

210

618 + 22 9

575-664

225 ± 10.9

200-239

South Africa

722 + 49.5

606-798

262 ± 17.9

223-291

655 ± 25.2

606-699

237 ±8.7

225-252

Russia (St Petersburg) 9

797 ± 68.7

738-857

308 ±9.8

299-316

Tahiti

636 ± 15.6

625-647

232 ±7.1

227-237

642 ± 25.6

614-676

205 ±5.4

196-214

Brazil

611 ± 42.7

567-703

237 ±17.8

217-282

548 ± 25,8

505-588

201 ± 12.1

183-217

Redescription

Adult female

Body width and length are given in Table 3.

Génital double-somite, second and third

abdominal somires densely ornamented with

distinct pits dorsally (Fig. 18A) and ventrally

(Fig. 18B) as figured. Génital double-somite

about as long as broad; séminal réceptacle divided

into narrow anterior and broad posterior lobes

(Fig. 1 SB). Row of spinules présent in anal clefr

either side ol midline (Fig 18A). Caudal rami

short, parailcl, with ornamentation comprising

rows of pits or cuticular dépréssions on ventral

surface (Fig. 18B). Length and width of caudal

rami from different populations given in Table 4.

Antennule 8-scgmented (Fig. 19C): setal

formula 8, 12, 6, 5, 2 + aesthetasc, 2, 2 +•

aesthetasc, 7 + aesthetasc. Third segment with

two partial suture lines ventrally. Coxobasis of

antenna with complex ornamentation on caudal

and frontal surfaces as figured (Fig. 19A, B) and

lacking spinular row near inner setae caudally

(arrowed in Fig. 19B).

Leg I inrercoxal sclerite ornamented with dense

spinules on anterior surface (Fig. 20B). Leg 2 with

outer apical spine of distal endopodal segment

stout (Fig. 20C); inrercoxal sclerite ornamented

wirh dense spinules ou anterior surlace, single row

small spinules on posterior (Fig. 21 A). Leg 3

(Fig. 21D. E) intercoxal sclerite ornamented with

two spinular rows on posterior surface; innef coxal

spine stouter than that ol I! jhnhnatui. Leg 4

(Fig. 2 IB. C) inner coxal spine stouter than in

P. ftmbriaius; intercoxal sclerite with two spinular

Table 4. — Caudal rami length (CL), caudal rami width (CW) and ratio ot length relative to width (L:W) of Paracyclops chiltoni from

different localities (measurements in pm). (N = number of specimens measured).

LOCALITY

SEX

(mean ± SD)

RANGE

(mean ± SD)

RANGE

L:W

New Zealand

79 ± 8.4

68-96

26 ± 1.4

25-30

England

95 ± 12.8

77-116

28 ± 1.6

26-30

3.4

South Africa

94 ± 10.8

78-115

27 ± 1.4

25-29

3.5

Russia (St Petersburg)

97 ± 5.9

88-101

25 ± 3.5

20-28

3.9

Tahiti

68 ± 14.1

58-78

23 ± 1.4

22-24

2.9

Brazil

80 ± 13.5

60-109

25 ± 1.6

23-29

3.2

Azores

94 ± 10

84-109

28 ± 1.5

26-30

3.4

Easter Island

2.5

594

ZOOSYSTEMA • 1998 -20(4)

Révision of Paracyclops fimbriatus -complex

rows on posterior surface (Fig. 21C). The outer

spinulose sera ot leg 5 was equal in length to the

inner spine in rnosi spécimens from New Zealand

and in ail specimens examined from orher

localities (Fig. 22A) but distinaly longer rhan the

inner spine in a lew spécimens lrorn New Zealand

(Fig. 18C).

Adult male

Urosomal somites ornamented with cuticular pits

dorsally and ventrally as figured (Fig. 23B, C).

Caudal rami about 2.3 times longer than broad.

Coxobasis of antenna with wcll-developed spi-

nular row ncar base ol inner setae (arrowed in

Fig. 23E): innermost seta of coxobasis more

strongly spinulate than female. First .segment ol

the antennule (Fig. 18D) with large seta and

modified by ornamentation of strong spinules in

proximal and midsections, tapering to fine point

distally (Fig. 18D). First segment with long

setiform aesthetasc (Fig. 18D). The setal éléments

similar to those of P. imminuta. Segmentai fusion

pattern as follows I-V, VI-VII, VIII. IX, X, XI,

XII, XIII, XIV, XV, XVI, XVII, XVI1I-XX,

XX1-XXII1, XXIV-XXVIII.

VariablLiiy

Unless stated the following variability applies to

the fcmales. The cuticular pits on the ventral

surface of the caudal rarni may not be as well

defined in sorne specimens from outside the

Palaearctic zoogeographical région. The outer

spinulose seta ol leg 5 was equal in length to the

inner spine in most specimens Iront New

Zealand and in ail specimens examined lrorn

other localities (Fig. 22A) but distinctly longer

than the inner spine in a few specimens from

New Zealand (Fig. 18C). The structure of rhe

Fig. 20. — P. chiltoni; adult S; A, intercoxal sclerite and coxa of leg 1, posterior; B, leg 1, anterior; C, leg 2, anterior. Scale bar: 50 pm.

ZOOSYSTEMA • 1998 - 20(4)

595

Karaytug S. & Boxshall G. A.

Fig. 21. — P. chiltonï, adult S; A, intercoxal sclerite and coxa of leg 2, posterior; B, leg 4, anterior; C, intercoxal sclerite and coxa of

leg 4, posterior; D, leg 3, anterior; E, intercoxal sclerite and coxa of leg 3, posterior. Scale bar: 50 pm.

596

ZOOSYSTEMA • 1998 -20(4)

Révision of Paracyclops fimbriatus-c omplex

Fig. 22. — P. chiltonr, A (New Zealand), adult 9. leg 5, ventral; B (U.K.). C (Brazil), adult 9, inner spine of leg 5, ventral; D. adult d,

E, adult ?, anal somite, dorsal (Tahiti); F, adult d ; G, adult 9, leg 4, intercoxal sclerite, posterior (Brazil); H, adult S, anal somite and

caudal rami, dorsal (Brazil); I, adult 9, caudal rami, ventral (St Petersburg). Scale bars: A-C. F, G, 25 pm; D, E, Fl, I, 50 pm.

ZOOSYSTEMA • 1998 - 20(4)

597

Karaytug S. & Boxshall G. A.

inner spine of Icg 5 was slightly different in some

specimensJrom Hngland (Fig. 22B) and front the

State of Sâo Paulo, Brazil (Fig. 22C). In material

from Tahiti, the nurnber of spinules in the anal

cleft, eitlier side of the midlinc (arrovvcd in

Fig. 22D, E), was very small in both sexes. In

specimens from the State of Sào Paulo, Bra/.il

(Fig. 221l) with relative])' longer caudal rami, the

terminal accessory seta (VI) is shorter rhan rhe

postérolatéral seta (III), and the spinular row at

the base of the antérolatéral seta (II) is slightly

curved anteriorly (arrowed in Fig. 22H). In some

specimens from New Zealand, England and

Russia (St Petersburg), the inner margins of the

caudal rami may he irregular, not smooth

(Fig. 221). The distal spinules on the posterior

surface of rhe intercoxal sclerite of leg 4 are

especiallv well-developed in females from the

State of Rio Grande in Brazil (Fig. 22G) and in

males from the State of Sâo Paulo in Brazil

(Fig. 22F).

Differential dlagnosis

P. chiltoni can be differentiated from other

Paracyclops species by the combination of the

following four characters: (1) the structure of

leg 5 (Figs I8G, 22A); (2) rhe presence of the

conspicuous, well-developed spinular row near

the base of two setae on the coxobasis of antenna

in the male (arrowed in Fig. 23E), rhis spinular

row is not présent in the female (arrowed in

Fig. 19B); (3) the obvious, dense, cuticular

ornamentation of pits on the dorsal and ventral

surfaces of the génital double-somite, and second

and third urosomal somites (Fig. 18A); (4) the

presence of similar cuticular dépréssions on the

ventral surface of the caudal rami (Fig. I SB) The

last character may be less pronounced in some

specimens from outside the Palearctic région and

should therefore be used with caution.

P. chiltoni also dîffets from P fimbriatus and

P. imminuta in the much shorter outer seta of

leg 5 (Figs 1SC). In the male of P. chiltoni,

aesthetasc G on the first antennulary segment is

présent (Fig. 18D) but this aesthetasc is absent in

P. fimbriatus.

Remarks

Paracyclops chiltoni (Thomson, 1882) was

originally dcscribed from New Zealand, Later,

Kiefer (1928) described P fhiitimus, which

mate h es well wirh P chiltoni described hcrcin, on

the basis of the very short caudal rami, only three

times as long as broad, and the short outer seta of

leg 5 whtch is about as long as the inner spine.

Kiefer (1931) expressed the opinion ihat

Thomson (1882) likewise had found P jirntimus

and therefore synonymized P chiltoni with

P finitinnts. However, according to the rules of

Zoological Nomenclature, P chiltoni takes

priority and consequently P finitinnts is here

ranked as a junior subjective synonym of P chil¬

toni. Dussart & Dcfaye (1985) had earlier accept-

ed PJinitimus as a synonym of P f. chiltoni.

DISCUSSION

Currently, seven species of Paracyclops occur in

the Palearctic région of which P baicalensis,

P dilatants, P. affinis and P poppci are each quite

distinct m their morphology (Karaytug 1998).

P fimbriatus , P. imminuta and P chiltoni are,

however, verv closely related and the failure to

separate P imminuta from the other two of the

species complet was the main reason behind the

taxonomie confusion concerning ail three. This

taxonomie problem could not hâve been solved

by using tradicional learures such as proportional

measurements of the caudal rami and the

terminal spines of the third endopodal segment

of leg 4, because fhese characters vary consi-

derably within populations as well as between

populations. Since there is extensive overlap in

variability expressed hetween these three variable

species, it is concluded that mosi early records are

unreliable.

The t)'pe material of none of the three species is

estant. In the absence of types it was concluded

that rhe best Option would be to redescribe

P fimbriatus front the vicinity of St Petersburg,

one of rhe listed type localities, on the basis of

newly collected topotypic material. Examination

of three unsorted santples from the vicinity of

St Petersburg rcvealed four distinct Paracyclops

species, P fimbriatus, P poppei, P chiltoni and

P. imminuta. It is interesting to note that the

sample from P. Dudergofka (St Petersburg)

598

ZOOSYSTEMA • 1998 - 20(4)

Révision of Paracyclops fimbriatus-comp\cx

Fig. 23. — P. chiltoni; adult 3; A, detail of leg 5 and leg 6, ventral; B. urosome, ventral; C, urosome, dorsal; D, body. dorsal; E.

antenna, coxobasis, caudal; F, terminal endopodal segment of leg 1. Scale bars: A, 50 pm; B, C, 100 pm; D, E, F, 25 pm.

ZOOSYSTEMA • 1998 • 20 (4)

599

Karaytug S. & Boxshall G. A.

concained four females of P. fimbriatus , one

female of P imminuta , and four females of

/? chiltoni. The sample from the River Igora

(St Petersburg) contained fourteen females of

P fimbriatus and one female of f! imminuta. The

sample from River Ravan (St Petersburg)

contained two females of P fimbriatus and scven

females, three males of P. poppei. A single female

of P fimbriatus from P. Dudergofka (St Peter¬

sburg, Russia) was selected as the neotype.

Dctailed redescriptions of P chiltoni and P poppei

left no doubt that they are distinct specie.s. The

fourth taxon présent in the St Petersburg samples

was ver)' probleinatic. It closely resembled the

rather variable P fimbriatus but coule! be disring-

uished on the basis of rhe arrangement of spinttlar

row on rhe posterior surface of coxa of leg 1 and

the présence of the spinular row near the base of

nvo inner sctac on the coxobasis of antenna.

Historically rhere hâve been some questions

concerning the validity of several of the named

species and subspecies of the Paraeyc/ops fimbriatus

complet. Gurney ( 1933) expressed certain doubts

about rite validity of P. abuobensis Kieler, 1929 and

Pfinitimus Kiefer, 1928. Rylov (1963) agreed with

Gurney's doubts and treated thèse two nominal

species as vjrieties of P fimbriatus. l.indberg

(1958) synonymized P fimbriatus forma imminuta

Kiefer, 1929 with P fimbriatus chiltoni which he

accepted as a valid subspecies. l.indberg (1958)

also synonymized the previously described species

and subspecies on rhe basis of similarities between

old descriptions, apparently without contparing

type material and without giving detailed

redescriptions.

Few workers hâve Considered the possibility that

the underlying reason for rhe gross variability

found in particular populations of Pfimbruttus is

the co-cxistencc of closely related symparric

species. A first indication was given by Frenzel

(1976) who studied a number of populations of

P. fimbriatus from locations in Germany and

Tenerife in which hoth “typical" and “ chiltoni-

type” spécimens co-occurred. Using the “Furcal

index” [Lcngth (L): Width (W) ratio of the

caudal rami in which W is measured halfway L]

he recognized a distinct discontinuity separating

two groups which vvere also distinguished — be it

to a lesser extent — on the basis of the distance

measured proximally between boih caudal rami.

Frenzel (1976) remarked that the P5 was

remarkably constant in the group with short

caudal rami and agreed in form and shape with

Lindberg’s (1958) illustrations of P fimbriatus

chiltoni. Because hoth groups still showed overlap

in several other morphometric parameters,

Frenzel (1976) left the matter undecided and

conduded that only breeding experiments or

more sophtsticated techniques such as

chromosome research could elucidate the

problem. In a later papet Frenzel (1977)

conducted various breeding experiments and

succeeded in proving that al leasr three

reproductive isolâtes (“Kreu/.ungsisolate”)

ocCurred in his samples, of which two could be

identifiai with already described species, i.e.

P. fimbriatus and P. chiltoni. The author also

found that for each of these the morphometry of

the caudal rami and the morphology of the P5

remained largely constant during successive

générations and température did nul seem to

bave any signiPteant inlluence on these characters.

Ail three had the sanie chromosome number in

the female (2n = 13) which is identical to that

fourni in P. djfinis (Braun, 1909). It lias to be

remarked herc rli.tt Frenzel did not stutly any

possible différences in chromosome structure.

Frenzel (1977) conduded rhatalthough rhe three

types were entirely intetsterile, clearcut

distinction on purely morphoJogical grounds was

as yer impossible silice an intermediate form was

also separated during rhe breeding experiments.

Fie recommended further cross-breeding

experiments with other species such as

P abnobertsis , P poppei and P andinus in order to

test rhe distinctiveness of the three German types.

In rhe absence of type material, it was impossible

to assign Fiscliér's (1853) Cyctops fimbriatus to

eichcr P fimbriatus or P imminuta or even to

P chiltoni. Thé two critical characters that best

separate P fimbriatus from the other two species

cannot he disçcrned from Fiscbers original

drawings. These characters are:

1. The spinular ornamentation on rhe frontal

surface of the coxobasis of the antenna of both

sexes.

2. The pattern of spinular ornamentation on the

posterior surface of the coxa of leg 1.

600

ZOOSYSTEMA • 1998 • 20 (4)

Révision of Paracyclopsfimbriatus-c omplex

The shape and length-width ratio of the caudal

ramus of P fimbriatus and P. irmninuta as well as

their geographical distributions overlap to a

certain extern. Unfortunatcly, I ischer’s original

drawings ol caudal rami fall within thaï degree of

overlap. For the sake of nomenclatural stability

the species that has been believed to be tnte

Paracyclops fimbriatus by the majority ol workers

dealing with Palaearctic materi.il, vvhich occlus in

St Petersburg and which could be represented by

Fischer’s figures, has been chosen here for redes¬

cription as typical P. fimbriatus.

The morphoiogy of P fimbriatus from Asia differs

slightly from that of European specitnens. Some

variation is noted in the arrangement and

number of spinular rows on the posterior surface

of the întercoxal sclerite of leg 4 as well as in the

shape of the inner coxal spine (slightly curved) on

leg 4 in some specitnens from the viciniry of Pake

Baikal (Fig. 611, I), Mongolia, Uzbekistan and

India (Fig. 6G, K). Material from India especially

showed rematkable variability on the inner spine

of leg 5 as shown in figure 6A-C. The inner spine

was usually longer than in other material

examined and spinules were présent at the base of

the middle seta in some specitnens (Fig. 6B, C).

It is possible that Asian specitnens of fimbriatus

tend to show a greater degree of variability in

these characters. Examination of more

P. fimbriatus material front Asia, and the

opportunity to carty out breeding experiments

between rhem, wottld provide valuable new

information on the taxonomy of P. fimbriatus in

this part of the Palaearctic.

P. fimbriatus and P. imminuta are, at présent,

distributed ptobably throughout rhe Palaearctic.

Ffowever, the biogeography of P chiltoni is

rematkable because it is the only species of the

genus with very wide distribution (Karayrug

1998). It occurs, for example, in very remote

locations, such as Easter Isl.md, as well as in

Europe. Breeding experiments between geogra-

phically isolat.ed populations would provide new

insights into rhe possible involvement of several

sibling species ol P. chiltoni.

Acknowledgcnicn ts

We would like to thank Prof. Victor Alekseev

(Russian Academy of Science) for collecting

Paracyclops material from St Petersburg, and Dr

Rony Huys (NHM) for translating the Gertnan

literarure and cominenring on the manuscript.

We are also grateful to Dr Danielle Dcfaye

(MNHN), Dr 1. Van de Velde (Royal Belgian

institute of Natural Sciences) and the anonymous

reviewer for their improvements. This research

was funded by Balikesir University, Turkey and

litrkish Highcr Edticational Council, grant to

S, Karaytugs postgraduate studies.

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(Copepoda; Cyclopoid;») front the U.S.A.

Zoo/ogiseber Anseiger 236: ! 09-118.

Rylov V. M. 1963. — Freshwater (ÿclopoida. Fauna of

USSR, Crustacea 3 (3): J-Vl, 1-314 (Translation by

Israël Progt.tm Scienr.Trans. ofV. M. Rylov, 1948],

Sars C. O. 1863. - Oversigt af de indc-nlandske

FerslcvandscopCpoder. Forhandlinger i Videntkabs-

selskabet i Kristiania (Jahr 1862): 212-262.

1913-1918. — An account of Crustacea of Norway.

6. Copepoda Cyclopoida. Bergen. 225 p.

Sthmeif O. 1892. — LIeutschlands freilebende

Siisswasser-Copepoden. 1. Teil: Cyclopidae.

Bibliothcca Zoologica 11: 1-191.

•Slamek-Husek R. 1939. — F.ine Wiederentdeckung

von Paracyclops fimbriarus f. imminuta Kiefer in

Bd h m en. Vestnik Ceskasltwenske Spolecnosti

Zoologicke. Praha 6. 7: 427-430.

Stcibu Ô. 1955. — Pfispèvck k poznânt buchanek

(Cyclopoida) i. Brncnska a jiztu Moravy. Beitrag zur

Kenntnis der Cyclopidcnfauna der L'mgchung von

Brünn und aus Süumâhren. Vestnik Ceskoslovenske

Spolecnosti Zoologicke. Praha 19: 354-364 [English

summaries].

T homson G. M. 1882. — On the New-Zealand

Copepoda, Transactions and Proceedings of the New

Zealand Itistituie 15: 93-116.

Uljanin V. N. 1875, — Rakoobraznyya, in Ecdchenko

A. P.(çd-), Puleshestvie v Turkestan, Izvestiya

imperatorskago Obscbchcstiva Lyubitelei Fsresivozna-

niya, Antropologii 1 Ftnografu pri imperatorskom

Moskovskom Universitete 11 (6): 1-63.

Van de Velde 1. 1984. — Révision of the African

species of the genus Mesocyclops Sars, 1914 (Cope¬

poda: Cyclopidae). Hydrobiologta 109 (1): 3-66.

Submitted on 4 March 1998;

accepted on 8 June 1998.

602

ZOOSYSTEMA • 1998 - 20(4)

A second species

of the genus Balssia Kemp, 1922

(Crustacea, Decapoda, Pontoniinae)

Alexander J. BRUCE

Queensland Muséum, P.O. Box 3300, South Brisbane,

Queensland, 4101 (Australia)

abruce @ 3D.com .au

Bruce A. J. 1998. — A second species of the genus Balssia Kemp, 1922 (Crustacea,

Decapoda, Pontoniinae). Zoosystema 20 (4): 603-611.

KEYWORDS

Balssia noeli,

Crustacea,

Decapoda,

Pontoniinae,

new species,

Mediterranean Sea,

taxonomy,

commensal ism.

ABSTRACT

A new species of commensal pontoniine shrimp, Balssia noeli n.sp., from the

vicinity of Banyuls, western Mediterranean, is described and illustrated. The

new species is readily distinguished from B. gasti (Balss, 1921), the only

other species of the genus, by irs lack of strong sculpruring on the carapace

and abdomen. The species is associated with the gorgonians Corallium,

Eunicella, Gerardia and Paramuricella.

MOTS CLÉS

Balssia noeli,

Crustacea,

Decapoda,

Pontoniinae,

espèce nouvelle,

Méditerranée,

systématique,

commensalisme.

RÉSUMÉ

Une deuxième espèce du genre Balssia Kemp, 1922 (Crustacea, Decapoda,

Pontoniinae). Une nouvelle espèce de crevette pontoniine, Balssia noeli n.sp.,

provenant de la région de Banyuls, Méditerranée occidentale, est décrite et

illustrée. La nouvelle espèce se distingue facilement de B. gasti, la seule autre

espèce du genre, par la sculpture beaucoup moins accentuée de la carapace et

de l’abdomen. L’espèce est associée aux gorgones Corallium, Eunicella,

Gerardia et Paramuricella.

ZOOSYSTEMA • 1998 • 20(4)

603

Bruce A. J.

INTRODUCTION

A single specimen of Amphipalaemon gasti, a

small shrimp associated with the precious coral,

Corallium rubrum (Linnaeus, 1758) from the

Gulf ol Naples, was first described and illustratcd

by Heinrich Balss in 1921. Kemp (1922) reco-

gnized iliat the species was a pontoniine shrimp

and did not belong in the gémis Amphipalaemon

Nobili, 1901, now considered a synonym of the

genus Anchistioides Paulson. 1875 in the family

Anchistioididae. The nevv genus Balssia was then

designated lor its réception, The species has since

been reporred from a wide range ol localities on

the European coasts of rhe western Méditer-

ranean and Adriatic Seas, Corsica, rhe Canary

Islands and sonie West African localities, and

from a variety of host animais (Manconi tic Mori

1990). The dtscovery of a second species in the

western Medirerranean suggests that some of rhe

earlier records should be re-examined ro confirm

rhat they do ail refer ro Balssia gasti (Balss,

1921). The specimens are deposited in the col¬

lection of the Muséum national d'Histoire natu¬

relle, Paris (MNHN), Nationaal Natuur-

historisch Muséum, Leiden (NNM), and

Queensland Muséum, Brisbane (QM).

I am most grateful to Dr Pierre Noël for the

opportunity to report upon these specimens.

Carapace length (CL) refers to the post orbital

carapace length.

SYSTEMATICS

Family Palaemonidae Rafinesque, 1815

Subfamily PONTONIINAE Kingsley, 1878

Genus Balssia Kemp, 1922

Balssia noeli n.sp.

(Figs 1-4)

Balssia sp. - Noël 1992:64.

MATERJAL EXAMINER. — Mediterrancan Sea.

Vicinity of Banyuls: 7 ovigerous $ $, 1 ovigerous $,

1 ovigerous 9 (Laboratoire Arago).

From “ancîent collections”, details of exact localities,

dates, depths and collectors noc available.

TYPES. — The ovigerous female holotype is deposited

in the collection of the Muséum national d’Histoire

naturelle, Paris, catalogue number MNHN-Na

13440, together with the dissected paratype female,

MNHN-Na 13441. A paratype is also deposited in

the collection of the N.uiona.iJ Natuurhistotisch

Muséum, Leiden, catalogue number RMN H D

4^887, and one in the Queensland Muséum,

Brisbane, number QM W 23107. The remaining spe¬

cimens arc also designated as paratypes.

MessüREMENTS. — Ovigerous female holotype, cara¬

pace length 4.0 mm: carapace and rostrum 6.0 mm;

total body length (appio*.) 15.5 mm; second pereio-

pods, major chela 1.6 mm; minor chela 1.6 mm; leng¬

th nfovum 0.5 mm.

COLORATION AND HOSTS. The specimens, ail front

gorgonacean coelenterates, were annotared as follows:

from lot #1, (1) white, on Eunieella Verrill, 1869;

(2) violer, on ParamuriceUa Kôlliker, 1865; (3) yellow,

on Gerardia Lacaze-Dultriers, 1864 (zoantharian of

the family Parazoanthidae); (4) red, on Corallium

Cuvier, 1826. No data available on colouration or

hosts of lots #2-#3.

EtyMOLOiJY. — Fhe species is named in honour of

Dr. Pierre Y. Noël, who Icindly provided the spéci¬

mens upon which this report is based.

Description (ovigerous females, Fig. 1)

Small stoutly builr pontoniine shrimps of gene-

rally subcylindrical body form (Fig. 1), cephalo-

thoracic région ventrally flattened.

Rostrum (Fig. 2A, C)

Slender, oval in section, without distinct carinae,

about 0.5 of carapace length, reaching to end of

intermédiare segment or middle of distal seg¬

ment of antennular peduncle distally acute, with

three small acute teeth dorsally, proximal reeth

large, acute, distal tooth small, ventrally unar-

med, few médian setae proximally only.

Carapace (Fig. 2A)

Swollen, globular, smooth, cervical groove feebly

indicated, small cervical and hepatic tubercles

usually discernible, with distinct postrostral cari-

na bearing rwo small teeth, posterior touth usual¬

ly obtuse, antertor tooth acute. small epigastric

tooth présent or absent; orbïc well-devcloped,

superior m.trgm carinate with three small acute

teeth, posterior margin mconiplete, latéral border

carinate with two subacute tubercles posteriorly,

604

ZOOSYSTEMA • 1998 • 20(4)

A second species of Balssia (Pontoniinae)

Fig. 1. — Balssia noeli n.sp., ovigerous holotype 9 f Banyuls. Scale bar: 2.0 mm.

continuons with large acute dorsolaterally angled

antennal spine, branchio-stegal sulcus présent

belovv latéral orbital margin, antérolatéral margin

of branchiostergite slighrly produced, rounded,

inferior orbital angle broadly triangular.

Abdominal segments (Fig. 2E. 1, 4K)

Smooth, thicd segment nor postetiorly produced,

non-carinate, first tbree pleura expanded, broad¬

ly rounded, lourth vvitb posterior margin feebly

bilobed, centrally fcehly carinate laterally, fifth

similar with uppcr lobe acutely produced, sixrh

segment 1.6 limes length of fifth, 2.0 times lon¬

ger than central depth, postérolatéral angle stron-

gly produced, acute, posteroventral angle

produced, subacute.

Telson (Fig. 21)

About 1.8 times length ol sixrh segment, about

3.0 times longer rhan anterior width, latéral mar-

gins straight, postetiorly convergent, with small

subequal marginal latéral spines at 0.5 and 0.9 ol

telson length, posterior margin (Fig. 4K) 0.3 of

anterior width, transversely truncate, without

médian point, latéral posterior spines similar to

dorsal spines, intermediate spines about 0.75 of

telson length, about 7.5 times longer than basal

width, submedian spines slender, setulose, shor-

ter than intermediate spines.

Ophrhalmic somite dorsally antero-posteriorly

bilobed, without pigment spot.

Antennular peduncle (Fig. 2G)

Normal, with short flagella; proximal segment

1.2 times longer than broad, latéral margin

expanded, angular, produced distally with small

rounded lobe reaching to middle of intermediate

segmenr length, with large acute slender distola-

teral tooth, exceeding intermediate segment

length, ventromedial margin with strong tooth at

0.5 of length, intermediate and distal segments

normal, subequal, combined length about 0.7 of

proximal segment length, intermediate segment

with small setose latéral lobe, upper flagellum

biramous with etght proximal segments fiised,

shorter free ramus with fout segments, with

about eighteen groups of aesthetascs, lower

ramus short, filiform.

ZOOSYSTEMA • 1998 • 20(4)

605

Bruce A. J.

Antenna (Fig. 2H)

With basicerite stout, with small blunt antéro¬

latéral tooth; carpocerire short stout, rcaching to

about half scaphocerite length, flattcncd, about

2.0 times longer than wide, flagellum short,

about 2.0 limes carapace length; scaphocerite

broad, about 1.6 times longer than central

width, latéral margin feebly convex, with small

acute distal tooth, at about 0.8 of lamellar length,

falling far short of anterior margin of lamella.

Eye (Fig. 2F)

Stout, with well-pigmented globular cornea, dia-

meter about 0.18 ol carapace length, without

accessory pigment spot, stalle length subequal to

width and corneal diameter, with minute antero-

dorsal tubercle.

Mouth parts

Mandible (Fig. 3) slender, without palp; incisor

process (Fig. 3C) slender, tapering, with four

small acute teeth distally, central pair smaller than

outer teeth; molar process (Fig, 3B) slender, obli-

quely truncate distally, with three small stout,

blunt teeth and two bands of setae, Maxillula

(Fig. 3D) with feebly bilobed palp, lower lobe

with single short seta; upper laçinia short, subeir-

cular, with several short simple spines distally and

numerous finely serrulate setae; lower lobe short

and broad, with numerous spiniform setae distal¬

ly and ventrally, Maxilla (Fig, 3E) wirh short

broad, non-setose palp, media) margin sinuous,

distal endite feebly bilobed, lobes subequal, upper

lobe wirh about ftfteen slender simple setae, lower

lobe with about ten. proximal endite obsolète,

médial margin feebly convex; scaphognathite

broad, about 3.4 times longer than central width.

First maxilliped (Fig 3F) with short, non-setose,

tapering palp (Fig. 3G), about 2.3 times longer

than proximal width. basal and coxal endites

completely fused, broad, about 1.9 times longer

than wide, broadly rounded distally, médial mar¬

gin distally sttaight, proximally concave, fringes

with numerous slender, feebly serrulate setae; exo-

pod w'ith flagellum greatly reduced, scarcely

exceeding caridean lobe, subcylindrical, with two

short plumose setae distally, caridean lobe large,

broadly expanded, 2.0 times longer than broad;

epipod large, triangular, feebly bilobed. Second

maxilliped (Fig. 3H) with endopod of normal

form, dactylar Segment narrow, médial margin

with numerous long serrulate spines, propodal

segment not antero-medially produced, ischiome-

rux and basis normal, with exopod greatly redu¬

ced, not exceeding basis, lamellar, non-setose;

coxa stout, without médial process, epipod well-

developed, triquetral, without podobranch. Third

maxilliped (Fig. 31) rcaching to about middle of

carpocerite length, ischiomerus and basis comple¬

tely fused, jonction indicat.çd by small médial

margin notch, about 3.0 rimes longer than proxi¬

mal width, tapering distally, sparsely setose alone

distal médial margin, penultimate segment about

0.5 of ischiomerus-basal segment, 2.3 times lon¬

ger than central width, uniform, with numerous

serrulate spiniform setae medially, distal segment

about 0.4 of ischiumeral-basal segment length,

2.4 times longer than proximal width, densely

spinulose medially and distally, with serrulate

spines; exopod as in second maxilliped; coxa with

small non-setose médial process, broad rounded

latéral plate and rudimentary arthrobranch pré¬

sent.

Thoracic stermtes (Fig. 2D)

Broad, first with small transverse, feebly bilobed

canna, third widest (so chat third maxilltpeds are

latéral to first pereiopods in ventral view), fourth

without médian process, with rounded latéral

lobes, posterior sternites broad, without spécial

features.

First pereiopods (Fig. 4A, B)

Slender, exceeding carpocerite by chela and car-

pus, chela (Fig. 4B) slender, palm subcylindrical,

tapering slighdy distally, about 3.6 times longer

than wide. fingets about 0.66 of palm length,

slender, with simple acute tips, cutting edges

entire, latéral, carpus about 1.25 times longer

than chela, 6,5 times longer than distal width,

subcylindrical, tapering siightly proximally;

merus about 0-9 of carpus length, of similar

width, ischium about 0.9 of chela length, basis

and coxa normal, coxa with small setose ventro-

medial process.

Second pereiopods (Fig. 4C-G)

Feebly developed, subequal, similar; exceeding

608

ZOOSYSTEMA • 1998 • 20(4)

Bruce A. J.

carpocerite by length of fingers; chela (Fig. 4D,

E) about 0.4 of carapace length, palm smooth,

oval in section, about 1.8 limes longer than distal

depth, fingers (Fig. 4F) subequal to palm length,

sparsely setose, gaping slightly proximally, dacty-

lus slender, curved, slightly compressed, tip

(Fig. 4G) with stour acute articulated tooth with

small, non-arriculaced toorh laterally. curring edge

latéral, entire, fixed finger similar but expanded

proximally, cutting edge présent distaUy, entire;

carpus short, stout, unarmed, 2.0 times longer

than distal width, tapered proximally, subequal to

palm length; merus 1.2 times palm length, about

3.0 times longer than wide. uniform, unarmed;

ischium subequal to palm length, unarmed, basis

and coxa normal, without spécial features.

AmbuLuary pereiopods (4H-J)

Robust, third (Fig. 4H) exceeding carpocerite by

length of dactylus, dactylus (Fig. 4J) about 0.45

of propod length, stout, simple, acute, strongly

curved, with feeblv demarcated unguis, about

0.5 of corpus length, 3.0 rimes longer than basal

width, corpus compressed, dorsal margin convex,

2.0 times basal deprh, with paired sensory setae

distaUy, sparse setae along dorsal margin; propod

(Fig. 41) about 0.4 of carapace length, 6.0 rimes

longer than depth, uniform, slightly bowed,

without spines, sparse setae distaUy, carpus about

0.3 of propod length, 1.6 times longer than dis¬

tal width, tapering proximally, unarmed, merus

0.65 of propod length, 2.6 times longer than

deep, unarmed, ischium 0.5 of propod length,

unarmed, basis and coxa normal, without spécial

features, coxa stout. Fourth and fifth pereiopods

similar, loutth with propod 0.8 of third propod

length, fifth subequal to third.

Uropod (Fig. 2J, 4L)

With protopodite with small blunt postérolatéral

lobe; exopod broad, about 2.1 times longer than

wide, latéral margin convex, unarmed, with small

acute distal tooth (Fig. 4L) with larger mobile

spinule medially; endopod reaching to distal end

of exopod, about 2.8 times longer than wide.

Ova very numerous, small.

Systematic: position

A well-illustrated account of Balssia gasti was

provided by Balss (1921), based on a single ovi-

gerous fernale specimen, and is therefore fully

comparable with the présent material.

Manconi & Mori (1990) State that, in B. gasti

the carapace length ranges from 3,2 to 4.0 mm

in ovigerous femalcs so that the rwo species are of

similar size.

The major characters that distinguish Balssia

noeli from B. gasti (Balss), the only other species

of the genus, are as follows; shorter rostrum, not

well exceeding antennular peduncle, with three

smailer dorsal teeth only; a short bidentate post-

rostral carina, with or without a small epigastric

tooth; cervical and hepatic spines reduced to

small rubercîes; first three abdominal terga

without médian carinae; fourth pleuron lacking

acute rooth posteriorly; anterodorsal eyestalk

tubercle minute; antérolatéral tooth of proximal

segment of antennular peduncle not well excee¬

ding intermédiare segment; maxilla with distal

endite feebly bilobed; fixed finger of second per-

eiopod with proximal half only expanded.

Remarks

In the original description of Balssia gasti Balss

noted particularly the strong sculpturing, of the

carapace, and abdomen, with a distinct dorsal

carina the absence of which most clearly séparâtes

his species from B. noeli. It is possible that the as

yec unknown males of B. noeli will show a more

strongly sculptured appearance than the females,

as is known to occur in some other ornate ponto-

niine associâtes of coelenterates such as Dasycaris

zanzibarica (Bruce, 1973) The différences are

not strongly marked in B. gasti us is shown by the

figures in Zariquiey Alvarez (1946, fig. 111).

Balssia gasti was first found in association with the

precious coral Corallium tubrum (L., 1758), an

alcyonacean host. Most subséquent records hâve

been from this host. There hâve also been a num-

ber of records from other hosts, reviewed by

Manconi & Mort (1990), including several from

gorgonians. including species of F.unicella,

Gerardia, Lopbogorgia Milne Edwards, 1857, and

ParamuriceUa. [’ossibly somc of these should be

referted to B. noeli. Mori et al. (1994) hâve confir-

med that their material, from Sardinia and asso-

ciated with the gorgonian host Paramuricella

clavata (Risso, 1826), belonged to Balss’ species

610

ZOOSYSTEMA • 1998 • 20(4)

A second species of Balssia (Pontoniinae)

sensu stricto, so rhcrc is no doubt that B. gasti can

occur vvith both alcyonacean and gorgonian hosts.

The associations reported with sponge hosts

(Ledoyer 1968) secm likely to bave been of acci¬

dentai origin due to the cxigcndes of collection

ntethods. Hovvever, the possibility ol associations

with sponges should be furthcr investigated before

its exclusion. The sélection of host animais of

B. noeli otherwise overlaps significandy with that

of B. gasti , but further collections may show that

each species has distinctive host préférences.

The spedmen illuscrated by Noël (1985) shows a

marked absence of strong sculpturing on the cara¬

pace and may therefore belong to the présent spe¬

cies. The live specimen was collected from

Banyuls in the late 1960’s or early 1970’s by Alain

Thiriot and photographed by Jean Lecomte. It

was a pure yellow colour, associated with a gorgo¬

nian host, and may hâve formed part of the mate-

rial upon which the présent study is based (P. Y.

Noël, pers. comm., January 1998).

The status of the various colour patterns and

host animais remains mysterious. It is quite pos¬

sible that colour pattern and host sélection are

genetically determined and thàt the various

colour “forms" of each “species” actually repre-

sent a compiex of related sibling species. Shrimp

coloration may be under physiological control.

There is ample scope for further study in these

fields. Knowlton & Relier (1985), and Duffy

(1996) hâve demonstrated that these complexes

exist in some commensal Alpheidae and their

occurrence in other caridean lantilies seems highly

likely. Some of the monospecific commensal

shrimp généra that are found in association with

a wide variety of different host types may well

consist of several sibling species. Knowlton

(1993) and Dufly (1996) consider that diversity

in the relevant taxa is probably several times

greater than presently recognized and that this is

correlated with a high degree ofhost specificity.

Acknowledgements

This study was facilitated by the support of the

Australian Biological Resources Survey.

REFERENCES

Balss H. 1921. — Ueber eine neue Pontoniide aus

dem Golf von Neapel. Mitteilungen aus der

Zoologiscben Station zu Neapel 22 ( 15): 523-526.

Bruce A. J. 1973- — Notes on some Indo-Pacific

Pontoniinae, XXIV Dasycaris zanzibarica n.sp.

from the western Indian Ôcean. with remarks on

other species of rhe genus Dasycaris Kemp. 1922

(Decapoda Narantia). Crustaceana 24 (3): 247-260.

Duffy J. E. 1996. — Species boundarics, spécialisa¬

tion, and the radiation of sponge-dwelling alpheid

shrintp. Biological Journal oftbe Linnean Society 58:

307-324.

Kemp S. 1922. —Noies on Crustacea Decapoda in

the Indian Muséum. XV. Pontoniinae. Records of

the Indian Muséum 24: 113-288.

Knmvlton N. 1993. — Sibling species in the sea.

Anrtual Review of Ecology and Systematics 24:

189-216.

Knowlton N. & Keller B. D. 1985. — Two more

sibling species of alpheid shrimps associated with

the Caribbean sea anémones Bartholomea annulata

and Heteractis lucida. Bulletin of Marine Science 37

(3): 893-904.

Ledoyer M. 1968. — Ecologie de la faune vagîle des

biotopes méditerranéens accessibles en scaphandre

autonome. (1) Introduction. Données analytiques

sur les biotopes de substrat dur. Recueil des Travaux

de la Station Marine d'JEndaume 56 (40) : 103-149.

Manconi R. & Mori M. 1990. — New records of

Balssia gasti (Balss, 1921) (Decapoda, Palaemo-

nidae) in the western Mediterranean Sea. Crustace-

ana 59 ( 1): 96-99.

Mori M., Mot! C. & Bianchi C. N. 1994. — Notes

on the life history of the Pontoniine shrimp Balssia

gasti (Balss, 1921). Oebalia 20: 129-137.

Noël P. Y. 1985. — Lhomochrornie et l’homorypie

chez les Crustacés. Bulletin de la Société entomolo-

giifue de France 90 : 1004-1015.

— 1992. — Clé préliminaire d identification des

Crustacea Decapoda de France et des principales

autres espèces d'Europe. Collections patrimoines

naturels 9, Secrétariat de la Faune et de la Flore,

Muséum national d’Histoire naturelle, Paris :

1-145.

Zariquiey Alvarez R. 1946. — Crustâceos Decâpodos

Mcditerraneos. Instituto Espanol de Estudios

Mediterrdneos 2: 1-181.

Submitted on 6Augustl997;

accepted on 29 January 1998.

ZOOSYSTEMA • 1998 • 20(4)

611

Evolution of the abdominal holding Systems

of brachyuran crabs (Crustacea, Decapoda,

Brachyura)

Danièle GUINOT & Jean-Marie BOUCHARD

Laboratoire de Zoologie (Arthropodes), Muséum national d’Histoire naturelle,

61 rue de Buffon, F-75231 Paris cedex 05 (France)

guinot@mnhn.Fr

bouchard@mnhn.Fr

Guinot D. & Bouchard J.-M. 1998. — Evolution of the abdominal holding Systems of bra¬

chyuran crabs (Crustacea, Decapoda, Brachyura). Zoosystema 20 (4): 613-694.

KEYWORDS

Abdomen,

Brachyura,

coaptation,

holding System,

homology,

locking mechanism,

microstructurc,

press-button,

phylogeny,

uropod.

ABSTRACT

Brachyuran decapod crustaceans are characterized by a reduced abdomen,

folded beneath the céphalothorax, and inserted between the pereiopods or in

a spécial cavity, which prevents it Irom impedîng movements. 1 he acquisition

of a retaining/locking mechanism may be regarded as a synapomorphy for the

Brachyura. In primitive podotrematous forrns (Dromiidae, Homolidae), the

structures involve the coxae c>f thoracopods. In liigher Brachyura, the struc¬

tures located on limbs hâve moved onto the sternal plate. The most common

holding sysreni is the press-button, consisring of a prominence always on the

fifth sternitc and a Socket always positioned on the sixth abdominal segment.

The innovation with a sternal différentiation appears to bave occurred inde-

pendently several times: on sternite 4 (Homoloidea: bomolid press-button),

on sternite 5 (Dynomenidae, Lyreidinae), on sternite 6 (Phyllotymolinidae,

Cyclodorippidae), and on sternite 5 (F.ubrachyura: typical press-button). In

the Heterotremata, only the Leucosiidae display a retaining System rhat dit-

fers from the typical press-button. A true Socket evolved within the

Podotremaia (Homoloidea, Lyreidinae), and in almost ail Eubrachyura. The

abdominal Socket is hypothesized to be homologous with the uropod. The

absence of any locking structures in the most highly carcinized thoracotre-

niatous crabs (for instance Oeypodinae pro parte) is considered to be the

resuit of a secondary loss, occurring at about the rime when terrestrial habi¬

tats replace the exclusively marine lite. The microstructures, discovered on

borh homolid and typical press-buttons, rnight be regarded as an indicator of

sex, puberty moult and âge of individuals, and are probably rcliable indica-

tors of phylogenetic affinities at different levcls.

ZOOSYSTEMA • 1998 • 20 (4)

613

Guinot D. & Bouchard J.-M.

MOTS CLÉS

Abdomen,

bouton-pression,

Brachytira.

coaptation,

homologie,

mécanisme d’accrochage,

microstructure,

phylogénie,

uropode.

RÉSUMÉ

Evolution de< systèmes de maintien de l'abdomen chez les Brachyoures (Crustacea ,

Decapoda, Brachyura). Les Crustacés l fëcapodes Brachyoures sonr caractérisés

par un abdomen réduit, replié sous le céphalothorax et inséré entre les péréio-

podes ou dans une cavité particulière, ce qui empêche toute gêne lors des

déplacements. L'acquisition d'un système de maintien ou de Fermeture de

l'abdomen est considérée ici comme une synapomorphie des Brachyoures.

Chez les formes primitives de l’odotremata (Promiidae, Hotnolidae) les

structures Font intervenir la coxa des thoracopodes. Chez le.s Brachyoures plus

avancés les structures se trouvent déplacées sur le sternum thoracique. Le sys¬

tème le plus répandu est le bouton-pression, à savoir une proéminence appar¬

tenant toujours au sternite thoracique 5, et une fossette toujours située sur le

segment abdominal 6. La novarion que constitue la différenciation sternale

serait apparue indépendamment plusieurs Fois : sur le sternite 4

(Hornoloidea : bouton-pression homolien), sur le sternite 5 (Dynomenidae,

Lyreidinae), sur le sternite 6 (Phyllotymolinidae, Cyclodorippidae), et sur le

sternite 5 (Eubrachyura : bouton-pression rypique)- Chez le.s Hetcrotremata,

seuls le.s Leuco.siidac offrent un système de maintien diffèrent du bouton-

pression typique. Une vraie fossette est apparue au sein des Podorremata chez

les Hornoloidea et les Lyreidinae, ainsi que pour l’ensemble des Eubrachyura.

La fossette abdominale est ici homologuée à l'uropode. L’absence de struc¬

tures chez les Thoracorremata les plus évolués (Ocypodinae pro parte par

exemple) est considérée comme le résultat d'une perte, lors du passage d'un

habitat exclusivement marin à un mode de vie amphibie. Les microstructures

decouvertes sur le bouton homolien et sur le bouton-pression typique repré¬

sentent des indicateurs du sexe, de la mue de puberté et de l’âge des indivi¬

dus ; de plus, elles sont susceptibles de refléter les affinités phylogénétiques à

divers niveaux.

INTRODUCTION

The term coaptation (front the Latin cum, with;

aptare , fit), that lor a long tinte referred to medical

language for the setting of a fracture or luxation,

was revived by the Frencb biologist L. Cuénot

(1921; 1925) ro name, in arthropods and more

especially in insects, the fitting berween rvvo diffe¬

rent and originally independent parts of the bodv,

with a more or less pronounced engagement.

Many types of coaptation are dcveloped in alrnost

ail brachyuran crabs ro retain their abdomen ben-

eath the céphalothorax. The niost wcll-known is

the bouton-pression or press-button (Cucnot 1941 ).

In the tollovving scheme used in insects (Corset

1931), threc main patterns, with sevcral degrees of

différentiation, may be recognized in the

Brachyura, at the binocular level (Table 1, p. 634):

1. Coaptation by juxtaposition. There is a relative

independence of both parts, a simple modalicy of

union, or a simple joining: the two parts are only

coupled, In rhe case when a spine or a projection

simply overhangs or overlaps the other part,

without inducing marked modifications, it may

be considered the first State of a coaptation.

2. Coaptation by engagement (in French engre¬

nage). Considered a more advanced degree: both

corresponding parts are more differentiated but

can be easily separated. A variety of configura¬

tions, front simple to more complex, exists.

3. Coaptation by assemblage (in French assem¬

blage). One of the two parts, which were origi¬

nally completely independent, is firmly engaged

in a Socket or a hollow, wtthin which it is held

614

ZOOSYSTEMA ■ 1998 -20(4)

Abdominal holding Systems (Brachyura)

fast most of rhe time. The two parts can only be

separated witli difliculty and displaced in a given

direction, often with a clicking sound. This is the

case of rhc typical boulon-pression and homolid

press-button,

In Brachyura the reducrion-folding ot the abdo*

men is probably the principal event in the process

of carcinization, i.e. the adoption of a crab-like

habitus. In contrast to long-bodied forms of

decapods in which the developed and extended

abdomen, with movable somites and large bira-

rnous pleopods, is used for locomotion, brachyu-

ran crabs are characrerized by a short abdomen

that is basically formed by six segments plus the

telson and lies in a flexed position. Brachyura

lack locomotory appendages and rail fan, as they

exist in “macruran” decapods lor exarnple: die

locomotory function of the abdomen hax been

lost. Either the brachvuran abdomen is sintply

folded forward between the pereiopods, or it may

be accomiuodaied by a thoracic dépréssion, the

sterno-abdominal cavity, which i.s more or less

hollowed and defined. Besides that, the pleon is

secured in a flexed position and sometimes tight-

ly held in place against the ventral surface by a

spécial System.

In most primitive crabs the abdomen is simply

held by rhc bases of varions thoracopods (Mxp2,

Mxp3, PI through P3), and coaptations by jux¬

taposition and engagement are developed. Beat, in

most crabs, the most coin mon means consists of

a “truc” locking mcchanism, the press-button.

The press-button System is a coaptation by

assemblage between structures that bclong to rwo

clearly defined somites ol two different parts of

the body: the fifth somite on the thoracic ster¬

num and rhe sixrh segment on rhe abdomen (or

pleomere 6). It consists of a pair of salient struc¬

tures on the sternal plate thar fits into a pair of

sockets on abdominal segment 6.

It is beyond the scope of this paper to reconstruct

phylogencric rclationships within the Brachyura

based on the abdominal holding apparatus.

However, analysis of rhe different dcvices or stra¬

tegies utilized by brachyuran crabs to retain or

lock their abdomen sheds new light on the évo¬

lution of the group. In tnrn, using this criterion

at different taxonomie levcls might perhaps open

a way for the reappraisal of the Brachyura.

MATERIALS AND MF.THODS

Our study is based on the large research collec¬

tions of the Muséum national d’Histoire naturelle,

which are essential for a sound, pragmatic basis

for compréhensive analysis. The observations

always involved dead specimens (alcohol preser-

ved), that were often manipulated. That is why

ail our suggestions on the functioning must be re-

considered with fresh specimens and living ani¬

mais. In rhe course of this work, thousands of

specimens were investigated, but it is clear that

our conclusions need to be completed and hypo¬

thèses are presented to serve as a basis for further

investigations and discussion.

Représentatives of several généra, belonging to

fifty-seven families available to us, bave been stu-

died and illustrated. Some results are taken from

the work in ptogress by one of us (J.-M B.) on

microstt uctures of the locking System with the

scanning électron microscope.

We follow the classification of Guinot (1977,

1978a, cf. also Guinot & Richer de Forges 1997).

The families, listed here in a certain order, do not

represent a classification, even if a phylogenetic

background is présent. The list presented here is

noc exhaustive. Regarding the components of

some families, the traditional arrangement was

sometimes conveniently preterred; the lamily

level was often preferred, because monophyly is

betrer supported (but not in Varuninae for

example). In any case we examinée! at least the

type genus, and we atlempied to extend our

observations to ail principal component généra.

Our description essentially concerns the males

sincc, gcncrally, structures are lost in females after

the puberty moult.

Carapace dimensions are given as carapace length

followed by carapace widih, the measures inclu-

ding any anrero-lateral teeth at rhe widest point,

cw, carapace width; cl, carapace length. Appen-

dages and segments frequently referred to in des¬

criptive accounts and figures are usually abbrevia-

ted as follows: P1-P5, pereiopods (Pl, chelipeds,

P2-P5, walking legs); Mxpl-Mxp3, maxillipeds;

Gl, First male pleopod; G2, second male pleo-

pod. The rerm rhoracopod is used for the appen¬

dages Mxpl-Mxp3 and P1-P5. We preterred

the terminology somite 4 to fourth somite or

ZOOSYSTEMA • 1998 -20(4)

615

Guinot D. & Bouchard J.-M.

abdominal segment 4 to fourth abdominal seg¬

ment, for example.

Abbreviations for the institutions are as follows:

BMNH The Natural History Muséum, London;

MNHN Muséum national d'Histoire naturelle, Paris

(MNHN-B used for catalogue number; B

means Brachyura);

SAM South African Muséum, CapcTown;

USNM National Muséum of Natural History,

Smithsonian Institution, Washington.

HISTORIC

The abdominal retaining apparatus, that is, a pair

of sternal prontinences and a pair of sockets on

the penultimate segment of abdomen, was first

reported and describcd by Duvernoy (1850) in a

chapter on the external reproductive organs in

the Decapoda. The first illustration was given in

a second paper on the same subjcct by Duvernoy

(1853), who wondcrcd that such a “simple

mechanism" (p. 141) was not previously obser-

ved. This author noticed its particular configura¬

tion in leucosiids and the loss of morphological

structures in Uca Leach, 1814 ( Gelasimus

Latreille, 1817).

It took nearly 80 ycars to again hear about the

abdominal retaining apparatus. Pérez (1928a,

1928b, 1929), another French scientist, created

the tenu bouton-pression (an allusion to its fonc¬

tion as a press-stud or a dôme fastener) and revie-

wed the retaining structures in diverse Brachyura.

He had a clear understanding about évolution of

the System, rcgarding the dromiid disposition as

basal, the System in Homola Leach, 1815 as inter-

mediàry (but not derived front Dm min Weber,

1795), the typical press-button présent in the

majority of crabs htit finally lost in Uca, and the

unusual arrangement in leucosiid form.s. From

this evolutionary scheme (1928b: 649) Pérez

hypothesized that the border of the Socket on

pleomere 6 was perhaps the last vestige of the

ancienr uropod. Pérez (1933a, 1933b) examined

the functional patterns with regard to sex, molt,

and maring, and discussed the pecuüar case of the

femini/.ation of the male abdomen in sacculinized

Pachygmpsns nuirrnoratus (Fabricius, 1787).

Dealing with the coaptations in Crustacea,

Macquart (1936) briefly mentioned the locking

mechanism in the Brachyura, suggesting that it

may be replaced by another phy.siologieal System

in advanced taxa such as Uca. Kollmann (1937)

.studied, with more detail, the coaptations bet-

ween rhe abdomen and céphalothorax and the

morphological parts involved in rhe retaining Sys¬

tems, following rhe configuration in primitive

crabs such as Dromia and Homola, then in dorip-

pids, in more advanced form.s such as Carcinus

Leach, 1814, Maja l.amarck, 1801, and in leuco¬

siids. Cuénot (1935: J 94 I: 209-215, figs 43, 44)

was interested by rhe problcmatics ol coaptated

organs and their fmality: the brachyuran press-

button was comparcd to Systems found in the

Cephalopoda and in heniipterid insec ts (Poisson

1922). But the typical locking mechanism device

ol brachyuran crabs was only noticed occasional-

ly (Bliss 1982).

Despite the filet that the diversity ol holding Sys¬

tems was demonstrated by Pérez as possibly

reflccting phylogcnctic rclationships, only a few

taxonomists hâve paid attention ro this character

in brachyuran crabs. Pearson (1908: 15, 19,

fi g. 3) briefly mentioned and tîgured the locking

arrangement in Cancerpagurus Linnc, 1758, with

two small "tubercles" or “papilla’’ on the lift h tho-

racic somite. Hoestlandt (1940; 1948) studied

the apparatus in Eriocbeir si riens is H. Mil ne

Edwards, 1854, and showed that its lunctionality

is lost beyond a certain si/.e of male individuals.

.Stauher (1945) analyscd its condition in dillerent

stages of pinnotherids. Hartnoll (1965) collected

information on grapsid crabs from Janvaica and

describcd rbosc spccies in which morphological

structures tend to he reduccd or lost. Guinot

(1976: 47, 52) ealled attention to the press-but-

ton in the family Belliidae: in Bellia picta

H. Milne Edwards, 1848, males are equipped

with a nonfunctional apparatus.

For I lartnoll ( 1975), who elegantly compared the

copulatory structure and fonction in primitive

Brachyura, three différent patterns of retaining

Systems may be recognized, exemplîfied by

Dromia, Homobi and Lyreidus. Bourne (1922: 69,

pl, 4, fig, 4) was the first author to point out the

System found in Lyreidus de Haan, 1841, that is,

two "small aliform processes’’ on abdominal seg¬

ment 6 and two small knobs on “pterygoid pro-

616

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

cesses” of stérilité 5. The same fearure was remar-

ked in rhe genus Lysirude Goeke, 1986 by

Feldmann ôt Wilson (1988: 478). The subfamily

Lyreidinae was established (Gninot 1993b) by

taking into account the presence of their locking

mechanism, that is unique in the family

Raninidae. In the Podotremata, abdominal réten¬

tion by pereiopods in the dynomenid

Pamdynomene luberadaui Sakai, 1963 and the

disposition in the fossil podotrematous family

Dakoticancridae Rathbun, 1917 were examined

(Guinot 1993a). McLay (1991; 1993) used the

size and shape of uropods, and their rôle in the

abdominal locking mechanism in dromioid

crabs. Tavares (1994, 1996, 1998) was the First

author to use the locking mechanism as a crite-

rion in a cladistic analysis and desctibed two spé¬

cial Systems: a “sliding System” in the family

Cyclodorippidae and a “stop sysrem' in Phyllo-

tymolinum Tavares, 1993, that allowed its érec¬

tion as type genus o( a new family, Phyllotymo-

linidae. Bellwood (1996: 175), who reported the

presence ol a press-button mechanism in the

Calappidac and Hcparidae, and the absence of

buttons in the Orithyiidae and Leucosiidae, also

used her results in a cladistic analysis of these crabs.

The présent paper tabes into account numerous

data given by Guinot (1978a; 1979a, cf. table 5)

who cxtensively investigated most of the

Brachyura in respect of abdominal holding struc¬

tural patterns with an evolutionary insight, and

preliminary results of a thesis on the same subjcct

that was undertaken in the Laboratoire de

Zoologie, Muséum national d’Histoire naturelle

(Bouchard 1996).

MODALITIES OF ABDOMINAL RETAI-

NING AND LOCKING SYSTEMS

PODOTREMATA Guinot, 1977

Guinot (1977, 1978a, 1979a, 1991; Guinot et<il,

1994; Guinot &: Richcr de Forges 1997) divided

the Brachyura into threc sections mainly on the

basis of the location of the sexual orifices. The

coxal positions of male and female gonopores,

with external fertilization, were referred to as

symplesiomorphies, which characterize the podo-

treine crabs. The presence in podotreme females

of rwo separate reproductive ducts. one for egg

laying (génital pore on P3) and one for sperma-

rophore laying (aperture of the spermarheca at

the end of the sternal suture 7/8) contrasts with

the single duct in hetcrotreme/rhoracotrenie

female crabs in wbicb the sternal vulvae are the

sole orifices used for egg laying, for rhe intromis¬

sion of male pleopods (that allows internai ferti-

lizarion) and tor rhe storage of the spermatozoas.

As a restilv of 18S rRNA analysis. Spears & Abele

(1988: 2A) renroved rhe Dromiidae from rhe

Brachyura, arguing rhat the dromiids "branch

very earlv, prior ro the Anomura”. Concluding

that “therc is no inolecular support f...] for the

division Podotremata (sensu Guinot), which

groups raninids and dromiids rogether on the

basis of a similar gonopore location", Spears,

Abele &c Kim (1992: 456) proposcd abandoning

the taxon Podotremata and placing the Raninidae

at the lower limit of the Brachyura. This inter¬

prétation of the molecular data was based on the

study of only four species of Podotremata,

wiihout considering any dynomenid, homolo-

dromiid, homolid, latreilliid, cymonomid or

cyclodorippid groups. But additional investiga¬

tions conducted by the same authors led to ano-

rher major change by considering the Dromiacea

as rrue brachyuran.s ro become the sister group of

the Homololdea (Spears & Abele 1996: I4bis,

and oral statement in 2nd F.uropean Crustacean

Conférence, Liège 1996).

The question of whether the Podotremata is a

monophyletic group remains topical. A parsimo-

ny analysis using mainly sperniatozoal ultrasiruc-

ture suggesrs that the Podotremata is a monophv-

Ietic taxon (Jàmjeson 1994). Investigations in

progress (Guinot & Tavares, in préparation) sup¬

port that on the one hand, Dromiacea and on the

other hand, Homoloidca arc basal podoirenies;

Raninoidca and Cyelodorippoidea sensu Tavares

1994 are more advanced podotremes, somc of

their characters resembling apomorphjes of the

higher Brachyura. Monophyly versus paraphyly

of the Podotremata and their possible placement

as rhe sister group of the hcterotrcme-thoracotre-

me assemblage remain open questions.

An cssentiaJ apomorphv of Brachyura is the loss

of the tail fan, which is présent in “Macrura” and

ZOOSYSTEMA ■ 1998 -20(4)

617

Guinot D. & Bouchard J.-M.

Anomura (Lichodidae and Lomoidea cxccptcd).

In die primitive crabs, i.c. rhe Podotremata, the

uropods can romain as dorsal places (Dynome-

nidae and most Dromiidae) or as ventral lobes

(Homolodromiidae and a few Dromiidae). üuch

platelike or lobelike vestiges are lost in ail other

families (Ilomolidae, Latreilliidae, Poupiniidae,

Cymonomidae, Cyclodorippidae, Phyllotymoli-

nidae, Raninidae; see Table 3, p. 683). Another

robust apomoi phy of rhe Brachyura is rhe abdo¬

men. whicli is rcdueed, iuvcrted, and folded

against rhe ventral sutface, with subset]tient cli-

minacion ol its locomotory fùnction. Il in certain

Podotremata dre abdomen is srill incomplercly

folded, in orhers the pleon acquircs its flexed

position against the thoracic sternum, h: is thus

not surprising to fine! that in Podotremata the

abdomen is retained by various devices.

Mosr Podotremata immobilize their abdomen by

means of a tubencle, knob, clongate (lange, serra-

ted ridge, projection, or spine chat arises from the

thoracopod coxae (Mxp3 and/or perciopods

P1-P2 and even 1*3). These structures overhang

the abdomen, or fit into complementary parts of

the abdomen (coaptations by juxtaposition and

by engagement), or act as a stop System. In ail of

these cases, rhe abdomen is prevemed from slip-

ping out. Additionally, in certain Podotremata

there is a coaptation by assemblage, consisting of

a fixation by abdominal soekets that fit over pro¬

jections isstied from thoracic sternum (sternite 4

for the homolid press-ballon and sternite 5 for

the lyreidine projection).

Various mechanisms can simultaneously be effec¬

tive in the same podotreme crab. A striking

example is shown by homolids (Tables 1, 2):

rétention is underraken by several structures that

originate in the coxae of various appendages, pro¬

bable' used together, and, in addition, by a very

firm press-button.

SUPFIU AMIIV Dijomiacea de Haan, 1833

On rhe basis of larval morpbology, some workers

had for a long rime excluded the Dromiacea from

the Brachyura, whereas Homoloidea and

Raninoidea were retained in the Brachyura as early

or pre-Brachyura (Williamson 1%5, 1982; Rice

1980, 1981a, b, 1983). The same authors attemp-

ted to explain the dromiacean paradox (that is,

anomuran features of the lame and brachyuran

featutes of the adults) by invoking horizontal gene

transfer (Williamson 1988, 1992; Williamson &

Ricc 1996). While investigations on the copulato-

ry structures and rnating hehavîoui showed that

the Dromiacea bave evolved towards rhe brachyu¬

ran condition (ftarrnoll 1975), the Dromiacea

continued to be regarded as' non-brachyuran on

general morphological grounds by a few carcinolo-

gists (de Saint Laurent 1979). The controvetsy vvas

incrcased by the firsr, fragmentary molecular data

(Spcars et al 1992; sec above). Scholtz & Richter

(1995: 321; 1996: 14) fourni évidence for rhe

monophyly of the Brachyura, including the

Dromiacea, from phylogenetic analysis. The first

spermntological analyses indicated that, ai: that

stage of imr investigations, il there is a distinctive

dromiacean groundplan vvhich morphologically

supports the group Dromiacea, sperm structure

does not clcarly disringuish the constituent fami¬

lies Homolodromiidae. Dromiidae and Dynome-

nidae (Jamieson et al 1995; Cuinot et al. 1998).

Considération may bave to be given to the fact

that the majoritv of, il not ail, the Dromiidae are

monophyletic together with the Dvnomenidae

and Homolodromiidae. These spcrmatological

rcsults, however, nced to be regarded wirh pruden¬

ce, awairing more extensive studies in additional

taxa.

Many characters arc exclusively sbared by

Homolodromiidae, Dromiidae and Dynome-

nidae, which endorse récognition of the mono¬

phyletic group Dromiacea. Dromiaceans are ple-

siornorphic by tbe persistence of a pair of appen¬

dages on eacli abdominal segment (in males ves¬

tigial, or sometimes lost, pleopods on pleo-

meres 3-5; in the two sexes, uropods on pleo-

rtiere 6) in contrast to other podotremes (i.e,

Homoloidea, Cyclodorippoidea, Raninoidea) in

which pleopods and uropods in males have been

losr on pleomcres 3-5 and on plcomcre 6, rcspcc-

tively, as in the Heterotrem.ua- 1 horacotremata

assemblage. It should be noted, however, that in

the présent paper we attempt to dentonstrate the

homology between the uropod and the abdomi¬

nal Socket, which implies that rhe uropod is prac-

tically never completcly lost in brachyuran crabs

(see Hypothesis about the homology uropod-

socket. Table 3).

618

ZOOSYSTEMA ■ 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Family Homolodromiidae Alcock, 1900

The family, reviewed and demonstrated as mono-

phyletic by Guinot (1995), shows a great number

of plesiomorphic characters, that support the

contention that it contains lhe most primitive

members of the Podotremara.

Scholtz & Richter (1995: 322) supported the pri¬

mitive status of the Homolodromiidae but wcnt

Fie. 1.— Retainlng apparatus in Homolodromiidae and In Hypoconcha Guérin-Méneville (Dromiidae); A, B, Oicranodromia lelderi

Martin, 9 23 x 14 mm, in the vicinity oi San Andrès, "Oregon”, stn 1908 (USNM)r C, Hypoconcha californiensls Bouvier, î 20.3 x

19,4 mm, syntype, Gult ol California (MNHN-B 22066): D. Hypoconcha arcuata Stimpson, 6 12.3 x 9.3 mm. Sombrero (MNHN-

B 22065). Abbreviations: a5, a6, abdominal segments 5, 6; b, bend al righl angles of abdominal segment 5; c, oatina: cxl, cx2, coxae

of PI, P2; g. granules; t, telson; tu, tubercle; 4, sternite 4.

ZOOSYSTEMA • 1998 - 20(4)

619

Guinot D. & Bouchard J.-M.

so far as to regard homolodromiids as die sister

group of ail otlier Bracliyura; however they hâve

since returned homolodromiids to the Dromia-

cea (Scholtz & Richter, oral statement in rhe 2nd

European Crusracean Conférence, Liège 1996).

We prefer not to separare rhe 1 Iomolodromiidae

front the Dromiidae, even if the combination of

their morphological characters is unique. The

close relatlonship betvveen homolodromiids, dro-

miids and dynomenids is complenienied by the

sperm characters, which show a mixture of fea-

tures of these three familles (Guinot ei al. 1998).

The family Homolodromiidae conststs of only

two généra, Homolodromia A. Milne Edwards,

1880 and Dicranodromta A. Milne Edwards,

1880, with no more than twenry living species.

But mimerons homolodromiid ancestors consd-

tute the dosely related, exclusively fossil, family

Prosopidae von Meyer, 1860, known from the

Middle Juras.sie or even earlier.

Homolodromiids are characterized by small,

exclusively ventral and lobiform uropods, vesti¬

gial pleopods ofteti présent on pleonteres 3-5 in

males, abdominal pleura sometimes markedly

extended, and by a very long telson. The abdo¬

men, which is inserred betvveen the inclined walls

forrned by the coxae of the pereiopods, lies in a

rather decp excavation betwecn the coxae, and

the pleonteres are somewhat shaped to the inter¬

nai surface of the corresponding coxae. Homolo¬

dromiids hâve sterno-coxal dépréssions at the

level of P2 and P3 (Guinot 1995).

In Homolodromia paradoxa A. Milne Edwards,

1880 the coxae of PI and P2 are smooth and the

abdomen xeems simply fiexed betvveen the per¬

eiopods. In most homolodromiids, however,

there are a few granules on the coxa of PI and

some more numerous and regulatly arranged gta-

nules on rhe coxa of P2: if they do not dearly

overhang the telson, at leasr they perhaps restrict

the sideways movemems of the abdomen. Jn H.

kai Guinot, 1993, Dïcrunodromui ko ru bar

Guinot, 1993 and D. nagaii Guinot, 1995, for

example, the vveakly granular coxae of Pl are

clearly hollowed at their hases and the telson

slides along beside them; on P2 sonie coxal sharp

granules are located close to the abdominal mat-

gin. In II. bolivien Doflein, 1904 the granules on

P2 are particularly acute and corne very close to

the telson, being perhaps effective. In males,

sometimes otherwise almost completely smooth,

these granules occur only on PI and P2 coxae.

The granules are lost in mature females, in which

the internai surfaces of the coxae become smooth

and modifiée! to receive the larger abdomen.

In only one species, Dïcranodromïa fdderi

Martin, 1990, where carinate ridges project over

the ventral margins of the ftrst pereiopods

(Martin 1990, (ïgs lb, le, 3A), a unique struc¬

ture has been discovered, which is similar in

females and males (Guinot 1995: 250, fig. 32b)

(Fig. 1 A, B). The inferior part of the coxae of the

chelipeds forms a distinct dépréssion, which is

surrounded by a thick, smooth, carinated and

horseshoe-shaped prominence. l'his structure

overhangs rhe extremity of the telson on both

sides, prevenring it iront being lifted up, and the

lel.son is retained by sliding through the déprés¬

sion. A few granules are présent on the Pl coxae;

more numerous and regularly arranged granules

are also présent on the P2 coxae.

Discussion

The carinate structure of D. felderi appears to be

similar to rhat in other Podotremata in having a

coxal projection front different legs that ove¬

rhangs the abdomen. It is, however, developed on

the whole coxa of Pl and it is smooth, not spi-

nose. The P1 coxa! keel of D. felderi evokes the

Pl and P2 coxal carinae of Hypuconcba califor-

niensis Bouvier, 1898 (Fig. 1C) (see below). the

rôle of which remains enîgmatic. if in the

Homolodromiidae the différentiation ai D. felde¬

ri seents to be an effective retaining structure, the

acute granules ou the coxae of the two first pe¬

reiopods, specially on P2, could be also ituerpre-

ted as structures involved in retaining the abdo¬

men (Bouchard 1996).

It wlll be valuable to look at the numerous fossil

prosopid forms to verify whether such structures

are présent.

Family Dromiidae de Haan, 1833

The extensive révision of rhe sponge crabs by

Mcl.ay (1993) did not support their exclusion

from rhe Brachyura and their association wirh the

Anomura. Evoking rhe complex evolutionary

relationships among more than hundred Atlantic,

620

ZOOSYSTÉMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Indian and Pacific species distributed in about

thirty gênera, McLay (1993) did not subscribe to

the paraphyly of the family. Only die placement

of the genus Hypoconcha Gucrin-Méneville,

1854, was considcred doubtful. The question is

to know il Hypoconchd is relatcd or not to another

dromiid, Conchoccetes Stimpson, 1858, which

shares with the Hypoconcha species an unusual

camouflage habit of carrying a bivalve shell

Analysis ol the problem suggests rhat Hypconcha

probably belongs in the Dromiidae (McLay

1993: 229).

We vvill lollow here the new generic arrangement

proposed by McLay (1993) where attention was

paid to “the size and shape of uropods in relation

to their rôle in the abdominal locking mecha-

nism”. In dromiids the still artîculated, sorne-

times mobile, uropods are (more or less) functio-

nal only when they to ns ist of dorsal plates, which

is the case In most genera/species. Very few dro¬

miid species are devoid of any apparent retaimng

structures.

if a truc sterno-abdominal cavity is lacking in

dromiids, the abdomen often serdes itself by its

posterîor part in the deep dépréssion formed by

the inclined sides ofthe coxae of P1 -P2. The uro¬

pods are markedly sexually dimorphic in shape

and size. In males and in young females uropods

are more or less laterally prominent and visible

dorsally, whereas in mature lemales they usually

seem transversally extended, intercalated between

the segment 6 and the telson, and somewhat

concealed in the general outlinc of the enlarged

abdomen. The telson also is sexually dimorphic,

being longer and narrower in adult males.

Dromiids display several means of abdomen

rétention, thaï involve the différentiation of one

or several paris of the peteiopods and abdomen.

In addition to the variable structures projecting

front the coxae of the rwo firsr pereiopods, the

vestigial uropods may lie invoived. The signifi-

cant rôle ol the uropods in dromiid males is cor-

relatedwifh their sideways projection bevond the

latéral ntargins of the abdomen.

Sonie examples of the dromiid devices are here

descri lied.

Structures on PI only

The case of the abdomen being retained only by

one pair of legs, in this case PI, is found in spe¬

cies of the shell-carrying genus Hypoconcha

Guérin-Méneville. in which the broad and angu-

lar abdomen lias distinct pleura and where the

uropods are reduced to two very minute plates

intercalated ventrally, not visible dorsally.

In H. califomiemis (Fig. 1C) thccoxa ofPl bcars

a group ol spinous tubercles, the strongest of

which overhangs the external border of the telson

in its half posterior part and liolds it. The mit of

the curious keels on the coxae of PI and P2

found in H. califomiemis is enigmatic and evokes

tlie case of Üicranodromia felderi (Fig. IA, B);

such marked carinate structures are not présent in

ail Hypoconcha species. In the genus Hypo-concha,

H. araiata Stimpson, 1858 (Fig. 1D) and

//. panamensis Smith, 1869 hâve no coxal projec¬

tions, and their System of abdominal rétention is

spécial. The male abdomen is short, with the tip

of the telson reaching the level of P2, and the

posterior part of segment 5, close to the articula¬

tion with segment 6, is bent at right angles so that

the posterior part of the abdomen (posterior part

of segment 5, segment 6, and telson) lies fiat on

the ventral surface.

In the dromiid Haledromia bicavcmosa (Zietz,

1887) (speetes not examincd), with a spécial

combination ofcharacters (in particular widened

carapace, small and concealed uropods, and, pro¬

bably, direct development), McLay (1993: 158,

table 2) described a retaining apparatus that

m volves a blunt “tooth on bases of first legs

against margin of penultimate (abdominal] seg¬

ment,”

Structures on PI and P2

Both structures functional.

In Lauridromia dehaani (Rathbun. 1923)

(Pig. 2A, B) the coxae of PI and P2 each Ivave a

denticulaied prominence which fus into a déprés¬

sion on the external margin ofthe telson and the

somite 6, tespecrivcly. In a ventral view of the

abdomen, along the external margin and just pos¬

terior to the uropod, a notch on the telson

receives a coxal prominence of PI while a deep

dépréssion receives a coxal prominence of P2. In

addition, the mobile uropod is invoived, being

engaged against the bases of rhe coxa of P2.

W hen the abdomen is extended, the pair of coxal

ZOOSYSTEMA • 1998 -20(4)

621

Guinot D. & Bouchard J.-M.

prominences on P2 acts as in full-lock with the

pair of uropods. Thus, liere, the uropod plays an

important rôle.

It is not clear why specimens of L. dehaam exa¬

mi ned by Dai &: Ng (1997: 757) are indicated as

having a serrated flange “obvious only on the

coxa of the First leg."

Structures un Pl supposed nonfunctional

In Lauridromia intermedia (Laurie, 1906)

(Fig. 2C) the abdomen settles tightiy bervveen the

bases of the first pereiopods, the coxa of which

bears a granular prominence, probably not func-

tional; on the external margin of the telson the

corresponding area is not notched. in compensa¬

tion, the well-developed coxal prominence of P2,

which bears a fine and regularly denriculated

crest, fits into a dccplv cxcavated dépréssion on

the latéral edge of abdominal segment 6. The nar-

row uropod find.x irs place perlectly in contact

with the coxal projection, and ihe abdomen is

firmly retained at dus level. The internui side of

the P3 coxa bears a nonfunctional minute granu¬

le. Additionally, on the sternum two areas are dit-

ferentiated, one with sparse granules at the level

of the P1 condylus and another one with a tuber-

culate crest at rbe Jevel of the P2 condylus.

Variations exist concerning rhe relative position

of the uropod in regard to rhe coxal projection of

P2. In Takedromia enstatipes (T. Sakai, 1969) the

high coxal prominence on P2 cornes into contact

with the uropod, which is movable.

In Dromia erythropus (G. Edwards, 1771)

(Fig. 2D) and in D. marmorea Forest, 1974, the

uropod lies ta the r far from the coxal prominence.

A similar organizarion is observed in D. monodi

Forest et Guinot, 1966 (Fig. 3A), which has spe-

cially mobile uropods and a coxal prominence on

P2 bearing 8-10 small denticles aligned on a high

oblique crest. Flic coxa of PI remains tubercula-

te but does not seem to be effective since lhere is

no contact with the toison. In D. monodi sonie

granules are also présent on the latéral part of the

sternite 4. In h is diagnosis of the genus Dromia

Weber, 1795, McLay (1993: 149) tnentioned the

uropods as “used in male abdominal locking

mechanism by fitting in Iront of serrated flange

on the bases of first legs”, a sentence that led

Dai &C Ng (1997: 757) to misinterpret ‘this ser¬

rated flange supposedly présent only on the coxa

of the first leg,”

Homatodromia coppingeri Miers, 1884 (Fig. 3B)

has longirudinally extended and pointed uro¬

pods, that are clearly mobile. On the Pl coxa

there is a small tuberculatc prominence and on

the P2 coxa a particuJarly high, cupuliform, and

denriculated prominence that is nearly horizontal

at rest and becames funcrional by the movement

of the leg, This prominence consritutes a full-lock

System with uropods: rhe long telson is not invol-

ved.

The structure on the Pl coxae of Dromia perso-

nata (I.inné, 1758) (Fig. 4B) and D. bollorei

Forest, 1974, which consists of a small granular

prominence far from the abdomens margin, is

not functionai. The salient granular prominence

on the P2 coxa cornes in contact with the border

ol segment 6 and is distant from the mobile uro¬

pod. The abdomen of the preserved specimens

examined was not retained.

Only ibe observation of living animais may lielp

to conclude how efficient the mobile uropods are

as a full-lock mechanism.

Structures on P2 and P3

When rhe abdomen is short and the telson’s tip

lies far from rhe level of the chelipeds, norhing

functionai is observed on the Pl coxa, and struc¬

tures on P2 and P3 are présent instead.

In Conchoecetes intermedius Lewinsobn, 1984

(Fig. 4A), a bivalve-cajrying crab, rhe P2 coxa

bears a strong and ornamented prominence that

acts as a lull-lock mechanism with a very dose,

salient uropod. The narrow epi.s terni te 5, which

remains exposed and visible between rhe coxae of

P2 and P3, is also provided with some granules.

In addition, a rounded tubercle (the internai one

of an aligned séries) on rhe P3 coxa cornes into

contact with an meurved part ol the latéral abdo¬

minal margin at the level of segment 5. Similar

granules on the P4 coxae (the legs that hold the

shell) are too small to be effective.

In Sphaerodromia Alcock, 1899, which is consi-

dered the most primitive known dromiid genus

by McLay (1993: 127) individuals carry large

pièces of sponge for camouflage. McLay (1991:

466) reported a rdatively “elaborate grasping

mechanism” in the genus. Ail preserved speci-

622

ZOOSYSTEMA • 1998 - 20 (4)

Fig. 2. — Rctaimng apparatus in Dromlidae. A. B. Lauridromla otedaan/(Ralhbun); A, 80.4 y 76.9 mm. Seychelles (MNHN-B 7760);

B, t 42x45 mm. Gulfol Suez (MNHN-B 21994): internai surface ol abdomen; C. Lauridromia irUermeüia (Laune), c5 52 4 • 48.1 mm,

New Caledonta (MNHN B 12754): O. Dromia erythropus (G. Edwards), <1 25.2 ■ 25.3 mm, Venezuela (MNHN-B 12713), E,

Sphaerodromla nux Alcock, Ç 58.7 - 67,1 mm, Madagascar (MNHN-B 6872); withou! abdomen. Abbreviatkms; a5, a6, abdominal

segments 5-6; ex 1 cx2 cx3, coxaeof Pi-P3; d. dépréssion; n. nolch, pr, projection oflhe coxa, t, telson; u. uropod; 4,5, sternites 4,

ZOOSYSTEMA • 1998 -20 (4)

623

Guinot D. & Bouchard J.-M.

mens that were examined did not hâve their

abdomens weJI retained. We did not observe any

specialized structures on the PI coxa of S. nux

Alcock, 1899 (Fig. 2E), S. ducoussoi McLay, 199!

and S. briznj>s McLay et Crosnicr, 1991. On the

coxae of P2 rhere is a disrinctiy granular promi-

nence that acts on anrerior part of the telson (the

dorsal uropods are not involved); on the coxa of

P3 there is a larger granular prominence, that is,

however, completely covered by abdominal seg¬

ment 5.

Structures on P2 only

In Hemispbaerodrornia rnonodus (Stebbing, 1918)

(Fig. 3C, D), which has a narrowed abdominal

segment 6 and mobile dorsal uropods, the whole

base of the P2 coxa forms a rounded and salient

prominence, that is serrated on irs infernal and

superior margins. This prominence is rigrhly

encircled in the space just behind the uropods

and fits into a dépréssion on the border of pleo-

mere 6, much thickened at this level.

In Cryptndromiopsis avtillensis (Srimpson, 1858)

(Fig. 4C), in which the dorsal uropods apparent-

ly do not play a rôle, a short but sufficiently deve-

loped spine that projects from the base of the P2

coxa overhangs segment (S anteriorly and fîrmly

holds the abdomen. It is noteworthy that a mar-

kedly visible granule is présent on the P3 and P4

coxae.

The three known species of Exodrornidia

Stebbing, 1905, E. spinosa (Studer, 1883), the

type spccics ot the genus, E. bicomis (Studer,

1883) and /:. spinosissima (Kensley, 1977), ail

Irom South Africa, were examined. They are cha-

racterized by vestigial male pleopods on seg¬

ments 3-5 (not discernible, however, in ont

small-si/.ed male of F. spinosa ) and hy uropods

that are only small ventral lobes and cannot play

any rôle in retaining the abdomen. Ail three show

a highly marked différentiation of the coxae of P2

(Guinot 1995: 187), a feature that was neglected

in the taxonomie descriptions. In E. bicomis

(Figs 4D, 5A), a ver)' 1 long, stout, smooth (and

white) spine projects from the coxa of P2 in a

direction parallel to the latéral margin of the

abdomen. In E. spinosissima (Fig. 5B, C) the 1*2

projection is relatively thicker and tubetculâte at

its base, and similarly dtrected. This projection

overhangs abdominal segment 5 and the poste-

rior portion o( segment 4 only when the P2 are

moved hackw-ards. The presence of such strong

coxal projections is unique in dromiids. Their

rôle in preventing the abdomen being lifred up is

évident, and significantly implies an active invol-

vement of the legs (i.e . P2) in order to place the

long spincs ahove the abdomen.

Exodrornidia spinosa (Studer, 1883) (Fig. 5D, E)

shows a rounded and granular prominence that

projects from the P2 coxa and fits into a deep

dépréssion on the margin of abdominal

segment 6. This segment is distinctly hollowed

along mosr of its thickened edge. The coxal pro¬

jection is visible on the illustration given by

Doflein (1904, pi. 6, fig. 2); the dépréssion on

plcomere 6 and the telson’s spine are not visible

in the sketches by Macpherson (1983: 13,

fïg. 6A). The modification of the external edge of

pleomere 6 resembles that of Hemispbaerodrornia

(Fig. 3C, D), but a différence lies in the facr that

mobile dorsal uropods are involved in

Hemisphaerodrotniti , whereas absolutely not in

F. spinosa (only small ventral lobes).

The three Exodrornidia species hâve the posterior

part of telson ending in an acutely sharp spine in

males (Figs 4D, 5A, B, D) but rounded in

females. The acute dp moves insidc a marked fur-

row' rhat is hollowed on sternitc 4 in males, whe¬

reas sternite 4 is norgrooved in females. The fur-

row' of sternite 4 could be particularly "useful”

when in E. bicomis and E. spinosissima the pair of

long coxal projections rnust overhang the abdo¬

men at times when free movement of the abdo¬

men is necessary.

E. bicomis, chelipeds of which are considerably

elongated, evidence of strong positive allotnetric

growah, is a peculiar dromiid: hits of rural are

attached to the long pseudorostral teeth, and the

whole body is concealed by fragments of sand or

Fig. 3. — Retaining apparatus in Dromiidae. A Dromia monodi

Forost et Guinot. -î 30 • 32 mm. Mission Casamance (MNHN-

B 21937): B. l-lom.ilodromia coppingefi Miors, < 5,2 s 5 3 mm,

Se/chéiles (MNHN B 26130): C. D Hemisph$ei't,i<iion)i/i mono-

clos ISlebtwog), 14 15.5 rnm. Fort-Dauphm (MNHN-

B 11204): m D detail of abdomen, ventral (above) and side

(below) vivws Abbieviaiions, a5 a6, abdominal tégiuuiits 5, 6;

cxl cx2. coxae ot Pi . P2; d. dépréssion; mxp3, external maxil-

liped; pr, projection; t, telson; u. Lropod: 4 sternite <1 Satae

were not figured. Scale bars: imm.

624

ZOOSYSTEMA • 1998 -20 (4)

Guinot D. & Bouchard J.-M.

mud among rhe stiff bristles and spines. Carrying

behaviour vvith the very short, but subchelifbrm,

last pereiopods is possible, but pcrhaps unneces-

sary (Guinot et al. 1995). E. spinosa also has long

chelipeds and the carapace covered vvith slill

bristles, but materials oi settlement wete not

observed on tbe body of the two specimens that

were examined.

No apparent structures

[n the species of Pseudodrnmiti Stimpson, 1858

that were examined, P. latent Stimpson, 1858 and

/.’ rotunda (Macleay, 1838), which are always inti-

mately associated with ascidians, and in the

monospecific genus Asadiopbilus Richters, 1880

(with A. caphymeforrnis Richters, 1880, a small-

sized species with the body tightly enclosed in a

Fig. 4. — Retainlng apparatus in Dromildae; A, Conchoecetas întermedius Lewinsohn, 6 16 x 17 mm, hololype, Madagascar

(MNHN-B 6891), B, Dromia personata (Linné), i 52 :■ 65 mm, Azores, Faial (MNHN-B 21969); C. Cryplodromiüpsis antillensis

(Stimpson). cl 18 x 18 mm, Frencn Guyana (MNHN-B 22029); D. Exodnomidia bicornis (Studsr), S 18 (wlthout pssudorostral

spines) x 19 mm. Souh Africa (SAM 10724), Abbreviations: a6, abdominal segment 6; cxl, cx2. cx3, coxae of P1-P3; f, furrow on

sternite 4; g, granule; mxp3, external maxllllpea; pr, promînence; sp, spme; t, telson; tu, tubercle; u, uropod; 3. 4, stérilités 3. 4.

62 6

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Fig. 5. — Retammg apparalus In the Ihree specips of Exodromidia Stebbing (Dromîidae) A, Exodromidla bicornis (Suider), ■ 18

(without pseudorostral spinas) • 19 mm, South Afnca ISAM Al0724): 0 C, Exodromidia spinosissima (Kensley). . 4,8 ■ 4 6 mm,

South Africa (SAM Al 7681 ): In C, detail of rlght and left PS coxae (the projections are broken at their tlps. at least on coxa of teft P2);

D, E, Exodromidia spinosa (Studer), ! 29 • 32 mm, Cape Point (BMNH 1929.12.1.234-235) in E, detail of abdomen in ventral

(above) and side (below) vîews. Abbreviatioris: a5, a6, abdominal segments 5. 6, cx2, coxae. ot P2; d.s., sterno-coxal dépréssion;

f, furrow; mxp3 oxlernal maxilliped; pl5, vestigial pleopod on abdominal segment 5; pr, projection, sp, spine; t, telson, u, uropod;

3, 4, sternites 3. 4. Setae were not figured. Scale bars: A, C, D, 1 mm; B, 0.5 mm; E, 2 mm.

ZOOSYSTEMA • 1998 - 20(4)

627

Guinot D. & Bouchard J.-M.

compound ascidian held by rhe prehensile P4

and P5, see Guinot 1995), rhe absence of an

abdominal rétention in males can be inferred by

the protection obtained by living inside rhe asci-

dian. In these highly specialized forms, the vesti-

gial pleopods on segments 3-5 are lost. The uro-

pods are only represented hy small ventral lobes

in Pseudadromia , whereas rhere is no trace of uro-

pods în Ascidiopbiins,

Tunedromia yamasbitai (Takeda et Miyaké,

1970), type species of rhe genus Tunedromia

McLay, 1993, and only known from lemales, is

indicated as lacking uropods (McLay 1993: 134,

table 1), but males need to be examinated.

Discussion

The case in which only PI i.s involved, rétention

of the abdomen ar the level of rhe telson charac-

terizes Hypoconcba ralifomiemis. In oiher cases,

the involvemenr ofother appendages (P2 and/or

P3) and of the dorsal uropods frees the PI from a

retaining funccion. In Exodromidia, which lacks

dorsal uropods, only P2 is involved. Only svhen

the coxal structure on PI plays a rôle, an involve-

ment of ihe telson was observed with a subsé¬

quent modification ot its edge, which can be

markedly thickened, notched or hollowed (coap¬

tation by juxtaposition or by engagement). In the

dromiid evoiutionary sériés, the loss of the retai¬

ning fonction of the abdomen by rhe chelipeds

seems correlated with rhe loss of rhe holding at

the level of the telson. Thus the only appendages

able to play this rôle, by nteans of a coxal projec¬

tion, are P2 and P3. The P4, which in dromiids

are reduced and along with P5 hold a camoufla¬

ge cap, cannot be used for abdominal rétention.

The P2, which develop rhe most complex and

probably the most efficient Systems, remain rhe

only appendages to be used. At this stage, the

sternal somite involved in retaining is the fifth

(corresponding to I’2) and rhe involved pleomere

is the sixth. In rhe case wbere rhe coxal promi-

nences on P2 and I’3 ( Spbaerodrornia ) are com-

pletcly covercd by the abdomen, these structures

are similar to those in the preceditig cases

although they are ineffective.

As in ail Brachyura the abdomen is used for the

protection of the eggs in female dromiids. In the

family Dromndae the sexual dimorphism of uro¬

pods concerns not only the size (uropods are

generally smaller in males) but also the shape and

orientation (uropods are generally more salient in

males), so that the rôle of rhe uropods remains

very efficient in adult males.

Among the numerous modalicies of abdominal

rétention shown by dromiids, we observed cases

in which the abdomen, simply applied fiat on the

sternum berween the legs, is more or less effi-

ciently immpbilized by a prominence that acts as

a stop, often with horizonrally expanded uropods

( Conchoecetes , Dromia personata , D. bollorei ,

D. monodt). In other cases ( Lauridromia ), the

abdomen, with almost vertical uropods, is inser-

red in a rather deep dépression berween the legs,

and the edges of rhe telson and segment 6 beco-

me highly modified complementary to the cor¬

responding coxae. Coaptations hy juxtaposition

and by engagement perform an cxtrcmely effi¬

cient way of retaining the abdomen. We suggest

to name the configuration in which the uropods

contact with coxal prominence», as the full-lock

System.

fhe case of Hetnisphaerodrotnia monodus com¬

bines a coaptation by engagement (the rhick bor¬

der of pleomere 6 is distinctly hollowed) and a

full-lock berween rhe serrared prominences and

the very mobile dorsal uropods. This configura¬

tion provides astrong System.

The uropods are very mobile iu many cases (for

example Dromia personata, D. bollorei, D. mono-

di. Lûuridramia dehaani, Homalodromia coppin-

geru Hemisphaerndromia monodus), and this is

probably correlated with their rôle in abdominal

rétention.

Other modalides are exemplified hy cases of

simple overhanging, without any coaptation bet-

ween the abdomen and the coxae of legs, and

without the involvement of uropods

(Cryptadromiopsis antillensis, Exodromidia). The

coxal P2 projection in /:. bicornis and E. spbiosis-

sima appears as the most devcloped amongst the

dromiids and perhaps in ail braçhyuran crabs.

The abdominal segment 6 nvay he markedly

modified in relation to its involvement in retai¬

ning the abdomen. Its thickened external edge is

distinctly hollowed in Lauridromia dehaani,

!.. intermedia, Hemisphaerndromia monodus and

Exodrot7iidia spinosa.

628

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Modifications of the telson in relation to the

retaining apparatus are sometimcs pronounced in

dromiids: an cxamplc is the deep notch on the

external margin in LauridrorHia débattui. In the

genus Exodroniidia, it seenis that the sexually

dimorphic telson may play a rôle when it moves

inside the furrow on sternite 4 to put the P2

coxae in an overhanging position. This déprés¬

sion of sternite 4, absent in females, also consti-

tutes a sexually dimorphic feature and, if it is rcal-

ly involved in facihtating the accommodation of

the telson spine, it is also an exceptional pheno-

menon in brachyuran crabs.

The cases with no apparent retaining structures in

dromiids are rare. As far as we know, it only exists

in species aSsociatcd with ascidians, viz. Pseudo-

drumia rotunda and Ascidiophilus caphyraeforrnis.

Hypoconcha an a cita and //. panamemis also lack

differenriated structures, but the)'' markedly differ

in having their abdomen lient at right angles, and

with its posterior hall particularly orienred to the

ventral surlace. This abdominal curvature, iess

pronounced in males than in females, seems to be

correlated with the inclination of the two last tho-

racic sternites relative to the preceding ones.

Family Dynomunidal Ortmann, 1892

The phydogenetic relationships of the family

Dynomenidae to both Homolodromiidac and

Dromiidae (exem pli lied by Dynomene tanensis

Yokoya. 1933. Homolodrnrnia kai Gtiinot, 1993,

and Sphaerodromia Idmellata C irosnier, 1994, res-

pectively), bave been examined usingspermatolo-

gical characters (Guinot et al. 1998). As in the

case of general morphology the ultrastructurc of

spermarozoa strongly endorses monophyly of the

Dromiacea, without a clear récognition of its

three familles, however.

The question of the existence of a retaining appa¬

ratus in dynomenids was first raised by Guinot

(1978a, 1979a, 1993a); see also McLay (1991).

In dynomenids the thoracic sternum is relatively

wider than in dromiids and forms a plate that

inclines posteriori)'. The only sternite that is pla-

ced in a plane other than the preceding ones is the

sternite 8, which corresponds to the last pair of

legs. The P5, which are reduced but are neither

dorsal nor specially movable (but their rnove-

ments are coordinated with those of the other per-

eiopods, C.. L. McLay pers. comm.), are always

intercalated between the postérolatéral margins of

the carapace and the bases of P4. A sterno-abdo-

minal cavity sensu stricto is not présent. T he abdo¬

men is veiy broad, even in males, relatively much

broader than in dromiids, flexible, and is not clo-

sclv held on the sternum. The dynomenid abdo¬

men is the most primitive one within the

Brachyura in maintaining pleopods on seg¬

ments 3-3 as (gcnerally) biramous vestiges. Dorsal

uropods are always well-developed, nevet projec-

ted, even in males, and they show sexual dimor-

phism, being larger in females. In preserved dyno¬

menids, ac leasi in aduJt individuals, the male

abdomen is never firmly held against the sternum,

resembling the lemale condition.

Apparatus on the coxae oj thoracopods

An example of an effective retaining apparatus is

shown by Acdnthodromia erinacea A. Milne

F.dwards, 1880, a spccics ornamented wirh many

spines. fn the single mature female examined, the

broad and curved abdomen covers the whole ster¬

num. and the developed telson covers the proxi¬

mal part of Mxp3 (Figs 6A, 7A, B). The coxae of

four thoracopods are involved. The eoxa of Mxp3

bears a particular Spinule, stronger than the sur-

rounding ones, which overhangs the posterior

part of the long telson. The coxae of the three

first perciopods (P1-P3) présent a latéral déprés¬

sion that props up the abdomen. In addition,

each coxa is equipped with two to three orna-

ments (spinules or granules) that overhang the

telson at P1-P2 level and the pleomere 6 with its

uropods at the P3 level. On episternite 5, which

is covcred by the telson, two small granules are

présent but they seem not to play any rôle. This

configuration in females is probably similar to

that tound in males, which werc not examined.

In other dynomenids, in which the abdomen is

much shorter and tends to not entirely cover the

sternal plate, sonie structures were observed

which can bc referred to as a retaining System,

that is perhaps inefficient at least in large adults.

According to species, it involves eidier P2 t P3

(never PI) or sternite 5, and two main types may

bc describcd: one analogous to Acanthodrornia

and another one (see bclow, Apparatus on the

sternum).

ZOOSYSTEMA • 1998 ‘20 14)

629

Guinot D. & Bouchard J.-M.

In two Dynornene species, two wcak and orna- gin, another coxal prominence cxists on P3, but

mented coxal prominences occur on P2 and P3, it is even smallcr. In a male individual from

respectively, in the adult males. In D. tancnsis Glorieuses lslands idenufied as Dynomçnc aff.

Yokova 1933 (Figs PB, 7C) there is a well-deve- detoineyi lakcda, 1977 (Fig. 6C), lhe coxae of P2

loped and tuberculace prominence locaied on the and P3 hear a Weak prominence, the ornamenta-

P2 coxa lhat cornes very close to the uropod mai- don of which is asymmetrical: on the left P2

Fig. 6. — Retainmg apparatus in Dynomenidae: A Acanthodramia erinacea A. Milne Edwards, ovigerous 14 7 > 12 mm. West of

Puerto Rico (USNM 124263}; B, Dynorvene tanensis Yokoya, 1 19.4 '.19 7 rtim, New Caledonîa (MNHN B 25583); C, Dynomene

aff. devaneyi TaKeda, 6 23.2'- 22.7 mm. Glorieuses Islands. Benthedi Exp. 1977 (MNHN B 22510); D, Dynomene hispida Guèrin-

Méneville. J 8.8 • 11,5 mm, New Caledonia (MNHN-B 22091). Abbrevialrons. a6, abdominal segment 6; cxl. cx2, cx3, coxae of

P1-P3; mxp3. extarnal maxllliped; pr. coxal, or sternal prominence- sp, spine, t, telson; u, uropod; 1-3. sterpites 1-3; 4, 5, sternites

4, 5.

630

ZOOSYSTEMA • 1998 - 20 (4)

Abdominal holding Systems (Brachyura)

mxp3

Fig. 7. — Retainlng apparatus in Dynomenldae: A B, Acanlhodromia ennacea A Milne Edwards, ovigerous i 14.7 x 12 mm. west

of Puerto Rico (USNM 124263): thoracic sternum with and without abdomen: C. Dynomene tanensis Yokoya, c 12.2 13 mm. New

Caledonia, Smtb 2. stn DW1 (MNHN), wilh detail ot the prominences on P2 and P3 coxae; D, Dynomene hisplda Guèrin-Mèneville,

6 6.9 x 8.8 mm. New Caledonia (MNHN 8 22091) Abbreviations a6, abdominal segment 6; cxl. cx2, cx3. coxae of P1-P3; d.s.,

sterno-coxal deprossion; mxp3, oxternal maxillipsd; pr, projection; sp, spinule; t, telson, u, uropod; 1-3, sternltes 1-3; 4, 5, sternites 4,

5. Setae were not figured. Scale bars; 1 mm.

ZOOSYSTEMA • 1998 -20(4)

Guinot D. & Bouchard J.-M.

coxa, the prominence is distinctly bifid; on thc

right P2 coxa and on the left P3 coxa, the promi¬

nence has only one tuburcle, and on the right P3

coxa the prominence shows a row of three

tubercles. The complété System appears not to be

effective. The présence of “blutât teeth” on the P2

and P3 coxae was noticed by Mcl.uy (1991: 466)

in D. devaneyi Takeda, 1977.

The case of Paradynomene tuberculata T. Sakai,

1963 (Guinot 1993a), the body of which is com-

pletely covered by large granules, is difficult to

analyse. The external margin of the abdomen

cornes close to P2 and P3 coxae, and only the

very exrernal part of the episternire 3 is visible

when the abdomen is flexed. The sets of granules

at the prominent bases of the P2 and P3 coxae

appear as possible rneans of restricting the latéral

movemcnts of the abdomen. The episternite 5 is

ornamented with a lew sparse granules which do

not appear to be effective.

Apparatus on lhe sternum

In male Dynomene bispida Guérin-Méneville,

1832, type species ol the genus Dynomene , the

différentiation is locatcd on the narrow exposed

episternite 5 (Pigs 6D, 7D). Sometimcs its surface

seenis to be granular or lormed hy more than one

tubercle or spine. Tliis prominence on sternite 3

lies close to the uropod; it is possible that in the

juvéniles it overhangs the uropod. But the pro¬

minence bccomes more minute in larger indivi-

duals, and it seems to be in relation to the uropod

(Guinot 1979a, pi. 21, figs 8, 9).

The same configuration of a sternal tubercle, that

is more or less prominent or acute but apparent-

ly ineffective, exists in tnany of the dynomenids

that were examined, such as D. fillmli Bouvier,

1894, D. pmedator A. Milne Edwards, 1879,

D. spinosa Rathbun, 1911, and D. ursula

Stimpson, 1860. These Dynomene species charac-

terized by a sternal différentiation are in contrast

to the rwo preeeding D. tauensis and D. alf. deva¬

neyi, as well as to the généra Acanthodromia and

Paradynomene , which hâve coxal différentiations.

Discussion

It is noteworthy that in certain dynomenids the

retaining structure does not involve the limbs any

more but takes place on the corresponding somi-

tc (sternite 5)» certainly a conséquence of the

broadening of the thoracic plate. Such a différen¬

ce of position indicates to us that diverse dyuo-

menid crabs belong to distinct généra instcad of

the tradicional single genus Dynomene , in other

words, that D. tauensis and D. alf. devaneyi must

be disiinguished in regard to thi.s character and

cannot belong to the same genus as the typical

species D. bispida. It is obvious that in the cases

where coxal structures are involved (Acantbodro-

mia, Paradynomene, D. tauensis and D. aff. deva¬

neyi, Figs 6A-C, 7A-C), the abdomen laterally

filLs thc wholc space berween the coxae and ante-

riorJy covers a large part of sternite 4. In the case

where sternal structures are involved ( Dynomene

bispida, Figs 6D, 7D) the thoracic sternum

appears latcrally and anteriorly more exposed.

Dynomenids show a pattern like that of dro-

niiids, with an involvemenr of rhoracopods and a

major rôle of P2. No drorniid crab, however, has

a structure locatcd on Mxp3 analogous to the

configuration in Acanthodromia. In regard to dro-

miids, a dynomenid innovation is, in many of the

Dynomene , thc sternal différentiation on thoracic

somite 5, corresponding to P2, just f'acing Lhe

large uropod. The question is to know if the

minute sriucture positioned on rhe P2 coxa or on

sternite 3, that just touches thc uropod (perhaps

overhanging thc uropodai margin in thc young

individuals) orquite apart in adults. is hencc in a

position to be effective in retaining. Wc can assu¬

me tlvat, in large males, thc abdomen is restricted

in its latéral movemcnts. The rudimentary struc¬

tures, either coxal or sternal, are placed in such a

manner that they may fill the same rôle.

Dynomenids djfter Iront dromiids in ntany

points: in dynomenids rhe dorsal uropods do not

appear to be modified and their sexual dirnor-

phism seems to bave no effect; the sexually

ditnorphic telson never seems to he modified. As

homolodromiids and most dromiids, dynonte-

nids hâve stcrno-coxal dépressions at the level of

P2, P3, and also P4,

The dynomenid apparatus, which disappears in

females after the puberty moult, should be stu-

died in detail, cspecially in living animais during

their growth. l he examination of rhe numerous

foïvsil dynomenids would be informative as well.

The discovery in ancestral dynomenids of sternal

632

ZOOSYSTEMA • 1998 -20(4)

Abdominal holding Systems (Brachyura)

mxp3

Fig. 8. — Retaining apparatus in Homolidae and Poupmiidae A, Homola orientalis Henderson, 6 34.8 x 23 mm, French Polynesia

(MNHN-B 22385); B, Paromola bathyalis Guinot al Richer de Forges, i 72 x 48 mm, paratype, New Caledonia (MNFIN-B 19898);

C. Homolomannia sibogao Ihle, rî 39.2 x 27 mm. Philippines. Musorstom 2 (MNHN-B 13825); D, Poupinia hirsula Guinot, £ 48 x

38 mm, holotype French Polynesia (MNHN-B 24346). Abbrevlations: a6, abdominal segment 6; cxl, cx2, coxae of Pi, P2; mxp3,

external maxîlliped; sp, splne; t, telson.

ZOOSYSTEMA • 1998

Guinot D. & Bouchard J.-M.

Table 1 . — Diagramatic représentation of various modes of abdominal holding. (In white; articles of appendages involved; stippled

zone: abdominal part: in grey: sternal sonnte 4). A simple juxtaposition wlthout any différentiation nor projection; B, juxtaposition by

a projection, without a notch on abdomen: C, coaptation by engagement involving a differentiated structure togethar wlth a notch on

abdomen, D-F various modes ot overhanging; G, H, overhanging and coaptation by engagement; I. coaptation by engagement

involving a differentiated structure togetherwth a notch on abdomen; J juxtaposition By a differentiated structure, without a notcli on

abdomen. The homolfd press-button (coaptation by assemblage), présent in ail généra. i$ represented by tig. K

634

ZOOSYSTEMA • 1998 -20(4)

Abdominal holding Systems (Brachyura)l

Table 2. — Modes of abdominal holding and various involved appendages in the généra of the family Homolidae.

Letters refer to table 1. The shape of the differentiated structures (spine, granule, tubercle, or others) was not given. Letters in brac-

kets indicate a nonfunctional structure. The coxa often has two distinct zones thaï may show a different mode of retaining. (Data for

the genus Gordonopsis are from Guinot & Richer de Forges 1995)

GENUS

Mxp2

coxae

Mxp3 articles

Thoracopods

coxae of

coxae

basis

ischion

Homola

Paromolopsis

(J)-D

(J)

Paromola

l-H

Moloha

l-F

Latreillopsis

l-F

(J)

Dagnaudus

B-D

llhopsis

l-G

Homolochunia

(E)

(A)

Yaldwynopsis

l-D

Hypsophrys

J-H

l-E

Gordonopsis

(1)

Homologenus

(D)

Homolomannia

C-G

(A)

Homolax

J-D

J-A

prominences used as projections overhanging the

uropods, instead of the acamhodromiid, parady-

nomenid and [pro parte) dynomenid coxal pro¬

jections, would he very signilicant.

SUPERFAMIIY HOMOLOIDEA de Haan, 1839

The superfamily Homoloidea has long been asso-

ciated with the Dromiacea, and certain authors

(Hartnoll 1975; Stevcic 1981; Bishop 1986) hâve

continued to subordinare Homoloidea to the

Dromiacea. But, at the présent tinte, the mor-

phological and spermatological data support the

proposition lhat the Homoloidea should be

rentoved from the Dromiacea and, ai a higher

level, their inclusion in a single grouping, the

Podotremata (Guinot et al, 1994; Guinot S l

Richcr de Forges 1995). Before the establishment

of the Poupiniidae, the I lomoloidea were charac-

terized as having the P5 dorsal, reduced in length

and subchelare or chelate (with the exception of

certain latreilliids). Discovery of Poupinia hirsuta

Guinot 1991, with P5 dorsal and movable, redu¬

ced in width but not in length and not subcheli-

form, changes this définition. In poupiniids the

P5 are relatively unmodifted but are cfearly diffe¬

rent from the preceding legs. So the homoloid

character of P5 different fioni P4 (instead of

similai P4 and P5 in the Homolodromüdae and

most Dromiidac) remains a valuable criterion.

The family Homolidae is charactei ized by a dor¬

sal linea bomolica , which is absent in the two

other families Latreilliidae and Poupiniidae.

F ami iv Homolidae de Haan, 1839

In homolids the abdomen is long, and ihe telson

engages between the coxae of third maxillipeds

and sometimes between second maxillipeds,

which in a way clasp the posterior part of the

pleon These crabs are remarkable due to the co¬

existence of two types of retaining and locking

mechanisms (Guinot 1979a; Guinot & Richet de

Forges 1995). On the one hand, a retaining Sys¬

tem: seyeral projections on the coxae of varjous

appendages (Mxp3, PI through P.3) are involved,

often acting sintultaneously (Tables 1,2). On the

other hand. a locking mechanism: differentiated

structures are présent on the sternal surface, with

a pair of spécial prominences (on sternite 4) fit-

ting into a pair of sockets in the internai part of

the abdominal segment 6.

Structures ou appendages

Mosc homolids immobilize their abdomen by

using either a deep dépréssion on the coxae that

ZOOSYSTEMA • 1998 -20(4)

635

Guinot D. & Bouchard J.-M.

Fia. 9. — Retainlng apparatus in Homolldae and Latreillïïdae; A. Homologenus /ewïGuinol et Richer de Forges, 6 10.2 ■ 6,5 mm (cl

without roslrum, cw wlthout latéral splnes), paralype, Cheslertleld Islands (MNHN-B 19883) B, Homolochuma kullarOrittln erBtown,

d 26.5 x 23 mm (cl withour roslrum), New Caledonia (MNHN-B 16662), wilh détail of cm and homolid press-button: C, Homola

ranunculus Guinot et Richer de Forges, ri <12 6 35.5 mm, paratype. New Caledonia (MNHN B 20272); O, Latreillla ail. valida de

Haan, <? 13.5 x 8.3 mm, New Caledonia, Hallpro 1 , sln CP 851 (MNHN), wilh detail ol tha homotrd press-button, Abbraviations,: a5,

a6, abdominal segments 5,6; cxl, cx2. c*5 coxae of Pi, P2, P5: d s,, slerno-coxal dépréssion: g. female gonopoie, h.b,, homolid

button; p, pénis; t, telson; 1, 2, 3, 4, sternites 1-4. Setae were not (igured. Scale bars: A, C, 5 mm; B, D, 1 mm.

636

ZOOSYSTEMA • 1998 *20 (4)

Abdominal holding Systems (Brachyura)

corresponds with complementaty surfaces on rhe

abdominal margin, or diverse structures (such as

granules, tubercles, spines, spinules, groups of

similar structures, or a sliding surface). The struc¬

ture situatcd on the coxae of Mxp3 correspond ro

the posterior part of telson, chose on PI coxae to

abdominal segment 6, chose on P2 coxae to

abdominal segment 5, and tliose on P3 coxae (a

structure probably nonlunctional) to abdominal

segment 4, respectively. These structures may fit

into complemenrary parts of the abdomen or

overhang it. Tables 1 and 2 summarize the action

of the various appendages involved, with rheir

respective modes ol coaptation,

Some exemples are briclly reviewed here.

In Homologenus A. Milne Edwards, 1888

(Fig- 9A), probably very primitive, the cxtrcmity

of the telson engages betwecn the bases of the

Mxp3. which props up. In addition, a remarkably

long and spinose prominence on PI coxa com¬

plété!)' overhangs the telson. The spines on P2

and P3 Coxae seem ineffective. In Lamoha Ng,

1 998 (= Hypsophrys Wood-Mason, 1892) the tel¬

son is propped up by Mxp2, overhung by Mxp3,

Fig. 10. — Homolid butlon and socket In Homolidae and Latrallllidae, A, C, Moloha majora (Kubo), <t 51 x 38 mm, Sagami Bay

(MNHN-B 24798); A. thoracic sternum withoul abdomen, C. ventral surface of abdomen; B, Lalreillia valida de Haan, cl 16 mm. Japan

(MNHN): thoracic sternum wlthout abdomen; D, Homola orientais Henderson, 6 31.6 -; 25.2 mm, Frenob Polynésie (MNHN-

B 22385): photographs with the scanning électron microscope ol the righl homolid butlon. internai side view. Abbreviations: a6. abdo¬

minal segment 6; cxl, cx2, coxae of PI, P2; h.b., homolid button; mxp3, exlernai maxilliped; so, socket; t, telson; 4, stermte 4. Scale

bars: D, 100 pm.

ZOOSYSTEMA • 1998 ■ 20 (4)

637

Guinot D. & Bouchard J.-M.

and every articular condyle of PI -P4 (more deve-

loped on P1-P2) on die sternum lus on a com-

plementary part of ihe abdominal margin.

Homolomannia Ihle, 1912 (Fig. SC) is characteri-

zed by a System that is c.hielly provided by the

internai margin of thc P2 and I’3 bases, which

cover part of the telson. Homolochunia Doflein,

1904 (Fig. 9B), an advanced genus with the last

pair of pereiopods rerminating in a rrue subchela

(Guinot & Rieher de Forges 1981), is characreri-

zedby a marked coaptation berween the posterior

part of the abdomen and the sternum, specially ar

the level of Mxp3 and Pl ; the Mxp3 coxa beats a

strong projection chat fits a cotnplementary

part on the telson; an extension of the Pl coxa

overhangs pleomere 6; there are no spinules on

the other appendages. In Homola orienta lis

Henderson, 1888 (Fig. 8A), H. ranunculus

Guinot et Rieher de Forges, 1995 (Fig. 9C) and

Paromota bathyalis Guinot et Rieher de Forges,

1995 (Fig. SB), die appendages involvcd are simi-

lar ( Fable 2).

Sternal structures

The latero-external borders of the abdomen of

homolids reacli the proximity of the articular

condyles of the coxaeof Pl on the corresponding

sternite, the srernite 4. In front and very close to

this condylus there is a particular sternal struc¬

ture, very prominent, higb. elongated, and not

smooth under the binocular; the pair of latéral

prominences of somite 4 fus into a pair of sockets

hollowed on abdominal segment 6 at its postéro¬

latéral angles [Fig. 10A, C; Moloha majora (Kubo,

1936)]. The sternal différentiation appears mar-

kedly serrulate, spinulate or tuberculate, with

complex microstructures (Fig. lüD) that explain

the strerigth of the locking mechanism.

The abdoinina! structures of Homola were called

crans de rétention by Pérez (1928b: 649; 1929;

1149). The whole apparatus was named bouton-

pression homolten (homolid press-button) by

Guinot (1978a, 1979a). This paired sternal Sys¬

tem, observed in ail homolids, latreilliids and

poupiniids, appears as a unique innovation of the

superfamily Homoloidea.

Family LATRP.iu.MDAE de Flaan, 1840

The sternal plate of latreilliids is broader than in

homolids and is not completcly covered by the

abdomen in males; the telson enters befween the

bases of maxillipeds. In the tewspeeimens exami-

ned. the Mxp3 coxae are never equipped with a

spine but, vvhen closed together, seem capable of

retaining the telson at its dp. On sternite 4 a pair

ol very high, cristiform, markcdly dcnticulated

and sometimes acutc projections, with a complex

apical texture, fus into a pair of sockets in the

postero-lateral angles of pleomere 6. In the

latreilliids in which the sternal plate is only a litt-

le broadened, as in latreillia valida de Flaan,

1839 (Fig. 10B), the sternal structure is positio-

ned near the latéral margin of thc plate, as obser¬

ved in the family Homolidae. But in the

latreilliids in which a larger part of the thoracic

sternum becomes lateralLy exposed and with a

broad sternite 4 extended in front of chelipeds,

the sternal structures protude nearer the médial

part of the plare (Fig. 9D; see also I,atreillia man-

ningi Williams, 1982, fig. 2d). It is noteworthy

that the sterno-coxal dépressions are completely

absent in ail latreilliids.

The family Latreilliidae has the sanie press-but-

ton apparatus as the family Homolidae. The

homolid press-button seems to remain the only

effective mechanism in latreilliids; the bases of

the pereiopods, far froni the abdominal margins,

are unused.The press-button in larreilliitls proves

to lie very effective, and it is definite that the sur¬

face of the dcnticulated hutton possess micro¬

structures that improve the mechanism.

Family Poutinudae Guinot, 1991

Males and females of Poupinia hirsuta Guinot,

1991 bear two lcinds of Systems, on iimbs and on

the sternum, which appareiltly operate simuita-

neously (Fig. 8D).

Two very acutc spines projeta front thc coxa of

Mxp3 and clearly overhang the telson, which

pénétrâtes far berween the maxüliped bases.

These spines are truly effective but require that

the Mxp3 move close together. A spinule located

at the base of the Pl coxa, however, appears

insufBcient to be opérative. A pair of ornamented

sternal prominences lies on the sternal plate (ster¬

nite 4), near the articulation with chelipeds and

fits into a pair of deep sockets hollowed on the

internai surface of abdominal segment 6.

638

ZOOSYSTEMA • 1998 -20(4)

Abdominal holding Systems (Brachyura)

The poupiniids, distinctive because of their ovoid

carapace, their P5 not adapted for carrying beha-

viour or (or grasping, and their high gill couru,

possess thc samc “homolid press-button” as

homolids and latreilliids.

Discussion

Projections on the basal parts of appendages arc

used in homolids (on Mxp3 and PI through P.3)

and poupiniids (on Mxp3), together with the

homolid press-button. In larreilliids, in which the

sternum is broadened, only the carin.ue and ser-

rulate prominence on sternite 4 seems to be used-

A significant resuit of our investigations is that

the “homolid press-button” represents a synapo-

morphy for thc homoloids, supporting thc inclu¬

sion of Homolidae, I atreilliidae and Poupiniidae

in the same group, the (lomoloidea. As a struc¬

ture, situated on the sternal plate, and as a

mechanism, rhe “homolid press-button" is simi-

lar to the typical press-button in eubrachyuran

crabs which is alwavs found on sternite 5. mstead

of sternite 4 as in homoloids. We agréé with Pérez

(1928b, 1929) who concludcd that Homola can-

not represent the passage betwecn dromiids and

true Brachyura, despire the fact that the socket

occupies the same pleomere, the sLxdr, as in the

Heterorremata-Thoracotrernuta assemblage.

Similar to rhe Dromiacea, homolids and poupi-

niids hâve deep sterno-coxal dépressions insidc

which the P2 and P3 coxae may slide. For us,

absence of sterno-coxal dépréssions in larreilliids,

which présent more advanced features (especially

along with a wider sternum) than homolids and

poupiniids, might be corrclatcd with the fact that

the pereiopods hâve no rôle in abdominal réten¬

tion.

SUPERFAMILY RaNINOIOEA de llaan, 1839

d’he superfam’dy Raninoidea, which contains the

single family Raninidae, or Irog crabs, was subdi-

vided into several sut)tamilies and assigned to the

Podotremata (Guinot 1978a, 1979a, 1993b).

The basal relationship ot raninoids to the hetero-

treme-thoracotreme assemblage, indicated by

some morphological features, spermatological

analysis of Ranirta ranina (Linné, 1758)

(Jamieson 1989) and molecular sequences

(Spears et al. 1992), was partly refuted by the

spetmatozoal uhrastructure in rwo other raninoid

taxa (Jamieson rf d/. 1994; |amîeson 1994). The

distinctive morphology of thc raninoids (elonga-

te body, thoracic pleura largcly exposed by rcduc-

tion ot the branchiostegite, elongate and rriangu-

lar buccal trame, spécial rtspiratory apparalus,

flattened propodi and dactyli ot pereiopods)

seems ro reflcct a high degree of specialization for

burrowing, that lias characterized the group

throughout its long géologie history, since its

appearance in rhe Harly Albian.

Tn raninids the sternal plate is narrow, xometimes

linear, and lacks sterno-coxal dépréssions, whiie

the abdominal segment 6 lacks appendagelilce

uropods (for the case of the Lyreidinae, see

below), the abdomen is very short and only part¬

ly and loosely flexed (leaving exposed a large part

of the thoracic sternum anteriorly), male pleo-

pods are short, and the P2 through P5 coxae lie

close together,

Family RANINIDAE de Haan, 1839

No retaining apparatus was observed in

Raninidae cxcept in thc subfamily Lyreidinae

Guinot, 1993. The extant members of thts subfa-

mily are distinguished m particular by an abdo¬

men that flexes segments 4-6 and entircly covc-rs

the thoracic sternite 6, and by the thoracic ster¬

num (Guinot 1993b: 1326, 1330, ftg, 4). Ail of

the examined représentatives of the généra

Lyre i dus de Haan, 1839 and Lysirude Goeke,

1986 hâve a pair of strong elongate projections

f'rom sternite 5 (“pterygoid processes" of Bourne

1922) that firrnly lits into a pair of deep sockets

in the latero-posterior extended angles of abdo¬

minal segment 6 (Fig. 11). The base of the pro¬

jection starts ar the level ol the articulât condylus

of P2 on sternite 5. The rip of the projection,

more or less strongly reculved and distally hook-

shaped, beats an additional subdistal hook. The

whole structure acts as a safety catch. The confi¬

guration may remain effective in ovigerous

females, even with a large egg mass: the projec¬

tions are always présent, but the hooks become

more or less blunt.

In / y indus brevifrtms T. Sakai, 1937 (Fig. 1 IA, B)

thc apparatus is somewhat similar to that of

L. tridentatus de Haan, 1841 (Guinot 1979a:

128, pl. 22, figs 6, 7; 1993, figs 6, 7). In L. ste-

ZOOSYSTEMA • 1998 - 20(4)

639

Guinot D. & Bouchard J.-M.

nops Wood-Mason, 1887, the projection also

ends in rwo hooks, distal and subdistal, similar to

those of Z, brevifrons and L. rridcntatus. A matu¬

re female, however, from the Philippines (36.4 X

21.9 mm, Musorstom 1980. stn 71) identificd bv

G. D. Goeke as L. stenops, was observed with tv?o

subdistal hooks in addition to the distal hooked-

tip.

In Lysirude, at least in L. channeri (Wood-Mason,

1885), the sternal projection is shorter rhan in

the examined Lyreidus species. It lies entirely on

the sternal surface. In juvénile fetnales and adult

males of Lysirude channeri (Fig. 11C) the projec¬

tion is lifted up ai the hooked tip and bears a sub¬

distal hook, giving an effective locking mecha-

nism. In mature temales (Fig. 11 D) the projec¬

tion becomes entirely applied on the sternal sur¬

face and becomes gradually stuooth (sometimes

cxcept lor a smail dcnticlc) and more sctose with

size. The abdominal Socket is obsolète or absent,

and locking is no longer possible.

Feldmann & Wilson (1988: 478) noriced that

“the males, in mature Récent species of Lyreidus

and Lysirude, possess straight bladed, raiscd

flanges; vvhereas the female possesses notably

reduced, ourwardly curved projections'. We

observed that the locking upparatus is sexually

dimorphic in lysirude channeri , but not so clear-

ly in Lyreidus species thaï vvere examined. The

dimorphic State noted by T. Sakai (1937: 170) in

Lyreidus tridentatus conccrns abdominal seg¬

ment 5 and not the rhoracic sternite 5 (sec

Feldmann & Wilson 1988).

Discussion

Only in Lyreidus and Lysirude is the abdomen

situated between P3-P4, ftxed against the ventral

surface, and pleomerc 6 bears a Socket. The diffé¬

rentiation which ensures rétention is located on

sternite 5, as the prominence in certain advanced

dynomenids (Figs 6D, 7D) and the typical press-

button of eubrachyran crabs (see Figs 13-17,

20-24).

A complété re-examtnaiion of ail the fossil rani-

noids, which diverstfïed in the Cretaoeous and

flourished in the Eoccne, should enable further

résolution of the abdominal holding apparatus,

especially with the examination of the lyreidines.

On the preserved sternal plate the processes ari-

sing from sternite 5 and the presence of hooks

must be checked in fossil crabs arnibuted to

lyreidus (L. succedamts Collins et Rasmussen,

1992, L. rosenkrantzi Collins et Rasmussen,

1992, and L. bupinulatus Collins et Rasmussen,

1992, ail front Upper Cretaceous-Lower Tcriiary

from West Gteenland). Rogetis orii Berglund et

Feldmann, 1989, Iront Lower Middle Eocene of

Oregon, distincrly shows a pair of projections: we

wonder whether they putyed a similar rôle to that

in extant l.yreidinae. The précisé configuration of

processes arising from sternite 5 would be noted in

another fossil raninoid subfamily, the

Palaeocorystinae Lôrenthey in Lôrenthey &

Beurlen, 1929, especially in Notopocatystes McCoy,

1849, the oldest known genus of rhe superfamily

Raninoidea (Collins 1997).

Superfamily Cyci.odorippoidea Ortmann,

1892

We use this category for convenience to regroup

the familics, Cyclodorippidac, Cymonomidac

and Phyllotymolinidac, despire possible paraphy-

ly (M. lavarcs pers. com.). As in Homoloidea

and Raninoidea (and ail Eubrachyura) pleo-

meres 3-5 lack vestigial pleopods, and abdominal

segment 6 lacks (dorsal or lobiform) uropods. A

character shared by ail Cyelodorippoidea is the

wide sternal plate. For each pair of pereiopods the

respective bases of the coxae are very far apart so

that there is no contact betrween the margin of the

abdomen and the bases of the legs. A very short

and more or less excavated sterno-abdominal

cavity is présent, with ils complété spart occu-

pied by stout pleopods. Complementary parts

beiween the borders of both this cavity and the

abdomen secure it in place. Additionally, a pecu-

liar holding may occur, that is different in the

familles Cyclodorippidac and PhyLlotymolinidae.

Family Cyclodorippiuak Ortmann, 1892

A sterno-abdominal cavity, which is deeply hollo-

wed but very short, is characreristic of the family

si net it occupies only the posterior part of the

thoracic sternum (stérilités 6-8). The short abdo¬

men, characteri/ed by the presence of a long pleo-

telson, is beru at right angles, its posterior haif

being applied against the sternal surface. In addi¬

tion, at the level of sternite 6, the latéral border of

640

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

che cavitv is m.trked by a groove into which the

thin borders of clic- pleotelson can engage by

moving l'urward. It was named “sliding System"

{système à glissière) bvTavares [(1994: 210; 1996,

fig. 9A-C, E; 1998: 116, figs 7A-C. 8 for

Clythrocerus nitidus (A. Milne Edwards, 1880)J.

There is a coaptation by engagement of the bor-

ders of both sternum and pleotelson, but without

any sockets.

Family Phyllotymounidae Tavares, 1998

There is a relatively shallow sterno-abdominal

cavity, without well-dcfmed borders. The abdo¬

men is short, the telson only reaching somite 5.

The sternite 6 bears a pair of oblique prorni-

nences in Pbyllütymolhutm Tavares, 1993 and a

pair of granular ridges in Genkaia Miyaké et

Takeda, 1970, structures with which the enlarged

base of the telson forms a block System, named

Fig. 11. — Locking apparatus îri Lyreidinae (Ramnîdae): A. B. Lyreidus brevifrons T. Sakaî, c' 49.4 x 23.4 mm, New Caledonia

(MNHN-B 24319): A, ventral surface with abdomen llexed In locking position; 8, without abdomen. C, D, Lysiruds channeri (Wood-

Mason), Philippines (MNHN-B 11562), C. ,t 25.5 ■ 17.4 mm; D. ■> 33.720.5 mm; photographs with the scanning élection micro¬

scope to show the pairs ol projections with hooks well developed in a male, blunl or lost in an old female. Abbrevialîons: a6, abdo¬

minal segment 6; cx2 cx3, coxae of P2, P3, g, gmglymus lor articular condylus of P2, h, hook; pr, projection with hooks; I. telson;

4, 5, sternites 4. 5. Scale bars: C, 100 pm; D, 1 mm.

ZOOSYSTEMA • 1998 -20(4)

641

Guinot D. & Bouchard J.-M.

système à butoir by Tavares (1994: 208, 209;

1998: 116, fig. IB).

Family Cymonomidae Bouvier, 1897

There is only a shallow sterne-abdominal cavity,

the stérilités 6-8 being only slightly concave, and

the abdomen is verv short. No spécial structure

for abdominal holding vvas observed (Tavares

1994). As in the Cyclodorippidae, the abdomen

is bent in its middle at rigbt angles forming two

sections. The posterior pleomeres thus remain

compietely applied against the sternum (M.

Tavares pers. corn.). It is reminescent ol the cases

of Hypoconchü nrcuata and H. panamémis in

which the abdomen is bent at right angles in the

posterior part of segment 3 (Fig. 11)).

Discussion

The two retaining Systems found in the cyclodo-

rippids (coaptation by engagement) and phyllo-

tymolinids (block System) were one character

used in the cladistic analysis by Tavares (1994)

that allowed the élection of the Phyllotymo-

linidae as a new Family (Tavares 1998).

In contrast to ail Podotrerrtata previously mentio-

ned, in the Cyclodorippoidea the sternal plate has

acquited a “normal” widih and is grooved by a

true stemo-abdominal cavity: the coxae of the

appendages cannot be involved in the rétention,

and sterno-coxal dépréssions are absent. In the

Cyclodorippidae there is a coaptation by engage¬

ment, whereas in Phyllocymolinidae an innova¬

tion is the cmergence of a pair of prominent ster¬

nal structures (on sternite 6, a vert' posterior ster-

nite). In borh cases no cortesponding sockets

exist on abdomen and it is only the telson which

is involved by slipping along the sternal grooves

(Cyclodorippidae) or by blocking at the sternal

prominences (Phyllotymolinidae).

Superh,amily Dakoticancroidea Rathbun,

1917

Family D.akoticancridae Rathbun, 1917

The exclusively lossil family Dakoticancridae,

from the Cretaceous of Norrh America, was

confirmed as belonging in Podotrentata because

of the large spermathecal orifices accompanying

the coxal female gonopores on P3 (Guinot

1993b). The dakoticancrids hâve a very broad

thoracic sternum, an abdomen without uropods

(at least in dorsal view), and a well-excavated ster-

no-ahdominal cavity, On sternite 5, along the

border of this cavity, a set of granules was obser-

ved at the level of abdominal segment 6, but

rather similar to that on the following sternites

( ibid.y figs 6, 8): their rôle in abdomen holding is

questionable.

HETEROTREMATA Guinot, 1977

The Heterotremata and Thoracotremata share a

sternal location of the female gonopores, with the

innovation of the vulvae openingon sternite 6 in

direct communication with the séminal récep¬

tacle. There is now only one female orifice for

reproduction. Anorher strong synapomorphy of

the heterotreme-thoracotreme assemblage is the

morphology of the firsi male pleopod, which is

compietely closed and provided with two distinct

basal foramina for the introduction of the pénis

and the second gonopod (Guinot 1979a: 239).

The monophvly ol the non-podotreme brachyu-

rans is well supported.

The Heterotremata generally show a coxal loca¬

tion of the male gonopores on the P5 coxae. But

ment bers of sorne tamilies (in the Dorippidae,

diverse Dorippinae and ail Ethusinae, and mem-

bers of the Leucosiidae and Goneplacidae) and

certain advanced heterotreme families (flymeno-

Sornatidae, Pinnotheridae, Cryptochiridae) exhi-

bit a coxo-stetnal organisation with the pénis

located in a sternal groove or potential channel,

and embedded under the sternal plates

(Guinot & Richer de Forges 1997). This requires

providing a more précise définition of the

Heterotremata, in contrast to the Thoraco-

Fig. 12 - Configuration of ventral surface in the Corystidae;

A, B. Corysles cassivelaunus (Pennant): A, î 35.2 x 24.6 mm,

Bay of Biscay (MNHN-B 5381); buttons absent; B, ? 31 x

24 mm, East Channel (MNHN-B 5385): vulvae not covered by

abdomen: C, Nautilocorystos ocetofus(Gray). L 39 5 » 34 mm.

Cape ot Good Hope (MNHN-B 3924S): butions prescrit (broken

on one sidel bul M' lions pleontefe 8, D, (Irtniozo hhxirnis Gray,

5 26 x 21.3 mm. Moluccas (MNHN-B 1/168): note lhe very

stiort abdomen; E Psouctocoryste s slonrlus (Pooppig) ' 50 x

44 mm. Chiie (MNHN-B 5407)' vulvae noi covered by abdomen.

Abbreviations: a, abdomen; b, buiton ol lhe press butlort: g. G1 :

m.l., médian line, s.s. sternal shleld (sterniies 1-3). I, telson;

v, vulva; 4, 5, 6. 7, sternites 4-7.

642

ZOOSYSTEMA • 1998 - 20(4)

ZOOSYSTEMA • 1998 *20 (4)

Guinot D. & Bouchard J.-M.

tremata, which attempts a better distinction

concerning the patterns lor male ducts.

Family Coryslidal Samouelle, 1819

In Corystes II ose, 1802, with a narrow, rather liât

and horizon tally metamerized thoracic sternum

(Guinot 1979a: 81, 132, pi, 9, fig. 1), and in

Pseudoctnystes 11 Milne Edwards, 1837 (Guinot

1979a: 132, fig. 20B) the bottom of the sternal

plate ts longirudinally divided by the médian line,

which is complété Front the sternites 4 through 7

(Corystes, Fig. I2A, B), or interrupted ( Pseudo-

corystes, Fig, 12E). There is an undefined sterno-

abdomina! cavity. The male abdomen is veiy

short and has the rvvo first plcomeres in a dorsal

position: its pleomere 6 lies posteriori}' and does

not go beyond sternite 7. Retaining structures are

absent in ail specimens examined. Abdominal

sexual dimorphism is weak. lt is noteworthy thaï

in these rwo généra the vulvae remain uncovered

by the abdomen which is very short, even in

females. Exposed vulvae (Fig. 12B, E) are rarely

observed in brachyuran crabs (Guinot 1979a,

pl. 25, Figs 4-6).

In Gonieza Gray, 1831 the sternal plate is very

narrow, well metamerized and longitudinally

grooved by the médian line that extends front

sternites 4 through 8, and the sterno-abdomin.il

cavity occupies a posterior location, being excava-

ted only on sternites 7 (partly) and 8. The abdo¬

men is very short It is only a tuft of long and stiff

setae front the sntall tclson, which helps close the

posterior part of the cavity and shelcers the sexual

pleopods. In G . bicornis Gray, 1831 (Fig. 12D)

the abdomen is spécial, with the somites laterally

expanded (as with pleural prolongations). At the

level of sontite 6, the latéral part bends vertically

and closes the cavity on each side. Structures of

the press-button type are absent. In females ol

Gomezü the abdomen remains very short and

laterally expanded, there is no true sterno-abdo-

minal cavity (only a lowering of the somites) and

the vulvae cannot be covered by the telson, which

lies far backwards. The egg mass is unprotected.

The configuration in Jouas Jacquinot et Lucas,

1853 is similar, with a relatively short abdomen.

The long first pleopods, however, are exposed in

grooves ol the sternal plate, outside the sterno-

abdominal cavity. Only a damaged male specimen

of/. distincte! (de I bon, 1835) was examined, and

no locking structures were observed,

Only two incomplète males (a suhadult and an

adtih with a damaged abdomen) and a juvénile

helonging to the genus Nautilocoiystes H. Milne

Edwards, 1837 (Guinot 1979a: 133, fig. 20C,

pl. 23, fig. 5) were examined. Its general organi-

zation (Fig, 12C) is similar to the preceding géné¬

ra: a sternal plate hoiizontaily metamerized and

longitudinally grooved, and an undefined sterno-

abdominal cavity. Jn comparison to Corystes, the

abdomen is longer, but the posrcro-lateral angles

of pleomere 6 remain far from sternal suture 5/6.

A pair of acute, reversed liooks is positioned at

die base of sternite 5, very dose to suture 5/6. No

sockets on the pleomere 6 could be observed.

There is such a large gap between the sternal

hooks and any part of pleomere 6 that contact

appears impossible.

Family Atklkcyci.ipae Ortmann, 1893

Rice (1981a) suggested that the family Atelecy-

clidae sensu lato was not a rnotiophyletic group.

Atelécyclus Leacli, 1814 (Guinot 1979a: 136, pl. 9,

figs 4, 5, pl. 23, fig, 9) has à very narrow and

medially grooved sierual plate and a weakly hollo-

wed sterno-abdominal cavity, Both parts of the

press-button svstcni are présent and functional. Iti

juvéniles and adule males (up to about 46 X

47 mm; fifteen specimens examined) of 'Aieleeyclus

rotundatus (Olivi, 1792) the developed tuhercles

and sockets form an effective locking rnechanism.

In the saine sample that was examined from the

Bay of Biscay, a male (35.4 X 36.8 mm; Fig, 13A)

was fourni with the rwo complementary parts

firmly assembted, rhus an effective locking mecha-

nism; and a slightly smaller male (33.9 X

35.2 mm; Fig. 13B) was fotind witltout an effec¬

tive rnechanism, the sockets in the postero-lateral

angles of pleomere 6 reaebing the suture 5/6 but

not the buttons. Ir looks as if the abdomen is too

short in rhis "abnormal” male.

In Peltarion Jacquinot, 1 847. about fifty indivi-

diKils of P. spinulosnm (White, 1843) were exami¬

ned. The sterno-abdominal cavity is excavated,

and the two parts of the apparatus arc wcll-diffc-

rentiated. There is a pair of sternal tubcrcles very

close to suture 5/6; sockets are defined on the

produced latero-posterior angles of abdominal

644

ZOOSYSTEMA • 1998 - 20(4)

Fig. 13. — Lockmg apparatus in lhe Atelecydidae: A, B. Alelecyclus rolundalus (Olivi). Bay ot Biscay (MNHN-B 5400)' A, cî 34 x

35.5 mm: buttons titting inlo sockets B, 3 33.9 •: 35.2 mm: buttons not fitting into sockets; C, Telmessus cheiragonus (Tïleslus), 9

35 x 46 mm. Japan (MNHN-B 5408): vulvae not covered by abdomen; note the notch on the edge ot pleomere 6, O, Trachycarcinus

alcocki (Doflem), 6 70.3 > 65.2 mm, Philippines (MNHN-B 11571): note the bulge on the side ot large cheliped and lhe right button

(at left) not fitting into Socket, Abbreviatlons, 36, abdominal segment 6; b. button ot |he press-button; cxl, coxae ot PI s.s., slernal

shield (sternites 1-3); t. telson; v, vulva; 4, 5, 6, 7, 8, sternites 4-8.

ZOOSYSTEMA • 1998 -20 (4)

645

Guinot D. & Bouchard J.-M.

segment 6, nor very deep, however. Locking is

possible but it does not seem very efficient, des-

pite the acuteness ol rhe buttons. In certain

males, either small (26.3 X 26.2 mm) or much

larger (53.9 x 56.5 mm and 54.8 X 57 mm),

there is a small gap between the pair of sternal

tubercles and the strongly pioduced postero-late-

ral angles of pleomete 6, i luit is enough to pre-

vent proper Fastening. Fhe two components of

the locking System are not lost in larger indivi-

duals, even if apparently not used, The figure

(Fig. 14A) shows a large male individual (54 x

57 mm; cl indicared by error as 52.6 mm in

Guinot 1979a; 135, fig. 2UD) in which no close

connection between the two parts of the appara-

tus was observed. The site at which the two parts

do not artach ro each other is variable. In females,

the coincidence between the two sternal and

abdominal parts exists in some individuals, but

the sockets do not seem to be defined. In females

front about 32 X 32 mm up, the buttons rend to

beconte smaller and disappear, The material in

the MNI1N identified as lî spinubsum shows a

strong vanability, notably concernîng the areola-

tion and granulation of the dorsal surface of cara¬

pace and the spLnularion of pereiopods. It would

be înteresting to verify wherher the différences

observed in the liaison between the two parts ol

the locking System might lie related to other mor-

phological features. It is noteworthv that

Peltarion displays various patterns, in patticular a

non-correspondencc even in young males, or an

asymmetry illustrated by a larger gap between the

two structures on die major cheliped side (a male

41.7x43.8 mm).

In Trachycarcinus Faxon, 1893, in which one of

the chelipeds becomes very large, the pair of

tubercles is siiuated in the middle of sternite 5

and the two components of the apparatus func-

cion well. But in two large specimens of T. alcoc-

ki (Doflein, 1903) (Fig. 13D) and of T. crosnieri

Guinot, 1986, a non-correspondence was obser¬

ved on rhe major cheliped side. The connection

only opérâtes on the minor cheliped side. This

asymmetry can be explaincd by the increased

development of PI muscles, which induces a

“bulge” of sternite 4 and modifies the general

organisation of rhe anierior part ol plastron, thus

displacing the tubercle on the side in question.

Family CHEIRACONIDAIî Ortmann, 1893

(= TELMF.SSINAU Guinot, 1977)

A familial tank is provisionally given here to this

small group, traditionally assigned to the

Atelecydinae within the family Atelecyclidae,

awaiting a révision of ail Corysroidea (Balss 1957:

1635; Guinot 1978a: 259; 1979a: 136, pl. 25,

Figs 1-3; 1989: 349; Rite 1981a; 293; Stevcic

1990: 33; Sasaki & Mihara 1993: 511). The

adult and zoeal features of Erimacrus Benedict,

1892 and Tebnesius White, 1846 do not obvious-

ly ally them witli any known group.

In both Erimacrus and Tc/messus, there is a relati-

vely narrow sternal plate, horizontally merameri-

zed and longitudinal ly grooved by the médial

line, with the buttons (very close to suture 5/6)

fïtting well inro marked sockets. In T cbcimgonus

(Tilesius, 1812) the large females (for example

35 x 43 mm), with devèloped pleopods, ahvays

bave acute buttons corresponding with sockets

that reniant visible but are partially masked by

setae. Most curions is the position ol the very

wide vulvac lving latéral ly on the exposed parts of

sternite 6, close to the P3 coxae, and appearing

even more exposed because of the strong notch

on the external margin ol pleontere 6 (Fig. I3C).

Family Thiipal Dana, 1852

The small genus Chia Lcach, 1815 (for ontoge-

netîc considérations, sec Ingle 1984) îs characte-

rized by a very narrow, transvefsally metamerized

and longitudinally grooved sternal plate, with a

parricularly long sternite 4, and by an abdomen

that fi Us rhe whole space between the legs front

P3 rhrough P5, with the telson lving only in a

small dépréssion on sternite 5. Tbete is a pair of

prominenccs, with an acute tip, close by sutu¬

re 5/6. The sockets are shallow but they seem to

be effective (Guinot 1979a: 83, 135, 136,

fig. 20A).

Family CancriDAK Latreille, 1803

The thoracic sternum is not very enlarged in

Cancer Linné, 1758. Ail the sutures of the ster¬

num are continuous and parallel, the médian line

extends until il te middle ol somite 4, and there is

a sterno-abdominal tuviry. 7 he button is relative-

ly small but acute, and the socket is deeply

646

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

Fig. 14. — Loclgog apparatus in AtelecycUdae (A, Peltarian Jacquinot) and Belliidae (B, C, Se/ffa H Mttne Edwards); A Peltarion spi-

nulosum (While). ' 54 57 mm, Cape Horn (MNHN-B .5405) die two parts of lhe lockmg apparatus are not in conespondence, but

sternal buttons and sockets are présent 1 After Gumot 1979a. fig. 20C, specimen mdtcated as cl 52.6 mrri); B. C, Bulha picta H. Milne

Edwards; B, i 50.5 / 45.5 mm, nolotype. Peru (MNHN-B 1 1 thoradc sternum and abdomen, withoul coincidence between the but¬

tons and the sixth abdominal segment: C c 29 > 25.4 mm. 'Tahiti'' (erroneous origin) (MNHN-B 2); note in this young female the

coincidence between the buttons and the sixth abdominal segment, and outtons locatad close ta the e.xpossd vulvae. Abbreviations:

a6, abdominal segment 6; b, button; s.a., sterna-abdominal cavrty t. telson, v vutua 1-6, sternltes 1 tbrough 6; 3/4, 4/5, 5/6, sternal

sutures 3/4, 4/5 and 5/6. The parts located under abdomen are drawn wlth a continuous line. Setae were not ligured.

ZOOSYSTEMA • 1998 -20 (4)

647

Guinot D. & Bouchard J.-M.

hollowed and lined by a thick ridge on the poste-

ro-lateral angles of pleomere 6.

Family Belliidae Dana, 1852

Inthe burrowing Beilia H. Milne Edwards, 1848

(Guinor 1976: 48, ligs 10B, 11A; 1979a: 134),

there is a relative!)' vveak sterno-abdominal cavity.

The large holotype of Beilia picta H. Milne

Edwards, 1848 (male 50.5 X 45.5 mm)

(Fig. I4B) bears a pair of small prominences at

the posterior part ot the sternitc ncar suture 5/6;

they cannot bc rcached. however, by the well-

defined sockets cil pleomere 6, which lie poste-

riorly. This would hâve scemed an aberrant indi-

vidual but Rathbun (1930: 175, pl. 79, fig. 3)

figured the saine configuration in a smaller male

(cl 28.5: esrimnted cw 25.3 mm, USNM 22066)

in which, there was also no coincidence between

the parts, the abdomen looking “too short”. In

another respect, B. picta exhibits a spécial confi¬

guration in females. The vulvae are completely

(only partly in small individualsj uncovered by

the abdomen and remain exposed on sternire 6,

latéral to pleomere 6 In two females (24,8 X

22.3 mm and 29 x 25.4 mm; Fig. 14C) rhe liai¬

son is normal. But in a large ovigerous female

(39.5 x 36 mm) the prominence is replaced wirh

a scar close ro the large vulva and the fitting is no

more effective (Guinot 1979a, fig. 39A-C). It is

noteworthy that the structures lock into each

other in a female of cw 22.3 mm but not in a

male ol cw 25.3 mm, but this can be expiained by

the markedly short male abdomen in Beilia picta

(Fig. I4B,C).

In the other généra of the family Belliidae,

Corystoides Lucas, 1844, Acanthocyclus Lucas,

1844 and Heterozius A. Milne Edwards, 1867,

the few examined specimens show a ‘normal”

configuration as well in the locking apparatus as

in the location ot vulvae (Guinot 1976: 50-55,

figs 10A, C, D, 11B-D: 1979a: 134).

Family Orithyiidae Dana, 1852

Orithyia Fabricius, 1798, monospecific with

O, sinica (Linné, 1771), was separated front the

family Galappidae by Ortmann ( 1892: 555, 559),

who was the first author to suggest a familial sta¬

tus for Orithyiidae within the Calappinea. Alcock

(1896: 138) subdivided the Calappinae in two

alliances, Calappoida and Orilhyioidu. thle

(1918: 178) recognized the relationshipx linking

Orithyia to the Dorippidae (better than to the

Galappidae). On the basis of thé larval stages des-

cribed by Hong (1976), Rico (1980; 317;" 1981a:

293) indicatcd the présence of a combination of

characters unknown in any other identified larv'ae,

although shovving some si milan fies with dorip-

pids, but perhaps “of little si gni fi can ce”. For a

long tinte, most carcinologists considercd the sub-

faniily Otithyiinae as a component of the

Galappidae inside the ancicnt Oxystom.ua (Balss

1957: 1611;T. Sakai 1976: 127, 143; Dai & Yang

1991: 101, 113). A separate family, Orithyiidae,

distinguished from the Galappidae, was resurrec-

ted by Guinot (1978a: 254). Bcllwood (1996:

185, fig. 4) défi ned a clade Dorippc + Otithyiinae

supportée! by nine synapomorphies and placed

the family inside the Dorippoidea, but many cha-

ractcrs, such as dusse of the abdomen and the vul¬

vae, were not taken into accouru,

’Fhe peculiar configuration of Orithyia was descri-

bed by Guinot (1979a: 133, pl. 14, figs 7-9) des¬

pi te poor and incomplète material. Il is now pos¬

sible to document this with addition,si specimens.

The thoracic sternum is very wide and circular,

with ail the sutures interrupted and the presence

of a médial line on sternites 7 and 8; a large part

of sternite 8 is exposed. There is a long sterno-

abdominal cavity, its smooth surface contrasting

with the granular rcmainder ol the plate. The first

two abdominal segments are gathered together

between the strong coxae of the subdorsal I J 5, and

pleomere I does not seem ro be movable. In

males, the abdomen is very short and fills only the

middle of the sterno-abdnnnnal cavity, its rip not

reaching the suture 6/7, and the pleomere 6 lies

very far front sternite 5. Neither buttons nor soc¬

kets were observed in our large-sized males

(Fig. 15A), the short first pleopods of which can

be completely sheltered by the abdomen. This

observation needs to be verified in young males.

In (emales (Fig 15B), the abdomen is longer and

fills a slightly larger part of the sterno-abdominal

cavity, and vulvae are completely cxreraval on ster¬

nite 6. They remain exposed not becausc of the

abdomens shormess but because of their latéral

location on the plate, outside the cavity.

648

ZOOSYSTÉMA • 1998 *20(4)

Fig. 15. — Configuration ol the ventral surface in Orithyiidae and Dorippidae: A B. Orithyia sinica (Linné): A. i 83 6 •: 73.6 mm,

China Sea (MNHNI-B 11612): sterno abdominal cavily and very short abdomen; B, 9 64 x60 mm, without origîn (MNHN-B 93S): vul-

vae not coversd by abdomen: C, D, Medorippe lanala (Linné), 3 26.3 x 32.1 mm. Morocco (MNHN-B 19861): C, without abdomen:

D, witti abdomen in a flexed position, Abbreviations a6, abdominal segment 6, b, button ol the ptess-button. pl, pleopod; s.a., stet-

no-abdominal cavity: t, telson: v. vulva: 5/6, sternal suture 5/6: 4. 5, 6, 7 8, sternites 4-8

Family Dorippidae Macleay, 1838

SuBFAMILY Dorippinae Macleay, 1838

Thèse Heterorremata are characterized by rhe last

two pairs of legs being dorsal, reduced and pre-

hensile, whicli allovvs carrying behaviour (Guinot

et al. 1995). The sternal plate is very wide, espe-

cially at tlte level ofsomite 5, ail sutures are inter-

rupted, the sterno-abdominal cavity is rather

deep, and the abdomen does not reach sutu¬

re 4/5. The First two pleomeres are dorsal and

clasped between the P4-P5 coxae. A typical press-

button is présent (Guinot 1979a, Fig. 28), but

strangely rhe prominence appears in a concavity

of the sternal suture 5/6 (Figs 15C, D, 16A, R).

In the nunierous généra attributed ro the subfa-

mily (Holrhuis & Manning 1990, figs 21 h, i,

37c, 4lb, c, 43h, i, 45h, i, 50h, i, 55i, 56i-k), it

ZOOSYSTEMA • 1998 -20(4)

649

Guinot D. & Bouchard J.-M.

looks as il in both sexes, this suture is sufficiently

curved backwards to include tbe nodules, which

face tbe sockets in the postero-lareral angles of

abdominal segment 6. Tbe extremely acute pro-

minences and rbe deep sockets provide an effi¬

cient locking mechanism. We consider the

unique course of the sternal suture 5/6 as a syna-

pomorphy of the subfamily Donppinae (not in

the Ethusinae, sec- below).

Guinot ( 1979a: 139, pl. 25, figs S, 9) observed in

Medorippc lanata (Linné, 1767) the persistence of

acute buttons and defïned sockets in posr-

pubertal and even ovigerous females, wondering

whether the mechanism commues to be functio-

nal. In dorippine females (Fig. )6A) the sterno-

abdominal cavity is peculiar. The abdomen,

which partly remains in the prolongation of the

carapace, does not lie in an excavation on the

sternal plate but occupies a postcrior location

against the almo.sr vertical stérilités 7 and 8. In

conséquence, the components of the holding Sys¬

tem (ai the level where the pleomere 6 (aces the

eurve ol the sternal suture 5/6, a curve présent in

females as in the males) are not affected by the

broadening of the different structures and the

modifications which occur on the poSterior

région affer the puberty moult in females. The

vtilvae lie very close to the locking promincnces.

Perhaps only the tbickness of the egg mass and

the long pleopodal setae prevent locking. In an

ovigerous female of Nendnrippe aft. callida

(Fabncius, 1798), from Australia, a firm closing

was found usïng efficient buttons and sockets.

The reexamination of ail the dorippine species

and observations of live animais are needed.

Fig. 16. — Locking apparatus in Dorippidae: Medorippe lanata (Linné), Dahomey (MNHN-B 20528); A, 9 22.6 x 30.5 mm: B, 3

19.1 x 23.3 mm, both with an effective apparatus. Abbreviations: b, button; so, socket; s.a., sterno-abdominal cavity; v, vulva; 3/4,

4/5, 5/6, sternal sutures 3/4, 4/5 and 5/6. Triangle shows the curve of the sternal suture 5/6 where the button is located. Setae were

not figured. Scale bars: 1 cm.

650

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

SUBFAMI1.Y F.niUSINAL Guinoi, 1977

The ethusine su b family differs in having rhe ster¬

nal suture 5/6 normally directed, without rhe

curve seen in the dorippinex. fn Ethusa Roux,

1830 there is a strong prominence. fn Etnttsina

Smith, 1884 males of ail the species that were

examined show a longitudinal!)' elongated pro¬

minence, with a wide basis, resulting in a good

locking with rhe large sockets. In mature teniales,

with a cavity normally hoJlowed along the

somites, both components ol the apparatus are

well-developed and permit effective locking.

Discussion

The distinct course of the sternal suture 5/6 with

the locking prominence positioned inside a notch

is an exclusive dorippine characrer, rhat allows a

séparation into the subfamilie.s Dorippinae and

Ethusinae. In orher respects, the persistence of

the System in ovigcrous females constitutes a

synapomorphy of dorippids and erhusids, i.e. the

family Dorippidae.

Family Retroplumidae Gill, 1894

This small family, with two extant généra, was

placed in the catometopous crabs despite the

male gonoporcs opening on a tubercle at the base

of P5 (AJcock 1899: 78, 79, Ptenoplacïdae). The

group, sometimes placed ncar the Paliddae (Balss

1957: 1662), was transferred to Dorippoidca

with much réservation (Guinot 1978a: 151),

then raised to supratamilia! level, without any

connections with other brachyuran families (de

Saint Laurent 1989). Several rctroplmnîd fossils

are known, which were assigned to Ocypodoidea

(Glaessner I960; Via Boada 1982) and/or in the

Thoracotremata (Collins et al. 1994; Vega &

Feld matin 1992).

Guinot (1979a: 148, fig. 30C, F), who observed

only one female of a Retmpluma species, indicated

the peculiar locking apparatus of the family and

emphasized that it was effective even in mature

fentales. The faet that the tetroplumid females

retain a functional and very efficient mechanism,

even in mature stages, was discussed (ibid .: 157).

In the révision by de Saint Laurent (1989), who

had at her disposai abundant matcrial and establi-

shed a new genus and several new species, a typical

press-button was attributed to the retroplumids.

In both sexes the abdominal tnargins are coapta-

ted with thoseof the scerno-abdominal cavity, the

respective parts of each complementing each

other. The external margin ol pleomere 6 distally

beats a characteristit latéral expansion, which is

more or less pointed, and its edge immediately

underneath is distinctly hollowed. Both abdomi¬

nal parts (expansion t dépréssion) perfectly fit

into complementarv parts (depressed and salient)

at the margins of the sterno-abdominai cavity. A

transverse ridge in the middle of sternite 5 ends

in a thickened and sometimes granular promi¬

nence at the cavity s border. The latéral expansion

of pleomere 6 fus into the dépression in front of

the prominence, which blocks the abdomen. This

results in a strict adjustment and immobility of

the abdomen, posterior to the telson.

In Retropluma serenei de Saint Laurent, 1989

there is a high and granular prominence, which is

more developed in female.s than in males. In

Bathypluma spmijer de Saint Laurent, 1989, in

addition to the prominence, there is, ncar

suture 4/5, a rounded nodule which is specially

developed in females (Fig. 17C). The facr that

this “button” is less marked in males (Fig. I7A,

B) than in females, which are weakly dimorphic,

is a unique adaptation among retroplumids.

The spécial structures shown by retroplumids

deserve to bc extenstvely studied. The fossil repré¬

sentatives of the family are similar to extant retro¬

plumids. For cxample, the Upper Cretaccous

Costapluma cancana Collins et Morris (1975,

pl. 9 r , figs 4, 8), the late Cretaceous C. salaman-

ca Feldmann et al. (1997, fig. 3), and the

Paleocene C. nord es tin a Feldmann et Martius-

Neto (1995, fig. 3), which bave their ventral sur¬

face preserved, show an abdomen embedded in a

deep abdominal cavity which isexcavated on the

wide sternal plate, and a pleomere 6 with angles

laierallv expanded in a notch of sternite 4 near

suture 4/5. It is obvious that the rétention was

efficient indeed.

Family Pai.iuoal Rathbun. 1898

( : CYMorüUlDAF. Faxon, 1895)

l’hc peculiar characters of the family, summari-

zed by Guinot (1978a. 249; 1979a: 111,

figs 30G, 31. pl. 19, fig. 5, pl. 24, fig. 9), indica¬

ted that, despite a male génital orifice opening in

ZOOSYSTEMA • 1998 - 20 (4)

651

Guinot D. & Bouchard J.-M.

a sternal position, it is an advanced heterotreme, and Crossotonotinae Moosa & Serène, 1981, and

with a coxo-sternal organisation. When establi- ereating four new généra and six new species,

shing two subfamilies, Palicinae Rathbun, 1898 Moosa &C Serène (1981) did not elucidate their

Fig. 17. — Locking apparatus in Fletroplumidae, Palicidae and Hexapodidae' A-C„ Sathypluma spinilerde Saint Laurent, Philippines

(MNHN-B7017):A, B, 6 12.4x 15.6 mm: thoracic sternum with and witbout abdomen; C '7 119 • 14.9 mm: thoracicsternum wlthout

abdomen, D. Pallcus cnronii (P Roux), c 6 >. 7 mm, Atlantic, "Le Talisman' (MMHN-B 16335): ventral sudace ol abdomen; E.

Hexapus sexpes (Fabricius), 9 10.6 * 18 mm. Persian Gui! (MNHN-B 10209): Ihoracic sternum without abdomen: note the persis¬

tent buttons. Abbreviations: a, abdomen; a6. abdominal segment 6; b, button ol the press-button: so, socket; s.a.. sierno abdominal

cavity; t, telson; v, vulva; 4, 5. 6, sternites 4-6.

ZOOSYSTEMA • 1998

Abdominal holding Systems (Brachyura)

status. The phylogenetic relationships between

palicids and dorippids remains questionable.

Nine palîcine and crüssotonotinc species were

exainined. in chc type spedes Palicus caronii

(R Roux, 1830) the sternite 5, located very ante-

riorly, beats a strong proniinence jusr below the

level of suture 4/5. The correspondlng socket

(Fig. 17D) is deeply hollowed along tlie cxternal

margin of pleomere 6, but in the wholc anterior

part of the segment instead of the usual postéro¬

latéral angle. The proniinence remains as a non-

functional corneous scat in ovigerous females.

Family Llucusiioal Samouelle, 1819

Many leucosiids are rypical heterotremes, having

the male gonopore opening on the eondylus of

the P5 coxa, which is the plesiomorphic condi¬

tion. Some advanced représentatives, however,

show sternal male gonopores (Guinot 1979a: 15,

ftgs 45A-C, 55B-D; Guinot & Richer de Forges

1997: 478). (Sec Fig. I9C, maie gonopore in a

coxo-sterna! position).

The sternal plate of leucosiids is rather wide and

ail the sutures are disconrinuous. The well-

defined episrernites do nor cover the condyli of

P1-P4 in primitive généra such as Iphiculus

Alcock, 1896 and Pariphiculus Adams et White,

1848, but in advanced forms such as Leucosia

Weber, 1795 or Philym Lcach, 1817, the condyli

of ail pereiopods are covered. The sterno-abdo-

minal cavity is very deep and often elongate as far

as the bases of Mxp3, and the anterior segments

of the abdomen are gathered between the P5

coxae. The first abdominal segments, instead

remaining dorsal as in many eubrachyurans

crabs, are Completely tolded under the body, the

first pleomere being covered by chc posterior

margin of carapace.

There arc several ways to allovv die plcon’s base to

bc firmly grasped. rhtse involve a connection

between the sternal plate, pleomere l and the

carapace; the transverse élongation of pleo-

mere(s) 1 and/or 2; the development of a dorsal-

ly visible expansion of sternite 8, which sur¬

rounds the eondylus of P5, and, additionally (as

in Philyra ), a dépréssion inside the sterno-abdo-

minal cavity at the surface of sternite 8, which

receives the anterior part of pleomere 3. The

pleopods themselves are coaptated with the bot-

tom and margins of the sterno-abdominal cavity

and with the abdomen (Fig. 19D). A perfect

coaptation exists in Leucosia where, in addition,

the Gl is situaied in such a mantier chat it pro¬

vides a place for the sternal locking structure.

Both male and female leucosiids are often found

with the abdominal margins so closely ftxed to

chc sternal plate that the abdomen is broken

when lilred. Ovigerous females are known to

hâve a brood chamber, somerimes dosed like a

box, because its margins are completely knitted

to the edge of the sterno-abdominal cavity.

Guinot (1979a: 145-148) rccognized two main

types of abdominal retaining among leucosiids. In

the two primitive généra Iphiculus and Pdriphi-

culus, which hâve been recentlv re-examined, nei-

ther prominences nor sockcts are présent, and the

highly movable telson sinks into chc bottom of the

cavity. In Randallia Stimpson, 1857 (Fig. 18) and

Ilia Leach, 1817 (Fig. 19A) chc sterno-abdominal

cavity is ver)' deep and markcdly tapering, with its

sides fttting the thick and sinuous edges of the

abdomen. There is a mixture of coaptations by jux-

Fig. 18. — Abdomen in Leucosiidae (Phylirinae). Randallia villo-

sa Chen, holotype, 8: segments 3 through 6 plus telson. (After

Chen 1989, fig. 11 d).

ZOOSYSTEMA • 1998 -20(4)

653

Guinot D. & Bouchard J.-M.

taposition and by engagement, which characrerizes

the subhimilies Fbaliinae Smnpson, 1871. Crypto-

cneminae Srimpson, 1858, and part of the

Philyrinae Rarhbun, 1937-

In the subfàmily Leucosiinae, Leucosia (the type

species L. cmniolaris (Linné, 1758) was not avai-

lable] shows a particular configuration, vvith a

sort of spur thaï projects inside the sterno-abdo-

minal cavity, at the level of sternal suture 4/5. In

/.. nnatum (Herbsr, 1783) and L. unidentata de

Haan, 1841 for example, the spur enters in a

large notch excavated in rhe thick edge of the

abdominal margin, and rhe abdomen becomes

completely included in the snrooth sternal sur¬

face (Fig. 19B-D). The endophragmal skeleton of

Leucosia shows, ar the level of endosternite 4/5, a

pair of apophyses, thaï exactly corresponds to the

pair ofspurs (Fig. I9E). In addition to the spur

of sternité 5, species of Leucosia show different

developnlents of sternites 6 and 7 along the sterno-

abdominal caviry, to which correspond comple-

mentarv curves on the abdomen mai'gins

(Fig. 19B). The Leucosia species display a more or

less acure spur on sternite 5.

A similar configuration is seen in the species exa¬

minée! of Philyra Leach, 1817, with various

expansions of sternites 6 and 7. In P laevidorsalis

Miers, I 881, in which the spur of sternite 5 is not

well developed, the sternite 6 prorudes into a cor-

responding notch on the abdominal margin. In

P. pismn de Haan. 1841 there arc- rwo granular

profusions oti sternites 5 and 6. In RaiidcilHa vil-

losa Chen, 1989 the border ot the sterno-abdo-

minal cavity bears scvetal protrusions with diffe¬

rent notches in the edge of the abdomen, parti-

cularly a circulât one at the junction of pleo-

meres 5 and 6 (Fig. 18). In ail of its characters

(carapace, plcopods) R. uilloia obviously docs not

belong to the genus Randallia , a fact confîrmcd

by the spécial configuration of the stemo-abdo-

minal margin. A re-evaluation of ail leucosiid

members is nceded.

Discussion

On the basis ol Leucosia ocellata Bell, 1855,

Bellwood (1996: 176, 191) indicated the absence

of “cohésion nodules” in leucosiids and recorded

these as absent in her matrix. Ir is clear, on the

one hand, that in Leucosia the abdomen is not

locked by a typicaJ press-button, which supports

rhe spécial category established by Guinot

(1979a: l46) for the leucosiine spur. On rhe

other hand, despite the presence of a hollow that,

like rhe usual sockct, (partly) bclongs to pleome-

re 6, there is no true socket but only a notch at

the junction of pleomeres 5 and 6. Endo¬

phragmal invaginations similar to the apophyses

of the endosternite 4/5 observed in Leucosia vvere

never fourni in relation to rhe buttons of the typi-

cal press-button.

Few brachyuran crabs exhibit, as in leucosiids,

such a üniquely attachée! abdomen, joined with

the resr of rhe sternal plate in such a way that the

abdomen is broken when one attempts ro lift it

(Fig. 19B). Pinnotherids also show stages in

which the abdomen is fiised with adjacent zones

of rhoracic sternites (sec below).

Both the sternal spur and abdominal notch are

visible externâlly at the junction berween the long

pleomcrc- 6 and the preceding segment. A charac-

teristic of the locking mechanism in Leucosia is

thar the sternal part involved ean be visible exter-

nally, in contrast to the typlcai press-button

where the sternal part is covercd by the folded

abdomen and normally not seen. Such a structu¬

re may be easily recognized in fossils for the iden¬

tification of Leucosia and allied généra. For

example, the well preserved Neogene leucosiids

srudjed by Morris 6i Collins (1991, figs 17b, 18,

24b) show the presence of a spur in a Leucosia

species, and its absence in two Ipbiçulus species.

Family Pinnotheridae de Haan, 1833

The sternal plate is more or less wide, with ail

sutures discontinuons, and the sterno-ahdominal

cavity is deeply hollowed. The male sexual orifice

of pea crabs, which opens on the sternum, has

gcnerally led to their assignment to the çatome-

topous crabs. Rice (1980: 315) regarded the

Pinnotheridae (with rhe Leucosiidae) as “the

most advanced catomevopous familles". But this

male configuration was reconsidered as coxo-ster-

nal and, consequently, the familv Pinnotheridae

Was transfered ro the Heterotremata, instead of

Thoracottemata (Guinot & Richer de Forges

1997).

It is difficult to derermine the morphological cha-

racteristics of the apparatus in pinnotherids

654

ZOOSYSTEMA - 1998 - 20(4)

Abdominal holding Systems (Brachyura)

because of the rareness of males and the weak- belonging to Ostracotheres tridacnae (Rüppell,

ness, loss or absence of specialized structures in 1830) and Pinnotheres pisurn (Linné, 1758), a

soft individuals. In hard-shelled individuals double locking mechanism has been observed: a

Fig. 19. — Leucosiidae (A. Ebalunae; B E, Leucosiinae), withour a typical aboominal locking apparatus; A. Ilia nucléus (Linné), S

27 x 26 mm, Mytllene, Pon Vera (MNHN-8 19975); B, C, Leucosia sp. (identified as L uniclentala de Haan), £ 29 27 mm, in the

vicinity of Tokyo (MNHN-B 16965) thoracic sternum with and witnout abdomen; D: ventral surface of abdomen ; E, Leucosia sp. (iden-

tified L. longilrons de Haan), £ 21 - 20 mm, New Caledonia (MNHN B 17525). endophragmal skeleton, with ttie pair ol endosterna!

apophyses. Abbrevialions: a5, a6, abdominal segments 5, 6; ap, endosternal apophysis, g Gl n notch; p, pénis (male gonopore in

a coxo-sternal position); s.a., sterno-abdominal cavity; sp, spur; t, telson; 4. 5, 6, 7, sternites 4-7.

ZOOSYSTEMA • 1998 - 20(4)

655

Guinot D. & Bouchard J.-M.

large knob on sternite 5 and a smaller one on

stemite 6 (Guinot 1979a: 148, fig. 24D, pl. 24,

figs 7, 8), Only one prominence on che lone ster¬

nite 5 was seen in Pinnixn tmnsversalis (H. Milne

Edwards et Lucas, 1843). In Pinnotheres pisum,

Atkins (1926: 476, figs 1, 2) indicated one pair of

prominences and sockets in a hard male stage,

and also in youngfeinales whidi are almosr indis-

tinguishable from males.

Stauber (1945- 274, 277, 278, 280, figs 15, 16)

studied living individuals of Znops ostreurn (Sav,

1817) (as Pinnotheres ostreurn, cf. Manning

1993a: 528; 1993b: 127), which is symbiotic

with tbe oyster Cmsostrea virginioi (Gmelin),

and recognized severaJ distinct stages berween die

invasive crab stage and the adult. In a hard male

stage (mean cw 2.13 mm) rwo pairs of knobs

were présent on thoracomeres 5 and 6 (shallower

and smaller on the sixth) to which eorresponded

rwo pairs of shelf-like projections on pleomeres 6

and 5, “making a close-fitting efficient locking

mechanism'’. In the first fcmalc stage (mean cw

1.83 mm), which invades the oyster, only the sets

on thoracomere 5 and on pleomere 6 were pré¬

sent. During these masculine and féminine

stages, it was impossible to lift the abdomen

without injuring it. Ehe sanie sec occurred in the

second fenialc stage, but the mechanism was wea-

kened by the thinner exoskeleton. For that stage

and for the subséquent féminine stages (the last

being the mature fcniale), which luid a very wide

abdomen and no knobs, Stauber concluded chat

it was an adaptation to living with in the mamie

cavity of the host. In a masculine second stage

(cw 2-4.8 mm), with a soft body, two pairs of

sternal knobs and only one pair of abdominal

structures on pleomere 6 were présent, so that the

abdomen was easily lilted (rom the sternum even

in living crabs [ibide. 280). Christensen &

McDermott (1958), who re-investigated the bict-

logy of Zcwps ostreurn with new information and

new terminology, and Christensen (1959) for

Pinnotheres pisum, did not mention the presence

or rôle of locking structures.

It is obvions that the locking System of pinnothe-

rids varies during their life historv. There is no

information available to détermine whecher the

remaining pinnorherid généra follow the same

pattern as Zaops ostreurn.

The prominence and Socket (Fig. 26A, B) of a

hard male of P. pisum were observed with the

scanning électron microscope.

Family Calappidae de Haan, 1833

The sternal plate is relatively narrow but

sutures 4/5 and 5/6 are discontinuons, being

separated by a very small gnp; the well-defined

epsiternites cover condyli of P2-P4. The abdo¬

men is rather long and lodged in a well-excavated

cavity. There is a typicai press-button, with pro-

minences (appearing granulous under the bino-

cular) close to sternal suture 4/5 (Guinot 1979a,

pl. 14, figs 1, 2). Bellwood (1996: 176, fig. 2A)

indicated sonie nodules located itnmediately

behind che sternal suture 4/5 in the calappids

that were examined.

Family MauIHDAE Macleay, 1838

The thoracie sternum is slightly enlarged, only the

suture 4/5 is discontinuons and che sterno-abdo-

niinal cavity occupies a very small part of

sternite 4. There is a typicai press-button, with

prominences not very far from the sternal sutu¬

re 5/6 (Guinot 1979a, pl. 14, figs 4, 5). Bellwood

(1996: 176, fig. 2C) indicated that the nodules

were immediately located above the sternal

suture 5/6 in the Matutidae that were examined.

FAMILIF.S Ap.THIUDAF. Dana, 1852 entend. AND

HF.PATIDAK Stimpson, 1871

Guinot (1966-1967) réhabilitated the ancient

Hepattnae Stimpson, 1871 (sec also Ortmann

1892: 556, 5(s9). Fhe genus Hep,uns Latreille,

1802, generally placed ncar Qdetppn Weber, 1795

and Matuta Weber, 1795 (Rathbun 193“: 234;

Balss 195“: 1611-1612), was separated. The pro-

visional désignation Parthenoxystomata was used

by Guinot to include, in addition to Heptttus,

some allied généra that were prcviuusly classified

in various high level taxa: Aothni l.eacb, 1816,

attributccl to the Parthenopidae (Rathbun 1925:

550; Balss 1957: 1629-1630; Gatth 1958; 434,

467); Osdchiln Stimpson, 1871. included in

Calappinae (T. Sakai 1965: 57) or in M.mitinae

(Rathbun 1937: 234; Balss 1957: 1612) or in

Aethrinae entend. (T. Sakai 1976: 288; Huang

1989: 347); lieputellu Smith, 1869, and Artaeo-

morpha Miers, 1878 (see Guinot 1978b), tradi-

656

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

tionaliy attributcd to chc Leucosiidae Ebaliinae.

The phylogenetic relationship of ail chese généra

was supported by a morphocline. The genus

Sakaila Manning et Holrhuis, 1981 was subse-

quently included in the parthenoxystomatous

group.

The affinities of these généra wirli the other

calappids, represented by CaLippu, and the par-

thenopids semu lata remained questionable (rhar

was the reason of rhe informai désignation

Parthenoxystomata), but rhe problem vvas ofren

ignored and never solved. The familial désigna¬

tion Aethridae, generally not accepted, was used

by Ng & Rodriguez (1986) \vho recognized four

f'amilies in the large parthenopid group semu lata:

Daldorfiidac, Dairidâe (see below), Partheno-

pidae s.s., and Aethridae.

Bellwood (1996: 185, 186) conlirmed the sepa-

rate status of Hepatus with a familial rank,

Hepatidae pro parte sensu Guinot, close to the

Calappidae, and considered the two fantilies

Hepatidae and Calappidae as sister group.s in the

supertamily Calappoidea. The crucial question of

the status of the parthenoxystomatous genus

Aetbra was ornitted fmm the cladistic analysis of

Bellwood. If further researchcs demonstratc that

Hepatus and Activa belong to the sa me elade, the

farnily name Aethridae Dana, 1852 has prior-

ity over Hepatidae Stimpson 1871. Osdchild

contains several spccies, somc of which hâve a

more aerhrid lacies, while others look more like

hepatids (Guinot 1966-1907: 836, 838).

In aethrids/hepatids the sternal plate is wider

than in calappids, and, in conirast to calappids,

the well-defined epsiternites do not cover the

condyli ol P2-P4. Plie sutures 4/5 and 5/6 are

medially separated,

In males of Hepatus the external margin of every

abdominal segment is sinuous and fits into a cor-

responding surface at the border of the very deep

sterno-abdominal caviry. Sterniie 5 afso beats,

posterior to a dépréssion, a prominence which

hangs on a well-delimited socket on the internai

surface of pleomere 6, I bis is a case where the

combination of a press-butron apparatus and of

multiple sternum/abdomen attachments allows a

powerful holding of the pleon. Bellwood (1996:

176, fig. 2B) defined ihe organization in hepa¬

tids, attribudng to sternal structures a location

half-way between the two sutures 4/5 and 5/6 in

hepatids, at leasr in the material that was exami-

ned. We hâve fourni that in Aetbra uruposa

(I inné, 1764), the type species of the genus, rhe

nodule is more posterior and that, in sonie

Osaebda species, the wide prominence can lie

located dose to suture 5/6,

In aethrid/hepatid females each sternite 6 and 7

beats a strong protubérance rhat fits into a notch

on abdominal margin. After the moult of puber-

ty a vestige of the socket can remain on rhe inter¬

nai surface of pleomere 6,

The males of Aetbra Leach, Osachila Stimpson

and Sakaila Manning et Holthuis présent a

remarkable indentation on the particularly thick

edge of the abdomen, to which complementary

sinuous surfaces of the sterno-abdominal cavity

correspond. I he sculptured and eroded orna¬

mentation ol diese régions provides a strong

attachment. A sternal press-button is présent. A

Socket is deeply excavated and defined by a strong

superior margin. In females, protubérances on

sternites (particularly at the base of the cavity) fit

into the corresponding edge ol the abdomen,

which allows a perfect closing.

Famiues PARTHENOPlDAt Macleay, 1838 AND

Daldori iidaf. Ng et Rodriguez. 1986

The fantilies Parthenopidae and Daldorfiidae are

dealt with separately, awaiiing a review ol Lheir

phylogenetic relationship.s and précisé status.

The margins of hoth sterno-abdominal cavity

and abdomen display remarkable corresponding

surfaces, with a deeply eroded complementary

ornamentation, in contrast to the smooth cavity.

The press-button is typical (Guinot 1979a: 138,

pl. 17, figs 1-4).

Family Dm ridai- Ng et Rodriguez, 1986

Guinot (1967) showed the close relationship bet¬

ween Daim de Haan, 1833 (previously regarded

as a xanthid) and Dainndes Stebbing, 1920. A

family was esrablished by Ng &c Rodriguez

( 1986: 90, 97) for these généra, specialized but

with plesiomorphic structures (notably G2 with a

long and wliiplike flagellum). The exact relation¬

ship with other parthenopids sensu lata needs to

be established.

ZOOSYSTEMA ■ 1998 • 20 (4)

657

Guinot D. & Bouchard J.-M.

Daira and Dnirodes hâve a similar locking appa-

ratus. In Dairoides knsei (T. Sakai, 1938) a perleci

coaptation was (ound in a male (cw 59 mm) but,

in a larger male (55 X 71 mm), the two promi-

nences did not fit into sockets since îhey were

external to thc abdomen (Guinot 19~9a: 138,

fig. 27A, pi. 17, fig. 9).

SUPERFAMILY Ma|OIDEA Samouelle, 1819 (seve-

ral families)

In the Majoidea, vvhich contains about 800 spe-

cies assigned to more titan 150 généra and in

which 50 suprageneric categories were listed

(Manning & I folthui.s 1981: 253). Garth (1958)

and Griffin 8e Thtnrer (1986) recognized seven

subfamilial taxa. In these two publications, fîve

subfantilics rcceived the sanie name: Majinae

Samouelle, 1819, Inachinae Macleay. 1838,

Mithracitiae Macleay, 1838, Pisinae Dana, 1851,

Oregoniinae Garth, 1958, whereas two had difle-

rent names: Epialrinae Macleay, 1838 (in

Griffin & Tranter = Acanthonychinae Stimpson,

1871, in Garth) and Tychinae Dana, 1851 (in

Griffin & Tranter = Ophthalmiinae Balss, 1929,

in Garth). On the basis of larvul characters six

majid subfamilies were considered by Ingle

(1976): Oregoniinae, Acanthonychinae, Inachi¬

nae, Pisinae, Majinae, Ophthalmiinae, Guinot

(1978) suggested that the majids may be raised to

superfaniily status, and Clark & Webber (1991)

proposed a new classification of the Majoidea,

distingnishing four major families: Majidac,

Inachidae, Oregonîidae, and Macrocheiridae

Dana, 1851. We proposed to consider a supple-

mentary category, the lnachoididae Dana, 1851

(Drach & Guinot 1982, 1983; Guinot 1984;

Guinot & Richer de horges 1997).

It is beyond thc sçope of this paper to investigate

the locking Systems in such a large group. It

appears that the majoid mechanism is a tvpical

press-button. In certain majoid représentatives,

for example in mosc Inachinae [Fig. 2üA, B:

Inachus dorsettensis (Tonnant, 1777)] and ail

Inachoidinae [Fig. 20C, D: Pyromain tuberculata

(Lockingtou, 1877); Fig. 20F,, F: Collodes lepto-

cheles Rathbun, 1894] the abdominal segment 6

is fused to telson in a pleotelson so that the soc-

kets are unusually located on die last element of

the abdomen. The button may be very acute,

often positioncd on oblique flanks of the sterno-

ahdominal cavity, and the socket exrremely deep

(see Figs 20B-D, F, 21A-F). In the majid genus

Thacdnùphrys Griffin et Tramer, 1986 botli but-

ton and socket may be persistent in adult lemales

(Fig. 21E, F).

The family Hymcnosomatidae Macleay, 1838

was recently affiliated with Majoidea by

Guinot & Richer de Forges (1997), who provi-

ded information on the locking Systems.

Srernites 1-3 form a sliield, more or less narrow

and prolonged between the Mxp3; thc plate

constitnted by xternites 4-7 is very widc, with ail

the sutures short, interrupted, and placed latcral-

ly. The undivided broad médial part of the plate

corresponds to thc absence of médial skeletal

phragma. The sterno-abduminal cavity of males

is generally very reduced in length, sometimes

shallow, othenvise deep, in contràst to the large

and often verv specialized hrood ohambers found

in lemales. The Hymenosomatidae difler from

the Lisual brachyuran pattern ol six abdominal

segments plus telson in never having more rhan

five pleomeres plus telson in both sexes. We

regard the fusion of abdominal segment 6 with

telson, that is the formation of a pleotelson, as an

hymenosomatid synapomorphy, In the advanced

species of the family. additional pleomeres may

be fused to that terminal pleotelson, which leads

to a weakly segmented abdomen, tor cxample ol

only three éléments (segments 3-5 lused to pleo¬

telson) in the case of male and fcmale Criistaenia

/udawanensis (Serène, 1971).

The configuration is of the press-button type, but

with sottie specialized featlires. As for vulvae, the

locking prominences are situafed on the undivi¬

ded part ol sternal plate, olten on the oblique

flanlcs ol the sterno-abdominal cavity; at first

sight rheir lielonging to stemlte 5 is not obvious

(Guinot 1979a: 1 H), 149, lig. 30A. pl. 20, fig. 8,

pl. 23, fig- 6). Dissection of the skeleton permit-

red us to deline the internai configuration of

phragma and the exact place of the sternal pro-

minences on somite 5 (Guinot & Richer de

Forges 1997, figs 2A, C, F, 6F). In hymenoso-

matids the sockets appear laterally located at the

pleotelson hase but belong to pleontere 6 as in ail

Brachyura.

658

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Fig. 20. — Locking apparatus in Majidae (A, B, Inachinae; C-F, Inachoidinae); A. B, Inacdus dorseltensis (Pennant),. d 20 :< 18 mm,

Saint-Martin-de-Ré (MNHN-B 17337)' A, Ihoracic sternum with abdomen; B, central Surface of abdomen, C, D, Pyromaia lubercula-

ta (Lockington). : 17* 12.2 mm. Lower California (MNHN-B 9290): C, thoracic sternum withoul abdomen; D, ventral surface of abdo¬

men; E, F, Collodes leplocheles Rathbun, i 17.5 * 13.2 mm, Gult of Mexico (MNHN-B 14055): E, thoracic sternum with abdomen:

F, ventral surface of abdomen. Abbreviations: a6, abdominal segment 6; b, button; pt, pleotelson; so, Socket; t, telson; 5, sternite 5.

ZOOSYSTEMA • 1998 - 20(4)

Guinot D. & Bouchard J.-M.

Anothcr characteristic fèature of hymenosoma-

tids is rhe présence of defined imercalated plate-

lets, eiiher complctely arciculated, movable as in

Odiormms pilosus (A. Mil ne Edwards, 1873)

(Fig. 27) and Amarinus angelicus (Holthuis,

1968), or fused, the suture rhat delimits rhem

being more or less discernible (Lucas 1980, fig. 7;

Guinot & Richer de Forges 1997, Figs 4A-E,

6B-E). We consider that the transformation sériés

of the movable imercalated plalelets into fused

plates, then rheir complété intégration to pleotel-

son, illustratc the stages towards the apomorphic

condition. The imercalated plarclets in the

Hymenosomaridae correspond to the sockets in

other brachyuran crabs and, as such, constitute

the complememary abdominal parts for locking

and are used for this purpose. When the delimi-

ted platelets hâve disappeared, only sockets

remain excavated on the ventral surface of pleo-

telson’s base (see Hypothesis about die homology

uropod-socket).

FAMILIES Portunidae Rafincsquc, 1815, PLATY-

XANTHIDAK Guinot, 1977, Eripiiiidae Ortmann,

1893 (= Menippidae Ortmann, 1893), Carpi-

i.iidae Ortmann, 1893, Xanthidae Macleay,

1838, Panopeidae Ortmann, 1893, Bytho-

GRAEIDAE Williams, 1980, TrapéKIIDAE Miers,

1886, PlLlJMNIDAE Samouelle, 1819, Humi.DO-

N1DAE Miers, 1879, GeRVONIDAL Colosi, 1923,

Goneplacidaf Macleay, 1838 sensu lato (tiot an

exhaustive list)

There is a typical press-button. The button and

the socket vary only according to their size,

shape, microstructures (Bouchard, in prépara¬

tion), and to their relative location on the surface

of the corresponding somite, always on the surface

of sternite 5 for the button and on the internai

surface of pleomere 6 for the socket. See text and

figures in Guinot 1979a.

The button and the socket (Fig. 26C, D) of the

hydrothermal crab Austinograea alayseae Guinot,

1990 were observed with the scanmng électron

microscope.

Families of freshwater crabs (DeCKENHDAE

Ortmann, 1897, Gecarcinucidae Rathbun,

1904, Parathelphusidae Colosi, 1920, Potami-

DAE Ortmann, 1896, POTAMONAUTJ DA F. Bott,

1970, PSE U DOT HELPHU Si DAE Ortmann, 1893,

SUNDATHEl PHUS1DAE Bott, 1969, TRICHO-

tlACm.IDAE H. Milite Edwards, 1853)

Ail the représentatives of tbese diverse brachyuran

families are mie freshwater crabs, without marine

larval stages: they are rescricted to freshwater

habitats and, apparently, no members Dor close

relative are marine. They ail hâve a male opening

that is located on the P5 coxa: they belong in the

Heterotremaia. The existence of diverse forms of

a penial groove, which is rudimentary in

Pseudothelphusidae and partially covered by ster¬

nal parts in Trichodactylidae, could be considered

as first stages in the process leading to coxo-ster-

na! orifices (Rodriguez 1992; Ng & Rodriguez

1995).

It is usual that the abdomen rétention is ensured

by a typical press-button, but the configuration

needs to be verified in ail taxa.

Family HkXAI'OOIIME Miers, 1886

A doubtful and ccinporary assignment of the

Family Hexapodidae to the i'horacotremata was

suggested by Guinot (1978a; 1979a: 114. 145,

fig. 32). Hexapodids, that are highly modified

(some species aie known to live in tubes of anne-

lids and on hydroids), appear to not belong ro the

catometopous group {ibid:, 215).Theanribution

of the Family to the Heierottemara (Guinor &

Richer de Forges 1997: 496, table 1) is in accor¬

dance with the ontogenetic data suggesting a

close relationship to Goneplacidae (Pereyra Lago

1988). The apparatus belongs to the press-button

type (Fig. 17F).

Family Crypiih hIRIDAE Panlson, 1875

The afFmities ot gall crabs with other brachyuran

groups rcmain unclcar, despitc the essemial Fin-

dings of Kropp (1988, 1989, 1990) and Kropp &

Manning ( 1985, 1987, 1996). The male orifice is

indieated as opening in a sternal position, but

possible States of this character need to be analy-

sed. A coxo-sternal organization is admitted, lea¬

ding to the inclusion of the family among the

advanced Heterotremata (Guinot &c Richer de

Forges 1997: 496, table 1).

660

ZOOSYSTEMA • 1998 - 20 (4)

Abdominal holding Systems (Brachyura)

Fig. 21. — Locking apparatus in three majid species of Thacanophrys Griffin efTranter: A, B, Thacanophrys albanyensis Ward (as

Chloriroides barunai Serdne), S 26 ; 14 mm holotype. Indoriesia (MNHN-B 17817): A. thoraciG sternum wjthout abdomen: B, ven¬

tral surface of abdomen; C. D. Thacanophrys harmandi (Bouvier) (as Acabthophrys harmandi), ,î 46 ;< 24.5 mm, holoiype in lhe vici-

nity of Tokyo (MNHN-B 22330): C, thotacic sternum witlioul abdomen D. ventral surface of abdomen: E- F, Thacanophrys hngispi-

na (de Haan) (as Chorinus longlspinus). Y .23 • 27.4 mm, Japan (MNHN-0 17853): E, thoracic sternum without abdomen: note tbe

persistent buttons; F. ventral surface ot abdomen, wrth persistent sockets, Abbrevia|ions. a6, abdominal segment 6; b, button; s.a.,

sterno-abdominal cavlty; so, Socket; t, telson; v, vulva; 5, sternite 5.

ZOOSYSTEMA • 1998 - 20(4)

661

Guinot D. & Bouchard J.-M.

The extensive studies of Kropp reveal a wide

range of configurations exhibited by the family,

in particular with regard to the thoracic sternum,

often illustrated in the females. The sternal pljte

varies from relativefy narrow ( Çryptocbinn Heller,

1861) to wide or very wide (Hapalocarciuus

Stimpsün, 1859, Pseudohapalocartinus Fize et

Serène, 1956). d ht sternal sutures 4/5 and 5/6

are always interrupfed, but in the posterior part

of the plastron the sutures are diversely arranged.

The sutures 6/7 and 7/8 remain continuous in:

Cryptvchirus, Litboscaptus A. Mil ne Edwards,

1862 pro parte, Zibrovia Kropp et Manning,

1996, Opecarcinus Kropp et Manning, 1987 pru

parte, Fiztuereneut Takeda ef l amura, 1980. Only

suture 7/8 may remain continuous, the 6/7 sutu¬

re being more or less largely interrupted, in:

Litboscaptus pro parte, Opecarcinus pro parte,

Hiroia Takeda et Tamura, 1980. The posterior

sutures, 6/7 and 7/8, are interrupted, so thaï ail

the sutures are interrupted on rhe largely undivi-

ded, broad sternal plate, in: Llapalocarcimis,

Pseudohapalocarcinus, Fungicola Serène, 1966,

Pseudocryptocbirus Hiro, 1938, Xynomaia Kropp,

1990 (not exhaustive lists).

The shape ol the sterno-abdominal cavity and the

précisé location of the abdomen on rhe ventral

plate are not well-kown in cryptochirids. The

existence of a locking appararus was never repor-

ted in males, always smaller than the females and

relatively rare. The abdomen seems to be ventral-

ly folded, withour spécial morphological devices

to hold it (R. K. Kropp pers. coin.). It is note-

worthy that absence of any apparent structures

occurs in forms linding protection within galls or

in open pits formed in corals.

THORAC.OTREMATA Guinot, 1977

The Thoracotremara, sterni trente for the fentale

orifice (vulvae on stérilité 6), show a constant

sternal location of the male pores, which is on

sternite 8 but in various places in relation to the

sternal suture 7/8, so that the sternal male ope-

ning constitutes a multistatc character (Guinot

1979a: 218, figs 53, 54, 55F-H, 56). The section

Thoracotremata includes only littoral, amphi-

bious and terrestrial brachyuran crabs (the genus

Planes Bowdich, 1825 excepted) in marine or

braclcish habitats, with diverse représentatives

adapted to freshwarer existence (Guinot &

Richer de Forges 1997: 496, table 1).

In ail représentatives of the Thoracotremata the

thoracic sternum is wide or very wide, the

sutures 4/5 through 7/8 are interrupted, the

general ly deep sterno-abdominal cavity is often

anteriorly delimited by a cristifornt mangin, the

abdomen forms a thin plate, and the male orifice

always opens on the sternum. A non-origination

ol the male gonopore and pénis from P5 coxa, i.e.

a direct sternal emergence, was observed in

Ocypode cursor (Linné 1758) in which the pénis

emerges near the end of thoracic suture 7/8

(Guinot 1979a, fig. 56). Regarding the gonopo-

re/pems organizadon. the ihoracotrematous cha¬

racter state appears eiiher with a pro.ximiry ro P5

coxa (Grapsinae, Plagusiinae, a part of Gecarci-

nidae and Sesarminae) or a location that is com¬

pletel)' sternal, for exemple in the Ocypodidae. It

is now obvious that che l horacoirernata sensu

Guinot (1977, I97S, 1979) display various

conformations of male ducts and openings: this

entphasizes the necessity of additional information

and further investigations (R. von Sternberg pers.

corn.).

Family Grapsidal Macleay, 1838

lu both subfamilies Grapsinae and Plagusiinae

there is a complété typical press-button appara-

tus, persistent and apparently functional throu-

ghout life. In the Plagusiinae the socket is located

in the postero-laieral and external angles ol abdo¬

minal .segment 6, while in the Grapsinae the Soc¬

ket is similarlv positioned in rhe postero-lareral

part but generally lar from the external ntargin of

the abdomen. In both subfamilies Varuninae and

Sesarminae, the locking mechanism is also of the

press-button type: eirher it is présent throughout

lite or tends to disappear in large crabs (even in

males) or it is contpletely lost.

SUBl’AMILV GRai'M\ak Macleay, 1838

Within the genus Grapsus Lamarck, 1801, as in

G. grapsus (Linné, 1758) and G. tenuicrustatus

(Herbsr, 1 T 83) (Fig- 22A), there is an acutc pro-

minence. persistent in males throughout lile

(Hartnoll 1965: 128), and a socket far Iront the

662

ZOOSYSTEMA ■ 1998 -20(4)

Abdominal holding Systems (Brachyura)

external margin of the sterno-abdominal cavity Cornuberr ( 10 S S ) In male /.' traits vers its (Gibbes,

(Guinot 1979a: 142, pl. 20, figs 1, 2). In Gonio- 1850) and P. gracüis (de Saussure, 1858)

psis cruentatn (Latreille, 1803) the locking appa- the mechanism is functional in ail sizes, then

ratus is functional only in immature females and throughout life, appearing in the first stages of

in ail males. post-larval existence and only regressing in matu-

In Pachygrapsus Randall, 1839 the press-button re females, which hâve a widened abdomen

was studied by Pérez (1933a, 1933b) and Vernet- (Hartnoll 1965: 135, 136).

Fig. 22. — Locking apparatus in Grapsidae, subfamily Grapsinae; A, Grapsus tenuicruslatus (Herbst). <5 65 ■ 72 mm. Sainte-Luce

(MNHN-B 11541); B-D, Metopograpsus latifrons (White), i 28.8 x 32.8 mm, type sériés of M.pictusk. Milne Edwards, probably New

Caledonia (MNHN-B 12882): B and C, thoracic sternum with and without G1 ; D, ventral surface of abdomen. Abbreviations: a6, abdo¬

minal segment 8; b, bütton; d, dépréssion to recelve the apex of G1 ; g, G1 ; pr, projectiorron sternite 6; so, SGcket; s.a., sterno-abdo¬

minal cavity; 5, 6, sternites 5 and 6.

ZOOSYSTEMA • 1998 -20(4)

663

Guinot D. & Bouchard J.-M.

In the marertal of the genus Geograpstts Stimpson,

1858 that was examinée!, for example G. lividus

(H. Milne Edwards, 1837), the press-button is

typical, with the Socket far front the external mar-

gin of the abdomen and sometimes horizontal.

In Metopogntpsus H. Milne Edwards, 1853, as in

M. frontalis Miers, 1 880 and M. océaniens

(Jacquinot, 1853). the button is very prominent

and the Socket wide, without defined borders. The

most striking case ts shown by M. latifrons

(White, 1847) (Fig. 22B-D: male of the type

sériés of M. pictusA. Milne Edwards, 1867). The

button on somite 5 is vert' elevated, transvcrscly

elongated, and offers an uneven surface. There is

a close coaptation between the button and the

male first gonopod: the T-shaped apex' ol G1

cornes in contact with the posterior portion of the

prominenee and is situated in a corresponding

dépréssion hollow'ed on the surface of die sterno-

abdominal cavity, just behind the button and at

the level ol suture 4/5. In addition, a rougit pro-

minencc on somite 6 appears to limit sideways

movement ol the adjacent pleopod. In

Metopograpsus messor (borskal, 1775) and M. tbu-

kuhar (Ow'en, 1839) the button is smaller, roun-

ded, not transvcrscly elongated, and the apex ol

G1 is not T-shaped. Ir is doubtful, however, that

ail these species actually belong ro the sanie genus.

In Planes Bowdich, 1825 the button is ver}' close

to the sternal suture 5/6 and the Socket is near the

external rnargin of the abdomen.

SUBFAMIIY PlaGUMINAE Dana, 1851

In ail the examined species of Phgusia Latreillc,

1804, P. depcessa (Fabricius, 1775), P immacula-

ta Lamarck, 1818 and P. glabra Dana, 1851

(Fig. 23C-D), the sternal prominenee is strong,

looking more or less corneous. Tliis corneous

aspect is duc to a scalv paving, observed with the

scanning électron microscope (Fig. 26K P. imma-

culatd) (Bouchard in préparation). The deep Soc¬

ket persists and fonctions throughout life, except

for post-puberty females wliere only a sternal scar

remains visible (Guinot 1979a: 144, pl. 18,

figs 9-11).

In Percnon Gistel, 1848, P. planissimum (Fferbst,

1804), P. ajfine (H. Milne Edwards, 1853),

P abbreviettum (Dana, 1851), and P. guinotae

Crosnier, 1965 were examined: the button has a

wide base, forming a large prominenee, and its

surface looks corneous; the Socket is deep and

outlined with a strong thickening (Guinot

1979a, pl. 23, figs 2, 3).

SUBFAMII Y VARUNINAI H. Milne Edwards, 1853

I he subfamily Varuninae, which includes more

than flfteen généra living in estuafme and bra-

ckisli or freshwater habitats, forms an artificial

grouping that needs re-evaluation. h similarly

appears heterogeneous in respect of the locking

System.

In Varuna littentta (Fabricius, 1798) (ej. Guinot

1979a: 142, pl. 20, fig. 5) (Fig. 23A) and in the

species of Gaeticc Gistel, 1835, Hemigrapsus

Dana, 1851, Bracbynatus de Haan, 1833,

Pyxidognathus A. Milne Edwards, 1878, and

Ptychogttatbus Stimpson, 1858 that were exami¬

ned, a tvpical press-button was présent and per¬

sistent throughouL the life in males, with a Socket

a little remote boni the abdominal rnargin. In

Euchirograpsus H. Milne Edw'.irds, 1853, P. ame-

ricanus A. Milne Edwards, 1853, E. liguriens

H. Milne Edwards, 1853 and E. timorensis

Tiirkay, 1975 that were examined h ad a specially

acute button and a Socket posteriori)' defined by

a sttongly calcified border, which suggests that

the genus is misplaccd in the Varuninae.

A sniail button and a Socket arc présent in

Eriocheir sinensis 11. Milne Edwards, 1854

(Fig. 23B), but the locking mechanism is no lon¬

ger efficient in males beyond a carapace width of

about 23 mm even if both morphological consti-

tuents are still présent (Eloestlandt 1940: 21,

fig. 8; 1948: 55). The zone bearing the acute but-

ton is covered by setae, and the Socket seems the

first to disappear, even when the button is not yet

lost (Guinot 1979a: !42, pl. 23, fig. 1). In the

male niaterial of E. japontea (de Haan, 1835) that

was examined, the button is s ma 11 but remains

présent, in a noteh loft by the setae of Gl, and the

Socket is weak. In spécimens of E. jormosa Cban,

Hung et Yu, 1995 (a species to bc referred to a

new genus, cf. Guo et al 1997) that were exami¬

ned, the button is always smafl, the setae on Gl

being in contact with the button; the diffuse Soc¬

ket seems to be ineffective.

In Armacopleura Stimpson, 1858 and Tetragrapsus

Rathbun, 1918, the button is blunt and the soc-

664

ZOOSYSTEMA - 1998 -20 (4)

Abdominal holding Systems (Brachyura)

ket weak, In Cyrtograpsus Dana, 1851, C. angula- Members of rhe genus Orcovita Ng et Tomasick,

tus Dana, 1852 was exarnined: ihe large males 1994, ali anchialine or cavernicolous, possess an

were devoid of locking structures; a male tneasu- apparatus of the press-button type, wirh a mar-

ring 14.b X 17.4 mm bore a pair of minute but- ked burton but only a diffuse socket (to be car¬

tons, covered by the Cil setae, but liad obsolète rected in the diagnosis given by Ng, Guinot &

sockets; in a sntalkr indiv'idual measurjng 11.9 X llifie 1996), so that the abdomen seems to be not

14 mm, however, the buttons were already lost. efficiently retained by this mechanism.

Fig. 23. — Locking apparatus in Grapsîdae, subfamilfes Varuninae and Plagusiinae; A, Varuna litterala (Fabricius), i 29 x 31 mm,

Madagascar (MNHN-B 16777); B, Eriocheir sinensis H. Milne Edwards, <5 71 x 75 mm. Escaut (MNHN-B 12581): button absent

(setae were partly removed from right G1 ); C, D, Plagusia glabra Dana, near Sydney (MNHN-B 11700): C, ovîgerous 9 35 x 38 mm:

button persistent as a scar; D, a 32 x 33.5 mm. Abbreviations: b, button; s, scar of the button; v, vulva; 5, 6, sternites 5, 6.

ZOOSYSTEMA • 1998 -20(4)

665

Guinot D. & Bouchard J.-M.

SubfamiiY Sfsarminaf. Dana, 1851

The subfamily Sestirminae, an artificial grouping,

is among thc l.irgest brachyuran subfamilies, with

représentatives thaï are imertidal, often mangro¬

ve dwellers, or adapced to (reshwaier, or very ter-

restrial (Ng 1988). In the scsaxmine species rhat

were examincd rhe burton was small and general-

ly did not work aller a certain size (sometimes at

a size well beiow thaï of puberty), or was obsolè¬

te or absent; il présent, the Socket was wcak.

In Sesarma Say, 1817 (type species 5. reticulatum

Say, 1817) and in al lied généra the configuration

is not constant. The sternite 5, often covered by

setae, may bc covered by Gl. In the male niate-

rial that was examincd rhe small-sized indivîduals

had a very small button, which could disappear at

a partieular molt, and a soeket whieh was imper¬

ceptible imder the binoettlar.

In Sesarma ( Chiromantes) huzardi (Desmarest,

1825) the locking mechanism seems to be func-

tional. A well-developed protninence was obser-

ved on sternite 5 as well as a small soeket in the

postéro-latéral corner ol pleomere 6. This appara-

tus remains apparent!)' constant and functional at

any (molting stage) size of the individuals. In

S. ( Perisesarma ) alberti Rathbun, 1921 a small

sternal prominencc is observable at any growing

stage ot the individuals despite thc soeket not

being well-dilfcrentiatcd.

In Sesarma (Chironutnles) gultatum A. Milnc

Edwards, 1869 the button disappears beyond the

size 12.9 x 16.2 mm. In S, (Holnmetopus) ricordi

H. Milne Edwards, 1853, small individuals (cl less

than 7 nim) show a normal locking apparatus, but

this becomes non-functional in larger males: only

vestigial rubercles remain on thc sternum and they

become smaller with incTeasing body size

(Hartnoll 1965: 133). Young individuals of

Neoepisesarma ( Neoepisesarma) mederi (H, Milne

Edwards, 1853) hâve a minute button near sternal

suture AI 5 and a small soeket, but these structures

tend to disappear at large sites, In Neosarrnatium

meinerti (de Man, 1887) the button is présent in a

male measuring 31 X 37 mm but is lost in a male

measurïng 36.5 x 42.7 mm. In Sarmatium Dana,

1851 the button is small or absent, and there is no

soeket.

In other sesarmine species the absence of buttons

even before the moult of puberty in males is pos¬

sible. In living Sesarma ( Sesarma) curacaoense de

Man, 1892, Hartnoll (1965: 139) nodeed the

absence of any abdominal locking apparatus.

S. (Sesarma) reticulantm, S. ( Chiromantes) haerna-

tûcheit (de Haan, 1833), 5. (Chiromantes) dehaa-

jiI H. Milne Edwards, 1853, S. (Chiromantes)

guttatum A. Milne Edwards, 1869, S. ( Geose-

sduna) maculation de Man, 1892, Parasesnrma

plicatum (Latreille, 1803) perhaps also, but an

examination ofspecimetis oF ail si/es is necessary.

No button was observed in Neusarmatiurn smithi

(H. Milne Edwards, 1853), N. meinerti (de Man,

1887), and N. trisptnosum Davie, 1994.

In the examinée! species of the généra fleliee de

Haan, 1835 and Pseudohelice K. Sakai, I iirkay et

Yang, 1996, only a very minute tubcrcle exists on

sternite 5 in males. It does not correspond ro any

(perceptible) soeket, and locking does not appear

possible. In H. erassa Dana, 1851 and

Pseudohelice leachii (Hess, 1865) the tubercle

disappears beyond a certain size in males. In ail

the examincd male individuals ol H. tridens de

Haan, 1835 a second minute granule is présent

on the posrerior portion of somitc 5.

In Cyclograpsus H. Milne Edwards, 1837 the but-

ton is small or lost and the soeket absent. The

apparatus is vestigial in small individuals of

C iuteger IL Milne Edwards, 1837 (Hartnoll

1965). The sternite 5 is covered by the long male

First gonopods in C. purlctütus 11. Milne Edwards,

1837. while in C. integer the gonopods are xhor-

ter and the sternite is free.

In MetapLtx El. Milne Edwards, 1852 the button

is absent or doubrful. and there is no apparent

soeket. In Chasmagnathus de Haan, 1833 the

button is small and there is no discernible soeket.

In Aratus pisonii (H, Milne Edwards, 1837) no

trace of button or soeket was lound, even in

males. Observing living animais front Jamaica,

Hartnoll (1965: 121. 145) recorded that thc loc¬

king mechanism is completel)' lost in this tree

climbing crab. lt was remarked rhat in sesarmine

species the disappearance ol the apparatus is not

accompanied by any overall change in the abdo¬

men appearance, while in Aratus H. Milne

Edwards, 1853, the male abdomen is large,

somewhat resembling that of mature fetnales.

Hartnoll suggesred that this “femini/.ation" could

perhaps account for the loss of the locking System.

666

ZOOSYSTEMA • 1998 > 20 (4)

Abdominal holding Systems (Brachyura)

FAMILY OcYPODIDAK Rafmesqac, 1815 topbthalmus (Palks, 1772), O. Sara tan (Porskâl,

1775), O. çtmlimtinus Latreillc, 1818, O. platy-

SUBFAMII Y OcYl’ODINAL Ralinesque, 1815 taras H. Milne Edwards, 1852, O. ajricana de

We examined most species of the genus Ocypode Man, 1881, O. wadagascanensis Crosnier, 1065.

Weber, 1795. The button is very small in O. cera- It is absent in O. atrsnr (Linné, 1758) (Fig. 24B),

Fig. 24. — Locking apparatus in Ocypodidae, subfamilies Macrophlhalminae and Ocypodinae: A. Macrophthamus (Venitus) latreillei

(Desmarest), n 38 « 51 mm, NW coast of Madagascar (MNHN-B 12614): bulton présent; B. Ocypode cursor (Linné), t 36 >. 45 mm,

Dakar (MNHN). bulton absent; C, Uca (Boboruca ) Ihayeri Rathbun. ' 20 - 31 mm, French Guiana (MNHN-B 12049): two horizontal

carinae on siernite 4; D Uca (AtrucaI tangeri (Eydoux), rî 26 x 39 mm, Dakar (MNHN-B 12027): a marked tuberde and additional

granules. Abbreviations: b, button; c, carina on sternite 4; g, G1 ; gr, granule: tu, tubercle; 4. 5, 6, sternites 4, 5, and 6.

ZOOSYSTEMA • 1998 -20(4) £,(pj

Guinot D. & Bouchard J.-M.

O. qiuulrata (Fabridus, 1787). O. gaudichaudii

H. Milne Edwards et Lucas, 1843. The socker is

vestigial ot absent. In O. pygoides Orrmann, 1894

there is only a vestigial button.

In the large gcnus Uca Leacb. 1814, which has

been subdividcd (Bott 1973; Grime 1975; von

Hagen 1976). the configuration is vaxiable.

According to the species, there is either a button,

small but acute, or a granular zone, or a longitudi¬

nal set of granules fwith the anterior one more

developetl) located under the abdomen, but no

differentiated Socket is discernible. In the case of a

unique button, it is ttot covered by the relatively

long pleopods which are more medially situated.

We use b}' convenience the nomenclature of Gratte

(1975) instead of Bons names (1973) (see von

Hagen 1976; Manning & Holthuis 1981: 224).

In liai (Afi-uca) Utngerï (Eydoux, 1835) there is

generally a marked tubercle jusr behind the ster-

nal suture 4/5, in a setose région, and somerimes

one or even two additional granules are in the

same row (Fig. 24D). In U. (Uca) maracoani

(Latreille 1802), along the external border of the

sterno-abdominal cavity, rhe sternite 5 beats a set

of arrangée! granules, which extend a little on to

sternites 4 and 6 (addirionaUy, a pair of small

carinae is présent on sternite 4, see below). In

Uca (U.) primeps (Smith, 1870) rhe granules are

larger and more numerous on the anterior part of

sternite 5, which appears slightly elevated.

Species belctnging mostly to the subgenus

Minuca Bott, 1954 are characrerized by an addi¬

tional structure, called crans d'arrêt (safety catch)

by Guinot (1979: 153). In Uni (Mimait) vocator

(Herbst, 1804), U. (M,) mordax (Smith, 1870),

U. (Al. ) rapax (Smith, 1870) (Fig. 26E) and in

U. (Boboruca ) thayeri Rathbun, 1900 (Fig. 24G)

there is a rather setose, somerimes minurely gra¬

nular area, a small tubercle well visible behind

sternal suture 4/5, and a spécial structure that

arises front a denticulated crest that encircles the

end of the sterno-ahdominal cavicy. On each side,

in the middle of sternite 4, tins crest is prolongée!

internally by two short and srnooth (under rhe

binocular) carinae, which are practically horizon¬

tal but directed towards rhe slope of the cavity

and penetrate inside the sterno-abdominal cavity.

This pair of carinae corresponds to the limit bet-

ween abdominal segment 6 and telson. It is sug-

gested thaï, when the ctab inserrs its telson in the

deep sterno-abdominal cavity' (animais preserved

in alcohol do ttoi show any muscular tonus), the

tluckened anterior fiait ot the telson is held b y

the carinae, which immobilises the abdomen. Ail

the Uca (Minuca) species that Were examined

posseSsed a pair of carinae.

Othet Uca species hâve a similar pair of carinae,

as in the case ol Uca ( Ccluca ) pugilator (Bosc,

1802) in which the carinae are thickened at their

distal end. In Uça (Ccluca) urugayensis Nobili,

1901 thèse carinae are markedly curved npwards.

SUBFAMILY HhLOECIINAU H. Milne Edwards,

1852 (= HelOECHNAE Türkay, 1983)

Heloecius Dana, 1851, considered as the mosc pri¬

mitive genns wirhin the Family Ücypodidae, has

been removed from the subfamily Ocypodinae to

be placed in the subfamily Fleloeciinae, in which

the genus Ucides Rathbun, 1897 wasalso (at least

provisionally') included (Türkay 1983; Fielder &

Greenwood 1985;Tavares 1990). In II. cordifor-

mts (FI. Milne Edwards, 1837), an intcrtidal and

burrowing crab active ai low water in the man¬

grove, and Ucides cordcilus (Linné, 1763). that

occurs snpratidally (Harrnoll, 1988), no button

or soeket was observed, In these two gênera, the

sterno-abdominal cavity is very deep and long,

the G 1 reach the level ol sternite 4 (then eovering

the sternite), and the somites with the usual loc-

king press-button (sternite 5 and pleomere 6) do

not correspond. Ucides is notahly distinguished

by its abdominal segments 5 and 6 fused in a

rigid plate, whereas Heloecius has a wider and

normally segmented abdomen.

Subi AMII Y Macrophthai minae Dana, 1852

In Macrophthahnus l atreille, 1829 (see Komai et

ai 1995) a part of the species examined showed a

tubercle, somerimes acute and always behind the

sternal suture 4/5, and a shallow socker in the

middle ol abdominal segment 6. Macrophthahnus

(Vendus) latrnllei (Desmaresi, 1822) (Fig, 24A)

represents an example of an acute button, proba-

bly providing an effective locking mechanism. In

M. (Macrophthahnus) dilatants de Haan, 1835,

the socker is posirioned in the anterior part of the

abdominal segment 6, and the apparatus seems to

fonction. In M. (Macrophthahnus) milloti

668

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Crosnier, 1965. M. ( Macrophthubnus) brevis

(Herbst, 1804), M. {Macrophthaïmus) consobri-

nus Nobiti, 1906 and M. (Mopsocarcinus) boscii

Audouin, 1826, the System seetus to be effective.

Other macrophthalmine speties bave a System

which appears weak and perhaps even inefficient.

In M. ( Hemiplax ) binipes (Jacquinot, 1853) there

is a button, but a Socket is not perceptible under

the binocular. In M. ( Marrons) depressus Riippell,

1830, and M. (Mareotis) hantai Wanda et K.

Sakai, 1989, rbc buttons become very minute.

M. [Mareotis) japonicus (de Haan, 1835) has lost

both parts, sternal and abdominal, of the locking

apparatus.

SUBFAMILY DOTILLINAK Stimpson, 1858

(= SCOI’IMI.RINAF Alcock, 1900)

In Dotilla Stimpson, 1858 the sterno-abdominal

cavity is very deep, with well-defined margins.

The abdomen is perfectly inserred in the cavity

and its ventral part is almost completely covered

by the swollen hindgut. In D. myctiroides

(H. Milne Edwards, 1852), D. fenestmta

Hilgendorf, 1869, D. wiebmar/ni de Man, 1892,

and D. tmthtbarica Nobilt, 1903 there is a mi¬

nute prominence, very close to (almosr on)

suture 4/5, but no socker is discernible, ar least

under a binocular. In D. sulcata (Forskal, 1775)

the suture 4/5 is thicketred and slïghtiy carinated,

without a button, and no socker is perceptible.

In the two examined speciesof Dolillopsh Kemp,

1919, D. brevilarsis (de Man, 1888) and D. pro¬

fita (Nobili, 1903), the anterior segments of the

abdomen are inserted in a relatively broad cavity,

and the pleon is markedly constricted and well-

inserted at the level of the basal part of seg¬

ment 5. The posterior end of the abdomen is

included in a deep and narrow sterno-abdominal

cavity. No button or Socket is visible.

In ail speeies of Scopirnent de Haan, 1833 thar

were examined, lhere is a small button behtnd

suture 4/5 and a Socket in the postero-lateral

angles of abdominal segment 6. The mcchanism

is functional.

Family Camptandriidae Stimpson, 1858

A subfamily level was proposed by Stimpson

(1858) but, for a long time, the genus

Camptandrium Stimpson, 1858, was referred to

diverse groups, being attributed to the Macro-

phthalminae wtthin the family Ocypodidae

(Balss 1957). Serène (1974) formally defmed the

subfamily Camptandriinae and included the

généra Leipocten Kemp, 1915 and Pameleisto-

tsoma de Man, 1895, whereas ar the sanie time

Pretzmann (1974) established the “riew” rribe

Camptandriini. Manning & Holthtiis (1981:

193, 348), who described fîve new West African

généra, recognizcd the subfamily. Reviewing the

genus Barumi Stebbing, 1904, Harminto & Ng

(1991) emphasized the unique eharacters of the

camptandriine group, in particular the unusually

recurved Gl. The taxonomy of the camptan-

driines, which consists ot nearly rwent) r généra,

was revised by Tan & Ng (1994) and Al-

Khayat & Jones (1996), and awaits being raised

to family status Camptandriidae (Ng 1998 and

pers, connu.). The camptandriid crabs usually

live in cstuarinc, especiaily mangrove habitats.

In Tylodiplax de Man. 1895 and Baruna

Stebbing, 1904 there is a typical press-but Ion,

persistent throughout life. In ,S bénins anonialus

(Shen, 1935) the sternal tubercte is acure, and the

Socket clearlv visible. In Paracteistostamit depres-

sum de Man, 1895, Serenelbt leachii (Audouin,

1826) and Hyagynais microcheimm (Tweedîe,

1937) the button, situated close to the sterno-

abdominal margîn, varies Irom prominent to flat-

tened, and the socket, situated on the edge and

the whole lengrh of abdominal segment 6, varies

front slïghtiy hollowed to fiat.

The locking apparatus seems functional in ail

examined Camptandriidae.

Family Gkcarcinjdae Macleay, 1838

The male abdomen forms a relatively thin plate.

On stertrite 5, often covered with numerous

serae, tltere is a prominence with an uneven sur¬

lace; no abdominal socket is discernible, however.

In Gecarcimis lateralis (de Fréminville, 1835)

three male individuels nieasuring 22 X 30 mm,

39 X 32 mm, and 42 X 34 mm hâve two granules,

one larger than the other, very close to suture 4/5;

in a male individual nieasuring 46 X 36 mm the

granules are fused and are not rcached by the

male First gonopods. In G. planatus Stimpson,

1860 (Fig. 25A, B) a large oblique prominence is

located in the middle of sternite 5, with an un-

ZOOSYSTEMA • 1998 • 20 (4)

669

Guinot D. & Bouchard J.-M.

even and setose surface; no Socket, however, is

perceptible on pléorrrcre 5.

In Cardhoma camijex (Herbsc, 1796), a male

measuring 59 X 68 mm shows 2-3 granules very

close to suture 4/5 in a seriferous zone; a male

measuring 83 x 69 mm shows a granulous pro-

minence plus a row ot granules, the apex of G1

being jusr behind the granules. DiiçopLix hirtipes

(Dana, 1852) lollows the same pattern.

Epigrapsttspülitus Heller, 1862 displays a relative-

ly strong and rugose buttoii in the middle of scer-

nite 5; rhe Socket is visible because of ils calcified

border.

Gecarcoitlea htlandii H. Mil ne F.dwards, 1837

(male individual measuring 50 X 67 mm) shows

a wide and rough prominence on rhe seriferous

sternire 5, probably with particular microstruc¬

tures, but no corresponding Socket.

Family Mictyridal Dana, 1851

In Mictyris Farreille, 1806 (type species: M. lon-

gicarpus Latreilie, 1806) rhe sterno-abdominal

cavity is not deeply hollowed and its margins are

not well-defined latérall.y. The abdomen is long,

particularly wide and expanded, except for the

small telson; its anterior segments are tightly

inserted between the P5 coxae and srernire 8

(Guinot 1979a, fig. 29C). In addition, in M. lon-

gicarpus an oblique protubérance on each side of

sternite 8, just behind the suture 7/8, props up

the pleomere 2.

The five species of Mictyris tliat were examined

do not hâve a sternal bution or an abdominal Soc¬

ket. In the males (no female was observed) ot

M. platychcles Fl. Milne Edwards, 1852. two long

latéral projections arise Irom a crest on sternite 4

(Fig. 25C, D). The latero-external parts of pleo¬

mere 6. with no visible differentiated zone, lean

on the pair of projections, and the telson loosely

moves between these two srrong structures.

The crest on sternite 4 is wirhout projections in

mictyrids, except in only one species, M. platy-

cheles. lt is évident thaï the pair of projections in

M. platycheles is not homologous with a locking

apparatus of the ptess-button type: the telson is

simplv received in the space between the two pro¬

jections. Apparetuly latéral displacements of the

abdomen will be prevented due to the pair of

projections that guides the course of the telson.

GENERAL DISCUSSION

When observing living crabs or specimens preser-

ved in alcohol and when referring to published

data, ir is rare, if not exceptional, to find indivi-

duals with rheir abdomen not fixed to the ventral

surface. A retained abdomen is the rule for males

and also for prcpubertal females, while the mature

females acquire a wide abdomen that (usually)

cannor be held to the thorax any longer.

Mau and femai.e abdomens

Male and female abdomens hâve initially six dis¬

tinct segments, ail looselv articulated, plus the

telson. There i.s a tendency toward fusion of two

or several pleoineres in a more tigid plate, some-

times with the sutures no longer visible.

The abdomen is capable of srrong flexion and its

ventral folding varies front weak to complété.

The firsr segments are not completely flcxed

under rhe céphalothorax even in certain advanced

crabs (Guinot 1979a, fig. 1 H, 1). I he abdomen

may be long, entering the bases ol the Mxp3 and

in grear part folded, as in the Homolidae, or very

short and inconrpletely flexed under, as in the

Raninidae. The ventral inflexion ol the abdomen

is not an exclusively brachyuran condition, since

it occurs in orher Decapoda with a crablike faciès

(Porcellanidae for example). In Decapoda an

elongated abdomen may be regarded as the ple-

siomorphic condition, versus a short and flexed

abdomen in the apomorphtc condition. But the

developed, unprotected and unsegmented abdo¬

men in paguroids, whieh contains alniost exclusi¬

ve!)' the reproductive System, di tiers not only

Irom that of otber anomurans but Irom that of

decnpodx in general (McEaughlin 1980: 150,

Figs 4?A, 50A). The long abdomens, as well as the

short, well-calcified (with symmetrieal or tnore or

less asymmetrical tergal plates) and folded abdo¬

mens, are not similar among the Repfantia.

In the Brachynra, plesiomorphic character States

are die narrow tergal part of abdominal segments

accompanied by elongated latéral pleura and a

long telson, botb tound in Homolodromiidae

(Guinot 1993a. 1995). For Scholtz & Richter

(1995), the “original" decapod telson is tiiangu-

lar and pointed posteriorly; it is wide, more or

670

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

less square-shaped and posteriorly rounded in Dromiidae [note that the telson is not square in

their Eureptantia, and it remains relatively wide Homolodromiidae], Scholtz & Richter conclu-

and rounded in primitive brachyuran taxasucb as ded (p. 298) that the triangular telson found in

Fig. 25. — Locking apparatus in Gecarcinidae and Mictyrldae; A. B, Gecarcinusplanatus Stimpson, <$ 53 x 77 mm, Lower California

(MNHN-B 13154): A, thoracic sternum, witltout abdomen: B, ventral surface ol abdomen, without discemible sockets; C, 0, Mictyris

platycheles H. Milne Edwards, <J 13.3 x 12.3 mm, Australia (MNHN-B 16086): C, ventral view; D, detail ot the crest on sternîte 4 with

two projections. Abbreviations: a6, abdominal segment 6; b, button; g, G1 ; pr, pair ol projections on sternite 4; s.a., sterno-abdomi-

nal cavity; t, telson; 4, 5, sternites 4 and 5.

ZOOSYSTEMA • 1998 - 20(4)

671

Guinot D. & Bouchard J.-M.

most crabs eau be regarded as “a secondary alte¬

ration perhaps related to the réduction of the

pleon and the loss of the tail fan.”

In Podotrenrata, the plesiomorphic nature of the

male abdomen is demonstrated by the presence

ofpleopodal (birarnous or uniramous) rudiments

on the somites 3-5 (most Homolodromiidae, ail

Dynomenidae, and a Icw Dromiidac). fhese

same threc familics are charaeterized by presence

of vestigial uropotLs, which are represented either

by dorsal plates (ail Dynomenidae, most

Dromiidac) or ventral lobes (ail Homolodro-

miidac, and a few Dromiidae) (McLay 19f>3;

Guinot 1993a, 1995) tn the other podotreme

groups (Homoloidea, Cyclodorippoidea, Rani-

noidea. Le. the Archaeobrachyura Guinot, 1977)

and in ail mena bers of the Heterotremata-

Thoracotremata assemblage, thcrc is neifher a

trace of vestigial pleopods on somites 3-5 in

males, nor a trace of appendagelike, platelike or

lobelike uropods in both sexes (Table 3). The per-

sistence of pleopods as rudiments on abdominal

somites 3-5 in males could be regarded as related

to the presence of vestigial uropods. In most of

the Brachvura, the male abdomen hecomes lim-

bless, except for gonopods 1 and 2.

In Brachyura, the rôle of the abdomen is very dif¬

ferent in the two sexes. Since the flrst rwo pairs of

pleopods change (relatively) little at puberty in

males, growth of the male abdomen is approxi-

mately isométrie throughout. The female abdo¬

men shows marked positive allometry in the juvé¬

nile phase and a large merease in size at the

puberty molt, which brings it to a functional size

when it is required. Compared to the chelae in

males which continue to grow, the female abdo¬

men is nor an independent effector but can ope-

rate only together with the sternum. Having

attained an effective size at the puberty molt, the

abdomen reduces its allometric growth, since any

further disproportionné merease would for

example make walking difficult (I lartnol! 1982:

173). The female abdomen, in conjunction with

the sternum, encloses a chamber that facilitâtes

the fixation of the eggs to the pleopods and their

protection during incubation. In many groups,

during incubation, ovigerous females do not hâve

their abdomen hc-ld closcly against the thorax

because it is displaced by the egg mass.

Sti rno-aboominai CAVtrv

In brachyuran crabs the rhoracic sternum main-

tains the ancestral metamery, at least in the région

formed by somites 4-8, that is, corresponding to

thoracopods 4-8 (= pereiopods 1-5). The sternal

plate, which shows an isométrie growth (Drach

1959; Guinot 1979a), constitutes an cssential

part of the organism lor phylogenetic studies. In

accordance with configuration of the sternal

pinte, two major groups may be disringuished:

the Podotremata and the Heterotremata-

Thoraeotremata assemblage.

In Podotremnra, the arrhrodial cavities of the per¬

eiopods are not aligned (but, as în ail Brachyura,

theyopen laterally, instead ofventrally in macru-

ran forms) and the sternum shows at least two

different planes, the posterait part being tilted.

The abdomen more or !ess fi Ils up the entire

width of the sternal plate, generally covering the

space berween the coxae of pereiopods, some-

tjntes extending over bases of the coxae, and is

rarely narrower than the sternal plate (Dynome¬

nidae, Latreilliidae). There is not a true sterno-

abdominal cavity, with defined margins, in tnost

podotreme males. In the family Dynomenidae, a

slight sternal dépréssion, without defined bor-

ders, receives the rather wide abdomen. Except¬

ions in the Podotremata are the Cyclodorip¬

poidea, which bave a relatively wide sternal plate,

and also the (fo.ssil) Dakoticancroidea.

In podotreme females, a strong modification of

the ventral surface occurs, wirh inclination of the

coxae and modification of their internai surface,

to form a wider space to receive the increased

abdomen 1 thîs coxal zone becomes completely

smooth and naked.

In the Heterotremata-Thoracotremata assembla¬

ge, the whole sternum tends to form a plate in the

same plane (the arthrodial -cavities of thoracic

pereiopods 1-5 arc- aligned) and to widen, some-

times extensively. Metamery varies froni having

ail sutures 4/5 through 7/8 parallel and complété

in a narrow plate (plesiomorphy) to being more

or less oblique and becoming partlv or complete-

ly interrupted in a broad plate (apomorphy). The

ventral surface is excavated by a spécial déprés¬

sion, with a sh.ipe that (more or less) corresponds

to rbe abdomen. The abdomen folds down on it,

completely covering the vulvae and pleopods,

672

ZOOSYSTEMA - 1998 - 20(4)

Abdominal holding Systems (Brachyura)

except for rare cases. In the Thoraco-tremata,

male sternal apùrtures also are covered.

Two synapomorphies of the Hecerotremata-

Thoracotremau assemblage are: présence of a

sterno-abdominal cavity wicb defined borders

(présent, however, in certain Cyclodorippoidea

and in Dakoticancroidea); présence of a press-

button, which is rypica! and remains practically

unchanged in ail eubrachyuran crabs.

Carcinization

In the process of carcinization (Borradaile 1916;

Tiirkay 1986; Maçpherson 1988; Lcmaitrc &

McLaugblin 1995; Poupin & Mcl.aughlin 1996;

McLaughlin 1997) which leads to rhe crab-like

form, i.e. rhe derivcd condition, diverse trends are

corrclated; réduction of the relative length of

céphalothorax, specially broadening ol die carapa¬

ce, front, buccal trame, branchial cavity, and ster¬

nal plate; réduction of the rostrum; tolding of the

antennules and antennae; general flattening of the

céphalothorax leading to cephalic condensation

with the sensorial organs in the sanie plane, and

with modification of orbits and eyestalks; réduc¬

tion and folding of die abdomen and fusion of

diverse pleomeres; partial loss ol metamery on

dorsal surface of the carapace and more signifî-

cantly on the ventral surface; loss of pleopods

(except for pairs I and 2) in males and of caudal

furca in borh sexes. Locomotion is no longer pro-

vided by rhe abdomen but assurned by the t h tira¬

de pereiopods. The strong réduction ol the abdo¬

men and its ‘ intégration’ to the body modifies the

center of graviry ol rhe animal. The thoracic ster¬

num broadens and torms a hard horizontal plate

in a sort of shield, rhat in a way parallels the dor¬

sal carapace. In the médial part of rhe sternal

plate, the excavation of a cavity, with delimited

margins to receive the abdomen, constitutes an

essential evolutionary step of carcinization.

Intercalation of a wide sternal plate between the

bases of the legs induces a grearer distance bet¬

ween the P5 coxae and gonopods, with the effect

that the cosal sexual male openings (and penes)

become far removed from the loramina at the

bases ol the gonopods. The moving (migration)

of male gonopores to a sternal position, close to

the médian axis in the vicinity of gonopods, is a

derived condition within the Brachyura as a

whole and also w ithïn certain familles.

Depending on the transformation of the cylindri-

cal shape to broad/compact and conical and

increasing ihoraçic muscular mass, the endo-

phragmal System significantly changes, 1rs new

organization contributing to the consolidation of

the whole thorax, with concentration at the apex

of its muscular attachaient to, or suspension

from, the carapace (Secretan in press). As a consé¬

quence of the réduction of rhe abdomen, the

functional removal of the impeding abdomen,

and as soon as ali internai readjustments were

complétée! with efficient reconstruction of tire

skeleton, niables a vast iinprovement in the

mechanism of locomotion. Thèse adjustments,

that were ovcrcome already in the Dtomiacea,

hâve produced the highly siiccessful brachyuran

group, wliose bodies are adapred for efficient wal-

king and running (Bauer 1989: 64, fig. 18). It is

a matter of tact, the Brachyura show strong adap¬

tative radiation and dispersai to terrestrial and

freshwater habitats (Glaessner 1957).

Brachyuran radiations

The three highest taxonomie units proposed by

Guinot (1977), which express by their naines the

change in the position of gonopores, correspond

grosso modo to the principal radiations in the

Brachyura and cati be documented paleontologi-

cally from the beginningof their existence at Icast

in the Earlv jurassic to the présent.

The Podoiremata hecame diversified in Middle

and Upper Jurassic sédiments. "Their actual repré¬

sentatives are ail marine, often inliabit deep

waters and show a commun tendency toward car-

rying behaviour vvith rhe last pair(s) of legs

(Guinot et al. 1995).

À remarkable synapomorphy distinguishes ail

otlier crabs, that is, the Heterotremata-

Thoracotremata assemblage: innovation of the

female gonopores on the sternal plaie, the vulvae,

and loss of isolated spermathecae (Tavares &

Secretan 1992).

The most diverse groups ol crabs, rhe Hctero-

tremata, most ol which survive to the présent, is

a product of Late Cretaceous-Early Tertiary.

Their actual members show full development of

ail locomotory pereiopods (except for the Dorip-

poidea and other small other groups such as

ZOOSYSTEMA • 1998 - 20(4)

673

Guinot D. & Bouchard J.-M.

retroplumids and palicids). They are generally

marine. Are excepted rhe true freshwater crabs:

the diverse families and ail représentatives are

heterotreme. Ir is to be noted that the families

Hymenosontatidae (with some metnbers adapted

to freshwater environmenrs),. Goneplacidae,

Hexapodidae, Pitinotheridae, and Cryptochiri-

dae were recently attributcd to the Heterotremata

(Guinot & Richer de Forges 1997, table 1).

The remai ning crabs belong to the Thoracotre-

mata, which are known frorn the Cenozoic. On

account ofthe last révision ol the classification by

Guinot & Richer de Forges (1997), they arc

almost entirely composcd ol littoral and terres-

trial speeies, ail distributed in five htmilies:

Grapsidae, Gecarcinidae, Camptandriidae, Ocy-

podidae, and Micrvridae. They are justly the

forms which hâve successhtlly exploitcd a varicty

of aerial environmenrs, vvith ail necessary adjust-

ments in physiology, behaviour, reproductive and

developntental strategies (BJiss 1968; Burggren &

McMahon 1988), They are rhe result of complé¬

té carcinization allowing advantageous locontoto-

ry r adaptation such as Fast tunning, prolonged

walking on land (migration), rree-climbing, bur-

rowing. Thus it is noteworthy that the classifica¬

tion based on morphological characters fits with

the ecological and behavioural (ethological)

habits. Because of their recent appearance the

Thoracorremata hâve had rclatively little tinte (or

radiation and probahly exemplily only the early

stages of évolution of their terresrrial compétence.

Abdominal rhtaining by the limbs

In the podotrematous families Homolodromii-

dae, Dromiidae, Homolidae, and Poupiniidae,

abdominal rétention (when existing) is achieved

by diverse means that involve various appendages

(Mxp3, P1-P2, sometimes even P3). The struc¬

tures overhang the abdomen or are coaptated

with its margin (Tables I, 2), prevent ir being lif-

ted up. In addition to rétention by the limbs, the

Homolidae and Poupiniidae display a press-but-

ton System (homolid press-button): both Systems

ntay act simukaneouslv. But, in the more advan-

ced homoloid family Latreilliidae, rétention by

the coxae of appendages was lost, and only the

press-button System is effective.

In the family Dynomenidae, only primitive

représentatives hâve their abdomen restricred in

its latéral rnovements by coxal structures of the

second pair of peteiopods. The more advanced

dynomenid speeies similarly restrict sideways

movenrents of their abdomen but by a sternal

srrut lute.

The Raninidae hâve no retaining apparatus. An

exception is the subfamily Lyreidinae, where the

peteiopods do nul play any rôle: instead a sternal

structure arising front stérilité 5 acts with a Socket

on pleomere 6.

In the Cyclodorippoidea, which possess a relati-

vely wide sternal plate and bave their abdomen

far removed Iront the coxae of the peteiopods, the

appendages are never involved. Spécial retaining

dcvices, with structures located on the sternum,

exisr in the htmilies Cyclodorippidae (coaptation

by engagement) and Phyllotymolinidae (block

System).

In specializing the brachyuran ntodel, the evolu-

tionary process has provided an abdomen that is

lar removed Iront the coxae of the pereiopods and

is completcly placed on the wide sternal plate.

This resuhs in a “migration' of retaining éléments

on to the thoracic sternum, and particularly front

P2 coxa on to sternite 5 (a transformation sériés

loLind within rhe family Dynomenidae). Thus is

explained the entergence of a pair of prominences

ou rhe sternite corresponding to P2, i.e. ster¬

nite 5.

In the advanced Podotremata and ail

Eubrachyura, the thoracopods are no longer

involved: they bccome free Iront ail functions

Other titan locomotion and ntay be active for

otber purposes.

In the Podotremata that are considered as the

most primitive, viz. Drontiacca and Homoloidea

(excepr l atreilliidae), the abdomen is retaincd by

thoracopods (Mxp3 through P3), rnovements of

which are very important in tltat retaining Sys¬

tem. At the level of P2-P3 the thoracic sternum

présents chnracterisric dépréssions, already obser-

ved in the Homolodromiidae. These stertto-coxal

dépréssions (Guinot 1995 ) aliow coxae of the legs

to eysily slip i nsi de during their movement and to

swivel up on their condylar articulation with

increasing amplitude.

Sterno-coxal dépréssions are présent in ail

674

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Homolodromiidae, Dromiidae (exceptions are

very rare: Pscudodrnmia , Ascidiophilus ), ail Dyno-

menidae, ail Homolidae, and in the Foupiniidae

(Figs 5D, 7A, 7D, 9C). It is remarkable chat thcy

are absent in the Latreilliidae, where the pcrcio-

pods do nor play any rôle and the fixation is

exclusively secured bv the press-button on sterni-

te 4 (Fig. 9D). Sterno-coxal dépréssions also lack

in ail brachyuran crabs in which perciopods are

no longer involved in retaining, that is in ail

Eubrachycira. A conclusion ntay be drawn: a

strict corrélation exista between the involvement

of pereiopods (P2-P3) and the presence of such

sterno-coxal dépréssions, exempliüed by the dif¬

ferent patterns found in the two related families

Homolidae and Latreilliidae.

Modifications occtir in the abdomen in relation

ro the retaining apparatus by the limbs.

Abdominal segment 6 may be markedly modi-

fied. On the one hand. in Podofremata it is

directly in contact with a differentiated structure

on the pereiopods: either it is remarkably

constricted, lot example in Dromia monodi

(Fig. 3A), or iis exlernal edge is thicketied and

hollowed for example in Lauridrurnia dehaani

(Fig. 2A, B), L. inWmedhi (Fig, 2C),

Hemispbaerodrornia manodiis (Fig. 3C, D), and

Exodromidia spttiosa (Fig. 5D). On the other

hand, a transformation occurs when the appen-

dagelike ttropod becomes a sexually dimorphic

dorsal plate (most Dromiidae, ail Dynomenidae)

and particularly when it has evolved into a Soc¬

ket, which is a major event (Homoloidea,

Lyreidinae, practically ail Eubrachyura) (see

Hypochesis about the homology uropod-socket).

The telson also may be significantly mndified: for

example the deep notch on les external edge in

the dromiid Ltiuridromia dehaani, the long and

sexually dimorphic spine at irs tip in the dromiid

genus Exodromidia , irs base (pleorelson) with a

very thin edge in the cyclodortppid genus

Clythrocerus. In the Homoloidea the telson plays

a rôle in abdominal rétention and is inserted bet¬

ween maxillipeds: tr is either overhung by

(Fig. 8D) or Pics into Mxp3 (sometimes there is a

combination of these two modaücies), with

modifications occurring in irs general shape and

on its external rnargin (Figs 8B, C, 9B).

Fasti-ninc; .and unfasthning

The musculature of abdomen, well stttdied in

Cancer pagurus (Pearson 1908), in Carcinus mae-

nas (Daniel 1931) and in diverse species

(Abrahamczio-Scanzoni 1942), consists of a pair

ot dorsal extensor muscles and a pair of ventral

flexor muscles for each segment. The musculature

pattern dépends on the number of segments,

either ail free or diversely fused, and also varies

with respect to sex. In crabs with a press-button,

the abdomen i$ attachée! at the level of

pleomere 6, which limits general movement of

preeeding segments, The telson alone, sometimes

very movable, is capable of frcely moving around

its articulation with pleomere 6, This configura¬

tion diables the telson to be moved alone, for

example while defecating, since the anus opens

on the telson (for a discussion on telson and anal

segment, see Schram 1986: 7).

In the literature concerning brachyuran crabs,

cases of tmfolded or extended abdomens are

genetally only menrioned during mating. In the

course of their numerous experiments on living

Plchygrapsus mnrmaratus and Carcinus marnas,

rcspectively, Vernet-Cornubert (1958) and

Demeusy (1958) (who took the locking Systems

into aeçount in the two sexes) do not indicate

cases ol unfolded abdomen except lor the mating

process. When the abdomen is expcrimentally

unfastened, action to refasten it on to buttons is

not immédiate; in Carcinus amenas it nceds a few

seconds to two hours (Kollmann 19.37: 80).

It is obvious that, in the narural posture, proba-

bly the resting position, the abdomen is closed

and attacher! to the thoracic sternum. The ques¬

tion is to know wherher mature males unfasten

their abdomen at a rime other than the mating

process (including pte- and posr-copulatory per-

iods) and also for the exuviation. The young male

and the female before the moult of puberty per-

haps do nur need to move their abdomen for any

purpose. In fact the abdominal flexing is poorly

studit'd in the Brachyura.

Abdomen flappingin males is, to our knowledge,

generally reported at the period of copulation

(Watson 1970, 1972; Hiatt, 1948; Hinsch

1968). Laboratory observations on mating beha-

viour of Chionoccetes bairdi indicate that the crab

may repeat several times the action ol lowering its

ZOOSYSTEMA • 1998 - 20(4)

675

Guinot D. & Bouchard J.-M.

abdomen, holding iropen for a few seconds and

chen flexing ir upward to a fully closed position

(Donaldson & Adams 1989: 44, tables 1, 5).

These authors report that males and femalcs nor-

mally llap their entire abdomen while defecating.

But a movemem of ihe telson alone might be sui-

ficient. Movements involving abdomen extension

are probably required during lhe procedures ol

species récognition and courcsnip. It is not sur-

prising that, at least in certain species, flapping is

an action performed during reproductive beha-

viour. For example periodic flexing ot the male

abdomen was observed during the grasping ol

females in Chiotwecetes opilio (Warson 1970:

1612). Ir was demonstrated that sacculinized

Rhithropanapeu* harrhii (Gould) showed an

abdominal flexing response when exposed to cer¬

tain peprides, similar to the pheromones relcased

by the cggs just beforc hatching (de Vrics et ai

1989). Rhizocephalan-parasitized Uemigrapsus

sanguineus (de Haan) llex their abdomen with a

waving action, while the sacculinid relcascs nau-

plii (Takahashi et al. 1997: 159).

Flapping Iras another tunction in mature and ovi-

gerous females (Bauer 1989: 66) than in males,

and spécial pheromones seem to induce abdomi¬

nal flapping in ovigerous females. It may be asked

whether abdomen fixation is permanent for pre-

pubertal males and females. It was hypothesized,

already by Pétez (1928a: 462), that females of

Carcintts nutenas naturally untasten the abdomen

during the moult, notably at puberty moult, just

Lrefore mating (the male attends the female until

she moults, aller whtch rhey mate immediarely).

This is eflectively possible in ail species in whicb

mating females mate only while soft, as in

Carcintts. ht the case where a hard female mares,

the mature female does not need to unfasten its

abdomen since it is not fixed. A similar behaviour

pattern, viz. an active involvement for abdomen

detachment, is necessary for inalcs of afl brachyu-

ran crabs since they copulate only while in the

fully hardened condition (Flartnoll 1969). The

only vvay to fasten and unfasten dre abdomen

seems to bc by a postero-anterior movement. In

living crabs the experimental unfastening is gene-

rally very difllculi.

In leucosiids, the abdomen adhères to the sternal

plate in males and, to the highest degree, in

mature females. Its detachment probably nécessi¬

tâtes a physiologica! mechanism when coptilating

with a hard female, perhaps a local décalcification

(Drach 1939, 1955, unpublished data). Hartnoll

(1969: 168) indicates that. in Pbilyra, copulating

femalcs are presumably not very reccntly rnoul-

ted: when the female evetuually excends ber

abdomen be fthe male] rapidly moves so as to

face her, and mating occurs."

The abdomen, in conjunction with the sterno-

abdominal cavity, serves to cover (with some rare

exceptions) the Inst rwo pairs of gonopods,

which act fbl' the sperm intromission during

copulation, and the female pleopods before the

puberty mole, it provides a protection lor the

apertures of spermathecae in the Podotremata,

for vulvae in the Eubrachyura (with somc rare

exceptions), and gcncrally for the pênes, cither

coxal or sternal. It is cleai that inclusion of the

abdomen between the perciopods or in a spécial

cavity obviâtes its possible impeding movements

during swimming, burrowing or walking. In the

great majority of specimens examined the sterno-

abdominal cavity was cleai). which certainly

proves a perferr seal. Brachyuran crabs lack gene¬

ral body cleatting but srill remain quile clean

(Bauer 1981, 1989).

Tllli CASE OF THE lAMll.YL hUCOSIIDAE

In leucosiids such as Leurosia a first considération

is the absence of any Socket on abdominal seg¬

ment 6 (Fig. 19D). This pleomere is very elonga-

te and lias its whole lengtli located on sternite 4.

The abdomen is fixed by a system that conspi-

cuously differs front the typical press-button. The

sternite 5 is prolongcd into a spur that pcrfectly

enters a notch of the abdominal margin between

segments 5 and 6. The spur is often completed by

anaJogous protrusions of sternites 6 and 7. The

leucosiine spur appears as att aurapomorphy for

the most advaneed leucosiid taxa. It would be

interesting to verify whether simple coaptations

by engagement (Fig. 18) lound in primitive

représentatives (Ebaliinae, Cryptocneminae,

Pltilyrinac pro parte) are corrclatcd to coxal male

gonopores and whether a coaptation by assem¬

blage with a spur tends to occur in advaneed

représentatives with coxo-sternal male gonopores

(Philyrinae pro parte , Leucositnae, see the pénis in

676

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

Fig. 19C, p). It is noteworthy thar this particular

kind of coaptation occurs in the advanced faniilv

Leucosiidae, characterizcd by a pronounced

cephalic condensation.

FAILURHS, LOSS or absence oe hoi.oinc,

Al'PARATUS

The absence of a morphological holding appara-

tus is rare in Brachyura.

Certain Podotrenrara do not présent any spécial

holding structures. Our investigations, however,

hâve shown the existence of sonie particular

devices. The shell-carrying Hypoconcha arcuata

and H. panumemh and the Gymonomidae hâve

their abdomen simply bent at tight angles, with

only its posterior part being normally applied

againsr the ventral surface. Burrowing Raninidae

hâve a very short abdomen: anterior segments are

in prolongation with the carapace and the post¬

erior ones not flcxcti, which does not allow a fas-

tening. Nevertheless a firm Ioclcing is found in

the subfamily Lyreidinae (Fig. 11), the abdomen

of which is a little longer, narrower, and better

included between the legs than in other raninid

subfamilies.

Some f leterotremata lack a locking press-button

or show a dysfuncrion of the mechanism

(Table 3). This essentially concerns crabs having

a narrow thoracic sternum, with sutures 4/5

rhrough 7/8 grosso modo parollel and continuons,

and with a devcloped médian line. Their sterno-

abdominal caviry is eirher absent or hardly/weak-

ly defined. This configuration appears to be rela-

ted to the character stare “short abdomen", thar

gives rise to a gap between the rwo parts usually

involved, t.e. srernire 5 and pleomere 6, A remar-

kable conséquence in females is that tbe vulvae

remain exposed on the srernire 6. The putative

conséquence manifests itself in diverse manners:

Case 1. No apparent structures (no buttons, no

sockets): Corystes , Pseudocorystes , Jouas , and

Gomeza (but, at least in G. bicomis, a very short

sterno-abdominal cavity is présent posteriorlv,

and closing is provided in particular by a bending

of the posterior parr of rhe abdomen). The.se four

généra belong in rhe family Corystidae (Fig. 12A,

B, D, E).

Case 2. Presence of acute and posteriorly directed

hooks close to suture 5/6, but located so far from

the abdomen that locking is not possible:

Nautilnmrystes (Fig. 12C), which bclongs in the

Corystidae, Absence of sockets in postiarval

forms and in juvéniles needs to be confirmcd.

Case 3- Presence of both differentiated sternal

tuhcrcles and abdominal sockets. But in certain

variably si/ed adulr individuals a non-correspon-

dence occurs between rhe rwo parts, that became

sépara ted by a gap: Atelecyclus rotu ridât us

(Fig. 13A, B), and Peltarion (Fig. 14A) The

mechanism was effective only in some of rhe

individuals. The loss was apparently not related

to si/.e as shown by examining about thirty male

specirtiens. Atelecyclus and Peltarion belong to the

primitive family Atelecyclidae, in which a true

Sterno-abdominal cavity, however, is présent

(unlike rhe Corystidae).

Case 4. Asymmetry in disposition caused by a

gap between the two structures on onc side of the

body. In Ttachycurcinus (Fig. 1.313) the mecha¬

nism generally functions, but a dyslunction

occurs in old male individuals with one heùvy

ckela.

In addition, the case 1 is also found in crabs in

which the thoracic sternum widens, with ail the

sutures interrupted: Orithyiidae. A sterno-abdo¬

minal cavity is présent, which is elongared but

not covered by the short abdomen, so that the

vulvae remain exposed in females.

The case 2 is also found in crabs having a wider

thoracic sternum, with ail the sutures interrup-

ted, but with a “too short” abdomen: Bellia pieta

(Fig. 14B, C) in the family Belliidac. Despitc the

presence of both prominences and sockets, there

is no possibility of locking. It is to be noted that

the mechanism worlcs in young females. The vul¬

vae arc always exposed in females of this species.

The non-coincidence of the locking structures

occurs in the shorter abdomen of Bellia, that is,

in the most primitive représentative of the family

Belliidae and a typical burrowing form. The two

other généra of the family (Autnthocyclus and

Ileterozim), with a well-excavated srerno-abdomi-

nal cavity, a longer abdomen, and “normal” vul¬

vae, hâve a functional press-button: they are more

carcinized and advanced forms, nor so speciafized

for burrowing activity (note that Heterozius is

free-living but often is found decply burrowed

under boulders, C. L. McLay pers. com.).

ZOOSYSTEMA ■ 1998 • 20 (4)

677

Guinot D. & Bouchard J.-M.

In the family Tliiidae, also rather primitive, tlie

abdominal locking does not seem to be very firm.

In conclusion, the primitive condition in ail the

preceding familles ts confirmed by the failure of

the lockîng System (complété or partial absence

of structures or non-correspondence between

sternal and abdominal structures), evcn at the

individual level. Thls plesiomorphic State seems

to mainly characterixe burying or burrowing

heterottematous fortns.

In the l’iniiotheridaea prcss-button vvas observcd

only in hard-shelled stages, that may bear a large

knob on stctnite 5 and a smaller one on sterni-

te 6, which is very unusuak the soft-shelled stage

seems to be devoid of anv morphological struc¬

tures. In the Cryptochindae, or gall crabs, no

trace of an apparatus is visible. That recalls the

commensal dromiids Pseudodroniia and

Ascidiophilus (their body is enclosed in an asci-

dian) where also lacks an abdominal rctaining

mechanism. Absence of any morphological appa¬

ratus seems to be related to the protection allor-

ded by the host.

A crucial question is to déterminé where the

press-buiton lias been established in the lietero-

treme lincagc, since sternal prominences combi-

ned with sockets are an innovation of the

Eubrachyura (for t he case of the Homoloidea and

the Lyreidinae, see below). An investigation of

corysrid and atelecyclid fossil généra probably

would give good rcsults concerning the common

ancestor. It is likely that the ancestral condition

in corystids was with a complété apparatus.

Within the Thoracotremata, the press-burton is

very efficient in diverse groups (Grapsinae,

Plagusiinae, Camptandriidae for example) but

tends to become reduced, obsolète and non-fonc-

tional (Varuninae, Sesarminae), or completel y

disappears in the most advanced taxa (essentially

in the Ocvpodidae). The abdomen is no longer

fastened but, however, remains applied against

the sternal plate in the rather deep sterno-abdo-

minal cavit)'.

The most primitive Thoracotremata are the grap-

sid subfamilies Grapsinae and Plagusiinae, with

the male gonopore adjacent to the P5 coxa

(Guinot 1979a, fig. 52A-C): they hâve an efficient

press-button. Ail ot them are truly marine forms,

either shorc crabs (rock crabs) or largely aquatic

( Pennon ), with the exception of some Geagrapsus

species which show terrestrial adaptations. The

monophyly of either of these rwo fàmilies is well-

supported by their locking Systems (the status of

certain grapsine généra perhaps needs to be re-

evalutated).

The Sesarminae, in whîch the proximity of the

gonopore varies front very close to P5 coxa to

rather far away (Guinot 1979a, fig. 52D, I, J),

hâve a press-button which is either normal and

effective or inefficient beyond a certain size, or

complété!}' lost. The Varuninae, in which the

gonopore is more or less far on sternite S (Guinot

1979a, fig, 52F-} I), obviously do not lorm a

monophyleric group; rnany représentatives show

a press-button winch tends to be lost. It would be

irtteresting to compare the rcduction/loss of the

locking Systems with regard to the Jifestyle

(Burggren & McMahon 1988).

In the Camptandriidae (Guinot 1979a,

fig. 33C), that ccrtainly is a monophyleric group,

the press-button configuration is very homoge-

neous, with an effective mechanism that persists

throughout life in males: note that their pleopods

3re not very chitinized.

The Gecarcinidae, with a gonopore either close

to the P5 coxa or only a little away (Guinot

1979a, fig. 54A-D), may show a button but no

defined Socket. It is notevvorthy that Ppigrapsus

politui, which has a less terrestrial habitat rhan the

test of the family, has a strong apparatus in

contrast with the others where the locking is not

effective.

The Ocvpodidae. in which the gonopore is Sternal

and in relation to suture 7/8 (Guinot 1979a,

figs 53A, B, D, 54E), are more advanced than

Grapsidae, lhe Macrophthalminae and rhe

Dotillinae (= Scopimerinae) hâve a press-button

which is more or less efficient, depending on gene-

ra/species. In contrast, the Ocypodinae, some of

which hâve vestigial but inefficient buttons, and

the Heloeciinae seem to never retain their abdo¬

men by this device. Hat also seem to hâve, at least

in the exatnined preserved material, their abdomen

only loosely applied. The pair of carinae on sterni¬

te 4, that characterizes the suhgenus Mirtuai for

example, constitutes a new kind of apparatus. In

Mictyridae, in which locking by a press-button is

678

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

no longer possible (Guinot 1979a, fig. 29), the develops a new kind of rétention on sternite 4 that

large abdomen seems to be not firmly retained, restricts latéral movement of the abdomen in its

even in the species (.Mictyris platycheles) which posterior part.

Fie. 26. — Photographe with the scanning électron microscope; A, B: Plnnotheres pisum (Linné), 3 4.7 ■ 4.6 mm. at the liard stage,

La Rochelle (MNHN-B 10604); A. button; B. Socket; C. D. Austinograea alayseae Guinot. i 25.3 ■ 39.6 mm, Pacllic Océan Lau

back-arc Basin, BL 10, 1750 m (MNHN-B 24055); C. button; D, sockel; E, detail of the granular zone on stemites 5 and 6 of Uca

(Minuca) rapÿ A'(Smith), 3 15>:23 mm, West Indies (MNHN-B 22540) 1 F, button of Plagusia Immaculata LamaroK, 3 29.1 x30.7min,

Pondichéry (MNHN B 11702). Abbrevtalions; a6, abdominal segment 6, t, telson 5, slêrnite 5. Scale bars: A, 10 pm; B Q F, 100 pm;

F, 1 mm.

ZOOSYSTEMA • 1998 -20(4) 679

Guinor D. & Bouchard J.-M.

This suggesrs rhe following remark, The Thora-

cotremata forms chat became independenr faim

the sea (induding chose occurring intertidally but

with frequent aerial exposure, or living supraiidal-

ly, or being adapted to life on land) show a trend

not to fix their abdomen by a press-button appa-

ratus and to replace it with another System.

Microstructure

Investigations with the scanning électron micro¬

scope are in progress (J.-M. Bouchard) with the

aim to point our the présence ol nticrostructures

on sternal prominences and sockets. At the pré¬

sent tinte, with a sample of représentatives taken

from about fifteen brachyuran families, a great

variety of microstructures has been discovercd on

the buttons, differing in relative size, orientation,

general shape, detailed morphological features,

ornamentation, and setae. These preliminary

results permit rhe following First continents.

The microstructures of the sternal prominences

might represent a valuable criterion at the generic

level (for example in the Lyreidinac they distin-

guish the two généra Lyreidus and Lysirude), and

hypothetically at higlter levels. They are an indi-

cator of sex (being présent in males and beco-

ming modifted itt temales). puberty moult (at

least in females), and the âge ol individuals. For

example in Lysirude ehanneri the hooks of the loc-

king projections arc wcll-dcvdopcd in ntales and

blunt or even obsolète in temales (Fig. I IC, D).

The variety of patterns shown by rhe promi-

nences is exemplified by Figs 10D (Hontolidae),

26A (Pinnotheridae), 26C (Bythogracidae), and

26F (Grupsidae Plagusiinae). It is obvious that

microstructures improve the contact hetween

involved surfaces, enhance the assemblage of

both parts and increase cfficiency of the locking

mechanism. U is tioteworthy that microstructures

were observed as well among the Podotremata, in

the Lyreidinae (sternal projection from srernite 5)

and in the Homolidae (homolid press-button on

sternire 4), as in the Heterotremata-Thoracotre-

rnata assemblage (typical press-button on stérili¬

té 5). Phese microstructures mighr be acceptcd as

valid indic.itor.s ol phylogenetic affinities, Le.

synapomorpliies, except if they are suspected of

reflecting convergence

Investigations concerning the microstructures of

the sockets might similarly be taken into accaunt:

different patterns in oriemarion, shape, and

depth hâve already been observed (Fig. 26B, D).

TI IL ABDÜM1NA1 RETAINING MECHANISM AS A

PI IYLOGliNETlC CRITERION

The détermination of what might be the primiti¬

ve or advaneed mode of abdominal reraining

mechanism among Brachyura is basically a matter

of funcLional character polarity assessment. The

method of outgroup analysis is not informative.

In advaneed Detapoda a general trend toward

protection of abdomen is observed. "Macruran”

forms and some galatheid possess strongly calci-

fied abdominal tergal plates; paguroids hâve their

long and soft abdomen sheltered inside a gastro-

pod shell or protecred inside wood; crablike por-

cellanids, lithodids and lomoids show abdominal

réduction and flexion benearh the hody, without

any apparent retainingstructures. In contrast, bra-

chyuran crabs f'old their reduced abdomen and

obviate its possible impeding movements in two

basic ways: a holding by the thoracopods and/or a

locking by a press-button sysrem (homolid press-

button and typical press-button).

In utilizing the method ofingroup analysis, hold¬

ing by the limbs would indicare rhar this mode of

operation is rhe most primitive. In Podotremata

the retaining type that is used. with its detail,

may offer a reliable criterion in recognizing podo-

rreme familles. Careful srudy of the évolution of

rhe form of both sockets and buttons, with uni-

fomiity in general shape, position, and function

indicates that the press-button is a very successfi.il

mode of abdominal holding. This uniformity

within the wide diversitv of bodics in the

Eubrachyura probably shows that the press-

button is the most efficient use of their cnergy

resources in order to achieve locomotion, repro¬

duction and other fonctions. The retaining or

locking mechanism is an al most constant feature

in Brachyura. Its absence in diverse advaneed

Thoracotremata is considercd to lie the resuit of

a sccondary loss: its replaecmenr by an alterna¬

tive method romains to be cxplained.

Iil the Podotremata a sternal différentiation cvol-

ved independtnteiy in several lineages: dynome-

nid, homoloid, lyreidine, cyclodorippoid (cyclo-

dorippid and phyllotymolinid).

680

ZOOSYSTEMA • 1998 • 20 (4)

Abdominal holding Systems (Brachyura)

Some of che dynomenids (. Acanthodromia > Para-

dynotnenc, Dyuomene pro parte cxcmplified by

D. tanemis ) bave coxal structures on P2 (plesio-

morphy), but several of them (typical Dynontene,

with D. hispida) liave acquired a sternal structure

(apomorphy). Dcspite the links Dromiidae/

Dynomeiiîdac within the Dromioidea, advanced

dynomenids display an innovation with regard to

dromiids in which the structures always dépend

on limbs. In these dynomenids the pair of sternal

différentiations is posirioned on thoracic somite 5

(more preciscly on epistcrnitc 5) jList facing the

large dorsal uropods. In dynomenids, ncither

abdomen nor uropods arc modificd (wberças in

dromiids the abdominal 01 uropodal border is

ventrally or laterallv modlfied), Modification of

coxal structures into sternal structures is a consé¬

quence of the significant broadcning of thoracic

sternum (Figs G, 7). Thèse coxal or sternal struc¬

tures are similar in their morphology and connec¬

tion with abdomen (in overhanging, touching or

lying quite apart from the abdominal margin)

and play the same rôle in restricting its latéral

movement at the very most. The dynomenid ster¬

nal structure has the same location, on srernite 5,

as the button in the typical press-butron of the

Eubrachyura, but the involvement of the abdo¬

men appears as quite different, in dynomenids it

is not a press-button System: the abdomen is only

loosely applied on the sternum and is' only res-

tricted in its sideways movements. An essential

point is that the abdominal part (uropod indu

ded) is never modified in the dynomenid confi¬

guration. Such a combination is not inconsistent

with the idea thaï sotkets (in Homoloidea and

Eubrachyura) are issued from uropods.

In brief, within the Brachyura a sternal structure

has appeared:

- Once in Dynomenidae (sternite 5), without

sockets.

- Once for ail the Homoloidea (sternite 4): the

homolid press-button is practically similar in the

Three families (llomolidae, Latreîlliidae. Poupini-

îdae) and exhibits microstructures. The pair of

sockets hollowed on abdominal segment 6 shows

a configuration similar to sockets of the typical

press-button in the Heterotremata-Thoracotre-

mata assemblage

- Once in the Raninoidea, but in a single subfa-

mily, the Lyreidinae (sternite 5), with sockets.

The concerned somite is the same as in dynome¬

nids and in the Heterotremata-Thoracotremata

assemblage, Locking is very efficient, even in

females with eggs.

- Iwjce in the Cyclodorippoidea (sternite 6),

probably separarely in the Phyllotymolinidae and

in the Cyclodorippidae. But no corresponding

sockets exist, the telson being involved in locking

by its movements along rhe sternal structures.

Locking is very efficient.

- Once in the Eubrachyura (sternite 5), with soc¬

kets.

Two synapomorphies for the Heterotrcmata-

Thoracotremata assemhlage are: presence of a

sterno-abdominal cavity, with dehmited borders

(présent, however, in Cyclodorippoidea and in

Dakoticaticroidea), but not evolved in the most

primitive families; presence ol a press-button,

which is typical and remains practically unchan-

gcd in ail eubrachyuran crabs.

The novelty with a sternal différentiation has

appeared independentiy several times within the

Brachyura: five times in the Podotremata, and

only once in rhe I leterorremata-Thoracotremata

assemblage. The question of a common origin for

both Lyreidinae and Eubrachyura nceds further

considération.

Within the Brachyura a socket has appeared three

tintes: in Homoloidea, Lyreidinae and Eubra¬

chyura. Again, the question of a common origin

for both Lyreidinae and Eubrachyura is highligh-

ted. The condition in Eubrachyura, with a typi¬

cal press-button on sternite 5, is nor regarded as

having evolved from a primitive stage as in dyno¬

menids or homoloids for example.

The ABDOMINAL RETAINING MECHANISM USF.D

Aï PALEONTOLOGICAL DATA

The nature of fossil ntaterial does not preclude

observation of the abdominal retaining/locking

Systems. Wlten the coxae of pereiopods and ame-

rior sternal plate are preserved. it could bc easy to

see if any process overhangs the telson and how

the uropods (if présent) may be involved. It can

be deduced wlieibet wc are dealing with a podo-

trematous crab ot not, and also to décidé wbether

it is a homolodromiid instead of a homolid.

ZOOSYSTEMA • 1998 • 20 (4)

681

Guinot D. & Bouchard J.-M.

Projections arising from stérilité 5 are présent in

several raninid fosiils, for example in the

Lyreidinae ( l.yreidus succedunus, L. rosenkntntzi ,

L. bispinulittus, and Rogais orrt), but information

is lacking on structures thaï miglit allow locking

with sockcts on pleomere 6, such as the hooks in

the extant Lyreidus and Lysirude. A careful com

parison with représentatives ol the exclusivclv fos-

sil subfamily Palacocorystinac, for example

Notopocorystes stokesii (Mantell, 1844), is neccssa-

• 7 -

In the case of an apparatus ol the press-button

type, the rwo parts are normally not visible when

the abdomen is tolded. But, in puhlishcd illustra¬

tions, apparent sternal buttons can bc clearly

observed when the abdomen is lost or displaccd.

In Ophthalmoplax stepbensoni Rathbun, 1935

(pl. 13, ftg. 14), indicated as a female, two

tubercles on sternite 5 seem to correspond to the

buttons.

Male specimens of Xanthopsis dufouni (H. Milne

Edwards, 1850) and Harpactncarcinus souverbiei

A. Milne Edwards 1862, Lnbocarcinus paidinn-

wurtembergen$is von Meyer, 1847, ail three from

the Lutetian, chat were examined, hâve visible

sternal promipences (Bouchard 1996, pl. 6,

fig. 3a-c).

Fossil représentatives of the family

Retroplumidae, for example Costaplurna concava ,

C. salarnanca, and C. nordestirui which hâve their

ventral surface preserved, show marked latéral

expansions on pleomere 6 and a notch on sterni¬

te 4, thus a mechanism similar to that of extant

species and probably efficient indeed.

Fossil leucosiids clearly exhibir the spécial fea-

tures of their abdomen, fused with the sternal

parts (Rathbun 1926, pl. 1 1). Morris & Collins

(1991, figs 17b, 18, 24b) hâve shown Neogene

leucosiids in which the spur on sternite 5 is clear¬

ly visible in a le tiens nt species, but absent in two

Iphicnhis species.

The spécial structures shown by fossil brachyuran

crabs deserve to be extensively studied.

HyPOTHI- SIS ABOU T THE HOMOl.OGY UROI’OD-

SOCKf.T

For Hessler (1983: 157, fig. 6) évolution of the

eumalacostracan (caridoid) abdomen from that

of a phyllocarid through fusion of the seventh

original abdominal segment to the sixth occurred

in concert with the évolution of the tail fan. He

suggested that the sixth pleopods have evolved

into uropods bccausc they were the most poste-

rior appendages. Uropods combined with telson

to form the tail fan are présent throughout the

Decapoda [for other crustacean groups see

Schram (1974) and Wilson (1996). Exceptions

are the Lithodidae (uropods lost in males and

females) and the Lomoidea (uropods reduced in

fcmalcs, lost in males) (Balss 1940; McLaughlin

1983a, b; Grutier 1993); ail of thèse bave essen-

tially a flcxed abdomen. Therc is obviously a

functional sigtiiflcancc to loss of uropods.

It is generally said that the uropods are lost in ail

Brachyura, cxcepr in the Dromiacea where they

remain as vestiges. In cstablishing the new taxon

Meiura (= reduced tail) to include the two mono-

phyletic groups Anomura and Brachyura, Scholz

& Richrer ( 1995) have not stressed the important

différence regarding the biramous uropods, pré¬

sent in (almost) ail Anomura and always absent

in Brachyura. Thus the charactcr furnished by

uropods in the mosr primitive crabs takes on

great importance, and their rôle in abdominal

rétention deserves considération. We attempt to

demonstrate hcre that the socket hoilowed on the

sixth abdominal segment in hr.icli)'uran crabs is

dircctlv derived from the uropod.

Ferez (1928b: 649; 1929; I 145) has noted the

rcsetnblance between pagurids, which grasp and

hold on to the shells columella with their develo-

ped uropods, and dromiids which liold their

abdomen with their vestigial dorsal uropods. He

pointedout that the condition of Dromia may be

a stage in évolution ol the higher brachyuran

condition. He concluded that in higher crabs the

socket which allows lastcniug is perhaps "le der¬

nier vestige de l’uropode évanoui ’ (last vestige of

the disappearing uropod). Harrnoll (1975: 16),

Guinot (1979a: 156) and Guinot & Richer de

Forges (1997: 472) adopted the idea of a homo-

logy between the uropod and the socket. Even if

the socket may be regarded as homologous with

the uropod, we do not think that it is directly

evolved from the dromiid dorsal uropod.

This homolngy is supported liy several argu¬

ments:

— Ail the Podotremata which have vestigial uro-

682

ZOOSYSTEMA • 1998 - 20(4)

Abdominal holding Systems (Brachyura)

Table 3. — Uropod patterns in Brachyura (in comparison to the

biramous uropods in other Decapoda).

Taxon

Uropod pattern

PODOTREMATA

Homolodromiidae

ventral lobes

Dynomenidae

dorsal plates

Dromiidae

dorsal plates

Exceptions:

- Hypoconcha

ventral lobes

- Exodromidia

ventral lobes

- Pseudodromia

ventral lobes

- Ascidlophilus

absent

Homolidae

sockets

Latreilliidae

sockets

Poupiniidae

sockets

Cynomomdae

absent

Cyclodorippidae

absent

Phyllotymolinidae

absent

Raninidae

absent

Exception :

Lyreidinae

sockets

HETEROTREMATA (*)

sockets

Exceptions:

Corystidae

absent

Orlthylldae

absent

Cryptochiridae

absent

Leucosiidae

absent

THORACOTREMATA

sockets

Exceptions:

Grapsidae Sesarminae ( pro parte)

absent

Grapsidae Varuninae ( pro parte)

absent

Gecarcinidae ( pro parte)

absent

Ocypodidae (pro parte)

absent

Mictyrldae

absent

* Note that certain généra in the family Hymenosoma-

tidae hâve dorsal platelets ventrally hollowed in deep

sockets.

pods (dorsal plates or ventral lobes) are devoid of

sockets (Dromiidac, Dynomcnidae, Homolodro-

miidae), Le. the Dromiacca.

- Ail the Homoloidcà, vvhich havc no trace of

appendagelike or dorsal or lobiform uropods,

possess sockets (honiolid press-but ton), as well in

the primitive family Poupiniidae as in the most

advanced onc, the Latreilliidae.

- Ail (or nearly ail) brachyuran crabs that always

lack uropods possèss sockets. Arnong the

57 familics revicwed here the press-button cha-

racterizes 46 families.

- Examples of brachyuran crabs devoid of both

uropods and sockets are very rare: the Raninidae

(Lyreidinae excepted) and the Cyclodorippoidea

(Table 3).

- Dorsal uropods, cven when utiiized in dromiids

for abdominal rétention, never beat a Socket ven¬

trally: they operate as a full-lock System and not

as a press-button.

As previously said, it is accepted that the dorsal

plates seen in the great majority of dromiids and

in ail dynomenids represent modifîed uropods.

The transformation uropod-soeket has necessari-

Iv rcquired a substantial modification.

Ventral lobes ol homolodromiids (and some dro-

miitis) (see Table 3) obviously appear as rudi¬

ments of the appendages of pleomere 6, without

a modification as important as tn the case of dor¬

sal uropods. The lobes resemble rudimentary

pleopods found on preceding pleomeres in ail

Homolodromia and most of the Dicranodromia

species (Guinot 1995: 179, figs 5, 25), but they

differ by their insertion and orientation.

Uropodal lobes, cither with two articles or weak-

ly bifîd, svere observed in certain homolodromiid

species ( ibid .). An argument to support that these

ventral lobes are vestiges of the si.Nth pair of

appendages is shown by their configuration in

the Dynomenidac. In dynomenids. which hâve

large dorsal uropodal plates (visible ventrally and

dorsally, and larget than in dromiids), vestigial

pleopods are alvvays présent on segments 3-5 in

males. One could expecr that, similarly, pleo¬

mere 6 might beat a ventral pair c)f pleopods 6: in

fact, this site is occupied by rhe triangular uro¬

pod, the ventral surface of which is not hollowed

(Figs 6B-D, 7C, D). It is our considered opinion

that rhe dromioid dorsal uropods are specialized

structures.

When. in the Brachyura, a Socket was présent on

the sixth segment, no vestigial appendage has ever

been observed on the third rhrough fifth abdo¬

minal segments.

Case uf the Hymenosornatidae

Within the Heterotremata, the Flymenosomati-

dae are particularly interesting because in the

nrost primitive members of the family, the soc¬

kets are situated on intercalary platelets. What is

more interesring is that rhe platelets may be

movable, for example in the généra Odiomaris Ng

ZOOSYSTEMA • 1998 -20(4)

683

Guinot D. & Bouchard J.-M.

et Richer de Forges, 1996, A mari) ut s Lucas, 1980,

and Hymeuosoma Dcsmarest, 1825, pro parte

(Guinot & Richer de Forges 1997).

Internai surface of rhe platelet is hollowed as a

deep socket, externally bordered by a thickened

calcificd margin and separated from the pleotel-

son by a membrane (Bouchard 1996). These

structures are clearly visible lor e.\3mple in

OrJiomaris piles us (A. Milne Fdwards, 1873) (bet-

ter known as F la mena pilota or Amarimts pilosns)

(Fig. 27). In establishing Atmtrinus angelicus ,

Holthuis (1968: 115) described “a xmall movable

segment attached to the base of the telson and the

postero-lateral angle of the sixth somite” and

Fig. 27 — Locking apparatus in Hymenosomatidae, for example in Odiomaris pilosus (A. Milne Edwards), 6 20 x 22.6 mm, New

Caledonia (MNHN-B 26146); A, B, abdomen in dorsal and ventral views; C, dorsal view of pleotelson; D, side view of the mobile inter-

calated platelet, witb socket (C, D, scanning électron microscope photographs). Abbreviations: a5, abdominal segment 5; m, mem-

branous zone; p, intercalated platelet; pt, pleotelson; so, socket. Scale bars: C, 200 pm; D, 100 pm.

ZOOSYSTEMA • 1998 -20(4)

Abdominal holding Systems (Brachyura)

remarked: "Such intcrcalatcd plates ta my know¬

ledge hâve xo l.it not been reported Irom an y bra-

chyuran other than Dromiiclae. Whether these

structures in the présent specics and in the next

[= O. pilosus] are homologous with thosc of the

Dromiids, I am unable to say."

In fact, the Hymenosomatidae show several pat¬

terns: platelets movable: platelets simple and no

longer arriculared, wirh more or less distinct

sutures (plesiomorphic condition); différentia¬

tions dorsalIv visible at the pleotelson base (for

example rhe trilobate pleotelson in the genus

Hymenicoides Kemp, 1917); prominences more

or less marked. When the platelets disappear dor-

sally, the sockets are situated at the sanie place,

wihout a visible cxtcrnal indication (apomorphic

condition). Ail these pièces, either platelets or

sockets, play the sanie rôle in covering the acute

buttons of the press-button apparatus and firmly

attaching the abdomen.

We will examine whether the morphological crite-

ria provided hère are sufficient for idenrifying uro-

pods and sockets as homologues (Hentlig 1966;

Wiley 1981; Wiigele 1996). The similarity of

position (topographie and position in relation to

other parts) is obvions. The socket is a more or less

deep dépression on abdominal somitc 6, delimi-

ted by a catcified margin; the most rhickcned part

of this margin is always latcro-posterior, allovving

an efficient locking with the button. In

Eubrachyura (Amarinus and Odinmdris excepted)

the socket is integrated into pleomere 6 and pré¬

sents varions forms: from rounded (Fig. 26B) to

more or less elongated (Fig. 26D), In dorsal view,

the movable hymenosornatid platelet signiftcantly

rccalls a dorsal uropod, but its internai surface is

hollowed as a deep socket. Thus, the hymenoso-

matid platelet ventrally looks like (and functions

as) a socket; dotsally, it looks like a dorsal plate.

In researching the criterion of spécial similarity

between uropods and sockets. one rteeds to inves-

tigate similarities of structure, for exaniple in his-

tological detail, and ontogenetic resemblance.

The criterion ol phylogenetic position may rein¬

force homologues. Ventral lobes of homolodro-

miids might probably' be considered as tratisitio-

nal passage between thedccapod uropods and the

sockets. In dromiids the uropods are often used

to retain the abdomen. Ventral lobes are présent

in sonie primitive dromiids and do not play any

rôle. The movable and artieulated dorsal uropods

of certain dromiid forms are apparemly more pri¬

mitive thaï the lused ones. The large and always

complctelv dorsal plates in dynomenids, that are

more advanced, appear as derived. We remark

that in rhe brachyuran evolutionary scheme,

where uropods are lost, sockets are tmiformly

developed in rhe same location, on abdominal

somite 6. As already remarked, rhe trench toward

loss of abdominal limbs are components of the

carcinization process in the Decapoda. The

noyelty represented by the socket may be regar-

ded as the eharaaer State présence of uropod, but

we ignore the process of différentiation from the

uropod into the socket.

The Brachyura which possess lobe-shaped ventral

uropods or platt-shaped dorsal uropods, are reco-

gnized as being more primitive (dromiids, dyno¬

menids) than thosc having sockets at the same

location (Homoloidea, I.yredinae, and F.ubra-

chyura).

The criterion of position is provided by a strict

and constant location of ventral lobes, dorsal

plates, platelets, and sockets on pleomere 6. F.ven

in the cases vvhere the sockets appear atypically

positioncd, they bclong to the same pleomere.

For example, when sockets are tound on the last

element ol abdomen (Hymenosomatidae Fig. 27,

lnachoidinae Fig. 20C-F. and in a part ol the

lnachinae Fig. 20A, B), this is easily explained by

rhe fusion as pleotelson of segment 6 with relson.

The criterion of continuance through intermé¬

diare forms may be provided by presence of

movable dorsal uropods in dromiids and movable

plaLelets in hymenosomatids. Dromiid uropods,

that are never used when applicd on a surface but

act in full-lock, laek ventral sockets, vvhile the

hymenosornatid platelets are ventrally hollowed

and operare as a press-button. Absence of sockets

in advanced Brachyura [pro parte Grapsidae,

Gecarcinidae, Ocypodidae) is considered to be

the resuit of a secondary loss.

In conclusion the appendages of the sixth abdo¬

minal segment are practically always présent in

Decapoda, but with different patterns (biraincms

liinb, rasp, lobe, plate, platelet, socket). Hence it

is probably not accurate to défi ne the Brachyura

by the absence of uropods.

ZOOSYSTEMA • 1998 -20(4)

685

Guinot D. & Bouchard J.-M.

Acknowledgements

Only at une of the major muséums, Iike the

Muséum national d'Ilistoire naturelle, Paris, a

large référencé collection, essenrial to such a

review ofall brachyuran groups, can be found.

We wish to express spécial gratitude to

S. Secretan (Muséum national d'Histoire naturel-

le/CNRvS, Laboratoire de Paléontologie) for

paleontological data and loan of fussil crabs,

M. Tavares (Universidade Santa Ursula, Rio de

Janeiro) for many (tours ol arguments and dis¬

cussion on the Cyclodorippoidea and Podotre-

mata in general, and R, B. Manning (National

Muséum of Natural History; Smithsonian

Institution) (or his encouragement. We are grate-

ful to R. K. Kropp (Battelle Océan Sciences),

P. K. L Ng (National Universiry of Singapctre)

and M. Türkay (Forschtmgsinstitut Senckenberg,

Frankfurt) for prpviding useful information.

We acknowledge: R. Cleva for providing excel¬

lent assistance with the scanning électron micro¬

scope; Mrs. M. van der Merwe (South African

Muséum, Cape Town) and P. Clark (The Natural

History Muséum, London) for the loan ol rare

dromiid.s; M. Judson (Muséum national d’His-

toire naturelle) for his kind help and advice;

C. Chardin for his support.

We are greatly indebred to P. Castro (California

State Polytechnic Universiry) who had carefully

and critically react rhrough an earlier draft of this

manuscript and improved it with constructive

comment*; and to L. B. Holrhuis (Nationaal

Natuurhistorisches Muséum, Leiden), R. B.

Manning and, particularly, C. L. McLay

(Universiry' of Canterbury, New Zealand) for

their valuable comment* and criricisms.

Photographs were taken over a period of years by r

Jacques Rebière. Original drawings are due, also

over a period of years, to Michèle Bertoncini. We

express our profond thanks for their invaluable

contribution, which make this work sound and

comprehensive.

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Submitted on 2 April 1998;

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694

ZOOSYSTEMA • 1998 - 20 (4)

The types of Recent and certain fossil

opisthobranch molluscs in the Muséum

national d’Histoire naturelle, Paris

Ângel VALDÉS & Virginie HÉROS

Laboratoire de Biologie des Invertébrés Marins et Malacologie,

Muséum national d’Histoire naturelle,

55 rue de Buffon, F-75231 Paris cedex 05 (France)

malaco@mnhn.fr

avaldes@casmail.calacademy.org

Valdés À. & Héros V. 1998. — The types of Recent and certain fossil opisthobranch mol¬

luscs in the Muséum national d'Histoire naturelle. Paris. Zoosystema 20 (4): 695-742.

KEY WORD.S

type specimens,

opisthobtanchs,

Mollusca,

MNHN, Paris.

ABSTRACT

Three hundred and fifty seven lots of Recent and certain fossil opisthobran¬

ch mollusc type-specimens deposited in the Laboratoire des Invertébrés

Marins et Malacogie of the Muséum national d’Histoire naturelle (MNHN)

are catalogued by original binomen and arranged alphabetically within fami-

lies. Most of the fossil type specimens are housed in the Laboratoire de

Paléontologie, and therefore are not included in this catalogue. The essential

bibliographical, geographical and taxonomie information is provided for each

taxon.

MOTS CLÉS

types,

opisthobninenes,

Mollusca,

MNHN, Paris.

RÉSUMÉ

Les types actuels et de quelques fossiles de mollusques opisthobranebes du Muséum

national d’Histoire naturelle , Paris. Trois cent cinquante-sept types d’opistho-

branches actuels et quelques fossiles déposés au Laboratoire de Biologie des

Invertébrés Marins et Malacologie du Muséum national d'Histoire naturelle

(MNHN) ont été identifiés et listés par ordre alphabétique d’espèces à l’inté¬

rieur de chaque famille. La plupart des types fossiles se trouvent au

Laboratoire de Paléontologie et ne sont pas mentionnés dans cette liste. Les

principales références bibliographiques, géographiques et taxonomiques

accompagnent chaque porte-nom.

ZOOSYSTEMA • 1998 • 20 (4)

695

Valdés Â. & Héros V.

INTRODUCTION

In recent years, opisthobranch systematists hâve

become increasingly interested in examining type

material as part ot their research. However, the

scarcity of updated tnuseum opisthobranch type

catalogues make this task arduous and rime

consuming. Most muséum opisthobranch type

collections consisr ot liistorically important spéci¬

mens for vvhiclt the identification ol type spcci-

mens, and the détermination ol their status, is

usually dilficult to résolve vvith certainty. Often,

old type specimens rernain unrecognized in the

general collections, which may bouse types of

species not expected to be présent at that

muséum.

The objective of this paper is to document the

présent collection and scope ol the Recent and

certain lossil opisthobranch name-bearing type

specimens in the Muséum national d Histoire

naturelle of Paris (MNHN). This will lacilitate

future systematic research based on this material.

A BRI EF HISTÔRY OF 1 HE COLLECTIONS

The Muséum national d’Histoire naturelle in

Paris (M.NHN) was founded in 1793, and short-

ly thereafter, the First opisthobranch type speci¬

mens werc deposited in ils collections. The ear-

liest major collections extant are those of die

French expéditions to exotic countries, carried

out during the eariy nineteenth century

{YAstrolabe, Y Uranie, Savigny expédition to

Egypt, d'Orbigny expédition to South America,

Péron expéditions to the Indo-Pacific), described

by the French classic authors: G. C.uvier,

J. C. Quoy & J. P. Gaimard, J. V. Audouin, A. de

Férussac, P. S. Rang, A. d’Orbigny, L. F. A.

Souleyet.

In the second half oF die nineteenth century, and

the beginning ol the twentieth century, the

French carried out a signifieant collecting effort

in their colonies in the Caribhean, Africa, Eastern

Asia and South-Wcst Pacific. This material,

added ro MNHN collections, was also mainly

described by French authors (G. Deshayes,

L. Morlet, A. T. de Rochebrunc, P. Fischer), At

about the same rime, L. Morlet and H, Crosse

described several opisthobranch fossil species

whose types are now in MNHN, and new océa¬

nographie expéditions (Cap Horn, Travailleur et

Talisman, Diguei expédition to Baja California,

French Antarctic expéditions. Gravier Expédition

to Djibouti) werc an additional source of type

specimens, moxtly described by A. I.ocard,

P. Fischer, J. B. M. Vayssicrc and J. Risbcc.

During the First and Second World Wars, unlike

several natural history muséums in Europe, the

MNHN collections remaincd untouched In

thcsc and in the Following years, many type spe-

cimens were added as a resuit i>f the work of

A. Pruvot-Fol who described material collected

abroad and From the French coasrs. 1 lowever the

types of many species that she described could

ne ver be found, possibly bccause the specimens

were dtscarded aFter dissection; the extant mate-

rial is mostly that of formai expéditions or

muséum holdings that were entrusted to her for

description. Material Front the Calypso expédi¬

tions was also described by Ev. Marcus. In the late

1950s, the Fischer Family shared «lie property of

Journal de Conchyliologie , E, Pischer-Piette, then

prolessor and director of the Laboratoire de

Malacologie, donared the type collection of the

Journal , conraining a number of opisthobranch

types. At that rime, the Muséum wer collections

Were stored in high glass containers, at the

Galerie de Zoologie, the main exhibit building.

When this gallery was closed to the public in

1965 the collections remaincd tbere, and many

specimens wetit dry, or werc otherwi.se damaged.

A new âge in the MNHN opisthobranch collec¬

tion, charactcrixed by the interiiationalization of

the scope of the type collection, started vvith the

arrivai of P. Bouchet on the staff in 1975. I le not

only added his own types, but also was instru¬

mental m encouraging authors from other coun¬

tries to deposit their type specimens in MNIIN.

Many signifieant types of Eumpean and African

species were conrributed through the Spanish

school (J. A. Ortea, J, C. Garda-Gémicz, and

orhers), and Indo-Pacific material collected by

French scietitists were described by W. B.

Rudman (some ol them srill on exrended loan).

In 1985, the MNHN scientific collections vvere

moved to a new, large underground storage buil¬

ding, and the type specimens of Recent and cer¬

tain fossil molluscs were separated and placed in

696

ZOOSYSTEMA • 1998 • 20 (4)

Opisthobranch mollusc type specimens in MNHN

a repository in thc Laboratoire de Biologie des

Invertébrés Marins et Malacologie. However, the

opisthobranch collection xtill reniai ned in need

of curation for several years. Lite older type spe-

cimens were stored Logethcr vvitb other tnaterial

in antiquated fashion, and no longer suitable

containers, and it was difficult to Pind spécifie

lots. In fact, there are .some examples in rbe recenr

literaturc of type specimens eonsidered lost

(W.ïgele 1985, 1990) that hâve recently appea-

red. For this reason, the First author of this paper

was kindly invited in 1995 to work at MNHN as

an assistant curator, to reorganize the opistho¬

branch collection, and separate the type speci¬

mens. This task, carried eut in collaboration with

the second author, was concluded ditring another

short-term stay in 1997, with the lilial séparation

and inventory of ail type specimens. During this

work, we hâve also found several types of species

described by authors whose type matcrial was not

expected to be in MNHN. Some of them are

replacement naines, names introduced under the

Article 70c (ICZN 1985), or specimens appa-

rently borrowed by A. Vayssière front other insti¬

tutions, never returned and later deposited at

MNHN.

Type Catai .oc.ue

The catalogue of Recent and certain fossil types

of opisthobranch molluscs in MNHN covers

three hundred and fifty-three lots of name-bea-

ring type specimens. However, it does not inclu-

de most ol the fossil species in MNFIN which are

stored in the Laboratoire de Paléontologie (e.g.

the Cossmann collection).

This catalogue is restricted to uame-bearing

types: holotypes, Ieuotypes, neotypes and syn-

types. Other type lots (paratypes and paralecto-

types) are in (or will be translerred to) the gene¬

ral collection, and are only included in the cata¬

logue as accessory matcrial of the name-bearing

types. Ail type specimens are listed by species-

group names and attanged alphabefically within

families. To facilite the search, the complété list of

names, arranged alphabetically, indicating the

family, higher category taxa and page number

where eacb naine bas been placed is included.

When the same lot is the type of two different

species-group names, they are listed separatejy,

unless if rhev are identically spelled. The essermal

taxonomie, geographicàl and bibliographical

information is provided for each species-gioup

name. These data include the name of the species

cited exactly as puhlished in the original descrip¬

tion, the author, date and bibliographie référencé,

the type locality as originally printed and its

modem geographïcal équivalent, thc category

and number of specimens, State of conservation,

accessory tnaterial (e.g, photographs, micro,scopy

slides, spawn or food of the specimens, paratypes

and pâralectotypes), and the name of the collec¬

ter when it is known. The old géographie names,

no longer used, are printed berween quotation

marks. The names of large géographie areas,

countrïes or archipelagos are translated into

Lnglish, but other names are printed in their

local spelling. The State of conservation includes

two main items: “specimen ”, which refers tu spe¬

cimens conserved in alcohol complète ol soft

parts, and “shell’, when only a shell remain in the

dry collection. When the type specimen held is

also known to be originally or subsequently figu-

red it is indicated. Ail shells (with the exception

ol four nominal species catalogucd while the

paper was already in press) are illustrated, even if

the)' hâve been prcviously ligured. Figured speci¬

mens are individually identificd in the collection.

Short remarks with the currently accepted taxo¬

nomie status of the species, lectoiype désigna¬

tions, status as type species of genus-group

names, or any other useful information are given.

No attempt lias been made to list the synonyms

of each species, but opinions on the taxonomie

status of several names hâve been included (only

when they arc bascd in the examination of the

type matcrial or are derived from an exhaustive

révision). Ail names bave been grouped in fami¬

lles on the basis of available literature. When it

was not possible we bave been obliged to adopt

oui own view, which obviously requires confir¬

mation by further xtudies. This is not a cricical

review of the material, but merely is an attempt

to make its usage casier.

The families arc enumerated in the order of the

classification followed by Rudtnan & Willan

(1998), with nonicnclatural adjustments

(Bouchet & Rocroi, pers. comm.).

ZOOSYSTEMA • 1998 -20(4)

697

Valdés Â. & Héros V.

CEPHALASPIDEA

Family AcTEONIDAE d’Orhigny, 1842

azoricus. Actaeon azorieus Locard, 1897: 85-87,

pl. 3, Figs 8-11. Type locality: Talisman (1883)

Expédition, stn 126, 38°37’N - 28°21‘W, bet-

ween Pieo and S. Jorge, Atones, 1258 m.

HOLOTYPE (by monotypy, figured by

Locard, 1897, pl. 3, Figs 8-11); shell (Fig, IA).

Remarks: I bis is the type species oF elle genus

Inopimdon Boucher, 1975, by original dési¬

gnation.

maltzani. Actaeon maltzani Dautzenberg, 1910:

10, pl. 1, figs 1, 2. Type locality: Baie de

Rufisque and Gorce, Sénégal. SYNTYPE

(Figured by Dautzenberg, 1910: pl, 1, figs 1,

2): 1 shell (Fig. IB). Remarks: Dautzenberg

(1910) placcd Tornatella senega/ensis Petit de la

Saussaye, 1851 in the genus Acteon de

Montfort, 1810 (citcd as Actaeon) and therefo-

re Actaeon senegalemis Maltzan, 1885 beeomes a

junior secondary honionym of A. senega/etisis

(Petit de la Saussaye, 1851). Thcrcfore,

Dautzenberg (1910) introdueed the replace¬

ment naine Actaeon maltzani for it. This new

species was based on Maltzan's material and

newly collected specimens studied by Dautzen¬

berg. Ail of them are syntypes of this species.

morelletorum. Actaeon morelletorum Gougerot et

Braillon, 1968: 200, pl, I, Fig. 3. Type locality:

Barisseuse, France (middle Eocene). HOLO-

TYPE (by original désignation, Figured by

Gougerot & Braillon pl. 1, fig. 3).

vagabunda. Tornatella vagabunda Mabille, 1885:

208. Type locality: Magallanes, Chile. SYN¬

TYPE: 1 shell (Fig. IC).

Family APLUSTRIDAK Gray, 1847

[= HydATTNIDAE Pilsbry, 1895]

eximia. Huila exmu a Deshayes, 1863: 55, 56,

pl. 7, figs 23, 24. Type locality: La Réunion

(Indian Océan). SYNTYPES: 3 shells (Fig. 1D).

guamensis. Bullaea guamrmis Quoy et Gaimard,

1824; 423-425, pl. 66, figs 10-12. Type locali¬

ty: Agana, Guam (Pacific Océan). SYNTYPE:

1 specimen, lcg. Quoy and Gaimard.

Remarks: Pruvot-Fol (1934b) placed this spe¬

cies in the genus Micromelo Pilsbry, 1894.

Family RlNGICULIDAK Philippi, 1853

iidmirabilis. Ringicula admirabilis Morlet, 1883:

203, 204, pl. 9, fig. 3. Type locality: Mediter-

rancan Sea. SYNTYPE: 1 shell, leg. Morlet

(Fig. IE). Remaries: This is the type species of

the genus Plicatm Nordsieck, 1972. by original

désignation. According to Ciccone Savona

(l l )82), this species is a junior synonytn of

Ringicula confortais Monterosato, 1877.

bourgeois i. Ringicula bourgeoisi Morlet, 1878b:

261,262, pl, 8, fig. 5. Type locality: Pontlevoy,

Paulniy, Ferrières l'Arçon, Manthelan (France,

middle Miocene), Mandillot (France, early

Miocène). SYNTYPES: 3 shells, leg. Crosse

(Fig. IF).

bourgnignati. Ringicula bourguignati de

Rochebrune, 1883: 178, Type locality:

Casamance, Sénégal. SYNTYPES: 2 shells

(Fig. IG)

calédonien. Ringicula calédonien Morlet, 1880:

154, I 55, pl, 5, fig. 1. Type locality: "Plage de

la Baie Pouen" (= Baie de Pritzbuer), New

Caledonia. SYNTYT’ES 14 shells, leg. Morlet

(Fig. JH),

crosses. Ringierula crosse/' Morlet, 1878b: 273,

274, pl. 7, fig. I I. Type locality; various locali-

ries in central and western Europe (middle and

upper Miocene). SYNTYPE: I shell, leg.

Crosse (Fig. 11),

gaudryana. Ringicula gaudryana Morlet, 1878b:

283, 284, pl. 7, fig. 12. Type locality: various

localities in central and western Europe (mid¬

dle Miocene to Pliocène). SYNTYPE: 1 shell,

leg. Crosse (Fig. 1 J).

maries. Ringicula maria Morlet, 1880: 152, 153.

Type locality: “Nossibé" G Nosy Be), Mada¬

gascar. SYNTYPES: 9 shells, leg. de Folin

(Fig. 2A).

minutula. Ringicula minauda Locard, 1897: 89,

90, pl. 14, figs 7-9. Type locality: Talisman

(1883) Expédition, stn 84, 22°54'N -

17°26’W, off Sahara, 860 m. SYNTYPE:

I shell (Fig. 2B).

noumeensis. Ringicula nourneensis Morlet, 1880:

155, 156, pl. 5, fig. 3. Type locality: Nouméa,

New Caledonia. SYNTYPES: 18 shells, leg.

Morlet (Fig. 2C).

oehlertiae. Ringicula oehlertiae Morlet, 1880:

698

ZOOSYSTEMA • 1998 - 20(4)

Opisthobranch mollusc type specimens in MNHN

Fig. 1. — A. holotype of Actaeon azoricus, 11.3 mm; B. syntype of Actaeon maltzani. 3.1 mm; C, syntype of Tornatella vagabunda,

8.2 mm; D, syntype of Bulla eximia, 10.2 mm; E, syntype of Ringicula admirabilis, 3.6 mm; F, syntype of Ringicula bourgeois!, 3.9 mm;

G, syntype of Ringicula bourguignali, 4.0 mm; H, syntype of Ringicula caledonica. 1.9 mm; I, syntype of Ringicula crossei, 3.7 mm;

J, syntype of Ringicula gaudryana, 12 mm.

ZOOSYSTEMA • 1998 • 20 (4)

699

Valdés Â. & Héros V.

156, 157, pl. 5, fig. 4. Type locality: Seas of

China and Japan. SYNTYPES: 3 shells, leg.

Morlet and Saint-John (Fig. 2D).

passieri. Ringicula passieri Morlet, 1880: 157,

158, pl. 5, fig. 5. Type locality: Cap Breton

Canyon, Bay of Biscay. SYNTYPF: 1 shell, leg.

de Folin (Fig. 2F.).

paulucciae. Ringicula paulucciae Morlet, 1878b:

266, 267, pl. 6, fig. 6, pl. 8, fig. 9. Type locali¬

ty: Saucats and Dax, France (earlv Miocene).

SYNTYPF,: I shell, leg. Crosse (Fig. 2F).

pirulina. Ringiculapirulina Locard, 1897: 87, 88,

pl. 14, figs 1-6. lype locality: Talisman (1883)

Expédition, srn 33, 32°3TN - 09 Ü 48'W, off

Atlantic coast of Morocco, 1350 m. SYN-

TYPF..S: 2 shells (Fig. 2G). Remarks:

According to Bouchet (1975b), this is a junior

synonym of Ringicula nitida Verrill, 1874.

pontelevicnsis. Ringicula pontelevieusis Morlet,

1878b: 274, 275, pl. 8, fig. 8. Type locality:

varions localities in France (rniddle Miocene).

SYNTYPF: 1 shell, leg. Crosse (Fig. 2H).

ptilchella. Ringicula pulchella Morlet [ ex Jeifreys

MS], 1880: 158, pl. 5, fig. 6. Type locality:

Atlantic European. SYNTYPES: 4 shells

(Fig. 21). Rcmarks: This is the type species of

the subgenus Ringaciaeon Nordsicck ( 1972) by

original désignation.

quadriplicata. Ringicula quadriplicata Morlet,

1878b: 286, 287, pl- 7 , fig. 1 . Type locality:

several localities in Europe (rniddle and upper

Miocene, and lower Pliocène). SYNTYPE:

I shell, leg. Crosse (Fig. 3A).

rosildae. Ringicula rosildae Rosso et Saubade,

1985: 305, 306, figs 8-11. Type locality:

14°36’30"N - 17°15'30”W, off Sénégal.

HOLOTYPE: said to bc deposited in MNHN

in the original publication, but never presenred

to the muséum.

roussellae. Ringicula roussellae Rosso et Saubade,

1985: 303, 304, fig. 2, Type locality:

14°22’40”N - 17°09’W, off Sénégal. HOLO-

TYTF: said to be deposited in MNHN in the

original publication, but never presented to the

muséum.

salleana. Ringicula salleana Morlet, 1880: 153,

154 Type locality: Cap Breton Canyon, Bay of

Biscay, SYNTYPE (probably): 1 shell, leg. de

Folin (Fig. 3B).

savignyi. Ringicula savignyi Morlet. 1878a: 117,

118, pl. 5, fig. F Type locality: Sue/., Fgypt

(Red Sea). SYNTYPES (one figured by

Bouchet & Danngal 1982, fig. 59): 10 shells,

leg. Savigny (Fig. 3C).

scblumbergeri. Ringicula schlumbcrgeri Morlet,

1883: 204-206, pl. 9, fig. 4. Type locality:

Mediterranean Sea. SYNTYPES: 3 shells, leg.

Morlet (Fig. 3D). Remarks: According to

Ciccone & Savon a (1982), this is a junior

synonym of Ringicula confondis Monrerosato,

1877-

senegalensis. Ringicula senegalensis Morlet, 1883:

202, 203, pl. 9, fig. 2. Type locality: Sénégal.

SYNTYPES: 2 shells, leg.'de Folin (Fig. 3£).

terquemi. Ringicula terquemi Morlet, 1880: 159,

160, pl. 5, fig. 7- Type locality: "Baie de

Sniyme" ( = Bay of Jzmir), Turkey. SYN¬

TYPES: 5 shells, leg. Terquem (Fig. 3F).

Remarks: According to Ciccone & Savona

(1982), Hais is a junior synonym ol Ringicula

aariculata (.Ménard de la Groye, 1811).

touruoueri. Ringicula toumoucri Morlet, 1878b:

287, 288, pl. 6, fig. 10. Type locality: vartous

localities in central and western Europe

(Neogene). SYNTYPE: 1 shell* leg. Crosse

(Fig. 3G).

Family CYUCHNIDAE H. et A. Adams. 1854

[= Tornatinidaf. P. Fischer, 1883]

abyssicola. Mamillocylicbna abyssicola Bouchet,

1975b: 349, 350, fig. 14. Type locality:

Biaçores Expédition, stn 251,

47"38’N - 08°56’W, Banc de la Chapelle,

3360-3600 m. HOLOTYPE (by original dési¬

gnation): shell (Fig. 4F).

crossei. Cytkbna crossei Bucquoy, Dautzenberg et

Dollfits, 1886: 526, 527, pl. 64, figs 9-11.

lype locality: not specified; cited front Canet,

France. Viareggio, Italy and Alger. SYNTYPES

(One figured by Bucquoy, Dautzenberg &

Dollfus 1886: 526: pl. 64, fig. 9): 2 shells front

_ Alger.

imperceptus. Meloscapbander imperceptus

Bouchet, 1975b: 341-343, figs 9, 10, pl. 3,

figs G-H. Type locality: Talisman (1883)

Expédition, off Mauritania. HOLOTYPE (by

original désignation, figured by Bouchet,

700

ZOOSYSTEMA • 1998 -20(4)

Fig. 2. — A. syntype of Ringicula mariei, 1.7 mm; B, syntype ot Ringicula minutula, 2.9 mm; C, syntype of Ringicula noumeensis,

3.7 mm; D. syntype of Ringicula oehlertiae, 3.9 mm; E. syntype of Ringicula passieri, 4.2 mm; F, syntype of Ringicula paulucciae,

3.4 mm; G. syntype of Ringicula pirulina, 6.8 mm; H. syntype of Ringicula ponteleviensis, 3.2 mm; I, syntype of Ringicula pulchella,

2.7 mm.

ZOOSYSTEMA • 1998 -20(4)

701

Valdés Â. & Héros V.

1975b, pl. 3. fig. H): shell (Fig. 4G) + 1 para-

type.

insperata. Bulla insperdta. P. Fischer in Locard

1897: 54, 55, pl. 1, ligs 19-22. Type locality:

Talisman ( 1883) Expédition, srn 76, 25°01’N -

16°55’\V, off Sahara, 2638 m. SYNTYPES:

2 specimens (1 of rhem dissecred). Remarks:

According to Bouchet (1975b), this is a junior

subjective synonym oi .S 'capbander mundus

Watson, 1886.

lemchei. Cylichna lemehei Bouchet et Warcn,

1979: 228, 229, figs 16, 18G, H. J-L. Type

locality: Norbi Expédition, stn CP02

(64°26’-64°19’N, 01°36'-01°44'E) Norwegian

Sea. HOLOTYPE (by original désignation,

figured by Bouchet 8e Warén 1979, là g. 18J-E):

shell (Fig. 4M).

millepunctata. Bulla millepunctata Locard,

1897: 52-54, pl. 2, ligs 3-6. Type locality: not

specified; cited Irom Talisman (1883)

Expédition, several stations, ofl Sahara,

Sénégal and Azurés, SYNTYPES: 1 specimen

and 2 shells (Fig. 41). Remarks: This is the

type species of the subgenus Bullocardia

Nordsieck, 1972, by original désignation.

According ro Bouchet ( 1975b), this is a junior

subjective synonym of Scaphander nobüis

Verrill, 1884.

mirabilis. Tornatina mirabilis Locard, 1897: 72,

73, pl 2, figs 20-24. Type locality; Travailleur

(1882) Expédition, stn 23, 38°2TN -

09°27’W, off Portugal, 2000 m. SYNTYPE:

1 shell (Fig. 5A).

morelletorum. Cylichna morelletorum Gougerot

et Le Renard, 1983: 82, 85, fig 1 2. Type loca¬

lity: Ferme de l’Orme, France (middle

Eocene). SYNTYPES: 24 shells.

olivaeformis. Tornatina olivaeformis Issel, 1869:

171, 172. Type locality: Golf of Suez, Egypt

(Red Sea). SYNTYPE (figured by Savigny

1817, pl. 6, fig. 25 and Bouchet & Danrigal

1982, fig. 56): 1 shell (Fig. 5E).

petiti. Cylirhmipetits Dautzenberg, 1923: 70, 71,

fig. rexr. Type locality; “Fênérive” (= Fenoarivo

Atsinanana), Madagascar. HOLOTYPE (by

monotypy, figured by Dautzenberg, 1923, fig

text); shell (Fig. 5B).

pusillina. Tornatina pustllina Locard, 1897:

75-77, pl- 2, ligs 29, 30. Type locality:

Travailleur (1880) Expédition, drag. 2,

43“46'N - 01°55’W, Cantabrian Sea, North of

Spahi, 1019 m. SYNTYPE: 1 shell (Fig. 5F).

scaphandr aides. Roxdnid scaphandroides Staadt

in Cossmann &c Pissarro 1913, pl. 55,

fig. 242-12. Type locality: Chenay, France (late

Paleocene). H01,0 TYPE (by monotypy, figu¬

red by Cossmann & Pissarro 1913, pl. 55,

fig. 242-12): shell (Fig. 5C).

villersii. Bulla villersii Audouin. 1826: 39. Type

locality: Egypt. SYNTYPES (one figured by

Savigny, 1817. pl. 5, fig. 4 and Bouchet &

Danrigal 1982, fig, 53): 2 shells (Fig. 5D).

voluta. Bulla voluta Quoy et Gaimard. 1833:

359, 360, pl. 26, figs 33-35. Type locality:

Guani (Pacific Océan). SYNTYPE: 1 shell, leg.

Quoy and Gaimard (Fig. 5G). Remarks:

Pruvot-Fol (1934b) placed this species in the

genus Acteocind J. E. Gray, 1847.

Family Rll'USIDAl Thiele, 1925

canariensis. Cyliclmina canariensis Nordsieck et

Garcta-Talavera, 1979: 178, pl. 44, fig. 46.

Type localitv: lenerife, Canary lslands.

SYNTYPE: 1 shell (Fig. 5H).

candiduUt. Cylichna candidttla Locard, 1892: 28.

Type locality; Atlantic, toast of France. SYN-

TYPE: 1 shell (Fig. 51).

desgenettii. Bulla desgcncttii Audouin, 1826: 39.

Type locality: Egypt. SYNTYPES (one figured

by Savigny 1817. pl. 5, fig. 6 and Bouchet &

Danrigal 1982, fig. 55); 5 shells (Fig. 5J).

dilatata. Retusa dilatata Pallary, 1904: 215, 216,

pl. 7, fig. 8. Type locality: Gulf of Gabès,

Tunisia. SYNTYPES: 11 shells (Fig. 5K).

Remarks: Pallary (1904) selected a 6 mm long

shell as the “type” of this species. However, in

MNHN collection there are rvvo vials with 4

and 7 shells respectively, ail of rhem similar in

length (about 6 mm), vvhich rherefore are

considered syntypes.

fourierii, Bulla fourierii Audouin, 1826: 39.

Type locality: Egypt. HOLOT YPE (by mono¬

typy, figured by Savigny, 1817, pl. 5, fig. 5 and

Bouchet & Danrigal 1982, fig. 54): shell

(Fig. 5L).

girardi. Bu/la girardi Audouin, 1826: .39. Type

locality: Egypt. HOLOTYPE (by monotypy,

702

ZOOSYSTEMA • 1998 - 20(4)

Opisthobranch mollusc type specimens in MNHN

Fig. 3. — A, syntype of Ringicula quadriplicata, 8.9 mm; B, probable syntype of Ringicula salleana, 5.0 mm; C, syntype of Ringicula

savignyi , 2.8 mm, D. syntype ot Ringicula schlumbergeri, 3.1 mm; E. syntype of Ringicula senegalensis, 2.3 mm; F, syntype of

Ringicula terquemi, 1.8 mm; G, syntype of Ringicula tournouefi, 4.3 mm; H, syntype oi Butta australis de Ferussac, 41.6 mm;

I, syntype of Bulla australis Quoy et Gaimard, 49.4 mm.

ZOOSYSTEMA • 1998 - 20(4) 703

Valdés Â. & Héros V.

Fig. 4. — A, holotype of Bulla mongii, 1.4 mm; B, syntype of Bulla orbignyana, 16.1 mm; C. syntype of Bulla ovoidea, 13.6 mm;

D, syntype of Haminea perrieri, 16.4 mm; E. holotype of Cylichnium waldae, 28.1 mm; F, holotype of Mamillocylichna abyssicola,

2.8 mm; G, holotype of Meloscaphander imperœptus, 19.3 mm; H, holotype of Cylichna lemchei, 7.1 mm; I, syntype of Bulla mille-

punctata, 41.0 mm.

704 ZOOSYSTEMA • 1998 • 20 (4)

Opisthobranch mollusc type specimens in MNHN

Fig. 5. — A, syntype of Tornatina mirabilis , 2.6 mm; B. holotype of Cylichna petiti, 6.5 mm; C, holotype of Roxania scaphandroides,

12.9 mm; D. syntype o( Bulla villersii, 1.3 mm; E, syntype of Tornatina olivaetormis. 3.3 mm; F, syntype of Tornatina pusillina, 2.0 mm;

G, syntype of Bulla voluta, 9.7 mm, H, syntype of Cylicbnina canariensis, 2.6 mm; I, syntype of Cylichna candidula , 4.2 mm;

J, syntype of Bulla desgenettii, 1.7 mm; K, syntype of Refusa dilatata, 5.7 mm; L, holotype of Bulla fourierii, 2.5 mm.

ZOOSYSTEMA • 1998 • 20 (4)

705

Valdés Â. & Héros V.

figured by Savigny 1817. pl. 5, fig. 3 and

Bouchet & Danrigal 1982, fig. 52): shell

(Fig. 6A). Remarks: This is the t)<pe species of

the genus Bullina Risso in Audouin 1826, by

monotypy.

simplex . Bulla simplex Locard, 1897: 55-57, pl- 2,

figs 7-9. Type loeaüty: 7 alisrnan (1883) Expédi¬

tion, stn 75, 25°01’N - 16*53'W, off Sahara,

2325-2518 m. SYNTYPES: 8 shells (Fig. 6B).

Remarks: Bouchet ( 1975h) transfetred this spe-

cies to the genus Rcliçbna Rudm.m, 1971.

tenerifensis. Cylichninti tenerijênsis Nordsieck et

Garcta-Talavera, 1979: 177. 178, pl. 44,

fig. 45. Type locaiity; Tcnerife, Canary Islands.

SYNTYPE: 1 shell (Fig. 6C). Remarks: In the

caption ot the Hg. 45 the name of this species

is spelled CyUchnina tenerijjàe.

truncatelLi. Cylichna truncjttella Locard, 1886:

533-534. Type locaiity: Cannes, France. SYN-

TYPES: 3 sJiells (Fig. 6D). Remarks:

According to Lemche (1948), this is a junior

subjective synonym ot Refusa truncatula

(Bruguière, 1792).

Family PhilinidaeJ. E. Gray, 1850

azorica. Philine azorica Bouchet, 1975b: 353,

354, fig. 17, pl. 4, figs F, 1. Type locaiity:

Biaçores Expédition, stn 167, 37°46’N -

25°48'W, off Azurés, Sâo Miguel, 140 m.

HOLOTYPE (by original désignation, figured

by Bouchet 1975b, fig. 17, pl. 4, figs F, I): spé¬

cimen (dried) + .shell (Fig. 6E) and dried frag¬

ments.

milneedwardsi. Philine milneedwardsi Locard,

1897: 35-37, pl. 1, figs 7-9. Type locaiity:

Talisman (1883) Expédition, stn 107, “Rade de

Porte Grande" (= Porto Grande), Sâo Vicente,

Cape Verde Islands, 75-90 m. SYNTYPES:

2 shells (Fig. 6F).

motiilifera. Philine manilifera Bouchet, 1975b:

354-356, fig. 18, pl. 4, figs D, E. Type locaii¬

ty: Biaçores Expédition, stn 41,

37°43’N - 29 D 04’W, oft Azores, Princesse Alice

Bank, 450-475 m. HOLOTYPE (by original

désignation)' shell (Fig. 6G) + dried fragments.

Family AGI.AJIDAF Pilsbry, 1895

hirundinina. Bulla hirundinina Quoy et

Gaimard, 1833: 367-369, pl. 26, figs 20-25.

Type locaiity: “Ilots aux Cerfs”, “île de France”

(= Mauritius), lndian Océan. SYNTYPES:

4 specimens (1 ol them dissected), leg. Quoy

and Gaimard. Remarks: This is the type spe¬

cies o( the genus Chdidonura A. Adams, 1850,

by monotypy.

quinza. Aglaja quinza Ev. Marcus, 1979: 132,

133, figs 1-4. Type locaiity: Calypso

Expédition, stn 115, 23°56’S - 44°17'W, bet-

ween Rio de Janeiro and Santos, Brazil, 23 m.

HOLOTYPE (by monotypy, figured by Ev.

Marcus 1979, figs 1-4): specimen (dissec-

red) t shell and male organ slides.

seurati. Doridium seurati Vayssière, 1926:

125-128. pl. 13. Type locaiity: “près de la

Skhirra" (= near Sakhira), Gulf of Gabès,

Tunisia. SYNT YPES: 11 specimens. Remarks:

Rudman (1972a) placed this species in the

genus Melanochlamys Cheeseman, 1881.

Family Hamineidae Pilsbry, 1925

africana. /lieras afrieana P. Fischer in Locard

1897: 62, 63. pl 2, figs 15-19. Type locaiity:

nor specified;: cited front Talisman (1883)

Expédition, stn 96, 19°19’N - 18 n 02'W, off

Mauritania, 2320-2333 m, and stn 101,

16°38’N - 18°24'W, Cape Verde Islands,

3200 m SYNTYPE: I shell (Fig. 7E) front

stn 96. Remarks: Bouchet (1975b) placed this

species in the genus Cytichnium Dali, 1908.

avachis, Bulla avachis Quoy et Gaimard, 1833:

361, pl. 26, figs 28-30. Type locaiity: ‘Port du

Roi-Georges” (= King Georges Sound),

Western Australia. SYNTYPE: 1 shell, leg.

Quoy and Gaimard (Fig. 7F). Remarks:

Pruvot-Fol (1934b) placed this species in the

genus Haminea Leach, 1820.

brevis. Bulla brevis Quoy et Gaimard, 1833: 358,

359, pl. 26, figs 36, 37. Type locaiity: “Port du

Roi-Georges” (= King Georges Sound),

Western Australia. SYNTYPES: 10 shells, leg.

Quoy and Gaimard (Fig. 7G). Remarks:

Pruvot-Fol (1934b) placed this species in the

genus Hrtminea Leach, 1820.

cymbalum. Bulla cymbalum Quoy et Gaimard,

1833: 362, pl. 26, figs - 26, 27. Type locaiity:

Baie d'Humata (= Umatac), Guarn (Pacific

706

ZOOSYSTEMA ■ 1998 -20(4)

Opisthobranch mollusc type specimens in MNHN

Fig. 6. — A, holotype of Bulla girardi, 2.4 mm: B, syntype of Bulla simplex, 10.7 mm: C, syntype of Cylichnina tenerifensis, 1.3 mm;

D, syntype of Cylichna Iruncatella, 2.5 mm; E. holotype of Philine azorica, 7.5 mm: F. syntype of Philine milneedwardsi, 23.1 mm;

G, holotype of Philine monilifera, 8.8 mm; H, lectotype of Creseis acicula. 12.6 mm: I, lectotype of Hyalaea aflinis, 11.6 mm;

J, syntype of Cuvieria astesana, 6.2 mm; K, holotype of Cleodora chaptalii Souleyet and Clio chaptaliiJ. E. Gray, 12.8 mm.

ZOOSYSTEMA • 1998 -20(4)

707

Valdés Â. & Héros V.

Océan). SYNTYPE: 1 shell (broken), leg.

Quoy and Gaimard (Fig. 7H). Remaries: This

is the type spccies of the genus Lamprohami-

noea Kuroda et Habe in I labe 1952, by origi¬

nal désignation.

fischeri. Acrnn fischeri I ncard, 1897:63-65, pl. 2,

figs 10-14. Type locality: Travailleur (1880)

Expédition, stn 10 , 43°39’N - 03 n 28’W, Bayof

Biscaye, I960 m. SYNTYPES; 2 shells (1 bro¬

ken) (Fig. 71). Remarks: In actingas firsr revi-

sor in the meaning of the Article 24 (ICZN

1985), Bouchet (1975b) determined that

Acents afrmina P. Fischer in Locard 1897 (pla-

ced in the genus Cylicbnium Dali, 1908) lias

precedence over its subjective synonym /lieras

fischeri Locard, 1897.

gallica. Bulla gallica Locard, 1905: 22, 23. Type

localitv: Atlantic and Mediterranean coast of

France. SYNTYPES: 4 .shells (Fig. 7J).

mongti. Bulla niougii Andouin, 1826: 39. Type

locality: Egypt. HOLOTYPI. (by monotypy,

figured by Savigny 1817, pl. 5, fig. 7 and

Bouchet & Danrigal 1982, fig. 58): shell

(Fig. 4A).

orbignyanu. Bulla orbignyana de Ferussac, 1822:

573. Type localitv: La Rochelle, France. SYN¬

TYPES: 4 shells (Fig. 4B).

ovoidea. Bulla ovoidea Quoy et Gaimard, 1833:

348, 349, pl. 26, figs 17-19. Type locality:

“Plage d’Humata" (= Umatac), Guam (Pacific

Océan). SYNTYPE: 1 shell. leg. Quoy and

Gaimard (Fig. 4C), Remarks: Pruvot-Fol

(1934b) placed this species in the genus

Haminea Leach, 1820.

perrieri. Haminea perrieri Morlet, 1889: 178,

179, pl. 6, fig. 7. Type localitv: “Golfe de

Siam" (= Gulf of Thailand).' SYNTYPE:

1 shell, leg. Pavie (Fig. 4D).

tvaldae. Cylichnium waldâe Bouchet, 1975b:

344-347, figs 11, 12, pl, 4, figs B, G. Type

locality: U ''aida Expédition, stn CY15.

22°53 S - 11°56’E. Sourheastern Atlantic,

1756 m. HO LO'l YPE (by original désigna¬

tion, figured by Bouchet 1975b, pl. 4, fig. B):

shell (Fig. 4E).

Family SMARAGDINELLIDAE Thiele, 1925

glauca. Bulla glauca Quoy et Gaimard, 1833:

352, 353, pl. 26, figs 10-12. Type locality:

“Havre Carteret", New Ircland, Bismarck

Archipelago. HOLOTYPE (by monotypy,

figured by Quoy Si Gaimard 1833, pl. 26,

figs 10-12): spécimen, leg. Quoy and Gaimard.

Remarks: In acting as firsr levisor in rhe mea¬

ning of the Article 24 (ICZN 1985), Pruvot-

Fol (1934b) determined rhaL Bulla tandis Rang

in Quoy Si Gaimard 1833 (placed in the genus

Smnragdinella A. Adams, 1848) has precedence

over tts sujective synonym Bulla glauca Quoy

et Gaimard, 1833.

lutea. Bulla lutea Quoy et Gaimard, 1833: 369,

370, pl. 26, figs 40-44. Type locality: “Port

Dorey (- Manolcwari), Irian Jaya, Tndonesia.

SYNTYPES: 4 specimens, leg. Quoy and

Gaimard.

viridis. Bulla viridis Rang in Quoy & Gaimard

1833. 350-352, pl. 26, figs 13-16. Type locali¬

ty: “Rade d’Humata" (= Umatac), Guam

(Pacific Occan). SYNTYPES: 22 specimens (6

of them dissccted) and 3 shells, leg. Quoy and

Gaimard. Rcmarks: 1 his is the type species of

the genus Striaragdinellt A. Adams, 1848, by

monotypy. According to Rudman (1972b),

this is a junior synonym of Smaragdine/la caly-

culata (Broderip et Sowcrby, 1829).

Family Bui.LIDAK J. E. Gray, 1827

australis. Bulla austmlis de Ferussac, 1822: 573.

Type locality: Port Jackson, Sydney, Australia.

SYNTYPE: 1 shell (Fig. 3H).

australis. Bulld australis Quoy & Gaimard 1833:

357. 358, pl. 26, ligs 38, 39. Type locality:

“port du Roi-Georges" ( = King Georges

Sound), Western Australia. SYNTYPES (one

figured by Quoy 6e Gaimard 1833, pl. 26,

figs 38-39): 11 shells, leg. Quoy and Gaimard

(Fig. 31). Remarks: This naine is preoccupied

by Bulla australis de Ferussac, 1822.

delorti. Bulla delorti Mabille, 1896: 116. Type

locality: Jap.m. SYNTYPES: 4 shells (Fig. 7A).

mabillei. Bulla mabillei Locard, 1897: 50, 51,

pl. 2, figs L 2. Type locality: Talisman (1883)

Expédition, stn 107 (no coordinates are speci-

fied), “Rade de Porte Grande’ (= Porto

Grande), San Vicente, Cape Verde Islands,

75-90 m. SYNTYPES (1 figured by Locard

708

ZOOSYSTEMA • 1998 -20(4)

Opisthobranch mollusc type specimens in MNHN

Fig. 7. — A, syntype of Butta delorti, 35.2 mm: B, syntype of Butta mabillei, 60.1 mm; C, syntype of Butta modesta, 8.1 mm;

D. syntype of Butta secunda, 33.2 mm; E, syntype of Aœras atncana, 15.8 mm; F. syntype of Butta arachis, 17.7 mm; G. syntype of

Butta brevis, 12.6 mm; H, syntype of Butta cymbalum, 26.6 mm: I, syntype of Aœras fischeri, 7.0 mm. J, syntype of Butta gallica,

19.6 mm.

ZOOSYSTEMA • 1998 - 20(4)

709

Valdés Â. & Héros V.

1897, pl. 26, figs 38, 39): 2 shells (Fig. 7B).

modesta. Bu IL/ modem Risso, 1826: 49, pl. 1,

Fig, 7. lypc locality: Nice, France, SYNTYPES:

3 shells (1 broken) (Fig. 70.

secunda. Bulla seconda Mabille, 1896: 112. Type

local ity: Japan. SYNTYPES: 6 shells (Fig. 7D).

Family RUNCINIDAE H. «A. Adams, 1854

coronata. Pelta coronata de Quatrefages, 1844:

151, 152, pl. 3, fîg- 6, pl. 5, fig. 7, pl. 6, figs 3,

6, 9, 15. Type locality: Ile de Bréhat, Bretagne,

France. SYNTYPES: 4 specimens, leg.

Quatrefages. Reinarks: T'his is the type species

of the genus Pelta Quatrefages, 1844, by origi¬

nal désignation. The genus Pelta has been sup-

pressed under plcnary powers bv Opinion 811

(ICZN 1967).

lentieula. Runehut lenticula Gofas, Ortea et

Rodriguez, 1991: 541-545, figs 1-6. Type loca¬

lity: Chapcu Armado, Namibe, Angola.

HOLOl’YPE (by original désignation): spéci¬

men + 10 paratypes.

macrodenticulata. Runcina macrodenticulata

Garda, Garcia-Gômez et Lôpez, 1990: 4-6,

fig. 1. Type locality: near Playa Benitez, Ceuta,

Moroccan side of Strait of Gibraltar. HOLO-

TYPE (by original désignation): specimen.

Fig. 8. — A, B, syntype of Prasina borbonica, 4.3 mm: C. syntype of Berthelinia elegans, 0.6 mm: D, holotype of Lobiger souverbii,

6.2 mm: E, syntype of Umbrella cumingi, 39.2 mm.

710

ZOOSYSTEMA • 1998 -20 (4)

Opisthobranch mollusc type specimens in MNHN

SACOGLOSSA

Family OXYNOEIDAE Stoliczka, 1868

sieboldii. Lophocercus sieboldii Krohn, 1847:

55-59, pl. 2, figs 5-9, 11. Type locality:

Messina, Italy. SYNTYPES: 2 speciniens (1 of

them lacking shell). Remarks: This is the type

species of rhe genus" Lophocercus Krohn, 1847,

by monotypy. Aher Môrch (1863b),

Lophocercus is currently considérée! a synonym

of Oxynoe Rafinesque, 1819.

souverbii. Lobiger souverbii F. Fischer, 1857: 273,

274, pl. Il, figs 7-10. Type localicy:

Guadeloupe (Caribbean Sea). 11ÜLOTYPF.

(by monotypy, figured by P. Fischer 1857,

pl.' Il, figs 7. 8): shell (Fig' 8D).

Family JULllDAE E. A. Smith, 1885

borbonica. Prasina borbonica Deshayes, 1863:

29, pl. 4, figs 4-8. Type locality: La Réunion

(Indian Océan). SYNTYPES: 2 shells

(2 valves), 1 shell (1 valve), leg. Petit de la

Saussaye (Fig. 8A, B). Remarks: This is the

type species of the genus Prasina Deshayes,

1863, by monotypy, Kay (1968) placed P. bor¬

bonica in the genus Julio Gould, 1862, and

suggested that it is probably a synonym of

J. exquhita Gould, 1862.

elegans. Benhelinia eleguns Crosse, 1875: 79, 80,

pl. 2, fig. 3. Type locality: lower calcareous,

Lutetian Stage (middle tocene), Courtagnon,

France. SYNTYPES (2 figured by Crosse,

1875, pl. 2, fig. 3): 7 shells (1 valve, considera-

bly damaged), leg. Bcrihclin (Fig. 8C).

Remarks: This is the type species of the genus

Berthelinia Crosse, 1875, by monotypy.

Family PLACOBRANCHIDAE J. E. Gray, 1840

schrammi. Tridachiti schrarnmi Môrch, 1863a:

41. Type locality: Guadeloupe (Caribbean

Sea). SYNTYPES: 4 specimens. Remarks:

Deshayes (1857) established the new genus

Tridachia Deshayes, 1857, but did not rame a

species. Môrch (1863a) introduced for the first

rime the binominal name Tridachia schrammi

in référencé to Deshayes’ description.

T. schrammi , currently considered a synonym

of Elysia crispata (Môrch, 1863), is the type

species of the genus Tridachia Deshayes, 1857,

by subséquent monotypy. Following Gosliner

(1 995), Tridachia is a junior .synonym of Elysia

Risso, 1818.

Family HeRMAEIDAF. H. et A. Adams, 1854

lierai. Stiliger lierai Ortea, 1982a: 188-191,

figs 8, 9. pl. L fig. B. Type locality: El Prix,

Tenerife, Canary Islands. HOLOTYPE (by

original désignation): specimen + spawn +

photo.

lozattoi. Ercolania lozanoi Ortea. 1982a:

194-196, fig. 13. Type locality: Palm-Mar,

Tenerite, Canary Islands. HOLOTYPE (by

original désignation, figured by Orrea 1982a,

fig. 13): specimen.

verticilata. Placida verticilata Ortea, 1982a:

191-194, figs 10, 1 l, 12b. Type locality: Punta

Hidalgo, Tenerife, Canary Islands.

HOl.OTYPE (by original désignation): spéci¬

men.

APLYSIOMORPHA [= ANASPIDEA]

Family Akeridae Mazzarelli, 1891

bicittcta. Bulla bicincta Quoy et Gaimard, 1833:

355, 356, pl. 26, figs 31, .32. Type locality:

“Porr du Roi-Georges” (= King Georges

Sound), Western Australia. SYNTYPES:

2 shells glued to cardboard, severeiy bru ken,

leg. Quoy and Gaimard.

elegans. Acera elegans Locard, 1886: 535. Type

locality: Quiberon, France. .SYNTYPES:

4 shells (3 broken) (Fig. 91). Remarks:

According ro Lemche (1948), this is a junior

subjective synonym of Akera bullata (Muller,

1776).

spiraui. Accra spirata Staadr in Cossmann &

Pissarro 19)3, pl. 55, fig. 244-2.Type locality:

Chenay, France (late Paleocene). HOLOTYPE

(by monotypy, figured by Cossmann &

Pissarro 1913, pl. 55, fig. 244-2): shell

(Fig. 9J).

Family Api.YSIIDAE Lamarck, 1809

alba. Aplysiella gravieri var. alba Vayssière,

ZOOSYSTEMA • 1998 -20(4)

711

Valdés Â. & Héros V.

1906b: 67, 68, pi. 4, figs 66-68. Type locality:

Obock, Djibouti. HOLOTYPE (by monoty-

py, b gu red by Vayssière 1906b, pl. 4,

figs 66-68): specimen (dissected) + sotne parts

prepared for SEM. Remarks' According to

Martinez (1996), rhis subspecific-rank taxon is

a junior synonym of Petalifern petalifern (Rang,

1828).

ascifera. Aplysia ascifera Rang, 1828b: 51, 52.

pl. 4, figs 7-9. Type locality: Saint Jean,

Cayenne, French Guîana. SYNTYPES: 2 .spé¬

cimens, leg. Richard. Remarks: According to

Engel & Humnielinck (1936), this is a junior

synonym of Dolabri/era dolabrifera (Rang,

1828).

brasiliana. Aplysia brasiliana Rang, 1828b: 55,

56, pl. 8, figs 1-3. Type localiry: Brazil.

SYNTYPES: 3 spécimens (2 of client dissec¬

ted), leg. Quoy and Gaimurd.

cirrbifera. Aplysia eirrhiféra Quoy et Gaimard,

1832: 311, 312, pl. 24, fig. 8. Type locality':

“îles aux Cerfs”, “île de France'’ (= Mauritius).

Indian Océan. SYNTYPF.S: 2 specimens, leg.

Quoy and Gaimard. Remarks: This is rhe type

species of the subgenus Barnardaclesia Cales et

Engel. 1935, by original désignation.

dactylo mêla. Aplysia dactylo mêla Rang, 1828b:

56, pl. 9. Type locality: “Saint-Yago de la

Praya” (= Sâo Tiago), Sâo Tiago Island, Praia,

Cape Verde Islands. SYNTYPE: 1 specimen,

leg. Rang. Remaries: 1 bis is the type species of

the subgenus Vania Paies, 1960. by original

désignation.

ecaudata. Aplysia ccaudata Rang, 1828b: 47.

pl. 2. Type localiry: “Waigiou” (= Waigeo) and

"Rawack” (= Garni, Irian Jaya, Indonesia.

SYN FYPES: 3 specimens (1 of them dissect¬

ed), leg. Quoy and Gaimard. Remarks: This is

probably a junior synonym of Dolabella auri-

cularia (Lightfoot, 1786).

gravieri. Aplysiella gravieri Vayssière, 1906b:

58-66, pl. 4, figs 56-65. Type localiry: Gulf of

Aden, Djibouti. HOLOTYPE (by monotypy,

figured by Vayssière 1906b, pl. 4, figs 56-65

and Martinez 1996, fig. 10): specimen (dissec¬

ted) + sotne parts préparer! for SEM. Remarks:

According to Martinez (1996), this is a junior

synonym of Petalifern petalifern (Rang, 1828).

inca. Aplysia inca d’Orbigny, 1835: 207-209

(1837), pl. 19, figs 1-3 (1835). Type locality:

between Callao and Isla de San Lorenzo, and

Puerto de Lima, Peni. SYNTYPE: I specimen

(dissected), leg. d’Orbigny.

juliana. Aplysia ju/iana Quoy et Gaimard, 1832:

.309, 310, pl 24, figs 5, 6. Type localiry: “île

de France" (= Mauritius), Indian Océan.

SYNTYPES: 1 specimen (dissected) and

I fragment of shell, leg. Quoy and Gaimard.

Remarks: Aplysia juliana has bcen placed in

the Official List of Spécifie Nantes in Zoology,

Opinion 1844 (ICZN 1996). I bis is the type

species of tbe subgenus lu lia Pruvot-Fol,

1934, by monotypy.

keraudrenii. Aplysia keraudrenii Rang, 1828b:

59, 60, pl. 13. Type locality: unknown.

HOLOTYPE (by monotypy, figured by Rang,

1828b, pl. 13): specimen (dissected), leg.

Lesson.

lessonü. Aplysia lesstmii Rang, 1828b: 60, pl. 14.

Type locality: Paita, Perd. SYN TYPE: I speci¬

men (dissected and considerahly dantitged),

leg. l.esson and Garnot. Remarks: According

ro F.ales (1960), this could be a junior syno¬

nym of Aplysia keraudrenii Rang, 1828.

longicautla. Aplysia longicauda Quoy et Gaimard,

1824: 421, 422, pl. 66, fig. 8. Type locality:

not specified; cited front “île de France’’

(= Mauritius), Indian Océan; "îles Sandwich”

(= Hawaii), “Baie des Chiens-Marins” (= Shark

Bay), Australia; Rio de Janeiro, Brazil. SYN¬

TYPE: 1 specimen.

maillardû Dolabrifera maillardi Deshaves, 1863:

53, 54, pl. 7, figs 20-22. Type localiry: La

Réunion (Indian Océan). SY'NTYPES:

3 shclls. Rcmarks: According ro Engel &

Elummelinck (1936), this is a junior Synonym

of Dolabrifera dolabrifera (Cuvier, 1817).

nudata. Aplysia nudata Rang, 1828b: 72, pl. 22,

figs 3-5. Type localiry: on floaring algae near

Hawaii. SYNTYPE: l specimen (dissected),

leg. Quoy and Gaimard.

oabottensis. Aplysia otihouensis Souleyet, 1852:

461, 462, pl. 25, 10-13. Type locality: Oahu,

“îles Sandwich” (= Hawaii). SYNTYPE: 1 spe¬

cimen, leg. Eydoux and Soule) T et. Remarks:

According ro Engel &. HummeJinck (1936),

this could bc a junior synonym of Dolabrifera

dolabrifera (Rang, 1828).

712

ZOOSYSTEMA • 1998 -20(4)

Opischobranch mollusc type specimens in MNHN

Fig. 9. — A, lectotype of Creseis clava, 10.0 mm; B. iectotype of Hyalaea gibbosa, 7.2 mm; C. syntype of Hyalaea globulosa and lec-

totype of Cavohna globulosa, 6.2 mm; D, syntype of Hyalaea levigata. 2.0 mm; E. lectotype of Hyalaea limbata, 7.9 mm; F, syntype

of Hyalea rangii. 8.3 mm, G. syntype of Creseis striata. 5.7 mm; H. lectotype of Hyalea uncinata, 7.6 mm: I, syntype of Acera ele-

gans, 21.6 mm; J, holotype of Acera spirata, 17.2 mm.

ZOOSYSTEMA ■ 1998 -20(4)

713

Valdés Â. & Héros V.

petalifera. Aplysia petalifera Rang, 1828b: 52,

pl. 5, figs 1-3. Type locality: Nice, France.

SYNTYPES: 2 specimens (dissected), leg.

Risso. Remarks: This is the Lype species of che

genus Petalifera ). F'. Gray, 1847, by absolute

tauconymy.

pleei. Aplysia pleei Rang, 1828b: 70, pl. 21- Type

localiry: Antilles. SYNTYPE: I specimen (dis¬

sected), leg. Plée. Remaries: According to

Eales & Engel (1935), this naine musc bc used

for a sübspecies of Bursatella leachii de

Blainville, 1817 occuring throughout the

Caribbean Sea.

protea . Aplysia protea Rang, 1828b: 56, 57,

pl. 10, figs 1-3. Type locality: Antilles.

SYNTYPES: 3 specimens (1 of them dissec¬

ted), leg. Richard; 2 specimens (dissected), leg.

Plée. Remarks: According to Paies (1960), this

is a junior synonym of Aplysia dactylornela

Rang, 1828. ’

rufa. Aplysia rufa Quoy et Gaimard, 1832: 314,

pl. 24, fig. 7. Type locality: “rade d’Umata”

(= Bay of IJmatac), Guam (Pacific Océan).

SYNTYPE: 1 specimen (dissected), leg. Quoy

and Gaimard.

striata. Aplysia striata Quoy et Gaimard, 1832:

315, 316, pl. 24, figs 9-11. lype locality: “Port

Dorey” (- Manokwari), Irian Jaya, Indonesia.

SYNTYPE: 1 specimen, leg. Quoy and

Gaimard. Remarks: In the original description

of this species, Quoy & Gaimard (1832) noted

that it is probably the same as Stylocbeilus lon-

gicauda (Quoy et Gaimard, 1824).

teremidi. Aplysia temnidi Rang, 1828b: 48, pl. 3,

figs 1-3. Type locality: not specified; cited ffom

Tahiti and Bora Bora, Society Islands (Pacific

Océan); Oualan, Caroline Islands (Pacific

Océan). SYNTYPE: 1 specimen, leg. l.esson

and Gamot. Remarks: This is prohably a

junior synonym of Dolabella auricularia

(Lightfoot, 1786).

tongana. Aplysia tongana Quoy et Gaimard,

1832: 305, 306, pl. 23, figs 6, 7. Type locality:

“Ile de Pangaï-Modou”, Tongatapu (Pacific

Océan). SYNTYPE: ! specimen (dissected),

leg. Quoy and Gaimard. Remarks: This is pro¬

bably a junior synonym of Dolabella auricula¬

ria (Lightfoot, 1786).

unguifera. Aplysia unguifera Rang, 1828b: 52,

pl. 5, figs 4-7. Type locality: Mediterranean

Sea. SYN TYPES: 2 specimens (1 of them dis¬

sected), leg. Risso. Remarks: According to

Engel & Hummclinck (1936), this is a junior

synonym of Petalifera petalifera (Rang, 1828).

NOTAS PI DEA

Family Umbraculidak Dali, 1889

cumingi. Umbrella cumingi Deshayes, 1863: 52,

53, pl. 8, figs 4, 5. Type locality: La Réunion

(lndian Océan). SYNTYPES: 3 shells

(Fig. 8E).

Family Pl.F.UROBRANCHlDAF. j. E. Gray, 1827

amboinei. Oscaniopsis ambatnei Vayssière, 1900:

9, 10. Type locality: Arnbon, Moluccas,

Indonesia. SYNTYPES: 2 specimens, one of

them dissected. Remarks: According to Ev.

Marcus & Gosliner (1984), this is a junior

synonym aïEuselenops lunieeps (Cuvier, 1817).

brocki. Berthella brocki Vayssière, 1897a:

120-122, pl. 5, figs 8-10. lype locality: not

specified; cited frotn Atnbon, Moluccas,

Indonesia; Edam, near Jakarta, Indonesia;

Jervis Bay, Austtalia. SYNTYPES: 8 specimens

(2 of them dissected),

eapensis. Plmrobrancbaea capemis Vayssière,

1900: 10, 11. lype locality: Cape of Good

Hope, Southern Africa. SYNTYPE: 1 speci¬

men (considcrably damaged), leg. Raynaud.

Remarks: According to Ev. Marcus &

Gosliner (1984), this is a junior synonym of

Pleurobmtichca tarda Verrill, 1880.

citrinus. Pleurobrancbus citrinus Rüppell et

Leuckart, 1828: 20, pl. 5, figs 1A-C. Type

locality: Suez, F.gypt. SYNTYPES: 2 specimens

(1 of diem dissected), leg. Rüppell. Remarks:

This species is currently placed in the genus

Bertbellina Gardiner, 1936.

crossei. Pleurobrancbus crossei Vayssière, 1897b:

353, 354, fig. 1. Type locaiity: Caribbean Sea.

HOLOTYPE (by monorypy, figured by

Vayssière 1897b, fig. 1): specimen (dissected),

leg. Plée.

digueti. Pleurobrancbus digueti de Rochebrune,

1895: 240. Type locality: Mogote, Bahta de La

714

ZOOSYSTEMA • 1998 - 20(4)

Opischobranch mollusc type specimens in MNHN

Paz, Mexico. SYNTYPES: 4 specimens, leg.

Diguet.

giardi. Pleurobrartchus giardi Vayssière, 1897b:

354-356, fig. 2. 'Iype locality: Camiguin,

Luzon, Philippines. SYNTYPE: 1 specimen

(dissected), leg. Semper.

maculatum. Pleurobruncbidium macidatum

Quoy et Gaimard, 1832: 301, 302, pl. 22,

figs 11-14. Type locality: Southern Australia,

SYNTYPES: 7 specimens (4 of them dissec¬

ted), leg. Quoy and Gaimard Remarks: Ev.

Marcus fit Gosliner (1984) placed this species

in the genus Pleurobranchaea Meckel in Leue

1813.

niant ilia tus. Pleurobranchus rnanùllatus Quoy et

Gaimard, 1832; 294-296, pl. 22, figs 1-6. Type

locality: Port Louis, “île de France”

(= Mauritius), Indian Océan. SYNTYPES:

2 specimens (dissected).

peronii. Pleurobranchus peronii Cuvier, 1804b:

275, 276, pl. 18, figs 1-6. Type locality: “Mer

des Indes" (= Indian Océan). SYNTYPE:

1 specimen. leg. Peron. Remarks: This is the

type species of the genus Pleurobranchus

Cuvier, 1804, by monotypy.

perrieri. Pleurobranchus perrieri Vayssière,

1897a: 126-128, pl. 4, figs 2-4. Iype locality:

not specified; cited front Philippines; Ambon,

Moluccas, Indonesia; Tahiti. SYNTYPES:

16 specimens ( 1 2 of them dissected).

vayssierei. Pleurobranchaea vayssierei Ev. Marcus

et Gosliner, 1984: 38, 39, fig. 19. Iype locali¬

ty: Alger, Algeria. HOLOTYPE (by original

désignation, figured by Ev. Marcus fie Gosliner

1984, fig. 19): specimen (dissected), leg.

Vayssière + radula, jaws and reproductive Sys¬

tem slides.

THECOSOMATA

Family Cavoliniidaf. J. E. Gray, 1850

acicula. Creseis aciculd Rang, 1828a: 317, pl. 17,

fig. 6. Type locality: “Merdes Indes” (= Indian

Océan). l.ECTOTYPE (selected by van der

Spoel 1976: 189): shell (Fig. 6H) -t 18 paralec-

totypes (broken).

affinis. Hyalaea ajfinis d’Orbigny, 1834: 91-93,

pl. 5, figs 6-10. Type locality: approximately

between 30° ro 34°S and 78" to 90"W (Pacific

Océan). LECTOTYPF, (selected by van der

Spoel, 1976: 194): shell. leg. d’Orbigny

(Fig. 61) + 1 paralcctotype. Rcmarks:

According to van der Spoel (1967; 1976), this

nominal species is merely the form affinis of

Cavolinia tridentata (Forsskâl in Niebuhr

1775).

angulata. Hyalaea angubita Souleyet, 1852: 152,

1 53, pl. 5, figs 1 -6. Type locality: not specified;

cited front Atlantic, Indian Océan and Sea of

China. SYNTYPE: 1 specimen, front Indian

Océan, leg. Eydoux and Soulever. Remarks:

Sec continents on Cavolina angulosa J. E. Gray,

1850.

angulosa. Cavolina angulosa). E. Gray, 1850: 8.

Iype locality: Indian Océan. LECTOTYPE

(selected by van der Spoel 1976: 193): speci¬

men, leg. Eydoux and Souleyet. Remarks:

Eydoux & Souleyet (1846-1849) figured this

species under the vernacular French nante

“Hyale angulée’'. J. E. Gray (1850) is the First

author who introduced the liante Cavolina

angulosa under binominal nomenclature,

without description, but referring to the

figures in the Atlas of Eydoux fit Souleyet.

However, J. E. Gray incorrectly made référen¬

cé to pl. 4, figs 1-6, whereas Eydoux fie

Souleyet (1846-1849) figured this species in

pl. 5, figs 1-6. This is certain to be a typogra¬

phies! error hecause ]. E. Gray explicitly men-

rioned the name "Hyale angulée”. l.ater,

Souleyet (1852) established Hyalaea angulata

as a new species. giving a description and a

référencé to tlie sanie figures. Therefore, tliis is

a case of independetu latinization of the sanie

liante. Van der Spoel et al. (1993) placed this

species in the genus Diacavolinaia van der

Spoel, 1987.

astesana. Cuvieria astesana Rang, 1829b: 498,

499, pl. 19, fig. B. Type locaïity: “l’Astésan”

(= Asti), Italy (Pliocène). SYNTYPE: 1 shell

(broken) (Fig. 6J).

chaptalii. Cleo dora cbaptalii Souleyet, 1852:

183, 184, pl. 7, figs 1-5. Type locality: Cape of

Good I lope (Southern Africa). HOLOTYPE

(by monotypy, figured by Eydoux fit Souleyet

1846-1849, pl. 7, figs 1-5): shell, leg. Eydoux

ZOOSYSTEMA • 1998 - 20(4)

715

Valdés Â. & Héros V.

and Souleyet (Fig. 6K). Remarks: See remarks

under ('lia chaptalii J. F.. Gray, 1850.

chaptalii. Clin chaptalii J. F. Gray, 1850: 14.

Type localtiy: Cape of Good Hope (Southern

Africa). HOLOTYPE (by monotypy, figured

by F.ydoux JY Souleyet 1846-1849, pi. 7,

figs 1-5): shell, leg. Eydoux and Souleyet

(Fig. 6K), Remarks: F.ydoux & Souleyet

(1846-1849, pl. 7, figs 1-5) figured this species

under the vernacular French name “Clêodore

de Chaptal”. [. E, Gray (1850) is the first

author who introduced the name Cltu chaptalii

under binominal nomenclature, without a des¬

cription, but referring to the figures in the Atlas

of F.ydoux Sè Souleyet. Latcr, Souleyet (1852)

descri bed Cleo dora chaptalii as a new species,

giving a description and a référencé ro the same

figures. Therefore, tins is a case ol independent

latini/ation of the sanie name. Since borh spe¬

cies are based on the same single specimen,

Cleodom chaptalii Souleyet, 1852 is an objecti¬

ve junior synonym of' Clin chaptalii J. E. Gray,

1850. In addition, they are secondary homo¬

nyms. According ro van der Spoel (1967,

1976), thts nominal species is just the form

major of Diacria trispinosa (Lesueur, 1821).

clava. Creseis clava Rang, 1828a: 317, pl. 17,

fig. 5. Type locality: Agulhas Bank. Southern

Africa. LECTOTYPE (selected by van der

Spoel, 1976: 189): shell (Fig. 9A) + 4 paralec-

totypes. Remarks: According ro van der Spoel

(1967: 59), this nominal species is merely a

form clava of Creseis ad.eu la Rang, 1828.

gibbosa. Hyalaea gihbosa Rang in d Orbigny

1834: 95, 96, pl. 5, figs 16-20. Type locality:

not specified; cited from approximately bet-

ween 34°N to 40"S and 26" to 43"W (Atlantic

Océan). LECTOTYPE (selected by van der

Spoel, 1976: 195): shell, leg. d'Orbigny

(Fig. 9B) + 3 paralectotypes. Remarks:

According to van der Spoel ( 1967; 1976), this

species belongs to the genus Cctvolinia

Abildgaard, 1791.

globulosa. Ihyalaea globulosa Rang in Souleyet

1852: 142,' 143, pl. 4, figs 20-24 Type locali¬

ty: not specified; cited (rom Pacific, Atlantic,

Indian Océan and Sea of China. SYNTYPES:

5 shells (Fig. 9C) front "Nouvelle Hollande’

(= Australia), leg. Rang. Remarks: See remarks

under Cavolinaglobulosa). F. Gray, 1850.

globulosa. Cavolina globulosa ). F.. Gray. 1850: 8.

lype locality: not specified; cited from Pacific,

Atlantic, Indian Océan and Sea of China.

LECTOTYPE (selected by van der Spoel,

1976: 196): shell (Fig. 90 -t 4 paralectotypes

from "Nouvelle Hollande" (= Australia), leg.

Rang. Remarks: Eydoux & Souleyet

(1846-1849, pl. 4, figs 20-24) figured this spe¬

cies under the vernacular French name "Hyale

globuleuse'. J. E. Gray (1850) is the first

author who introduced the name Cavolina glo¬

bulosa under binominal nomenclature,

without a description, but citing to the figures

in the Atlas of Eydoux Soulevée. Latcr, Rang

in Souleyet (1852) dcscribcd Hyalaea globulosa

as a new species, giving a description and a

reference to the same figures. Therefore, this is

a case of independent latini/ation of rhe same

name. Both nominal species Cavolina glolntlo-

sa). F. Grav, J 850 and Hyalaea globulosa Rang

itt Soulever 1852 are secondary homonyms.

inflala. Cleodora influa Soulever, 1852: 188,

pl. 7, figs 17-19, Type locality: Atlantic Océan.

SYNTYPE: 1 specimen, leg. Eydoux and

Souleyet. Remarks: Sce remarks on Balantium

inflation). E. Gray. 1850.

inflatum. Balantium inflatura). E. Gray, 1850:

15. Type locality: Atlantic Océan. SYNTYPE:

1 specimen, leg. Eydoux and Souleyet.

Remarks: Eydoux & Souleyet (1846-1849,

pl. 7, figs 17 19) figured this species under the

vernacular french name "CIcodorc renflée”.

J. E. Gray ( 1850) is the first author who intro¬

duced the name Balantium inflatum under

binominal nomenclature, without a descrip¬

tion, but citing to the figures in the Atlas of

Eydoux & Souleyet. Later, Souleyet (1852)

described Cleodora influa as a new species,

independenly giving a description and a réfé¬

rencé to the same figures. Therefore, this is a

case ol independent latini/ation of the same

name. Since both species are based on the same

type material, Cleodom influa Souleyet, 1852

is an objective junior synonym of Balantium

inflatum). E. Gray, 1850. In addition, rhey are

secondary homonyms. According to van der

Spoel (1967: 76), this could be a synonym of

Clio recurva (Children, 1823).

716

ZOOSYSTEMA • 1998 -20(4)

Opisthobranch mollusc type specimens in MNHN

levigata. Hyalaea levigata d’Orbigny, 1834: 110,

111, pi. 7, figs 15-19. Type locality: not speci-

fied; cjted from approximately 20°S, 87°W

(Pacific Océan). SYNTYPE (figured by

d’Orbigny 1835, pi. 7, figs 15-19); 1 shell

juvénile, leg. d Orbigny (Fig. 9D). Remarks:

According to van dcr Spocl et al. (1993), this is

probablv a junior synonym of Diacavolinia

Longiroftns (Lesueur, 1821 ),

limbata, Hyalaea /tnt bat a d’Orbigny, 1834:

101-103, pl. 6, figs I 1-15. Type localiry: not

specified; cited from approximately beeween

30° to 33°S and 80° to 90°W (Pacific Océan),

and approximately between 12°N to 24°S and

25° to 28°W (Atlantic Océan). LECTOTYPE

(selected by van der Spoel, 1970: 193): shell,

from Atlantic Océan, leg. d’Orbigny

(Fig. 9E) + 2 paralectotypes. Remarks: Van der

Spoel et al. (1993) placed this species in the

genus Diacavolinia van der Spoel, 1987.

longicaudattis. Cita longicaudatus Souleyet,

1852: 286-288, pl. 14, figs 17-21. Type locali-

ty: approximately 10°N * 21°W (Atlantic

Océan). SYNTYPE: 1 specimen, leg. Eydoux

and Souleyet. Remarks: This is the type spe-

cies of the genus Clionina Pruvot-Fol, 1924, by

monotypy.

oblonga. Clio oblonga Rampai, 1997: 175-185,

Pl. 1, figs A-l. Type locality: Tyrrhenian Sea,

39°36’N - 13°49'E, 2103 m (Quaternary).

HOLO’lYPE: 1 shell (figured by Rampai,

1997: pl, 1, figs A-D) -t 3 paratypes.

orbignii. Hyalea orbignii Rang, 1827: 383. Type

locality: Saint-Paul-les-Dax (lower Miocene:

Burdigalian, France). HOLOTYPE; van der

Spoel (1976: 192) records the holotype to be

présent in MNHN, but this specimen is cur-

rently untraceable,

rangii. Hyalea rangii Deshayes, 1831: 310. Type

locality: coast ol Africa. SYNTYPF.S: 5 shells

(Fig. 9F). Remarks: According to van der Spoel

(1967), this is a junior synonym of Cavolinia

tridentata (Forsskâl in Niebuhr 1775).

striata. Creseis striata Rang 1828a: 314, 315,

pl. 17, fig. 3. Type locality': not specified; cited

from Atlantic and Indian Océans. SYN-

TYPES: 9 shells (broker») (Fig. 9G). Remarks:

This is the type species of the genus Hyalocylis

de Folin, 1875, by monotypy.

uncinata. Hyalea uncivata Rang, 1829a. 114.

Type locality: unknown. I.ECTOTYPE (selec-

ted by van der Spoel, 1976: 195): shell

(Fig. 9H) + 11 paralectotypes. Remarks: Van

der Spoel (1987: 78) placed this species in the

genus Cavolinia Abildgaard, 1791.

Family CY.VIBU1.IIDAE J. E. Gray, 1840

ovata. Cymbidia ouata Quoy et Gaimard, 1833:

373, 374, pl. 27, figs 25-30. Type locality:

Amboine, Indonesia. LECTOTYPE (selected

by van der Spoel, 1976: 198): specimen, Icg.

Quoy and Gaimard + 7 paralectotypes.

Remarks. Van der Spoel ( 1976: 41 ) placed this

species in the genus Corolla Dali, 1871.

NUDIBRANCH1A-DORIDINA

Family GONIODORIDIDAE H. et A. Adams, 1854

darwini. Thecacera darwini Pruvot-Fol, 1950:

49-51, figs 1-4. Type localiry: “Baie Orange",

Bahîa de Nassau, Chile. SYNTYPES: 7 speci¬

mens (1 of them dissected).

pila sa. Hopkimia pilosa Bouchet et Ortea, 1983:

227-231, figs 1-7. Type locality: Nguetu Reef,

New Caledonia. HOLOTYPE (by original

désignation): specimen + 1 pararype.

polycerelloides. Bermudella polycerelloïdes Ortea

et Boucher, 1983: 50-54, figs 1,2. Type locali-

ty: Los Cristianos, Tenerife, Ganary Islands.

HOLOTYPE (by' original désignation): speci¬

men + 1 paratype.

savignyi. Goniodoris savignyt Pruvot-Fol, 1933:

117, 118, pl. 2, figs 23-26. Type locality: Gulf

of Suez, F.gypt. HOLOTYPE (by monotypy,

figured by Pruvot-Fol, 1933, pl. 2, figs 23-26):

specimen (dissected), leg. Dollfus. Remarks:

This is the type species of the subgenus Gottio-

doridclla Pruvot-Fol, 1933, by monotypy.

violacea. Goniodoris violacea Risbec, 1928:

177-179, fig. 53. pl. 7, fig. 5. Type locality:

Pointe de l’Artillerie, Nouméa, New

Caledonia. HOLOTYPE (by monotypy, figu¬

red by Risbec, 1928, fig. 53, pl. 7, fig. 5): spe¬

cimen (dissected). Remarks: In the figure cap-

don of pl. 7, fig 5, this is cited as Chromodoris

violacea.

ZOOSYSTEMA • 1998 -20(4)

717

Valdés A. & Héros V.

Family ONCHIDOR1D1DAE Gray, 1827

reticulata. Onchidoris miculata Ortea, 1979b:

169-173, 11g. 2. Type locality: Concha de

Artedo, Asturias, Spain. 1 lOLOTYPE (by ori¬

ginal désignation, figured by Ortea 1979b,

fig. 2): Specitnen t photo.

tridactila. Onchidoris tridactila Ortea et

Ballesteros, 1982: 241-246, figs 2, 3 (in part),

7 (in part), 8 B, C. Type locality: Verdicio,

Asturias, Spain. HOLOTYPE (by original

désignation): specimen.

Family POFYCF.RIDAF Adler et Hancock, 1845

Julgurans. Plocamopherus fulgurans Risbec, 1928:

206-208, fig. 63, pl. 4, fig. 4. Type locality:

Nouméa and Bourail, New Calcdonia. SYN-

TYPE: 1 specimen.

gulo. Plocamopherus gulo Ev. Marcus, 1979:

134-136, figs 7-12. Type locality: Calypso

Expédition, stn 145. 26°34’S - 47°22’W, bet-

ween Parana and Santa Catarina, Brazil,

100 m. HOLOTYPE (by monotypy, ligured

by Ev. Marcus 1979, figs 7-12): specimen

(considerably damaged) + radula and jaws

slide.

picta. Polycera picta Risbcc, 1928: 200-202,

fig. 61, pl. 7, fig- 6. Type locality: Pointe de

l’Artillerie (Nouméa) and Bourail, New

Caledoma. SYNTYPES: 2 spccimens.

Family GyMNODORIDIDAE Odhner, 1941

ceutae. Tambja ceulae Garcia-Gômez et Ortea,

1988: 302-305, figs 1-7, pl. 1. Type locality: El

Pineo, Ceuta, Spain. HOLOTYPE (by original

désignation, figured by Garcia-Gômez &

Ortea 1988, figs 1-7, pl. 1): specimen (dissec-

ted) + radula slide.

europaea . Roboastra europaea Garcia-Gômez,

1985: 169-174, figs 1-5. Type locality: Tarifa,

Spain. HOLOTYPE (by original désignation):

specimen.

perlucens. Trevelyana perlucens Risbec, 1928:

185-189, fig. 57, 57bis, pl. 1, fig. 12, pl. 5,

fig. 1. lype locality: New Calcdonia. SYN¬

TYPES: 3 specimens.

suggens. Trevelyana suggens Risbec, 1928:

190-193, fig. 58, pl. B, fig. 1, pl. 5, figs 3, 6,

pl. 7, fig. 10. Type locality: “Tembia”

(= Timbia) and Nouméa, New Caledonia.

SYNTYPES: 2 specimens.

Family Vayssifrfidaf Thiele, 1931

caledonica. Vayssierea calédonien Risbec, 1928:

290-292, fig. 98, pl. 12, fig. 8. Type locality:

New Caledonia. SYNTYPES: 6 specimens.

Remarks: This is the type spccies of the genus

Vayssierea Risbec, 1928, by monotypy.

Family HexaBRANCHIDAF Bergh, 1891

lacera. Doris lacera Cuvier, 1804a: 452-465, 473,

pl 73, figs 1-3. Type locality: Timor,

lndonesia. SYNTYPES: 2 specimens (di.ssec-

ted), leg. Péron and Lesueur. Remarks:

Pruvot-Fol (1934a) placed this species in the

genus Hexabranchus Ehrenberg, 1831.

sandwichiensis. Doris sandwichiensis Souleyet,

1852: 451,452, pl. 25, figs 1 -4. Type locality:

“îles Sandwich" (= Hawaii). SYNTYPE: 1 spe¬

cimen (dissected), leg. Eydoux and Souleyet.

Remarks: According to Thompson (1972),

this is a junior synonym of Hexabranchus san-

guineus (Rüppell et Leuckart, 1828).

Family DORIDIDAF. Rafinesque, 1815

aspera. Staurodoris aspera Risbec, 1928: 101,

102, fig. 20, pl. 2, fig. 8. Type locality: Baie de

Plum and Nouméa, New Caledonia. SYNTY¬

PE: l specimen.

immonda. Platydoris immonda Risbec, 1928: 84,

85, fig. 12, pl. 1, fig. 4. Type locality: New

Calcdonia. SYNTYPE: 1 specimen. Remarks:

According to Brodic & Willan (1993), this is a

junior synonym of Siraius nucleold (Pease,

1860).

lopbatus. Thorybnpus lophatus Bouchet, 1977:

43-46, figs 11-13, pl. 1, fig. D. Type locality:

Biaçores Expédition, stn 241, 37°37N -

25°32W. of'f Santa Maria, Azores, 395-465 m.

HOLOTYPE (by original désignation, figured

by Bouchet, 1977, figs 11-13, pl. 1. fig. D):

specimen (dissected). Remaries: This is the

type species of the genus Tboiybopus Bouchet,

1977, by original désignation.

718

ZOOSYSTEMA • 1998 - 20(4)

Opisthobranch moliuse type specimens in MNHN

Family ArtCHinORininAF. Bergh, 1892

antarctica. Archidoris tuberculata var. antarctica

Vayssière, 1917: 16, 17. Type loealiry: beeween

“Ile Jenny” and “Terre Adélaïde”, Marguerite

Bay (Antarctica). SYNTYPES: 3 specimens

(dissected). Remarks: According to Wagele

(1993), this subspecific-rank taxon is a junior

synonym oF Austrodons kerguelenensis (Bergh,

1884).

carinata. Doris carinata Quoy et Gaimard, 1832:

254, pl. 16, figs 10-14. Type locality: “rivière

Tamise” (= Thames), New Zeaiand. SYNTYPE:

1 specimen. Remarks: This is the type species of

the genus Atagema J. E. Gray, 1850, by mono-

typy.

flava. Guynnia flava Risbec, 1928: 103, 104,

fig. 21, pl. 3. fig. 6. Type locality: New

Caledonia. SYNTYPE: 1 specimen.

granuLatissima. Archidoris grctnulatissima

Vayssière, 1917: 17-19, pl. 4, Figs 43, 44. Type

locality: "Dragage IX” (coordinates unknown),

Adelaide Island, and “Baie de l'Amirauté",

King Georges Island, Antarctica. SYNTYPES:

4 specimens (dissected). Remarks: According

to Wagele (1993), this is a junior synonym of

Austrodoris kerguelenensis (Bergh, 1884).

odonoghuer. Archidoris odonoghuei Pruvot-Fol,

1933: 11 S-120, pl, 2, figs 18-22. Type locality:

Gulf of .Suez, Egypt SYNTYPES: 3 specimens

(dried), leg. Dollftts.

paagoumenei. Phlegmodoris paagoumenei Risbec,

1928: 87-90, fig. 15, pl, B, fig. 3, pl. 3, fig. 1.

Type locality: Paagoumène, New Caledonia.

HOLOTYPE (by monotypy): radula.

schembrii. Doris scheinbrii Verany, 1846: 97,

101, 102. Type locality: Gulf o( Genova

(Méditerranean Sca). SYNTYPES: 2 speci¬

mens. Remarks: l his is the only specimen of

the Verany opisthobranch collection which

could be located. It seems to hâve been borro-

wed by Vayssière and iater was deposited in

MNHN. The remainder of the Verany collec¬

tion is probably lost.

vayssierea. Archidoris vayssierea O’Donoghue,

1929: 812-814, fig. 223. Type locality: Suez

Canal, Egypt. SYNTYPE (figured by Vayssière,

1912, pl. 2, figs 27-29): 1 specimen (dissec¬

ted), leg. Gravier. Remarks: O’Donoghue

(1929) introduccd the naine Archidoris vayssie¬

rea based on one specimen reported by

Vayssière (1912) from the Suez Canal (under

the name Archidoris staminea Basedow et

Hedley, 1905) and deposited in MNHN, and

another specimen of his own collection from

this area. Both specimens are syntypes of this

species.

Family BapïODORIDIDAE Odhner, 1926

boucheti. Carrninodoris boucheti Ortea, 1979c:

579-582. figs 5-7. Type locality: Concha de

Artedo, Asturiax, Spam. HOLOTYPE (by ori¬

ginal désignation): specimen.

ceneris. Paradons ceneris Ortea, 1995: 21-25,

figs IA, 1E, 13-15. Type locality: Puerto de

Arrccife, Lanzarote, Canary Islands. HOLO¬

TYPE (by original désignation, figured by

Ortea 1995, figs IA, 1E, 13-15): specimen

(dissected).

fongosa. Baptodoris fongosa Risbec, 1928:

111-115, fig. 25, pl. 2, fig. 9, pl. 3, fig. 9. Type

locality: New Caledonia. SYNTYPES: 2 speci¬

mens + spawn.

inversa. Paradons inversa Ortea, 1995: 14-17,

figs IC. 6-8. Type locality: El Médano,

Tenerife, Canary Islands. HOLOTYPE (by

original désignation, figured by Ortea 1995,

fig. 6A): specimen.

mollis. Paradons mollis Ortea, 1995: 18-21,

figs IB, 9-12. Type locality: Adeje, Tenerife,

Canary Islands. HOLOTYPE (by original

désignation, figured by Ortea 1995, figs IB,

9-12): specimen (dissected).

peresi. Baptodoris perezi Liera et Ortea in Ortea

et al. 1982: 24-29, figs 9-1 1, pl. 1, fig. C. Type

locality: Playa de San Juan, Tenerite, Canaiy

Islands. HOLOTYPE (by original désigna¬

tion): specimen.

Family DISCODORIDIDAE Bergh, 1891

ascitica. Taringa ascitica Ortea, Pérez-Sânchez et

Liera, 19S2: 34-38, figs 15-17, pl. 2, fig. H.

Type locality: La lslcta, Lanzarote, Canary

Islands. HOLOTYPE (by original désignation,

figured by Ortea et al 1982, figs 15-17, pl. 2,

fig. H): specimen (dissected).

bacalladoi. Geitodoris bacalladoi Ortea, 1990:

ZOOSYSTEMA • 1998 -20(4)

719

Valdés Â. & Héros V.

116-119, figs 12-14. Type locality: Agua

Duke, ’T’enerife, Canary Islands. HOLOTYPE

(by original désignation): specimen + 1 para-

type.

bacalladoi. Taringa bacalladoi Ortea, Pérez-

Sanchez et Liera, 1982: 42-45, figs 20-22.

pl. 1, fig. D. Type locality: Arrccife, Lanzarote.

Canary Islands. HOLOTYPE (by original

désignation): specimen + 1 radula slide proba-

bly of rhe holoty|>e.

bonosi. Geitodoris borxosi Ortea et Ballesteros,

1981: 337-341, figs 1-3. Type locality: Cabo

Botafoch, Ibiza (Mediterranean Sea).

HOLOTYPE (by original désignation): speci¬

men.

confusa. Discodoris confus a Ballesteros, Liera et

Ortea, 1984: 236-244. figs 6-8, 10C. Type

locality: Las Galetas, Lanzarote, Canary

Islands. HOLOTYPE (by original désigna¬

tion): specimen.

edwardsi. Discodoris edwardsi Vayssière, 1902:

232-234, pl. 9, figs 4-9. Type locality:

Talisman (1883) Expédition, off Cape Ghir,

Morocco, 2100 m. HOLOTYPE (by mono-

typy): specimen.

erythraeensis. Discodoris erythraeensis Vayssière,

1912: 23, 24, pi. 10, figs' 150-152, Type loca¬

lity: “Nord d’Amboulé”, Golfe de Tadjoura,

Djibouti. HOLOTYPE (by monotypy, figured

by Vayssière 1912, pl. 10, figs 1 50-1 52): speci¬

men (dissected), leg. Gravier. Retnarks: The

name Discodoris erythraeensis was first inrrodu-

ced by Vayssière (191 1) without description

(nomen nudum).

fava. Taringa fana Ballesteros, Liera et Ortea,

1984: 246-252, figs 11-14. Type locality:

Cubellas, Spain. HOLOTYPE: said to be

deposited in MNHN in the original publica¬

tion, but never presented to the muséum.

lutea. Archidoris mandata var. lutea Vayssière,

1919: 57-65, pl. 4. figs 1-11. Type locality:

Carry, Golfe de Marseille, France. SYN-

TYPES: 2 spécimens (dissected). Remarks:

Thompson & Brown ( 1984) ptaced rhis taxon

in the genus Doris Linnaeus, 1758, as a syno-

nym of Doris sticta (Iiedale et O’Donoghue,

1923), and therefore it becomes a junior

secondarv homouym of Doris lutea Risso,

1818.

nayarita. Peltodoris nayarita Ortea et Liera,

1981: 47-51, figs 1 -4. Type localiry: Isla lsabel,

Pacific coast of Mexico. HOLOTYPE (by ori¬

ginal désignation, figured by Ortea &i Liera

1981, figs 1-4): specimen (dissected).

Remarks: In the original description this spe-

cics" is indistinedy cîted as Peltodoris nayarita

and Anisodoris nayarita.

notiperda. Discodot is notiperda Risbec, 1956: 15,

16, pl. 9, figs 43-49, pl. 10, figs 50, 51. Type

locality: Port Dayot and NhaTrang, Vietnam.

SYNTYPES: 9 specimens (3 of thenr dissec¬

ted).

oleica. Taringa oleica Ortea, Pérez-Sânchez et

Liera, 1982: 29-33, figs 12-14, pl. 2, fig. E.

Type locality: Melenara, Las Palmas, Canary

Islands. HOLOTYPE (by original désigna¬

tion): specimen.

perfossa. Geitodoris perfossa Ortea, 1990:

109-115, figs 8-11. Type locality: Los

Cancajos, I a Palma, Canary Islands. HOLO¬

TYPE: said to be deposited in MNHN in rhe

original publication, but never presenred ro the

muséum.

rosi. Discodoris rosi Ortea, 1979c: 575-579,

figs 1-4. Type locality: Ovinana, Asturias, Spain.

! 10 LOI AT E (by original désignation): speci¬

men.

rubens. Discodoris rubens Vayssière, 1919: 65-67,

pl. 6, figs 39-45. Type locality: Carry, Golfe de

Marseille, France. SYNTYPES: 3 specimens

(dried).

sordida. Doris sordida Quoy et Gaimard, 1 832:

266, pl. 19, figs 12, 13. Type locality: “îlots

aux Cerfs”, “île de France” (= Mauritius),

Indian Océan. SYNTYPE: l specimen (dissec¬

ted), leg. Quoy and Gaimard. Remarks: This

name is preoccupied by Doris sordida Rüppell

et l.euckart, 1828 (see sordidata). Pruvot-Fol

(1934b) placed this species in the genus

Discodoris Bergh, 1877.

sordidata. Doris sordidata Abraham, 1877: 206.

Type locality: “îlots aux Cerfs , “île de France”

(= Mauritius). Indian Océan. SYNTYPE:

1 specimen (dissected), leg. Quoy and

Gaimard. Remarks: This is a replacement

name for Doris sordida Quoy et Gaimard,

1832, preoccupied by Doris sordida Riippell et

Leuckart, 1828.

720

ZOOSYSTEMA • 1998 • 20 (4)

Opisthobranch mollusc type specimens in MNHN

tritorquis:. Taringa tritorquis Ortea, Pérez-

Sanchez et Liera, 1982: 38-41, figs 18, 19,

pl. 2, fig. G. Type lotality: Puerto del Carmen,

Lanzarote, Canary Islands. HOLC3TYPE (by

original désignation): specimeri.

Family KhnTRODORIDIDAE Bergh, 1891

marchadi. Jorunna rnarchadi Rjsbec, 1956: 16,

17, pl. 11, Figs 54-57. Type locality: Patrie

Island, Paracels Islands (Southern Sea of

China). HOLOTYPE (by monotypy, figured

by Risbec, 1956. pl. 11, fïgs 54-57): specimen

(dissected).

onubensis. Jorunna onubensis Cervera, Garcia-

Gômez et Garcia, 1986: 119-128, figs 9-16,

pl. 1. Type locality: El Portil, Huelva, Spain.

HOLOTYPE (by original désignation): speci¬

men (dissected) + radula slide.

Family AsTERONOTtDAF. Thiele, 1931

solea. Doris soleu Cuvier, 1804a: 452-466, pl. 74,

figs 1, 2. Type locality: “île de France”

(= Mauritius), Indian Océan. SYNTYPE (figu¬

red by Pruvot-Fol 1934a, pl. 2, figs 1, 2):

1 specimen. Remarks: Pruvot-Fol (1934a) pla-

ced this species in the genus Asteronotus

Ehrenberg, 1831.

Family PLATYDORIDIDAE Bergh, 1891

cannata. Platydoris carinata Risbec, 1928: 85,

86, fig. 13, pl. 3, fig 10. Type locality: île

Hugon, New Caledonia. SYNTYPES: 2 speci¬

mens + spawn.

cruenta. Doris cruenta Quoy et Guimard, 1832:

260, 261, pl. 18, figs 5-7. Type locality: New

Guinea, Indonesia. SYNTYPF2 ! specimen

(dissected). Remarks: Pruvot-Fol (1934b) pla-

ced this species in the genus Platydoris Bergh,

1877.

laminea. Platydoris laminea Risbec, 1928: 86,

87, fig. 14, pl. 1, fig. 11. Type locality: New

Caledonia. SYNTYPES: 2 specimens.

macula ta. Platydoris mandata Bouchet, 1977:

37-41, figs 6-8, pl. 2, figs B, C. Type locality:

Thalasset (1970) Expédition, srn W413,

43°50N - 06*09' W, Bay of Biscay, 500-540 m.

HOLOTYPE (by original désignation): speci¬

men + 1 paratypc.

noumeae. Platydoris noumeae Risbec, 1928:

80-82, fig. 10, pl. 2, fig. 2. Type locality: New

Caledonia, SYNTYPES: 2 specimens.

scabra. Doris scabra Cuvier, 1804a: 466. Type

locality: Timor, Indonesia. SYNTYPE (figured

by Pruvot-Fol 1934a, fig. 3): I specimen (dis¬

sected), leg. Peron. Remarks: Pruvot-Fol

(1934b) placed this species in the genus

Platydoris Bergh, 1877.

spongilla. Platydoris spongilla Risbec, 1928:

82-84, fig. 11, pl. 5, fig. 12. Type locality:

Rocher à la Voile and Cale de Halage,

Nouméa, New Caledonia. SYNTYPE: 1 speci¬

men.

stomascuta. Platydoris stomascuta Bouchet, 1977:

35-37. figs 4, 5, pl. 2, fig. D. Type locality:

Biaçores Expédition, stn 41, 37°43N -

29 ,5 04W, ofi Azores, Princesse Alice Bank,

450-475 m. HOLOTYPE (by original dési¬

gnation): specimen + 2 paratypes.

Family MlAMIRIDAF. Bergh, 1891

rugosa. Gravieria rugosa Vayssière, 1912: 30-32,

pl. 10, figs 144-149. Type locality: “îles

Musha”, Golfe deTadjoura, Djibouti. HOLO¬

TYPE (by monotypy, figured by Vay r ssière

1912, pl. 10, figs 144-149): specimen (dissec¬

ted), ieg. Gravier. Remarks: The nante

Gravieria rugosa was firstly introduced by

Vayssière (1911), withour description {nomen

nudum). This is the type species of the genus

Gravieria Vayssière, 1912, by monotypy.

Family Al.nLSlDAF Odhner, 1939

expleta. Aldisa expleta Ortea, Pérez-Sânchez et

Liera, 1982: 10-13, figs 2, 3, pl. 1, fig. A. Type

localityc Playa del Carbon, Gran Canaria,

Canary Islands. HOLOTYPE (by original

désignation): specimen. Remarks: According

to Mtllen & Goslmer (1985) this species is a

junior synonym of Aldisa banyulensis Pruvot-

Fol, 1951.

nbatrangensis. Aldisa nhatrangensis Risbec,

1956: 14, 15, pl. 20, fig. 109, pl. 22, upper

riglu figure. Type locality: Hon Lon, Nha

Trang, Vietnam. HOLOTYPE (by original

ZOOSYSTEMA • 1998 -20(4)

721

Valdés Â. & Héros V.

désignation, figurcd by Risbec 1956, pl. 20,

fig. 109. pi. 22): specimen (dissected).

smaragdina. Aidiin smaragdina Ortea, Perez-

Sanchez et Liera, ] 982; 14-18, figs 4, 5, pl. 1,

fig. B. Type locality: L os Cancajos, La Pal ma,

Canary Islands. HQLOTYPF (by original

désignation): specimen. Remarks: According

to Millen & Gosliner (1985) this species is a

junior synonym of Aldisa binotata Pruvot-Fol,

1953.

Family Chromodoiudidae Bergh, 1892

aeruginosa. Glossodoris aeruginosa Rudman,

1995: 12-16, figs 6D, 11-13. Type locality:

Banc Gail, between Nouméa and Ile Ouen,

New Caledonia. HOLOTYPE (by original

désignation, figured by Rudman 1995,

figs 6D, 11-13): specimen (dissecied), leg.

Laboute.

albofimbria. Dunnllcdoris albofnnbria Rudman,

1995: 22-25, figs 6F, 19-21. Type locality;

Passe de Koumac, New Caledonia. HOLO¬

TYPE (by original désignation, figured by

Rudman 1995, figs 6F, 19-21): specimen (dis-

sectéd).

atromarginata. Doris atromarginata Cuvier,

1804:1: 473, pl. 74, fig. 6. Type locality: “Mer

des Indes” (— Indian Océan). SYN TYPES:

2 specimens, leg. Pérou. Remarks: This is the

type species of the genus Doriprismalica

d’Orbîgny, 1839, by subséquent désignation

by Herrmannsen (1846). Following Rudman

(1984) Dtmprinnatica is a junior synonym of

Glossodoris Ehrenberg, 1831.

auréola. Glossodoris auréola Rudman, 1995:

16-19, figs 6F., I0B, 14, 15. Type locality:

Passe Deverd, off Koumac, New Caledonia.

HOLOTYPE (by original désignation, figured

by Rudman 1995, figs 6E, 10B, 14. 15): .speci¬

men (dissected), leg. von Cosel.

boucbeti. Cbromadoris boucbeti Rudman, 1982:

190-193, figs IB, 4, 5. Type locality: northern

entrance to Longogoni Passage, Mayotte,

Comoro (Indian Océan). HOLOTYPE (by

original désignation, figured by Rudman 1982,

fig. 1 B): specimen + 3 para types.

britoi. Cbromadoris britoi Ortea et Pérez-

Sânchez, 1983: 62-65, figs 1-3, pl. 1, fig. 3.

Type locality: Agita Duke, Tenerife, Canary

Islands. HOLOTYPE (by original désigna¬

tion): specimen.

caledonicum. Ceratosoma caledonicum P. Fischer,

1876: 92, 93. Type locality: New Caledonia.

SYNTYPES: 2 specimens. Remarks: Accord¬

ing to Rudman (1988), this is a junior syno¬

nym of Ceratosoma trilobatum (|. E. Gray,

1827),

cantabrica. Hypsebdoris cantabrica Bouchet et

Ortea, 1980: I 18-121, figs 1-6, 13. Type loca¬

lity: Hendaye, France. HOLOTYPE (by origi¬

nal désignation): specimen + 11 paratypes.

ciminoi. Hypsebdoris ciminoi Ortea et Valdés in

Ortea et al, 1996: 134-136, figs 12B, 104,

105. Type localiry: Bomfin, Angola. HOLO¬

TYPE (by original désignation, figured by

Ortea étal. 1996, figs 12B, 104, 105): speci¬

men (dissected), leg. Rolan.

elavata. Cbromadoris clavata Risbec, 1928: 151,

152, fig. 42, pl. 7, fig. 2. Type locality; Baie de

l'Orphelinat, Nouméa, New Caledonia. SYN-

TYPE: 1 specimen. Remaries: According to

Rudman (1982), this is a junior synonym of

Cbromadoris striatella Bergh, 1877-

coi . Chromodoris coi Risbec, 1956: 9, 10, pl. 11,

figs 58, 59, pl. 22, lower figure. Type locality:

Hon Lon, Nha Trang, Vietnam. SYNTYPE:

1 specimen. Remaries: Rudman (1987) placed

this species in the genus Chromodoris Aider et

Hancock ( 1855).

decorata. Chromodoris decorata Risbec, 1928:

152-154, fig, 43, pl. 7, fig. 4, Type locality:

Baie de l’Orphelinat, Nouméa, New

Caledonia. SYNTYPE: 1 specimen (dissected).

Remarks: According to Rudman (1986b), this

is a junior synonym of Hypsebdoris macubsa

(Pease, 1871)-

dollfusi. Glossodoris dollfusi Pruvot-Fol, 1933:

126-128, pl. 1, figs 7, 8, pl. 3, fig. 40. Type

locality: Dell fus expédition (29°49’-29M5’N,

32"30'-32°27’E) 31 m, Red Sea, Egypt.

HOLOTYPE (by monotvpy, figurc-d by

Pruvot-Fol 1933, pl. 1, figs 7, 8, pl. 3, fig. 40):

specimen (dissected), leg. Dollfus.

elegans. Doris elegMS Quoy éi Gainïard, 1832:

273, 274, pl. 20, figs 12-14. Type locality:

Tongatapu (Pacific Océan). SYNTYPE: 1 spe¬

cimen, leg. Quoy and Gaimard. Remarks:

722

ZOOSYSTEMA • 1998 • ZO (4)

Opisthobranch mollusc type specimens in MNHN

Pruvot-Fol (1934b) placed this species in the

genus G/ossodoris Ehrenberg, 1831.

espinosai. Hypselodoris espinosai Ortea et Valdés

in Ortea et ni, 1996: 139-142, figs 12E,

109-1 I 1 Type locality: Puerto Morelos,

Quintana Roo, Mexico. IIOLOTYPE (by ori¬

ginal désignation, figured by Ortea et ai,

1996, fig. 109): specimen.

flavomarginata. Hypselodoris flavomarginatu

Rudni.m, 1995: 31-34, figs 6H, 25A, 27-29,

Type locality: Grand Coude, Bon rail, New

Caledonia. HOLOTYPE (by original désigna¬

tion): specimen, Icg. Faucomprc + 1 paratype.

fontandraui. Glossodoris fontandraui Pruvot-Fol,

1951: 24, 25, fig. 11, pl. 1, figs 4, 5, 26. Type

locality: Blancs, .Spam. NEOTYPE (designated

by Ortea et ni. 1996: 65): specimen. Remarks:

Ortea et ni (1996) placed this species in the

genus Hypselodoris Srimpson, 1855.

jrancoesii. Ccrntosoma frttncoesii de Rochebrune,

1894: 55. Type locality: Nouméa, New

Caledonia. SYNTYPE: 1 specimen, leg.

François. Remarks: According to Rudman

(1988), this is a junior synonym of Cemtosoma

tenue Abraham. 1876.

francoisae. Chromodoris francoisae Bouchet in

Bouchet &t Ortea 1980: 123-125, figs 11, 12,

14. Type locality: “Le Virage”, between Ngor

and Yof (Sénégal). HOLOTYPE (by original

désignation): specimen t- 1 paratype.

Remarks: Ortea et ni (1996) placed this spe¬

cies in the genus Mexicbromis Bertsch, 1977.

fi-ancoisi. Risbecia fmncoisi Odhncr, 1934: 248,

249. Type locality: Baie de l'Orphelinat,

Nouméa, New Caledonia. SYNTYPE: 1 speci¬

men. Remarks: Odhner ( 1934) established the

new genus Risbecia with Cemtosoma fmncoisi

de Rochebrune, 1894, in the sense of the tnisi-

dentificarion of Risbec ( 1928), as the type spe¬

cies. Under Article 70c (ICZN 1985). with

this nomenclatural act Odhner (1934) intro-

duced rbe new species Risbecia fmncoisi

Odhner, 1934, which is the type spccies ol the

genus Risbecia by original désignation. The

syntypes of this spccies are the specimens stu-

died by Risbec (1928).

gasconi. Hypselodoris gasconi Ortea et Valdés in

Ortea et al. 1996: 120-123, figs 12A, 90-92.

Type locality: Calvi, Corsica (Mediterranean

Sea). HOLOTYPE (by original désignation):

specimen.

geometrica. Chromodoris geometrica Risbec,

1928: 148-151. fig. 41, pl. 6, fig. 10. Type

locality: Anse Vata and Rocher à la Voile,

Nouméa, New Caledonia. SYNTYPE; I speci¬

men.

gibbosum. Cemtosoma gibbosum de Rochebrune,

1894: 55. Type locality: “Dead Joiand", Torres

Srrait, Northern Australia. SYNTYPE: I speci¬

men, leg. Lix. Remarks According to

Rudman (1988), this is a junior synonym of

Cemtosoma nilobatum (J. E. Gray, 1827).

gofasi. Hypselodoris gofasi Ortea et Valdés' in

Ortea et ai 1996: 123-129, figs 93-99. Type

locality: Santa Maria, Beitguela, Angola.

HOLOTYPE (by original désignation, figured

by Ortea et al 1996, fig. 93): specimen, leg.

Gofas.

goslineri. Chromodoris goslineri Ortea et Valdés

in Ortea et al. 1996: 143-146, figs 112-114.

Type locality: Villa do Porto, Santa Maria,

Azotes. HOLOTYPE (by original désignation,

figured by Ortea et al. 1996, fig. 112): speci¬

men (dissected), leg. Gofas.

hikuerensis. Rosodoris hikuerensis Pruvot-Fol,

1954: 23-27, fig. 9. Type locality: Hikueru,

Tuamotu (Pacific Océan). I IOLOTYPE (by

monotypy, figured by Pruvot-Fol 1954, fig. 9):

specimen (dissected). Remarks: This is the

type spccies ol the genus Rosodoris Pruvot-Fol,

1954, by original désignation. According to

Rudman (1986a), Rosodoris is a junior syno¬

nym ol Glossodoris Ehrenberg, 1831.

birsuta. Cadlinella hirsuta Rudman, 1995: 2-9,

figs 1-5, 6B, 7. Type locaiity: Baie du Prony,

New Caledonia. HOLOTYPE (by original

désignation): specimen, Icg. Laboure + 1 para-

type.

jousseaumei. Cemtosoma joussedumei de

Rochebrune, 1894: 55. Type locality: Red Sca.

SYNTYPES: 2 specimens (1 ol thetn dissec¬

ted). Icg. Botta. Remarks: According to

Rudman (1988), this is a junior synonym of

Cemtosoma tenue Abraham, 1876.

boumacensis. Hypselodoris koumacensis Rudman,

1995: 34-37, figs 6C, 24B, 30, 31. Type loca¬

lity: Grande Récif de Koumac, New

Caledonia. HOLOTYPE (by original designa-

ZOOSYSTEMA • 1998 -20(4)

723

Valdés Â. & Héros V.

tion, figured by Rudman 1995, figs 6C, 24B,

30, 31): specimen (dissected).

laboutei. Nouméa laboutei Rudman, 1986c:

399-401, figs 1 H, 15-17- Type locality: îlor

Gi, New Caledonia. HOLOTYPE (by original

désignation, figured by Rudman 1986c,

figs 1H, 15-17): specimen (dissected), leg.

Laboute.

lacteola. Hypselodoris lacteola Rudman, 1995:

28-30, figs 25B, 26. Type locality: east entrance

to Canal Woodin, New Caledonia. HOLOTY¬

PE (by original désignation, figured by Rudman

1995, fig. 25B): specimen, leg, Laboute.

lata. Chromodoris lata Risbec, 1928: 154-156,

fig. 44, pi. 8, fig. 7- Type locality: île Hugon

and îlot Maître (Nouméa), New Caledonia.

SYNTYPES: 4 specimens.

lemniscata. Doris lemniscata Quoy et Gaimard,

1832: 268, 269, pi 19, figs 8-11. Type locali¬

ty: Port Louis, "île de France” (= Mauritius),

Indian Océan. SYNTYPE: 1 specimen, leg.

Quoy and Gaimard (dissected). Remarks: This

is thc type species of the genus Durvilkdoris

Rudman, 1984, by original désignation.

lineata. Doris lineata Souleyet, 1852:453, pl 25,

figs 5-9. Type locality: “îles Sandwich"

(= Hawaii). SYNTYPES: 5 specimens, leg.

Eydoux and Souleyet. Remarks: Eliot (1905)

placed this species in the génies Chromodoris

Aider et Hancock, 1855

lixi. Ceratosoma lixi de Rochebrune, 1894: 55.

Type locality: “Dead Joland", Torres Strait,

Northern Auscralia. SY"NTYPES: 5 specimens,

leg. Lix. Remarks: According to Rudman

(1988), this is a junior synonym of Ceratosoma

trilobaturn (J. E. Gray, 1827).

magnifica. Doris magnifica Quoy et Gaimard,

1832. 270-272, pl. 20, figs 1-4. Type locality:

New Guinea, Indoncsia. SYNTYPES: 2 spéci¬

mens-, leg. Quoy and Gaimard. Remarks: This

is the type species of the genus Chromodoris

Aider et Hancock, 1855, by monotype.

malacitana. Hypselodoris malacitana L.uque,

1986: 550-557, figs 42-44 Type locality: La

Hetradura, Màlaga, Spain. NEOTYPE (desi-

gnated by Ortea et al. 1996; 109): specimen.

molloi. Mexichroniis molloi Onea et Valdés in

Ortea étal. 1996: 152-155. figs 12F, 118-120.

Type locality: Isla Picuda, Mochima,

Venezuela. HOLOTYPE (by original désigna¬

tion. figured by Ortea et al. 1996, figs 12F,

118-120): specimen (dissected).

montrouzieri. Vhorunria montrvuzieri Rudman,

1995: 37-40, figs 6G, 32-34. Type locality:

Passe de Koumac, New Caledonia. HOLO¬

TYPE (by original désignation, figured by

Rudman 1995, figs 6G, 32A, 33, 34); speci¬

men (dissected).

muniaini. Hypselodoris muniaini Ortea et Valdés

in Ortea ét al. 1996: 136-139, figs 106-108.

Type locality: Santo Antonio, Principe, West

Africa. HOLOTYPE (by original désignation,

figured by Ortea et al. 1996, figs 106-108):

specimen (dissected), leg. Roldn and

Fernandes. Remarks: In the Leader of the ori¬

ginal description of this species tt is misspelled

as Hypselodoris muniani.

odhneri. Risbecia odhneri Risbec, 1953: 94. Type

locality: Baie de l’Orphelinat. Nouméa, New

Caledonia. SYNTYPE: 1 specimen. Remarks:

This species was introduccd by Risbec (1953)

for the specimens cited by Risbec (1928) under

the name “ Ceratosoma frdncoisi Rochebrune”.

These specimens arc also die type material of

the species Risbecia francoisi Odhner, 1934,

introduced under Article 70c (ICZN 1985).

Therefore, Risbecia odhneri and Risbecia fran¬

coisi are objective synonyms.

nmatissima. Cadlina ornatissima Risbec, 1928:

163-165, fig. 47, pl. 8, fig. 4. Type locality:

Baie de 1 Orphelinat, Nouméa, New

Caledonia. HOLOTYPE (by monotypy, figu¬

red by Risbec, 1928, fig. 47, pl. 8, fig. 4): spe¬

cimen (dissected). Remarks: This is the type

species of the genus Cad!niella Thiele, 1931,

by monotypy.

or sût ii. Doris orsinii Verany, 1846: 96, 100, 101.

Type locality: Sicily, ftaly. NEOTYPE (desi-

gnated by Ortea et al.. 1996: 58): specimen.

Remarks: Ortea et al. (1996) placed this spe¬

cies in the genus Hypselodoris Stimpson, 1855.

picta. Doris pic ta Sehultz in Philippi 1836: 105.

Type locality: Catania, Sicily, Italy. NEOTYPE

(designated by Ortea étal. 1996: 43): specimen.

Remarks: Ortea étal. (1996) placed this species

in the genus Hypselodoris Stimpson, 1855.

pullata. Glossodorh pidlata Rudman, 1995:

10-12, figs 6A, 8, 9, 10A. Type locality: Passe

724

ZOOSYSTEMA • 1998 - 20(4)

Opisthobranch mollusc type specimens in MNHN

d’Ouraï, near La Foa, New Caledonia.

HOLOTYPE (by original désignation, figured

by Rudman 1995, figs 6A, 8, 9, 10A): spéci¬

men (dissected), Icg. Faucompré + 1 paratype.

punicea. Hypsclodons punicea Rudman, 1995:

25-28, figs 6J, 22-24A. Type locality: Passe de

Koumac, New Caledonia. HOLOTYPE (by

original désignation, Figured by Rudman 1995,

figs 6J, 22-24A): specimen (dissected).

pustulosa. Dans pustnlosa Cuvier, 1804a: 452,

473. Type locality: "Mer des Indes” (= Indian

Océan). SYNTYPE (figured by Pruvot-Fol

1934a, fig. 2, pl. 2, figs 3-7): I specimen (dis¬

sected), leg. Péron. Remarks: Pruvot-Fol

(1934a) placed this species in the genus

Ceratosoma J. F.. Gray, 1850.

ransoni. Glossodoris ransoni Pruvot-Fol, 1954:

18-20, fig. 6. Type locality: Hikueru, Tuamotu

(Pacific Océan). HOLOTYPF. (by monotypy):

specimen (dissected). Remaries: According to

Rudman (1987), this is a junior synonym of

Chromodons kuniei Pruvot-Fol, 1930.

reticulata. Dons reticulata Quoy et Gaimard,

1832: 272, 273. pi- 20, figs 9-11. Type locali¬

ty: Tongatapu (Pacific Océan). SYNTYPE:

1 specimen, leg. Quoy and Gaimard.

Remarks: Risbec (1956) placed this species in

the genus Risbecia Odhner, 1934.

rhopalicum. Ceratosoma rhopalicum de

Rochebrunc, 1894: 55. Type locality: Red Sea.

SYNTYPE: 1 specimen, leg. Botta. Remarks:

According to Rudman (1988), this is a junior

synonym of Ceratosoma tenue Abraham, 1876.

rorneri. Nouméa romeri Risbec, 1928: 165-167,

fig. 48, pl. 5, fig. 5. Type locality: Nouméa,

New Caledonia. SYNTYPES: 2 specimens

(one of them dissected). Remarks: This is the

type species of the genus Nouméa Risbec,

1928, by subséquent désignation by Baba

(1937).

symmetricHS. Glossodoris symmetricus Rudman,

1990: 275-279. figs IC, 6D, 8A, 9D (in part),

10A-D, 11, 12. Type locality: St. Gilles Reef,

La Réunion (Indian Océan). HOLOTYPE (by

original désignation, figured by Rudman 1990,

figs IC, 6D, 8A, 10B, 11A-F): specimen (dis¬

sected) , Icg. Jay.

tricolor. Doris tricoLor Cantraine, 1835; 383,

384. Type locality: Accitrezza, Sicily

(Mediterranean Sea). NEOTYPE (designated

by Ortea et al. 1996. 32). specimen. Remarks:

Ortea et al. (1996) placed this species in the

genus Hypselodaris Stimpson, 1855.

trouil/ati. Ghrontodoris trouilloti Risbec. 1928:

146, 147, fig. 39, pl. 8, lig. 5. Type locality:

Pointe de l’Artillerie, Nouméa, New

Caledonia. SYNTYPES: 2 specimens (dissec¬

ted).

undulata. Glossodoris undubita Pruvot-Fol, 1954:

21-23, fig. 8. Type locality: Hikueru, Tuamotu

(Pacific Océan). SYNTYPES: 3 specimens

(dissected). Remarks: According to Rudman

(1986a). tins is a junior synonym of Glossodoris

si bogue (Rergh, 1905).

vercanifnrma. Nouméa verconiforma Rudman,

1995: 19-22, figs 61, 16-18. Type locality; NW

sidc ol Rccif de l’Infernet, New Caledonia.

HOLOTYPE (by original désignation, figured

by Rudman 1995, figs 61, 16-18): specimen

(dissected).

versicolor. Cbromodoris versimlor Risbec, 1928:

147, 148, fig. 40, pl. B, fig. 2, pl. 6, fig. 3. Type

locality: Baie de l'Orphelinat, Nouméa, New

Caledonia. SYNTYPE: 1 radula. Remarks:

Risbec (1953) placed this species in the genus

Risbeda Odhner, 1934.

villafranca. Doris viUajranca Risso, 1818: 370.

Type locality: Blancs, Spain. NEOTYPE (desi¬

gnated by Ortea et al. 1996: 22): specimen.

Remarks: Ortea et al. (1996) placed this spe¬

cies in the genus Hypsdodoris Stimpson, 1855.

xicoi, Hypsdodoris xicoi Ortea et Vailles in Ortea

et al. 129-133, figs 12C, D, 100-103.

type locality: Esprainha, Sâ'o Tomé. West

Africa. HOLOTYPE (by original désignation,

figured by Ortea et ai 1996, figs 12D, 100B,

101 B): specimen, leg. Roldn and Fernandes.

Family Df.NDRJÇOORIUI DAF O'Donoghue, 1924

angolensls. Dendrodoris angolensis Valdés et

Ortea in Valdés et al 1996; 20-23, figs 2F, 11,

1 2C, Type locality: Luanda, Angola. HOLO¬

TYPF (by original désignation): specimen, leg.

Gofas + 2 pararypes.

aurea. Doris aurea Quoy et Gaimard, 1832:

265, pl. 19, figs 4-7. type locality: fervis Bay,

Australia. SYNTYPES: 5 specimens (2 of

ZOOSYSTEMA • 1998 -20(4)

725

Valdés Â. & Héros V.

thein dissected). Remarks: Pruvot-Fol

(1934b) placed this species in the genus

Dendrodoris Ehrenberg, 1831.

commuais. Doridopsis commuais Risbec, 1928:

67-69, fig. 7, pl. A, fig. 2, pl. 1, fig. 6. Type

localiiy: Pointe de l'Artillerie, Nouméa, New

Caledonia. SYNTYPES: 2 specimens.

Remarks: According vo Brodie et al. (1997),

this is a junior synonym of Dendrodoris fiima-

ta (Riippell et I euckart, 1828).

fosseti. Doridopsis fosseti Risbec, 1928: 64, 63.

pl. 5, fig. 4. Type locality: Pointe de l’Artillerie,

Nomnca, New Caledonia. SYNTYPE (ligured

by Risbec 192S, pl. 5, lig. 4): 1 specimen.

grandiflora. Doris grandijlora Rapp, 1827: 520,

521, tig. 3. Type locality: Strait of Gibraltar,

36°09’N - 06°09’W, Spain. NEOTYPE (desi-

gnated by Valdés et al. 1996: 9): specimen.

Remarks: Doris grandiflora has been placed in

the Official List o( Spécifie Namcs in Zoology,

Opinion 1805 (ICZN 1995). Valdés et al.

(1996) placed this species in the genus

Dendrodoris Ehrenberg, 1831.

herytra. Dendrodoris herytra Valdés et Ortea in

Valdcs étal. 1996:25-27. figs 2G, H, 12B, 14.

Type locality: Madeira. HOIOTYPE (by ori¬

ginal désignation): specimen, leg. Wirtz.

limbaui. Doris limbata Cuvier. 1804a: 468. 469,

pl. 74, fig. 3. Type locality: Marseille, France.

NEOTYPE (designated by Valdés et al. 1996:

5): specimen. Remaries: Valdés et al. (1996)

placed this species in the genus Dendrodoris

Ehrenberg, 1831.

minima. Dendrodoris minima Pruvot-Fol, 1951:

47. Type locality: Muros de Na Ion, Asturias,

Spain. NEOTYPE (designated by Valdés &

Ortea 1997: 249): specimen, leg. Rodriguez.

Remarks; Valdés & Ortea (1997) designated

the sanie specimen as neorype of Doriopsilla

pelseneen d’Olivcira, 1895, and therefore both

namcs become objective synonyms.

mollis. Doridopsis mollis Risbec, J 928: 65, 66,

fig 6, pl 2, fig. 4. Type locality: New

Caledonia. SYNTYPES: 2 specimens.

Remarks: According to Brodie et al. (1997),

this is a junior synonym ol Dendrodoris nigra

(Stimpson, 1855).

nigropunctatu. Doriopsis nigropunctata Vayssière,

1912: 77, 78, pl. 7, fig. 109. Type locality:

Djibouti. HOLOTYPE (by monotypy, figured

by Vayssière 1912, pl. 7, fig. 109): specimen

(dissected), leg. Gravier. Remarks: I bis species

probably must be placed in the genus

Dendrodoris Ehrenberg, 1831.

pelseneeri. Doriopsilla pelseneeri d'OIiveira,

1895: 12, 13. Type locality: Muros de Nalon,

Spain. NEOTYPL (designated by Valdés &

Ortea 1997: 249): specimen, leg. Rodriguez.

punctata . Doris punctata Quoy et Gaimard,

1832: 262, pl. 18. figs 8-10. Type locality: “Le

Havre Carreret”, New freland, Bismarck

Archipelago. SYNTYPE: I specimen, leg.

Quoy and Gaimard. Remarks: This naine is

preoccupied by Doris punctata Rüppell et

I.euckart, 1828 (see rufopunetuta). Pruvot-Fol

(1934b) placed this species in the genus

Dendrodoris Ehrenberg, 1831.

racemo.su. Dendrodoris racemosa Pruvot-Fol,

1951: 47. Type locality: Muros de Nalon,

Spain. NEOTYPE (designated by Valdés &

Ortea 1997: 249): specimen, leg. Rodriguez.

Remarks: Valdés & Ortea ( 1997) designated

the sanie specimen as neorype of Doriopsilla

pelseneeri d’OIiveira, 1895, and therefore both

liâmes become objective synonyms.

rosea. Doriopsis rosea Vayssière, 1912: 82, 83,

pl. 1, fig. 2, pl. 10. fig. 153. Type localiiy: near

Obock, Golfe de Tadjoura, Djibouti. SYN¬

TYPES (one figured by Vayssière 1912, pl 1,

fig. 2, pl. 10, fig. 153): I specimen (dissected),

leg. Jousseaume; I specimen (dissected) leg.

Gravier. Remarks: Plie name Doriopsis rosea

was firstly introduced by Vayssière (1911)

without description (nornen nudurri).

According to Brodie et al. (1997). this is a

junior synonym of Dendrodoris fit mata

(Rüppell et Leuckart, 1828).

rufopunctata. Dendrodoris rufopunctata Pruvot-

Fol, 1934b: 60, 61. Type locality: “Le Havre

Carreret’, New Ireland, Bismarck Archipelago.

SYNTYPE: 1 specimen, leg. Quoy and

Gaimard. Retnarks: This is a replacement

name for Doris punctata Quoy et Gaimard,

1832, preoccupied by Doris punctata Rüppell

et Leuckart, 1828.

senegalensis. Dendrodoris senegalcnsis Bouchet,

1975a: 124-127, ligs 3, 4, pl. 1, fig. 2. Type

locality: “Le Virage” between Ngor and Yofi

726

ZOOSYSTEMA • 1998 - 20(4)

Opisthobranch mollusc type specimens in MNHN

Cap Vert Peninsula, Sénégal. HOLOTYPE (by

original ilesignarion): specimen + 2 paratypes.

Remarks: Bouchet (1975a), originallv desi-

gnated a “Lectotype” for this species.. However,

following Article 73a (i) (ICZN 1985), if an

author when cstabli.shing a new nominal spe¬

cies States thaï une specimen is “the type”, or

uses some équivalent expression, that specimen

is the holorype by original désignation.

tuberculosa. Doris tuberculosa Quoy et Gaimard,

1832: 248, 249, pl. 16. figs 1,2. Type locality:

“Port Dorey” (= Manokwari), Inan Jaya,

Indonesia. SYNTYPE: I specimen (dissected).

Remarks: Pruvot-Fol (1934b) placed this spe¬

cies in the genus Dmdrodoris Ehrenberg, 1831.

Family PHYl.LimiUAK Rafinesquc, 1814

albonigra. Phyllidia albonigra Quoy et Gaimard,

1832: 291, 292, pl, 21, figs 26, 27. Type loca¬

lity: Tongatapu (Pacific Océan). LECTOTYPE

(selected by Brunckhorst 1993, figured by

Quoy & Gaimard 1832, pl. 21, figs 26, 27):

specimen. Remaries: Accordmg to Brunckhorst

(1993), this is a junior synonym of Bbyllidiella

pustulosa (Cuvier, 18(14).

aurata. Phyllidia au rata Pruvot-Fol, 1952:

408-411, figs 1-9. Type locality: Cannes,

France. HOLOTYPE (by monotvpy, figured

by Pruvot-Fol 1952, figs 1-9): specimen (dis¬

sected). Rcmarks: According to

Brunckhorst & Wïlkin (1989), this is a junior

synonym o (Phÿllidin fUtva A ta d as, 1847.

bayi. Fryeria bayi Bouchet, 1983: 65-68, figs 1,

2. Type locality: Puma Bianca, Corsica

(Mediterrànean Sea). HOLOTYPE (by origi¬

nal désignation, figured by Bouchet 1983,

fig. 1): specimen ■» 1 pararype.

berghi. Phyllidiopsis berghi Vayssière, 1902:

237-242,'pl. 9, figs 10-16, pl 10, fig. 17. Type

locality: Talislnan (1883) Expédition, stn 141,

45°59.00’N - 04°09.46'W, Bay ôf Biscay,

1480 m. HOLOTYPE (by monorypy, figured

by Vayssière 1902, pis 9, 10 and Valdés &

Ortea 1996, fig. 1 B); specimen (dissected).

borbonica. Phÿllidin borbonica unavailable.

Remarks: Brunckhorst (1993) studied one

specimen deposited in MNHN labelled

“Phÿllidin borbonica Cuv.”, and pointed out

that it is a large specimen of P. varicosa . Ar the

sainte tinte, he indicated that the species

Phÿllidin borbonica was inrroduced by Cuvier

( 1804c), and listed this name in the synonymy

ol P. varicosa. However, in that paper Cuvier

dnl not introduce any new species, but just

remarked that he was the finit author who des-

cribed the genus Phyllidia based on a single

specimen collected from “lie Bourbon’ (= La

Réunion). Therefore, Brunckhorst (1993) is

the first author who introdttced the mamrs-

cript nante P. borbonica. but in synonymy, so

that it is unavailable (ICZN 1985, Article

1 le). For addirional information on this speci¬

men see also the remarks on Phyllidia trilinea-

ta Cuvier, 1804 and Phyllidia varicosa

Lamarck, 1801.

bouebeti. Phyllidiopsis boucheti Valdés et Ortea,

1996: 5, 6, figs 1 D, 5 (in part), 6. Type locali-

tv: Funta de la Rasca, Tènerile, Canary Islands.

HOLOTYPE (by original désignation, figured

by Valdés tSi Ortea 1996, fig. 1D): specimen +

I pararype.

catena. Phyllidia catena Pruvot-Fol, 1956b:

70-72, figs 6, 7. Type locality: Mayotte and

Mauritius (Indian Océan). SYNTYPES [one

figured by Pruvof-Eol 1956b, figs 6 (in part),

7]: 2 specimens, leg. Mathieu; 1 specimen (dis¬

sected), leg. Cloué. Remarks: According to

Brunckhorst (1993), this is a junior synonym

of PbyllitiieUa zeylanka (kelaart, 1859).

dautzenbergi. Phyllidia dautzenbergi Vayssière,

1912: 85-87, pi, 1, figs 14, 15. Type locality:

“Nord d Ambouli , Golfe de ladjoura,

Djibouti. HOLOTYPE (by monotvpy, figured

by Vayssière 1912, pl. 1, figs 14, 15): .speci¬

men, leg. Gravier. Remarks: The name

Phyllidia dautzenbergi was firstly inrroduced by

Vayssière (1911) without description {nomeri

nciditm). Brunckhorst (1993) placed this spe¬

cies in the genus Phyllidiopsis Bergh, 1875.

flava . Phyllidia /lava Aradas, 1847: 121. Type

locality: XJendi, Gozo, Malta. NEOTYPE

(designated by Brunckhorst S: Willan 1989:

209, figured by Brunckhorst & Willan 1989,

fig. 1 ): specimen + photo.

gofasi. Peticulidia gofasi Valdés et Ortea, 1996: 7,

8, figs 1E 4C, 5 (in part), 8. Type loca¬

lity: Seamount 1 Expédition, stn DW61,

ZOOSYSTEMA • 1998 - 20(4)

727

Valdés Â. & Héros V.

36°40.02’N - l4 o 16.00'W, Joséphine Bank,

NorrJi Eastern Adantic, 200-205 m.

HOLOTYPE (by original désignation, figured

by Valdés & Orrea 1996, fig. 1F): specimen +

2 paratypes.

gyneitopla. Phyllidiopsis gynenopla Boucher,

1977: 50-53, figs 18, 19, pl. 2, fig. E. Type

locality: Biaçore. r Fxpedition, stn 159,

37°26'N - 25 t> 5TW, oll Santa Maria, Azotes,

525-600 m. HOLOTYPE (by original dési¬

gnation, figured by Bouchet 1977, figs 18, 19,

pl. 2, fig. E and Valdés &t Ortea 1996, fig. 1C):

specimen (di.ssecied). Remarks: According to

Valdés 6c Ortea (1996), thi.s is a junior syno-

nym of Phyllidiopsis bergh i Vayssière, 1902.

honloni. Pbyllidia hantant Risbec, 1956: 22, 23,

pl. 14, figs 71-75, pL 15, figs 79-81. Type loca¬

lity: Hon Lon, Nha Trang, Vietnam.

LECTOTYPE («elected by Brunckhorst,

1993): specimen 4 3 paralectotypes. Remarks:

According to Brunckhorst (1993), this is a

junior synonym of Phyllidia varicosa Lamarck,

1801.

krempfi. Phyllidiopsis krempfi Pruvot-Fol, 1957:

120, 121, figs 41-49, pl. 1, figs 7, 8. Type loca¬

lity: Nha Trang, Vietnam. HOLOTYPE (by

monotypy, figured by Pruvot-Fol 1957,

figs 41 -49, pl- 1, figs 7, S): specimen.

ttigra. Phyllidia nigm van Hasselt, 1824: 244.

Type locality: Cu Lao I lon “île Poulo Cecir de

Mer", Vietnam. NEOTYPE (designated by

Brunckhorst, 1993: 55): specimen. Remarks:

Brunckhorst (1993) placed this species in the

genus Phyllidiella Bergh, 1869.

ocellata. Phyllidia ocellata Cuvier, 1804b: 269,

pl. 18, fig. 7. Type locality: Timor, Indonesia.

LECTOTYPE (selected by Brunckhorst 1993,

figured by Pruvot-Fol 1956b, fig. 2): specimen

(dissected), leg. Pérou and Lcsueur.

pulitzeri. Phyllidia pulitzeri Pruvot-Fol, 1963:

566-569, figs 1-6, pl. 1. Type locality:

Portofino, Genova, Italy. HOLOTYPE (by ori¬

ginal désignation, figured by Pruvot-Fol 1963,

figs 1-6, pl. I): specimen, leg. Pulitzer.

NEOTYPE (designated by Wagele, 1985: 65,

figured by Brunckhorst & Willan 1989, fig. 1):

specimen + photo front Xlendi, Gozo, Malta.

Remarks: Wagele (1985) designated a neotype

of this species because at that time the holoty-

pe was considered lost. Latcr, Brunckhorst &

Willan (1989) selected the same specimen as

neotype of Phyllidia flapa Aradas, 1847, to fix

the synonymy between both nominal species.

1 lowever. we hâve rcdiscovered the holotype of

Phyllidia pulitzeri in MNHN, and therefore

this case should be STibmitted to the

Commission (1CZN 1985: Article 75h).

pustulosa. Phyllidia pus tu lu s a Cuvier, 1804b:

268, pl. 18, fig. 8, lype locality: Timor,

Indonesia. LECTOTYPE (selected by

Brunckhorst 1993, figured by Cuvier 1804b,

pl. A, fig. 8 and Brunckhorst 1993. fig. 27A):

specimen, leg. Pérou and Lesueur. Remarks:

This is the type species of the genus Phyllidiella

Bergh, 1869, by subséquent désignation by

Brunckhorst (1993)

serenei. Phyllidia screuct Risbec, 1956: 24, 25,

pl. 16, figs 82-84, pl. 17, figs 86-89. Type loca¬

lity: “île Poulo Cecir de Mer" (= Cu Lao Hon),

Vietnam. l.F.CTOTYPFl (selected by

Brunckhorst, 1993): specimen + 3 paralecto¬

types. Remarks: Brunckhorst (1993) designated

the lectotype of Phyllidia serenei as neotype of

Phyllidiella nigra (van Hasselt, 1824), and there¬

fore both names become objective synonyms.

trilineata. Phyllidia trtlineata Cuvier, 1804b:

268, pl. 18, figs 1-6. Type locality: La

Réunion, Indian Océan SYNTYPE (figured

by Cuvier 1804b, pl 18, figs 1-4): 1 specimen

(dissected) (Fig. 10B) NEOTYPE (designated

by Brunckhorst 1993: 27, figured by

Brunckhorst 1993, fig. 23): specimen, leg.

Drivas and Jay. Remarks: Cuvier (1797) esta-

blished the generic name Phyllidia for a single

specimen collected front "île Bourbon" (= La

Réunion), but did not name rhe species.

Lamarck (1801) erected the spécifie name

Phyllidia varicosa for the specimen seen by

Cuvier (1797). Cuvier (1804b) commented

that the species on which he himself based the

genus Phyllidia must be named Phyllidia trili¬

neata instead of P. varicosa because the former

name is more appropriare. At the saine urne,

he described and figured P trilineata based on

the original specimen (front La Réunion) and

rwo additiçmaJ individuals (considerably smal-

ler in size) collected by Pérou front “Mer des

Indes”. The examination of the specimen

728

ZOOSYSTEMA • 1998 - 20(4)

Opisthobranch mollusc type specimens in MNHN

Fig. 10, — A, Holotype of Phyllidia varicosa and syntype of

Phyllidia trilineala, 60.4 rnm; B. Photocopy of Cuvier’s (1804b,

pl. 18, fig. 1) drawlng ot Phyllidia trilineala.

which Brunckhorst considérée! the holotype of

the unavailable species P borbonica (see

remarks on Phyllidia borbonica) reveals that h

is one of the syntypes of P trilineata figured by

Cuvier (1804b, pl. 18, figs 1-4), see Figs 10A,

B. As remarked above, this specimen is labelled

“Phyllidia borbonica Cuv.", which unequivo-

cally indicates that it was collccted from La

Réunion. In addition, this specimen is very

large, so evidently it is the specimen on which

Cuvier (1797) based the genus Phyllidia (the

specimens collccted larer were considerably

smailer), and therefore the holotype of P vari¬

cosa. In the ancient catalogue of specimens in

alcohol of MNHN this specimen is registered

within the types of Phyllidia ocellata Cuvier,

1804 and Phyllidia pas tu/osa Cuvier, 1804,

both collccted from Timor by Péron and

Lessucur. The other syntypes of trilineata are

not registered and it is probable that they hâve

never been deposited in MNHN collections.

Brunckhorst (1993) selected a neotype for

P. varicosa and P. trilineata, but since the origi¬

nal type matcrial of this species has been redis-

covered, rhis case musr be submitted to the

Commission (ICZN 1985: Article 75h).

tuberçulata. Phyllidia tuberculata Risbec, 1928:

59, 60, fig. 3, pl. A, fig. 1, pl. 1, fig. 2. Type

locality: Baie de l’Orphelinat, Nouméa, New

Caledonia. SYNTYPE: I specimen (disseeted).

Rcmarks: According ro Brunckhorst (1993),

this is a junior synonym of PhylHdiopsis cardi-

tialis Bergh, 1875.

varicosa . Phyllidia varicosa Lamarck, 1801: 66.

Type locality: La Réunion, Indian Océan.

HOLOTYPE (by monotypy, figured by

Cuvier 1804b, pl. 18, figs 1-4): specimen (dis-

sected) (Fig. 10A). NEOTYPE (designated by

Brunckhorst 1993: 27, figured by Brunckhorst

1993, fig. 23): specimen, leg. Drivas and Jay.

Rcmarks: As indicated above (see remarks on

Phyllidia borbonica and Phyllidia trilineata),

the syntype ol P trilineata figured by Cuvier

(1804b, pl. 18, figs 1-4), is acrunlly in MNL1N

collections. This specimen labelled " Phyllidia

borbonica Cuv.” is the holotype by monotypy

of P. varicosa. Brunckhorst (1993) selected a

neotype for P. varicosa and P trilineata , but

since the original type materia) has been redis-

covered, rhis case must be submittçd to the

Commission (ICZN 1985: Article 75h).

lNCERTAE SED1S

alboranica. Doris (?) alboranica Bouchet, 1977:

29-34, figs 1, 2. Type locality: Polymède 2

Expédition, stn 66, 36°05N - 4°52W, Alboran

Sea, 910 m. HOLOTYPE (by original desi-

gnarion, figured by Bouchet 1977, figs I, 2):

specimen (disseeted).

rigida. Spongiodoris rigida Pruvot-Fol, 1933:

1.31 -133, pl. 2, figs 12-17. Type locality: Gulf

of Suez, Egypt. SYNTYPE: 1 specimen, leg.

Dollfus. Remarks: This is the type species of

the genus Spongiodoris Pruvot-Fol, 1933, by

monotypy.

NUDIBRANCHIA-DENDRONOTINA

Family TriTONIIDAE Lamarck, 1809

episcopalis. Tritonia episcopalis Bouchet, 1977:

ZOOSYSTEMA • 1998 ,20(4)

729

Valdés Â. & Héros V.

55-57, figs 22, 23, pl. 1, figs E, F, pl. 3, figs C,

D. Type locality: ïhalmsu (1973) Expédition,

stn Z409, 47 u 43N - 8°04W, off Brittany,

France, 1035-1OS0 m. HOLOTYÎ’E (by origi¬

nal désignation): specimen r 2 paratypes.

gravieri. Triamiopsis gravi fri Vayssière, 1912:

90-95, pl. 6. figs 78-80. Type locality: Golfe de

Tadjoura, Djibouti. HOI.OTYPF (by mono-

typy, figured by Vayssière 1912, pl. 6,

figs 78-86): specimen (dissected), lcg. Gravier.

Remarks: Tbe name Tritoniapsis gravieri was

firstly iritroduccd by Vayssière (1911) without

description (nnrnen nttdum).

hombergii. Tritonia hombergii Cuvier, 1802:

483-494, pis 1, 2. Type locality: Le Havre,

France SYNTYPES; 3 specimens, leg.

Homberg. Remarks: Tbis is rhe rype species of

the genus Tritonia Cuvier, 1797, by désigna¬

tion under plenary powers, Opinion 668

(ICZN 1963).

poirieri. Microlophus poirieri Mabille et de

Rochebrune in de Rochebrune & Mabille

1889: 1 1, 12, pl. 6, fig, 1. Type locality: "Baie

Orange”, Punta Arenas, Chile. SYNTYPE:

1 specimen. Remarks: This is the type species

of Microlophus Mabille et de Rochebrune,

1889, by monotypy.

Family DOTlDAEj. E. Gray, 1853

arteoi. Doto arteoi Ortea, 1978: 389-392,

figs A-D. Type locality: Conclut de Artedo,

Asturias, Spain. HOLOTYPE (by original

désignation): specimen.

cervicenigra. Doto cervicenigm Ortea et Bouchet,

1989: 265, 266, figs 5-7, 9. Type locality: La

Revellata, Calvi, Corsica (Mediterranean Sea).

HOLOTYPE (by original désignation, figured

by Ortea & Bouchet 1989, figs 5-7, 9): speci¬

men.

cindyneutes. Doto cindyneutes Bouchet, 1977:

57, 58, fig. 24. Type locality: Thalassa (1973)

Expédition, srn 7.435, 48“40N - 09"53W, off

Brittany, France, 1050 in. HOLOTYPE (by

original désignation): specimen + 1 paratype.

fluctifraga. Doto fluctifraga Ortea et Pérez-

Sânchez, 1982: 79-83, figs 1-4.Type locality: La

Garita, Cran Canaria, Canary Islands. HOLO¬

TYPE (by original désignation): specimen.

Jragaria. Doto fraparia Ortea et Bouchet, 1989:

262-264, figs 1-4. 8. Type locality: near La

Revellata, Calvi, Corsica (Mediterranean Sea).

HOLOTYPE (by original désignation, figured

by Ortea & Bouchet 1989, figs 2, 3, 8): speci¬

men + spawn + prey.

furvtt. Doto fuma Garcia-Gômez et Ortea. 1984:

208-211, figs 1, 2. pl. 1, fig. A. Type locality:

Tarifa, Spain. HOLOTYPE (by original dési¬

gnation): specimen.

racemnsa. Dota nteemosa Risbec, 1928: 269,270,

fig. 90, pl. 11, fig. 3. type locality: Rocher à la

Voile, Nouméa, New Caledonia. SYNTYPE:

1 specimen.

ussi. Doto ussi Ortea, 1982b: 1 -6, figs 1 -3. Type

locality: Mayotte, Comoros (Indian Océan).

HOLOTYPE (by original désignation): speci¬

men + photo + 1 paratype.

Family SCYLLAEIDAE Aider et Hancock, 1855

rosea. Melibe rosea Rang, 1829a: 130, pl. 3,

fig. 3. Type locality: Cape of Good Hope,

Southern Africa. SYNTYPE: 1 specimen, leg.

Rang.

NUDIBRANCHIA-ARMININA

Family ARMIN1DAE lredale et O’Donoghue, 1923

digueti. Armina digueti Pruvor-Fol, 1955:

464-466, figs 8-10. Type locality: Baja

California, Mexico. HOLOTYPE (by original

désignation, figured by Pruvot-Foi 1955, figs

8-10): specimen (dissected), leg. Diguet.

Family ZEPHYR!NIDAE lredale et

O'Donoghue, 1923

praeclara. Antiopella praeclara Bouchet, 1975a:

I27-L30, fig. 5, pl. I, fig. 3. Type locality: île

aux Serpents, near Dakar, Sénégal. HOLOTY¬

PE (by original désignation): specimen +

2 paratype$. Remarks: Bouchet (1975a) origi¬

nal])’ designated a 25 mm long specimen the

“Lectorype" for this species. I lowever, follo-

wing the Article 73a (i) (ICZN 1985), if an

author when establishing a new nominal spe-

730

ZOOSYSTEMA • 1998 • 20 (4)

Opisthobranch mollusc type specimens in MNHN

des States that one spedmen is “the type”, or

uses some équivalent expression, that spedmen

is the holotype by original désignation.

N UDI B RANCH IA-AEOLID INA

Family FlABEI.LlNIDAE Bergh, 1889

baetica. Flabellina baetica Garcia-Gômez, 1984:

61-64, pis 1, 2A. Type locality: Tarifa, Spain.

HOLOTYPE (by original désignation): sped¬

men.

barentsi. Coryphelbi barintsi Vayssière, 1913:

2-5, figs 1-6. Type locality: mouth of river

Rogacheva, Belush’ya Cuba Bay, SW coast of

Novaya Zemlya, Russia. HOLOTYPE (by

monotypy, figured by Vayssière 1913, figs 1-6):

specimen (dissected).

Family EüBRANC IIIDAI Odhner, 1934

arci. Eubrnnchus arci Ortea, 1980: 170-174,

figs 1, 2. Type locality: I’unta Hidalgo,

Tenerife, Cailary Islands. HOLOTYPE (by

original désignation): specimen.

Itnensis. Eubrançhus linensis Garcia-Gômez,

Cerverâ et Garcia, 1990: 585-591, figs 1-6, 11.

Type locality: Tarifa, Spain. HOLOTYPE (by

original désignation): specimen (dissected).

prietoi. Ettbraiickus prietoi Liera et Ortea, 1981 :

266-270, figs 1-3, pl. 1. Type locality:

Verdicio, Asturias, Spain. HOLOTYPE (by

original désignation, figured by Liera & Ortea

1981, pl. 1, figs D, E): spedmen.

Family AeOLIDIIDAE Gray, 1827

annulant. Eolidia annulant Quoy et Gaimaxd,

1832: 287, 288, pl. 21, figs 15-18. Type loca¬

lity: “Port Dorey” (= Manokwari), Irian Jaya,

Indonesia. SYNTYPE (figured by Quoy &

Gaimard 1833, pl. 21, figs 15-18): 1 spedmen.

Remarks: Pruvoc-Fot (1934b) placed this spe-

cies in the gcnus Cerberilla Bergh. 1873.

cryoporas. Baeolidia cryoparm Bouchet, 1977:

60-63, figs 26, 27. Type locality: Biogas III

Expédition, stn CV24, 47°33'N - 08\34’W,

Bay ot Biscay, 2110 m. HOLOT YPE (by ori¬

ginal désignation, figured by Bouchet 1977,

figs 26, 27): specimen (dissected).

paradoxa. Eolidina paradoxa Quatrefages, 1843:

31. Type locality: Saint-Vaast, France. SYN¬

TYPE: 1 spedmen.

quoyi. Baenlidui quoyi Pruvot-Fol, 1934b: 56,

fig. 19. Type locality: unknown. HOLOTYPE

(by monotypy): specimen (dissected), leg.

Quoy and Gaimard.

ransoni- Aenlidiopsis ratuoni Pruvot-Fol, 1956a:

228-231, figs l-l 1. Type locality: île Kaukura,

Tuamotu (Pacific Océan). SYNTYPE: 1 speci¬

men (dissected), leg. Ranson. Remarks: This is

the type species of the genus Aeolidiopsis

Pruvot-Fol, 1956, by original désignation.

Family Gl aucidaeJ. E. Gray, 1827

albti. Algaruiti alba Garcia-Gômez et Cervera,

1990; 734-740. figs 1-9. Type locality: Baia da

Baleeira, Sagres, Portugal. HOLOTYPE (by

original désignation), spedmen. Remarks:

This is the type species of the gcnus Atgarvia

Garcia-Gômez et Cervera, 1989, by original

désignation.

anadoni. Rioselleolis anadoni Ortea, 1979a:

132-138, pis 1-3. Type locality: Ribadesella,

Asturias, Spain. HOLOTYPE (by original dési¬

gnation): specimen. Remarks: This is the type

species of the genus Rioselleolh Ortea, 1979, by

original désignation, considered by Rolin et al.

(1991) as a junior synonym ot Bctbakina Roller,

1973.

bourailli. Aeolidia boit raidi Risbcc, 1928:

254-256, fig. 83, pl. 9, fig. 2. Type locality: île

Nou (Nouméa) and Bourail, New Calcdonia.

SYNTYPE: 1 specimen. Remarks: Rudman

(1980) placed this species in the genus

Phi d bina J. E. Gray, 1850.

cornuta. Aeolidia cornuta Risbec, 1928: 235,

236, fig. 71, pl. Il, fig. 5. Type locality: Pointe

de l’Artillerie (Nouméa) and Bourail, New

Caledonia. SYNTYPES: 3 specimens.

dangeri. Aeolidia daugert Risbec, 1928: 252-254,

fig. 82, pl. 9, fig. 1. Type locality: Rocher à la

Voile. Nouméa, New Caledonia, SYNTYPE:

1 specimen.

diffusa. Aeolidia diffusa Risbcc, 1928: 242-244,

fig. 75, pl. D, fig. 2, pl. 12, fig. 2. Type locali-

ZOOSYSTEMA • 1998 -20(4)

731

Valdés Â. & Héros V.

ty: New Caledonia. SYNTYPE: 1 specimen.

ducrosi. Aeolidia ducroù Risbec, 1928: 232, 233,

fîg. 69, pl. 11, fîg. 7, Type locality: Rocher à la

Voile, Nouméa, New Caledonia. SYNTYPE:

1 specimen.

joubint. Aeolidia joubini Risbec, 1928: 233-235,

fig. 70, pl. 10, fîg. 2. Type locaJiry: Nouméa,

New Caledonia. SYNTYPE: 1 specimen.

poindimiei. Aeolidia poindimiei Risbec, 1928:

246, 247, fig. 78, pl. 9, fîg. 3. Type localiry:

Poindimié and île N’Du Kué, New Caledonia.

SYNTYPE: 1 specimen. Reniarks: Rudman

(1991) placed this species in the genus

Pbyllodesmium Ehrenberg, 1831.

trunca. Aeolidia irunca Risbec, 1928: 236-238,

fig. 72, pl. 9, fig. 8. Type localiry: Rocher à la

Voile and Pointe de l'Ariillerie, Nouméa, New

Caledonia. SYNTYPE: 1 specimen.

violacea. Aeolidia vitüacea Risbec, 1928: 251,

252, fig. 81, pl. 11, fig. 2. Type locality: île

Nou and Rocher îi la Voile, Nouméa, New

Caledonia. SYNTYPE: 1 specimen.

vitreus. Favori nus vitreus Ortea, 1982c: 45-48,

figs 1. 2. Type locality: Los Cristianos,

Tenerife, Canary Islands. HOLOTYPE (by

original désignation): specimen + photo.

Family T HRGLI’KOIDAF. Bergh, 1889

francaisi. Guyvalvorui francaisi Vayssière, 1906a:

147. Type locality: “ÎJe Wandel" (= Renaud

Island), Atitarctica. HOLOTYPE (by monoty-

py): specimen (dissected). Remarks: This is

the type species of the genus Guyvalvoria

Vayssière, 1906, by monotypy.

Family FlONlDAL J. E. Gray, 1857

longicauda. Eotidia longicauda Quoy et

Gaimard, 1832: 288-290, pl. 21, figs 19, 20.

Type locality: Cook Strait, New Zealand.

SYNTYPES: 6 specimens. Remarks: This is

the type species of rhe genus Dolicbeolis Finlay,

1926. by monotypy.

Acknowledgements

This paper would hâve not been possible without

the invaluable help of Serge Gofas during the

inventory of the ty pe collection and élaboration

of the catalogue, his intimate knowledge of the

collection and of the literature was essential to the

completion of rhe work. Mr. Jean-Pierre Rocroi

helped in the hard ta.sk of fînding old papers in

the central library of MNHN. Philippe Bouchet,

Rudo von Cosel, Alan R. Rabat, Stefano Palazzi

and Anne-Marie Ohler reviewers made construc¬

tive commenrs on the manuscript. Raymond

Favia and Philippe Maestrati assisted during the

curatorial work. Mrs. Grasset (Centre Inter¬

universitaire de Microscopie Electronique,

Jussieu, Paris) produced the SEM micrographs.

Photographs hâve been Ljken by Pierre Lozouet

and printed by Christine Reynes (MNHN).

This project was made possible by two short-term

positions of rhe first author as visiting curator in

MNHN.

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ZOOSYSTEMA • 1998 -20(4)

735

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Submitted on 30 September 1997;

accepted on 22 April 1998.

738

ZOOSYSTEMA • 1998 -20(4)

Opisthobranch mollusc type specimens in MNHN

INDEX

Index of names including family and higher category taxa.

Page number in italics refer to illustrations.

Abbreviatjons

Aeo Aeolidina,

Arm Armina.

Aply Aplysiomorpha,

Ceph Cephalaspidea,

Den Dendronorina,

abyssicola, Mmnillacylichna - Cylichnidae (Ceph) . 7 00 , 704

acicula, Creseis- Cavoliniidae (Thec) ..... 707, 715

ACTBONIDAE ,... 698

admirabilis, Hingicula - Ringiculidae (Ceph) .698, 699

AEOLIDIIDAE . 731

AEOLIDINA . 731

aerugitwsa, Okssodoris - Chromodorididac (Nudi-Dor) .... 722

affinis, Hyttlaca - Cavnliniidae (Thec) . 707, 715

afriaina, Aeeras - Hamincidâe (Ceph) ..706, 709

AGIAJIDAË.706

AKER1DAË „. 711

itlba , Algarvia Claucidae (Nudi-Aeo) ..... 731

alba, Aplytie/la gmnirri - Aplysiidae (Aplv) .„.711

albojimbna, Punulledorts - Chromodorididae (Nudi-Dor) 722

albonigra, Pbyllidia - Phyllidiidae (Nudi-Dor) .727

albomùca, Paris (?) - Incvrut» sedis (Nudi-Dor) ................ 729

Al.DISIDAI- . 721

amboimi, Oscaninpsis - Pleurobranchidac (Nota) .71 4

anadom. Rioselltolis - Claucidae (Nudi-Aeo) .. 731

ANASPIDLA ..„.711

angoUnsis , Dntdrodori v - Deiidrodorididac (Nudi-Dor) .... 725

angulata, Hyalaea - Cavoliniidae (Thec) ... 715

angulosa, Qtvolim - Cavoliniidae (Tliec) .715

annulata, halidia - Acolidiidae (Nudi-Aeo) ... 731

antarctica, Anhidnm lulartytlnld — Arcliidundidae (Nudi-Dor) ....

APLUSTRIDAE .698

APLYSÏÏDAE .711

APLYSIOMORPI IA.„.711

arachis, Huila H.miineid.ie (Ceph) .706, 709

ARCHIDORIDIDAE.719

uni, Eubranthus - Lubranehidae (Nudi-Aeo) .731

ARMIN1DA1-: . ..„.730

ARM1NVNA .-.....730

arteoi, Potn - l lotnidae (Nudi-Den) . 730

asafera, Aplysia - Aplysiidae (Apiy) .. 712

asàtiui, Tmng/ 1 - Discodorididac (Nudi-Dor) ..719

nspera, Stauradoris — Dorididae (Nudi-Dor) .718

ASTERONOTIDAL . 721

astesana, Cttiàcria - ( ..ivolinridae ( Phec) . 707, 715

atromargwaia. Paris Chromodorididac (Nucli Dor) .722

auraut, Hhyllhiia Phyllidiidae (Nudi-Dor) . 727

aurai, Paris - Dendrndorididac (Nudi-Dor).725

auréola, (Utisstkloris- ChtOlîlodorididae*(Nudi-Dor) .......... 722

austnilis, de 1 moue Huila — Bulhdac (Ceph) ... 703, 708

australis, Quoy et C.iinitwd Huila - Bullidac (Ceph) .. 703, 708

azorica, Philme — Philinidae (Ceph) .706, 707

Dor Doridina,

Nudi Nudibranchia,

Nota Notaspidea,

Saco Sacoglossa,

Thec Thecosomata.

azoricus, Actaeo ii - Acteonidae (Ceph) ...698, 699

ihualladm, Geitodarh — Discodorididac (Nudi-Dor) .719

bacalbulai, Taringa - Discodorididac (Nudi-Dor) ...720

bdetica, ELibellitta - Elabeilinidae (Nudi-Aeo) .731

BAPTODORID1DAE .....719

rmrrnüi, Cttryphrlbt - 1-labellinidae (Nudi-Aeo) ... 731

bilyi, làyeritl - Phvllidiidae (Nudi-Dor) ....727

herghi, Phth/liupsu Phyllidiidae (Nudi-Dor) ...727

bii'incra, Hulin Aherid.it* (Aply) ...711

boHust. dénudons - Discodorididac (Nudi-Dor) .720

btnhtnma, l'hyltidia - Phyllidiidae (Nudi-Dor) .727

boruomca , l'rasina Juliidae (Saco) ... ~10 ,711

houe heu, (arvunudui i, - Raptodorididac (Nudi-Dor) .719

Imehtii, Cbrwiindoris - Chromodorididac (Nudi-Dor).722

bouchai, Phyltidiopsh Phyllidiidae (Nudi-Dor) ...727

bntiruil/i, I ,ulifbu - Claucidae (Nudi-Aeo) .731

huurgeolit. Uingtcida - Ringiculidae (Ceph) .— 698, 699

boujgaiguan, RinghruLi - Ringiculidae (Ceph) . 698. 699

bmiham . Aplysia - Aplysiidae (Aplv) .712

brtoii. Huila - Hamineidae (Ceph) ...706, 709

britoi, Chromodoris — Chromodorididae (Nudi-Dor) .722

brocki, OerrheUa - Plcurohrancbidae (Nom) ...714

BU1I.1DAE .708

calednniça , RijigiiuLt Riogiculidae (Ceph) .698,699

calédonien, Vayuiant - Vayssieridac (Nudi-Dor) .718

i aledonnum, Gmilôxrma - Chromodorididac (Nudi-Dor).. 722

ciwarintsh, Cyliclmina - Retusidae (Ceph) .702. 705

candiduk, Cyliehm - Retusidae (Ceph) .... 702, 705

rantahnra. Hypsebulorù Chromodorididae (Nudi-Dor) .... 722

capeniis i l'teurobramhueu - Pleurobraiuliidae (Nota) .714

carinata, Paris - Archidorididje tNudi-Dor) ...,, 719

carinata, idatydons - Plalydorididae (Nudi-Dor) . 721

tittemt, Phyllitlia - Phyllidiidae (Nudi-Dor) .727

CAVOl.rNlfDAI: .«715

mûris, Ibimdnru - Baplodoridldae (Nudi-Dor).719

CEPHALASPIDEA . 698

ccrmenipa, Dota Dovoidae (Nudi-L>en) .730

crutae, Jamb/a — Cyimrodorididae (Nudi-Dor) ... 718

ibaptalh. rlieadoni - Cavoliniidae (Thec) ... 707 ,715

ehaptalii, Clin — Cavoliniidae (Thec) . 707, 716

CHROMODORIDIDAE „„.,...-.722

limitait, Hypseltulorts Chromodorididae (Nudi-Dor) ...... 722

nnhneuta, Polo - Dotoidae (Nodi-Dcn) .730

rirrbijira. Aplysia - Aplysiidae (Aply) . 712

cïtrmus, l'ieurobranchus — Pleurobranchldae (Nota) .714

ZOOSYSTEMA • 1998 - 20(4)

739

Valdés Â. & Héros V.

dam, Creseis - Cavoliniidae (Ttieç).. 713, 716

clavOta, C-hromssdoris - Chromodorididac (Nudi-Dor) .722

coi, Chromodoris - Chroniodorididae (Nudi-Dor) ..722

commuais, Drindopsis - Dendrodorididue (Nudi-Dor).726

confina, Discwforis Discodoridîdae (Nudi-Dor) .720

comuia, Acalidia - Glaucidae (Nudi-Aeo) .731

commua, Pefio- Runcinidae (Ceph) ...710

crossei, Cy//Y/w«-Cylidinidac-(Ceph) .....700

crossei, Plesirubranchus - Pleurobranchiduc (Nota) .714

crosses, RingicuU Ringittllidac (Ceph) .. 698, 699

cruenta , Lions - Platydtirldidac (Nudi-Dor) .721

cryoporos , Hacolidiss - Acolidiidae (Nudi-Aeo) ..731

cumingi, Urnbrclla UmbiiLulidae (Nota) . 710, 714

CYLICHNIDAE .700

cymbalum, Bullit - Hamincidae (Ceph) .—.. 706, 709

CYMBUL.IIDAF..717

dactylomela, Aplysia- Aplyriidae (Aply).712

dangert, Aeolidia - Glaucidar (Nudi-Aeo) .731

darwini, Thecacera - Goniodorididae (Nudi-Dor)...717

dautzenbergi. Phyllidns - Phylhdiidac (Nudi-Dor) .727

decorata , Chwniodtu'u Chromodorididac (Nudi-Dor) ...... 722

delorti, Huila - Bullidac (Ceph) .... 708, 709

DF.NDROPORI DI DAF .725

DENDRONOTINA . 729

desgenettu , Huila - Rctusiduc (Ceph) .702, 705

diffusa, Aeolidia Glaucidae (Nudi-Aeo)... 731

digueli, Anmna - Arminidàe (Nudi-Arm) .730

digueti, Pletirobranclno — Pleurobranchidac (Nota) .714

dilatata, Rerusa - Rctusidac (Ceph) .... 702, 705

D1SCODORIDIDAF. .719

dollfrssi , Olossodorss — Chroniodorididae (Nudi-Dor) .722

DORI DI DAF . 718

DORID1NA .. 717

DOT1DAE . 730

ducrosi, Acolidùi - Glatiddae (Nudi-Aeo) .732

ecaudata, Aplysia - Aplysiidae (Aply) .... 712

edwardsi, Disctsdssris - Dîscodorididae (Nudi-Dor) .720

elegans, Accra • AJccridac (Aply) .....711 ,713

elegans, Bershcllttia - Juliidae (Saco) . 710, 711

elegans, Dori, - Chrontodondidae (Nudi-Dor) . 722

episcopalis, Priiania - Triiotiiidae (Nudi-Pcn) . 729

erytbraeensis, Disi'iidinir - Dtscodorididac (Nudi-Dor) ....720

espinosai, Hypstdndori, - Clmimadotididac (Nudi-Dor) ..723

F.UBRANC i HI DAF. .-.731

europaea, Raboassra - Cîymnodorididae (Nudi-Dor) .718

exirnta. Huila - Aplnsrridae (Ceph) . .. 698, 699

expleta , Aklisa Aldisidue (Nudi-Dor) .721

fana, Tarsnga - Discodorididae (Nudi-Dor) .720

FTONIDAF, .732

fischeri. Accrus — Hamineid.tr (Ceph) .... 708, 709

FLABELLLNI DAF .731

flava, Guyovia - Arehidorididae (Nudi-Dor) ..-.719

flava, l’hyllsdia - l'hyllidiid.ie (Nudi-Dor).727

flavosnargin.ua, Hypselodons — Chromodorididac (Nudi-Dor) ....

flucufrag/s. Dosa - Dotoidae (Nudi-Den)..730

fiusgosa, Haptodoris - Raptodorididae (Nudi-Dor) ..719

fontandrauu Glmodom - Chroniodorididae (Nudi-Dor) .... 723

fosseti, Doridopsis — Dendrodorididae (Nudi-Dor) .726

Jbisrierij, Brilla - Retusidae (Ceph) ...702, 705

frsigana, Dons Dotoidae (Nudi-Den) .730

frasicaisi, (liiyvalvnna - Tergipedidae (Nudi-Aeo) .732

fisuicoenl, C.rmfmina - Chromodondidae (Nudi-Dor) .723

Jnmcotsac, Cbrotntsdoris - Chromodorididac (Nudi-Dor) .... 723

fronçons, Risheeia- Chromodorididac (Nudi-Dor) —.723

fiilgurant Mmaiiwphrrus l'olyceridue (Nudi-Dor) ..718

fuma, Dotss - Dotoidae (Nudi-Den) ..-...730

g/llliCsh Huila - Haniineidae (Ceph) .708, 709

gylsconi, Hypiclod/iris - Chromodorididac (Nudi-Dor) .723

gaudryams, Rsrtgliula - Kingiculidac (Ceph) .698, 699

gcomctrica, Chromodoris - Chroniodorididae (Nudi-Dor) .... 723

gistrdi, Plesnohranchns - l’Ieurobranchidae (Nota) .715

gibbons, .Hyaltsea - Cavuliniidae ( I liée) . 713, 716

gibbnsssm, Ccratosonus - Chroniodorididae (Nudi-Dor) .723

gsrttrdi, Huila Reunidae (Ceph) .....702, 707

gtaiKn, MU Nnaragdincllidar (Ceph) .708

GLAUC1DAF.731

glabalma, Gioalina - Cavoliniidac l l'hec) . 713, 716

ghtbnJifSu, Hyatsea -Cavoliniidae (Thcc). 713, 716

gafasi, Hypselodons Chraniodorididau(Niidi-l)or) .723

gojasi. HtlicUlidU - l’hyllidiidae (Nudi-Dor) —.-,.727

GONIODORIDIDAE .-.717

gsnliricrs, Chmmodorh - Chroniodorididae (Nudi-Dor) .723

gmiidlfloM, Dons Dendrodorididae (Nudi-Dor).726

grrtnnLuissIma, Archtdoris - Arehidorididae (Nudi-Dor) .... 719

graviers, AplysielU - Aplysiidae (Aply) .. 712

graviers. Trsnsiinspiis - Tritoniidae (Nudi-Den) .730

gtuimerisb, HulUea — Aplustridae (Ceph) .698

gain, Plociimoplicnu - Polvceridae (Nudi-Dor) .. 718

GYMNODOR1D1DAF .718

gytscnnpla, Pbyllidtofijis~ Phyllîdiidae (Nudi-Dor) .728

HAMINFIDAl: .706

HFRMAH1DAF . 711

breytra, Pcssdrudoru - Dendrodorididae (Nudi-Dor).726

H FXA R RANCI Fil DA F ...718

hilmereilsis, Rosôdclns - Chroniodorididae (Nudi-Dor) .723

hirstila. (.sillstlciLi —Chroniodorididae (Nudi-Dor) .723

hmsndimmt. Huila Aglajidae (Ceph) . 706

bomber gu, TnhsnisS - Tritoniidae (Nudi-Den) ...730

hon/imi, Phyllidia - Phyllîdiidae (Nudi-Dor) . 728

HYDATINIDAE ...698

immondes, PUtydoris - Dorididae (Nudi-Dor) . 718

impenepnis, Meloscaphtotder - Cylichnidae (Ceph) .... 700, 704

usai. Aply,so - Aplysiidae ( Aply 1 .712

inflata. Cleodora - Cavoliniidae CI hcc) .716

injhsttun, RaDriuiiiii - Cavoliniidae (Tlïec) .716

inspemm, Ris lia - Cylichnidae (Ceph) .702

inversa. Paradons - Raptodorididae (Nudi-Dor) .719

joubini. Acalidia - Glaucidae (Nudi-Aeo) .732

joumaumei, Ceratosoma - Chromodorididac (Nudi-Dor) .. 723

julUria Aplysia Aplysiidae (Aply) ..712

JUU1DAF. . 711

KENTRODOR1DIDAH .721

kerassdressis, Aplysia - Aplysiidae (Aply) .. 712

kaumacensh, Hypscbsdorh - Chromodorididac (Nudi-Dor) 723

krempfr, Pbyllidsopsïs — Phyllidiidae (Nudi-Dor) . 728

740

ZOOSYSTEMA • 1998 -20(4)

Opisthobranch mollusc type specimens in MNHN

labo utei, Nouméa - Chromodorididae (Nudi-Dor) ..724

lacera, Doris- Hcxabranchidac (Nudi-Dor) .-.718

lacteola, JJyp.se/otlons - Chromodorididae (Nudi-Dor) .724

laminea , Phitydoris - Platydorididac (Nudi-Dor) ...721

lata, Chromotloris - Chrorinxlorididae (Nudi-Dor) .724

lemchei, Cyliriwa - CyJîcknid.ie (Ceph) ....702, 704

lemniscata . Dons— Chromodorididae (Nudi-Dor).724

lenlicula , Runcinn - Rnncimdae (Ceph) .710

lessonii, Aplysia - Aplysüdac (Aply) ...., H .. .712

levigata , HyaUea - Oivoliuiidao (Thcc).7/J, 717

limbata, Dorjs Dendrodorididae (Nudi-Dor) .726

limbaia, Hyolaea - Gàvrdiniûîae (Thec) .7/J, 717

lineata . Doris - Chromodorididae (Nudi-Dor) .................. 724

linensis, Eubnwchw - Eubranchidac (Nudi-Aeo) .731

lixi , Ceratosoma - Chromodorididae (Nudi-Dor) .724

lierai , Siiliger- Hermaeidae (Saco) . ... 711

hngtcauda, Aplysia - Aplvsiküe (Aply) .712

longicauda\ Èolidui - lionidae (Nudi-Aeo) .732

longicaudami, Clio - Cavoliniidae (Thcc) ...717

lophàtus j Thorybû[>uj - Dorididac (Nudi-Dor).718

lozatwi , Ercabini.j — Hermaeidae (Saco).......... 711

lutea, Arrimions mandata - Discodorididae (Nudi-Dor) .... 720

lutea, BuÜa - SmarHgdinellidae (Ceph) .. —708

mabiüei , 2fa/4x - Bullidae (Ceph) . 708, 709

macroden tien lata, Runcina — Runcinidae (Ceph) .710

maculata, Plalydoris - Platydorididae (Nudi-Dor) .721

maculatum, Pieuwbmnchidhsm - Plcurobranchidac (Nota) 71S

mügnijîcd , Doris - Chromodorididae* (Nudi-Dor) .724

maillardi , Dolabrifera — Aplvsiidae (Aply) ..712

malacitatia, Hypsriodons - Chromodorididae (Nudi-Dor) ..724

maltzam, Actueon Actconidac (Ceph) ..698, 699

matnillatus , Pltumbranrints — Pleurobranchidae (Nota) ...... 715

marebadi , Jonwnu — Kemrodorididae (Nudi-Dor) .721

manei , Ringiada — Ringiculidae (Ceph) ...698, 70/’

MIAMIRIDAE ......721

mïllepunctata , — Cylichnidae (Ceph) .. 702, 704

milncedwardsi . Philine - Philinidae (Ceph) .706, 797

minimOy Drndrodnns - Dendrodorididae (Nudi-Dor) .726

minutula , R ingrat la - Ringiculidae (Ceph) .698, 701

mirabilis, Tornatma Cylichnidae (Ceph) .702, 705

modesta . Bullidae (Ceph)... . 709 , 710

mollis. DoïîdopiU - Dendrodorididae (Nudi-Dor) .726

mollis , Paradons - Baptodorididae (Nudi-Dor) ».. 719

molloi, Mextchromis - Chromodorididae (Nudi-Dor) .724

mongii , Bulbt Hanuncidat (Ceph) ..794, 708

mouilifera, P/jJlirte— Philinidae (Ceph) .706, 707

momrouzim, l'horunm - Chromodorididae (Nudi-Dor) .. 724

morelletorutn, ActHOH - Aclcomdæ (Ceph).698

more lie tory nu Cyliriwa — Cylichnidae (Ceph) .702

muniaini, Hypselodoris - Chromodorididae (Nudi-Dor) .... 724

nayarita, Pdtodoris - Discodorididae (Nudi-Dor) .. 720

nbatrangemis, AUlisa -Aldistdae (Nudi-Dor).721

nigra, Phyllidia - Phyllidiidae (Nudi-Dnr) ...728

mgropunctuta , Doriùpsis - Dendrodorididae (Nudi-Dor) .... 726

NOTASP1 DEA .. 714

nouperda, Discodoris- Discodorididac (Nudi Dor) ...720

noumeae , PLitydons — Plarydorididae (Nudi-Dor) .721

noumeensis, Rmgkula Ringiculidae (Ceph) .. 698, 701

Hüdata, Aplysia Aplysüdae (Aply) .712

NUBIBRANCHIA ..717

oahouensis. Aplysia - Aplvsiidae (Aply).712

oblangdi Clio - CavoJintdae ..717

octlLua. Phyllidia - Phyllidiidae (Nudi-Dor) .728

udhneri , Risbeuct - Cchromodorididae (Nudi-Dor) .724

odanugbaei. An'bidons - Arehidorididae (Nudi-Dor) .719

uchlcrtùie. Ringiculn— Ringiculidae (Ceph) .698, 701

olriea, laringa - Discodorididac (Nudi-Dor) ... 720

ubvaefbn/iiy. Toniatinu - Cylichnidae (Ceph) . 702, 705

ONCH1DORID1DAF. .. 718

omtbemis Jorunna - Kentrodorididae (Nudi-Dor) .721

orbignii, Hyalra - Cavoliniidae ( 1 hec) .717

orbignyamt. Bulla — Hamineidac (Ceph) . 704, 708

oniiuissinni , Cadlina - Chromodondidae (Nudi-Dor) .724

orsifiii, Doris - Chromodorididae (Nudi-por) . 1 . n .. vlv ... l .... 724

ovata, Cymbidia - Cymbuliidae (Thec) ...717

avotdea , Buda - Hamineklae (Ceph) . 704, 708

OXYNOEIDAE . ...... 711

paagoumeneu Phlepnodnris - Archidorididae (Nudi-Dor) ..719

pantdoxu . Eoltdina- Aeolidiidae (Nudi-Aeo).731

passieri , Ringiatfa Ringiculidae (Ceph) ... 700, 701

paulucciae , Ringiettla - Ringiculidae (Ceph) .700, 701

peheneeri , Doriopsil/a - Dendrodorididae (Nudi-Dor) .726

pered\ Baptodorh - Baptodorididae (Nudi-Dor) 719

perfvssa. Geitodons - Discodorididac (Nudi-Dor) .720

perluct'ttis Trevdyano - Gymrtodorididae (Nudi-Dor) ........ 718

peronii, Plsurnbranrims Plcurubfyncludae (Nota) .715

perriniy Hammea - Hamineîdae {Ceph) .. 704, 708

perrieri. Pleurobramhns - Pleurobranchidae (Nota) .715

pctalijem , Aplysia - Aplyislidae (Aply) ..714

petiri, Cyfichna Cylichnidae (Ceph) . ..702, 705

PHILINIDAE.706

PHYLLIDIIDAE .727

prêta, Doris - Chromodorididae (Nudi-Dor) .. 724

prêta, Polycent - Polyccridae (Nudi-Dor) .718

pilota, Hüpkimia - Goniodorididac (Nudi-Dor) .. 717

prrulina, Rmgh ttLi - Ringiculidae (Ceph) . 700, 701

PLACOBRANCHIDAE . . 711

PLATYDÜRIDIDAE . 721

pleei, Aplysia - Aplvsiidae (Aply) ...... 714

PLEUROBRANCHIDAE . 714

pohidsmïri , Aeolîdia — Glaucidae (Nudi-Aeo) .732

poirierù Mrcrolopbus - Triconiidac (Nudi-Den) .“30

polyceniloides, BermudelLi - Gomodondidac (Nudi-Dor) ..717

PÔLYCRRIDAE ____ TTtt _...... 718

poiUelevicasii, Rnigicubi - Ringiculidae (Ceph) .700. 701

pmechra , AntwprlU - Xepltyritiidarc (Nudi-Arm) .730

pris toi, Etibranchus - Eubnuichidae (Nudi-Aeo) ..731

protea. Aplysia - Aplysitdæ (Aply) . 714

pulcbclla , Ringkula - Ringiculidae (Ceph) .700, 701

ptdirzeri, Phyllidia - l'hyllidiidae (Nudi-Dor) .... 728

pulùua , Gloswdurh Chroinodorîdidac (Nudi-Dor) .724

punctata, Dori' - Dendrodorididae (Nudi-Dor) _726

punicea , Hypselodoris - Chromodorididae (Nudi-Dor) .725

puiitLBift t Ibmot/na -«Cylichnidae (Ceph) .. 702, 705

pusrulosa, Doris - Chromodorididae (Nudi-Dor) ..725

pustulusa , Phyllidia - Phyllidiidae (Nudi-Dor) .728

quadriplicata, Ringuubt - Ringiculidae (Ceph) .700, 703

quin&a, Agluja - Aglajidae (Ceph) ...... 706

quoyi, Baeolidia - Aeolidiidae (Nudi-Aco) .731

ZOOSYSTEMA • 1998 -20 (4)

741

Valdés Â. 8t Héros V.

racemosa, Dendrodmis - Dei'uliudûruiid.u- (Nudi Dur) .726

racemosa, Dota - Dotoidae (Nudi-Den) .. .......... 730

rnngii, Hyalea - Cavolifùidae (Thcc) . 713, 717

ransum, Artslidvthfà — Acoliiiiniaé (NucpeAeo) .731

ransoni, Glosstsdpris - Chitvmodorididae (Nudi-Dor) ..725

reticulata, Dirrh - Chroinodorididac (Nudi-Dor) .725

reticukta, Onchidaris — Onel.idoriufdac (Nudi- Dor) .718

RETUSIDAE .!....!.1.702

rbopalicum , Ctmtlasoma — Chromodorididae (Nudi-Dor) .. 725

rigida, Spongiodoris - Incmse sedis (Nudi-Dor) .,.729

R1NGICUI IDAE .698

romeri, Nouméa - Chromodoiididae (Nudr-Dor) .. 725

rosea, Doriopsis - Dcndrodorididae (Nudi-Dor) .726

rosea, Mellhe - Scyllacidac (Nudi-Den) .73.0

rosi, Discodoris - Discodorididae (Nudi-Dor) .720

rositdae, Hingtntbt - Ringitulidae (Ceph.) .700

roussellae, JUngtcuhl - Ringitulidae (Ceph) .700

rubens, Discodtnn - f liscotio iidui.il- (Nudi-Dor) .720

rufa, Aplysia Aplysiidae (Aply) ... ... 714

rufopuncuthi, Dendmkris - Dendtodortdidae (Nudi-Dor) 726

rugosa, (mtruTi., - MiatTmtd.it (Nudi-Dor) ...................... 721

RUNCIN1DAE .710

SACOGLOSSA ...... 711

salleana, Ptngtiula - Ringitulidae (Ceph) ...700

sandwichiensis, Dons — Hexabranchidac (Nudi-Dor) .718

savignyi, Goniodoris Goniodorididae (Nudi-Dor) ... 717

savignyi, Ringtatht - Ringitulidae (Ceph) .700, 703

seabra, Dorh — Platydorididae (Nudi-Dor) .721

scaphandroidcs. Ro.xania - Cylichnidae (Ceph) . 702, 705

schembni. Dons - Afchtdorididae (Nudi-Dor) ..719

schlurnbergcri . Riugicuhi Ringitulidae (Ceph) .700, 703

schrammi, Tridachut Placobrandudae (Saco).711

SCY1.LAEIDAE ........730

secundo, Huila - Bullidne (Ceph) . 709, 710

senegalensis, Dmdrodorh - Dendrodorididac (Nudi-Dor) .. 726

senegalensis, Ringiatk - Ringitulidae (Ceph) 700, 703

strata, Phyllidia Phyllidlidac (Nudi-Dor) .728

seurati, Dnrtdium - Aglajidae (Ceph) ........,706

sieboldii, Lapkoccrcus - Oxynoeidae (Saco) ..711

simplex, Ihtlbt Rerusitlae (Ceph) . 706, 707

srmragdina, Aidiso - Aldisidae (Nudi-Dor) ,.... 722

SMARAC.DINEL1.1DAE .708

solea. Dons - Asieronoridac (Nudi-Dor) ..721

sordida, Dons - Discodorididae (Nudi-Dor) . 720

sordidata, Dum- Discodorittidae (Nudi-Dor)...720

souverbii, Lobiger- Oxynoeidae (Saco) ... 710, 711

spirata, Accra - Akcridar (Aply) .711 ,713

spongilla, Pbttydoris - Plarydorididae (Nudi-Dor) .721

stomascuta, Pbttydoris - Platydorididae (Nudi-Dor) .721

striata, Aplysia - Aplysiidae (Aply) . 714

striata, Crvsris Cavoliniidae (Thcc) .. 713, 717

suggnts, Tnve/yaiui - Cymnodtaididae (Nudi-Dor) ............ 718

symmetrietts, Gbssodons - Chromodorididae (Nudi-Dor) .. 725

tentrifemis, Cytichnim - Rctusidac (Ceph) .706, 707

termtidi, Aplysia Aplvsiidae (Aply) ..... 714

Tfc'RCIPEDIDAE .732

trnftnrm , RingtcuLt Ringitulidae (Ceph) .700, 703

TH ECOSOMATA . 715

tongttna, Aplysia - Aplvsiidac (Aply) .... 714

TClRNATINlDAF. . 700

tottmtmm . Ringiatlt- Ringitulidae (Ceph) ...700, 703

trkolor. Dons - Chromodorididae (Nudi-Dor) .725

trid/utila. On,bidons - Onchnlorididae (Nudi-Dor) .718

trilhmM , Phyüidia - l’hvilidiidae (Nudi -Dor) . 728, 729

TRITÛNUDAE ..'..729

triforquis, Taringa - Discodorididae (Nudi-Dor) .721

trimiiluti. Clmmtudiiris - Chromodorididae (Nudi-Dor) .... 725

trurtca, Aeulidia - Glaueidac (Nudi-Aen) .. 732

truneaiMi, Cylichna - Rctusidae (Ceph).706, 707

tubercubtta, Phyüidia— Phyllidlidac (Nudi-Dor) .729

tuberculosa, Doris Dendrodorididac (Nudi-Dor).727

UMBRACULIDAE . 714

utninuta, Hyaleti - i avoliniidae (Thec)... 713, 717

undu/ata, Glossodnris — Chromodorididae (Nudi-Dor) . 725

ungtiifera , Aplysia - Aplysiidae (Apiv) ..... 714

ussi, Dow — Doroidae (Nudi-Den) ....... 730

vagubunda, Tonutklln - Aerconidae (Ceph) .. 698, 699

t ant nsa, Phyltidia - Phyllidlidac (Nudi-Dor) ..729

vuyssùrrtti, Arcbuloris - Archidorididae (Nudi-Dor) ..719

tuyssierei, Plturobmnchaeu - Pleurobranchidae (Nota) .715

VÀYSSIEREIDAE .. 718

vercouifonmu Nouméa - Chromodorididae (Nudi-Dor) .... 725

verihsaiur, Chtvmodoris - Chrouiodurididac (Nudi-Dor) .... 725

prnicilaia, Plocidu Hcrm.icidjc (Sacn) .711

vilhijrancti, Doris - Cluomodmididat (Nudi-Dor) .725

ri!Irait. Huila - Cylichnidae (Ceph) ..... 702, 705

uiolaceu, Aeoliditl - GUuçidae (Nudi-Aen) .732

viobtera, Gtmiodorts ■ Gûiiindorididac (Nudi-Dor) .717

viridis , Huila - SmaragdinelJidae (Ceph) ..... 708

vitreus, Pavomtus - Glaucidae (Nudi-Aeo) .732

valu ta. Huila - Cylichnidae (Ceph) ... 702, 705

waldne, Cylichttium Hamineidac (Ceph) . 704, 708

xicoi, Hypselodoris- Chromodorididae (Nudi-Dor) .725

ZEPHYRINIDAE.730

742

ZOOSYSTEMA • 1998 - 20(4)

Remerciements/Acknowledgements

La Rédaction remercie vivement tous les rapporteurs, dont la liste est donnée ci-dessous, qui ont

accepté de donner temps et compétence pour l’évaluation des manuscrits publiés dans Zoosystema au

cours des années 1997 et 1998.

The Edi tors gratcfidly ücknowledge thc follouiing référées who provided their time and expertise in review-

ingpapers publhhed in Zoosystema during theyears 1997 and 1998.

Adam son M.

Vancouver

Canada

FIendler G.

Los Angeles

USA

Alcara L.

Madrid

Espagne

FIennache A.

Clères

France

AME74ANE N.

Paris

France

Holthuis L. B.

Leiden

Pays-Bas

Amiet j.-L.

Nyons

France

Hughes D. J.

Oban

Royaume Uni

Anderson R.

Guelph

Canada

Jones D.

Perth

Australie

Arnaud P. M.

Marseille

France

JUDSON M.

Paris

France

Baker J.

Cambridge

Royaume-Uni

Kabat A. R.

Washington

USA

Barre N.

Maison-Alton

France

Kensley B.

Washington

USA

Partie y D. M.

Rome

Italie

Kii.burn R. N.

Pietermaritzburg

Afrique du Sud

Beuwood O.

Townsvillc

Australie

Kinzelbacii R.

Rostock

Allemagne

Betsci I J. M.

Brunoy

France

Konishi K.

Sapporo

Japon

Beveridge I.

Victoria

Australie

Laubiër L.

Marseille

France

Blanc Ch. P.

Montpellier

France

Laurent R. F.

Tucuman

Argentine

Bock W.

New York

USA

Ledoyfh M.

Marseille

France

Bouchet Ph.

Paris

France

Lemajtrf. R.

Washington

USA

Bouix G.

Montpellier

France

LEvi C.

Paris

France

Bourgoin T.

Paris

France

Manning R. B.

Washington

USA

BourneW. R. P.

Aberdeen

Royaume-Uni

Marshau B.

Wellington

N.-Zélande

Boury-Esnault N.

Marseille

France

MaSSIN C.

Bruxelles

Belgique

BrescianiJ.

Charlottenlund

Danemark

Maisukuma A.

Fukuoka

Japan

Buckeridge J. S.

Auckland

N.-Zélande

MauchlineJ.

Oban

Royaume-Uni

CARU50 D.

Catania

Italie

MciAUGHI in P. A.

Washington

USA

Cassagneai) P.

Toulouse

France

MCIjAY C. L.

Christchurch

N.-Zélande

ChaCE F. A.

Washington

USA

Minelu A.

Padoue

Italie

Clark P. F.

London

Royaume-Uni

Moravec F.

C. Budcjovice

Rep. Tchèque

Cleva R.

Paris

France

NajtJ.

Paris

France

Cloudsley-ThompsonJ. L.

London

Royaume-Uni

Newman W. A.

La Jolla

USA

Combes P.

Montpellier

France

NG P. K. L.

Singapour

Conan C.

Saint-Denis

France

Ohi.kr A.

Paris

France

Condé B.

Nancy

France

Paueay G.

Mangilao

USA

Crosnier A.

Parcs

France

Pawson D.

Washington

USA

DavieP. J.F.

South Brisbane

Australie

PericartJ.

Montercau

France

Deharvenc L.

Toulouse

France

Poore G. C. B.

Victoria

Australie

D'HondiJ.-L.

Paris

France

Raven R. J.

South Brisbane

Australie

D'Udekem d’Acoz M. C.

Braine-l'Alleud

Belgique

Reiswig H. M.

Montréal

Canada

DUSSART B.

Le Bugue

France

Remaudiere G.

Paris

France

Fei.der F,

Lafayette

USA

RiceA. h,

Southampton

Royaume-Uni

ForestJ.

Paris

France

Roijard C,

Paris

France

Fransen C.

Leiden

Pays-Bas

Saint Girons H.

Paris

France

Gaui. B. S.

Fiaifa

Israël

Saint Laurent df. M.

Parcs

France

GascJ.-P.

Paris

France

Schëutr U.

Jârpâs

Suède

Gibson D. I.

London

Royaume-Uni

Seiichi I.

Tokyo

Japon

Gopas S.

Paris

France

Sissom D.

Canyon

USA

Guinot D.

Paris

France

Taiti S.

Florence

Italie

ZOOSYSTEMA • 1998 • 20 (4)

743

Remerciements/Acknowledgements

Tavares M.

Rio de Janeiro

Brésil

Williams A. B

Washington

USA

Vax ni Vei.uk I.

Bruxelles

Belgique

Young P. S.

Rio de Janeiro

Brésil

Van Hllsdingen P. J.

Leiden

Pays-Bas

Zapparoli M.

Viterbo

Italie

Vaucher C.

Genève

Suisse

Zevina G. B.

Moscou

Russie

VlLLEMANT C.

Paris

France

744

ZOOSYSTEMA

1998 • 20(4)

Instructions aux auteurs

La ligne éditoriale

Zoosystema est une revue consacrée à l'inventaire,

l'analyse er l'interprétation de la biodiversité anima¬

le. Elle publie des résultats originaux de recherches

en zoologie, particulièrement en systématique et

domaines associés : morphologie comparative,

fonctionnelle ci évolutive ; phylogénie ; biogéogra¬

phie ; taxinomie et nomenclature...

Un numéro de Zoosystema pourra être consacré à un

thème particulier sous la responsabilité d'un éditeur

invité.

Les auteurs devront suivre le Code International de

Nomenclature Zoologique. Il est recommandé que le

matériel-type soit, au moins en partie, déposé dans

les collections du MNHN.

Les manuscrits, dont le nombre de pages n'est pas

limité a priori , devront suivre rigoureusement les

recommandations aux auteurs (voir ci-dessous), et

seront adressés il la revue:

Service des Publications Scientifiques du Muséum

Zoosystema

57 rue Cuvier

F-75231 Paris cedex 05

Tel : (33) 01 40 79 34 38

Fax: (33) 01 40 79 38 58

e.mail : bulletin@mnhn.fr

Tout manuscrit non conforme à ces instructions

sera retourné pour mise au point. Chaque manus¬

crit est évalué par deux rapporteurs, ou plus.

Instructions aux auteurs

Chaque manuscrit soumis (y compris les illustra¬

tions) doit être présenté en trois exemplaires au for¬

mat A4, avec un double interligne et des marges

d’au moins 3 cm ; chaque page sera numérotée. Les

illustrations originales seront jointes au manuscrit

définitif, ainsi qu’une disquette 3.5” de format

Apple Macintosh ou compatible IBM (traitement

de texte Word de préférence), qui devra contenir

également les tableaux et éventuellement les illus¬

trations (Adobe Illustrator, Photoshop : Deneba

Canvas).

Le format

Les manuscrits, écrits en français ou en anglais, doi¬

vent être structurés comme suit :

- titre, si possible bref, en français et traduit en

anglais : un litre courant doit être proposé ;

- nom(s) et prénom(s) de(s) autcur(s) suivis de

leur(s) adresse(s) professionnellc(s), en indiquant si

possible le numéro de télécopie et l'adresse électro¬

nique ;

- résumés écrits en français cl en anglais (800 signes

au maximum chacun), suivis des mots clés er « key

words •• ;

- dans le texte courant, utiliser les italiques pour

tous les noms en latin : taxons de rangs générique

et spécifique (ex Pseudolaureola Kwon, Ferrara et

Tairi, 1992), étal , e,g. ... :

- dans le texte courant, les références aux auteurs

seront en minuscules, ex. Dupont (2001), Dupont

(2001, 2002), (Dupont 2001 ; Durand 2002),

(Dupont & Durand 2003, 2005), Dupont (2001 :

1), Dupont (2001, fig. 2) ;

- dans le texte courant, les références aux illustra¬

tions et aux tableaux de l’article seront présentées

ainsi : (Fig. 1), (Fig. 2A, D). (Fig. 2A-C), (Figs 3,

6), (Figs 3-5) ; (Tableau 1) ;

- les remerciements seront placés à la fin du texte,

avant les références bibliographiques ;

- les références bibliographiques doivent suivre les

exemples donnés ci-dessous ;

- indiquer dans la marge l'emplacement des illustra¬

tions ;

- donner les légendes des figures sur une feuille

séparée.

Les illustrations

Les illustrations au trait doivent être réalisées à

l’encre de Chine ou être fournies en impression

laser. Les photographies, bien contrastées, seront

sur fond noir ou blanc. Elles pourront être regrou¬

pées, et dans ce cas, identifiées par une lettre en

capitales (A, B, C. .). Les planches photogra¬

phiques, de préférence placées dans le corps de

l’article er non regroupées à la fin de celui-ci, doi¬

vent être traitées et numérotées comme des figures.

ZOOSYSTEMA • 1998 • 20(4)

745

Instructions aux auteurs

Les illustrations pourront être assemblées sur une

colonne (70 X 190 mnt) ou sur toute U largeur de la

justification (144 X 190 mm). Si possible, les

légendes (et lettrages) ne devraient pas figurer sur les

originaux. Ils seront disposes, alors, sur un calque

joint à chaque figure, la rédaction se chargeant de

les placer. Chaque figure doit compotier une échelle

métrique, sans aucun coefficient multiplicateur. Les

tableaux et graphiques, à inclure dans le manuscrit,

doivent pouvoir être imprimés sur une page et rester

lisibles après réduction éventuelle. Des photogra¬

phies en couleur pourront être publiées moyennant

une participation financière de ou des auteurs.

Références bibliographiques

Hoeg J. T. & Liitzen J. 1985. — Comparative

morphology and phylogeny of the family

Thompsoniidae (Cirripedia: Rhizoccphala:

Akenrrogonida) wirh description of ihree new

généra and seve-n new species. Zoologica Scripta

22: 363-386.

Rôckel D., Korn W. & Kohn A. J. 1995. —

Manuctl of the living Conidae, volume 1: lndo-

Paciflc région. Christa i lemmen, Wiesbaden,

517p.

Schwarier T. D. 1985. — Population structure of

black riger snakes, Nolechis ater niger, on offsho¬

re islands ol South Australia: 35-46, in Grigg G.,

Shine R. & Ehmann H. (eds), Biology of

Australasian Frogs and Reptiles. Surrey Beatty and

Sons, Sydney.

Épreuves et tirés à part

Les épreuves seront adressées à fauteur ou au pre¬

mier auteur (sauf indication contraire) et devront

être retournées corrigées sous huitaine. Les correc¬

tions, autres que celles imputables à la rédaction ou à

l’imprimeui, seront à la charge des auteurs. Le(s)

auteur(s) recevront gratuitement vingt-cinq tirés à

part pour les - articles jusqu'à cinquante pages (au-

delà, consulter la rédaction) ; les tirés à part supplé¬

mentaires seront à commander sur un formulaire

joint aux épreuves.

Soumettre un article pour publication dans

Zoosystema implique que celui-ci n’air pas été sou¬

mis dans une autre revue. Les droits de reproduc¬

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réservés à la revue. La reproduction de tout ou par¬

tie de l’article doit faire l'objet d'une demande écrite

préalable, adressée à la rédaction.

Scope of the Journal

Zoosystema is a journal devoted to the inventory,

analysis and interprétation of animal biodiversity. lr

puhlishes original results of zoological research,

particularly jn systemarics and relared fields; com¬

parative, functional and evolufionary morphology,

phylogeny, bîogeography, taxonomy and nomen¬

clature...

A complété issue of Zoosystema may be devoted to

several papers on a single topic tindcr the responsa¬

bilité of an invited ediror.

Papers should follow the International Code of

Zoological Nomenclature. We recommend that the

authors should deposit in the MNHN collections,

ar lcasr a part of the type matcrial.

Manuscripts, wilhout limitation ot the number of

pages, must conform strierly vvitli the instructions

to authors and will be sent to rhe Editor:

Service des Publications Scientifiques du Muséum

Zoosystema

57 rue Cuvier

F-75231 Paris cedex 05

Tel : (33) 01 40 79 34 38

Fax : (33) 01 40 79 38 58

e.mail: bullclin@mnhn.fr

Instructions to authors

Manuscripts must be submitred in triplieate in A4

format, double spaced, vvith margins of at least

3 cm and ail pages numbered. The original ligures

should be sent vvith the revjscd manuscript, as well

as a 3.5 diskei te Apple Macintosh or IBM-compa-

rible (Word, Word Perfra... ) format, which will

also contain tables and possibly figures (Adobe

lllustraror, Photoshop ; Deneba Canvas).

Format

Papers are to be written in simple and concise

Fccncli or English. They should be organized as fol-

lows:

- a brief tille in English and French;

- a Suggested running head;

- name(s) ofauthor(s), followed by their full profes-

sional address(es) and, if possible. Fax number and

e-mail;

- abstracts (in English and French) noi cxcceding

800 signs each, with key words and “mots clés";

- text with itaiieized words for Latin: taxa o! generic

and spécifie ranks (e.g. Pseudolaureola Kwon,

Fcrrara et Taiti, 1992), et al., ...;

746

ZOOSYSTEMA • 1998 • 20(4)

Instructions aux auteurs

- references to authors in main text should be pre-

sented as follnws: Smirh (2001), Smith (2001,

2002), (Smith 2001), (Smith 2001; Cary 2002),

(Smith & Cary 2003, 2005), Smith (2001: 1),

Smith (2001, fig. 2);

- references to illustrations and tables should be

indicated as follows: (Fig. 1), (Fig. 2A, D), (Fig.

2A-C), (Figs 3, 6), (Figs 3-5); (Table 1);

- keep acknowledgements short and place them at

the end of the text;

- give captions to illustrations on a separate sheet;

Illustrations

The éditorial board will pay spécial attention to the

quality and relevance of illustration.

Line drawings must be in Indian ink or high quality

laser printouts; high contrast photographs, placed

on white or black backgrounds, are required. These

can be grouped into Figures and their éléments and

identified by letters A, B, C... Plates are not placed

at the end of the article: they will be considered as

figures and numbered as such. Arrange Figures to fit

one or two columns (70 X 190 mm or 144 x

190 mm). No diagram or table is to exceed one

page. Letters, numbers, etc., for each figure, are to

be indicated on an accompanying overlay, not on

the original figure. A scalc bar is needed for each

figure (without magnification factor).

References

Hoeg J. T. & Lützen J. 1985. — Comparative

morphology and phylogeny of the family

Thompsoniidae (Cirripcdia: Rhizocephala:

Akentrogonida) with description of three new

généra and seven new spccics. Zoologica Scripta

22: 363-386.

Rockel D„ Korn W. & Kohn A. J. 1995. —

Manual of the living Conidae, volume 1: Indo-

Paciftc région. Christa Flemmen, Wiesbaden,

517 p.

Schwaner T. D. 1985. — Population structure of

black tiger snakes, Notechis ater tiiger, on offsho¬

re islands of South Australia: 35-46, in Grigg G.,

Shine R. & Ehmann H. (eds), Biology of

Australasian Frogs and Reptiles. Surrey Beatty and

Sons, Sydney.

Proofs and reprints

Proofs will be sent to the first author for correction

and must be returned within eight days by express

mail. Author(s) will receive twenty-fivc offprints

free of charge (for paper up to 50 pages; for paper

exceeding 50 pages, consult the editors); furrher

offprints can be ordered on a form supplied with

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The submission of a manuscript to Zoosystema

implies that the paper is not being olfered for

publication elsewhere. Copyright of published

paper, including illustrations, becomes the property

of the journal. Requests to reproduce material from

Zoosystema should be addresscd to the editor.

ZOOSYSTEMA • 1998 • 20(4)

747