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BULLETIN DU MUSÉUM NATIONAL D HISTOIRE NATURELLE

4' sér., 17, 1995, section A (Zoologie, Biologie et Écologie animales), n° 1-2,

SOMMAIRE — CONTENTS

Ph. Geniez & A. Foucard. — Un nouvel Acanthodactyle en Algérie : Acanthodactylus

taghitensis n. sp. (Reptilia, Sauria, Lacertidae). 3

A new Fringe-toed Lizard from Algeria: Acanthodactylus taghitensis n. sp. (Reptilia,

Sauria, Lacertidae).

P.-A. CROCHET, A. Dubois, A. Olher & H. Tunner. — Rana ( Pelophylax ) ridibunda

Pallas, 1771, Rana ( Pelophylax ) perezi Seoane, 1885 and their associated klepton

(Amphibia, Anura): morphological diagnoses and description of a new taxon. 11

Rana (Pelophylax) ridibunda Pallas, 1771, Rana (Pelophylax) perezi Seoane, 1885

et leur klepton associé (Amphibia, Anura) : diagnoses morphologiques et description

d’un nouveau taxon.

M. SEGONZAC & W. Vervoort. — First record of the genus Candelabrum (Cnidaria,

Hydrozoa, Athecata) from the Mid-Atlantic Ridge : a description of a new species

and a review of the genus. 31

Premières observations du genre Candelabrum (Cnidaria, Hydrozoa, Athecata) sur

des zones hydrothermales de la dorsale médio-atlantique : description d’une nouvelle

espèce et revue du genre.

B. Ben Slimane & M.-Cl. Durette-Desset. — Sur les Oswaldocruzia (Nematoda,

Trichostrongylina, Molineoidea), parasites d’amphibiens et de lézards de Cuba et de

Porto Rico . 65

On Oswaldocruzia spp (Nematoda, Trichostrongylina, Molineoidea), parasites of am¬

phibians and lizards from Cuba and Puerto Rico.

B. Ben Slimane, M. Verhaagh & M.-Cl. Durette-Desset. — Oswaldocruzia peruensis

n. sp. (Nematoda, Trichostrongylina, Molineoidea), parasite d’un nouvel Iguanidae

du Pérou . 77

Oswaldocruzia peruensis n. sp. (Nematoda, Trichostrongylina, Molineoidea), parasite

of an Iguanidae from Peru.

M. Judson. — Identity of the Antartic mite Gainia nivalis Trouessart (Acari, Nanor-

chestidae) . 83

Identité de l’acarien antarctique Gainia nivalis Trouessart (Acari, Nanorchestidae).

R. BOSMANS. — Description de Bordea, nouveau genre d’araignées des Pyrénées (Araneae,

Linyphiidae). 87

Description of Bordea, a new endemic spider genus from the Pyrénées (Araneae:

Linyphiidae).

W. Lourenço. — Description de trois nouveaux genres et quatre nouvelles espèces de

scorpions Buthidae de Madagascar. 95

Description of three new genera and four new species of Buthidae scorpions from

Madagascar.

A. Thiéry, J. Brtek & Ch. Gasc. — Cyst morphology of European Branchiopods

(Crustacea, Anostraca, Notostraca, Spinicaudata, Laevicaudata) . 107

Morphologie des œufs de Branchiopodes européens (Crustacea, Anostraca, Notos¬

traca, Spinicaudata, Laevicaudata).

A. G. Humes. — Three new species of Hemicyclops (Copepoda: Poecilostomatoida:

Clausidiidae) from northwestern Madagascar . 141

Trois nouvelles espèces d ’Hemicyclops (Copepoda: Poecilostomatoida: Clausidiidae)

du nord-ouest de Madagascar.

B. A. FOSTER & J. S. Buckeridge. — Barnacles (Cirripedia: Thoracica) of seas off the

Straits of Gibraltar. 163

Cirripèdes (Cirripedia: Thoracica) récoltés dans les earn du détroit de Gibraltar.

J. K. LOWRY & H. E. Stoddart. — A new species of Didymochelia from New Caledonia

(Crustacea: Amphipoda: Didymocheliidae) . 193

Une espèce nouvelle de Didymochelia de Nouvelle-Calédonie (Crustacea: Am¬

phipoda: Didymocheliidae).

P. J. F. Davie. — Two new species of Nanocassiope from the Western Pacific (Crustacea,

Brachyura, Xanthidae). 201

Deux nouvelles espèces de Nanocassiope de T Indo-Pacifique (Crustacea, Brachyura,

Xanthidae).

N. Ngoc-Ho. — Une espèce nouvelle de Neocallichirus des îles Tuamotu, Polynésie fran¬

çaise (Crustacea, Decapoda, Thalassinidea). 211

A new species of Neocallichirus from the Tuamotu Islands, French Polynesia

(Crustacea, Decapoda, Thalassinidea).

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 3-9.

Un nouvel Acanthodactyle en Algérie :

Acanthodactylus taghitensis n. sp. (Reptilia, Sauria, Lacertidae)

par Philippe GENIEZ et Antoine FOUCART

Résumé. — Un nouvel Acanthodactyle du groupe scutellatus découvert en Algérie, dans la région de Béni

Abbés, est décrit sous le nom de Acanthodactylus taghitensis, de Taghit, localité-type. Cette espèce se caractérise

surtout par la différence de taille entre les écailles dorsales et celles des flancs, par la possession de quatre

supralabiales en avant de la suboculaire et par un patron de coloration particulier.

Mots-clés. — Acanthodactylus taghitensis, Lacertidae, systématique, Sahara, Algérie.

A new Fringe-toed Lizard from Algeria : Acanthodactylus taghitensis n. sp.

(Reptilia, Sauria, Lacertidae)

Abstract. — A new Fringe-toed Lizard of the scutellatus group found in Algeria, in the Beni Abbés area,

is described under the name of Acanthodactylus taghitensis, from Taghit, type locality. This species is especially

characterized by the size difference between dorsal and lateral scales, by the possession of four supralabials in

front of the subocular and by an original colour pattern.

Keywords. — Acanthodactylus taghitensis, Lacertidae, systematics, Sahara, Algeria.

Ph. Geniez, Laboratoire de Biogéographie et Écologie des Vertébrés, EPHE, Université de Montpellier II, F- 34095 Montpellier

Cedex 05.

A. Foucart, CIRAD-PRIFAS, BP 5035, F- 34032 Montpellier.

INTRODUCTION

Au cours d’un trajet de liaison, lors d’une mission d’étude sur le Criquet pèlerin Schistocerca

gregaria (Forsskâl, 1775) programmée par le CIRAD-PRIFAS, l’un de nous (A. F.) a prélevé,

36 km après Taghit en direction de Béni Abbés, un Acanthodactyle répondant à des caractéris¬

tiques morphologiques (écaillure, coloration) qui, lorsqu’elles sont combinées, ne permettent d’at¬

tribuer cet animal à aucune espèce du genre Acanthodactylus décrite jusqu’alors. Par ailleurs,

un spécimen pouvant être rapporté à la même espèce a été photographié par M. Geniez, 5 km

après Taghit en direction de Béni Abbés. L’existence de deux animaux semblables et aussi par¬

ticuliers en regard des autres représentants du genre Acanthodactylus , provenant de deux localités

éloignées de quelque 30 km, suggère fortement, malgré la faiblesse de l’échantillon, que nous

avons affaire à une espèce nouvelle.

— 4 —

DESCRIPTION

Les deux Acanthodactyles de la région de Taghit, des femelles, appartiennent au groupe

scutellatus (sensu SALVADOR, 1982; ARNOLD, 1983): quatre rangées d’écailles autour des

doigts, écaille suboculaire ne touchant pas la lèvre, quatrième orteil fortement pectiné, écailles

ventrales disposées en séries transversales obliques, museau pointu, coloration pâle et peu

contrastée. Leur coloration générale est d’un beige roussâtre pâle ou orangé clair. Le dos

est parsemé de petites taches sombres (occupant chacune une à trois écailles) très espacées

et grossièrement disposées en cinq rangées longitudinales (fig. 1). Ces taches sont reliées

par des traits pâles qui s’estompent plus ou moins dans le tiers postérieur du dos. Les écailles

labiales et la suboculaire sont uniformément pâles. La face ventrale est blanc pur. En ce qui

concerne l’écaillure, on note 1 ou 2 granules insérés entre les pariétales, 1 rangée complète

de granules supraciliaires, se dédoublant en arrière de la troisième supra-oculaire, 4 labiales

en avant de la suboculaire, cette dernière étant en contact avec les labiales 4 et 5 (fig. 2).

Les écailles dorsales et latérales sont carénées et tectiformes, celles du dos étant environ

deux fois plus grandes que les latérales. Le passage des écailles du dos aux latérales s’effectue

brusquement (fig. 3). On compte 30 séries obliques transversales d'écailles ventrales dispo¬

sées en 14 rangées à mi-corps.

Fig. 1. — Vue dorsale de l’holotype d'Acanthodaetylus taghitensis n. sp.

Dorsal view of the holotype Acanthodactylus taghitensis n. sp.

— 5 —

Fig. 2. — Profil, côté droit, de la tête de l’holotype d’Acanthodactylus taghitensis n. sp.

Right head profile of the holotype of Acanthodactylus taghitensis n. sp.

FlG. 3. — Zone dorsolatérale de Fholotype d ’Acanthodactylus taghitensis n. sp., montrant le passage brusque des grandes écailles

du dos aux petites des flancs.

Dorsolateral view of the holotype of Acanthodactylus taghitensis n. sp. showing sharp demarcation between large scales on

back and smaller ones on flanks.

— 6 —

COMPARAISON AVEC LES AUTRES REPRÉSENTANTS

DU GROUPE SCUTELLATUS

U Acanthodactyle de Taghit présente des caractères communs avec d’autres taxons du groupe

scutellatus (tabl. I et II) : 4 écailles labiales en avant de la suboculaire (comme chez Acantho-

dactylus aureus ); taches dorsales sombres reliées longitudinalement par des traits clairs (comme

chez A. aureus)', faible nombre (44 et 47) de rangées d’écailles dorsales à mi-corps (comme

chez A. dumerili dumerili et certains A. aureus) ', faible nombre (21) de pores fémoraux (comme

chez A. d. dumerili, A. d. exiguus et A. longipes).

Il se distingue aisément des autres représentants du groupe scutellatus par les caractères

suivants (cf. tabl. I) : 4 labiales en avant de la suboculaire (fig. 2) (5 chez A. d. dumerili, A. d. exi¬

guus, A. scutellatus, 5 ou 6 chez A. longipes); faible nombre (44 et 47) de rangées d’écailles

dorsales à mi-corps (plus de 52 chez A. s. scutellatus, A. dumerili exiguus et A. longipes, ex¬

ceptionnellement de 46 à 49 chez A. scutellatus hardyi) ; écailles ventrales disposées obliquement

en 14 rangées longitudinales à mi-corps (12 perpendiculaires au corps de l’animal chez A. d.

dumerili, 16 à 18 obliques chez A. longipes) ; nombre de pores fémoraux (21) plus élevé que

chez A. d. dumerili (11 à 20); nombre de gulaires comptées longitudinalement (25) plus faible

que chez A. longipes (de 28 à 43).

Certains caractères morphologiques externes de l’Acanthodactyle de Taghit, lorsqu’ils sont

combinés deux à deux, permettent d’originaliser ce lézard sans ambiguïté. De plus, ce taxon

Tableau I. — Comparaison de Acanthodactylus taghitensis avec les autres Acanthodactyles à 4 rangées d’écailles autour des

doigts (d’après Arnold, 1983; 1986a; 19866; Bons & Girot, 1962; Salvador, 1982; Leviton & Anderson, 1967; obs.

pers.).

Taxons

Acanthodactylus taghitensis sp. nov.

44-47

A. aureus Günther, 1903

38-60

17-26

21-24

A. s. scutellatus (Audouin, 1809)

57-87

26-39

16-26

21-28

A. s. hardyi Haas, 1957

46-62

12-14

A. d. dumerili (Milne Edwards, 1829)

38-47

11-20

17-22

A. d. exiguus Lataste, 1885

53-69

16-25

15-21

5(6)

A. longipes Boulenger, 1918

60-97

16(18)

28-43

22-29

19-26

(5)6

A. cantons Günther, 1864

26-41

12-14

23-30

19-23

20-24

A. blanfordi Boulenger, 1918

30-50

12-14

28-35

14-22

20-23

A. schmidti Haas, 1957

105

32-54

13-18

17-23

21-25

A. arabicus Boulenger, 1918

27-37

14(16)

24-32

19-24

20-24

A. haasi Leviton & Anderson, 1967

36-44

12(14)

23-25

19-20

26-27

A. gongrorhynchatus Lev. & And., 1967

56-77

10-12

34-44

20-25

24-29

A. tilburyi Arnold, 1986

41-47

15-19

29-37

20-25

A. grandis Boulenger, 1909

103

39-64

L/C

14-18

22-35

15-24

19-25

4 ou 5

A. masirae Arnold, 1980

34-37

29-33

20-22

23-24

1 : Longueur tête-corps maximale (en mm). 2 : Nombre de rangées d’écailles dorsales à mi-corps. 3 : Écailles

dorsales carénées (C) ou lisses (L). 4 : Nombre de rangées longitudinales d’écailles ventrales à mi-corps. 5 :

Disposition des séries transversales d’écailles ventrales (O = oblique; P = perpendiculaire au corps de l’animal).

6 : Nombre d’écailles gulaires comptées dans le sens de la longueur. 7 : Nombre de pores fémoraux de chaque

côté. 8 : Nombre de lamelles sous le 4 e orteil. 9 : Nombre de supralabiales en avant de la suboculaire.

— 7 —

Tableau II. — Comparaison d'Acanthodactylus taghitensis avec les autres Acanthodactyles à 4 rangées d’écailles autour des

doigts (d’après Arnold, 1983; 1986a; 1986fc; Bons & Girot, 1962; Salvador, 1982; Leviton & Anderson, 1967;

obs. pers.).

Taxons

Remarques

Répartition géographique

Groupe

Acanthodactylus taghitensis

D 2 fois plus grandes que

Sahara algérien, région de Taghit

scutellatus

sp. nov.

A. aureus Günther, 1903

D ± de même taille que L

Région littorale atlantique du Sahara

scutellatus

A. s. scutellatus (Audouin,

D ± de même taille que L

Sahara, Palestine

scutellatus

1809)

A. s. hardyi Haas, 1957

D ± de même taille que L

Arabie Saoudite, Koweït, Irak

scutellatus

A. d. dumerili (Milne

D ± de même taille que L

Moitié sud de la Mauritanie,

scutellatus

Edwards, 1829)

A. d. exiguus Lataste, 1885

D ± de même taille que L

Sénégal

Tiers occidental du Sahara

scutellatus

A. longipes Boulenger. 1918

D + de même taille que L

Sahara

scutellatus

A. cantoris Günther, 1864

D beaucoup plus grandes

Afghanistan, N. de l’Inde, N.E. du

cantoris

A. blanfordi Boulenger, 1918

que L

D plus grandes que L

Pakistan

N. Oman, S. Iran, Pakistan,

cantons

A. schmidti Haas, 1957

D 2 fois plus petites que L

Afghanistan

Péninsule d’Arabie, Jordanie,

cantoris

A. arabicus Boulenger, 1918

D 2 fois plus grandes que L

S. W. Iran

Yémen du Sud, Yémen du Nord

cantons

A. haasi Leviton & Ander-

D plus grandes que L

Dharan, Sakaka (Arabie Saoudite),

cantons

son, 1967

A. gongrorhynchatus Lev. &

D un peu plus grandes

Oman

E. de la péninsule d’Arabie

cantons

And., 1967

A. tilburyi Arnold, 1986

que L

D ± de même taille que L

Riyadh, Al Jawf (Arabie Saoudite)

cantoris

A. grandis Boulenger, 1909

D ± de même taille que L

N. Arabie, Palestine, Irak, S. W.

grandis

A. masirae Arnold, 1980

D ± de même taille que L

Iran

Yémen du Sud, Oman

yemenicus

D = écailles dorsales ; L = écailles latérales.

Remarques : Toutes les espèces citées possèdent des écailles temporales carénées à l'exception de Acanlhodactylus

grandis et A. masirae chez lesquels elles sont lisses ; la suboculaire ne touche pas la lèvre supérieure à l’exception

de A. masirae ; le passage des écailles dorsales aux ventrales s’effectue sans transition ou presque sauf chez A.

gongrorhynchatus.

possède un critère qui semble unique au sein du groupe scutellatus ; les écailles dorsales sont

environ deux fois plus grandes que les latérales et le passage des dorsales aux latérales s’effectue

sans transition (fig. 3). Cette particularité pourrait le rapprocher de certains Acanthodactyles du

groupe cantoris (sensu Salvador, 1982; Arnold, 1983) qui, cependant, présentent des rangées

transversales d’écailles ventrales disposées perpendiculairement au corps de l’animal et non obli¬

quement, possèdent des écailles dorsales plus fortement carénées et s’élargissant progressivement

vers l’arrière du dos, et un patron de coloration tout à fait différent.

Acanthodactylus taghitensis n. sp.

Holotype. — Femelle adulte; 36 km au sud-sud-ouest de Taghit (30°41’ N, 2°07’ W), région de Béni

Abbés, Algérie, 3 novembre 1990. MNHN n° 1995-1201. Récolte : A. Foucart.

Etymologie. — taghitensis , de Taghit, localité près de laquelle ont été observés les animaux.

— 8 —

Diagnose

Acanthodactyle de taille relativement faible, présentant 4 séries d’écailles autour des doigts,

4 labiales en avant de la suboculaire, des écailles dorsales carénées et relativement peu nom¬

breuses (44 et 47 rangées à mi-corps), deux fois plus grandes sur le dos que sur les flancs et

sans zones de transition entre les grandes et les petites; 14 rangées longitudinales d’écailles

ventrales disposées en séries transversales obliques ; coloration dorsale beige orangé avec 5 ran¬

gées longitudinales de points sombres reliés entre eux par des traits clairs longitudinaux.

Description de lholotype

Dimensions : longueur tête-corps : 55 mm; longueur de la queue (intacte) : 103 mm; lon¬

gueur de la patte antérieure : 20 mm; longueur patte postérieure : 34 mm; longueur du piléus :

13,1 mm; largeur de la tête : 8,9 mm; hauteur de la tête : 6,8 mm. Pholidose : quatre rangées

d’écailles autour des doigts; écaille suboculaire ne bordant pas la lèvre; 4 labiales en avant de

la suboculaire, celle-ci en contact avec les supralabiales 4 et 5 ; 2 granules insérés entre les

pariétales ; 4 supra-oculaires à droite et à gauche, 1 rangée complète de granules supraciliaires

à droite et à gauche, se dédoublant en arrière de la troisième supra-oculaire; 44 rangées d’écailles

autour du dos à mi-corps ; 30 séries transversales obliques complètes de ventrales disposées en

14 rangées longitudinales à mi-corps; 25 écailles gulaires comptées le long d’une ligne médiane

longitudinale; 21 pores fémoraux à droite et à gauche. Les écailles du dos sont tectiformes et

carénées; elles sont environ deux fois plus grandes que les écailles des flancs; le passage des

écailles dorsales aux latérales s’effectue brusquement.

Coloration : face dorsale beige sable clair; face ventrale blanc pur; le dos est parsemé de

petites taches sombres plus ou moins alignées dans le sens de la longueur et reliées dans la

moitié antérieure du dos par des traits blanchâtres longitudinaux; les labiales et la suboculaire

sont uniformément blanches.

ÉCOLOGIE

Les deux échantillons provenant de deux localités permettent de donner une description

sommaire de l’habitat d 'Acanthodactylus taghitensis : alternance de zones rocheuses et de petites

dunes. Le spécimen désigné comme holotype a été capturé dissimulé sous une pierre, au bord

d’un petit oued asséché, sur du reg sablonneux avec Acacia raddiana et Launaea arborescens,

à quelque 500 m de la bordure du Grand Erg occidental. Autres reptiles observés dans la deuxième

station (5 km au sud-sud-ouest de Taghit) : Mesalina g. guttulata (Lichtenstein, 1823), Uromastyx

acanthinurus Bell, 1825 et Cerastes sp. (traces) (M. GENIEZ comm. pers.). Les deux premières

espèces citées ne sont pas inféodées au sable.

CONCLUSION

La découverte d’une nouvelle espèce de reptile dans la région de Béni Abbés, pourtant

relativement bien connue d’un point de vue herpétologique (c/. Gauthier 1966, 1967) montre

que l’Algérie est un pays incomplètement prospecté. Des recherches dans la région de Taghit

permettraient de trouver d’autres spécimens d 'Acanthodactylus taghitensis et de connaître la mor¬

phologie des mâles et des juvéniles.

— 9 —

Remerciements

Les auteurs sont heureux de pouvoir remercier Michel Geniez qui a photographié le deuxième spéci¬

men d ’Acanthodactylus taghitensis et nous a transmis ses observations, M. J. Bons qui nous a fait part

de son expérience des Acanthodactyles du groupe scutellatus , et J.-Y. Quero pour les photos de l’holotype.

RÉFÉRENCES

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by John Gasperetti. Proc. Calif. Acad. Sci., 4 e sér., 35 (9): 157-192.

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Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 11-30.

Rana (Pelophylax) ridibunda Pallas, 1771,

Rana (Pelophylax) perezi Seoane, 1885

and their associated klepton (Amphibia, Anura) :

morphological diagnoses and description of a new taxon

by Pierre-André CROCHET, Alain DUBOIS, Annemarie OHLER & Heinz TUNNER

Abstract. — Specimens of green frogs from Southern France belonging to the species Rana (Pelophylax)

perezi Seoane, 1885 and to its associated klepton were identified by enzyme electrophoresis. These specimens,

as well as specimens of Rana (Pelophylax) ridibunda Pallas, 1771 from Poland, were submitted to a morphometri-

cal analysis, which allowed to find diagnostic measurements and ratios for the three taxa. These allow to confirm

the identification of the lectotypes of Rana fortis Boulenger, 1884 (currently considered a subjective synonym

of Rana ridibunda, but which might prove to be a distinct taxon) and of Rana esculenta perezi. Both these

lectotypes are redescribed. A Latin scientific name and a description are also provided for the klepton associated

in Southern France and North-Eastern Spain with Rana perezi, and which was until now known as “Graf’s frog”

or “ Rana kl. RP”.

Keywords. — Green frogs, Rana ridibunda. Rana perezi, klepton, Graf’s frog, electrophoresis, morphometry,

diagnostic measurements and ratios.

Rana (Pelophylax) ridibunda Pallas, 1771, Rana (Pelophylax) perezi Seoane, 1885

et leur klepton associé (Amphibia, Anura) : diagnoses morphologiques

et description d’un nouveau taxon

Résumé. — Des spécimens de grenouilles vertes du sud de la France appartenant à l’espèce Rana (Pelo¬

phylax) perezi Seoane, 1885 et à son klepton associé ont été identifiés par électrophorèse de protéines. Ces

spécimens, ainsi que des spécimens de Rana (Pelophylax) ridibunda Pallas, 1771 de Pologne, ont été soumis à

une analyse morphométrique, qui a permis de trouver des mesures et des rapports de mesures diagnostiques pour

les trois taxons. Ces critères permettent de confirmer l’identification des lectotypes de Rana fortis Boulenger,

1884 (nom actuellement considéré comme un synonyme subjectif de Rana ridibunda, mais qui pourrait s’avérer

représenter un taxon distinct) et de Rana esculenta perezi. Ces deux lectotypes sont redécrits. Un nom scientifique

latin et une description sont aussi proposés pour le klepton associé dans le sud de la France et le nord-est de

l’Espagne avec Rana perezi, et qui était jusqu’à présent connu sous les noms de «grenouille de Graf» ou de

«Rana kl. RP».

Mots-clés. — Grenouilles vertes. Rana ridibunda, Rana perezi, klepton, grenouille de Graf, électrophorèse,

morphométrie, mesures et rapports diagnostiques.

P.-A. Crochet, Station biologique de la Tour du Valat, Le Sambuc, 13200 Arles, France, and CEFE-CNRS, B.P. 5051, 34033

Montpellier Cedex, France.

A. Dubois & A. Ohler, Laboratoire des Reptiles et Amphibiens. Muséum national d’Histoire naturelle, 25, rue Cuvier, 75005

Paris, France.

H. Tunner, Institut für Zoologie, Universitat Wien, Althanstrafie 14, 1090 Wien, Austria.

- 12 -

INTRODUCTION

The systematics of European green frogs had a bursting development in the last decades

(Berger, 1973; Hotz, 1974; Dubois, 1977; Günther, 1979; Graf & Polls Pelaz, 1989;

GÜNTHER, 1990; Dubois & Ohler, 1995 a-b). The high level of variation occurring among these

frogs has been observed for a long time, a fact which is reflected in the very heavy synonymies

in this group (Dubois & Ohler, 1995 a-b). Nevertheless, the biological status of the described

taxa started being understood only in the beginning of the seventies (Tunner, 1973), This group

contains not only biological species, but also special hybridogenetic taxons, known as kleptons

(Dubois & Günther, 1982; Dubois, 1991).

A distinct green frog taxon from the Iberian Peninsula and Southern France was described

as Rana esculenta perezi by SEOANE in 1885. It was long considered to be a subspecies of Rana

ridibunda (MERTENS & WERMUTH, 1960). An electrophoretic study by Graf et al. (1977) showed

not only that it is a distinct species (Hotz, 1974), but also that it is associated in Southern

France with a particular hybridogenetic taxon, which has been named “Graf’s frog” (Dubois,

1982) or “ Rana kl. RP” (Graf & Polls Pelaz, 1989). The genotype of this klepton is composed

of one genome from Rana perezi and one genome from Rana ridibunda (Graf & Polls Pelaz,

1989). The association between Rana perezi and Graf’s frog was later shown to occur also is

the north-eastern part of Spain (Uzzell & Tunner, 1983; Arano et al., 1995).

The electrophoretic discrimination between Rana perezi and its associated klepton raises

no particular problem. However, the morphological determination of these two taxa is not cur¬

rently possible. For field studies and for studies of fixed collection specimens, it would appear

very useful to know morphological characters that would allow to discriminate the two forms.

If such characters can be found, they will also allow to check the status of old type-specimens

and to solve nomenclatural problems (Dubois & Ohler, 1995 a-b).

In order to explore this question, we carried out a morphometrical study of specimens of

green frogs from several localities in Southern France belonging to the species Rana perezi and

to its associated klepton. These specimens had been ascribed to their respective taxa on the

basis of enzyme electrophoresis. The morphometrical analysis allowed to discover several dis¬

criminant morphometrical characters for the two taxa. These characters will allow to ascribe old

specimens, including types of ancient nominal taxa, to either of the two biological taxa.

For sake of comparison, we took measurements on a series of specimens of Rana ridibunda

from Poland collected and determined by Leszek Berger, and we provide information on diagnostic

morphometrical characters between this species and the two other green frogs mentioned above.

Rana esculenta perezi was described upon a series of syntypes and the name perezi is cur¬

rently used by specialists of European green frogs for the Iberian and Southern French biological

species, i.e. the non-hybrid taxon. We had the opportunity to study two of the syntypes of this

nominal taxon, including the recently designated lectotype (Dubois & Ohler, 1995a).

For the associated klepton, we first assumed (Dubois & Ohler, 1995a) that there was a

name available: Rana maritima Risso, 1827. This name was based on green frogs from Southern

France (“Alpes-Maritimes”, as defined at the beginning of the nineteenth century), a region where

both Rana perezi and Graf’s klepton were believed to occur according to the maps published

by Graf & Polls Pelaz (1989) and Castanet & Guyétant (1989). Since then, we carried

-13-

out field investigations in order to clarify the biological status of frogs in this part of France:

these researches showed the occurrence of Italian taxa in this area (Dubois & Ohler, 19956;

CROCHET et ai, in preparation). As a consequence, the name Rana maritima Risso, 1827 is

available for Italian green frogs, but not for Graf’s frog. Resolution of this nomenclatural problem

leaves no more name available for Graf’s frog, so that we are led to name it in this paper.

A description of the lectotype of Rana fords Boulenger, 1884 (DUBOIS & Ohler, 1995a)

is also included in this paper. The taxon Rana ridibunda as currently understood is probably

heterogeneous (GÜNTHER, 1982). Mating calls of Rana ridibunda from the Caspian Sea region

(type-locality) were recorded and studied (Schneider & Egiasarjan, 1991) and compared with

those from various East European and West Asian regions (Schneider & SlNSCH, 1992), and

no significant differences were found. However, no such comparisons were made involving mat¬

ing calls recorded in Central, Northern and Western Europe. Obvious differences in general aspect,

colours and size, between frogs from the latter areas and those from East Europe and Asia point

to the possible existence of two distinct species. Future works, using various methods (mor¬

phology, protein electrophoresis, caryology, bioacoustics, etc.) will have to address this question.

If a distinct species had to be recognized for the European frogs, it should bear the scientific

name Rana (Pelophylax) fortis (see DUBOIS & OHLER, 1995a). The specimens from Poland that

we examined in this study are typical for the Central and Western European populations currently

referred to Rana ridibunda, that include also the Berlin frogs for which the name Rana fortis

had been proposed.

MATERIAL AND METHODS

Abbreviations. — MCZ: Museum of Comparative Zoology, Cambridge, Massachusetts, USA; MNHN :

Muséum national d'Histoire naturelle, Paris, France; SVL : Snout-vent length.

Specimens studied. — The specimens listed below were measured. Specimens were also

studied by electrophoresis, except those whose number is followed below by an asterisk (*).

Juveniles, followed below by the sign #, were used only for the computation of values shown

in Fig. 2, but not of values given in Table II.

Rana perezi Seoane, 1885. — France: (A) Pyrénées-Orientales: Banyuls-sur-Mer (42°28’N,

3°07’E): MNHN 1993.3377-3381 (2 adult males and 3 adult females), coll. Jean-Daniel Graf,

March 1976; (B) Bouches-du-Rhône: (1) Faraman (43°25’N, 4°43’E), 2.5 km west of Salin-de-

Giraud: MNHN 1991.77, 1991.83, 1991.88 and 1991.94 (adult males), 1991.84#, 1991.87#,

1991.89-93# and 1991.95-96# (5 juvenile males and 4 juvenile females), coll. Alain Dubois,

8 September 1977; (2) Fumemorte (43°28’N, 4°42’E), 6 km north of Salin-de-Giraud: MNHN

1991.677 (adult male), 1991.676# and 1991.678# (juvenile male and juvenile female), coll. Alain

Dubois, 8 September 1977; (3) Saint-Bertrand (43°27’N, 4°39’E), 7 km north-west of Salin-de-

Giraud: MNHN 1991.675 (adult female), coll. Alain Dubois, 8 September 1977; (4) Collocation

(43°26’N, 4°41’E), 3.5 km north-west of Salin-de-Giraud: MNHN 1991.680 (adult male),

1991.679# (juvenile male), coll. Alain DUBOIS, 8 September 1977; (5) Astouin (43°32'N, 4°24’E),

10 km north of Saintes-Maries-de-la-Mer: MNHN 1991.97# (juvenile female), coll. Alain Dubois,

5 September 1977; (6) Bac du Petit Sauvage (43°29’N, 4°25’E), 5 km north-west of Saintes-

— 14

Maries-de-la-Mer: MNHN 1991.100#, 1991.670# and 1991.672-673# (3 juvenile males and 1 ju¬

venile female), coll. Alain Dubois, 5 September 1977. — Spain: Galicia: La Coruna: (1) La

Coruna (43°22’N, 8°23’W): MCZ 6832* (adult female), coll. D. Victor Lopez Seoane, lectotype

of Rana esculenta perezi', (2) Cabanas (43°24’N, 8°09’W): MNHN 1889.596* (adult female),

coll. D. Victor LOPEZ SEOANE, paralectotype of Rana esculenta perezi.

Rana “kl. RP” (Graf & Polls Pelaz, 1989). — France: (A) Gard: Gorges de l’Aiguillon,

between Lussan (44°09’N, 4°22’E) and La Bastide: MNHN 1993.3374-3376 (adult females), coll.

Jean-Daniel Graf, March 1976; (B) Bouches-du-Rhône: (1) Faraman (43°25’N, 4°43’E), 2.5 km

west of Salin-de-Giraud: MNHN 1991.76 and 1991.78 (adult females), 1991.79-80 and 1991.82

(subadult females), 1991.85 (juvenile male) and 1991.86 (adult male), coll. Alain Dubois, 8

September 1977; (2) Astouin (43°32’N, 4°24’E), 10 km north of Saintes-Maries-de-la-Mer:

MNHN 1991.98-99# (juvenile males), coll. Alain Dubois, 5 September 1977; (3) Bac du Petit

Sauvage (43°29’N, 4°25’E), 5 km north-west of Saintes-Maries-de-la-Mer: 1991.671# and

1991.674# (juvenile males), coll. Alain Dubois, 5 September 1977.

Rana ridibunda Pallas, 1771. — Germany: Berlin (52°32’N, 13°24’E), lake-like expansion

of the river Spree: MNHN 1884.254* (adult female), 1884.255* and 1884.256* (adult males),

coll, fisherman NOACK, spring 1884, respectively paralectotype, lectotype and paralectotype of

Rana fortis Boulenger, 1884. — Poland: Poznan Debina: MNHN 1982.2340* (adult female),

1982.2349* (adult male), 1982.2372-2373* (subadult females), 1982.2379* (subadult male), coll.

Leszek Berger, 20 August 1968.

Electrophoresis. — The specimens of Rana perezi and of its associated klepton were

identified by electrophoresis, except for old type-specimens. Those from Gard (Gorges de l’Ai¬

guillon) and from Pyrénées-Orientales (Banyuls-sur-Mer) were part of those studied and reported

upon by Graf et al. (1977), for which the following enzyme loci were studied: LDH-B, CK,

AAT-1 and AAT-2. Those from Bouches-du-Rhône (Camargue) were identified on the basis of

LDH alone, which is clearly diagnostic for the distinction of the two taxa; the method used for

this electrophoresis is described in TUNNER (1980: 261-262).

Measurements. — Twenty-six measurements (Table I) were taken from all specimens with

caliper or binocular microscope. Except for Rana ridibunda , only electrophoretically determined

adults and subadults were included in the morphometrical analysis which led to values given in

Table II. Since the aim of this work was to find diagnostic measurements, we discarded in the

results those measurements and ratios which did not show significant differences between the

three frog taxa studied.

Statistics. — Despite the low numbers of specimens studied, these numbers are enough

for a statistical analysis using non-parametric tests with a level of significance of .001 (see e.g.

Dubois, 1984). Mean, standard deviation and range were computed for all variables of all groups

on a personal computer with SPSS program (NORUSIS, 1992). The standard non-parametric Mann-

Whitney U test (Zar, 1984) was used to compare the three groups pairwise.

To compare ratios from adult-subadult and juvenile specimens, we used boxplot sketches

(Norusis, 1992). These boxplots display summary statistics for the distribution and plot the

median, the 25th percentile, the 75th percentile and values more than 1.5 box-lengths from 75th

percentile (outliers).

— 15 —

TABLE I. - Description of measurements.

Description des mensurations.

Eye length

Distance from eye to nostril

FLL

Forelimb length (from elbow to base of outer palmar tubercle)

FOTL

Fourth toe length (from base of outer metatarsal tubercle)

Head length (from mandibular articulation to tip of snout)

Head width

IBE

Distance between posterior edges of eyes

Distance between choanae

IFE

Distance between anterior edges of eyes

IMT

Length of inner metatarsal tubercle

IMTH

Height of inner metatarsal tubercle

Intemarial space

ITL

Inner toe length (from distal edge of inner metatarsal tubercle)

Distance between proximal edges of vomerine ridges

MBE

Distance from mandibular articulation to posterior edge of eye

MFE

Distance from mandibular articulation to anterior edge of eye

Distance from mandibular articulation to nostril

SVL

Snout-vent length

TFL

Third finger length (from base of proximal subarticular tubercle)

Tibia length

TYD

Greatest tympanum diameter

TYE

Distance from tympanum to back of eye

VRL

Length of vomerine ridge

VTL

Distance between distal edges of vomerine ridges

WOFF

Webbing between fourth and fifth toe (from base of outer metatarsal tubercle)

WOTF

Webbing between third and fourth toe (from base of outer metatarsal tubercle)

Nomenclature of kleptons. — For the nomenclatural treatment of kleptons, we follow

the proposals of Dubois & Günther (1982) and Dubois (1991), according to which kleptons

are taxa of the species-group that are nomenclaturally of the species rank but that have biological

properties different of those of “biological species” (“normal”, bisexual species). Kleptons receive

scientific Latin names similar to those of species, but which can be distinguished from the latter

by the insertion of the sign “kl.” between the genus-group name(s) and the species-group name(s).

We follow Dubois & Ohler (1995a) in their placement of European green frogs in the subgenus

Pelophylax Fitzinger, 1843 of the genus Rana Linnaeus, 1758.

Descriptive methods. — In order to facilitate comparisons, the detailed descriptions of

the three type-specimens given below follow the same plan. Webbing formula is presented ac¬

cording to Myers & Duellman (1982).

RESULTS

Morphometrical comparisons of Rana perezi, Rana ridibunda and Graf’s frog

Electrophoretic data obtained from the specimens collected in Bouches-du-Rhône, Gard and

Pyrénées-Orientales allowed to refer clearly these specimens to two taxa: Rana perezi and its

associated hybridogenetic klepton (see Graf et al., 1977; Graf & POLLS Pelaz, 1989). For

— 16 —

the research of diagnostic measurements, we pooled all specimens electrophoretically character¬

ized, irrespective of their sexes, ages and localities: thus we had 12 adult and subadult specimens

of Rana perezi and 9 adult and subadult specimens of Graf’s klepton.

Thirty-seven ratios were tested pairwise with the Mann-Whitney U test. Fourteen ratios

were found to be diagnostic between Rana perezi and Graf’s frog (Table II). These measurements

are from various parts of the body: head, foot and webbing. Particularly interesting and diagnostic

are the measurements which involve the vomerine ridges. The ratios IV/VRL, IV/IC, IV/SVL

and also WOTF/FOTL show highly significant differences between our two samples. Rana perezi

has a greater distance between vomerine ridges (Fig. 1), which appears in the ratios involving

IV. It also has a longer foot and a less developed webbing.

The sample of the hybridogenetic klepton is statistically different from Rana ridibunda for

16 ratios (Table II). The measurements involve also the different parts of the body. The ratios

TYD/IN, TYD/SVL, TYE/SVL and VRL/IC show rather high significant differences between

the two samples. Rana ridibunda's tympanum is relatively smaller than in both Rana perezi and

Graf’s frog. The distance between vomerine ridges is somewhat intermediate. Rana ridibunda

shows the highest development of webbing of the three forms.

Twelve ratios describing the two groups representing Rana ridibunda and Rana perezi show

statistically significant differences (Table II). Seven of these ratios (HW/SVL. IV/IC, IV/VRL,

TYD/SVL, TYD/TYE, WOTF/FOTL, WOTF/SVL) show rather high significant differences be¬

tween the two samples. Rana perezi has a clearly narrower head than Rana ridibunda , a larger

tympanum and tympanum-eye distance, and less webbing on feet. Vomerine ridges and associated

features are more separated than in Rana ridibunda.

Five ratios show significant differences between all three groups (IV/IC, IV/SVL, IV/VRL,

TYD/SVL, TYD/TYE). They concern vomerine ridges’ position and tympanum size. Foot

measurements of Rana kl. RP show intermediate values between Rana ridibunda and Rana perezi.

But the values for vomerine teeth morphology are intermediate in Rana ridibunda and extreme

in the hybrid.

Some ratios, which are clearly diagnostic between Rana perezi and Graf’s frog in adults,

can also be used cautiously for determination of juveniles of these two forms, as shown here

in Fig. 2 for the ratio IV/VRL: out of 42 specimens studied, only one showed an ambiguous

value for this ratio.

Of the ratios diagnostic for the complex Rana lessonae - Rana kl. esculenta - Rana rid¬

ibunda (Berger, 1973), only one, TL/SVL, shows a significant statistical difference between

Rana ridibunda and Rana perezi.

Study of two syntypes of Rana esculenta perezi Seoane, 1885

We were able to study and measure two of the original syntypes of Rana esculenta perezi

and we computed the ratios shown above to allow morphological distinction of the three taxa

studied. Table III shows that the values of most of these ratios fall within the ranges of perezi.

We conclude that these two syntypes are morphologically similar to the electrophoretically charac¬

terized specimens of Rana perezi. Designation by Dubois & Ohler (1995a) of one of these

two syntypes as lectotype of perezi was therefore appropriate to stabilize definitely the nomen-

clatural status of this name. A detailed description of this lectotype is provided below.

-17-

DESCRIPTIONS OF TYPE-SPECIMENS

Rana (Pelophylax) kl. grafi kl. nov.

(Figs la, 3)

Holotype. — MNHN 1993.3374, adult female (SVL 111.0 mm), collected by Jean-Daniel Graf in the Gorges

de l’Aiguillon, between Lussan (44°09’N, 4°22’E) and La Bastide, Gard, France, in March 1976.

Paratypes. — MNHN 1993.3375-3376, adult females (SVL 94.5 mm and 56.9 mm), same collection data

as the holotype. MNHN 1991.76, adult female (SVL 71.9 mm), 1991.78, adult female (SVL 66.5 mm), 1991.79-80,

subadult females (SVL 91.0 and 71.2 mm), 1991.82, subadult female (SVL 55.2 mm), 1991.85, juvenile male

(SVL 39.2 mm) and 1991.86, adult male (SVL 33.9 mm), collected by Alain Dubois at Faraman (43°25’N, 4°43’E),

2.5 km west of Salin-de-Giraud, Bouches-du-Rhône, France, on 8 September 1977. MNHN 1991.98-99, juvenile

males (SVL 27.6 mm and 36.5 mm), collected by Alain Dubois at Astouin (43°32’N, 4°24’E), 10 km north of

Saintes-Maries-de-la-Mer, Bouches-du-Rhône, France, on 5 September 1977. MNHN 1991.671 and 1991.674,

juvenile males (SVL 29.9 mm and 24.9 mm), collected by Alain Dubois near the Bac du Petit Sauvage (43°29’N,

4°25’E), 5 km north-west of Saintes-Maries-de-la-Mer, Bouches-du-Rhône, France, on 5 September 1977.

Note on paratypes. — Paratypes are morphologically similar to the holotype.

Etymology of species-group name. — We dedicate this frog to Jean-Daniel Graf who discovered this new

klepton and kindly provided us with some of the specimens he had studied by electrophoresis.

Diagnosis

A large-sized green frog. It can be distinguished morphologically from Rana perezi by more

webbing on feet and higher metatarsal tubercle. It differs from Rana ridibunda by its smaller

head width, eye-tympanum distance smaller than half of tympanum diameter, shorter fourth toe

and lesser webbing. Of both taxa, Graf’s frog is distinguished by its smaller intervomer distance,

longer vomerine teeth ridges and smaller ratio of vomerine ridge length by intervomer distance.

Electrophoretic characteristics of this taxon were published by Graf et al. (1977).

Description of holotype

Specimen with tissue sampling, snout-vent length 111.0 mm. Head slightly longer (35.2 mm)

than broad (34.0 mm); snout moderately pointed, slightly protruding beyond mouth, slightly

shorter (8.2 mm) than diameter of eye (8.6 mm); canthus rostralis rounded, loreal region abrupt,

concave; interorbital space flat, smaller (4.3 mm) than both internarial distance (5.1 mm) and

upper eyelid width (6.8 mm); nostril slightly nearer (7.4 mm) to eye than to tip of snout (8.2 mm);

tympanum present, its diameter (6.9 mm) more than two third of diameter of eye, its distance

to eye (5.1 mm) roughly three fourth of its diameter; vestige of pineal eye absent; tongue ovally

elongated, emarginated behind, finely granulated. Vomerine teeth between choanae in two ridges

forming an angle of 150°; ridges long (6.3 mm), 63.0 times the separation between them (0.1 mm).

Arm short, forearm (19.8 mm) shorter than hand (23.5 mm); finger length, shortest to

longest: II < I < IV < III; length of third finger 13.8 mm; tips of fingers pointed; three metacarpal

tubercles, moderately developed; supernumerary tubercle on base of finger III; subarticular

tubercles rounded, moderately developed.

Hind limbs moderately long, shank three times longer (48.2 mm) than broad (15.1 mm),

slightly shorter than thigh (50.9 mm); shank and thigh both shorter than distance from base of

inner metatarsal tubercle to tip of toe IV (51.6 mm); toes moderately long, length of IV (30.6 mm)

— 18 —

Table II. — Results of Mann-Whitney U-test comparing some morphometric ratios of adults and subadults of Rana perezi (P;

n = 13), Rana kl. graft (G; n = 8) and Rana ridibunda (R; n = 5). Ranges, mean and standard deviation are given for each

species. Significance level : *** 0.001; ** 0.010; * 0.050.

Résultats de la comparaison par le test U de Mann-Whitney de quelques rapports morphométriques chez des adultes et

subadultes de Rana perezi (P ; n = 13), Rana kl. grafi (G ; n = 8) et Rana ridibunda (R ; n = 5). Pour chaque espèce sont

données les valeurs extrêmes du rapport, la moyenne et l'écart-type. Coefficients de risque : ***0.001 ; ** 0.010 : *0.050.

Ratio

Rana perezi

Rana kl. grafi

Rana ridibunda P x R

P x G

R x G

FOTL/SVL

0.560 ± 0.053

0.479 - 0.632

0.536 ± 0.042

0.455 - 0.588

0.582 ± 0.015

0.556 - 0.595

HW/SVL

0.341 ± 0.012

0.322 - 0.359

0.351 ± 0.024

0.306 - 0.393

0.376 ± 0.015

0.353 - 0.393

IBE/SVL

0.236 ± 0.021

0.192 - 0.268

0.314 ± 0.017

0.179 - 0.230

0.221 ± 0.017

0.204 - 0.239

I C/S VL

0.099 ±0.010

0.082 - 0.115

0.086 ± 0.007

0.071 - 0.097

0.096 ± 0.005

0.091 - 0.102

IMT/SVL

0.051 ± 0.006

0.038 - 0.059

0.054 ± 0.006

0.048 - 0.065

0.047 ± 0.006

0.044 - 0.057

IN/SVL

0.069 ± 0.009

0.053 - 0.084

0.058 ± 0.007

0.046 - 0.069

0.062 ± 0.006

0.056 - 0.069

IV/IC

0.11 ± 0.03

0.066 - 0.171

0.03 ± 0.02

0.013 - 0.060

0.06 ± 0.01

0.046 - 0.073

***

IV/SVL

0.011 ± 0.004

0.006 - 0.018

0.003 ± 0.002

0.001 - 0.005

0.006 ± 0.001

0.004 - 0.007

***

IV/VRL

0.346 ± 0.139

0.192 - 0.692

0.008 ± 0.004

0.003 - 0.016

0.158 ± 0.032

0.123 - 0.200

***

THL/SVL

0.494 ± 0.029

0.448 - 0.559

0.517 ± 0.022

0.467 - 0.534

0.781 ± 0.023

0.457 - 0.509

TL/SV L

0.507 ± 0.034

0.445 - 0.557

0.531 ± 0.037

0.459 - 0.574

0.551 ± 0.018

0.525 - 0.566

TYD/IN

1.34 ± 0.30

1.00 - 2.00

1.86 ± 0.19

1.61 - 2.22

1.11 ± 0.10

0.973 - 1.19

TYD/SVL

0.091 + 0.008

0.081 - 0.108

0.106 ± 0.005

0.099 - 0.111

0.069 ± 0.003

0.066 - 0.072

TYD/TYE

2.86 ± 0.71

1.79 - 4.25

2.16 ± 0.21

1.83 - 2.39

1.88 ± 0.22

1.68 - 2.25

TYE/SVL

0.034 ±0.011

0.021 - 0.049

0.050 ± 0.006

0.044 - 0.059

0.037 ± 0.005

0.029 - 0.042

VRL/IC

0.34 ± 0.07

0.246 - 0.480

0.43 ± 0.04

0.365 - 0.482

0.36 ± 0.02

0.336 - 0.373

VRL/SVL

0.047 ±0.018

0.026 - 0.073

0.080 ± 0.023

0.057 - 0.132

0.069 ± 0.007

0.059 - 0.074

WOFF/SVL

0.339 ± 0.034

0.291 - 0.400

0.364 ± 0.034

0.312 - 0.405

0.384 ± 0.012

0.368 - 0.402

WOTF/FOTL

0.59 ± 0.06

0.435 - 0.648

0.67 ± 0.03

0.617 - 0.715

0.68 ± 0.02

0.638 - 0.697

***

WOTF/SVL

0.330 ± 0.035

0.284 - 0.397

0.358 ± 0.026

0.312 - 0.386

0.394 ± 0.010

0.380 - 0.405

WOTF/WOFF

0.976 ± 0.046

0.899 - 1.05

0.987 ± 0.033

0.945 - 1.06

1.03 ± 0.020

1.01 - 1.06

— 19 —

Fig. 1. — Vomerine teeth position in (a) Rana kl. graft (MNHN 1993.3374, holotype) and (b) Rana perezi (MNHN 1993.3377).

Scale: 5 mm. For the meaning of abbreviations, see Table I.

Position des dents vomériennes chez (a) Rana kl. grafi (MNHN 1993.3374, holotype) et (b) Rana perezi (MNHN 1993.3377).

Échelle : 5 mm. Pour la signification des abréviations, voir le tableau 1.

Fig. 2. — Boxplot sketches displaying summary statistics for the ratio IV/VRL in adult-subadult and juvenile specimens of Rana

kl. grafi and Rana perezi. The median, the 25th percentile, the 75th percentile and outliers are plotted. N: number. For the

meaning of other abbreviations, see Table I.

Diagramme « boxplot » montrant les paramètres statistiques principaux pour le rapport IV/VRL chez des spécimens adultes-

subadultes et juvéniles de Rana kl. grafi et Rana perezi. La médiane, le 25 e percentile, le 75 e percentile et les valeurs ex¬

centriques («outliers ») sont figurés. N: nombre. Pour la signification des autres abréviations, voir le tableau I.

less than one half of length from base of tarsus to tip of toe IV (72.6 mm): tips of toes rounded;

webbing moderately developed (I 1/2 - 1 II 1/2 - 1 1/2 III 1/2 - 1 1/2 IV I - 1/2 V); inner

metatarsal tubercle moderately developed, its length (5.3 mm) 2.9 times in length of toe I

(15.5 mm); glandular ridge on tarsus prominent.

Dorsum with large flat warts, also between eyes; two broad glandular dorsolateral ridges;

ventral surfaces smooth.

Colour in alcohol: Brownish with few large darker spots; a slightly clearer mediodorsal

line; dorsolateral folds darker brown; forelimbs without darker spots; hind limbs with few darker

bands; ventral surface cream and brown marbled, blurred; back of thigh dark brown with brown

spots.

3. — Rana (Pelophylax) kl. graft kl. nov., MNHN 1993.3374 (holotype): (a) dorsal view of head; (b) roof of mouth; (c)

ventral view of right foot.

Rana (Pelophylax) kl. grafi kl. nov.. MNHN 1993.3374 (holotype): (a) vue dorsale de la tête; (b) plafond buccal: (c) vue

ventrale du pied droit.

4. Rana (Pelophylax) perezi Seoane, 1885, MCZ 6832 (lectotype): (a) dorsal view of head; (b) roof of mouth- (c) ventral

view of right foot.

Rana (Pelophylax) perezt Seoane, 1885, MCZ 6832 (lectotype): (a) vue dorsale de la tête; (b) plafond buccal; (c) vue

— 23 —

Rana (Pelophylax) perezi Seoane, 1885

(Figs lb, 4)

Lectotype (by designation of Dubois & Ohler, 1995a) of Rana esculenta perezi Seoane, 1885: 171. —

MCZ 6832, adult female (SVL 63.6 mm), collected by D. Victor Lopez Seoane near La Coruna (43°22’N, 8°23'W),

Galicia, Spain, from Fernand Lataste’s collection.

Paralectotype of Rana esculenta perezi Seoane, 1885: 171. — MNHN 1889.596, adult female (SVL

71.5 mm), from Cabanas (43°24’N, 8°09'W), Galicia, Spain, donation of Victor Lopez Seoane to the Paris Museum

on 9 November 1889.

Note on paralectotype. — The paralectotype studied is similar to the lectotype, except that it shows an

asymmetry in its vomerine teeth. Other paralectotypes may possibly be found in the future, in other Museums

where Seoane or Lataste may have deposited some of the original syntypes of this taxon.

Diagnosis

A medium-sized green frog. It can be distinguished morphologically from Rana kl. grafi

by less webbing on its feet and its smoother metatarsal tubercle. It differs from Rana ridibunda

by its smaller head, tympanum closer to eye, shorter hand and foot. It has a larger intervomer

distance than both Rana ridibunda and Rana kl. graft.

Description of lectotype

Well-preserved specimen, snout-vent length 63.6 mm. Head slightly longer (22.9 mm) than

broad (21.8 mm); snout rounded, not protruding beyond mouth, longer (10.4 mm) than diameter

of eye (7.7 mm); canthus rostralis rounded, loreal region abrupt, concave; interorbital space flat,

smaller (1.9 mm) than both internarial distance (4.2 mm) and upper eyelid width (4.6 mm); nostril

slightly nearer (4.8 mm) to eye than to tip of snout (5.3 mm); tympanum present, its diameter

(4.1 mm) more than half of diameter of eye, its distance to eye (1.4 mm) roughly one third of

its diameter; vestige of pineal eye absent, tongue ovally elongated, deeply emarginated behind,

finely granulated. Vomerine teeth between choanae in two ridges forming an angle of 140°;

ridges long (1.94 mm), 3.1 times the separation between them (0.62 mm).

Arm short, forearm (11.2 mm) shorter than hand (14.2 mm); finger length, shortest to longest:

I < II < IV < III; length of third finger 8.5 mm; tips of fingers pointed; metacarpal tubercles

not distinct, subarticular tubercles rounded, moderately developed.

Hind limbs relatively short, shank almost three times longer (28.3 mm) than broad

(11.3 mm), slightly shorter than thigh (29.1 mm); shank and thigh both shorter than distance

from base of inner metatarsal tubercle to tip of toe IV (34.6 mm); toes moderately long, length

of IV (19.2 mm) less than one third of length from base of tarsus to tip of toe IV (46.9 mm);

tips of toes pointed; webbing moderately developed (I 1/2 - 2 II 1 - 2 III 1 - 2 IV 2 - 1 V);

inner metatarsal tubercle moderately developed, its length (3.6 mm) 2.3 times in length of toe

I (8.3 mm); glandular ridge on tarsus flat.

Dorsum with large flat warts, except on top of head; two broad glandular dorsolateral ridges;

ventral surfaces smooth.

Colour in alcohol: Brownish with few large darker spots; a slightly clearer mediodorsal

line; dorsolateral folds slightly darker brown; forelimbs with few darker spots; hind limbs with

darker spotlike bands; ventral surface cream and brown marbled, very homogeneous; back of

thigh blackish with few clearer spots, a dark line separating ventral from dorsal surface.

5. — Rana (Pelophylax) ridibunda Pallas, 1771, MNHN 1884.255 (lectotype of Rana forth Boulenger, 1884): (a) dorsal

view of head; (b) roof of mouth; (c) ventral view of right foot.

Rana (Pelophylax) ridibunda Pallas, 1771, MNHN 1884.255 (lectocype de Rana fortis Boulenger, 1884): (a) vue dorsale de

la tête: (b) plafond buccal: (c) vue ventrale du pied droit.

— 25 —

Rana (Pelophylax) ridibunda Pallas, 1771

(Fig. 5)

Lectotype (by designation of Dubois & Ohler, 1995a) of Rana fortis Boulenger, 1884: 220. — MNHN

1884.255, adult male (SVL 71.4 mm), collected by fisherman Noack in a lake-like expansion of the river Spree,

near Berlin, Berlin, Germany.

Paralectotypes of Rana fortis Boulenger, 1884: 220. — MNHN 1884.254, adult female (SVL 70.5 mm)

and 1884.256, adult male (SVL 70.5 mm), same collection data as the lectotype.

Note on paralectotypes. — Both paralectotypes studied are similar in size and colour to the lectotype.

Other paralectotypes will certainly be found in the future in other Museums, as Boulenger (1884) described this

species using a series of 85 syntypes.

Diagnosis

A large-sized green frog. Webbing of foot more important than in Rana perezi and Rana

kl. grafi. Head larger than in these two taxons, tympanum-eye distance larger than half of tym¬

panum diameter, vomerine teeth position intermediate.

Description of lectotype of Rana fortis Boulenger, 1884

Slightly dried alcohol fixed specimen, snout-vent length 71.4 mm. Head longer (28.3 mm)

than broad (26.0 mm); snout rounded, not protruding beyond mouth, longer (10.4 mm) than

diameter of eye (7.6 mm); canthus rostralis rounded, loreal region abrupt, concave; interorbital

space flat, smaller (2.3 mm) than both internarial distance (4.0 mm) and upper eyelid width

(4.9 mm); nostril slightly nearer (5.5 mm) to eye than to tip of snout (5.3 mm); tympanum present,

its diameter (5.3 mm) more than half of diameter of eye, its distance to eye (2.9 mm) roughly

one half of its diameter; vestige of pineal eye absent, tongue ovally elongated, deeply emarginated

behind, finely granulated. Vomerine teeth between choanae in two ridges forming an angle of

130°; ridges long (1.59 mm), 4.9 times the separation between them (0.34 mm).

Arm short, forearm (16.3 mm) shorter than hand (18.4 mm); finger length, shortest to

longest: I = II < IV < III; length of third finger 9.9 mm; tips of fingers pointed; metacarpal

tubercles not distinct, subarticular tubercles rounded, moderately developed.

Hind limbs relatively short, shank four times longer (36.1 mm) than broad (8.3 mm), longer

than thigh (29.6 mm); shank and thigh both shorter than distance from base of inner metatarsal

tubercle to tip of toe IV (41.2 mm); toes moderately long, length of IV (24.0 mm) more than

one third of length from base of tarsus to tip of toe IV (56.2 mm); tips of toes pointed; webbing

moderately developed (I 1/2 - 1 1/2 II 1/2 - 2 III 1 - 2 IV 2 - 1/2 V); inner metatarsal tubercle

moderately developed, its length (3.4 mm) 3.4 times in length of toe I (11.6 mm); glandular

ridge on tarsus flat.

Dorsum with large flat warts, except on top of head; two broad glandular dorsolateral ridges;

ventral surfaces smooth.

Colour in alcohol: Grayish with few large darker spots; a slightly clearer mediodorsal line;

dorsolateral folds slightly clearer; forelimbs with few darker spots; hind limbs with darker bands;

ventral surface whitish marbled with black, very homogeneous; back of thigh blackish with few

clearer spots. Vocal sacs blackish, nuptial pads on finger I blackish.

— 26 —

Table III. — Some morphometric ratios of type-specimens, allowing comparison with electrophoretically studied material. Values

followed by an asterisk * fall outside the range of variation for the taxon given in Table II. (-): measurement not possible

due to injuries. H: holotype; L: lectotype; PL: paralectotype.

Quelques rapports morphométriques chez quelques spécimens-types, permettant une comparaison avec les spécimens étudiés

par électrophorèse. Les valeurs suivies d’un astérisque tombent en dehors de l’intervalle de variation donné pour le taxon

dans le tableau II. (—) : mensuration impossible en raison de blessures. H: holotype; L: lectotype; PL: paralectotype.

Rana perezi

Rana kl.

grafi

Rana forth

Ratio

MCZ 6832

L (adult 9 )

MNHN

1889.596

PL (adult 9 )

MNHN

1993.3374

H (adult 9)

MNHN

1884.255

L (adult d)

MNHN

1884.254

PL (adult 9 )

MNHN

1884.256

PL (adult d)

FOTL / SVL

0.511

0.502

0.481

0.577

0.580

HW / SVL

0.343

0.351

0.332

0.364

0.363

0.369

IBE / SVL

0.218

0.234

0.179

0.200

0.217

0.203

IC / SVL

0.0933*

0.0909*

0.0712

0.0862*

0.101

0.0938

IMT / SVL

0.059 1

0.0643

0.0478

0.0476

0.0511

0.0440

IN / SVL

0.0622*

0.0573

0.0460

0.0672

0.0644

0.0634

IV / IC

0.0997

0.0923

0.0127

0.0612

0.0436*

0.0470

IV / SVL

0.0093

0.0084

0.00097

0.0052

0.0044

0.044

IV / VRL

0.073

0.231

0.0282

0.183

0.127

0.172

THL / SVL

0.446

0.464

0.467

0.458

0.507

TL / SVL

0.459

0.464

0.459

0.505*

0.504*

0.509*

TYD / IN

1.225*

1.342*

1.373

1.104

1.171

1.073

TYD / SVL

0.0762*

0.0769*

0.0681

0.0754

0.0754*

0.0680

TYD / TYE

2.882*

2.75*

1.353

1.822

1.907

1.762

TYE/SVL

0.0264*

0.0280*

0.0331

0.0409

0.0395

0.0386

VRL / IC

0.367

0.379

0.482

0.296*

0.345

0.273*

VRL/SVL

0.0342

0.0344

0.0568

0.0253*

0.0349

0.0256*

WOFF / SVL

0.311

0.305

0.331

0.377

0.392

0.369

WOTF / FOTL

0.598

0.618

0.674

0.695

0.691

0.667

WOTF / SVL

0.306

0.311

0.324

0.389

0.401

0.380

WOTF / WOFF

0.984

1.020

0.979

1.032

1.018

1.030

— 27 —

DISCUSSION

Morphological distinction between Rana perezi and Ran a kl. grafi

In our view, one of the goals of contemporaneous systematists should be to reconcile modern

and traditional methods of biology to ensure continuity in science. Studies of genotypes allow

understanding of genetic mechanisms involved in the evolution of taxa. But such methods cannot

be used without heavy investigations and should be paired with study of external characters and

morphology. On the basis of studies linking morphological and genetic information, keys based

on purely morphometrical characters can be worked out. Such keys are necessary for work on

collection material, field work, and preparation of distribution maps. Until now, most information

collected in the past by ancient authors on the distribution of green frogs is obsolete, as the

level of determination depended on the worker. To prepare well-founded keys would permit to

go further in our knowledge about biogeography and evolution of green frogs.

The morphologies of Rana perezi and Rana kl. graft had not been compared until now.

This paper is a first step in this respect. Even if the sample is small, clear (significant) mor¬

phometrical differences were found between two electrophoretically determined groups of frogs.

These characters permit identification of frogs for which no electrophoretic data are available.

The morphometric ratios traditionally used for species identification in the complex lessonae

- esculenta - ridibunda do not work in the complex perezi - grafi - ridibunda. In the latter

complex, the differentiation is more important in foot and vomerine teeth morphologies than in

tibia, inner metatarsal tubercle, and inner toe lengths. This shows clearly that morphological

studies should not only include characters that were valid for certain groups of species (“good”

characters), but that the analysis should embrace the whole morphology, including measurements

that concern all parts of the body.

Because of the small number of electrophoretically determined specimens available, in this

study we pooled specimens from different populations, of the two sexes and of various ages

and sizes. We excluded young specimens, because usually in frogs morphological changes are

very important during ontogeny, mostly as a result of allometric growth. In the future, it will

be interesting to investigate about variation between different populations and to include more

taxa for comparison. But, to correctly study such a problem, close attention should be paid to

the important intrapopulational variations in frogs due to sexual dimorphism and allometric

growth.

In some of the studied ratios, Rana kl. grafi is intermediate between the two non-hybrid

species. This might be a consequence of the hybrid origin of the klepton. In other ratios, Rana

kl. grafi shows distinct new characters. Vomerine teeth position is not like in any of the two

non-hybrid species, nor is it intermediate. It is clearly a new character.

Nomenclatural problems

A major aim of this study was to solve nomenclatural problems concerning green frogs

from Southern France and Iberian Peninsula using the most parsimonious way. In a first step

we defined morphometrical differences between the two taxa occurring there. Then we used

these characters for phenotypical determination of the lectotype and one paralectotype of Rana

— 28 —

esculenta perezi Seoane, 1885: this study confirmed that both specimens are members of the

non-hybrid taxon. This determination is consistent with the fact that only non-hybrid specimens

have been found in Western Spain until now (UZZELL & TUNNER. 1983; Arano et al. , 1995).

This solves the first nomenclatural problem. The second is solved by the proposal in this paper

of a new scientific name, Rana kl. graft, for the klepton found associated with Rana perezi in

some areas.

Historical considerations

European green frogs seem to be a monophyletic group closely related to East Asian green

frogs (Uzzell, 1982), with which they constitute the subgenus Pelophylax (Dubois, 1992).

Within the Western green frogs, Rana perezi and Rana saharica seem to form a monophyletic

group (Uzzell, 1982; Beerli, 1993). This group probably was isolated on the African continent

when the Mediterranean Sea was formed. Immigration to the Iberian Peninsula was made possible

by the closing of the Straits of Gibraltar. Estimated divergence time between Rana perezi and

Rana saharica corresponds to the date of reopening of the Straits of Gibraltar (Beerli, 1993).

Currently, the Northern half of France is occupied by the L-E system (Uzzell & Berger,

1975), i.e. mixed populations of Rana lessonae and Rana kl. esculenta , while the southern part

of the country is occupied by the P-RP system (Graf & Polls Pelaz, 1989) or P-G system,

i.e. mixed populations of Rana perezi and Rana kl. grafi. Besides, there is evidence of occurrence

of isolated populations of Rana ridibunda in various parts in France (Castanet & Guyetant,

1989; Crochet, Dubois & Ohler, unpubl.). The origin of the latter frogs is unknown: some

populations (near universities, etc.) are probably introduced (see e.g. Dubois, 1982), while others

are so isolated that such an origin seems unlikely. These frogs have not yet been compared with

Rana ridibunda of Central and Northern Europe.

Rana perezi seems to have been a rather recent invader of Europe coming from the South.

During its expansion it may have come in contact with the more Northern and Eastern European

taxa, possibly in Southern France. Rana kl. grafi would have appeared through hybridization in

this contact zone. There may have been primarily direct contact between Rana perezi and Rana

ridibunda, or the ridibunda genome may have been transmitted from esculenta to grafi through

hybridization between perezi and esculenta (Dubois & GÜNTHER, 1982: 296).

Other hypotheses can be considered. As the estimated age of European green frogs is much

older than the estimated age of immigration of the perezi-saharica stock into the Iberian

Peninsula, there might have existed a pr e-perezi frog in South-Western Europe. This might have

been the L-E system, progressively replaced by the P-G system, with a contact/hybrid zone that

would move northwards with progression of the latter. This would require competitive superiority

of the P-G system over the L-E system in this area, which could be experimentally tested. A

detailed study of the composition, structure and dynamics of green frog populations in areas

where the P-G system probably meets the L-E system (e.g. Charente in Western France, Rhone

valley. Southern Massif central) might also throw interesting lights on this question. Another

hypothesis would be that a ridibunda- like species might have been replaced in the Iberian

Peninsula by the P-G system. Some of the morphometrical particularities of grafi might indicate

that the second parental species was somewhat different from the current European ridibunda.

— 29 —

Acknowledgements

We warmly thank the following colleagues for the help they provided for this work: Jean-Daniel

Graf (Genève) deposited some of his electrophoretically determined specimens in the Paris Museum col¬

lections; Leszek Berger (Poznan) gave a large collection of Polish frogs to the Paris Museum; Jose

P. Rosado (Harvard, Cambridge) sent us in loan the syntype of Rana esculenta perezi kept in the collections

of the Museum of Comparative Zoology; Roger Bour (Paris) provided helpful comments and suggestions;

Laurent Bessol (Paris) kindly prepared the macrophotographs of Fig. 1.

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Abstract. — Two species of Candelabrum (Cnidaria, Hydrozoa, Athecata) have been discovered and col¬

lected at two hydrothermal areas of the Mid-Atlantic Ridge with French and American submersibles during five

cruises in 1988, 1993 and 1994. The first, Candelabrum serpentarii nov. sp., has been found at the Snake Pit

site (23°N, 3500 m depth) and represents an undescribed species. The second. Candelabrum phrygium (Fabricius,

1780), collected at the Lucky Strike site (37°N, 1700 m depth), is known to have a circumarctic distribution. In

spite of great differences in depth of occurrence, considerable morphological similarities exist between the new

species and the intertidal species C. cocksii (Vigurs, 1849). This discovery has given rise to a review of the

genus Candelabrum. In addition, ecological remarks on these three species are presented.

Keywords. — Hydrothermal vents, Mid-Atlantic Ridge, Hydrozoa, Candelabrum, biogeography, plate tec¬

tonics.

Premières observations du genre Candelabrum (Cnidaria, Hydrozoa, Athecata)

sur des zones hydrothermales de la dorsale médio-atlantique :

description d’une nouvelle espèce et revue du genre

Résumé. — Deux espèces de Candelabrum (Cnidaria, Hydrozoa, Athecata) ont été découvertes et récoltées

sur deux aires hydrothermales de la dorsale médio-atlantique grâce aux sous-marins français et américains au

cours de cinq missions en 1988, 1993 et 1994. La première. Candelabrum serpentarii nov. sp., récoltée sur le

site du Snake Pit (23° N, 3 500 m), est une espèce non décrite. La seconde. Candelabrum phrygium (Fabricius,

1780), récoltée sur le site Lucky Strike (37° N, 1 700 m), est connue pour sa répartition circumarctique. En dépit

des différences de profondeur des habitats, on remarque de fortes ressemblances morphologiques entre l’espèce

non décrite et l’espèce littorale connue C. cocksii. Ces découvertes donnent lieu à une revue des espèces du

genre Candelabrum. Quelques remarques écologiques sur ces trois espèces sont présentées.

Mots-clés. — Sources hydrothermales, dorsale médio-atlantique, Hydrozoaire, Candelabrum, biogéographie,

tectonique des plaques.

M. Segonzac, IFR EM ER. BP 70. 29280 Plouzané (France).

W. Vervoort, Nationaal Natuurhistorisch Museum, PO Box 9517, 2300 RA, Leiden (Netherlands).

INTRODUCTION

The first Atlantic hydrothermal communities found in the axial valley of the Mid-Atlantic

Ridge: the TAG (Trans-Atlantic Geotraverse) area at 26°N (Fig. 1) and 3700 m depth, and the

Snake Pit area at 23°N and 3500 m depth, have been described respectively by Galkin et al.

(1990) and SEGONZAC (1992). At the Snake Pit area, an undescribed species of Candelabrum

— 32 —

was collected by the submersible Nautile during the French cruise Hydrosnake in June 1988

and several other specimens were seen next to the active edifices. In June 1993, another specimen

was collected at the same site with the submersible Alvin during the American-French cruise

MAR 93. In situ, this organism, belonging to the class Hydrozoa, appeared as a whitish gelatinous

pen, undulating according to the water movements. Fixed on pillow lava or sulfide rock, it is

about 10 cm long and presents, on its base, pure white, clustered granulations (gonophores). It

was the first time that such organism, known from shallower waters, had been observed among

the hydrothermal communities.

Sampling of the hydrothermal fauna of the newly found Lucky Strike area (SW Azores

Islands, 37°N, 1700 m depth), realized in May 1993 with the submersible Alvin (Langmuir et

al., 1993), allowed the collection of another species of hydroid, known as Candelabrum phrygium

(Fabricius, 1780), among a sample of the mussel Bathymodiolus sp. During the French cruise

Diva 2 (June 1994), several other specimens were observed and collected at the same location

with the submersible Nautile.

In the Hydroida (hydropolyps-hydromedusae) of the Hydrozoa, the subclass Athecatae (An-

thomedusae) is principally characterized by the absence of a distinct hydrotheca around the polyp.

It comprises c. 50 families that are mostly marine. Among these families, the Candelabridae has

only two genera: Candelabrum de Blainville, 1830, and Monocoryne Broch, 1910; Candelabrum

has 14 species. The species of Candelabrum discovered at the Snake Pit is described here. It

presents surprising morphological resemblances with the species Candelabrum cocksii (VlGURS,

80°\V 60° 40° 20° 0°

Fig. 1. — Map of the northern Atlantic showing the position of the Mid-Atlantic Ridge and hydrothermal sites Snake Pit (locality

for Candelabrum serpentarii nov. sp.) and Lucky Strike [locality for Candelabrum phrygium (FABRICIUS, 1780)].

Carte de l’Atlantique Nord indiquant la position de la dorsale médio-atlantique et des sites hydrothermaux du Snake Pit

(localité de Candelabrum serpentarii nov. sp.) et de Lucky Strike [(localité de Candelabrum phrygium (Fabricius, 1780)].

— 33 —

1849), well known from intertidal areas. The species Candelabrum phrygium (FABRICIUS, 1780),

collected at Lucky Strike, is known to have a circumarctic distribution. These new data prompted

us to review the genus, while in the course of our investigation it became imperative to compare

the specimens from the Mid-Atlantic Ridge with the intertidal species. Ecological observations

were added where appropriate.

Discussion of the trophic behaviour and hypothesis about the distribution of these two hy¬

drothermal species on active sites are presented.

MATERIAL

Specimens of Candelabrum serpentarii nov. sp. were obtained from the Snake Pit hydrother¬

mal vents area (Mid-Atlantic Ridge, 23°23’N-47°56’W, 3500 m depth), first during the French

cruise Hydrosnake, submersible Nautile/N. O. Nadir, June 12-July 14, 1988, chief scientist:

Catherine MÉVEL (Université Paris-VI); second by American cruise MAR 93, submersible

Alvin/ R. V. Atlantis II and J. Charcot , June 5-30, 1993, chief scientists: Cindy VAN DOVER

(WHOI) and Aline Fiala (Université Paris-VI). The photograph of that species (Fig. 3A) orig¬

inates from the same locality and was provided by the French cruise Gravinaut, NautilefN.O.

Nadir, September 7-October 4, 1993, chief scientist: Jacques Dubois (Institut de Physique du

Globe, Paris).

The specimens of Candelabrum phrygium were obtained from the Lucky Strike hydrothermal

vents area (Mid-Atlantic Ridge, 37°17’N-32°16’W, 1626-1700 m depth), first by the American

cruise Lucky Strike, Alvin! R. V. Atlantis II, May 27-June 4, 1993, chief scientist: Charlie Lang¬

muir (Lamont-Doherty Earth Observatory); second by the French cruise Diva 2, NautilefN.O.

Nadir, June 2-July 4, 1994, chief scientists: Daniel Desbruyères and Anne-Marie Alayse

(IFREMER).

In addition, both species and their environment were observed in situ, on videotape and

on photographs taken by the submersible Nautile.

Some specimens of Candelabrum cocksii were placed at our disposal by Dr A. Castric-Fey

(Collège de France, Concarneau) and came from her private collection. Specimens of Candela¬

brum phrygium and some of the other species of Candelabrum were studied by one of us (W. V.)

in the collections of The Natural History Museum, London.

REVIEW OF THE GENUS

CANDELABRUM de Blainville, 1830

Candelabrum de Blainville, 1830: 284; type, by monotypy: Lucernaria phrygia Fabricius,

1780 (= Arum Vigurs, 1849: 90, type, by monotypy: Arum Cocksii Vigurs, 1849; Myriothela M.

Sars, 1851: 126, type, by monotypy: Myriothela arctica M. Sars, 1851; Spadix Gosse, 1853b:

125, type, by monotypy: Spadix purpurea Gosse, 1853b; Acandela Stechow, 1920: 45, type, by

monotypy and original designation: Myriothela mitra Bonnevie, 1898).

— 34 —

Description

Solitary hydroids of worm-like appearance; length varied, between 10 and 300 mm total

body length. Body divisible into three regions: a basal region (foot, hydrorhiza) serving attach¬

ment of the specimen, a blastostyle bearing region and a distal body portion, usually tapering,

having a small, circular mouth at its extreme end. Shape and development of foot differing in

the various species and with age: flattened and lobed to a varied extent or root-shaped, being

in that case a conical, pointed part of the body. Attaching filaments occur on both lobed and

root-shaped types of foot, usually with chitinous discs for attachment to solid substrates. Whole

or part of foot in some species with chitinous sheath, also including the attaching discs and

occasionally of considerable thickness. Blastostyle region only distinguishable in sexually mature

individuals, usually swollen compared to distal part of body; blastostyles are tubular to conical

prolongation of the body wall bearing male and/or female cryptomedusoid gonophores; body

cavity may continue in blastostyle; these arranged in one or several whorls or irregularly dis¬

tributed. Dioecious, but in C. cocksii and C. serpentarii nov. sp. monoecious with hermaphroditic

blastostyles. Development of larva to actinula in female gonophore, one or more may be present.

Feeding tentacles (claspers) attaching to gonophore with developing egg described for one species

(C. cocksii), may also be present on others. Distal part of body (trunk) as long as or much

longer than blastostyle bearing region, usually set with numerous small, capitate tentacles. Oc¬

casionally tentacles continue downwards into the blastostyle bearing region and/or occur on the

blastostyles. Some species have modified tentacles on blastostyle or foot. Enteron with folds of

endoderm. Cnidome composed of (usually two types of) desmonemes, haplonemes (probably

atrichous), heteronemes and stenoteles.

Remarks

The type species was considered by Fabricius (1780) to be a species of stalked jellyfishes

(Stauromedusae) and described as Lucernaria phrygia. De Blainville (1830, 1834) placed it

near the genus Sipunculus, now in the phylum Sipunculida, instituting for its reception the genus

Candelabrum. M. Sars (1851) redescribed the species as Myriothela arctica; in 1857, M. Sars

reported on the rediscovery of some of Fabricius’s material in Copenhagen and referred

FABRICIUS ’s species to Myriothela, this generic name being preferred by M. Sars (1871) and

ALLMAN (1874). L. AGASSIZ (1860-1862) regarded Candelabrum de Blainville, 1830 and Myrio¬

thela M. Sars, 1851, as being congeneric, and also recognized Candelabrum as the oldest available

name. The genus name Arum was used by ViGURS (1849) for his species Arum Cocksii, later

on redescribed by Gosse (1853b) as Spadix purpurea. Spadix Gosse, 1853(b) thus is a junior

subjective synonym of Arum Vigurs, 1849. There are only two main characteristics separating

Arum from the species of Candelabrum (cf. Stechow, 1922: 144; 1923: 38): the development

of the foot and the presence of claspers. The morphology of the foot in several species of My¬

riothela (= Candelabrum) has been studied by Manton (1940). It seems clear that a foot sur¬

rounded by a continuous and conspicuous sheath of chitinous periderm begins with the

development of such perisarc surrounding the place of contact between modified tentacles and

the substrate, the tentacles being in many instances lobed extrusions of the foot (our observations).

The chitinous perisarc, at least in C. cocksii, may ultimately fuse to form a continuous sheath

— 35 —

covering the foot, thinning out distally and merging with the thin cuticle covering the ectoderm.

It thus appears that the morphology of the foot is dependent upon the age of the specimen and

most likely also on the substrate and considering the limited number of species that has adequately

been studied, in this respect it is, in our opinion, unsuitable for generic distinction. The use of

the generic name Candelabrum has recently been reintroduced by Hand & Guilliam (1951),

PREVOT (1959) and CORNELIUS (1977).

C. cocksii so far is the only species in which the presence of claspers has been established;

they definitely seem to be absent from Candelabrum phrygium, the only remaining species in

Candelabrum of which the life history has to some extent been studied. All other species (with

the exception of course of C. cocksii) have been studied from preserved material. The presence

of claspers, considered by Manton (1940) to represent modified tentacles (for attachment) cannot

altogether be excluded in such insufficiently known species of Candelabrum. It seems illogical

to separate Arum from Candelabrum on the morphology of the foot, which is dependent upon

development and substrate, and the presence or absence of claspers, a character which cannot

be properly evaluated in all species of Arum and Candelabrum. We have therefore sunk Arum

Vigurs, 1849, into the synonymy of Candelabrum de Blainville, 1830, the latter having priority.

We thus follow CORNELIUS (1977) in referring Arum, Myriothela and Spadix to Candelabrum.

The genus Acandela Stechow, 1920, was instituted by STECHOW for Myriothela mitra Bon-

nevie, 1898, the only diagnostic character being the absence of (clavate) tentacles on the distal

portion of the body. Bonnevie’s Myriothela mitra may be based on a single specimen (the

number is not stated in the description), moreover it was obtained from deep water (2220 m),

so it probably had a rough assent in a trawl net. The possibility that the specimen consequently

was damaged, resulting in the loss of tentacles, cannot be excluded and has in fact been described

in other species of Candelabrum ( e.g. C. austrogeorgiae). We have relegated Acandela to the

synonymy of Candelabrum.

REVIEW OF THE SPECIES AND DESCRIPTIONS

Candelabrum arcticum (M. Sars, 1851)

Myriothela arctica M. Sars, 1851: 126, 131, 134 [= Candelabrum phrygium 1 (Fabricius,

1780)].

Candelabrum australe (Briggs, 1928)

Myriothela australis Briggs, 1928: 307-312, PI. 32, PI. 33 Fig. 3, PI. 34 Figs 1-4: Briggs,

1929: 244-264, Figs 1-4, Pis 42-44; Briggs, 1939: 10; Manton, 1940: 280 et seq.. Fig. 8a;

Dakin, Bennett & Pope, 1948: 208; Ralph, 1966: 158, 162.

Candelabrum australe — Hand & Gwilliam, 1951: 208.

Locality. — Found “on the lobes of the thallus of a seaweed thrown up on the sandy beach of Maroubra

Bay near Sydney, N. S. W.” (Briggs, 1928).

1. See discussion of synonymy of this species.

— 36 —

Remarks

Description and studies on gonophore development based on forty specimens from the lo¬

cality given above; no accurate depth record of the living specimens is given. Length ranging

from 4 to 30 mm, body elongated, basally cylindrical; blastostyle bearing region narrowed,

“marked by a series of well defined longitudinal furrows with finer transverse striations”. Distal

portion of body, above blastostyle region, covered by capitate tentacles. Blastostyles close to¬

gether, abundant, unbranched, elongate, cylindrical and clavate at distal extremity, there bearing

a bundle of tentacles differing from those on body by trumpet-shaped head and larger size,

having a long, slender, cylindrical stem. Male and female blastostyles on separate individuals

(dioecious). Female blastostyle with 3-4 mature and 6-8 immature gonophores on proximal por¬

tion; apex with 8-10 tentacles. Male blastostyles smaller and more numerous, some 15 being

present; apex with 6-9 tentacles. “Male and female gonophores have an apical opening repre¬

senting the velar aperture. The proximal end of the hydranth is truncated and is attached to the

substrate by a number of tentacle-like filaments which constitute the hydrorhiza. At the truncated

end of each of these short rooting processes is a small, circular, chitinous disc of dark brown

colour” (Briggs, 1928). There are no claspers. Additional specimens from Port Phillip Bay, near

Melbourne, Australia, are mentioned by Ralph (1966); length of preserved specimens 20-35 mm.

Candelabrum austrogeorgiae (Jaderholm, 1904)

Myriothela austro-georgiae Jaderholm, 1904: ii; JÀDERHOLM, 1905: 4, 6-9, 38, Pis 1-2, PI.

3 Figs 1-3; Billard, 1906: 2, 4-9, Figs 1-3; Hickson & Gravely, 1907: 19; Ritchie, 1909:

67, 69-70; Stechow, 1909: 37, 66; Vanhôffen, 1910: 272, 277, 339; Briggs, 1928: 315, 1939:

10; Manton, 1940: 281, 282; Rees & Thursfield, 1965: 45; STEPAN’YANTS, 1972: 63-64, Fig.

9, 1979: 27, PI. 5 Fig. 5, PI. 25 Fig. 2.

Candelabrum austrogeorgiae — STECHOW, 1922: 144; Stechow, 1923: 45; Hand & GwiL-

liam, 1951: 208.

Gonostyle of Siphonophore, Thompson, 1904: 19, PI. 1.

Localities. — “Vor Cumberland, Sth Georgia, 252-310 m, 5. vi. 1902; Stn 81, Bransfield Strait, 849 m,

25. xi. 1902; Stn 90, Bransfield Strait, 719-726 m, 05. xii. 1902” (Jaderholm, 1905).

Flanders Bay, Graham Land, 15. ii. 1904; Booth-Wandell Island, 26/30. ix and 28. x. 1904, low tide (Billard,

1906).

“Scotia Bay in the South Orkneys, 10 fms, iv. 1903; 9-10 fms, v. 1903; among mud and pebbles, 18. xii.

1903”. “On surface of the water, in a hole which had been cut in the ice. The depth of the water at that place

was 20-30 fathoms, the temperature was 29° F” (Ritchie, 1909).

“Observatory Bay, Kerguelen, 05. i. 1902, 10 x 4 mm” (Vanhôffen, 1910).

Davis Sea, Antarctica, near station Mir, 15-18 m; panantarctic species (Stepan’yants, 1979),

Remarks

Original description, by JÀDERHOLM (1904, 1905), based on unknown number of specimens

from South Georgia. Body worm-like, up to 300 mm long, composed of foot, blastostyle bearing

region and distal part exclusively bearing tentacles. Foot up to 20 mm long, basally with 3-4 mm

long projections bearing filaments for attachment, flattened at the tip. There is no perisarc. Blas¬

tostyle bearing region c. half length of distal region, with blastostyles and capitate tentacles.

— 37 —

Blastostyles slender, tubular, with a single terminal tentacle or a number of smaller tentacles,

bearing male or female gonophores; species is dioecious. Female gonophores usually 1-3, oc¬

casionally up to 6; male gonophores up to 10, of smaller diameter. Blastostyle bearing region

not wider than remaining, distal part of body, which tapers gradually and is covered by numerous

small capitate tentacles without noticeable arrangement.

The nematocysts are described to some extent by Billard (1906), who distinguishes two

types, viz. stenoteles and desmonemes; no measurements are given. Some additional morphol¬

ogical details are given in the description of Ritchie (1909) of specimens from the South Ork¬

neys; on these specimens Thompson’s (1904) description of the gonostyle of an unknown giant

Siphonophore is based, placed by Ritchie in the synonymy of the present species. The species

is also redescribed by Stepan’yants (1972, 1979).

Candelabrum capensis (Manton, 1940)

Myriothela capensis Manton, 1940: 276-287, Figs 7, 8b, 9, PI. 1 Figs 12, 13, PI. 3 Fig. 27;

Millard, 1957: 186, 1966: 437; Day, Field & Penrith, 1970: 12; Bouillon, 1974: 143; Mil¬

lard, 1975: 45, Figs 7D, 18A, F, G, 1978: 195 et seq., 1980: 130.

Candelabrum capensis — PREVOT, 1959: 98.

Localities. — “Aquarium Rocks, East London, Sth Africa, 17 & 19. vii. 1937, 8-17 m” (Manton, 1940).

False Bay, 34° 08.5’ S-18° 34.5’ E, 27 m, female specimen, 6.5 mm (Millard, 1957).

West coast Cape Peninsula, Kommetje, 34° 08.5’ S-18° 19.4’E, almost mature male attached to weed.

Ludertitz Bay, South-West Africa, 26° 38’ S-15° 09.3’ E, two mature male specimens and two young specimens

attached to crustacean appendage, largest 16 mm. Lamberts Bay, West coast Cape Peninsula, 32° 04.7’ S-18°

18.2' E, 17 m, mature female specimen attached to weed (Millard, 1966). According to Millard (1975) maximum

body length c. 25 mm.

Remarks

Body c. 25 mm long, cylindrical, slowly tapering from base onward; basal portion (foot)

c. one tenth of body length, attached, usually to algae, by means of 20-30 adhesive processes

capped by chitinoid discs (Millard, 1975: 45). Blastostyles up to 4 mm long in single whorl

of c. 20 above foot; unbranched, up to nine gonophores in proximal region and 4-7 capitate

tentacles in distal region; species dioecious. Female gonophores releasing up to three actinulae.

Distal part of body bearing many (400-600) densely packed, capitate tentacles. Cnidome ade¬

quately described by Millard (1966, 1975), composed of desmonemes (7.8-16.8 x 5.5-12.6

pm), stenoteles (9.9-11.4 x 7.2-8.1 pm), heteronemes (11.7-19.2 x 3.6-6.0 pm), haplonemes (10.8

x 9.9 pm) and probably also atrichous isorhizas (18.0 x 6.0 pm).

Candelabrum cocksii (Vigurs, 1849)

(Fig. 2c-d, Table 1)

Arum Cocksii Vigurs, 1849: 90.

Arum Cocksi(i) — COCKS, 1849: 90, 1852: 22, 1853a: 34, PI. 3 Figs 7-12; M. Sars, 1857:

195; Stechow, 1922: 144; Rees, 1956: 116; Marine Biological Association of the United King¬

dom, 1957: 39; Rees, 1957: 487, Fig. 39A, B; Prevot, 1959: 97, PI. 1 Fig. 1; Bruce, Colman

— 38 —

& Jones, 1963: 48; Teissier, 1965: 11; Van de Vyver, 1968: 349, Figs 16, IV, V; Fey, 1970:

390; Castric-Fey, 1970: 20; Cornelius, 1977: 521 et seq; Van de Vyver, 1980: 110, 112;

Castric & Michel, 1982: 79, Fig.

Spadix purpurea Gosse, 1853b: 126-127; COCKS, 1853b: 365.

Spadix purpurea p.p. FORBES, 1854: 31 (excl. synonyms).

Spadix cocksii — GOSSE, 1853b: 386: M. Sars, 1857: 195; G. O. Sars, 1877: 28, note.

Myriothela cocksi(i) — G. O. Sars, 1874: 96, 130, 135; Hincks, 1874: 136, 137; Storm,

1882: 7, 28, 30, at least part of specimens belong to Monocoryne gigantea (Bonnevie, 1898,

fide Swenander, 1904); Pennington, 1885: 58, PI. 3 Fig. 4; Bonnevie, 1899: 9, 31-34, 37;

Swenander, 1904: 4, 6; Jàderholm, 1905: 8; Billard, 1906: 5, 9; Broch, 1910: 194, 233,

238; Bedot, 1911: 212; Billard, 1912: 460, 1921: 12-17, Fig. 1; Benoît, 1923a: 1836-1838,

1923c: 507-510, Figs 1-4; Prenant & Teissier, 1924: 26; Benoît, 1925: 89 et seq., 113-193,

Figs 1-35; Chadwick, 1926: 51; Weill, 1926: 1244 et seq.; Billard, 1927: 513-514; Marine

biological Association of the United Kingdom, 1931: 69; Weill, 1934a, b: 44, 106, 124, 125,

355, 373-375, 444, Figs 114-116; Moore, 1937: 40; Bruce, 1939: 12; Eales, 1939: 38, PI. 3

Fig. 8; PÉRÈS, 1939: 539, PI. 25 Fig. 1; Bassindale, 1941: 147; Manton, 1941: 143 et seq..

Figs 1-2; Bruce, 1948: 45; Teissier, 1950: 11; Barrett & Yonge, 1958: 47, PI. 1; Eales,

1961: 34, PI. 3 Fig. 8, 1967: 34, PI. 3 Fig. 8; FIarvey, 1969: 14; Robins, 1969: 329; Nichols,

Cooke & Whiteley, 1971: 9, Fig.; Bouillon, 1974: 143; Hiscock, 1974: 23; Beigel, 1976:

121, Fig. 1, Pis 1-4; Beigel-Heuwinkel, 1982a: 225 et seq., 1982b: 199-210, Figs 1-22. 1984:

273, 1988: 57-66, Figs 1-12.

Candelabrum cocksi — Kramp, 1938: 66; Hand & Gwilliam, 1951: 208; Cornelius,

1977; Cornelius & Ryland, 1990: 116, Fig. 4.6.

Myriothela arctica — Wright, 1858: 433; Wright, 1859: 108; Hincks, 1861: 157-158;

Allman, 1864a: 411, 1864c: 63.

Myriothela arctica p.p. FORBES, 1854: 31 (excl. synonyms).

Candelabrum arcticum p.p. L. AGASSIZ, 1862: 341 (excl. synonyms).

Myriothela phrygia — HINCKS, 1868: 77, PI. 12 Fig. 3 (excl. synonyms in part). [Not My¬

riothela phrygia (Fabricius, 1780)].

Myriothela phrygia — ALLMAN, 1874: 317-321, 1875a: 135; BOURNE, 1889: 5, 1890: 392;

Hardy, 1891: 505 et seq., Pis 36-37; Garstang, 1894: 223; Crawford, 1895: 259 ( Myriothela

phrygia ); Gamble, 1896: 132; Browne, 1897: 243; Pruvot, 1897: 584, Tab. 22; Blackburn,

1899: 58 et seq., PI. 8; Labbé, 1899: 4 et seq., PI. 1 Figs 1, 4-9, PI. 2 Figs 13, 17, 21; Beaumont,

1900: 756, 766; Browne, 1904: 162, 188; Haeckel, 1904: PI. 6 Fig. 12; Hartlaub, 1904:

100; BOULENGER, 1908: 360; MÜLLER, 1908: 73; Boulenger, 1910: 775. [All not Myriothela

phrygia (Fabricius, 1780)].

Myriothela phrygia p.p. ALLMAN, 1872: 168, 382 (excl. synonyms in part). [Not Myriothela

phrygia (Fabricius, 1780)].

Myriothela — Allman, 1875b: 250 et seq.; (De) Korotneff, 1878: 363-365, 1879: 187-190.

Myriothela p.p. (De) Korotnev, 1880: 5-37, Figs 1-29, Pis 1-4.

Myriothèle — (De) Korotneff, 1888: 21 et seq.. Pis 1-2.

Material examined. — Three complete specimens and two damaged individuals from Glénan Islands, off

the Atlantic coast of France, collected in 1964 and 1965, in the tidal zone and upper subtidal zone; depth up to

17 m.

— 39 —

Diagnosis

Body composed of foot, blastostyle region and trunk. Foot large, only slightly shorter than

blastostyle region, with a number of slender prolongations that attach body to substrate, basis

of each prolongation with chitinous perisarcal disk; chitinous perisarc gradually extending up¬

wards and covering whole foot, externally more or less spinous. Monoecious, blastostyles slender,

with male and female gonophores and occasionally with some developing eggs attached by

claspers; dispersed capitate tentacles also occur. There is no terminal circle of tentacles. Trunk

in our specimens 15-20 mm long, collapsible, covered by small, capitate tentacles with ovoid

to globular capitulum; nematocysts dispersed.

Description

The more noticeable difference with both C. serpentarii and C. phrygium is in the develop¬

ment of the foot, which in the present species represents a conspicuous part of the body, attached

to the substrate by means of finger-shaped to lobed prolongations, that terminally have a distinct

chitinous disk, by means of which the specimens are attached to the front of algae, to calcareous

algae or to fragments of rock. The chitinous perisarc surrounding the attaching disks reaches

upwards to cover the whole of the foot with a layer of yellowish-brown perisarc, externally

rough to spinous. Development of this chitinous sheath is different in the various specimens.

The region bearing the blastostyles is c. 8 mm long; the blastostyles number 5 to 8 and are long

and thin, carrying male and female gonophores in various stages of development, the males vary

in diameter between 0.30 and 0.42 mm; the ripe female gonophore (with ovum) measures 0.60 mm

in diameter. In addition some of the blastostyles have a developing egg attached by means of

one or several claspers, resembling tentacles with a disc-shaped apical portion attached to the

hyaline egg membrane. Inside is a developing larva (actinula); diameter of whole structure

c. 1 mm. The state of preservation of the material does not permit further, more detailed obser¬

vation. The trunk portion of the body, in the present specimens, is 15-20 mm long, tube-shaped

and quite weak, which may be largely the result of inadequate fixation followed by transportation

of the specimens. No mouth could be found; the exterior of the trunk is covered with small,

capitate tentacles, pedicel 200-250 pm long, capitulum (often elongated ovoid) 130-200 pm

diameter. The nematocysts are dispersed over the exterior of the capitulum and have been studied

in squash preparations. There are two size classes of desmonemes, as well as haplonemes and

stenoteles. Large and small desmonemes are about equally abundant, ovoid and slightly asym¬

metrical because of the development of a slight elevation just besides the top of the capsule.

Large desmonemes 13.8-14.0 x 8.8-9.1 pm; small desmonemes 6.5-8.5 x 4.5-4.9 pm. They contain

a thick thread irregularly coiled inside capsule. Haplonemes present in small number, slenderer

than in the other two species and more asymmetrical, occasionally slightly banana-shaped, 18.8-

20.5 x 6.2-6.6 pm. The longitudinal portion of the thread is clearly visible but the obliquely

transverse coils are difficult to see. Stenoteles found in considerable numbers, ovoid with flattened

top, perfectly symmetrical, 9.8-10.7 x 8.2-8.6 pm; shaft visible with folded barbs inside.

Remarks

The principal area of distribution of this species is in the intertidal zone of the English

Channel coasts of Great Britain and France. It is definitely known to occur also at the Glénan

— 40 —

TABLE 1. — Synoptic table differentiating between three described species of Candelabrum

C. cocksii

C. phrygium

C. serpenter ii

Size

1-2 cm

4-40 cm

8-15 cm

Locality

Intertidal zone of NE At¬

lantic.

Circumarctic and at active

hydrothermal vents in the

NE Atlantic in water of

c. 11°C loaded with sulfu-

reous compounds.

Restricted to an area

around hydrothermal

vents in water of 2.4° C.

Food

Probably small Crustacea

(Copepoda and Amphipoda).

Shrimps and small Crustacea

(Amphipoda, Copepoda)

Probably Crustacea

Reproduction

Monoecious; claspers pre¬

sent; young polyp devel¬

oping into actinula;

dispersal consequently re¬

stricted.

Dioecious; no claspers ob¬

served; development of

young polyp unknown.

Monoecious; no claspers

observed; development

of young polyp unknown.

Nematocysts*

Desmonemes (in pm)

13.8-14.0 x 8.8-9.1 (large)

6.5-8.5 x 4.5-4.9 (small)

12.5-13.0 x 9.0-9.8

(large)

8.2-9.0 x 6.4 x 6.6

(small)

13.0-14.5 x 9.8-10.5

(large)

small type not observed

Haplonemes (in pm)

18.8-20.5 x 6.2-6.6

19.7-20.5 x 8.2-9.9

16.5-18.0 x 7.8-8.2

Stenoteles (in pm)

9.8-10.7 x 8.2-8.6

10.6-11.5 x 8.2-9.8

9.8-11.5 x 9.5-10.5

* It should be borne in mind that all observations in the present material are based on observations of unexploded

nematocysts; the identifications of the various types, particularly the haplonemes, are tentative.

Archipelago (where some individuals were observed in May 1994) in the northern Bay of Biscay,

at the Scilly Islands, at the Isle of Man and in the Bristol Channel. The records from Norway

(e.g. Kramp’s, 1938, record from “Norway S. of Lofoten”) are exclusively based on G. O. Sars’s

remarks on the occurrence of this species in deep water (100-200 m) off Aalesund, Norway;

there are no recent records from the Norwegian coast. However, one of us (W. V.) has seen an

undubitable specimen in an intertidal collection from the Bay of Cadiz, Atlantic coast of southern

Spain, made by Dr M. D. Medel, Huelva, Spain.

It is quite a problem to state accurately the geographical distribution of this, apparently

not quite rare, intertidal species because of its frequent confusion with Candelabrum phrygium.

Since 1874, the differences between both species have been pointedly worded by G. O. Sars;

his publication in Norwegian apparently escaping the notice of many later scientists. The con¬

fusion resulting from the tangled synonymy of the two species unfortunately has been aggravated

by Cornelius’ 1977 paper, in which both species are considered conspecific and the fact is

overlooked that Rees (1956) was fully aware of their specific differences.

(De) KOROTNEV’s (1880) lengthy Russian description of Myriothela refers partly to Can¬

delabrum cocksii which he studied at Roscoff; many details and some of the drawings have

41 —

Fig. 2. — a, b, nematocysts of Candelabrum serpentarii nov. sp. (a, desmoneme and haploneme; b, six desmonenes, three stenoteles,

one out of focus); c, d, nematocysts of Candelabrum cocksii (VlGURS, 1849) (c, four haplonemes, one large, five small

desmonemes, and one stenotele; d, eleven large and two small desmonemes, two stenoteles, one partly); e, f, nematocysts

of Candelabrum phrygium (Fabricius, 1780) (e, stenotele and haploneme; f, eight large and three small desmonemes, one

stenotele). All nematocysts have been photographed with the aid of Nomarski interference contrast; x 750.

a, b, nématocystes de Candelabrum serpentarii nov. sp. (a, desmonème et haplonème; b, six desmonèmes, trois sténotèles,

un hors cadre); c, d, nématocystes de Candelabrum cocksii (VlGURS, 1849) (c, quatre haplonèmes, un grand et cinq petits

desmonèmes, un sténotèle; d, onze grands et deux petits desmonèmes, deux sténostèles, un vu partiellement); e, f, nématocystes

de Candelabrum phrygium (Fabricius, 1780) (e, sténostèle et haplonème; f, huit grands et trois petits desmonèmes, un sté-

nostèle). Tous les nématocystes ont été photographiés à l’aide du microscope à contraste interférentiel Nomarski; x 750.

— 42 —

been taken from Allman’s (1876) paper. It is not clear whether or not Naumov’s (1960: 241-242,

Figs 130, 131) notes on Myriothela phrygia refer to his own observations or those listed by

(De) KOROTNEV. Certainly does NAUMOV’s figure 130 refer to Candelabrum cocksiv, it is taken

from one of (De) Korotnev’s plates and the claspers are distinctly visible. Naumov’s figure

131, as he indicates, is taken from Allman (1876) and refers to the actinula of Candelabrum

cocksii [we have tacitly assumed that the major portion of detailed observations on Myriothela

phrygia in Naumov’s paper is based on solid observations (remarks on geographical distribution,

bathymetrical record, life cycle, etc.)].

Candelabrum giganteum (Bonnevie, 1898)

Myriothela gigantea Bonnevie, 1898: 468, 490-491, PI. 27 Figs 46-47; BONNEVIE, 1899: 9,

11, 37, 38, PI. 4 Fig. 1; Jâderholm, 1905: 7; Broch, 1910: 194, 233, 236; Manton, 1941: 143.

Candelabrum giganteum — STECHOW, 1922: 144, 1923: 45; Hand & G WILLI AM, 1951: 208.

Locality. — Based on two more or less complete specimens and several fragments from deep water

(2195 m) of the North Atlantic, 75°12’N-03°20’E.

Remarks

Body elongated and thin, c. 300 mm long, basally thickest and slightly swollen, gradually

tapering distally and terminating in fine filament. Below swollen or thickened basal portion a

short, pointed foot with fine attaching filaments (no perisarc mentioned in original description).

Blastostyles distributed over lower half of body; female blastostyles 10-20 mm long, with 1 or

2 big gonophores; male blastostyles shorter, 6-7 mm with many gonophores; both female and

male gonophores have some terminal tentacles. Clavate tentacles on upper half of body, ap¬

parently also occurring in small numbers between blastostyles. Cnidome unknown. Curious fila¬

mentous appearance of distal part of body may be the result of inadequate fixation.

The species has not been rediscovered since the original description by Bonnevie of speci¬

mens collected by the Norwegian North Atlantic Expedition 1876-1878.

Candelabrum harrisonii (Briggs, 1928)

Myriothela harrisonii Briggs, 1928: 312, Fig. 1, PI. 33 Figs 1-2, PI. 34 Fig. 5; BRIGGS,

1930: 5-14, Fig. 1, Pis 1-3, 1931: 270-278, Figs 1-3, 1939: 10.

Candelabrum harrisoni — HAND & GWILLIAM, 1951: 208.

Locality. — Characterized as a “shallow water form, on underside of rocks below low-water mark at

Bulli, 40 miles south of Sydney, N. S. W.” (Briggs, 1928); number of individuals not stated.

Remarks

Body cylindrical, elongated, divisible into foot, blastostyle bearing region and distal trunk.

Foot set transversally towards length axis of body, with slender rooting processes, covered by

translucent, chestnut-brown perisarc. Distal region of body cylindrical, slightly narrowing towards

— 43 —

blastostyle bearing region, with many capitate tentacles, diminishing in number towards blas-

tostyle bearing zone. Blastostyles in single transverse row on swollen blastostyle region; that

portion of body with fairly deep longitudinal furrows and fine transverse striae. Blastostyle with

irregularly lobed base and a small number of gonophores (2-3 mature and 3-4 immature male

gonophores; number in female unknown but less) with apical opening representing velar aperture.

There is a single apical tentacle per blastostyle, often pushed aside by the developing gonophore.

The size of the specimens is nowhere given in the description, nor can it be deduced from the

figures. It probably had about the same size as Candelabrum australe with which it was simul¬

taneously described.

Nematocysts described by BRIGGS (1930): desmonemes 10-12 x 8-9 pm; haplonemes 15-21 x

6-9 pm.

Candelabrum meridianum (Briggs, 1938)

Myriothela meridiana Briggs, 1938: 9-10, PI. 15 Fig. 3; MILLARD, 1971: 399-401, Figs 1-2;

Stepan’yants, 1979: 26-27, PI. 4 Fig. 4.

Localities. — “Six specimens 12-30 mm high attached to stones below low water, Macquarie Island”

(Briggs, 1938). “Eight perfect or near-perfect individuals and about nine damaged specimens and fragments from

below rock in the littoral region in Transvaal Cove, Marion Island”, up to 27 mm long (Millard, 1971).

Remarks

Original description by Briggs, 1939, supplemented by Millard, 1971. Species with ten¬

dency for subdivision of basal part of body, bi- or tripedal, fusion to single tubular distal region

at c. one-third of height; body covered with small, capitate tentacles, continuing downward into

blastostyle region at lower end of body. Base of body naked, occasionally ridged, “attached to

substratum by a number of short adhesive tentacles each capped by a flat disc of brownish

perisarc” (Millard, 1971). Blastostyles closely set, c. 2 mm long, occasionally branched once

or twice, bearing 3-10 oval gonophores. Male gonophores sessile; female gonophores with short,

thick stalk, larger than males, largest with 4-5 actinulae. Dioecious species. Cnidome described

in detail by Millard (1971), composed of two size classes of oval desmonemes (11.3-14.9 x

8.7-11.3 and 6.2 x 5.2 pm), microbasic euryteles (14.9-16.5 x 5.7-6.1 pm), and other heteronemes

(possibly stenoteles, 10.4-10.8 x 6.2-7.2 pm).

Candelabrum minutum (Bonnevie, 1898)

Myriothela minuta Bonnevie, 1898: 468, 489-490, PI. 27 Fig. 44; Bonnevie, 1890: 9, 35,

37, PI. 3 Fig. 6a, b, PI. 4 Fig. 4; JÀDERHOLM, 1905: 7; Broch, 1910: 194, 233, 236; Manton,

1941: 143.

Candelabrum minutum — Stechow, 1922: 144, 1923: 45; Hand & Gwilliam, 1951: 208.

Locality. — Tromso, northern Norway; no depth record.

— 44 —

Remarks

Based on unknown number of specimens from Trornso, where the species was found by

M. Sars, apparently in the middle of the last century. Body small, cylindrical, thickest in middle

(c. 2 mm), basally with pointed foot bearing attaching filaments; capitate tentacles only found

in small number on extreme distal part of body and surrounding mouth. Blastostyles small, oc¬

curring on major part of body, bearing a single large (female) gonophore, the latter with a small,

rudimentary tentacle. Cnidome unknown. The species has not been rediscovered since the original

description.

Candelabrum mitra (Bonnevie, 1898)

Myriothela mitra Bonnevie, 1898: 468, 489, PI. 27 Fig. 43; BONNEVIE, 1899: 9, 11, 33,

37, 38, 40, PI. 3 Fig. 6c-e, PI. 4 Fig. 3; Jàderholm, 1905: 7; Manton, 1941: 143.

Acandela mitra — STECHOW, 1920: 45, 1922: 144, 1923: 47.

Candelabrum mitrurn — Hand & GWILLIAM, 1951: 208.

Locality. — Based on a (male ?) specimen (or specimens) from deep water (2222 m) of the North Atlantic

(63°22’N-05°29’W).

Remarks

Body conical, basally widest, there 10 mm diameter, tapering distally, there 1-2 mm. Basal

part of body with narrowly pointed, 10-20 mm long foot bearing rooting filaments; no perisarc

described. Blastostyles on basal third to fourth of body, in many irregular whorls. Blastostyles

conical, top with several capitate tentacles, curved; gonophores dispersed over blastostyle. There

are no tentacles on rest of body.

The atentaculate condition of the body occasioned Stechow (1920: 45) to institute a separate

genus, Acandela, for its reception. This atentaculate condition may well result from damage

sustained by the specimen studied by Bonnevie. The number of specimens available to Bonnevie

is not unambiguously stated and may very well have been one single specimen, obtained in a

haul from great depth. Moreover, in her 1898 paper Bonnevie complains about the bad preser¬

vation of her specimen(s): “Das Ektoderm des Polypen hat eine eigenthiimliche Struktur; und

ich beklage, dass seine Konservirung nicht gut genug ist, um eine genauere Untersuchung zu

gestatten, etc.” (: 489). Loss of tentacles due to damage is also described by Jàderholm (1905)

for Myriothela (= Candelabrum) austrogeorgiae.

Candelabrum penola (Manton, 1940)

Myriothela penola Manton, 1940: 256-276, Figs 1-6, PI. 1 Figs 10, 11, 14, PI. 2 Figs 15-21,

PI. 3 Figs 22-26, 28, PI. 4 Figs 29-34; BOUILLON, 1974: 143.

Candelabrum penola — BOUILLON, 1974: 143.

Locality. — Based on two specimens, a mature female 850 mm long and an immature male of 55 mm

body length, both found attached to the axis of a pennatulid and found floating alongside the research vessel

Penola in a creek of the Argentine Islands, Graham Land, Antarctica.

— 45 —

Remarks

Manton describes the species as being dioecious. The following notes are based on the

female specimen. Basal sixth of body, c. 100 mm long with a diameter of 12 mm, without ten¬

tacles and bearing numerous lobed blastostyles. Adhesive tentacles (rooting filaments) spring

from basal part of body and some of proximal blastostyles and attach polyp to substrate; they

are capped by a chitinoid disk; there is no perisarc. Female blastostyles 20-25 mm long, irregu¬

larly lobed or branched, with short, capitate tentacles (and on proximal blastostyles with some

adhesive tentacles). Gonophores distributed over blastostyle, numbering up to 10; usually only

one develops to maturity and is then quite large, 7.2 mm in diameter, considerably swollen by

development of large actinula. Distal five-sixths of body, length c. 650 mm, diameter at oral

end 7.5 mm, is covered by 0.5-2.5 mm long capitate tentacles; number estimated by Manton

at about 330,000.

The male specimen is quite young and will not be described here. The nematocysts are

described by Manton and consist of desmonemes of variable size (9-18 pm), haplonemes (13

x 9 pm) and heteronemes (10-18 x 7-13 pm).

Candelabrum phrygium (Fabricius, 1780)

(Figs 2e-f, 3E-F, Table 1 )

Lucernarici phrygia Fabricius, 1780: 343; Gmelin, 1791: 3151.

Candelabrum [phrygium] — DE Blainville, 1830: 284, 1834: 318.

Candelabrum phrygium — L. AGASSIZ, 1862: 341; ALLMAN, 1864b: 358 ( Candelabrum

Phrygia ); A. Agassiz, 1865: 186, 225, 226; Stechow, 1922: 144; 1923: 45; Kramp, 1932a: 5,

26, 1932b: 68, Tab. 1; 1943: 42; Hand & Gwilliam. 1951: 208; Cornelius, 1977: 521 et seq.;

Stepan'yants, 1985: 85; Antsulevich, 1987: 27; Stepan yants, 1989: 412 et seq.; Cornelius

& Ryland, 1990 116; Antsulevich, 1991: 40; Cairns et al. , 1991: 16.

Myriothela phrygia — G. O. Sars, 1873: 86, 119; HlNCKS, 1874: 136; G. O. Sars, 1874:

130, 140-142; Lütken, 1875: 188; G. O. Sars, 1877: 26, note; M. Sars, 1877: 23, PI. 2 Figs

29-36; StORM, 1879: 27; D’Urban, 1880: 255, 257, 258; HlNCKS, 1880a: 257; Storm, 1880:

122; Winther, 1880: 270; Storm, 1882: 8, 28, 30; Allman, 1888: xxi, xliv; Driesch, 1890:

154; Hardy, 1891: 505-537, Figs 36-37; Levinsen, 1893: 150; Vanhôffen, 1897: 245; Bon-

nevie, 1898: 491, 1899: 9, 11, 31, 33, 35, 37, 38, PI. 4 Figs 5-6; Blackburn. 1899: 58-63,

PI. 8; Whiteaves, 1901: 20; Broch, 1903: Tab.; Swenander, 1904: 4-6; Stephens, 1905: 40;

Billard, 1906: 5; Jàderholm, 1908: 192, 233, 237; Broch, 1910; 192, 233, 237; Deryugin,

1915: 304; Broch, 1916: 19-21, PI. 1 Figs 3, 8; Hartlaub, 1916: 110, Figs 38-39; Fraser,

1918: 332, 341, 1921: 148, Fig. 18; Svarchevskii, 1923: 99; Chadwick, 1926: 51; Manton,

1941: 143; FRASER, 1944: 88-89 [not PI. 15 Fig. 63 = Candelabrum cocksii (Vigurs, 1849)];

Berezina, 1948: 50, PI. 14 Fig. 1; Rees, 1956: 116; Naumov, 1960: 241-243, Figs 130-131;

Calder, 1972: 222, PI. 1 Fig. 5; Campbell, 1974: 151, Fig. 9D; Petersen, 1990: 203.

Myriothela phrygia p.p. Allman, 1872: 382 (excl. synonyms).

Corymorpha phrygia — MÔRCH, 1857: 24.

— 46 —

Myriothela arctica M. Sars, 1851: 126, 131, 134; ALDER, 1853: 35 ( Myristhela phrygia)',

Gosse, 1855: 20, Fig. 25; M. Sars, 1857: 192, 194; Wright, 1858: 433, 1859: 108; M. Sars,

1860 (German translation): 342; Hincks, 1861: 157; M. Sars, 1861: 693; Allman, 1864a: 411,

1864c: 63; Parfitt, 1866: 5; Verrill, 1879: 19; G. O. Sars, 1877: 26, note.

Myriothela arctica p.p. FORBES, 1854: 31 (excl. synonyms).

Candelabrum arcticum — Allman, 1864b: 358.

Candelabrum arcticum p.p. L. AGASSIZ, 1862: 341 (excl. synonyms in part).

Material examined. — All specimens were obtained from the Lucky Strike hydrothermal vents area:

— two from the Lucky Strike cruise (site Sintra, marker 3, 37°17.50’N-32°16.47’W, 1622 m depth; dive

2606, June 1st, 1993; collected by Meg Tivey); well preserved male specimens, both with developing male gono-

phores on blastostyles; one specimen attached to rock fragment (Fig. 3F), the other to a 20 mm long living

specimen of mussel Bathymodiolus sp;

— seven from the Diva 2 cruise:

— two fixed and preserved in formalin (one female with large, mature eggs attached to blastostyles), ob¬

tained at dive PL 02, site Sintra, June 4, 1994; collected by Philippe Crassous;

— two fixed in Bouin and tranferred to ethanol 70%, from dive PL 09, site Tour Eiffel (37°17.31’N-

32°16.5rW; 1690 m depth), June 11, 1994; collected by Luis Saldanha. Also a basal part of a male specimen

with many blastostyles with mature male gonophores, attached to a sulfide rock, and two parts of male specimen,

the basal part with rock fragments and some developing gonophores, the second part a segment of the trunk

with a complete, more or less digested shrimp' inside. The organic debris found in association with the three

specimens of Candelabrum phrygium contains remnants of an amphipod, of a small shrimp, of a calanoid copepod,

a complete healthy looking Ectinosomid (Copepoda Harpacticoida) and some unidentifiable animal remains;

— three (two females and a male: a 40 mm long female, eggs developing in gonophores; also a 65 mm

long female with some large gonophores and rest of amphipod attached to tentacles; in addition, c. 50 mm long

male, proximal part of trunk decomposed; blastostyle bearing region about as long as rest of trunk, with a great

number of blastostyles bearing many developing and mature gonophores) preserved in formalin, from dive PL

10, site Tour Eiffel, June 12, 1994; collected by Marie-Claire Fabri.

2. This is probably a specimen of a newly described crustacean Chorocaris fortunata (MARTIN & CHRISTIANSEN, 1995,

L. B. Holthuis, pers. comm.); TL c. 10 mm. diameter 2-3 mm.

Fig. 3. — A: Candelabrum serpentarii nov. sp. (length c. 10 cm). Snake Pit area, next to Élan site, 3515 m, on sulfide rock,

surrounded by polychaete tubes; B: Candelabrum serpentarii nov. sp. (length c. 8 cm). Snake Pit area, next to Les Ruches

site, 3505 m, on sulfide rock, surrounded with tubes of polychaeta Chaetopteridae and Zoantharia (Cnidaria); C: Candelabrum

serpentarii nov. sp. (paratype, length c. 7.5 cm), Snake Pit area, next to Les Ruches site, 3523 m, on pillow lava; D: The

same specimen, freshly collected, natural colours (however, one can notice the difference of colour with in situ organism of

photo A); the black spots are sulfidic metal particles precipitated on the animal in the slurp gun box; E: Candelabrum

phrygium (Fabricius, 1780), length 7-9 cm. Lucky Strike area, La Pagode site, 1626 m, on flange mineral formation covered

with white silica; one mussel (Bathymodiolus sp.) is visible; F: Candelabrum phrygium (Fabricius, 1780), Lucky Strike area,

Sintra site, 1622 m, on flange block.

A; Candelabrum serpentarii nov. sp. (L = env. 10 cm), zone du Snake Pit, près du site L’Élan, 3515 m, sur un bloc de

sulfure, entouré de tubes de polychètes; B: Candelabrum serpentarii nov. sp. (L = env. 8 cm), zone du Snake Pit, près du

site des Ruches, 3 505 m, sur un bloc de sulfure, entouré de tubes de polychètes Chaetopteridae et de zoanthaires (Cnidaires);

C: Candelabrum serpentarii nov. sp. (paratype, L = env. 7,5 cm), zone du Snake Pit, près du site des Ruches, 3 525 m, sur

des laves en coussin; D: le même spécimen, fraîchement récolté, couleurs naturelles (on notera toutefois la différence de

couleur avec l'organisme in situ de la photo A); les taches noires sont des particules de sulfures métalliques précipités sur

l’organisme dans le collecteur du système d'aspiration; E: Candelabrum phrygium (FABRICIUS, 1780), L - 7-9 cm, zone de

Lucky Strike, site de La Pagode, 1626 m, sur formation minérale “flange", couverte de silice blanche; une moule (Bathy¬

modiolus sp.) est visible; F: Candelabrum phrygium (Fabricius, 1780), zone de Lucky Strike, site Sintra, 1622 m, sur un

bloc de “flange".

— 48 —

Diagnosis

Body composed of foot, blastostyle bearing region and trunk. Foot: a lobed, flattened part

of body, attaching animal to rock or substrate (bivalve molluscs), without chitinous adhesive

portion. Blastostyle bearing region forming lower third to fourth of body, with a large number

(10 to 15) of big, tubular blastostyles, bearing male gonophores in various stages of development

and dispersed, capitate tentacles; apex of each blastostyle with circle of 4 or 5 tentacles. Re¬

mainder of body forming elongated, tubular trunk, completely covered by capitate tentacles.

Mouth distinct, at end of trunk.

Description

All specimens available are attached to rock fragments, one male is attached to the exterior

of a living bivalve. In all specimens the foot attaches the body to the substrate; there are no

chitinous adhesive disks or chitinous portions of the foot. In the specimen on the mollusc, the

foot is more distinctly lobed than in the others. Directly above the foot is the blastostyle bearing

part of the body, 12-15 mm high and bearing a considerable number (10 to ca. 50 in the male,

usually less in the female) of tube-shaped blastostyles, the body cavity continuing into the blas¬

tostyles. Male and female blastostyles are found on separate polyps; the species consequently

is dioecious. Each male blastostyle is 8-10 mm long and carries many developing male gono¬

phores, with dispersed, capitate tentacles in between. The apex of each blastostyle carries a

circle of 4 or 5 capitate tentacles. Male gonophores 0.40-0.80 mm in diameter, attached by means

of thin tissue strand and easily detached in the preserved specimens, apparently filled with

developing spermatocytes. The female blastostyles are shorter and thinner than the males and

have less gonophores. In the specimens inspected each blastostyle has gonophores in various

stages of development, the youngest in the basal part, the mature gonophores at the top; there

are only few tentacles. The smallest gonophores measured are 0.40 mm, the largest, apparently

mature gonophores are 1.40 mm. The gonophores that have been supposed to be mature contain

a single, large egg; it is attached to the blastostyle by means of a thin strand of tissue, being

the continuation of a thin layer of tissue covering the egg. Trunk vermiform, diameter c. 2.5 mm,

in the preserved specimens 25-30 mm long, completely covered by short, capitate tentacles. Each

tentacle has a 0.35-0.50 mm long pedicel and a capitulum of 0.17-0.25 mm diameter; nematocysts

dispersed over capitulum.

Nematocysts studied in squash preparations of capitulum of trunk tentacles, composed of

two size classes of desmonemes, haplonemes and stenoteles.

Haplonemes slightly longer than those of C. serpentarii , but of the same general shape:

elongated ovoid and slightly narrowed apically, as a result more or less pyriform, 19.7-20.5 x

8.2-9.9 pm, found more frequently than in capitulum of C. serpentarii. Internal structure fairly

obscure, but a longitudinally descending shaft and oblique coils of the thread could be observed.

Desmonemes of two size classes occur in profusion, the larger being almost identical with

those of C. serpentarii , broadly ovoid, slightly asymmetrical apically and there with a slight

elevation just outside middle of top, 12.5-13.0 x 9.0-9.8 pm. The smaller type is slightly more

elongated, 8.2-9.0 x 6.4-6.6 pm. Both types with a thick thread in irregular coils; in desmonemes

in perfect lateral view part of thread parallel to internal wall of basal portion. Both types were

found to occur in almost equal numbers.

— 49 —

Stenoteles scarce and apparently badly preserved, as internal structure was quite obscure;

they could best be recognized by the flattened top (when in good position), 10.6-11.5 x 8.2. x

9.8 pm. Details of shaft and thread could not be discerned. It was difficult to estimate the oc¬

currence of stenoteles because of bad preservation: they could only with certainty be distinguished

from desmonemes when in good position to see apical flattening.

Remarks

The various locality records are not specified here. The species is circumarctic, occurring

in both Atlantic (Bonnevie, 1898-99: Norwegian North Atlantic Expedition, Stn 303, 75°12'N-

03°02'E, 2195 m) and Pacific (Naumov, 1960) parts of the Arctic seas. In the Atlantic at least

it also penetrates boreal waters, though there usually at greater depths. It is now also known to

occur in deep water of the NE Atlantic.

The Lucky Strike specimens were generally found at the base of the active edifices, attached

to organic support (shell of living bivalve) or, more frequently, to mineral formations called

“flanges” and composed of a mixture of pyrite, marcasite and baritine, with sometimes some

white silica at the surface. The specimens are frequently observed at the site Sintra , but they

occur also at the site Tour Eiffel. At the Pagode site 7 or 8 individuals were observed together

on 0,5 m 2 (Fig. 3E). At that locality, a few animals (mussels, crabs and shrimps) are present,

but generally the surrounding faunal community is largely composed of bivalved Mytilidae

(Bathymodiolus) (Van DOVER et ai, submitted), shrimps (Alvinocarididae), crabs (Bytho-

graeidae) and several smaller Crustacea (Copepoda Siphonostomatoida, Amphipoda) living in a

mixture of sea water and hydrothermal fluid (loaded with sulfurous compounds), mean tempera¬

ture ca. 11° C. Other animals observed at the Lucky Strike hydrothermal vents area include

sponges ( Cladorhiza sp.) and hydroids (Eudendrium sp.), both found on the top of the inactive

edifices. In many cases these animals carry white, filamentous bacteria.

Candelabrum serpentarii nov. sp.

(Figs 2a-b, 3A, B, C, D, 4, Table 1)

Material examined. — One complete specimen, total body length 75 mm, and a 40 mm long upper part

of body of second specimen, both from the Hydrosnake cruise at the Snake Pit hydrothermal area (site Les

Ruches , 23°23’N-47°57’W, 3489 m depth, dive HS 10, June 28, 1988; collected by M. S.). Specimen with large

female and smaller male gonophores and presently slightly deteriorated because of frequent inspections; chitinous

covering of foot lost; remains in National Museum of Natural History, Leiden (paratype, RMNH Coll. No 27111).

Well preserved specimen in three parts, total body length c. 80 mm, composed of basal body region with

gonophores (in two parts) and trunk, from the MAR 93 cruise next to the site Elan in the Snake Pit area, about

30 m to the west of the site Les Ruches (dive 2619; 3525 m depth, June 20. 1993; collector: Jean-Paul Truchot).

This specimen was fixed in Bouin and later on transferred to ethanol 70%, now preserved in Muséum national

d'Histoire naturelle, Paris (holotype, MNHN Hy No 1133).

Etymology. — serpentarii , from the Latin serpentarium, snake pit.

Diagnosis

Large Candelabrum; total body length 60-80 mm, attached to solid substrate by means of

laterally flattened, basal foot covered by thick, curved, brownish-black perisarcal sheath. Rest

— 50 —

of body composed of proximal region bearing large blastostyles each bearing a number of male

and female gonophores in various stages of development, and an elongated trunk completely

covered with short, capitate tentacles. Capitulum of each tentacle with distal layer of nematocyst

bearing ectoderm. Cnidome (of capitate tentacles) composed of desmonemes (predominant), large

haplonemes and two size classes of stenoteles, of which the smaller class has only occasionally

been observed.

Description

Since none of the specimens is fully intact, parts of the following description are the result

of reconstruction.

The c. 60 mm long body can be divided into a foot, a blastostyle bearing region and the

apical trunk. The foot has only been observed in the paratype (Fig. 2), where it is a laterally

flattened, rounded portion of the body, attaching the polyp to the hard substrate (pillow lava)

by a considerable portion of its surface. The foot was covered by a thick, brownish-black, c. 1 mm

thick perisarcal shield, at first firmly attached to the foot, but later on spontaneously loosening

itself. No perisarcal threads with or without apical button, attaching the foot to the rock have

been observed.

The blastostyle bearing zone of the body has about the same diameter as the trunk (c. 7 mm),

the body cavity continues in that zone and in the blastostyles, that represent lateral, tentacle

bearing elevations of the body wall. Blastostyles 3-5 mm long, with c. 10 male and 3-5 apparently

fully mature female gonophores. The tentacles observed on the body between the blastostyles

as well as those on the blastostyles (between the gonophores and at the apex) are indistinguishable

from those on the trunk. Male gonophores c. 1-1.5 mm in diameter, apparently filled with develop¬

ing spermatocytes. The female gonophores are large, 4-5 mm in diameter, attached to the bla¬

stostyle by means of a short, stubby neck and surrounded by an opaque layer of ectodermal

cells; no nematocysts having been observed. The large egg can be dislodged by carefully cutting

the ectodermal covering; the egg in the well preserved specimen from the Snake Pit appeared

to be entirely filled with yolk with many fat goblets and was surrounded by a hyaline membrane.

No structure could externally be observed and it is presumed that the eggs, at least in this speci¬

men, are still unfertilized. No traces of claspers or aberrant tentacles could be found in the

blastostyle bearing zone.

The trunk is a tube-like expansion of the body externally fully covered by capitate tentacles.

The lumen of the gastral cavity is considerable; the mouth at the end of the trunk is closed.

The tentacles consist of a short, 0.6-0.8 mm long stalk and a globular capitulum of 0.4-0.6 mm

diameter; the apex of the capitulum is covered by a semiglobular layer of ectodermal cells with

FlG. 4 — Candelabrum serpentarii nov. sp., holotype, from Snake Pit area; a, distal part of body, completely covered with small,

capitate tentacles; b, close up of the capitate tentacles; c, proximal part of body with capitate tentacles on distal zone and

blastostyles, with male and female gonophores, as well as capitate tentacles, on the basal zone; d, close up of one of the

gonophores.

Candelabrum serpentarii nov. sp., holotype, provenant du Snake Pit; a, partie distale du corps, complètement recouvert de

petits tentacules capités; b, détail des tentacules capités; c, partie proximale du corps avec des tentacules capités au niveau

de la zone distale et des blastostyles, avec des gonophores mâles et femelles sur la partie basale; d, détail des gonophores.

— 52 —

many nematocysts. The tentacular stalks, as can be seen from the photographs of living specimens,

are hightly contracted in the preserved specimens, the capitulum standing off a distance of several

mm from the body surface.

The nematocysts have been studied in squash preparations of the capitulum of tentacles of

the trunk. There appear to be three types of nematocysts: haplonemes, stenoteles and desmonemes.

Haplonemes elongated ovoid, slightly narrowed distally and as a result also a bit pyriform

and slightly asymmetrical, 16.5-18 x 7.8-8.2 pm, rather uniform of size and shape. They have

not been found in exploded condition, consequently the characters of the thread could not be

studied. In unexploded condition the thread is seen to descent from the apex down to the bottom

in a straight line; the rest of the thread is coiled in obliquely transverse coils.

Stenoteles almost globular, in perfect position observed to be broadly ovoid, perfectly sym¬

metrical, with indistinctly flattened top. Shaft and barbs forming a central column in the unex¬

ploded capsule and about three-fourth the height of the capsule; thread in indistinct transverse

coils in the basal third of the capsule. Size 9.8-11.5 x 9.5-10.5 pm. A second type of stenotele

has occasionally been observed, but only in exploded condition, the capsule measuring 5.4 x

7.4 pm. Details of barbs and thread could not be observed.

Desmonemes broadly ovoid, but for the apical protrusion almost perfectly symmetrical, 13-

14.5 x 9.8-10.5 pm. Apical protrusion low, slightly besides middle of apex; thread thick, forming

a small number of loose coils following the internal desmonemal wall.

In the tentacular capitulum the desmonemes predominate; haplonemes and (large) stenoteles

forming c. 5 % of the capsular number. The smaller stenoteles have only occasionally been ob¬

served.

Remarks

In the Snake Pit field the occurrence of this species is restricted to an area of several

meters from an active structure and it is not observed outside a radius of some twenty or thirty

meters. The species may thus be considered to indicate the proximity of hydrothermal activity,

though one individual has been observed at inactive vents. Depending upon the proximity of

the active sites the specimens are attached to pillow lava (Fig. 3C) or sulfide rocks (Fig. 3A-B).

They are quite irregularly distributed, occasionally 2 or 3 individuals are found within one meter

distance. At this level there are probably no thermal anomalies, the temperature at those abyssal

depths being generally 2.4° C. Surrounding fauna is scarse (Segonzac, 1992), composed of some

fishes (Pisces Synaphobranchidae), galatheides (Crustacea Galatheidae), and occasionally one or

two isolated shrimps (Rimicaris exoculata), Williams & Rona, 1986 (Crustacea Alvinocarididae).

Candelabrum tentaculatum Millard, 1966

Myriothela tentaculata Millard, 1966: 437-440, Fig. 2; BOUILLON, 1974: 143; MILLARD.

1975: 46-48, Figs 17C, 18B-E; 1978: 195 et seq., 1979: 134.

Locality. — Based on five specimens from off Slangkop on west coast of Cape Peninsula, 34°09.3’S-18°

17.5’E, 24 March 1959, 43 m depth.

-53-

Remarks

Body up to 31 mm long, attached to encrusting Bryozoa by means of nine short, adhesive

processes capped by chitinoid discs springing from irregularly shaped basal portion. Blastostyles

in single whorl of 17, reaching 20 mm length, with c. 25 rather poorly developed capitate tentacles

on distal region and scattered amongst 4-6 gonophores on proximal 5 mm. Only male gonophores

known, species apparently dioecious. Rest of body densely covered with capitate tentacles. Cni-

dome adequately described by Millard (1975) and composed of heteronemes (up to 45 pm

long!), two size classes of desmonemes, stenoteles and atrichous isorhizas.

Candelabrum verrucosum (Bonnevie, 1898)

Myriothela verrucosa Bonnevie, 1898: 468, 490, PI. 27 Fig. 45; BONNEVIE, 1899: 9, 37,

PI. 4 Fig. 2.

Candelabrum verrucosum — STECHOW, 1922: 144, 1923: 45; HAND & GwiLLIAM, 1951: 208.

Locality. — No locality mentioned in original description (Bonnevie, 1898), but in Bonnevie’s 1899 paper

the species is mentioned in a table and Hammerfest (Norway) is given as the locality; there is no (definite)

depth record.

Remarks

Small species, total body length c. 40 mm, diameter 1-2 mm, thickest just under mouth.

Attached by means of attaching filaments springing from basal part of body. Lower fifth of

body bearing blastostyles, rest of body covered with capitate tentacles. Blastostyles short, with

one or two gonophores and with tentacles distally. Gonophores with dispersed clusters of ne-

matocysts over their surface. The species may be based on a single specimen (number of speci¬

mens or variability not stated); it should be recognizable by the clusters of nematocysts on the

gonophores, though according to Bonnevie these are difficult to perceive !

Candelabrum sp. 1

Myriothela (?) HICKSON & GRAVELY, 1907: 18-19, PI. 3 Fig. 18.

Locality. — Based on single specimen from Winter Quarters of Discovery expedition. Hut Point, McMurdo

Sound, Ross Sea, 13 Octoberl902 (Hickson & Gravely, 1907).

Remarks

Composed of c. 8 mm long hydrocaulus, 2 mm in diameter, basally with numerous filaments

attaching specimen to debris of sponge spicules. Body of hydranth c. 6 mm long, spindle shaped,

thickening from hydrocaulus and tapering apically to form conical hypostome. Distal half of

hydranth covered with short, thick capitate tentacles. No blastostyles or gonophores developed

on proximal part of body. Might turn out to be a juvenile specimen of one of the antarctic

species.

— 54 —

Candelabrum sp. 2

Candelabrum spec. HAND & GWILLIAM, 1951: 207, 208.

Candelabra sp. AUSTIN, 1985: 46 (sic).

Locality. — Three specimens were found in a pholad hole on the undersurface of a rock at mean lower

water at Pigeon Point, San Mateo County, California, USA, 6 May 1950.

Remarks

Based on three not fully mature specimens. The description by Hand & Gwilliam (1951)

is repeated here verbatim: “Hydranth: Not branched, solitary, naked, and arising from a creeping

hydrorhiza or possibly a disc. Hydrorhizae invested with perisarc. Exclusive of hydrorhizae, polyp

divisible into two zones; a distal tentacle-bearing zone and a proximal tentacle-free blastostylar

zone. The tentacle-bearing zone composes five-sixths or more of the polyp length and bears

approximately 500 tentacles in the adult; cylindrical, approximately the same diameter throughout.

Tentacles densely packed, short, capitate and not arranged in any discernible pattern. Mouth

terminal. Blastostylar zone swollen, of a slightly greater diameter than the tentacle bearing zone.

Sometimes separated from tentacle-bearing zone by a constriction; at the proximal end tapering

sharply to hydrorhiza. Structures referred to by Allman (1875) as claspers not present on speci¬

mens examined. Blastostyles giving rise to more than one gonophore. Largest specimen (pre¬

served) 2.5 cm long by 0.15 cm in diameter (including the tentacle)”.

Compared by Hand & Gwilliam with Candelabrum harrisoni Briggs, which it resembles

closely.

Candelabrum sp. 3

Dr Chad Hewitt and Dr Goddard (University of Tennessee, USA) recently informed us

that they will describe a species of Candelabrum from intertidal waters of the Pacific coast of

Oregon.

ECOLOGICAL REMARKS

1 — Trophic behaviour.

As stated above, a fairly large (c. 10 mm TL), partly decomposed shrimp ( Chorocaris for-

tunata Martin & Christiansen, 1995) was found in the enteron of a specimen of Candelabrum

phrygium, demonstrating their ability to capture large preys, as also indicated by their consider¬

able armament. This, and the presence of other small Crustacea (Amphipoda and Copepoda Si-

phonostomatoida) in their immediate vicinity, makes it likely that they can be considered to

prey upon the Lucky Strike faunal community.

Candelabrum serpentarii , on the contrary, does not actually live in a hydrothermal habitat.

It was found outside but close to the active sites, in an area deprived of visible fauna and

— 55 —

consequently less rich in prey. The absence (noticed during the submersible explorations) of

C. serpentarii outside an area of 20 or 30 m radius of the active sites leads to the consideration

that the species, directly or indirectly, benefits from the active sites communities. It has been

observed that an important bacterial production (both free and associated with invertebrates)

gives rise to a community largely dominated by the shrimp Rimicaris exoculata (SEGONZAC et

al, 1993). Though the type of food consumed by C. serpentarii so far has not been observed

directly, it seems reasonable to suggest that, as in C. phrygium, it consists mainly of shrimps

and other small Crustacea. The metabolism, in this animal, might be adapted to the occasional

capture of (large) preys, as is the case in many abyssal predators.

It should be remembered that in Candelabrum the number of tentacles is considerable, the

nematocysts being concentrated in the tentacular capitulum. As indicated above the considerable

amount of nematocysts allows the capture of large preys, for which action the haplonemes, ste-

nosteles and heteronemes, if present, are responsible. The numerous desmonemes may serve for

the attachment of preys, that can either be transported to the mouth or swallowed after curvature

of the trunk towards the place of attachment. Many athecate hydroids have an extensible mouth

capable of devouring a prey of considerable size. The Candelabrum species probably are no

exception in this respect.

2 — Biogeography.

Most Candelabrum species live in intertidal waters. Geographical distribution in the mem¬

bers of this genus is usually restricted by the absence of a planktonic stage and by the necessity

to be attached to hard substrates. The circumarctic distribution of C. phrygium, which has a

non-planktonic actinula larva, is not in agreement with the ecological status observed on the

Lucky Strike site. Indeed, its distribution there is restricted to areas of active venting (living in

a mixture of sea water and hydrothermal fluid loaded with sulfurous compounds, mean tempera¬

ture: 11° C). Moreover, direct observations and video analysis indicate the absence of individuals

outside the sites. It is thus difficult to understand why this species, is strongly linked to such

peculiar and very confined physical and chemical conditions.

The status of C. serpentarii is likewise difficult to explain, but it is different. As stated

above, this species has affinities with C. cocksii, a species only known from the NE Atlantic

intertidal zone. Both species have low dispersal abilities (the fertilized eggs develop into the

actinulae before they separate from blastostyles); they were never observed or collected on the

abyssal plains.

Considering those facts, and bearing in mind the difficulties in understanding the coloni¬

zation processes without fossil records, the isolation of the various Candelabrum populations

could be explained in terms of plate tectonics viz. the movement of oceanic plates. For example,

it could be hypothesized that Candelabrum serpentarii results from allopatric spéciation (with

the preservation of several morphologic features and the mode of reproduction) from an ancient

intertidal stock of C. cocksii (or, more likely, a common ancestral form) that became isolated

as a result of the successive events that led to the rifting and further spreading of the Atlantic

Ocean floor. One of the populations may progressively have adapted to the deep hydrothermal

environment where the hard substrate and the trophic conditions represented favourable living

conditions.

— 56 —

In order to answer questions concerning the present distribution of these species of Can¬

delabrum , several kinds of studies should be carried out:

1) a profound study leading towards a better understanding about their life-cycle;

2) a study of the deep water circulation pattern in this part of Atlantic Ocean;

3) the exploration of other ridge segments to gain additional records of species of Can¬

delabrum.

Technical aid

All C. cocksii, C. phrygium and C. serpentarii nematocysts (Fig. 2) have been photographed by

Dr J. C. den Hartog (Nationaal Natuurhistorisch Museum, Leiden). Fig. 3D-F and Fig. 4 were made from

photographs taken by Patrick Briand (IFREMER). The map shown in Fig. 1 was realized by Violaine

Martin (IFREMER). The pictures Fig. 3A, B, C and E, partly resulting from videotape enlargements, were

placed at our disposal by Gérard Vincent and Valérie Baty (Picture library, IFREMER).

Acknowledgements

The authors wish to express their gratitude to the chief scientists: A.-M. Alayse, D. Desbruyères,

J. Dubois, A. Fiala, C. Langmuir, C. Mével and C. Van Dover, as well as the crews of both the

research vessels and the submersibles, that participated in the cruises, Dr A. Castric-Fey (Concarneau)

who kindly placed specimens of C. cocksii at our disposal, Dr J. C. den Hartog (Leiden), Patrick Briand,

Violaine Martin, Gérard Vincent and Valérie Baty (IFREMER) for their technical aid, and the authorities

of the National Museum of Natural History where research on the specimens was partly carried out.

The specimens of C. serpentarii and C. phrygium originate from cruises Hydrosnake, Lucky Strike,

MAR 93, Gravinaut and Diva 2 organized by INSU, IFREMER and WHOI institutions within the framework

of the FARA program (French American Ridge Atlantic).

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(Essays on Hydra’s. I. Hydra baicalensis n. sp. II. On the question of Hydra’s phytogeny). Sb. Trud. Prof.

Prepod. gos. Irkutsk. Univ., 4: 90-102 [Russian],

Swenander, G., 1904. — Über die Athecaten Hydroiden des Drontheimsfjordes. K. norske Vidensk. Selsk. Skr.,

1903 (6): 1-18. PI. 1.

TEISSIER, G., 1950b. — Inventaire de la faune marine de Roscoff. Cnidaires et Cténaires. Trav. biol. Roscoff,

n. sen, suppl. 1: 1-43.

—1965. — Inventaire de la faune marine de Roscoff. Cnidaires-Cténaires. Trav. Stn biol. Roscoff, 16: 1-53

[64],

Thomson, J. A., 1904. — Scotia collections. Note on the gonostyle of two antarctic Siphonophora. Proc. R.

phys. Soc. Edinb., 16 (1): 19-22, PL 1.

Van DE Vyver, G., 1968a. — Étude du développement embryonnaire des hydraires athécates (Gymnoblastiques)

à gonophores. 2. Formes à actinulas. Archs Biol., Paris, 79: 327-363, Figs 1-7, i-v.

Van DE Vyver, G., 1968b. — Étude du développement embryonnaire des hydraires athécates (Gymnoblastiques)

à gonophores. 3. Discussion et conclusions générales. Archs Biol., Paris, 79: 365-379.

VAN DE VYVER, G., 1980. — A comparative study of the embryonic development of Hydrozoa Athecata. In: P.

TARDENT & R. Tardent, eds.. Developmental and cellular biology of coelenterates. Proceedings IV. In¬

ternational Coelenterate Conference, Interlaken, 4-8 Sept., 1979: 109-120, Figs 1-6, Tab. Elsevier/N. Holland

Biomed. Press, Amsterdam, etc.: i-xxvi, 1-499, Figs, Tabs.

Van Dover, C. L., D. Desbruyères, M. Segonzac, T. Comtet, L. Saldanha, A. Fiala-Médioni &

C. LANGMUIR, submitted. — Biology of the Lucky Strike Hydrothermal Field. Deep-sea Research.

VANHÔFFEN, E., 1897. — Die Fauna und Flora Gronlands. Gronland Expedition der Gesellschaft fiir Erdkunde

zu Berlin, 1891-1893, unter Leitung von E. VON DRYGALSKI, 2 (1): 1-383, Pis 1-8.

— 1910. — Die Hydroiden der Deutschen Siidpolar-Expedition 1901-1903. Deutsche Siidpolar-Expedition, 11

(= Zool. 3): 269-340, Figs 1-49.

VERRILL, A. E., 1879. — Preliminary check-list of the marine Invertebrata of the Atlantic coast, from Cape Cod

to the Gulf of St. Lawrence: 1-32. New Haven, Connecticut.

VlGURS, 1849. — In: Cocks W. P. 1849: 90.

WEILL, R., 1926. — Une catégorie spéciale de nématocystes commune aux seuls Hydrides, Gymnoblastides et

Siphonophores. C. r. hebd. Séanc. Acad. Sci. Paris, 182 (20): 1244-1247, Figs 1-4.

— 1934a.— Contribution à l’étude des Cnidaires et de leurs nématocystes. I. Recherches sur les nématocystes.

Morphologie - Physiologie - Développement. Trav. Stn zool. Wimereux, 10: 1-347, Figs 1-208.

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Sars]. Proc. R. phys. Soc. Edinb., 1: 432-433.

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Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 65-76.

Sur les Oswaldocruzia (Nematoda, Trichostrongylina,

Molineoidea), parasites d’amphibiens et de lézards

de Cuba et de Porto Rico

par Badreddine Ben Slimane et Marie-Claude DURETTE-DESSET

Résumé. — L’examen d'Oswaldocruzia de Cuba et de Porto Rico montre l’existence de quatre espèces

différentes. O. lenteixeirai Vigueras, 1938, dépourvue d’ailes cervicales, possède des côtes bursales 8 chevauchées

par les côtes 6 dans leurs deux tiers proximaux et des côtes bursales 5-6 jointives ; mâle avec 32-38 crêtes

cuticulaires au milieu du corps ; vestibule pourvu de formations distales. O. moraveci n. sp. (= O. lenteixeirai

sensu Barus et Moravec, 1967, nec Vigueras, 1938) se différencie de la précédente par des côtes bursales 8

chevauchées par les côtes 6 dans leur partie médiane et des côtes bursales 5-6 divergentes à leur extrémité ;

vestibule dépourvu de formations distales. O. anolisi Barus et Coy Otero, 1968, pourvue de grandes ailes cervi¬

cales, possède des côtes bursales 8 chevauchées par les côtes 6 dans leur partie médiane et des côtes bursales

5-6 jointives ; mâle avec 50 crêtes cuticulaires au milieu du corps ; vestibule dépourvu de formations distales.

O. barusi n. sp., pourvue d’ailes cervicales peu développées, possède des côtes bursales 8 chevauchées par les

côtes 6 dans leur partie médiane et des côtes bursales 5-6 jointives ; mâle avec 38 crêtes cuticulaires au milieu

du corps; vestibule dépourvu de formations distales.

Mots-clés. — Oswaldocruzia , Nématodes, Trichostrongylina, Amphibiens, Lézards, Antilles.

On Oswaldocruzia spp (Nematoda: Trichostrongylina-Molineoidea), parasites of amphibians

and lizards from Cuba and Puerto Rico

Abstract. — Four different species of Oswaldocruzia have been identified from Cuba and Puerto Rico.

O. lenteixeirai Vigueras, 1938, without cervical alae, with rays 8 joined to rays 6 along the proximal two thirds

and with rays 5 and 6 close together; male with 32-38 cuticular crests in the mid body ; vestibule with distal

formations. O. moraveci n. sp. (= O. lenteixeirai sensu Barus et Moravec, 1967, nec Vigueras, 1938) differen¬

tiated from the former by rays 8 overlapped by rays 6 along their median part and by the divergency of the

extremities of rays 5 and 6; vestibule without distal formations. O. anolisi Barus et Coy Otero, 1968, with

developed cervical alae; with rays 8 overlapped by rays 6 along their median part and with rays 5 and 6 close

together; male with 50 cuticular crests in the mid body; vestibule without distal formations. O. barusi n. sp.,

with poorly developed cervical alae; with rays 8 overlapped by rays 6 along their median part and with 5 and

6 rays close together; male with 38 cuticular crests in the mid body; vestibule without distal formations.

Keywords. — Oswaldocruzia, Nematoda, Trichostrongylina, Amphibians, Lizards, West Indies.

B. Ben Slimane et M.-Cl. Durette-Desset, Laboratoire de Biologie parasitaire, Protistologie, Helminthologie, Muséum national

d’Histoire naturelle, 61, rue Buffon, F-75231 Paris Cedex 05.

INTRODUCTION

Le genre Oswaldocruzia (44 espèces connues) est un parasite cosmopolite d’Amphibiens

et de Reptiles. Il a été créé par TRAVASSOS (1917) avec pour espèce type O. subauricularis.

Deux espèces ont été décrites et identifiées de diverses îles des Antilles : O. lenteixeirai Vigueras,

1938, à Cuba et Porto Rico et O. anolisi Barus et Coy Otero, 1968, à Cuba. Cependant, en

— 66 —

1968, Barus & Coy Otero ont mis O. anolisi en synonymie d’ O. lenteixeirai en se basant

sur le travail de Moravec & Vojtkova (1975) car ces auteurs estimaient que la présence ou

l’absence d’ailes cervicales n’était pas un bon caractère spécifique. Depuis cette époque, plusieurs

publications (Durette-Desset, Nasher & Ben Slimane, 1992, Ben Slimane, Durette-Desset

& Chabaud, 1993) se fondant sur des coupes transversales du corps montrent que, en plus de

leurs dimensions, les ailes cervicales peuvent être constituées par un nombre spécifique de crêtes

cuticulaires et avoir une forme spécifique.

Il nous a donc paru utile, en utilisant ce caractère, de reprendre les différents travaux concer¬

nant ces deux espèces.

Grâce à l’obligeance du Pr Moravec et du Dr Lichtenfels, que nous remercions très

vivement, nous avons obtenu les différents spécimens qui sont étudiés ci-dessous.

MATÉRIEL ET MÉTHODES

Les spécimens, conservés dans l'alcool à 70°, sont étudiés dans de l’eau ou du lacto-phénol

plus ou moins dilué. Lorsque le matériel est suffisamment abondant, les coupes transversales du

corps et la dissection des spicules sont réalisées selon les techniques de DURETTE-DESSET (1985).

La nomenclature utilisée pour la bourse caudale est celle de Durette-Desset & Chabaud

(1981), pour l’étude du trajet des côtes 8 par rapport aux côtes 6, celle de Durette-Desset et

al. (1992). La nomenclature du synlophe dans la région œsophagienne et des spiculés sont celles

de Ben Slimane et al. (1993).

REDESCRIPTION DES ESPÈCES

Oswaldocruzia lenteixeirai Vigueras, 1938

Matériel de redescription. — 1 mâle en deux morceaux et 1 femelle United States National Museum,

Helminthological Collections, n° 73235.

Hôte : Eleutherodactylus portoricensis Thomas, 1966 (Leptodactylidae).

Localisation : intestin grêle.

Origine géographique : Cupey, Porto Rico.

Collection : Julio Garcia.

Identification : G. D. Schmidt.

Fig. 1. — Oswaldocruzia lenteixeirai Vigueras, 1938 chez Eleutherodactylus portoricensis. (Matériel de Schmidt et Wittaker,

1975.) A-E, femelle : A, extrémité antérieure, vue latérale droite; B, C, vulve, vues latérale droite et ventrale; D, détail des

deirides et du pore excréteur, vue ventrale ; E, ovéjecteur, vue latérale droite ; F, mâle, synlophe au milieu du corps ; G, id.,

bourse caudale, vue ventrale avec disparition des crêtes cuticulaires ; H, femelle, queue, vue latérale droite ; I, femelle, extrémité

caudale, vue ventrale, disparition des crêtes cuticulaires. Toutes les coupes sont orientées comme la figure 1 F. Échelle : A,

E : 70 pm ; B-D, G, H : 50 pm ; F, I : 30 pm.

Oswaldocruzia lenteixeirai Vigueras, 1938 front Eleutherodactylus portoricensis. (Original material from Schmidt & Wiitaker,

1975.) A-E, female: A, anterior extremity, right lateral view; B. Vulva, right lateral view : C, id., ventral view; D, deirids

and excretory pore, ventral view ; E, ovejector, right lateral view ; F, male, synlophe at mid-body ; G, id. caudal bursa,

ventral view showing disappearance of cuticular ridges ; H, female, tail, right lateral view ; 1, female, caudal extremity,

ventral view, disapperance of cuticular ridges. All the body sections are orientated as fig. 1 F. Abbr.: d. = dorsal side ; dr.

= right side ; v = ventral side. Scale: A, E: 70 pm ; B-D, G, H: 50 pm ; F, I: 30 pm.

— 68 —

Description

Nématodes de grande taille, déroulés; pore excréteur situé postérieurement à la jonction

œsophago-intestinale ; deirides triangulaires, postérieures au pore excréteur (fig. 1 D); glandes

excrétrices bien développées. Ailes cervicales absentes.

Tête : présence d'une vésicule céphalique et d’une grosse dent œsophagienne dorsale

(fig. I A).

Synlophe (étudié chez le mâle en coupe transversale) : dans les deux sexes, corps parcouru

longitudinalement par des crêtes cuticulaires continues pour la grande majorité d’entre elles.

Certaines crêtes sont interrompues et remplacées par la naissance d’une autre crête. Chez le

mâle, les crêtes cuticulaires disparaissent au niveau de la bourse caudale (fig. 1 G) et au niveau

de la pointe caudale chez la femelle (fig. 1 H. I).

Chez le mâle, le nombre de crêtes est de 33 (16 crêtes dorsales, 17 crêtes ventrales) à

300 pm en avant de la bourse caudale. Toutes les crêtes sont orientées perpendiculairement à

la paroi du corps, espacées régulièrement et de taille équivalente (fig. 1 F).

Mâle : morceau antérieur long de 2 000 pm, postérieur de 3 400 pm sur 160pm de large

dans sa partie moyenne. Vésicule céphalique haute de 92 pm sur 40 pm de large. Anneau nerveux,

pore excréteur et deirides situés respectivement à 155 pm, 360 pm et 375 pm de l’apex. Œsophage

long de 340 pm.

Bourse caudale de type 2-3 à tendance 2-1-2 : l’extrémité des côtes 4 étant coudée vers

l'avant est plus proche de celle des côtes 3 que de celle des côtes 5. Côtes 2-3 d’une part et

côtes 5-6 d’autre part jointives. Côtes 8 naissant à la racine de la côte dorsale et chevauchées

par les côtes 6, sauf dans leur extrémité postérieure (type III). Côte dorsale divisée dans son

quart postérieur en trois rameaux, les rameaux externes (côte 9) se détachant avant la division

de la côte dorsale. Gubernaculum absent, cône génital de forme triangulaire haut de 30 pm sur

30 pm de large dans sa partie proximale, portant sur sa lèvre antérieure une large papille 0 et

deux minuscules papilles 7 sur sa lèvre postérieure (fig. 1 G).

Spicules non disséqués, mais conformes à la descrition de SCHMIDT & WlTTAKER (1975) :

ailés, longs de 180pm, à pointes complexes se divisant au tiers proximal de leur hauteur en

trois branches principales; chaque branche est en forme de lame, chaque lame se termine en

plusieurs pointes distales en forme de peigne.

Femelle : longue de 6900 pm sur 190 pm dans sa partie moyenne. Vésicule céphalique haute

de 85 pm sur 45 pm de large. Anneau nerveux, pore excréteur et deirides situés respectivement

à 155 pm, 320 pm et 335 pm de l'apex. Œsophage long de 300 pm (fig. 1 A).

Didelphie, la vulve s’ouvre à 2 800 pm de la pointe caudale. Vagina vera long de 48 pm

divisant le vestibule long de 270 pm en deux parties éqivalentes (fig. 1 E). Sphincter et trompe

de chaque branche longs respectivement de 40 pm et 20 pm, branche utérine antérieure longue

de 1 050 pm contenant 16 œufs au stade morula, branche utérine postérieure longue de 1 300 pm

contenant 12 œufs au stade morula. Œufs hauts de 88 pm sur 45 pm de large. Queue longue de

240 pm sur 70 pm de large au niveau de l’anus, la longueur de l’épine caudale ne peut être

donnée, cette dernière étant cassée (fig. 1 H, I).

— 69 —

Oswaldocruzia barusi n. sp.

Matériel type. — Mâle holotype, femelle allotype. Institute of Parasitology. Academy of Sciences of the

Czech Republic, Helminthological Collections n° 646, a ; 2 mâles, 2 femelles paratypes, Institute of Parasitology,

Academy of Sciences of the Czech Republic, Helminthological Collections n° 646, b.

Hôte : Bufo empusus Cope, 1862 (Bufonidae).

Localisation : intestin grêle.

Origine géographique : jardin botanique La Habana, Cuba.

Collection : Dr. V. Barus.

Identification : « O. lenteixeirai » V. Barus.

Description

Nématodes de grande taille présentant un tour de spire dans leur partie antérieure, le reste

du corps étant déroulé; pore excréteur de position variable, mais toujours situé dans le tiers

postérieur de l’oesophage ; deirides postérieures au pore excréteur et de forme triangulaire, glandes

excrétrices bien développées. Ailes cervicales présentes.

Tête : présence d’une vésicule céphalique et d’une petite dent œsophagienne dorsale. En

vue apicale, on observe 6 papilles labiales externes, 2 amphides, 4 papilles céphaliques. Bouche

triangulaire arrondie aux angles (fig. 2 B).

Synlophe (étudié chez un mâle et une femelle paratypes) : dans les deux sexes, corps par¬

couru longitudinalement par des crêtes cuticulaires continues à l’exception de quelques crêtes

interrompues de façon irrégulière. La quasi-totalité des crêtes chez le mâle, les quatre cinquièmes

chez la femelle naissent dans la région œsophagienne ; les crêtes disparaissent au niveau de la

bourse caudale chez le mâle et au niveau des phasmides chez la femelle.

Au niveau de la jonction œsophago-intestinale, le nombre de crêtes est de 37 (20 dorsales,

17 ventrales) chez le mâle (fig. 2 C), de 39 (20 dorsales, 19 ventrales) chez la femelle (fig. 2

E). Au milieu du corps, le nombre de crêtes est de 38 (20 dorsales, 18 ventrales) chez le mâle

(fig. 2 D) et de 48 (25 dorsales, 23 ventrales) chez la femelle (fig. 2 F).

Les crêtes sont espacées de façon régulière et sont de taille équivalente sauf dans la région

œsophagienne où les deux crêtes latéro-ventrales proches des cordons latéraux sont légèrement

plus grandes que les autres crêtes et forment des ailes cervicales visibles seulement en coupe

transversale; celles-ci naissent en arrière de la vésicule céphalique et disparaissent au niveau de

la jonction œsophago-intestinale.

Toutes les crêtes sont orientées perpendiculairement à la paroi du corps, sauf les ailes cer¬

vicales dont la pointe est orientée ventralement.

Mâle holotype : long de 4 800 pm sur 125 jim de large dans sa partie moyenne. Vésicule

céphalique haute de 75 pm sur 35 pm de large. Anneau nerveux, pore excréteur et deirides situés

respectivement à 150 p, 200 pm et 210 pm de l'apex. Œsophage long de 380 pm (fig. 2 A).

Bourse caudale de type 2-3 à tendance 2-1-2 : l’extrémité des côtes 4 étant coudée vers

l’avant est plus proche de celle des côtes 3 que de celle des côtes 5. Côtes 8 naissant à la racine

de la côte dorsale et chevauchées par les côtes 6 dans leur partie médiane (type II). Côte dorsale

divisée dans son quart postérieur en trois rameaux, le rameau externe (côte 9) se détachant avant

la division de la côte dorsale. Gubernaculum absent, cône génital de forme triangulaire haut de

— 70 —

30 pm sur 30 pm de large dans sa partie proximale, portant sur sa lèvre antérieure une large

papille 0 et deux minuscules papilles 7 sur sa lèvre postérieure (fig. 2 M).

Spicules ailés, longs de 120 pm, à pointes complexes se divisant au tiers proximal de leur

hauteur en trois branches principales chacune en forme de lame; chaque lame se termine en

plusieurs pointes distales en forme de peigne (fig. 2 J-L).

Femelle allotype : longue de 7 700 pm sur 120 pm dans sa partie moyenne. Vésicule cé¬

phalique haute de 70 pm sur 35 pm de large. Anneau nerveux, pore excréteur et deirides situés

respectivement à 175 pm, 230 pm et 250 pm de l’apex. Œsophage long de 420 pm.

Didelphie, la vulve s’ouvre à 2 850 pm de la pointe caudale. Vagina vera long de 45 pm

divisant le vestibule long de 540 pm en deux parties équivalentes. Sphincter et trompe de chaque

branche longs respectivement de 30 pm et de 20 pm (fig. 2 H). Branche utérine antérieure longue

de 1 750 pm contenant 19 œufs au stade morula, branche utérine postérieure longue de 1 750 pm

contenant 25 œufs au stade morula. Œufs hauts de 60 pm sur 40 pm de large. Queue longue de

230 pm sur 50 pm de large au niveau de l’anus, pointe caudale longue de 19 pm (fig. 2 I).

O. anolisi Barus & Coy Otero, 1968

Matériel. — 2 mâles, 1 femelle, Institute of Parasitology, Academy of Sciences of the Czech Republic,

Helminthological Collections, n° N-376.

Hôte : Anolis equestris Merrem, 1820 (Iguanidae).

Localisation : intestin grêle.

Origine géographique : Cuba.

Collection : Dr. V. Barus.

Identification : «O. anolis » V. Barus.

Description

Nématodes de grande taille et déroulés; pore excréteur situé très antérieurement (fig. 3 A);

deirides au même niveau, filiformes et bien développées. Grandes ailes cervicales.

Tête : présence d’une vésicule céphalique et d’une petite dent œsophagienne dorsale.

Synlophe (étudié en coupe transversale chez un mâle) : dans les deux sexes, corps parcouru

longitudinalement par des crêtes cuticulaires continues à l’exception de quelques crêtes inter¬

rompues de façon irrégulière. Les crêtes dorsales naissent en arrière de la vésicule céphalique

Fig. 2. — Oswaldocruzia barusi n. sp. chez Bufo empusus. A, mâle, partie antérieure, vue latérale gauche ; B, femelle, tête, vue

apicale ; C, D, mâle, synlophe : C, au niveau de la jonction œsophago-intestinale ; D, au milieu du corps ; E-G, femelle,

synlophe : E, au niveau de la jonction œsophago-intestinale ; F, au milieu du corps. G, au niveau du vestibule ; H, femelle,

ovéjecteur, vue latérale gauche ; I, femelle, queue, vue latérale gauche ; J-L, mâle, spicule gauche disséqué : J, vue dorsale ;

K, vue externe; L, vue ventrale; M, mâle, bourse caudale, vue ventrale. Toutes les coupes sont orientées comme la figure

2 C. Échelle : A, I, M : 80 pm ; B-G, J-L : 30 pm ; H : 100 pm.

Oswaldocruzia barusi n. sp. from Bufo empusus. A, male, anterior extremity, left lateral view; B, female, head, en face view;

C, D, male, synlophe: C, at the œsophago-intestinal junction level; D, at mid-body; E-G, female, synlophe: E, at the œso-

phago-intestinal junction level; F, at mid-body; G, at vestibule level; H, female, ovejector, left lateral view; I, female, tail,

left lateral view; J-L, male, dissected left spicule: J, dorsal view; K, externo-lateral view; L, ventral view; M, male, caudal

bursa, ventral view. All the body sections are oriented as fig. 2 C. Abbr.: d. = dorsal side, dr. = right side, g = left side,

v. = ventral side. Scale: A, I, M: 80 pm; B-G, J-L: 30 pm; H: 100 pm.

— 72 —

mais, sauf sur coupe transversale, elles ne deviennent visibles qu’au niveau de la jonction

œsophago-intestinale. Les crêtes ventrales apparaissent à environ 180pm en arrière de la fin de

l’œsophage.

Le nombre de crêtes est de 13 (11 dorsales et 2 ailes latérales) dans la partie antérieure

(fig. 3 B) et de 50 (25 dorsales, 25 ventrales) au milieu du corps (fig. 3 C). Les ailes cervicales

naissent à la base de la vésicule céphalique et sont longues de 850 pm chez le mâle et de 950 pm

chez la femelle. Chaque aile est composée d’une seule crête en position latéro-ventrale, haute

de 28 pm dans sa partie la plus large. Les crêtes dorsales et ventrales sont orientées perpendi¬

culairement à la paroi du corps, espacées régulièrement et de taille équivalente. Les ailes cer¬

vicales sont légèrement orientées dorso-ventralement.

Mâle : long de 8 350-9 700 pm sur 185-200 pm de large dans sa partie moyenne ; vésicule

céphalique haute de 85-85 pm sur 50-60 pm de large. Anneau nerveux, pore excréteur et deirides

situés respectivement à 140-150 pm, 190-200 pm et 210-220 pm de l’apex. Œsophage long de

440-480 pm (fig. 3 A).

Bourse caudale de type 2-3 à tendance 2-1-2 : l’extrémité des côtes 4 étant coudée vers

l’avant est plus proche de celle des côtes 3 que de celle des côtes 5. Côtes 8 naissant à la racine

de la côte dorsale, chevauchées par les côtes 6 dans leur partie médiane (type II). Côte dorsale

divisée dans son quart postérieur en trois rameaux, le rameau externe (côte 9) se détachant avant

la division de la côte dorsale. Gubernaculum absent, cône génital de forme triangulaire haut de

20 pm sur 20 pm de large dans sa partie proximale, portant sur sa lèvre antérieure une large

papille 0 et deux minuscules papilles 7 sur sa lèvre postérieure (fig. 3 E).

Spicules ailés, longs de 195-200 pm, à pointes complexes se divisant au tiers proximal de

leur hauteur en trois branches principales, chacune en forme de lame ; chaque lame se termine

en plusieurs pointes distales en forme de peigne.

Femelle: Longue de 13 100 pm sur 230 pm dans sa partie moyenne. Vésicule céphalique

haute de 85 pm sur 55 pm de large. Anneau nerveux, pore excréteur et deirides situés respec¬

tivement à 170 pm, 260 pm et 280 pm de l’apex. Œsophage long de 430 pm.

Didelphie, la vulve s’ouvre à 4 400 pm de la pointe caudale. Vagina vera long de 50 pm

divisant le vestibule, long de 600 pm, en deux parties équivalentes. Sphincter et trompe de chaque

branche longs respectivement de 40 et 30 pm (fig. 3 D). Chaque branche utérine longue de

2 200 pm contient 30 œufs au stade morula hauts de 75 pm sur 40 pm de large. Queue longue

de 280 pm sur 70 pm de large au niveau de l’anus, pointe caudale longue de 18 pm (fig. 3 F).

Fig. 3. — Oswaldocruzia anolisi Barus & Coy Otero, 1968 chez Anolis equestris. A, mâle, partie antérieure, vue ventrale; B-C,

mâle, synlophe : B, au niveau le plus large des ailes cervicales ; C, au milieu du corps ; D, femelle, ovéjecteur, vue latérale

droite ; E, mâle, bourse caudale, vue ventrale ; F, femelle, queue, vue latérale gauche. Toutes les coupes sont orientées comme

la figure 3 B. Échelle : A 100 pm; B, C : 50 pm; D : 90 pm E : 80 pm; F : 70 pm.

Oswaldocruzia anolisi Barus & Coy Otero, 1968, from Anolis equestris. A, male, anterior extremity, ventral view; B-C, male,

synlophe: B, at the highest wide level of cervical alae; C, at mid-body; D, female, ovejector, right lateral view; F, female,

tail, left lateral view. All the body sections are orientated as fig. 3 B. Abbr.: d. = dorsal side, dr. = right side, g. = left

side, v. = ventral side. Scale: A: 100 p m; B, C: 50 p m; D: 90 pm; E: 80 pm; F: 70 pm.

— 74 —

DISCUSSION

Les Oswaldocruzia parasites d’amphibiens et de reptiles des Antilles n’ont été décrits ac¬

tuellement que de Cuba et de Porto Rico. Us sont caractérisés par des spicules à trois branches

principales, chacune en forme de lame, chaque lame se terminant en plusieurs pointes distales

en forme de peigne. Us constituent donc un groupe très homogène si bien que, depuis la mise

en synonymie d ’O. anolisi, une seule espèce, O. lenteixeirai est reconnue par tous les auteurs.

Cependant, l'étude morphologique effectuée ci-dessus indique des différences importantes carac¬

téristiques de chacun des lots examinés. U est donc nécessaire de passer en revue les publications

faites à ce sujet pour tenter d’éclaircir le problème.

— En 1938, la description princeps de VlGUERAS porte sur des parasites de Hyla insula

(= Hyla septentrionalis) à Cuba. L’espèce est caractérisée par:

(1) l’absence d’ailes cervicales, tout au moins sur l’animal examiné sans coupe transversale

du corps;

(2) des côtes 6 et 8 dont la disposition est de type III ; des côtes 8 complètement chevauchées

par les côtes 6 en vue ventrale, sauf à leur extrémité distale ;

(3) un vestibule très long (850-870 pm) avec une formation (?) glandulaire aux deux ex¬

trémités.

— En 1967, Barus & Moravec pensent retrouver l’espèce chez le même hôte et dans la

même région. Cependant, plusieurs caractères diffèrent de ceux de l’espèce précédente :

(1) les côtes 6 et 8 ont une disposition de type II, les côtes 6 chevauchant ventralement

les côtes 8 uniquement dans leur partie médiane ;

(2) les côtes 5 et 6 ne sont pas jointives, mais leurs extrémités divergent. En outre, les

côtes 5, 6 et 8 sont de longueur équivalente;

(3) le vestibule est dépourvu de formations particulières à ses extrémités;

(4) la vulve est entourée de deux becs cuticulaires, l’un antérieur, l’autre postérieur.

Ces Nématodes nous paraissent donc différents de ceux de VlGUERAS et nous proposons

de les nommer Oswaldocruzia moraveci n. sp. (= O. lenteixeirai sensu Barus & MORAVEC,

1967, nec VlGUERAS, 1938).

— En 1968, Barus & Coy Otero décrivent O. anolisi chez Anolis equestris de Cuba. Le

matériel décrit ci-dessus correspond au matériel original, et les caractéristiques morphologiques

correspondent à celles indiquées dans la description originale; plusieurs éléments permettent de

différencier O. anolisi des deux autres espèces :

(1) présence d’ailes cervicales simples et bien développées;

(2) présence de becs cuticulaires périvulvaires, l’antérieur important, le postérieur faible;

(3) deirides allongées et bien développées chez O. anolisi, arrondies et petites chez O. len¬

teixeirai ;

(4) crêtes cuticulaires ventrales absentes dans la partie antérieure du corps chez O. anolisi,

présentes chez O. lenteixeirai ;

(5) côtes bursales 8 chevauchées par les côtes 6 dans leur portion médiane (type II) chez

O. anolisi et non dans leur deux tiers proximaux (type III) chez O. lenteixeirai.

La validité de l’espèce O. anolisi semble donc ne faire aucun doute.

— 75 —

— En 1969, Barus & Coy Otero et, en 1970, Coy Otero signalent la présence d'O. ano¬

lisi chez de nombreux lézards Iguanidae et Teiidae de Cuba.

— En 1972, BARUS identifie à O. lenteixeirai les parasites de différentes espèces d 'Eleu-

therodactylus de Cuba, mais signale que la longueur des spicules (87-133 pm) est inférieure à

celle des parasites de Hyla ou de Bufo (124-182 pm).

— En 1973, Barus identifie à O. lenteixeirai les parasites de différentes espèces de Bufo

de Cuba, et précise quelques éléments morphologiques (ailes cervicales très petites, synlophe

avec 34 à 36 crêtes cuticulaires).

— En 1975, SCHMIDT & WHITTAKER identifient O. lenteixeirai chez un Eleutherodactylus

de Porto Rico et fournissent quelques éléments morphologiques supplémentaires. C’est leur ma¬

tériel que nous avons utilisé ci-dessus pour redécrire l’espèce. Il eut été plus satisfaisant de

s’appuyer sur le matériel de Barus et Moravec de 1967 puisque l’hôte et la géographie coïn¬

cident avec ceux de la description originale, mais nous savons qu’un même hôte peut héberger

plusieurs espèces différentes d’ Oswaldocruzia. En outre et surtout, les différences morphologiques

entre la description de Vigueras et celle de Barus & Moravec sont considérables alors qu’au¬

cun élément important ne sépare le matériel de Schmidt & Whittaker de la description de

Vigueras.

En 1978, Barus & Coy Otero rapportent la présence d'Oswaldocruzia chez plusieurs ser¬

pents de Cuba, mais bien que ces parasites aient tous de grandes ailes cervicales, les auteurs

proposent la synonymie de leur espèce O. anolisi avec O. lenteixeirai ce qui, comme il est indiqué

plus haut, nous semble inexact.

Enfin, dans le présent article, des spécimens parasites de Bufo empusus de Cuba, qui avaient

été rattachés primitivement à O. lenteixeirai nous paraissent s’en différencier par les ailes cer¬

vicales, la bourse caudale, les spicules et la structure du vestibule et nous avons proposé de les

nommer O. barusi n. sp.

En conclusion, nous estimons que les Oswaldocruzia de Cuba et de Porto Rico, qui sont

les seuls étudiés actuellement aux Antilles, comportent au moins quatre espèces distinctes qui

peuvent être reconnues par les caractères suivants.

O. lenteixeirai Vigueras, 1938 : ailes cervicales absentes; 32-38 crêtes cuticulaires au milieu

du corps chez le mâle, 52-58 chez la femelle; bourse caudale avec des côtes 8 naissant à la

racine de la côte dorsale et chevauchées par les côtes 6 dans leurs deux tiers proximaux (type III)

côtes 2-3 d’une part et côtes 5-6 d'autre part jointives ; longueur des lames des spicules subégales ;

languettes vulvaires absentes ; vestibule pourvu de formations distales.

O. moraveci n. sp. (= O. lenteixeirai sensu BARUS & MORAVEC, 1967) : ailes cervicales

absentes; nombre de crêtes cuticulaires inconnu; bourse caudale avec des côtes 8 naissant sur

la côte dorsale et chevauchées par les côtes 6 dans leur partie médiane (type II) ; côtes 2-3

jointives; côtes 5-6 divergentes dans leur tiers distal ; longueur des lames des spicules subégales;

présence de languettes vulvaires; vestibule dépourvu de formations distales.

O. anolisi Barus & Coy Otero, 1968 : ailes cervicales très larges; absence de crêtes cuti¬

culaires ventrales dans la partie antérieure du corps; 50 crêtes cuticulaires au milieu du corps

chez le mâle; côtes 2-3 séparées et côtes 5-6 jointives; côtes 8 naissant sur la côte dorsale et

chevauchées par les côtes 6 dans leur partie médiane (type II) ; longueur des lames des spicules

subégales; présence d’une languette prévulvaire; vestibule dépourvu de formations distales.

— 76 —

O. baritsi n. sp. (= O. lenteixeirai sensu Barus, 1973) : ailes cervicales peu développées,

indiquées seulement par les deux paires de crêtes latérales légèrement plus grandes que les autres;

38 crêtes cuticulaires au milieu du corps chez le mâle, 48 chez la femelle; la disposition des

côtes 6 et 8 est de type II ; lame externe des spicules plus longue que les deux lames internes ;

languettes vulvaires absentes; vestibule dépourvu de formations distales.

Remerciements

Nous remercions vivement le Dr F. Moravec de l’Institut de Parasitologie de l'Académie des Sciences

de la République Tchèque et le Dr R. Lichtenfels de l’United States Department of Agriculture, Agri¬

cultural Research Service, Biosystematic Parasitology Laboratory pour le prêt des spécimens ayant servi

à cette étude.

RÉFÉRENCES

BARUS, V., 1973. — Nematodes parasitizing hosts of the genus Bufo (Amphibia) in Cuba. Folia Parasitologica

(Praha), 20: 29-39.

Barus, V., & A. COY Otero, 1968. — Freitasia teixeirai gen. n. et sp. n. and Other Nematodes Parasitizing

Anolis equestris (Squamata: Iguanidae). Folia Parasitologica (Praha), 15: 41-45.

Barus, V., & F. MORAVEC, 1967. — Systematic studies of parasitic worms, found in the hosts Lepisosteus

tristoechus (Ginglymodi, Lepisosteidae) and Hyla insulae (Ecaudata, Hylidae) from Cuba. Vestnik Cs. spot.

Zool. (Acta Soc. Zoo/. Bohemoslov.), 31: 1-14.

BEN SLIMANE, b., M.-Cl. Durette-Desset & A. G. CHABAUD, 1993. — Oswaldocruzia (Trichostrongyloidea)

parasites d’Amphibiens des Collections du Muséum de Paris. Ann. Parasit. Hum. Comp.. 68 : 88-100.

COY OTERO, A., 1970. — Contribucion al conocimiento de la helmintofauna de los saurios cubanos. Ciencias

(Univ. Habana), 4 : 1-50

DURETTE-DESSET, M. Cl., & A. G. CHABAUD, 1981. — Nouvel essai de classification des Nématodes Trichos¬

trongyloidea. Ann. Parasit. Hum. Comp., 56 : 297-312.

DURETTE-DESSET, M.-Cl., 1985. — Trichostrongyloid Nematodes and their Vertebrate Hosts: Reconstruction of

the Phylogeny of a Parasitic Group. Adv. Parasit., 24: 239-306.

Durette-Desset, M.-Cl., A. K. Nasher & B. BEN Slimane, 1992. — Oswaldocruzia arabica n. sp. (Nematoda,

Trichostrongyloidea) parasite d’un Bufonidae de la péninsule arabique et remarques sur des espèces proches.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 14 : 693-703.

MORAVEC, C. F., & L. Vojtkova, 1975. — Variabilitât von zwei Nematodenarten Oswaldocruzia filiformis

(Goeze, 1782) und Oxysomatium brevicaudatum (Zeder, 1800), der Gemeinsamen parasiten der Europâischen

Amphibien und Reptilien. Scripta Fac. Sc. Nat. Ujep Brunensis, Biologia 2, 5 : 61-76.

Schmidt, G. D., & F. H. WHITTAKER, 1975. — Nematode parasites of Puerto Rican tree frogs, Eleutherodactylus

spp: two new species and a proposal of Poekilostrongylus gen. nov. (Trichostrongylidae). Parasitology, 70:

287-294.

TRAVASSOS, L., 1917. — Trichostrongylinas brazileiras (5 e nota previa). Oswaldocruzia n. gen. Brazil medico,

31: 9.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 77-82.

Oswaldocruzia peruensis n. sp. (Nematoda,Trichostrongylina,

Molineoidea), parasite d’un Iguanidae du Pérou

par Badreddine BEN SLIMANE, Manfred VERHAAGH

et Marie-Claude DURETTE-DESSET

Résumé. — Description A'Oswaldocruzia peruensis n. sp., parasite de Stenocercus roseiventris du Pérou.

L'espèce appartient aux Oswaldocruzia néotropicaux caractérisés : (1) par des spicules à trois branches principales

(lame, fourche et sabot) avec division de la fourche en deçà de son tiers distal et division de la lame en «peigne»

à son extrémité distale ; (2) par une disposition relative des côtes 6, 8 et dorsale de type II. La seule espèce

proche, Oswaldocruzia vaucheri Ben Slimane & Durette-Desset, 1993, parasite d’un Leptodactylidae d’Équateur,

s’en distingue par l’existence de crêtes et non d’ondulations cuticulaires et par la forme des ailes cervicales.

Mots-clés. — Oswaldocruzia peruensis n. sp., Nematoda, Trichostrongylina, Molineoidea, Iguanidae, Ste¬

nocercus , Pérou.

Oswaldocruzia peruensis n. sp. (Nematoda, Trichostrongylina, Molineoidea),

parasite of an Iguanidae from Peru

Abstract. — Description of Oswaldocruzia peruensis n. sp. parasite of Stenocercus roseiventris from Peru.

O. peruensis belongs to the group of Oswaldocruzia spp. characterized by : (1) three spicular branches (blade,

fork and shoe) with the fork subdivided at its distal extremity and the blade subdivided into many processes ;

and (2) by a type II bursal pattern of rays 6, 8 and dorsal. The only closely related species, O. vaucheri Ben

Slimane & Durette-Desset, 1993, a parasite of leptodactylids in Ecuador, differs by the presence of cuticular

crests which undulate and by the shape of the cervical alae.

Keywords. — Oswaldocruzia peruensis n. sp., Nematoda, Trichostrongylina, Molineoidea, Iguanidae, Ste¬

nocercus, Peru.

B. Ben Slimane et M.-Cl. Durette-Desset, Laboratoire de Biologie Parasitaire, Protistologie, Helminthologie, Muséum national

d’Histoire naturelle, 61, rue Bujfon, F - 75231 Paris Cedex 05.

M. Verhaagh, Staatliches Museum für Naturkunde Karlsruhe, Erbprinzenstrasse, 13 (Friedrichsplatzl, Postfach 6209, Karlsruhe

1, 76042 Allemagne.

INTRODUCTION

Au cours d’un travail de terrain sur les fourmis à la station biologique de Panguana (9°37’S,

74°56’W), Rio Yuya pichis, Departamento Huanuco, Pérou, l’un de nous a capturé un Stenocercus

roseiventris (Iguanidae) dans un vaste piège posé en lisière de la forêt primaire humide. Outre

des Nématodes Ascarides non encore étudiés, trouvés dans l’estomac, l’intestin grêle et le colon,

nous avons trouvé 14 Nématodes Trichostrongles dans le rectum. Il s’agit d’une espèce non

connue A'Oswaldocruzia Travassos, 1917, que nous décrivons ci-après.

— 78 —

MATÉRIEL ET MÉTHODES

L’estomac et l’intestin de l’hôte ont été séparés du corps et fixés, non ouverts, à l’alcool

à 70° au moment de la capture en octobre 1984. Les parasites ont été extraits au printemps

1993 et conservés dans l’alcool à 70°. Le matériel étudié est déposé dans les collections du

Muséum national d’Histoire naturelle de Paris (MNHN) et du Staatliches Museum für Naturkunde

Karlsruhe (SMNK). Le synlophe est étudié selon la méthode de Durette-Desset (1985); la

nomenclature utilisée pour la bourse caudale est celle de Durette-Desset & Chabaud (1981);

concernant la description des côtes 6, 8 et 9 (côte dorsale), celle de Durette-Desset, Nasher

& Ben Slimane (1992); concernant les spicules celle de Ben Slimane, Durette-Desset &

Chabaud (1993).

La nomenclature utilisée au-dessus du groupe famille est celle de Durette-Desset & Cha¬

baud (1993). Les mensurations de Tholotype et de l’allotype sont données en premier suivies

de celles des extrêmes des paratypes entre parenthèses.

DESCRIPTION

Oswaldocruzia peruensis n. sp.

Matériel-type. — Mâle holotype, femelle allotype, MNHN 758 MDa; 3 mâles, 5 femelles paratypes, MNHN

758 MDb. 2 mâles, 2 femelles paratypes SMNK - Ver 172.

Hôte : Stenocercus roseiventris Duméril et Bibron, 1837 (Iguanidae).

Localisation : rectum.

Origine géographique : Panguana. Pérou.

Collecteur: M. Verhaagh ; date de récolte: 7 octobre 1984.

Nématodes ne présentant pas d’enroulement du corps. Vésicule céphalique présente; pore

excréteur situé dans le tiers postérieur de l'œsophage, deirides peu développées situées en arrière

du pore excréteur, glandes excrétrices bien développées. Présence d’ailes cervicales. Synlophe

formé d’ondulations dépourvues de soutien chitinoïde.

Tête : vésicule céphalique simple. En vue apicale, on observe deux amphides, six papilles

labiales externes dont les deux latérales sont accolées aux amphides, quatre papilles céphaliques

et une dent œsophagienne dorsale. Bouche triangulaire (fig. 1 B).

Fig. 1. — A : mâle, partie antérieure, vue latérale gauche. B : mâle, tête, vue apicale. C-F : synlophe ; C : mâle, coupe transversale

en arrière des deirides ; D : femelle, id. ; E : femelle, coupe transversale au milieu du corps ; F : mâle, id. G : mâle, détail

du pore excréteur et des deirides, vue ventrale. H : femelle, ovéjecteur, vue latérale droite. I : femelle, queue, vue latérale

droite. J : mâle, cône génital, vue ventrale. K-M : spicule gauche disséqué ; K : vue extemo-latérale ; L : vue interne ; M :

vue ventrale. N : mâle, bourse caudale, vue ventrale. Toutes les coupes transversales sont orientées comme la fig. 1 C.

Abréviations : d. = dos, dr. = droite, g. = gauche, v. = ventre. Échelle : A, H, I, N = 40 (im ; B, E, J, K, L, M = 30 pm;

C, D, F, G = 50 |im.

A : male, anterior part, left lateral view. B: male, head, en face view. C-F: synlophe; C: male, transversal section posterior

to the deirides; D : female, id.; E: female, synlophe at mid-body; F: male, id. G: male, excretory pore and deirids, ventral

view. H: female, ovejector, right lateral view. I: female, tail, right lateral view. J: male, genital cone ventral view. K-M:

dissected left spicule; K: extemo-lateral view; L : intemo-lateral view; M: ventral view. N: male, caudal bursa, ventral view.

All the body sections are orientated as the fig. 1 C. Abbreviations: d. = dorsal side; dr. = right side; g. = left side; v. =

ventral side. Scale: A, H, I, N = 40 | im; B, E, J, K, L, M = 30\im; C, D, F, G = 50 \im.

— 80 —

Synlophe (étudié, en coupe transversale, chez un mâle et une femelle) : dans les deux sexes,

corps parcouru longitudinalement par des lignes cuticulaires sous forme d’ondulations visibles

seulement en coupe transversale, sauf dans la partie antérieure où les ondulations dorsales sont

plus fortes, donc visibles in toto. Les ondulations dorsales naissent à la base de la vésicule

céphalique tandis que les ventrales n’apparaissent qu’à environ 300 pm en arrière de la vésicule

céphalique chez le mâle et 270 pm en arrière de la jonction œsophago-intestinale chez la femelle.

Toutes les ondulations disparaissent au niveau de la bourse caudale chez le mâle et au niveau

des phasmides chez la femelle.

Les ailes cervicales, arrondies en coupe transversale, naissent à 90-130 pm (mâle), 70-

150 pm (femelle) en arrière de la vésicule céphalique et sont longues de 410-540 pm (mâle),

340-510 pm (femelle). En coupe transversale, la hauteur maximale de l’aile se situe entre le

niveau de l’anneau nerveux et celui du pore excréteur. Cette hauteur est de 12,5 pm chez le

mâle (fig. 1 C) et 13,8 pm chez la femelle (fig. 1 D). Les ailes sont en position latero-ventrale

et orientées vers la face ventrale.

Au milieu du corps, le nombre d'ondulations est de 37 chez le mâle (18 dorsales, 19 ven¬

trales) (fig. 1 F) et de 43 chez la femelle (22 dorsales et 21 ventrales) (fig. 1 E).

Mâle: long de 6 950 (7 500-8 500) pm sur 100 (110-130) pm de large dans sa partie

moyenne. Vésicule céphalique simple, haute de 90 (90-100) pm sur 40 (50-50) pm de large.

Anneau nerveux, pore excréteur et deirides situés respectivement à 200 (220-240) pm, 350 (390-

410) pm et 370 (410-430) pm de l’apex. Œsophage long de 520 (530-620) pm (fig. 1 A).

Bourse caudale de type 2-3 à tendance 2-1-2, c’est-à-dire que les extrémités des côtes 4

étant dirigées vers l’avant sont plus proches de celles des côtes 3 que de celles des côtes 5.

Côtes 2-3 d’une part et 5-6 d’autre part, jointives; disposition relative des côtes 6, 8 et 9 (côte

dorsale) de type II, les côtes 8 naissant apicalement sur la côte dorsale et étant chevauchées

ventralement par les côtes 6 sur leur partie médiane. Les côtes 9 se détachent de la côte dorsale

avant la division de celle-ci en deux rameaux dont l’interne est le plus long (fig. 1 N). Guber-

naculum absent, cône génital haut de 20 sur 20 pm de large dans sa partie proximale, portant

sur sa lèvre antérieure une grosse papille 0 et deux minuscules papilles 7 sur sa lèvre postérieure

(fig. 1 J).

Spicules subégaux, longs de 180 (200-250) pm, divisés en deçà de leur tiers proximal en

trois branches : lame, fourche et sabot; la lame se divise en «peigne» d’abord en 3 rameaux

qui se ramifient en 9 pointes (fig. 1 K). Sur les trois spicules disséqués, la fourche se divise à

23, 25 et 26% de la hauteur totale des spicules, en 2 branches de longueur inégale et de forme

différente, l’interne étant plus courte et ramifiée (fig. 1 M).

Femelle : longue de 10 100 (8 700-13 550) pm sur 120 (100-150) pm de large dans sa partie

moyenne. Vésicule céphalique haute de 90 (80-100) pm sur 40 (40-50) pm de large. Anneau

nerveux, pore excréteur et deirides situés respectivement à 220 (200-250) pm, 390 (350-410) pm

et 410 (370-450) pm de l'apex. Œsophage long de 550 (510-620) pm.

Didelphie. La vulve s’ouvre à 3500 (2 800-4 450) pm de la pointe caudale. Vagina vera

long de 50 (50-50) pm divisant le vestibule long de 370 (320-390) pm en deux parties inégales,

l’antérieure étant plus grande de 30 (10-30) pm. Sphincter, trompe et branche utérine antérieurs

longs respectivement de 30 (30-30) pm, 30 (25-30) pm et 2 000 (1 750-2 650) pm. Sphincter,

trompe et branche utérine postérieurs longs respectivement de 30 (30-30) pm, 30 (25-30) pm et

— 81 —

2 100 (1 500-2 700)pm. Les œufs, embryonnés, hauts de 90 (80-85) pm sur 40 (40-45) pm de

large, sont au nombre de 40 (20-45) dans la branche utérine antérieure et 38 (20-50) dans la

postérieure (fig. 1 H). Queue longue de 180 (160-240) pm sur 60 (60-75) pm de large au niveau

de l’anus, se terminant par une pointe caudale longue de 20 (16-20) pm (fig. 1 I).

DISCUSSION

Les spécimens de l’iguane présentent les principaux caractères du genre Oswaldocruzia Tra-

vassos, 1917 et sont caractérisés, comme la grande majorité des espèces néotropicales de la

sous-zone guyano-brésilienne, par des spicules à trois branches principales dont la fourche est

divisée en deçà de son tiers postérieur et dont la lame se termine en «peigne».

Parmi ces espèces, seule O. vaucheri décrit par Ben Slimane et DURETTE-DESSET, 1993,

parasite d’un Leptodactylidae d’Équateur, possède, comme nos spécimens, une vésicule cépha¬

lique simple, une disposition des côtes 6, 8 et 9 (côte dorsale) de type II et des ailes cervicales.

Elle s’éloigne de l’espèce du Pérou par la présence de crêtes cuticulaires bien marquées, même

si elles sont dépourvues de soutien chitinoïde et par des ailes cervicales en forme de demi-lune.

Nous séparons donc les spécimens de l’Iguanidae dans une nouvelle espèce que nous pro¬

posons de nommer Oswaldocruzia peruensis n. sp.

Remerciements

Le second auteur est reconnaissant au Pr H. W. Koepcke et à sa fille le Dr J. Diller de

l’avoir autorisé à rester au Panguana pendant plus de deux ans et au Directeur général de la

Forêt et de la Faune de Lima de lui avoir accordé l’autorisation de faire des recherches au

Pérou. Le travail au Panguana a été financé par une subvention de la DAAD (German Academie

Exchange Service).

RÉFÉRENCES

BEN SLIMANE, b., & M.-Cl. Durette-Desset, 1993. — Quatre nouvelles espèces du genre Oswaldocruzia

Travassos, 1917 (Nematoda : Trichostrongyloidea) parasites d’Amphibiens d’Equateur. Rev. S. Zool., 100 :

113-136.

Ben Slimane, B., M.-Cl. Durette-Desset & A. G. Chabaud, 1993. — Oswaldocruzia (Trichostrongyloidea)

parasites d’Amphibiens des Collections du Muséum de Paris. Ann. Parasil. Hum. Comp., 68 : 88-100.

DURETTE-DESSET, M.-CL, 1985. — Trichostrongyloid Nematodes and their Vertebrate hosts : reconstruction of

the phylogeny of a parasitic group. Adv. Parasitol ., 24 : 239-306.

DURETTE-DESSET, M.-Cl. & A. G. CHABAUD, 1981. — Nouvel essai de classification des Nématodes Trichos¬

trongyloidea. Ann. Parasit. Hum. Comp.. 56 : 297-312.

— 1993. — Nomenclature des Strongylida au-dessus du groupe famille. Ann. Parasit. Hum. Comp.. 68 :111-112.

DURETTE-DESSET, M.-CL, A. K. Nasher & B. Ben slimane, 1992. — Oswaldocruzia arabica n. sp (Nematoda,

Trichostrongyloidea) parasite d’un Bufonidae de la péninsule arabique et remarques sur des espèces proches.

Bull. Mus. natl. Hist. nat. Paris, 4 e sér., 14 : 693-703.

TRAVASSOS, L., 1917. — Trichostrongylidae brazileiras. Oswaldocruzia n. gen. Brazil Medico, 31 : 3-4.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2: 83-86.

Identity of the Antarctic mite Gainia nivalis Trouessart

(Acari, Nanorchestidae)

by Mark JUDSON

Abstract. — The types of Gainia nivalis Trouessart, 1914, from Petermann Island (Antarctic Peninsula),

have been re-examined and a lectotype designated. Gainia Trouessart, 1914, is confirmed to be a junior subjective

synonym of Nanorchestes Topsent and Trouessart, 1890. Nanorchestes nivalis (n. comb.) is a senior subjective

synonym of N. gressitti Strandtmann, 1982 (n. syn.). Authorship of the family name Nanorchestidae is attributed

to Berlese, 1913.

Keywords. — Acari, Nanorchestes, Gainia, Antarctic.

Identité de l’acarien antarctique Gainia nivalis Trouessart

(Acari, Nanorchestidae)

Résumé. — Les types de Gainia nivalis Trouessart, 1914, provenant de Tile Petermann (Péninsule Antarc¬

tique), ont été réexaminés et un lectotype désigné. Gainia Trouessart, 1914, est confirmé comme synonyme sub¬

jectif de Nanorchestes Topsent et Trouessart, 1890. Nanorchestes gressitti Strandtmann, 1982, est un synonyme

subjectif de N. nivalis (n. comb., n. syn.). La paternité du nom de famille Nanorchestidae est attribuée à Berlese,

1913.

Mots-clés. — Acariens, Nanorchestes, Gainia, Antarctique.

M. Judson, Muséum national d'Histoire naturelle. Laboratoire de Zoologie (Arthropodes), 61, rue Buffon, F-75231 Paris Cedex

05.

INTRODUCTION

Mites of the cosmopolitan genus Nanorchestes Topsent & Trouessart (Nanorchestidae) form

an important and characteristic component of the Antarctic fauna. Of the thirty-or-so species

known world-wide, fourteen have been recorded from the Antarctic and sub-Antarctic regions

(Pugh, 1993) 1 , including the southernmost-living terrestrial arthropod (Gressitt, 1967a).

The genus Gainia was created by Trouessart (1914) for a new species, Gainia nivalis

Trouessart, collected from Petermann Island off the coast of the Antarctic Peninsula. Trouessart

placed it in the Alycinae, which at that time contained the genera Alycus C. L. Koch, Michaelia

Berlese, Nanorchestes and Hybalicus Berlese, stating that the new genus was closest to Michaelia

(- Bimichaelia Thor). WlLLMANN (in Thor & WlLLMANN, 1941) suggested that Gainia might

1. This high proportion of Antarctic species is probably due to the fact that relatively little attention has been paid to

the systematics of Nanorchestes species from other regions of the world.

— 84 —

be a synonym of Nanorchestes, a view reiterated by Strandtmann (1967) and Pugh (1993).

THERON (1975) listed Gainia as a synonym of Nanorchestes, without comment.

Because of the inadequacy of the original description, the identity of Gainia nivalis has

remained doubtful and there have been no further records of this species. Gressitt (1967 b:

382) suggested that G. nivalis might be identical with Nanorchestes antarcticus Strandtmann.

In fact the latter has since been shown to be a polyphyletic assemblage of species (Strandtmann,

1982). Fortunately, Trouessart's types are still present in the Muséum national d' Histoire na¬

turelle, Paris, and a re-examination of this material shows that it is conspecific with Nanorchestes

gressitti Strandtmann, 1982, one of the species previously confused with N. antarcticus.

Family NANORCHEST1DAE Berlese, 1913 2

Genus NANORCHESTES Topsent & Trouessart, 1890

Nanorchestes TOPSENT & TROUESSART, 1890: 892 (type species Nanorchestes amphibius

Topsent & Trouessart, 1890, by monotypy).

Gainia TROUESSART, 1914: 13-14 (type species Gainia nivalis Trouessart, 1914, by mono¬

typy); Thor & Willmann, 1941: 148. Synonymized by Theron, 1975: 1.

Remarks

Although Gainia is clearly a synonym of Nanorchestes as currently recognized, it might

perhaps be revalidated in the future. Nanorchestes is a relatively large genus, including about

30 species, and might usefully be split into smaller generic or subgeneric taxa. McDaniel &

Bolen (1981) named a closely related genus, Neonanorchestes, for two North American species

with clavate trichobothria nb. The validity of Neonanorchestes was questioned by BOOTH (1984),

but the recent discovery of alternating calyptostasy in this genus (Kethley, 1991) suggests that

it may be monophyletic. This leaves, however, the question as to whether Nanorchestes itself

represents a natural group. As both the clavate trichobothria and the alternating calyptostasis of

Neonanorchestes probably represent autapomorphies, it is possible that the recognition of this

genus leaves Nanorchestes paraphyletic. At least one species currently placed in Nanorchestes

-N. siculus (Berlese, 1910)- has clavate trichobothria, though its nymphs are as yet unknown.

Nanorchestes nivalis (Trouessart) n. comb.

Gainia nivalis TROUESSART, 1914: 14, Figs. 8-9; THOR & WILLMANN, 1941: 148-149,

Fig. 206; Dalenius, 1965: 421; Gressitt, 1967a: 5, Table 2; Gressitt, 1967b: 382; Strandt¬

mann, 1967: 79; Block, 1992: 161, 167; Pugh, 1993: 339.

Nanorchestes antarcticus (not Strandtmann), Strandtmann, 1967: 7, Fig. 14g (in part,

“variant 4” from South Orkney Islands).

2. Although authorship of the family Nanorchestidae has traditionally been accredited to GRANDJEAN (1937), the tribe

Nanorchestini was proposed by BERLESE (1913: 17), who is therefore the author of the co-ordinate family name (ICZN

Code, article 36).

— 85 —

Nanorchestes gressitti Strandtmann, 1982d: 253-256, Figs 1-17; CENXUAN & SHIMADA,

1991: 326, 330, Fig. ID; Pugh, 1993 : 338. New synonymy.

Material examined. — Lectotype 3 (MNHN, slide 63B9), paralectotypes 2 3,1$ (MNHN, slide 45C13)

and 8 other specimens (MNHN. slides 45B2 and 45C12), all labelled “Charcot, sur neige rouge, I. Petermann,

Antarctique.” The original identification labels read : “Gainia nivalis n. g.- n. sp. (Rostre. Type)” [45B2]; "Gainia

nivalis Trt. (Téguments)” [45C12]; and “Gainia nivalis Trt. (Profil)” [4503]; each label is also marked with a

cross in red pencil.

Remarks

Although only one of the three original slides was labelled “type”, it is clear that all of

the specimens are from the original collection and should be regarded as syntypes. The name

‘Charcot’ on the labels refers to Dr J.-B. Charcot, leader of the Second French Antarctic Ex¬

pedition (1908-1910). Petermann Island (65°10’S 64°10’W) was visited by the expedition in

March 1909, where the mites were collected by the expedition’s biologist, L. Gain (Trouessart,

1914).

The types were in poor condition when Trouessart received them, and have deteriorated

since. They were strongly compressed beneath the coverslips and the mountant had dried out

around them, so that little other than their silhouettes can be seen. The four specimens (3 6,

1 5) on slide 45C13 were temporarily de-mounted and examined in lactic acid; one of these

males has been selected as the lectotype and is now mounted on a separate slide (63B9).

Although a complete examination was not possible, sufficient details became visible to show

that the types are conspecific with Nanorchestes gressitti Strandtmann, known from Anvers Island

(Antarctic Peninsula), South Shetland Is, South Orkney Is, South Sandwich Is and South Georgia

Is. Apparent discrepancies between the descriptions of Trouessart (1914) and Strandtmann

(1982) are due to misinterpretations by Trouessart. In particular, the form of the chelicerae,

trichobothria and the apoteles of the legs are all typical of Nanorchestes. The types of nivalis

agree with Strandtmann’s description (as N. gressitti ) in the following :

1. Size : Trouessart gave the body length as being 0.5 mm, presumably including the

gnathosoma. The idiosomal length of the 12 specimens (probably mostly adult) varies from about

320 to 450 pm, though most are between 360 and 400 pm long. STRANDTMANN (1982) records

a range of 270-400 pm for adults. Allowing for the compression of the types of nivalis, these

figures are reasonably similar;

2. “Wings” of naso narrow and widely separated;

3. Trichobothrium na long (80 pm), strongly ciliated on distal third; nb long (65 pm), finely

ciliated, without apical branches;

4. Cheliceral seta cha bifurcate, branches roughly equal in length;

5. Anterior seta of telofemur I with only a few, reduced cilia at apex;

6. Tarsus III with 8 setae.

These characters, at least in the case of the specimens examined in detail, rule out the

possibility that types of nivalis are conspecific with N. berryi Strandtmann, a species recorded

as being sympatric with N. gressitti at two localities on the nearby Anvers Island by STRANDT¬

MANN (1982). However, as it is not possible to state with certainty that the remaining types are

all conspecific, it seems prudent to designate a lectotype for nivalis.

— 86 —

Acknowledgements

My thanks are due to: M. H. Naudo (MNHN, Paris) for help in locating the types of

Gainia nivalis and permission to remount some of them; Dr P. J. A. PUGH (British Antarctic

Survey, Cambridge) for providing relevant literature; and Prof. R.W. Strandtmann for com¬

menting on the manuscript.

REFERENCES

BERLESE, A., 1913. — Acarotheca Italica. Fasciculi l“ s et II US , Praemisso Acarorum systemate genera omnia in

suas Familias referente. M. Ricci, Florence: 221 pp.

BLOCK, W., 1992. — An annotated bibliography of Antarctic invertebrates (terrestrial and freshwater). British

Antarctic Survey, Cambridge: 263 pp.

BOOTH, R. G., 1984. — A new species of mite in the genus Nanorchestes (Acari : Prostigmata) from Signy

Island, South Orkney Islands. Br. antarct. Surv. Bull., 63: 111-116.

CENXUAN, P., & K. Shimada, 1991. — Cold hardiness of four Antarctic terrestrial mites in the active season

at King George Island. J. Insect Physiol., 37: 325-331.

Dalenius, P., 1965. — The acarology of the Antarctic regions. In MlEGHEM, J. van & OYE, P. van (eds).

Biogeography and ecology in Antarctica. Monographiae Biologicae, 15: 414-430. Dr W. JUNK, The Hague.

GRANDJEAN, F., 1937. — Le genre Pachygnathus Dugès (Alycus Koch) (Acariens). Cinquième et dernière partie.

Bull. Mus. natl. Hist. nat. Paris (2) 9 : 262-269.

GRESSITT, J. L., 1967a. — Introduction. In: GRESSITT, J. L. (ed.), Entomology of Antarctica. Antarct. Res. Ser.,

10: 1-33, American Geophysical Union, Baltimore.

— 19676. — Notes on arthropod populations in the Antarctic Peninsula-South Shetland Islands-South Orkney

Islands area. In: GRESSITT, J. L. (ed.), Entomology of Antarctica. Antarct. Res. Ser., 10: 373-391, American

Geophysical Union, Baltimore.

KETHLEY, J., 1991. — Calyptostasic nymphs of Neonanorchestes (Nanorchestidae): a third example of alternating

calyptostasy. In: DUSBÂBEK, F. & Bukva, V. (eds). Modern Acarology, vol. 2: 279-282. SPB Academic

Publishing, The Hague.

McDaniel, B., & E. G. BOLEN, 1981. — A new genus and two new species of Nanorchestidae from Padre

Island, Texas (Acari: Prostigmata). Acarologia, 22: 253-256.

PUGH, P. J. A., 1993. — A synonymic catalogue of the Acari from Antarctica, the sub-Antarctic Islands and the

Southern Ocean. J. nat. Hist., 27: 323-421.

STRANDTMANN, R. W., 1967. — Terrestrial Prostigmata (trombidiform mites). In: GRESSITT, J. L. (ed.). Ento¬

mology of Antarctica. Antarct. Res. Ser., 10: 51-80. American Geophysical Union, Baltimore.

— 1982. — Notes on Nanorchestes. V. Two new species of Nanorchestes (Acari: Nanorchestidae) from the

Antarctic Peninsula and South Atlantic islands. Pac. Insects, 24: 252-258.

THERON, P. D., 1975. — Two new species of the family Nanorchestidae (Acari: Endeostigmata) from pasture

soil in South Africa. Wet. Bydraes Potchefstroom Univ., (B) 63: 1-9.

THOR, S., & C. Willmann, 1941. — Acarina. Eupodidae, Penthalodidae, Penthaleidae, Rhagidiidae, Pachyg-

nathidae, Cunaxidae. Tierreich, 71a: i-xxviii, 1-186, Walter de GRUYTER & Co., Berlin.

TOPSENT, E., & E.-L. TROUESSART, 1890. — Sur un nouveau genre d’Acarien sauteur ( Nanorchestes amphibius)

des côtes de la Manche. C.r. Acad. Sci., Paris, 61: 891-892.

TROUESSART, E.-L., 1914. — Acariens. In : Deuxième expédition antarctique française (1908-1910), commandée

par le Dr Jean Charcot. Sciences naturelles : documents scientifiques : 1-16. Masson et Cie, Paris.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 87-94.

Description de Bordea, nouveau genre endémique

d’araignées des Pyrénées (Araneae : Linyphiidae)

par Robert BOSMANS

Résumé. — Le nouveau genre Bordea, endémique des Pyrénées, est créé pour les espèces Lepthyphantes

cavicola (Simon) et Lepthyphantes negrei Dresco. Il se distingue de Lepthyphantes principalement par les den-

ticules du tibia de la patte-mâchoire du mâle et le crochet long, non plié de la femelle. Les synonymes et combi¬

naisons suivants sont proposés : Lepthyphantes coijfaiti Denis, 1953 = Lepthyphantes cavicola (Simon, 1884) n.

syn. = Bordea cavicola (Simon) n. comb. Lepthyphantes ecclesiasticus Denis, 1959 = Lepthyphantes negrei Dresco,

1951 n. syn. = Bordea negrei (Dresco) n. comb.

Mots-clés. — Linyphiidae, Lepthyphantes , Pyrénées.

Description of Bordea, a new endemic spider genus

from the Pyrénées (Araneae : Linyphiidae)

Abstract. — To receive the species Lepthyphantes cavicola (Simon) and Lepthyphantes negrei Dresco, the

new genus Bordea is created, endemic to the Pyrénées, distinguished from Lepthyphantes by the tibial tubercles

in the male palp, and by the long, unfolded scape in the female. The following new synonyms and combinations

are proposed: Lepthyphantes coijfaiti Denis, 1953 = Lepthyphantes cavicola (Simon, 1884) n. syn. = Bordea

cavicola (Simon) n. comb.; Lepthyphantes ecclesiasticus Denis, 1959 = Lepthyphantes negrei Dresco, 1951 n.

syn. = Bordea negrei (Dresco) n. comb.

Keywords. — Linyphiidae, Lepthyphantes, Pyrénées.

Robert Bosmans, Laboratorium voor Ecologie der Dieren, Zoogeografie en Natuurbehoud. K. LI. Ledeganckstraat 35, B-9000

Gent, België.

INTRODUCTION

Considéré polyphylétique, le genre Lepthyphantes Simon compte actuellement environ

440 espèces. Saaristo & Tanasevitch (1992) ne distinguèrent pas moins de 31 groupes d’es¬

pèces. Malgré la création de nouveaux genres pour certains groupes (Eskov & Marusik, 1992 :

Parawubanoides ; TANASEVITCH, 1992 : Himalaphantes, Incestophantes, Herbipliantes et Cris-

piphantes; WUNDERLICH, 1993 : Megalepthyphantes) ou une meilleure délimination de certains

complexes d’espèces (Helsdingen et al., 1977 : groupe tenuis; Thaler, 1994: groupe man-

suetus; Thaler et al., 1994 : groupe annulatus ), la systématique du genre Lepthyphantes reste

problématique.

En révisant les Lepthyphantes de France, nous avons trouvé deux espèces «oubliées» par

Saaristo & Tanasevitch (ap. cit.) qu’ils ne classèrent dans aucun de leurs groupes. La mor¬

phologie des organes sexuels secondaires de ces espèces est très différente de celle des autres

groupes. Nous proposons la création d’un nouveau genre pour ces deux espèces.

— 88 —

Abréviations : MNHNP : Muséum national d’Histoire naturelle de Paris; MNCNM : Museo

nacional de Ciencias naturales de Madrid.

DESCRIPTION

BORDEA n. gen.

Espèce-Type. — Bordea cavicola (Simon).

Étymologie. — Le genre est dédié à L. Borde en reconnaissance.

Diagnose

Linyphiidae de petite taille (1,8-2,5 mm). Couleur: céphalothorax, sternum, chélicères et

pattes jaune orange ; abdomen blanchâtre.

Céphalothorax sans protubérances ; yeux normalement développés, presque équidistants.

Chélicères pourvu de 3 denticules antérieurs et 5 postérieurs. Organe de stridulation composé

de 20 stries peu évidentes.

Chétotaxie : fémur I pourvu d’une épine prolatérale, fémora II-IV mutiques ; tibia muni de

deux séries de 3 épines, chaque série composée d’une épine dorsale, une prolatérale et une

rétrolatérale ; métatarse pourvu d’une épine dorsale.

Pédipalpe (fig. 1-4, 9-11) : crin dorsal de la patella et du tibia faible, celui du tibia plus

long, celui de la patella à peine plus long que le diamètre; tibia présentant au côté externe

quelques denticules noirs; cymbium pourvu d'une courte saillie conique rétrolatérale; paracym-

bium pourvu dans sa concavité d’une carène ou d’une dent; apophyse supratégulaire se terminant

en deux denticules ; lamelle volumineuse, composée de deux branches inégales ; embolus pointu,

accompagné par un lobe latéral; tube séminifère pourvu d’une glande de Fickert composée de

deux ampoules, l’une ovale, l’autre arrondie, puis se dirigeant tout droit vers l’embolus.

Épigyne (fig. 5-7, 12-14) pourvue d’un crochet très long ni plié ni aplati, non divisé en

languette extérieure et languette intérieure comme chez Lepthyphantes; crochet beaucoup plus

long que les plaques latérales; plaque médiane proéminente. Vulve (fig. 8, 15) : spermathèques

ovales, canaux spermatiques se dirigeant d’abord vers le milieu de la base du crochet, puis tout

droit vers son sommet, où se situe l’orifice de copulation.

Le genre est très voisin de Lepthyphantes et des autres genres cités ci-dessus classés au¬

paravant dans Lepthyphantes. Il en diffère seulement par les organes sexuels secondaires. Les

denticules du tibia du pédipalpe, la forme remarquable de la glande de Fickert, et le crochet de

l’épigyne non plié ou aplati sont les caractères principaux qui distinguent ce nouveau genre de

l’espèce-type de Lepthyphantes.

Bordea cavicola (Simon) n. comb.

(Fig. 1-8)

Porrhomma cavicola Simon, 1884 : 355 (descr. S, 9).

Lepthyphantes cavicola Simon, 1929: 736; DRESCO, 1951 : 364.

— 89 —

Fig. 1-8. Bordea cavicola n. comb. 1, pédipalpe mâle, vue latérale; 2, idem, vue dorsale: 3, apophyse supratégulaire ; 4,

division embolique; 5, épigyne, vue ventrale; 6, idem, vue latérale; 7, idem, vue dorsale; 8, vulve. Echelle : 0,25 mm.

Bordea cavicola n. comb. 1, male palp, lateral view ; 2, idem, dorsal view; 3, suprategular apophysis; 4, embolic division;

5, epigyne, ventral view; 6, idem, lateral view; 7, idem, dorsal view; 8, vulva. Scale: 0.25 mm.

— 90 —

Lepthyphantes coiffaiti Denis, 1953 : 264 (descr. <3). n. syn.

Matériel étudié et citations. — FRANCE : Basses-Pyrénées : Arudy, vallon de Molarode, holotype 6 de

L. coiffaiti (Denis, 1953; non examiné). Arudy, bois du Bager, paratype â (Denis, 1953). Pyrénées-Orientales :

col de Mentet, 1 600 m, 1 S sous une pierre, 10 juin 1982, R. Bosmans leg. (collection R. Bosmans). - ESPAGNE :

Navarra : Alsasua, Cueva de Orobe, lectotype $ de Porrhomma cavicola, par présente désignation. Paralectotypes :

1 2 appartenant à L. cavicola , 1 2 appartenant à L. negrei (MNHNP 4983).

Distribution. — Pyrénées espagnoles et françaises.

Diagnose

L’espèce est facilement reconnaissable par les grands denticules du tibia et la lamelle très

allongée du pédipalpe chez le mâle, par le crochet aussi long que large de l’épigyne chez la

femelle.

Discussion

Dans la description originale, Simon (1884), figure le pédipalpe du mâle et l’épigyne de

la femelle. Pourtant, un mâle n’est pas présent dans la série type, comme déjà constaté par

Dresco (1951). Selon lui, cette série type était composée de cinq femelles, mais nous n’en

avons retrouvé que trois. Dresco écrit : «Une des femelles a un crochet différent des cavicola

typiques...». Il la prend pour une forme anormale, mais nous croyons qu’il s’agit ici d’un individu

appartenant à L. negrei.

En 1953, Denis décrivit L coiffaiti. Dans sa diagnose, il indique que l’espèce est très voisine

de L. negrei, traitée ci-dessous, elle-même très proche de L. cavicola. Les types mâles de L. ca¬

vicola et de L. coiffaiti ne sont pas disponibles, mais d’après Simon (1884 : 356, fig. 128) et

Denis (1953 : 265, fig. 3), et d’après un individu mâle capturé par nous-même, les deux espèces

sont identiques, surtout en ce qui concerne la forme typique de la lamelle. Nous les considérons

donc comme des synonymes.

Bordea negrei Dresco n. comb.

(Fig. 9-15)

Lepthyphantes negrei Dresco 1951 : 365 (descr. S, 9); DENIS, 1967 : 44, 47; DRESCO &

Hubert, 1968 : 493.

Lepthyphantes cavicola Machado 1940: 9 (identification incorrecte).

Lepthyphantes ecclesiasticus Denis 1959: 223; DRESCO 1962: 185. n. syn.

Lepthyphantes ictericus Simon 1929 : 599, 737 (descr. 3, non 9).

Matériel étudié et citations. — FRANCE : Ariège : gouffre de Peillot, 1 6, 1 2, 15 juillet 1945, série

type de L. negrei (MNHNP); Montagagne, gouffre de Soulisquet, 1 2,4 juin 1959 (MNHNP L398) ; étiquetté

L. ecclesiasticus. mais cité par Denis (1967) comme L. negrei. Haute-Garonne : Montmaurin, grande grotte de

la Seygouade (Denis, 1967). Hautes-Pyrénées : Bas-Nistos, grotte de l’Église, holotype femelle de L. ecclesiasticus

(Denis, 1959); grotte de l'Ours (Dresco, 1951); grotte Soulabère (Dresco, 1951); grotte de la Pale, Saint-Pré-

de-Bigorre, 1 2, octobre 1963 (Dresco & Hubert, 1968). — ESPAGNE: Navarra: Larraun, Cueva de Mar-

tinchurrito, 1 2, 22 août 1919 (sub. L. cavicola, MNCNM 53; Machado, 1940); Alsasua, Cueva de Orobe, 12

paralectotype de L. cavicola (MNHNP 4983).

— 91 —

Fig 9-15. — Bordea negrei n. comb. 9, pédipalpe mâle, vue latérale; 10, idem, vue dorsale; 11, division embolique; 12, épigyne,

vue ventrale; 13, idem, vue latérale; 14, idem, vue dorsale; 15, vulve. Échelle : 0,25 mm.

Bordea negrei n. comb. 9, male palp, lateral view; 10, idem, dorsal view; 11, embolic division; 12, epigyne, ventral view,

13, idem, lateral view; 14, idem, dorsal view; 15, vulva. Scale: 0.25 mm.

— 92 —

Diagnose

Très voisine de B. cavicola. Les mâles diffèrent par les denticules du tibia moins prononcés,

la dent de la branche distale du paracymbium non obtuse mais aiguë et la lamelle moins allongée ;

les femelles par le crochet beaucoup plus long que large en vue ventrale et par la plaque ventrale

moins proéminente.

Discussion

Bordea negrei fut excellemment décrite et illustrée par DRESCO (1951) qui indique que

l'espèce est voisine de B. cavicola et dont il donne également une figure de l’épigyne.

En 1959, Denis décrit des Pyrénées une femelle qu’il nomme Lepthyphantes ecclesiasticus.

Déjà Dresco (1962) douta de la validité de cette espèce, se demandant s’il ne s’agissait pas

d’un synonyme de L. negrei. En effet, les figures de Dresco (1951) de l’épigyne de L. negrei

et de Denis (1959) de l’épigyne de L. ecclesiasticus sont presque identiques. On a d’ailleurs

retrouvé au MNHNP un individu identifié par Denis portant une étiquette «L. ecclesiasticus»,

mais cité par lui en 1967 comme L. negrei. Il est probable que DENIS ait constaté la synonymie,

mais sans jamais la publier lui-même.

Un autre cas intéressant et énigmatique est celui d’un mâle et de deux femelles, décrits

comme Lepthyphantes ictericus par SIMON (1929) de la grotte de l’Ours dans les Hautes-Pyrénées.

SIMON figura l’épigyne de la femelle ressemblant beaucoup à un Centromerus. L’examen de

cette femelle a confirmé ce point de vue. Le mâle fut perdu accidentellement (Simon, 1929 :

599, en note). Son pédipalpe n’a pas été représenté mais dans la clé, les deux espèces sont

distinguées :

— Tibia, vu de profil, élevé en cône vertical, pourvu, sur sa pente antérieure (assez

loin de l’angle), d’un petit denticule noir obtus; crin dressé subapical; paracym¬

bium noir, pourvu dans sa concavité, vers le milieu, d’une petite apophyse

comprimée tronquée; tarse gros, offrant au bord externe (avant l’échrancure nor¬

male) une courte saillie conique . L. ictericus

— Tibia, vu de profil, légèrement convexe, armé, à l’angle apical, d’un petit den¬

ticule fauve, conique, dirigé en avant, et, un peu plus en arrière, de quelques

petites granulations noires piligères. Paracymbium très fortement courbé en de¬

mi-cercle, mutique au bord interne de sa concavité, sa branche antérieure chi-

tinisée rouge passant au noir à l’extrémité. Lanière du bulbe vue de profil

d’abord dirigée en avant puis coudée presque verticalement, enfin coudée en

avant en forme de Z. L. cavicola

Nous pensons que cette description de L. cavicola correspond à celle de L. negrei, autre

habitant de la grotte de l’Ours (Dresco, 1951). Le mâle et la femelle de L. ictericus ne sont

donc pas conspécifiques. La femelle appartient à une espèce voisine des Centromerus, et le mâle

est celui de B. negrei.

Le mâle retrouvé au MNCNM sous le nom Lepthyphantes cavicola et cité sous ce nom par

MACHADO (1940) concerne également Bordea negrei.

Dans la série type, ont été retrouvés dans le même tube des individus de Bordea cavicola

et de B. negrei, provenant de la Cueva de Orobe, la seule localité citée par Simon. Les deux

espèces vivraient donc ensemble dans la même grotte. Ceci reste pourtant à confirmer, car Simon

avait l’habitude de mélanger des récoltes de différentes localités. Il se peut que des spécimens

de B. negrei capturés par Bolivar en 1919 et cités beaucoup plus tard par Machado (1940)

comme B. cavicola, aient été envoyés par Bolivar à Simon qui les aurait incorporés dans sa

collection.

Remerciements

Nous remercions vivement C. Rollard et J. Heurtault (MNHNP) et A. G. Valdecasas (MNCNM)

pour le prêt de spécimens.

RÉFÉRENCES

DENIS, J., 1953. — Quelques captures d’Araignées Pyrénéennes. Bull. Soc. Hist. nat. Toulouse , 88 : 259-266.

— 1959. — Quelques Araignées cavernicoles des Pyrénées. Annales de Spéléologie, 14 : 219-231.

— 1967. — Quelques captures d’Araignées Pyrénéennes. Bulletin de la Société d'Histoire naturelle de Toulouse,

103 : 32-50.

DRESCO, E., 1951. — Sur quelques Lepthyphantes cavernicoles et description d’une espèce nouvelle. Bulletin

Muséum national d'Histoire naturelle de Paris, 23 : 363-367.

— 1962. — Araignées capturées en France dans des grottes ou des cavités souterraines. Annales de Spéléologie,

17 : 177-193.

DRESCO, E., & M. HUBERT, 1968. — Araneae speluncarum Galliae. Annales de Spéléologie, 23 : 483-500.

ESKOV, K. Y., & Y. M. MARUSIK, 1992. — On the mainly Siberian spider genus Wubana, Wubanoides gen. n.

and Poeciloneta (Aranei: Linyphiidae). Arthropoda Selecta, 1 : 21-38.

HELSDINGEN, P. J. Van, K. Thaler & C. DELTSHEV, 1977. — The tenuis group of Lepthyphantes Menge

(Araneae, Linyphiidae). Tijdschrift voor Entomologie, 120 : 1-54.

Machado, A. de Barros, 1940. — A colecçao de Aranhas cavernicoles do Museu nacional de Ciencias naturais

de Madrid. Ciencias, 7 : Bulletin de la Société de Zoologie de France, 64 : 60-70.

SAARISTO, M. L, & A. V. TANASEVITCH, 1993. — Notes on the systematics of the spider genus Lepthyphantes

Menge (Aranei Linyphiidae Micronetinae). Arthropoda selecta, 2: 55-61.

SIMON, E., 1884. — Les Arachnides de France, V (2) : 180-420. Paris.

— 1929. — Les Arachnides de France, VI (3) : 533-772. Paris.

TANASEVITCH, A. V., 1992. — New genera and species of the tribe Lepthyphantini (Aranei Linyphiidae Micro¬

netinae) from Asia (with some nomenclatorial notes on linyphiids). Arthropoda selecta, 1 : 39-50.

THALER, K., 1994. — Vikariante Verbreitung im Artenkreis von Lepthyphantes mansuetus in Mitteleuropa

(Araneida : Linyphiidae). Entomologia Generalis, 18: 171-185.

THALER, K., P. j. Van HELSDINGEN & C. DELTSHEV, 1994. — Vikariante Verbreitung im Artencomplex von

Lepthyphantes annulatus in Europa und ihre Deutung (Araneae, Linyphiidae). Zoologischer Anzeiger, 232:

111-127.

WUNDERLICH, J., 1993. — Beschreibung der neuen Spinnen-Gattung Megalepthyphantes aus der Familie der

Baldachinspinnen und einer bisher unbekannten Art aus Griechenland (Arachnida: Araneae: Linyphiidae).

Entomologische Zeitung, 103 : 168-171.

Bull. Mus. natl. Hist, ncit., Paris, 4 e sér., 17, 1995

n° 1-2 : 95-106.

Description de trois nouveaux genres et quatre nouvelles espèces

de scorpions Buthidae de Madagascar

par Wilson R. LOURENÇO

Résumé. — Trois nouveaux genres, Pseudouroplectes, Microcharmus et Neogrosphus, et quatre nouvelles

espèces, Pseudouroplectes betschi, Microcharmus cloud sleythompsoni, Tityobuthus guillaumeti et Tityobuthus po-

cocki appartenant à la famille des Buthidae sont décrits de Madagascar. La découverte des trois nouveaux genres

représente une intéressante contribution pour la clarification des origines et des affinités de la faune malgache.

Mots-clés. — Scorpion, Madagascar, Buthidae, nouveaux genres.

Description of three new genera and four new species of Buthidae Scorpions from Madagascar

Abstract. — Three new genera, Pseudouroplectes, Microcharmus and Neogrosphus and four new species,

Pseudouroplectes bestchi, Microcharmus cloudsleythompsoni, Tityobuthus guillaumeti and Tityobuthus pococki

belonging to the Buthidae family are described from Madagascar. These three new genera represent an interesting

contribution to our understanding of the origins and affinities of the fauna of Madagascar.

Keywords. — Scorpion, Madagascar, Buthidae, new genera.

W. R. Lourenço, travail réalisé au Laboratoire de Zoologie (Arthropodes) du Muséum national d'Histoire naturelle, 61, rue

Buffon, F-75005 Paris.

INTRODUCTION

Dans le cadre d’une étude biogéographique sur les scorpions de Madagascar, j’ai été amené

à examiner les collections des scorpions provenant de Madagascar et déposées au Muséum na¬

tional d’Histoire naturelle, Paris.

Une grande partie des collections de Paris ont déjà servi à la description de nouveaux genres

et de nouvelles espèces notamment par Gervais (1844), Simon (1880) et Kraepelin (1901).

À la suite de la révision monographique de Fage (1929) et de la revue des Arachnides de

Millot (1948), la faune scorpionique de Madagascar semble assez bien étudiée et peu de nou¬

veaux taxa ont été ajoutés. Quelques compléments sont apportés par VACHON (1969, 1979).

La découverte de trois nouveaux genres, dont deux peuvent être classés parmi les micro-

Scorpions, augmente significativement le taux de taxa endémiques de la grande île, mais apporte

surtout des éléments nouveaux, très précieux, sur les affinités de la faune scorpionique malgache.

Dans le présent travail, je me limiterai à décrire les nouveaux taxa avec quelques commen¬

taires sur leurs probables positions phylogénétiques. L'ensemble du problème biogéographique

— 96 —

sera traité dans l’étude mentionnée ci-dessus qui sera présentée lors du Colloque international

de biogéographie de Madagascar qui se tiendra à Paris en septembre 1995.

DESCRIPTION

Genre PSEUDOUROPLECTES n g.

(Fig. 1, 4-8)

Diagnose

Scorpions de petite taille; longueur totale des adultes d’environ 22-25 mm. Coloration gé¬

nérale jaunâtre avec quelques taches rougeâtres esquissées sur le mésosoma; anneau V du mé-

tasoma et telson d’une couleur davantage rougeâtre clair. La morphologie générale rappelle celle

du genre Uroplectes Peters, 1861. Granulation faiblement marquée. Carènes des pédipalpes dis¬

crètes ; corps lisse avec une carène médiane ; anneaux du métasoma avec des carènes bien mar¬

quées; les dorsales avec un granule postérieur spiniforme. Anneaux I et II avec 10 carènes; III

et IV à 8 carènes ; V avec 5 carènes ; les ventrales plus discrètes. Peignes sans fulcres ; absence

Fig. 1-3. — Exemplaires holotypes en vue dorsale. 1, Pseudouroplectes betschi (femelle); 2, Tityobuthus guillaumeti (mâle); 3,

Tityobuthus pococki (mâle).

Dorsal view of holotypes. 1, Pseudouroplectes betschi (female); 2, Tityobuthus guillaumeti (male); 3, Tityobuthus pococki

(male).

— 97 —

d’une dent interne plus développée. Deux éperons basitarsaux à la quatrième patte. Chélicères

avec la dentition caractéristique des Buthidae (Vachon, 1963); les deux dents basales sont très

discrètes, difficiles à individualiser. Tranchant des doigts mobiles des mains des pédipalpes avec

7 séries de granules. Trichobothriotaxie du type A avec la disposition a pour les trichobothries

de la face dorsale du fémur des pédipalpes (Vachon, 1963, 1965).

Le nouveau genre Pseudouroplectes rappelle le genre africain Uroplectes. Il peut cependant

être facilement distingué d ’Uroplectes par l’absence de fulcra aux peignes, l’absence de la dent

interne plus développée et la présence d’un sternum à mi-chemin entre la forme subtriangulaire

et subpentagonale.

Pseudouroplectes betschi n. sp.

(Fig. 1, 4-8)

Matériel-type. — Holotype femelle : plateau Mahafaly, province de Tuléar, Madagascar, 28 juillet 1967

(J.-M. Bestch) (90 m ait., zone à dolines, près de l’Andramanoetse Be; haut fourré arbustif à Euphorbes et

Didiéréacées ; chasse à vue, sous bois mort et pierres). Déposé au Muséum national d’Histoire naturelle, Paris,

MNHN RS-8512.

Étymologie. — Le nom spécifique est attribué en hommage au Pr Jean-Marie Betsch du Laboratoire d’Éco¬

logie du Muséum national d’Histoire naturelle, Paris.

Description

Coloration générale jaunâtre avec présence de taches rougeâtres longitudinales esquissées

sur le mésosoma. Anneau V du métasoma et telson rougeâtre clair. Face ventrale des prosoma

et mésosoma jaunâtre pâle. Le tubercule oculaire et les yeux latéraux sont noirs.

Morphologie

Prosoma : plaque prosomienne avec le bord antérieur concave. Tubercule oculaire antérieur

par rapport au centre de la plaque prosomienne; yeux médians séparés par un diamètre oculaire.

Trois paires d’yeux latéraux. Carènes du prosoma et du mésosoma très discrètes; seule une carène

médiane est bien perceptible sur le mésosoma. Plaque prosomienne et tergites du mésosoma

dépourvus de granulation, pratiquement lisses.

Métasoma ; tous les anneaux avec des carènes bien marquées ; les dorsales avec un granule

postérieur spiniforme; anneaux I et II avec 10 carènes; III et IV à 8 carènes; anneau V arrondi

avec 5 carènes; les ventrales sont plus discrètes que chez les autres anneaux. Vésicule lisse

fortement allongée. Aiguillon dépourvu d’épine ventrale. Sternites à stigmates linéaires aplatis.

Peignes avec 19-18 dents (paratype avec 16-15); absence de fulcres et d’une dent interne plus

développée. Deux éperons basitarsaux à la quatrième patte.

Pédipalpes lisses avec des carènes pratiquement imperceptibles. Tranchant des doigts mobiles

de la main avec 7 séries longitudinales de granules fins, disposés en ligne droite légèrement

oblique. Chélicères avec la dentition caractéristique des Buthidae. Les deux dents basales sont

peu individualisées et semblent être fusionnées en une seule. Trichobothriotaxie du type A-a,

orthobothriotaxique (Vachon, 1973, 1975). Un paratype femelle avec les mêmes données que

pour l’holotype.

— 98 —

Fig. 4-8. — Pseudouroplectes betschi, paratype femelle. 4, peigne, opercule-génital et sternum ; 5, fémur, vue dorsale ; 6, chélicère ;

7, tarse ; 8, anneau V du métasoma et telson.

Pseudouroplectes betschi, female paratype. 4, pectines . genital operculum and sternum; 5, femur, dorsal view; 6, chelicerae;

7, tarsus; 8, segment V of metasoma and telson.

Mensurations (en mm) de l'exemplaire décrit : plaque prosomienne, longueur 2,6; largeur

antérieure 1,5 ; largeur postérieure 2,5. Anneau caudal I, longueur 1,4; largeur 1,3. Anneau caudal

V, longueur 3,3; largeur 1,1; hauteur 1,1. Telson, largeur 0,7; hauteur 0,7. Pédipalpes : fémur,

longueur 2,1; tibia, longueur 2,7; pince, longueur 3,6; doigt mobile, longueur 2,7.

Genre MICROCHARMUS n. g.

(Fig. 9-11)

Diagnose

Scorpion de petite taille : longueur totale des adultes d’environ 20 mm. Coloration générale

jaunâtre; seuls les yeux sont de couleur noire. La morphologie générale rappelle celle du genre

Charmus Karsch, 1879 et, pour certains caractères, tels la forme des peignes et celle du sternum,

le nouveau genre paraît se rapprocher de Microtityus K. W., 1966. Cependant, dans ce deuxième

cas, il ne peut s’agir que d’une convergence. Granulation générale peu marquée, faible ou

moyenne selon les régions du corps. Carènes des pédipalpes et du corps très faiblement marquées ;

celles des anneaux du métasoma bien marquées sur les quatre premiers anneaux ; les dorsales

avec un granule distal spiniforme ; carènes ventrales peu marquées. Peignes très petits, sans ful-

cres ; un éperon basitarsal à la quatrième patte. Chélicères avec la dentition caractéristique des

Buthidae (Vachon, 1963); les deux dents basales sont très petites, peu marquées et difficiles

à distinguer. Tranchant des doigts mobiles des mains des pédipalpes avec 6 (7) séries de gra¬

nulations légèrement obliques. Trichobothriotaxie du type A avec la disposition a pour les tri-

chobothries de la face dorsale du fémur des pédipalpes (VACHON, 1973, 1975).

Le nouveau genre Microcharmus rappelle le genre Charmus, connu uniquement de l’Inde

et du Sri Lanka, par la morphologie de son sternum nettement subpentagonale et par la disposition

des trichobothries de la face dorsale du fémur des pédipalpes. Le nouveau genre peut néanmoins

être distingué du genre Charmus, par l’absence de fulcres et un nombre très réduit de dents aux

peignes (10-10).

Microcharmus cloudsleythompsoni n. sp.

(Fig. 9-11)

Matériel-type. — Holotype femelle : Zangoa, Nord Ouest, Madagascar, 1947 (J. Millot). Déposé au

Muséum national d’Histoire naturelle, Paris, MNHN RS-4811.

Etymologie. — Le nom spécifique est attribué en hommage au Pr John L. Cloudsley-Thompson, University

of London.

Fig. 9-11. — Microcharmus cloudsleythompsoni, holotype mâle. 9, anneau V du métasoma et telson; 10, peignes, opercule-génital

et sternum ; 11, fémur, vue dorsale.

Microcharmus cloudsleythompsoni, male holotype. 9, segment V of métasoma and telson: 10, pectines, genital operculum

and sternum; 11, femur, dorsal view.

— 100 -

Description

Coloration générale uniformément jaunâtre. Face ventrale des prosoma et mésosoma jaunâtre

pâle. Le tubercule oculaire et les yeux latéraux sont noirs.

Morphologie

Prosoma : plaque prosomienne avec le bord antérieur légèrement concave. Tubercule oculaire

antérieur par rapport au centre de la plaque prosomienne ; yeux médians séparés par un diamètre

oculaire environ. Trois paires d’yeux latéraux. Carènes du prosoma et du mésosoma très faible¬

ment marquées. Granulation de la plaque prosomienne et des tergites du mésosoma moyennement

à faiblement marquée.

Métasoma : carènes bien marquées sur les quatre premiers anneaux ; les dorsales avec un

petit granule distal spiniforme; les ventrales plus discrètement marquées; anneau V arrondi avec

des carènes moins marquées que chez les quatre autres anneaux. Anneaux I et II avec 10 carènes;

III et IV à 8 carènes ; anneau V avec 5 carènes. Vésicule lisse très allongée. Aiguillon dépourvu

d’épine ventrale. Stemites à stigmates linéaires aplatis. Peignes très petits avec un nombre de

dents particulièrement faible, 10-10. quasiment atypique chez un Buthidae, mais plutôt caracté¬

ristique des quelques micro-Scorpions; absence de fulcres. Sternum subpentagonal, caractéristique

de certains Buthidae tels Charmus et Microtityus.

Pédipalpes très lisses; carènes très faiblement marquées. Tranchant des doigts mobiles des

mains des pédipalpes avec 7 (6) séries de granulations légèrement obliques. Chélicères avec la

dentition caractéristique des Buthidae ; les deux dents basales sont néanmoins très petites, peu

marquées et difficiles à distinguer et semblent être fusionnées en une seule (Vachon, 1963).

Trichobothriotaxie du type A-a, orthobothriotaxique. Pas de paratypes.

Mensurations (en mm) de l’exemplaire décrit : plaque prosomienne, longueur 1,6; largeur

antérieure 1,1 ; largeur postérieure 1,8. Anneau caudal I. longueur 0,9; largeur 1,0. Anneau caudal

V, longueur 1,9; largeur 0,8; hauteur 0,7. Telson, largeur 0,5; hauteur 0,6. Pédipalpes : fémur,

longueur 1,3; tibia, longueur 1,7; pince, longueur 2,5; doigt mobile, longueur 1,7.

Genre NEOGROSPHUS n. g.

(Fig. 12)

Lors de la description de Grosphus griveaudi Vachon (Vachon, 1969), celui-ci soulève

plusieurs remarques sur le classement de cette espèce dans le genre Grosphus Simon. Ainsi,

Vachon (1969) affirme : «La détermination des spécimens qui ont permis la création de cette

espèce nouvelle nous a posé maints problèmes. Car G. griveaudi possède des caractères qui

l’isolent nettement des six autres espèces connues de Grosphus». Il affirme plus loin : «Il est

donc fort possible que l’espèce griveaudi appartienne à un sous-genre nouveau ou à un genre

nouveau. »

Au cours de la présente étude, après examen de la totalité de la série typique, constituée

de nombreux spécimens, j’en arrive à la conclusion que cette espèce doit être placée dans un

genre différent, pour lequel une nouvelle diagnose est donnée ci-dessous.

— 101 —

Fig. 12. — Vue dorsale d’ensemble de la femelle type de Neogrosphus griveaudi (Vachon), espèce-type du genre (d’après Vachon,

1969).

Dorsal view of the female type of Neogrosphus griveaudi (Vachon), type species of the genus (from Vachon, 1969).

Diagnose

Scorpions de taille petite et moyenne ; les mâles ont environ la moitié de la taille des femelles

(24 mm contre 45 mm pour les femelles). Le nouveau genre se particularise aussi par un nombre

de séries de granulations au tranchant des doigts mobiles des pinces des pédipalpes (8-9) bien

inférieur à ceux retrouvés sur les espèces de Grosphus (de 11 à 14). Puis, ainsi que le souligne

Vachon (1969), une caractéristique encore plus importante est la possession, sous la dent ter¬

minant le doigt mobile, de trois granules externes alors que tous les autres Grosphus en ont

quatre. Par ailleurs, la dent terminale est très développée et pointue chez Neogrosphus. Enfin,

la position de la trichobothrie V 2 de la face ventrale de la main est particulièrement ventrale,

caractéristique déjà observée pour le genre Parabuthus (Vachon, 1969), tandis que la languette

tarsale est très réduite comparée à celle observée chez les espèces de Grosphus.

Le travail de Vachon (1969) illustre déjà fort bien ces différents points. La totalité du

matériel que j’ai pu étudier est répertoriée dans ce même travail.

— 102 —

Tityobuthus guillaumeti n. sp.

(Fig. 2, 13-17)

Matériel-type. — Holotype mâle : Iaraka, baie d’Antongil, Madagascar, 17 novembre 1969 (J.-M. Bestch)

(700 m alt., forêt dense humide de moyenne altitude). Déposé au Muséum national d’Histoire naturelle, Paris,

MNHN RS-7268.

Etymologie. — Le nom spécifique est attribué en hommage au Dr Jean-Louis Guillaumet de l'ORSTOM,

Paris.

Description

Coloration générale uniformément jaunâtre. Face ventrale du prosoma et du mésosoma jau¬

nâtre pâle. Le tubercule oculaire et les yeux latéraux sont noirs.

Morphologie

Prosoma : plaque prosomienne avec le bord antérieur légèrement concave. Tubercule oculaire

antérieur par rapport au centre de la plaque prosomienne ; yeux médians séparés par trois quarts

d’un diamètre oculaire. Trois paires d’yeux latéraux. Carènes du prosoma et du mésosoma pra¬

tiquement imperceptibles. Granulation du corps en général très faible ou inexistante.

Métasoma : tous les anneaux avec des carènes faiblement marquées ou absentes. Carènes

ventrales de l’anneau V dentillées. Vésicule lisse très allongée. Aiguillon avec une épine ventrale

Fig. 13-17. — Tityobuthus guillaumeti, holotype mâle. 13, fémur, vue dorsale; 14, peigne; 15, tarse; 16, chélicère; 17, anneau

V du métasoma et telson.

Tityobuthus guillaumeti, male holotype. 13, femur, dorsal view; 14, pectine; 15, tarsus; 16, chelicerae; 17, segment V of

métasoma and telson.

103 —

bien développée. Sternites à stigmates plutôt linéaires aplatis. Peignes très grands avec 15-14

dents, absence de fulcres. Deux éperons basitarsaux à la quatrième patte.

Pédipalpes très lisses. Seules trois dents sont remarquées sur la face interne du tibia. Tran¬

chant des doigts mobiles de la main avec 7 séries longitudinales de granules très fins disposés

presque en ligne droite, l’une derrière l’autre. Chélicères avec la dentition caractéristique des

Buthidae, cependant les deux dents basales se distinguent très peu l’une de l’autre et semblent

quasiment fusionnées en une seule (Vachon, 1963). Trichobothriotaxie du type A-a, orthobo-

thriotaxique (Vachon, 1973, 1975). Pas de paratypes.

Mensurations (en mm) de l’exemplaire décrit : plaque prosomienne, longueur 2,1 ; largeur

antérieure 1,4; largeur postérieure 2,1. Anneau caudal I, longueur 1.1 ; largeur 1,2. Anneau caudal

V, longueur 2,7; largeur 1,0; hauteur 1,1. Telson, largeur 0,8; hauteur 0,8. Pédipalpe : fémur,

longueur 2,2; tibia, longueur 2,3; pince, longueur 3,5; doigt mobile, longueur 2,1.

Cette nouvelle espèce est voisine de Tityobuthus baroni (Pocock), par sa taille et sa mor¬

phologie générale. Cependant elle peut être distinguée de T. baroni par l’absence de fulcres aux

peignes. L’étude d'une cinquantaine d'exemplaires de T. baroni de la collection du Muséum a

démontré que les fulcres sont toujours présentes, y compris dans tous les exemplaires immatures.

Vachon (in litt.) m’avait suggéré que cette espèce pourrait être placée dans un genre nouveau,

cependant il me semble plus prudent de retarder une telle décision.

Tityobuthus pococki n. sp.

(Fig. 3, 18-22)

Matériel-type. — Holotype mâle : W. Bekopaka Antsingy, Madagascar, juillet 1970 (P. Griveaud). Déposé

au Muséum national d’Histoire naturelle, Paris MNHN RS-5418.

Étymologie. — Le nom spécifique est attribué en hommage à R. I. Pocock, créateur du genre Tityobuthus.

Description

Coloration générale globalement jaunâtre, avec la présence de quelques taches brunâtres

sur la plaque prosomienne. Le tubercule oculaire et les yeux latéraux sont noirs. Deux taches

longitudinales brunâtres sur le mesosoma. Ensemble de la face ventrale et sternites jaunâtres.

Métasoma : trois premiers anneaux jaunâtres; IV et V rougeâtres. Vésicule rougeâtre.

Morphologie

Prosoma : plaque prosomienne avec le bord antérieur légèrement concave. Tubercule oculaire

antérieur par rapport au centre de la plaque prosomienne ; yeux médians séparés par moins d’un

diamètre oculaire. Trois paires d’yeux latéraux. Carènes du prosoma et du mesosoma discrètes;

seule une carène est bien perceptible sur les tergites I à VI du mesosoma. VII avec 5 carènes.

Plaque prosomienne et tergites du mesosoma avec une granulation moyenne.

Métasoma : tous les anneaux avec des carènes bien marquées; les dorsales avec un granule

postérieur spiniforme; anneaux I et II avec 10 carènes; III avec 8 carènes; anneaux IV et V

avec des carènes très discrètes; les carènes ventrales sont absentes sur tous les anneaux; anneau

V très arrondi. Vésicule lisse, allongée. Aiguillon très long avec une épine ventrale aiguë. Ster-

— 104 —

FIG. 18-22. — Tityobuthus pococki, holotype mâle. 18, fémur, vue dorsale; 19, tibia, vue dorsale; 20, pince, vue latérale; 21,

section de la quatrième patte avec éperon tibial ; 22, plaque prosomienne.

Tityobuthus pococki, male holotype. 18, femur, dorsal view; 19, tibia, dorsal view; 20, chelae, lateral view; 21, section of

fourth leg with tibial spur; 22, carapace.

nites à stigmates linéaires aplatis. Peignes avec 22-23 dents. Epéron tibial très développé, ce

qui différencie cette espèce des autres du genre (Vachon, 1979).

Pédipalpes avec des carènes moyennement marquées. Fémur avec 5 carènes ; tibia à carènes

peu définies et à face interne avec des granules spiniformes; pince plutôt lisse. Tranchant des

doigts mobiles de la main avec 8 séries longitudinales de granules fins, disposés en ligne droite

légèrement oblique ; présence de granules accessoires plus gros. Chélicères avec la dentition

caractéristique des Buthidae. Les deux dents basales sont peu individualisées. Trichobothriotaxie

du type A-oc, orthobothriotaxique (Vachon, 1973, 1975). Pas de paratypes.

Mensurations (en mm) de l’exemplaire décrit : plaque prosomienne, longueur 2,9; largeur

antérieure 2,1 ; largeur postérieure 3,0. Anneau caudal I, longueur 2,0; largeur 1,7. Anneau caudal

V, longueur 3,5; largeur 1,5; hauteur 1,5. Telson, largeur 0,9; hauteur 1,0. Pédipalpes ; fémur,

longueur 2,8; tibia, longueur, 3,7; pince, longueur 6,0; doigt mobile, longueur 4,1.

Cette nouvelle espèce se situe, par sa taille, dans une position intermédiaire entre les trois

espèces connues du genre Tityobuthus ; elle est plus grande que Tityobuthus baroni (Pocock) et

plus petite que Tityobuthus gracilis (Fage). Elle peut être facilement distinguée des espèces citées

par la présence d’éperons tibiaux bien développés, très discrets ou absents chez les autres espèces

(VACHON, 1979). Enfin, elle se distingue de Tityobuthus guillaumeti par la présence de fulcres

aux peignes.

— 105 —

CONSIDÉRATIONS PRÉLIMINAIRES SUR LA PHYLOGÉNIE

DES NOUVEAUX TAXA DÉCRITS

Dans une optique tout à fait préliminaire, il me semble nécessaire de souligner l’importance

de la découverte de trois nouveaux genres à Madagascar pour la clarification des origines et

affinités de la faune scorpionique actuelle.

# Microcharmus cloudsleythompsoni

® Pseudouroplectes bctschi

O Tityobuthus guillaumeti

♦ Tityobuthus pococki

Y Neogrosphus griveaudi

Pieds

Mètres

13.123

4.000

6.562

2.000

3.281

—

1.000

1.640

—

500

656

200

—

Lactmp.

250 km

150 mi

Fig. 23. — Carte avec l’indication des stations typiques des genres et des espèces nouvelles décrites dans le présent travail.

Map with indication of the typical localities of the new genera and new species.

— 106 —

La position de Pseudouroplectes semble être relativement simple à préciser car ses affinités

avec le genre africain Uroplectes peuvent être facilement admises. Le genre Uroplectes est lar¬

gement réparti dans la région afrotropicale, à l’exception du Zaïre et d’une partie de l’Afrique

occidentale recouvrant le Cameroun et le Gabon (Lamoral, 1979).

La découverte du nouveau genre Microcharmus est particulièrement intéressante, car ses

affinités avec le genre Charmus démontrent la possible existence d'un lien entre la faune scor-

pionique malgache et celle de la région indo-malaise; le genre Charmus n’était jusqu’à présent

connu que de l’Inde et du Sri Lanka.

Le genre Neogrosphus est sans aucun doute associé à Grosphus, et sa différenciation a

certainement eu lieu plus récemment que celle d’autres genres malgaches. La création des trois

genres nouveaux, associée à la description de quatre espèces nouvelles augmente d’une manière

significative la diversité de la faune scorpionique de la grande île et son taux d’endémicité.

Ces quelques conclusions préliminaires seront discutées dans un article actuellement en cours

de rédaction sur la biogéographie de Madagascar. Dans cet article global, seront clarifiées les

questions des origines, affinités et différenciation de la faune malgache actuelle.

Remerciements

Je suis très reconnaissant à la direction du Laboratoire de Zoologie, Arthropodes, du Muséum national

d’Histoire naturelle, Paris, de m’avoir facilité l’étude du matériel utilisé dans le présent travail. Mes re¬

merciements vont aussi tout particulièrement à M. J. Rebière pour sa contribution à la réalisation de

plusieurs dessins illustrant le présent travail.

RÉFÉRENCES

Fage, L., 1929. — Les Scorpions de Madagascar. Faune des Colonies françaises 3. Soc. Edit. Géogr. Mûrit.

Colon., Paris : 637-694.

Gervais, R, 1844. — Remarques sur la famille des Scorpions. Archs. Mus. Hist, nat., Paris, 4 : 201-240.

KRAEPELIN, K., 1901. — Catalogue des Scorpions des collections du Muséum d’Histoire naturelle de Paris. Bull.

Mus. Hist. nat. Paris, 7 : 265-274.

LAMORAL, b. H., 1979. — The scorpions of Namibia (Arachnida: Scorpionida). Ann. Natal Mus., 23 (3): 497-784.

MlLLOT, J., 1948. — Revue générale des Arachnides de Madagascar. Mém. Inst. Sci. Madagascar., sér. A, 1 (2).

[Mém. Soc. Biogéogr. (1953) : 127-145].

SIMON, E., 1880. — Descriptions de genres et espèces de l’ordre des Scorpions. Ann. Soc. Entom. Fr., 10 (5) :

377-398.

VACHON, M., 1963. — De l’utilité, en systématique, d’une nomenclature des dents des chélicères chez les Scor¬

pions. Bull. Mus. natn. Hist, nat., Paris 2 e sér., 35 (2) : 161-166.

— 1969. — Grosphus griveaudi, nouvelle espèce de Scorpion Buthidae Malgache. Bull. Mus. natn. Hist, nat.,

Paris 2 e sér., 4 (2) : 476-483.

— 1973. — Etude des caractères utilisés pour classer les familles et les genres de Scorpions (Arachnides). 1.

La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les Scor¬

pions. Bull. Mus. natn. Hist, nat., Paris, 3 e sér., n° 140, Zool., 104 : 857-958.

— 1975. — Sur l’utilisation de la trichobothriotaxie du bras des pédipalpes des Scorpions (Arachnides) dans

le classement des genres de la famille des Buthidae Simon. C. r. Acad. Sci., Paris, sér. D, 281 : 1597-1599.

— 1979. — Remarques biogéographiques sur la faune des Scorpions de Madagascar à propos de l’utilisation

de caractères trichobothriotaxiques permettant la distinction des genres Odonturus Karsch, 1879 et Tityobu-

thus Pocock, 1893. C. r V e Coll. Arach., IX : 217-224.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2: 107-140.

Cyst morphology of European branchiopods

(Crustacea: Anostraca, Notostraca, Spinicaudata, Laevicaudata)

by Alain THIÉRY, Jàn BRTEK and Charles GASC

Abstract. — A comparative analysis of cyst (= resting egg) size and shell architecture by scanning electron

microscopy is reported in 40 species of “large branchiopods”, belonging to 21 genera from all the families

inhabiting inland waters of Europe. This paper follows one on the geographic distribution of the European

branchiopods where 72 species and subspecies belonging to 22 genera and 13 families were listed (Brtek &

Thiéry, 1995). Particular attention has been paid to cyst shape and morphology of the external shell surface.

The results are discussed in the light of existing literature. When the taxonomic value of the external shell

surface pattern shows its limits, the combination size/surface pattern seems more useful, being sometimes spe¬

cies-specific, while in other cases (such as in Branchipus, Tanymastix , Cyzicus, Imnadia) it appears to reflect

taxonomic relationships of higher rank. In other cases, a third character, the structure of the cyst envelope,

observable in fractured cysts, must be taken into account to separate cysts (as in Artemia sp. and Branchinectella).

“Large branchiopods” eggs can provide new taxonomic information that is useful in the definition of natural

groups.

Keywords. — Cyst morphology, resting egg, Europe, Crustacea, Anostraca, Notostraca, Spinicaudata, Lae¬

vicaudata, scanning electron microscopy, taxonomy.

Morphologie des œufs de branchiopodes européens

(Crustacea: Anostraca, Notostraca, Spinicaudata, Laevicaudata)

Résumé. — La morphologie des œufs (= œufs de résistance) de 40 espèces de branchiopodes (Anostraca,

Notostraca, Spinicaudata, Laevicaudata) appartenant à 21 genres représentant toutes les familles peuplant les eaux

stagnantes continentales d’Europe, est décrite au moyen du microscope électronique à balayage. Cette étude fait

suite à l’article de Brtek & Thiéry (1995) qui dresse l’inventaire actuel de ces crustacés branchiopodes en Europe

et illustre la distribution géographique des 72 espèces et sous-espèces répertoriées, représentantes de 22 genres

et 13 familles. La forme des œufs et la morphologie externe de leur enveloppe sont décrites, les résultats étant

confrontés aux données bibliographiques. Dans de nombreux cas, les ornementations de surface sont des critères

fiables permettant une identification spécifique ou générique. Lorsque ces ornementations ne présentent pas de

caractéristiques suffisantes pour établir une discrimination des œufs, il est alors nécessaire de considérer la combi¬

naison diamètre/ornementations de surface qui peut être spécifique ou parfois générique comme c’est le cas pour

les genres Branchipus , Tanymastix, Cyzicus et Imnadia. Dans quelques cas, un troisième caractère, la structure

interne de l’enveloppe de l’œuf, observable en coupe, doit être considéré pour séparer des œufs d’aspect identique

mais d’espèces différentes (cas pour Artemia sp. et Branchinectella media). La morphologie des œufs de bran¬

chiopodes, ayant une incontestable valeur taxonomique, devrait être prise en compte pour compléter les études

sur la phylogénie de ces crustacés.

Mots-clés. — Morphologie, œufs de résistance, Europe, Crustacea, Anostraca, Notostraca, Spinicaudata,

Laevicaudata, microscope électronique à balayage, taxonomie.

A. Thiéry & Ch. Gasc, Laboratoire de Biologie animale-Hydrobiologie, Faculté des Sciences, Université d'Avignon, 33 , rue

Louis-Pasteur, F-84000 Avignon.

J. Brtek, Hornonitrianske Muzeum, Hlinkova 44, 971 01 Prievidza, Slovakia.

— 108 —

INTRODUCTION

“Large branchiopods” - fairy shrimps (Anostraca), tadpole shrimps (Notostraca) and clam

shrimps (Spinicaudata + Laevicaudata) - are crustaceans living in inland temporary freshwaters,

or sometimes saltwaters. As they live in physically and chemically fluctuating astatic habitats,

with recurrent phases of complete drying (temporary ponds, ditches, salterns...), these crustaceans

present adaptations to desiccation, the main one being the production of only one sort of egg

- thick-walled resting eggs (= cysts). The morphology of cysts has been described since the

work of Daday (1910 to 1927) and in several cases specific distinctness has been hypothetized.

During the mid- and late 1980s, the micromorphology of cysts has been a central topic in “large

branchiopods” biology. Scanning electron microscopy has been used to obtain accurate images

of fine surface structures and increasing knowledge has led a number of authors to consider and

discuss the taxonomic importance of the micromorphology and size of the cysts (review in Thiéry

& Gasc, 1991; De Walsche et al . , 1991; Mura, 1992a, b). While for several countries cyst

morphology has been studied, (Spain: Alonso & Alcaraz, 1984. all the orders; Italy: Mura,

1986. 1992a, both only anostracans; France: Thiéry & Gasc, 1991, all the orders), no global

view on the European scale is available. The present study is the first one to attempt a synthesis.

MATERIAL AND METHODS

Identification

Species identifications were made using adult morphology (second antenna and penes of

the male; female ovisac), using Daday (1910 to 1927), Linder (1941), Brtek (1959, 1964,

1966), Longhurst (1955), Straskraba (1965a, b; 1966). Branchiopod taxonomy follows Fryer

(1987) and Brtek & Thiéry (1995).

The surveyed area is shown in Fig. 1.

SEM STUDIES

Cysts were removed from female ovisacs, kept in 10% formaldehyde, then fixed in a bath

of KAAD (30% kerosene, 60% absolute ethanol, 5% acetic acid, 5% dioxane) for 24 h (see

THIÉRY & Gasc, 1991). From preserved females, we chose only individuals with well formed

complete egg-shells. The egg-shell pattern changes from fertilization to time of deposition (see

THIÉRY, 1985 for Triops\ Mura, 1992b for Chirocephalus diaphanus). After COt critical point

drying, cysts were coated with gold and then observed using a Cambridge Stereoscan 360 at an

acceleration voltage of 20 KV and a working distance varying from 9-10 mm for the smaller

cysts (clam shrimps) to 25 mm for the bigger ones (Branchinecta and tadpole shrimps). Some

technical data are given on Fig. 4c.

Measurements

Cyst diameter and surface ornamentation (spines, ridges...) were measured with a SEM

numerical point-point system (see example in Figs 4b - crosses, and 4d - vertical lines) and

— 109 —

FIG. 1. — Area surveyed in the study of cyst morphology of “large branchiopods”.

Aire géographique prise en compte pour l’étude de la morphologie des œufs des « grands Branchiopodes ».

sometimes on calibrated SEM photonegatives. Diameters cited from the literature were measured

from published calibrated photographs. To study the variability of cyst diameter we use a Variation

Index (V. I.) defined as: maximum diameter-minimum diameter/mean diameter, in per cent.

Material examined

ANOSTRACA

Anemia parthenogenetica Bowen & Sterling, 1978: saltworks of Salin-de-Giraud, Camargue, S. France, 6,

5. 1989, coll. A. Thiéry (Figs 7h, 8a).

Anemia tunisiana Bowen & Sterling, 1978 (= bisexual strain): coastal saltworks, Sète-Villeroy, S. France,

5. 5. 1989, coll. A. Thiéry (Fig. 8b).

Branchinecta ferox (Milne-Edwards, 1840): Hurbanovo, 17. 4. 1956, and Palarikovo, 22. 4. 1969, SW

Slovakia, both in coll. J. Brtek (136) (Figs 4b, c).

Branchinecta minuta Smirnov, 1948: Dnepropetrovsk-district, Ukraine, 6. 4. 1937, coll. S. Smirnov (Figs 4f,

g; 5b-d).

Branchinecta orientalis Sars, 1901: Illmitz, Lake Kirchsee, Austria, 26. 5. 1963, coll. J. Vornatscher

(Fig. 4e).

Branchinecta paludosa (Müller, 1788): High Tatra mountains, Horne Furkotske Lake, N. Slovakia. (Figs 4a;

5a).

Branchinecta tolli (Sars, 1897): Laptevis Sea, Stolbovoj Island, Russia, 29. 2. 1985, and Tiksi, Yakutia,

Russia, 1. 8. 1985, coll. N. Vekhoff (Figs 4d; 5e-g).

Branchinectella media (Schmankevitch, 1873): Troick District, Kudaj-Sugur lake, Kazachstan, 11. 4. 1932,

coll. S. Smirnov (Figs 7e; 8e).

Branchinella spinosa (Milne-Edwards, 1840): salterns of Salin-de-Giraud, Camargue, Southern France, 6.

5. 1989 (Fig. 8c).

Branchipus blanchardi Daday, 1908: High Alps, type locality on the Cristol Plateau, France, 24. 8. 1988,

coll. A. Thiéry (Fig. 6j, k).

Branchipus schaefferi Fischer, 1834: Tekovské Luzany (Sec. Kertész 1956 as “Nagysallo”, SW Slovakia,

27. 5. 1972, coll. J. Brtek (1513) syn. visnyai (Fig. 6h), and Rivesaltes, 19. 11. 1987, and Plain of Crau, 26. 3.

1994, S. France, coll. A. Thiéry (Fig. 6i).

Branchipus laevicornis* Daday, 1910: Eregli, Asia Minor, coll. E. Daday, 1912 MNHN Bp273 (Fig. 7d).

Chirocephalus bairdi* (Brauer, 1877): Berekhat Hitin, Israel, 24. 3. 1983, coll. R. Ortal (Fig. lOa-c).

Chirocephalus chyzeri (Daday, 1890): Maly Hores, 24. 4. 1970, and Vel’ky Kamenec, SE Slovakia, 24. 4.

1970, coll. J. Brtek (745, 752) (Fig. 9c, d).

Chirocephalus carnuntanus (Brauer, 1877): Gbelce, 28. 4. 1966, and Hurbanovo, 14. 4. 1967, SW Slovakia,

coll. J. Brtek (228) (Figs 9i; 1 Oi l).

Chirocephalus diaphanus Prévost, 1803: Saint-Maximin, S. France (Provence), 7. 2. 1988, coll. A. Thiéry

(Fig. 8j).

Chirocephalus diaphanus carinatus (Daday, 1910): Musala, Bulgaria, alt. 2300 m, 8. 8. 1962, coll. J. Gulicka

(Fig. 9b).

Chirocephalus josephinae (Grube, 1853): Ust’-Kut, Asian Russia, 4. 6. 1925, coll. S. Smirnov (Figs 9h;

lOh).

Chirocephalus orghidani Brtek, 1966: Planitza, Romania, 18. 4. 1960, coll. P. Banarescu (Figs 8i; 10e).

Chirocephalus pelagonicus Petkovski, 1986: Golemo Konjari, Macedonia, 4. 5. 1985, coll. S. Petkovski

(Fig. 9a).

Chirocephalus salinus (Daday, 1910): “Trepadoules” Porto Vecchio, SE Corsica, 28. 1. 1988, coll. A. Thiéry

(Figs 8k; lOd).

Chirocephalus shadini (Smirnov, 1928): Bol'-Zatin, 25. 4. 1970, and Zatin, 15. 4. 1969, SE Slovakia, coll.

J. Brtek (755, 399) (Figs 9g; lOf).

Chirocephalus slovacicus Brtek, 1971: Jesenské, 12. 4. 1968, Janice, 22. 4. 1970, and Lenartovce-Vlkyna,

21. 4. 1981, S. Slovakia, coll. J. Brtek (365, 718, 1873) (Figs 9e; 10g).

Chirocephalus spinicaudatus Simon, 1886: France, MNHN Bpl57, coll. E. Simon (25. 96), E. Daday det.

1909 (Fig. 9f).

Drepanosurus biro stratus (Fischer, 1851): Tomsk, Russia, 25. 4. 1919 (Fig. 8h: immature cyst).

Drepanosurus hankoi (Dudich, 1927): Kralovsky Chlmec, SE Slovakia, 15. 4. 1969, coll. J. Brtek (398)

(Figs 6a, b; 7a).

Linderiella massaliensis Thiéry & Champeau, 1988: Saint-Maximin, S. France, 7. 2. 1988, coll. A. Thiéry

(F igs 7f; 8d).

Species living outside the study area, but described for comparison.

— Ill —

Polyartemia forcipata Fischer, 1851: Murmansk, Russia, 1898 (No 228) (Fig. 7b).

Siphonophanes grubii (Dybowski, 1860): Moravia, coll. J. FIhrabé 27. 4. 1958, MNF1N Bp 161, and Slovakia

mer-occid. Sv. Jur. 25. 4. 1958 J. Brtek leg., MNHN Bp 260 (Figs 6f, g; 7c).

Streptocephalus torvicornis (Waga, 1842): Malé Trakany, 19. 7. 1969, and Dobra, 19. 7. 1969, SE Slovakia,

coll. J. Brtek (545, 552) (Fig. 6c-e).

Tanymastix stagnalis (Linnaeus, 1758): Fontainebleau forest, France, 9. 8. 1988, coll. A. Thiéry (Fig. 8f).

Tanymastix stellae Cottarelli, 1968: “Trepadoules” South of Porto Vecchio, Corsica, 28. 1. 1988, coll. A.

Thiéry (Fig. 8g).

NOTOSTRACA

Lepidurus apus (Linnaeus, 1758): Kamenicnà, SW Slovakia, 14. 5. 1970, coll. J. Brtek (840); Gavoti, 21.

2. 1988, and Rochefort du Gard, S. France, both in coll. A. Thiéry (Figs lie; 12a, d, f-h).

'Lepidurus arcticus (Pallas, 1793); Sachanika west Coast of Novaya Zemlya, Russia, 12. 9. 1925, coll. Zool.

Inst. AN USSR (Fig. 12b, e).

Lepidurus couesii Packard, 1875: Ulan Bator, Mongolia, 7. 7. 1970, coll. Z. Peregi (Fig. 12c).

Triops cancriformis (Bose, 1801): Komàrovce, SE Slovakia, 17. 7. 1969, coll. J. Brtek; Opoul, 26. 3.

1988, and Plain of Crau, 26. 3. 1994, S. France, coll. A. Thiéry (Fig. lla-d).

SPIN1CAUDATA

Limnadia lenticularis (Linnaeus, 1761): Komarno, 26. 7. 1969, and Kava, 30. 5. 1973, SW Slovakia, coll.

J. Brtek (602) (Fig. 13a-c).

Imnadia yeyetta Hertzog, 1935: Sulany, 15. 5. 1970, and Medvedov, 16. 5. 1970, SW Slovakia, coll. J. Brtek

(893) (Fig. 13d, e), and La Galère, Plain of Crau, S. France, 26. 3. 1994, coll. A. Thiéry.

Cyzicus tetracerus (Krynicki, 1830): Dobra, 3. 6. 1970, and Jenkovce, 31. 5. 1970, SE Slovakia, coll.

J. Brtek (984, 1065); Gavoti. Provence, S. France, 21. 2. 1988, coll. A. Thiéry (Fig. 13f, g, i).

Eocyzicus orientalis Daday, 1913: Astrahan, Russia, 3. 6. 1968. (Fig. 13h, j).

Leptestheria dahalacencis (Riippel, 1837): Sul’any, 15. 5. 1970, and Nové Zàmky, 29. 5. 1973, SW Slovakia,

coll. J. Brtek (866, 1779) (Fig. 14a-e).

Eoleptestheria ticinensis (Balsamo-Crivelli, 1859): Novosad-Jastrabie, 1. 6. 1970, and Zempllnske Jastrabie-

Oborin, 1. 6. 1970, SE Slovakia, coll. J. Brtek (1005, 1009) (Fig. 14f).

LAEVICAUDATA

Lynceus brachyurus Müller, 1776: Kralovsky Chlmec, 3. 6. 1970, SE Slovakia, coll. J. Brtek (1043)

(Fig. 14g, h).

RESULTS

Size-Diameter

Results are presented in Table I and Fig. 2. In Table I, the species are listed in taxonomic

order; in Fig. 2 by increasing size. For each species, our original data (labelled “this study”)

have been supplemented by data from the literature. In all cases, except where diameters are

related to erroneous scale bars, as in Alonso & Alcaraz (1984) for Branchinecta ferox and B.

cervantesi or in Mura (1992, PI. 2. p. 237) for six species of Chirocephalus, the cysts are of

more or less consistent diameter. In most cases, our results agree with data in the literature.

In most cases, the Variation Index (VI) ranges from 3 to 30%. However, in several species,

such as in Branchipus schaefferi, Tanymastix stagnalis, Streptocephalus torvicornis, Chiro¬

cephalus diaphanus, and Artemia, the index is higher, from 30 to 55%. The lesser values of the

Variation Index, indicating a relative constancy in cyst diameter whatever the geographical origin

of the population, are found in species with restricted areas of distribution, while values of V. I.

— 112 —

Leptestheria dahalacensis

Lynceus brachyurus

Eocyzicus orienlalis

Eoleptestheria llclnencis

Cyzicus grubei

Cyzicus tetracerus

Cyzicus buchetl

Llmnadia lenticularis

Imnadia yeyetta

Leptestheria mayeti

Branchipus schaefferi

Branchinectella media

Rrtemia parthenogenetica

Branchipus intermedius

Rrtemia tunisiana

Branchinecta orientalis

Chirocephalus breulpalpis

Streptocephaius toruicornis toruicor

Branchinella spinosa

Linderietla sp. (Southern Spain)

Tanymastix stagnalls

Chirocephalus carnuntanus

Linderiella massaliensis

Streptocephaius toruicornis buchetl

Branchipus cortesi

Chirocephalus diaphanus

Chirocephalus josephinae

Chirocephalus ruffol

Chirocephalus chyzeri

Chirocephalus spinlcaudatus

Chirocephalus kerkyrensis

Branchinecta paludosa

Chirocephalus diaphanus carinatus

Chirocephalus orghidani

Chirocephalus shadinl

Branchipus blanchardi

Branchinecta minuta

Chirocephalus sibyllae

Chirocephalus pelagonicus

Orepanosurus hankoi

Chirocephalus slouacicus

Branchinecta tolli

Siphonophanes grubii

Triops cancriformis cancriformls

Chirocephalus salinus

Branchinecta ferox

Tanymastix motasi

Chirocephalus marchesoni

Tanymastix stellae

Triops cancriformis simplex

Lepidurus apus

Lepidurus couesii

Triops cancriformis mauritanicus

Polyartemia forcipata

Lepidurus arcticus

Triops numidicus

Fig. 2. — Diameters in pm (low and high values from this paper and from available literature) of the cysts of European branchiopods

ranged from the smaller to the larger. Open circles indicate single measurements.

Diamètres en \im (valeurs inférieures et supérieures d’après les résultats de cette étude et les données bibliographiques) des

œufs des Branchiopodes d’Europe classés par ordre croissant de taille. Les cercles indiquent des mesures isolées.

are greater in common and widely distributed species (for geographical distribution in Europe,

see Brtek & ThiéRY, 1995).

The V. I. might be also linked with the diversity of measurements (number of authors or

precision of the measuremts, as shown in Fig. 3 which illustrates the correlation between the

Variation Index and the number of citations in the literature: V. 1. (%) = 5.558 number of citations

+ 0.39 (n = 31, r 2 = 0.633, P < 0.05).

The smallest cysts are those of spinicaudatans and laevicaudatans, ranging from 95 to

209 pm. Those of anostracans range from 198 to 527 pm and of notostracans from 350 to 780 pm.

The Arctic species Polyartemia forcipata and Lepidurus arcticus have the largest cysts of the

European “large branchiopods”.

— 113 —

Fig. 3. — Relation between variation index (%) and number of citations in the literature. V. I. % = 5.558 citations + 0.39 (n =

31, r 2 = 0.633, P < 0.05).

Relation entre l'indice de variation (en %) et le nombre de références dans la littérature. V I. (%) = 5,558 citations + 0,39

(n = 31, r 2 = 0,633, P < 0,05).

Cyst morphology

Anostraca (Table I and Figs 4 to 14): except for the cysts of the genus Tanymastix which

are lenticular, all those of anostracans are more or less spherical. Their surface may be smooth,

as in Artemia and Branchinectella media , or, more usually ornamented by ridges, spines, de¬

pressions or crests.

Polyartemiidae (Fig. 7b): Polyartemia forcipata has a large spherical cyst, with a rugose

surface, with rounded spines shorter than 10 pm in length, as described by Mura (1992b. Plate

5.3).

Branchinectidae (Figs 4, 5): except for those of Branchinecta tolli which are slightly rugose

(Figs 4d; 5e, f, g), the cysts of the European species of Branchinecta are characterized by low

ridges which delineate large polygons (Figs 4a-c, e-g; 5a, b). (Gilchrist, 1978; Mura & Thiéry,

1986; Mura, 1991a, and Maeda-Martinez et al, 1992, 1993). The cysts of Branchinecta

paludosa and B. ferox are similar in appearance and of the same size, but the polygonal areas

tend to be larger in the former. Likewise the cysts of Branchinecta orientalis and B. minuta are

very similar (Fig. 4e, f, g). They are smaller than those of B. paludosa and B. ferox, and their

ridges are more rounded and delineate smaller polygonal areas. No objective character distin¬

guishes them. Concerning the internal structure of the egg-shell, its spongy aspect in Branchinecta

— 114

Fig. 4. — Cysts of anostraca: a) Branchinecta paludosa; b), c) B. ferox; d) B. tolli; e) B. orientalis ; f)> g) B. minuta, (common

scale bar in Jim).

Œufs d'Anostraca : a) Branchinecta paludosa; b), c) B. ferox; d) B. tolli ; e) B. orientalis ; f), g) B. minuta ( échelle commune

en \im).

minuta (Fig. 5c, d), agrees with the findings of GILCHRIST (1978) for B. ferox. In B. tolli, the

cyst wall is thin with thick low roots (Fig. 5f). No data are available for B. orientalis.

Artemiidae (Figs 7, 8): cysts of Artemia are spherical and smooth (Figs 7h; 8a, b), as de¬

scribed in many studies (Mazzini, 1978; Gilchrist, 1978; Mura & Thiéry, 1986; Spotte &

— 115 —

Fig. 5. — Cysts of Anostraca: a) details of Branchinecta paludosa, b) idem, B. minuta, c) idem, outer membrane cracked showing

spongy structure; d) detail of spongy structure; e) Branchinecta tolli', f) Branchinecta tolli, detail of a cracked part of egg

shell; g) B. tolli, detail of surface showing little rounded pits, (scale bars in pm).

Œufs d’ Anostraca : a) détail, Branchinecta paludosa; b) idem, B. minuta; c) B. minuta, la membrane externe déchirée permet

de voir la structure spongieuse ; d) B. minuta, détail de la structure spongieuse ; e) Branchinecta tolli; f) B. tolli, détail de

la partie déchirée de l'enveloppe ; g) B. tolli, détail de la surface montrant les petits tubercules (échelles en \lm).

— 116 —

Fig. 6. — Cysts of Anostraca: a), b) Drepanosurus hankoi ; c), d), e) Streptocephcilus torvicornis\ f), g) Siphonophanes grubii\

h) Branchipus schaefferi syn. visnyai ; i) Branchipus schaefferi; j), k) Branchipus blanchardi (common scale bar in pm).

Œufs d’Anostraca : a) et b) Drepanosurus hankoi ; c), d), e) Streptocephalus torvicomis ; f), g) Siphonophanes grubii ; h)

Branchipus schaefferi syn. visnyai ; i) Branchipus schaefferi ; j), k) Branchipus blanchardi (échelle commune en \im).

— 117 —

Anderson, 1988; Thiéry & Gasc, 1991). Except for a small difference in size, no objective

distinction exists between the cysts of the parthenogenetic and bisexual strains. The vitelline

envelope is thin and spongy (Fig. 7h).

Branchipodidae (Figs 6, 7, 8): this family presents two very different patterns: lenticular

cysts in Tanymastix (Figs 7g; 8f, g) (only small and often unreliable differences between the

species) and wrinkled and angulated in Branchipus (Fig. 6h-k). The cysts of Branchipus schaefferi

and B. blanchardi are shown in this paper while that of B. cortesi is illustrated in ALONSO &

Jaume (1991, Fig. 4b, c, p. 227). As in Tanymastix, whose cyst pattern can be considered as

generic, the angulated shape of Branchipus cysts is so similar in the four European species that

we suggest that it could be generic also. This is confirmed by the cyst of Branchipus laevicornis,

the fifth species of the genus, living in Asia minor, which is very similar in shape to that of

the three species studied (Fig. 7d) and that studied by Alonso & Jaume (1991). The differences

between these species are hardly distinguishable without a large series permitting study of the

morphology and size variability. This pattern is found also in a thamnocephalid, Dendrocephalus

spartaenovae (Mura, 1992b, Plate 6.3), but this is a South American species, so no confusion

is possible in field studies. In section the alveolar layer of the cyst wall of Branchipus blanchardi

and B. laevicornis (both unpublished), are spongy as in B. schaefferi (Kupka, 1940, Figs 6-11;

Gilchrist, 1978, Figs lc, d, 2a-d). In addition, the outer surface has numerous pores in B.

blanchardi (THIÉRY & GASC, 1991, Fig. 32), B. laevicornis (unpublished) and B. schaefferi

(Kupka, 1940, Fig. 5; Gilchrist, 1978, Fig. lb; Mura & Thiéry, 1986, PI. IV.B).

Thamnocephalidae (Fig. 8): the cyst of Branchinella spinosa is spherical with a surface

showing an irregular polygonal pattern formed by low mound-like ridges. These ridges are more

or less pronounced (in relation to the maturation of the egg shell). Cyst size is close to that of

Artemia (which always has smooth cysts), with which this species could coexist in coastal saline

waters as in the Camargue and Sardinia. The B. spinosa pattern is constant for all populations

observed (Alonso & Alcaraz, 1984, Fig. 2.d; Mura, 1986, Plate 5.a; 1992b, Plate 1.5; Mura

& Thiéry, 1986, Plate II.a).

Streptocephalidae (Fig. 6): cysts of Streptocephalus torvicornis are well known (Valousek,

1952; Alonso & Alcaraz, 1984; Mura & Thiéry, 1986; Thiéry, 1987; Thiéry & Gasc,

1991; De Walsche et al., 1991; Mura, 1992b). They have a “folded look” with polygonal

cells irregularly shaped by raised ridges (Fig. 6c-e). Cysts of Streptocephalus torvicornis bucheti

are similar to those of S. t. torvicornis (Alonso & Alcaraz, 1984, and Mura & Thiéry,

1986).

Linderiellidae (Figs 7, 8): the cysts of the genus Linderiella are spiny (Fig. 8d), even with

acute spines ( 15-20 pm long) in the French Linderiella massaliensis (Fig. 7f), or with flat trumpet

tips in the Spanish form (Alonso & Alcaraz, 1984; Thiéry & Champeau, 1988). In all cases

the spines are different from those of the spiny cysts of some species of Chirocephalus, where

they arise from the crests of honeycomb-like surface of the shell. In Linderiella they arise directly

from the shell.

Chirocephalidae Chirocephalinae (Figs 8, 9, 10): at first glance cysts of Chirocephalus spe¬

cies present great heterogeneity. Some are smooth, e.g. Chirocephalus marchesonii (MURA et

ai, 1978, PI. 1; MURA, 1986, PI. 4), some spiny, e.g. C. ruffoi (MURA, 1986, PI. 12), C. diaphanus

carinatus (Fig. 9b) and C. carnuntatus (Fig. 9i), some more or less bumpy as C. chyzeri (Fig. 9c, d),

— 118 —

Fig. 7. — Details of the cysts of Anostraca: a) Drepanosurus hankoi — see low ridges (arrows); b) Polyartemia forcipata —

see small spines (arrows); c) Siphonophanes grubii ; d) whole cyst, Branchipus laevicomis\ e) internal view of envelope,

Branchinectella media (see the pillars — arrow); f) spines of Linderiella massaliensis (those marked with an arrow are

broken); g) section, Tanymastix stagnalis, showing spongy structure; h) detail of the envelope, Artemia (a.l. — alveolar

layer). (Scale bars in pm.)

Détails des œufs d’Anostraca : a) Drepanosurus hankoi — voir les rides (flèches) ; b) Polyartemia forcipata — noter les

petites épines (flèches); c) Siphonophanes grubii; d) œuf de Branchipus laevicornis ; e) vue interne de l’enveloppe chez

Branchinectella media (noter les fdaments — flèche); f) épines de Linderiella massaliensis (celles marquées d’une flèche

sont cassées ; g) coupe transversale de l’œuf de Tanymastix stagnalis montrant la structure spongieuse ; h) détail de l’enveloppe

d’ Artemia (a. I. — couche alvéolaire). (Échelles en \im.)

— 119 —

Fig. 8. — Cysts of Anostraca: a) Artemia (bisexual strain); b) idem, cracked cyst; c) Branchinella spinosa; d) Linderiella mass-

aliensis’, e) Branchinectella media’, f) Tanymastix stellae; g) Tanymastix stagnalis’, h) immature cyst, Drepanosurus birostratus’,

i) Chirocephalus orghidanr, j) Chirocephalus diaphanus; k) Chirocephalus salinus (common scale bar in fim).

Œufs d’ Anostraca : Artemia (souche bisexuée); b) idem, œuf déchiré; c) Branchinella spinosa; d) Linderiella massaliensis ;

e) Branchinectella media; f) Tanymastix stellae; g) Tanymastix stagnalis; h) œuf immature, Drepanosurus birostratus; i)

Chirocephalus orghidani; j) Chirocephalus diaphanus; k) Chirocephalus salinus (échelle commune en pm).

— 120 —

C. slovacicus (Fig. 9e), C. spinicaudatus (Fig. 9f), C. shadini (Fig. 9g), and C. josephinae

(Fig. 9h), or have thin and high ridges in C. orghidani (Fig. 8i), C. diaphanus (Fig. 8j), C. salinus

(Fig. 8k), and C. pelagonicus (Fig. 9a). As far as grouping distinctions are concerned (see Brtek

& Thiéry, 1995), a typical pattern seems to be dominant in each Chirocephalus group. In the

bairdi- group (4 species), three species (C. orghidani - this paper, Fig. 8i; C. brevipalpis -

PETKOVSKI 1991a, Fig. le; and C. kerkyrensis - MURA et al, 1978, PI. 4; Mura, 1986, PI. 10;

1992b. PI. 2.1, 2) have a pattern of polygonal areas formed by high and thin crests. This pattern

is also present in Chirocephalus bairdi (Fig. lOa-c), a species of the Levant. Cysts of the last

Fig. 9. — Cysts of Anostraca: a) Chirocephalus pelagonicus ; b) Chirocephalus diaphanus carinatus ; c), d) C. chyzeri\ e) C.

slovacicus ; f) C. spinicaudatus (note the verucose aspect under the external crust, arrows); g) C. shadini ; h) C. josephinae ;

i) C. camuntanus (common scale bar in pm).

Œufs d'Anostraca : a) Chirocephalus pelagonicus; b) Chirocephalus diaphanus carinatus; c), d) C. chyzeri ; e) C. slovacicus;

f) C. spinicaudatus ( noter l’aspect verruqueux, flèches) ; g) C. shadini ; h) C. josephinae; i) C. camuntanus (échelle commune

en \im).

— 122 —

species, C. vornatscheri, are unknown. In the diaphanus- group (8 species), a cyst surface pattern

of the type already described for the bairdi- group, is found in four species (C. diaphanus, C.

salinus, C. pelagonicus - this paper, Figs 8j, k; 9a, and C. sibyllae - MURA, 1986, PI. 9.b, and

1992b, PI. 2.7). A spiny appearance is observed in two species (C. diaphanus carinatus - this

paper, and C. rujfoi - Mura, 1986, Pis 12 and 13.a, and 1992b, PI. 2.8, PI. 5.6), while only

one has a smooth aspect (C. marchesonii - MURA, 1986, PI. 4, 1992b, PI. 2.9). Cysts of C. reiseri

are unknown. The pristicephalus- group (4 species) is more heterogeneous with one spiny cyst

(C. carnuntanus. Figs 9i; lOi, j, k, 1), one slightly verrucose (C. shadini. Fig. 9g), or sometimes

more verrucose (Fig. lOf). and a cyst clearly bumpy (C. josephinae, Figs 9h, lOh). The spines

of Chirocephalus carnuntanus differ from those of C. diaphanus carinatus, Linderiella massal-

iensis, and C. rujfoi (see Mura, 1986, Pis 12.b, 13. a) by their length and their “trident” aspect

in their distal border (Fig. lOi-k). Cysts of the fourth species, Chirocephalus ripophilus, remain

unknown. In the spinicaudatus-group (6 species), three cysts have a verrucose aspect (C. spini-

caudatus, Fig. 9f; C. chyzeri. Fig. 9c, d; and C. slovacicus, Figs 9e; 10g); those of C. croaticus,

horribilis, and robustus, are unknown.

Chirocephalidae Eubranchipodinae (Figs 6, 7, 8): cysts of Drepanosurus hankoi are spherical

with irregular, unconnected low ridges or vertical lamellae, 10-15 pm high, non converging and

situated without any regularity (Fig. 7a). The surface is also rugose. This pattern is described

by Mura (1992b, Plate 5.2). As we could study the cyst of only one of the three European

species of Drepanosurus, nothing can be said about the generic pattern, but we can note that

the pattern found in D. hankoi agrees with Daday’s description (1910, Fig. 33g, 1, p. 244) of

D. birostratus whose cysts are ova superficie spinulosa, spinalis bacilliformibus. The cysts of

D. birostratus described in the present study (Fig. 8h) are immature. No data are available for

D. vladimiri.

Cyst of Siphonophanes grubii somewhat resemble those of Drepanosurus hankoi but with

thin wrinkled low ribs, connected to one another (Fig. 6f, g), and arranged in a dense complex

coral-like network delineating small irregular areas. The wrinkled ribs are 14-17 pm high

(Fig. 7c). (see earlier descriptions made by Thiéry & Gasc, 1991, Fig. 14, and Mura, 1992b,

Plate 5.1.

Chirocephalidae Branchinectellinae (Figs 7, 8): cysts of Branchinectella media are smooth

(Figs 7e; 8e; Alonso & Alcaraz, 1984, Fig. 2.e) and superficially similar to those of Artemia.

The structure of the shell, however, is clearly different, with an inner alveolar layer which consists

of vertical strands approximately 15 pm long and 1-2 pm thick (Fig. It), while in Artemia it is

only spongy (Fig. 7h).

Notostraca. Triopsidae (Figs 11, 12): cysts are spherical with a smooth or finely rugose

surface. Only the diameter can be used in identification but variation within each species (V. I.

from 13 to 26%), makes distinction difficult, particularly between Triops cancriformis and Lepidu-

rus couesii or between Lepidurus couesii and L. apus.

Notostracans have the largest resting eggs with spherical cyst up to 400 pm in diameter

(Fig. 12). Triops cancriformis, Lepidurus couesii and L. apus have the smallest cysts with re¬

spectively diameters of 350-400 pm, 425 pm and 430-520 pm. Lepidurus arcticus cysts are larger

(603-621 pm) and have a rough surface while those of Lepidurus apus, L. couesii and Triops

cancriformis are smooth (Figs 11, 12). The alveolar envelope is also thinner than in other

— 123 —

Fig. 11. — Cysts of Notostraca: a), b) Triops cancriformis\ c) idem, detail of surface; d) idem, cysts stuck on gravel; e) Lepidurus

apus, cyst stuck on gravel (g). (Scale bars in (im.)

Œufs de Notostraca : a), b) Triops cancriformis ; c) idem, détail de la surface ; d) idem, œuf collé à des graviers ; e) Lepidurus

apus, œuf collé sur un gravier (g). (Échelles en | xm.)

notostracans (from 25 to 70 pm) while the alveolar layer reaches sometimes 110 pm. As reported

by Thiéry (1985) and Fryer (1988) cysts are mostly covered with sediments and gravels

(Fig. lid, e).

Cysts of the Afro-Asian Triops numidicus, penetrating Europe only in Sicily and Majorca,

is not illustrated here. A description is given in Thiéry (1985, 1995), and Thiéry (in press)

(spherical and smooth cyst, diameter 610-740 pm).

Spinicaudata (Figs 13, 14). Cyzicidae (Fig. 13): cysts of Cyzicus species are spherical,

smooth with a very thin membrane (< 2 pm) covering a network of thin radiating and entangled

setules (Fig. 13f, g, i) approximately 0.4 pm thick and 10-15 pm long (= the vitelline envelope

sensu Tommasini & Scanabissi Sabelli, 1989). The membrane is sometimes cracked (Fig. 13f,

i) allowing the setose envelope to be observed without any treatment. As discussed by Thiéry

& Gasc (1991) and Thiéry (1995), this egg shell structure can be considered generic. For other

descriptions of Cyzicus cysts see Daday (1914: Cyzicus tetracerus)\ Alonso & Alcaraz (1984:

Cyzicus grubei), Thiéry (in press: Cyzicus gihoni), and Thiéry (in prep.: Cyzicus bucheti).

Eocyzicus has a different egg shell pattern. The external surface of E. orientalis cyst is

bumpy (Fig. I3h, j), as described by Daday (1914: Ova membrana concinne granulata tecta).

Fig. 12. — Cysts of Notostraca: a) Lepidurus apus', b) surface of the cyst of Lepidurus arcticus with numerous craters (cr.); c)

Lepidurus couesii ; d) cysts of Lepidurus apus glued together by cement (ce.); e) L arcticus, cross section of wall showing

alveolar layer (a.l.), embryonic cuticle (e.c.), crater (cr.); f) L. apus, section of cyst wall with alveolar layer (a.l.), inner layer

(i.l.); g), h) surface of L. apus cysts showing in h) the obliterated craters (arrows). (Scale bars in pm.)

Œufs de Notostraca : a) Lepidurus apus; b) Lepidurus arcticus, noter la présence de cratères (c.r.) en surface ; c) Lepidurus

couesii; d) œufs de Lepidurus apus agglutinés par un cément (ce.); e) L. articus, coupe transversale de la paroi montrant

la couche alvéolaire (a.l.), la cuticule embryonnaire (e.c.) et les cratères (cr.); f) L. apus, coupe transversale de la paroi,

couche alvéolaire (a.l.), couche interne (i.l.); g), h) surface des œufs de L. apus montrant (en h)) des cratères oblitérés

(flèches). (Échelles en [lm).

Fig. 13. — Cysts of conchostracans (Spinicaudata): a), b), c) Limnadia lenticularis (b), c): same scale); d), e) Imnadia yeyetta

(same scale); f), g) Cyzicus tetracerus\ h) Eocyzicus orientalis (f), g), h): same scale); i) cracked egg shell, C. tetracerus-,

j) outer surface, E. orientalis. (scale bars in pm).

Œufs de conchostracés (Spinicaudata) : a), b), c) Limnadia lenticularis (b), c) : même échelle) ; d), e) Imnadia yeyetta (même

échelle); f), g) Cyzicus tetracerus; h) Eocyzicus orientalis (f), g), h): même échelle); i) enveloppe externe déchirée de

C. tetracerus; j) surface externe de E. orientalis (échelles en p/n).

— 126 —

A similar pattern has been described for other species of the genus by Samyiah et al. (1985:

= Caenestheriella), and THIÉRY (1987; 1995). Cysts of Eocyzicus propinqus, E. skorikowi and

E. tadei were not available for our study.

Leptestheriidae (Fig. 14): cysts of Leptestheria and Eoleptestheria are spherical and smooth

with no clearly distinct differences (Fig. 14a-d). Species and genera of this family may be sep¬

arated by adult characters only. The cyst pattern of several species of the Leptestheriidae - small

with a smooth surface - has already been presented by Botnariuc (1947, PI. I, Fig. 7: Leptes¬

theria intermedia ), ThiÉRY (1987, 1995: Leptestheria mayeti), TOMMASINI & SCANABISSI SABELLI

(1989, Figs 11, 12: Leptestheria dahalacencis), and BRENDONCK et al. (1993: Leptestheria aegyp-

tiaca).

Fig. 14. — Cysts of conchostracans (Spinicaudata and Laevicaudata): a)-c) Leptestheria dahalacencis ; d) idem, not fully mature

cyst deformed by preparation for scanning; e) idem, surface; f) detail of surface, Eoleptestheria ticinensis\ g) Lynceus brachy-

urus ; h) same species, detail of the surface showing depressions (arrows). (a)-d), g): same scale.) (Scale bars in pm.)

Œufs de conchostracés (Spinicaudata et Laevicaudata) : a)-c) Leptestheria dahalacencis; d) idem, œuf immature déformé

par la préparation pour le scanning ; e) idem, surface ; f) détail de la surface chez Eoleptestheria ticinensis ; g) Lynceus

brachyurus; h) même espèce, détail des dépressions de surface (flèches). (a)-d), g): même échelle.) (Échelles en \lm.)

— 127 —

Limnadiidae (Fig. 13): cysts of Limnadia and Imnadia are now well known (Thiéry &

Gasc, 1991). The present material supplies additional pictures but no new information. In Lim¬

nadia, the cyst is clam-shell like and biconcave with ridges converging to an apical point

(Fig. 13a-c), so that Martin (1989) called them “twisted eggs”. No objective difference could

be observed between the cysts of American populations described by Martin (1989) and those

of European populations described by Sars (1896), Daday (1925), Zaffagnini (1967), and

Thiéry & Gasc (1991).

Imnadia cysts have a spiral aspect (Fig. 13d, e), as illustrated by Botnariuc (1947), Btrek

(1957, Fig. 2h), and THIÉRY & Gasc (1991). In both Limnadia and Imnadia the cyst patterns

are unique in the “large branchiopods” and can be considered as generic.

LAEVICAUDATA (Fig. 14): cysts of Lynceus brachyurus are spherical and look like a small

golf-ball with more or less angular depressions (Fig. 14g, h). They are among the smaller cysts

of European phyllopods (approximately 120 pm). Cysts of Lynceus andronachensis illustrated

by Botnariuc (1947, PI. I, Fig. 15) resembles those of L. brachyurus.

DISCUSSION

From our observations, complemented by reliable data in the literature, it appears that the

species present a relatively constant morphology of the mature cysts over their whole distribu¬

tional area. However, the fact that the surface pattern and size sometimes presents relative var¬

iability is hardly discussed by Mura (1991a, 1992a, b). In the case of phyllopods it is true that

several species, occupying a large distribution area, present more or less important differences

in size between populations of different habitats particularly between those in lowlands and high

mountains (Thiéry, 1987, in prep.; Belk et al., 1990; Mura, 1991b). In these cases the size

is statistically different but the external pattern remains quite similar.

When species have cysts with similar surface patterns, their size usually allows their iden¬

tification. When the two characters are unable to allow a clear distinction, it is useful to consider

the structure of the shell, as in the case of Artemia and Branchinectella media (spongy vs roots/pil¬

lars). Only a small number of species have markedly similar cysts ( Leptestheria and Eoleptes-

theria in the Leptestheriidae, Branchinecta paludosa and B. ferox, the different species of the

genus Tanymastix, and several notostracans). In the case of Branchinecta orientalis and B. minuta,

the fact that the cysts closely resemble each other in pattern and size raises the question of an

eventual synonymy, as was the case for Branchipus blanchardi and B. alpinus (see the respective

cysts in Mura, 1986, PI. 13b, and Thiéry & Gasc, 1991, Figs 16, 28).

For about 80% of the European species, the combination of surface pattern/size is a valid

means of species identification. In some other cases, e.g. Tanymastix, Branchipus, Cyzicus, cyst

morphology presents only a generic peculiarity.

As observed in most of species of Anostraca (Hellstrôm & Nauwerck, 1971; Thiéry,

1987), the number of resting eggs is related to the length of the female. Except some cases

(Belk et al., 1990), the size of cysts is usually approximately constant from the first to last

batches (Belk, 1977; Mura, 1991b). Considering the absolute value of the size of the cysts

within the Anostraca, Notostraca, Spinicaudata and Laevicaudata, our study allows us to

— 128 —

distinguish three groups: notostracans with eggs larger than 450 pm, conchostracans with eggs

less than 200 pm, and, between these groups anostracans with cysts from 250 to 450 pm. This is

in accord with the gradient found by Thiéry (1987) for Moroccan “large branchiopods”, by Thiéry

& Gasc (1991) for French species, and by Brendock et al. (1993) for tropical African species.

Our study needs, however, to be completed by the description of cysts of species which

are lacking in this paper (lack of material such as Chirocephalus ripophilus, C. wbustus , C.

reiseri, C. horribilis, or immature cysts such as in the case of Drepanosurus birostratus). Also,

even though egg morphology seems to be a useful species- or genus-specific character, the full

extent of its taxonomic value in some cases remains limited. Some cysts of different species

are very similar. Despite cases which are the subject of conflicting opinions, cyst morphology,

shape, and size taken together often represent a useful adjunct to identification which should

be included along with classical species descriptions, as was done for example by Maeda-

Martinez et al. (1992. 1993), and Smith (1992). However, the choice of cysts (maturity level)

and measurements need stringent attention, to avoid the confusions or errors (erroneous scale

bars) which cause some data to be unusable. (Errors and omissions found in all references about

cysts are noted in Table I).

Cyst data also provide additional support for the close relationship recognized between cer¬

tain anostracans on other grounds by Daday (1910). For example, in the Chirocephalidae, if

the cyst morphology pattern of different species follows in part the groups determined on classical

characters of the male (antennae, penes), it is sometimes discordant ( e.g. the heterogenous Pris-

ticephalus-grovip). In consequence it will be interesting to study large series of cysts from different

populations to prove or disprove statistically that the size of cysts is more or less constant and

not correlated with the length/age of the female. It will be also useful to consider whether size

is linked with altitude, as in Triops cancriformis and Chirocephalus diaphanus in Morocco

(Thiéry, 1987), or in Chirocephalus in Italy (Mura, 1991b).

Finally, the morphological study of cysts has reached its limits and needs to be completed

by biochemical studies which may provide insights into the formation of so great a variety of

egg shells in the “large branchiopods” (spiny, bumpy, lenticular, spiral, twisted, smooth, etc.).

Research dealing with this subject, previously discussed by Thiéry (1987, p. 338), is in progress.

We think also that isoelectric focusing data could help to clarify the taxonomy of several families,

the Chirocephalids for example. This technique could also measure the genetic distance between

the five endemic species of Branchipus, the genetic distance between the Linderiellidae and

Chirocephalidae, between Leptestheria and Eoleptestheria, and help determine a hierarchy in

characters (importance of the cyst vs penes, or antennae). We will use biochemical methods

which were initiated in anostracan studies by SlEDEL & SIMPSON (1984), NAVARRO et al. (1987),

Requintina & Simpson (1987), Fugate (1992), Thiéry & Fugate (1994), for more extensive

research.

Acknowledgements

We thank Dr G. Fryer for the loan of specimens of Lepidurus arcticus from Iceland, Dr N. A.

Akatova (Saint Petersbourg), Pr G. Mura (Roma), Dr S. Petkosvki (Skopje), and Dr N. Vekhoff (Mos¬

cow), for several preserved anostracans, and Dr D. Defaye (MNHN Paris) for access to the Muséum

collections. The first author thanks Dr D. Belk and Pr H. J. Dumont for their useful remarks.

— 129 —

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Thiery (1995) which are not in this table have cysts which remain unknown, n = number of measured cysts, V. I. = variation

index, S = standard error, x = mean. The symbol /?/ in the column of measurements indicates that the value is doubtful,

corresponding most of the time to a scale-bar error.

Liste récapitulative des mesures des œufs des espèces de « phyllopodes » actuellement connues en Europe. Les œufs des

espèces citées dans la liste de BRTEK & ThiÊry (1995) qui ne sont pas dans ce tableau ne sont pas connus, n = nombre

d'œufs mesurés, V. 1. — indice de variation (en %), x - moyenne, S ~ erreur standard. Le symbole /‘I/ dans la colonne des

mesures indique que la valeur tirée d’une référence bibliographique est douteuse, la plupart du temps suite à une erreur

d'échelle.

SPECIES

CYST MEASUREMENTS

[mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

ANOSTRACA

Polyartemia forcipata

527 pm (n = 1)

This study

Russia

(V. I. = 15.76 %)

0.45 mm

Hellstrôm & Nauwerck

(1971)

Sweden

Branchinecta ferox

386 - 412 pm (n = 6)

This study

Slovakia

(V. I. = 24.61 %)

352 - 383 pm (n = 3)

This study

Slovakia

0.36 - 0.42 mm

Alonso (1985)

Spain

error in scale

Alonso & Alcaraz (1984)

Spain

438 pm

Mura & Thiéry (1986)

Morocco

342 pm

Petkovski (19916)

Yugoslavia

0.241 mm

(cysts clearly immature)

César (1989)

MHN Paris

without reference

Branchinecta minuta

290 - 305 pm (n = 3)

This study

Russia

— 134 —

SPECIES

CYST MEASUREMENTS

(mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

Branchinecta orientalis

277 - 280 urn (n = 2)

This study

Austria

(V. 1. = 37.40 %)

219 pm

Alonso & Alcaraz (1984)

Spain

0.22 - 0.32 mm

Alonso (1985)

Spain

263 pm

Petkovski (19916)

Yugoslavia

Branchinecta paludosa

394 - 442 |im (n = 3)

This study

Slovakia

(V. I, = 47.64%)

264 urn /?/

Mura (1991a)

North America

320 - 325 pm (n = 15)

This study

Russia

0.396 mm

César (1989)

MNHN Paris

without reference

0.27 - 0.38 mm

(x = 0.35 mm, n > 30)

Belk (1977)

USA-Arizona

Branchinecta tolli

(V. I. = 7.43 %)

311 - 335 pm (n = 7)

This study

Russia

Artemia parthenoge-

netica

216 - 269 pm [x = 240.3 (am,

SD = 18.43, n = 101

This study

France

(V. I. = 27.51 %)

220 - 270 pm

Thiéry & Gasc (1991)

France

241.2 (im

VlDELA & CASTELA BRANCO

(1987)

Portugal

259 (im

VANHAECKE & SORGELOOS

(1980)

France

266 (am (n = 50)

Vieira & Am at (1985)

Portugal

238.8 (im

Thiéry & Robert (1992)

France

251.8 - 263.1 (im

Castritsi - Catharios et al.

(1987)

Greece

260.1 - 264.7 (am (n = 500)

Abatzopoulos et al. (1987)

Greece

270.1 - 279.3 pm

(in laboratory culture)

Abatzopoulos et al. (1987)

Greece

208 - 234 |am (n = 14)

Thiéry (in press)

Saudi Arabia

15.3 - 28.9 (am /?/

(x = 24.1 (am)

Majic & Vukadin (1987)

Yugoslavia

259.6 (am

Vanhaecke & Sorgeloos

(1980)

France (Aigues-Mortes)

276.3 (im

Vanhaecke & Sorgeloos

(1980)

France (Lavalduc)

264.4 (am

Vanhaecke & Sorgeloos

(1980)

France (Salins de Giraud)

257.8 (im

Vanhaecke & Sorgeloos

(1980)

France (Salins d’Hyères)

253.6 (am

Vanhaecke & Sorgeloos

(1980)

Spain

284.9 (am

Vanhaecke & Sorgeloos

(1980)

Italy

— 135 —

SPECIES

CYST MEASUREMENTS

[mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

0.20 - 0.25 mm

Mathias (1937)

France

255.39 pm (n = 655)

Triantaphyllidis et al.

(1993)

Greece Lesbos Island-

Kallon

269.71 pm (n = 777)

Triantaphyllidis et al.

(1993)

Greece Lesbos

Polychnitos

Artemia tunisiana

208 - 286 |im (n = 5)

This study

France (Sète)

(bisexual)

231.2 - 268.0 [xm

VlDELA & CASTELA BRANCO

(1987)

Portugal

(V. I. = 31.57 %)

245 |im

Thiéry & Gasc (1991)

France (Sète)

245.3 )lm

Thiéry & Robert (1992)

France (Sète)

222-234 pm

(probably a New World bi¬

sexual strain-not A. tunisiana)

Maillard & Baudet

(1980)

France (Guérande)

Branchipus blanchardi

288 - 290 pm (n = 3)

This study

France

(Alps-Cristol plateau)

(V. I. = 19.18 %)

300 pm

Thiéry & Gasc (1991)

France

(Alps-Cristol plateau)

# 0.35 mm

Alonso (1989)

France (Alps)

291 pm

Mura ( 1986)

Italy (Alps)

Branchipus cortesi

0.25 mm

Alonso & Jaume (1991)

Spain

Branchipus inlermedius

200 pm

Orghidan (1947)

Romania

Branchipus schaefferi

228 - 270 pm (n = 5)

This study

France (Languedoc)

(V. I. = 42.36 %)

205 - 250 pm (n = 4)

This study

France (Crau)

238 - 253 Jim (n = 4)

syn. visnyai

This study

Slovakia

254 pm - syn. visnyai

Mura (1986)

Italy

195 pm

Kupka (1940)

not indicated

0.25 mm

Gilchrist (1978)

not indicated

0.2 - 0.3 mm

Alonso (1985)

Spain

0.2 - 0.28 mm

Mathias (1937)

not indicated

320 pm /?/

Mura (1986)

Italy

253 pm

Mura & Thiéry (1986)

Morocco

260 - 289 pm (n = 10)

Thiéry ( 1995)

Saudi Arabia

Tanymastix motasi

0.35 mm

Petkovski (1995)

Macedonia

Tanymastix stagnalis

300/313 -192/195 pm (n = 5)

This study

France

(V. I. = 55.52 %)

0.233 / 0.164 mm

Garreau de Loubresse

(1982)

France

337.8 pm

Alonso & Alcaraz (1984)

Spain

0.40 - 0.43 mm

Alonso (1985)

Spain

— 136 —

SPECIES

CYST MEASUREMENTS

[mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

0.43 mm

Al-Tikrity & Grainger

(1990)

Ireland

0.35 - 0.40 mm

Petkovski (1995)

Macedonia

Tanymastix stellae

370 pm

Champeau & Thiéry

(1990)

Corsica

485 pm /?/

Mura (1986)

Sardinia

Branchinella spinosa

250 - 260 pm

Mura (19926)

Italy

(V. I. = 15.35 %)

260.1 pm

Thiéry & Gasc (1991)

France (Camargue)

223 pm

Mura & Thiéry (1986)

Morocco

224 pm

Mura (1986)

Italy

225 - 235 pm

Thiéry (in press)

Saudi Arabia

Streptocephalus torvi-

coniis torvicornis

263 - 279 pm (n = 3)

This study

Slovakia

(V. I. = 42.76 %)

0.3 mm

Valousek (1952)

Czechoslovakia

0.22 - 0.33 mm

Alonso (1985)

Spain

250 pm

Alonso & Alcaraz (1984)

Spain

240 pm

Mura (1992a)

Origin not indicated

231 - 239 pm (n = 8)

Thiéry (1995)

Saudi Arabia

0.22 mm

Zograf (1907)

Russia

Streptocephalus ton’i-

cornis bucheti

244.3 pm

Alonso & Alcaraz (1984)

Spain

(V. I. = 12.29 %)

276.3 pm

Mura & Thiéry (1986)

Morocco

Linderiella massaliensis

(V. I. = 8.0 %)

240 - 260 pm

Thiéry & Gasc (1991)

France

Linderiella sp.

(V. I. = 12.24 %)

0.23 - 0.26 mm

Alonso (1985)

Spain

255 pm

Alonso & Alcaraz (1984)

Spain

Chirocephalus brevi-

palpis

(316 pm from fig.l.e,

but the text indicates

0.22 - 0.24 mm)

Petkovski (1991a)

Yugoslavia

(erroneous scale bar in the

figure, Petkovski, ab ora)

Chirocephalus carnun-

tanus

240 - 260 pm (n = 4)

This study

Slovakia

0.154 mm (immature cyst)

Zograf (1907)

not indicated

Chirocephalus chyzeri

259 - 310 pm (n = 5)

This study

Slovakia

Chirocephalus diapha-

nus

370 - 429 pm (n = 6)

This study

Slovakia

(V. I. = 52.08 %)

360 - 420 pm

Thiéry & Gasc (1991)

France

340 pm

Alonso & Alcaraz (1984)

Spain

— 137 —

SPECIES

CYST MEASUREMENTS

[mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

287 (im

Mura et ai (1978)

Italy

0.25 - 0.32 mm

Alonso (1985)

Spain

0.43 mm

Hall & Mac Donald

(1975)

England

270 pm

Poisson et al. (1946)

France

0.25 mm

Mathias (1937)

not indicated

error in scale

Mura (1992b)

Italy

Chirocephalus diapha-

nus carinatus

270 - 288 (am (n = 9)

This study

Bulgaria

Chirocephalus josephi-

nae

331 - 345 pm (n = 3)

This study

Russia

0.25 mm

ZoGRAF (1907)

Russia

Chirocephalus kerky-

rensis

266 - 331 |am

Mura (1986)

Italy

(V. I. = 21.48 %)

error in scale

Mura (1992b)

Italy

311 (am (n = 50)

Mura et al. (1978)

Italy

Chirocephalus marche -

soni

428 (am (n = 50)

Mura et al. (1978)

Italy

363 (am

Mura (1986)

Italy

error in scale

Mura (1992b)

Italy

Chirocephalus orghi-

dani (V. I. = 5.99 %)

275 - 292 (am (n = 3)

This study

Roumania

Chirocephalus pelago-

nicus

295 - 296 (am (n = 2)

This study

Yugoslavia

(V. 1. = 12.10 %)

333 |am

Petkovski (1986)

Yugoslavia

Chirocephalus ruffoi

252 |am

Mura ( 1986)

Italy

no scale

Cottarelli & Mura (1984)

Italy

error in scale

Mura (1992b)

Italy

Chirocephalus salinus

345 - 363 |am (n = 4)

This study

Corsica

341 - 360 (am (n = 4)

Thiéry & Gasc (1991)

Corsica

355 (am

Champeau & Thiéry

(1990)

Corsica

256 |am /?/

Mura et al. (1978)

Sardinia

362 (am

Mura (1986)

Sardinia (code 600)

256 (am /?/

Mura (1986)

Sardinia (code 300)

error in scale

Mura (1992b)

not indicated

Chirocephalus shadini

275 - 280 (am (n = 6)

This study

Slovakia

Chirocephalus slovaci-

cus

311 - 325 (am (n = 3)

This study

Slovakia

(V. I. = 10.52 %)

306 - 332 (am (n = 5)

This study

Slovakia

329 - 340 (am (n = 3)

This study

Slovakia

— 138 —

SPECIES

CYST MEASUREMENTS

[mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

Chirocephalus spini-

caudatus

264 pm

This study

France (MNHN Paris

Bp. 157)

Chirocephalus sibyllae

290 [im

Mura (1986)

Italy

error in scale

Mura (1992b)

Italy

Drepanosurus birostra-

trus

173 (tm, but immature cyst

This study

Russia

Drepanosurus hankoi

341 - 358 (tm (n = 5)

This study

Slovakia

300 pm

Mura (1992b)

origin not indicated

Siphonophanes grubii

353 (tm (n = 1)

This study

Slovakia

(MNHN Paris Bp. 260)

(V. I. = 16.23 %)

0.30 - 0.33 mm

Buchholz (1864)

Germany

312 ptrn

Mura (1992b)

not indicated

Branchinectella media

198 - 211 pm (n = 3)

This study

Russia

(V. I. = 6.36 %)

208.6 (im

Alonso & Alcaraz (1984)

Spain

0.2 mm

Alonso (1985)

Spain

NOTOSTRACA

Lepidurus apus

436 - 485 (im (n = 5)

This study

France (Provence) - dried

(V. I. = 26.08 %)

432 - 495 (im (n = 12)

This study

South-France (Gavoti)

455 - 471 (im (n = 4)

This study

France (Languedoc)

496 (tm (n = 1)

This study

Slovakia

0.4 - 0.5 mm

Alonso (1985)

Spain

479 (im

Alonso & Alcaraz (1984)

Spain

0.6 mm

Réau de la Gaignonnière

( du) (1908)

France (Angers)

440 |im

Arnoult (1951)

France (Toulouse)

0.4 - 0.6 mm

Mathias (1937)

not indicated

430 - 520 (tm

Thiéry & Gasc (1991)

France

Lepidurus couesii

420 - 425 (tm (n = 3)

This study

Mongolia

Lepidurus arcticus

603 - 621 (tm (n = 3)

This study

Russia

(V. I. = 14.78 %)

0.7 mm

Braem (1893)

not indicated

700 (tm

Fryer (1988)

Iceland

Triops cancriformis

350 - 380 (tm (n = 3)

This study

Slovakia

(V. I. = 16.21 %)

368 - 372 (tm (n = 6)

This study

France (Opoul)

355 - 375 (im (n = 4)

This study

France (Crau)

0.37 mm

Gilchrist (1978)

England

360 - 400 (im

Thiéry & Gasc (1991)

France (Languedoc)

0.4 - 0.6 mm /?/

Mathias (1937)

France

0.34 mm

Tommasini et al. (1989)

Italy

139 —

SPECIES

CYST MEASUREMENTS

[mini - maxi or mean (n = )]

REFERENCE

IN LITERATURE

LOCALITY

Triops cancriformis

mauritanicus

423 pm

Alonso & Alcaraz (1984)

Spain

(V. I. = 18.98 %)

442 - 510 pm

Thiéry (1987)

Morocco

Triops cancriformis

simplex

391 |im

Alonso & Alcaraz (1984)

Spain

(V. I. = 5.52 %)

370 pm

Thiéry (1987)

Morocco

Triops numidicus

1 mm

Ghigi (1921)

not indicated

625 - 750 |im

Thiéry (1987)

Morocco

SPINICAUDATA

Limnadia lenticularis

185 - 209 |im (n = 2)

This study

Slovakia

(V. I. = 12.18 %)

199 - 208 pm (n = 2)

This study

Slovakia

200 - 208 pm

Thiéry & G asc (1991)

not indicated

0.15 - 0.22 mm

Lereboullet (1866)

not indicated

0.15 - 0.17 mm

Zaffagnini (1967)

Italy

Imnadia yeyetta

180/180 - 185/190 pm (n =

This study

Slovakia

(V. I. = 11.23 %)

165/150 - 175/160 urn (n =

This study

France (Crau)

185/182 urn

Thiéry & G asc (1991)

not indicated

Cyzicus tetracerus

146 - 161 pm (n = 7)

This study

Slovakia

(V. I. = 14.30 %)

141 - 153 pm (n = 5)

This study

Slovakia

145 - 150 urn

Thiéry & Gasc (1991)

France (Provence)

139.5 pm

Alonso & Alcaraz (1984)

Spain

0.15 - 0.16 mm

Mathias (1937)

France

Cyzicus grubei

124 pm

Alonso & Alcaraz (1984)

Spain

Cyzicus bucheti

150 - 190 pm

Thiéry (1987)

Morocco

Eocyzicus orientalis

119 - 122 pm (n = 2)

This study

Russia

(V. I. = 2.49 %)

Leptestheria dahala-

cencis

95 - 103 pm (n = 3)

This study

Slovakia

(V. I. = 14.63 %)

103 - 110 pm (n = 8)

This study

Slovakia

100 |xm

SCANABISSl SaBELLI &

Tommasini (1992)

Italy

Leptestheria mayeti

179 pm

Thiéry (1987)

Morocco

Eoleptestheria ticinensis

125 - 130 pm (n = 5)

This study

Slovakia

(V. I. = 6.35 %)

122 - 126 |im (n = 4)

This study

Slovakia

127 pm

Thiéry & Gasc (1991)

France (Camargue)

LAEVICAUDATA

Lynceus brachyurus

114 - 119 |J.m (n = 2)

This study

Slovakia

(V. I. = 4.29 %)

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 141-162.

Three new species of Hemicyclops

(Copepoda: Poecilostomatoida: Clausidiidae)

from northwestern Madagascar

by Arthur G. HUMES

Abstract. — Three new species of the copepod genus Hemicyclops (Clausidiidae) from Nosy Bé, north¬

western Madagascar, are described, from burrows of unknown origin, probably those of crustaceans, one from

the scleractinian Favia sp., and one from a gastropod shell occupied by a hermit crab. Hemicyclops acanthophorus ,

new species, may be recognized by a pair of posterolateral spiniform processes on the genital double-somite of

the female, and by the triangular form of the free segment of leg 5. Hemicyclops apiculus , new species, is

characterized by its elongate genital double-somite with anterior rounded lateral expansions, its subquadrate caudal

ramus, and the small sharply pointed lateral projections on the first segment of leg 5. Hemicyclops vicinalis,

new species, may be distinguished by the absence of setules on the inner side of the caudal ramus and on the

outer margin of the second segment of the antenna proximal to the seta.

Keywords. — Copepoda, Poecilostomatoida, Clausidiidae, Hemicyclops , Madagascar.

Trois nouvelles espèces d ’Hemicyclops (Copepoda : Poecilostomatoida : Clausidiidae)

du nord-ouest de Madagascar

Résumé. — Trois espèces nouvelles du genre Hemicyclops (Copepoda : Clausidiidae) de Nosy Bé, au nord-

ouest de Madagascar, sont décrites, l’une provenant de terriers d’origine inconnue, probablement de crustacés,

une autre provenant du scléractiniaire Favia sp., et la troisième d’une coquille de gastéropode occupée par un

bernard-l’hermite. Hemicyclops acanthophorus n. sp. peut être identifiée par une paire de prolongements spini-

formes postérolatéraux sur le somite génital de la femelle et la forme triangulaire du segment libre de la cinquième

patte. Hemicyclops apiculus n. sp. se caractérise par un somite génital allongé avec des expansions latérales

antérieures arrondies, des rames caudales subcarrées et la présence de petites projections pointues latérales sur

le premier segment de la cinquième patte. Hemicyclops vicinalis n. sp. peut être identifiée par l’absence de

sétules sur le bord interne de la rame caudale et au bord externe du deuxième segment de l'antenne.

Mots-clés. — Copépodes, Poecilostomatoida, Clausidiidae, Hemicyclops, Madagascar.

A. G. Humes, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts, 02543, USA.

INTRODUCTION

Since the revision of Hemicyclops by VERVOORT & RAMIREZ (1966), in which 23 valid

species were recognized, several congeners have been described, extending the range of this

clausidiid genus to Hong Kong (Boxshall & Humes, 1987), Korea (Ho & Kim, 1990, 1991;

Kim & Ho, 1992), Panama (HUMES, 1984), Curaçao (STOCK, 1992), Japan (ITOH & NlSHlDA,

1993), and Brazil (Kihara & da Rocha, 1994).

In addition to the species recognized in the work of Vervoort & Ramirez (1966), the

following species have been described : Hemicyclops perinsignis Humes, 1973, from the sponge

Agelas sp. in Madagascar; H. columnaris Humes, 1984, from the scleractinian coral Pontes lobata

- 142 —

Dana on the Pacific coast of Panama; H. mortoni Boxshall & Humes, 1987, from burrows of

the echiuran Ochetostoma erythrogrammon Leuckhart & Rüppell at Hong Kong; H. ctenidis Ho

& Kim, 1990, from the polychaete Neanthes japonica (Izuka) in Korea; H. gomsoensis Ho &

Kim, 1991, from burrows occupied by the crab Macrophthalmus japonicus de Haan in Korea;

H. saxatilis Ho & Kim, 1991, from burrows occupied by the crab Heteropanope (Pilumnopeus)

makinana (Rathbun) in Korea; H. geminatus Stock, 1992, from the hermit crabs Calcinus tibicen

(Herbst), Paguristes grayi Benedict, and Dardanus venosus (H. Milne Edwards) at Curaçao;

H. caissarum Kihara & da Rocha, 1993, from burrows of the callianassid Callichirus major (Say)

in Brazil; H. japonicus Itoh & Nishida, 1993, from dredged material in Tokyo Bay, Japan; and

H. sebastiani Kihara & da Rocha, 1993, from the thalassinid Callichirus guassutinga (Rodriguez)

in Brazil.

In this paper, three new species of Hemicyclops, all from northwestern Madagascar, are

described. The number of valid species in the genus now stands at 36.

MATERIAL AND METHODS

The copepods from intertidal burrows were collected at low tide by means of a small bilge

pump. In other cases, the host invertebrates were washed in a dilute solution of ethanol in sea

water (approximately 5%) and the copepods recovered from the sediment obtained after passing

the wash water through a fine net.

Lactic acid was used to clear the copepods for measurements and dissection. The length

of the body does not include the setae on the caudal rami. The segments of the antennule were

measured along their posterior nonsetiferous margins. In the formula for legs 1-4. Roman numer¬

als indicate spines, Arabic numerals represent setae.

POECILOSTOMATOIDA Thorell, 1859

Family Clausidiidae Embleton, 1901

HEMICYCLOPS Boeck, 1872

Hemicyclops acanthophorus n. sp.

(Figs 1-4)

Type material. — 13 9 2,5 $ â from large burrows in intertidal sand, Befotaka, Nosy Bé, northwestern

Madagascar, 29 April 1964. Holotype 2 (MNHN - Cp 1052), allotype â (MNHN - Cp 1053), and 12paratypes

(9 2 2, 3 ââ) (MNHN - Cp 1054) deposited in the Muséum national d’Histoire naturelle, Paris. Remaining

paratypes (dissected) and copepodids (3 2 2, 1 S ) in the collection of the author.

Etymology. — The name of the species is formed from the Greek words akantha , a thorn or spine, and

phorein, to bear or carry, alluding to the pair of small spiniform processes on the genital double-somite of the

female.

Description of female

Body (Fig. la) relatively slender. Length 1.35 mm (1.25-1.45 mm) and greatest width

0.56 mm (0.50-0.61 mm), based on 10 specimens. Greatest dorsoventral thickness 0.37 mm.

Epimera of somites bearing legs 1-4 flaring outward and pointed. Somite bearing leg 1 fused

— 143 —

Fig. 1. — Hemicyclops acanthophorus, n. sp., female, a, dorsal (scale A); b, urosome, dorsal (B); c, spiniform process on genital

double-somite, lateral (C); d, anal somite and caudal ramus, dorsal (D); e, median part of cephalosome, showing rostrum,

labrum, postoral area, and area between maxillipeds and first pair of legs, ventral (B); f, antennule, posteroventral (E).

Ai = antennule, A 2 = antenna, MXPD = maxilliped. Pi = leg 1.

Hemicyclops acanthophorus, n. sp., femelle, a, vue dorsale (échelle A); b, urosome, vue dorsale (B); c, processus spiniforme

sur le double somite génital, vue latérale (C) ; d, somite anal et rames caudales, vue dorsale (D) ; e, partie médiane du

céphalosome, montrant le rostre, le labre, la région postorale et la zone située entre les maxillipèdes et la première paire

de pattes, vue ventrale (B); f, antennule, vue postéro-ventrale (E). Ai = antennule, A 2 = antenne, MXPD = maxillipède.

Pi = première paire de pattes.

with cephalosome. Ratio of length to width of prosome 1.66:1. Ratio of length of prosome to

that of urosome 1.80: 1.

Somite bearing leg 5 (Fig. lb) 101 x 218 pm. Genital double-somite 112 x 177 pm, wider

than long, in dorsal view with gently rounded lateral margins and bearing pair of small post¬

erolateral, unequally bifid, spiniform processes about 15 pm long (Fig. lc), perhaps part of mod¬

ified leg 6. Genital areas located laterally but no evidence of setae. Three postgenital somites

78 x 127, 75 x 103, and 88 x 86 pm.

Caudal ramus (Fig. Id) unornamented, elongate, 75 x 26 pm, ratio 3: I. Outer lateral seta

30 pm, dorsal seta 52 pm, outermost terminal seta 40 pm. innermost terminal seta 65 pm, all

smooth. Two long median terminal setae 180 pm (outer) and 450 pm (inner), both with delicate

short lateral setules.

Body surface without visible sensilla.

Egg sac not seen.

Rostrum (Fig. le) broadly rounded posteroventrally. Antennule (Fig. If) 308 pm long.

Lengths of 7 segments : 13 (44 pm along anterior margin), 39, 36, 60, 39, 39, and 39 pm, re¬

spectively. Formula for armature: 4, 15, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete.

All setae smooth. Distal seta on segment 4 long, 250 pm.

Antenna (Fig. 2a) 4-segmented, with armature 1. 1,4, and 7. First segment with long distal

seta having lateral setules. Second segment with shorter smooth seta. Third segment with 3 slender

setae, one of them with setules, and 1 swollen almost clawlike seta with recurved tip. Fourth

segment with 6 smooth setae and 1 seta with setules.

Labrum (Fig. le) with marginal row of spines and pair of rows of small submarginal spines.

Postoral area with 2 rows of spines (Fig. 2b).

Mandible (Fig. 2c) terminally with 2 stout elements and 2 setae. Paragnath (Fig. 2b) elongate

lobe, 48 pm, with few minute setules. Maxillule (Fig. 2d) bearing 8 setae. Maxilla (Fig. 2e) with

first segment bearing 2 long setae with setules and 1 very small setule; second segment with

3 setae and large bifurcate element. Maxilliped (Fig. 2f) 4-segmented, with 2 setae on first seg¬

ment, 2 setae on second segment, and 6 setae on fourth segment, one of them slightly bent and

having lateral setules.

Ventral area between maxillipeds and first pair of legs as in Fig. le.

Legs 1-4 (Figs 2g, h, 3a, b) segmented and armed as follows :

coxa 0-1

basis l-I

P 2

coxa 0-1

basis 1-0

P 3

coxa 0-1

basis 1-0

P-l

coxa 0-1

basis 1-0

exp 1-0;

1-1;

1,1,6

enp 0-1;

0-1;

1,5

exp 1-0;

1-1;

11,11,5

enp 0-1;

0-2;

1,11,3

exp 1-0;

1-1;

11,1,5

enp 0-1;

0-2;

1,11,3

exp 1-0;

I-1 ;

1,11,5

enp 0-1;

0-2;

1.11,2

Intercoxal plate of leg 1 with slender setules on ventral margin; intercoxal plates of legs 2-4

with marginal spines.

— 145 —

Fig. 2. — Hemicyclops acanthophorus, n. sp., female, a, antenna, posterior (scale D); b, postoral area and paragnaths, posterior

(D) ; c, mandible, posterior (C); d, maxillule, posterior (C); e, maxilla, posterior (C); f, maxilliped, posterior (C); g, leg 1

and intercoxal plate, anterior (E); h, leg 2 and intercoxal plate, anterior (E).

Hemicyclops acanthophorus, n. sp., femelle, a, antenne, face postérieure ( échelle D) ; b, région postorale et paragnathes,

vue postérieure (D); c, mandibule, vue postérieure (C) ; d, maxillule, vue postérieure (C); e, maxille, vue postérieure (C) ;

f maxillipède, vue postérieure (C) ; g, Pi et plaque intercoxale, face antérieure (E) ; h, P 2 et plaque intercoxale, face antérieure

(E) .

— 146 —

Leg 1 with inner spine on basis 50 |im. Other details of ornamentation as illustrated.

Leg 5 (Fig. 3c) with free segment triangular, 57 pm along inner side, 68 pm along outer

side and 65 pm wide, armed with 3 spines and 1 small seta, these elements from outer to inner

47, 50, 22, and 42 pm. Segment ornamented near insertion of outermost seta with group of

spinules. Dorsal seta 70 pm, with few spinules near its insertion.

Leg 6 not identified.

Living specimens in transmitted light opaque gray, eye red.

Description of male

Prosome slightly broader (Fig. 3d) than in female. Length 1.05 mm (1.00-1.08 mm) and

greatest width 0.44 mm (0.43-0.46 mm), based on 5 specimens. Greatest dorsoventral thickness

0.28 mm. Internal sclerotizations visible along sides of prosome as in Fig. 3e. Somite bearing

leg 4 having somewhat different form (Fig. 3d) than in female. Ratio of length to width of pro-

some 1.61:1. Ratio of length of prosome to that of urosome 1.40:1.

Segment bearing leg 5 (Fig. 4a) 70 x 153 pm. Genital somite 146 x 180 xm, shape similar

to that of female, but lacking pair of spiniform processes. Four postgenital somites 65 x 112,

60 x 91, 40 x 81, and 42 x 74 pm.

Caudal ramus like that of female but smaller, 61 x 25 pm.

Body surface unomamented.

Rostrum similar to that of female. Antennule like that of female, but extra seta added on

segments 3 and 4. Antenna like that of female.

Labrum, mandible, paragnath, maxillule, and maxilla resembling those of female. Maxilliped

(Fig. 4b, c) with long seta on first segment. Second segment expanded, with 2 inner setae and

2 rows of spines, those in one row stout, those in other row slender. Claw 148 pm long with

slight protuberance on concave margin, and bearing 2 small proximal setae.

Postoral area as in female.

Leg 1 (Fig. 4d) lacking inner spine on basis (as in many congeners), but otherwise as in

female. Legs 2-4 like those of female.

Leg 5 (Fig. 4e) with free segment elongate, 91 x 47 pm, ornamented with spinules along

both sides; armature as in female.

Leg 6 (Fig. 4f) consisting of posteroventral flap on genital somite bearing 1 seta 26 pm.

Spermatophore not seen.

Color as in female.

Remarks

Hemicyclops acanthophorus may be distinguished from all other species in the genus by

the presence of a pair of small thornlike posterolateral spiniform processes on the genital double¬

somite of the female. The triangular shape of the free segment of leg 5 in the female is also

characteristic.

Although the occupants of the burrows in which the copepods were found is not known,

it is likely that they were crustaceans. Several species of Hemicyclops are known from burrows

of crustaceans at Nosy Bé, Madagascar: 11. axiophilus Humes, 1965, and //. amplicaudatus

— 147 —

Fig. 3. — Hemicyclops acanthophorus, n. sp., female, a, leg 3 and intercoxal plate, anterior (scale E); b, leg 4 and intercoxal

plate, anterior (E); c, leg 5, dorsal (C). Male, d, dorsal (F); e, side of prosome showing internal sclerotizations, dorsal (F).

Ai = antennule, A 2 = antenna, MD = mandible, MX 2 = maxilla, MXPD = maxilliped.

Hemicyclops acanthophorus, n. sp., femelle, a, P3 et plaque intercoxale, face antérieure (échelle E) ; b, P4 et plaque inter-

coxale, face antérieure (E) ; c, P5, vue dorsale (C). Mâle, d, vue dorsale (F) ; e, bord du prosome montrant des sclérotisations

internes, vue dorsale (F). Ai = antennule, A 2 = antenne, MD = mandibule, MX 2 = maxille ; MXPD = maxillipède.

— 148 —

Fig. 4. — Hemicyclops acanthophorus, n. sp., male, a, urosome, dorsal (scale B); b, maxilliped, posterior (D); c, maxilliped,

anterior (D); d, leg 1 and intercoxal plate, anterior (E); e, leg 5, ventral (D); f, genital somite, ventral (E).

Hemicyclops acanthophorus, n. sp., mâle, a, urosome, vue dorsale (échelle B); b, maxillipède, vue postérieure (D) ; c, maxil-

lipède, vue antérieure (D) ; d, Pi et plaque intercoxale, vue antérieure (E) ; e, P 5 , vue ventrale (D) ; f, somite génital, face

ventrale (E).

— 149 —

Humes, 1965, from burrows of the thalassinidean macruran Axius (Neaxius) acanthus (Axiidae);

H. acanthosquillae Humes, 1965, from the body of the stomatopod Acanthosquilla sp. (= A.

humesi Manning) dug from intertidal sand; and H visendus Humes, Cressey, and Gooding. 1958,

from the body surface of the thalassinidean macruran Upogebia (Upogebia) sp. (Callianassidae)

in burrows (Humes, 1965).

Hemicyclops apiculus n. sp.

(Figs 5-7)

Type material. — 3 2 2 from washing of one colony of the scleractinian coral Favia sp., in 3 m, Pointe-

à-la-Fièvre, Nosy Bé, northwestern Madagascar, 26 December 1963. Holotype (MNHN - Cp 1055) and 1 paratype

(MNHN - Cp 1056) deposited in (he Muséum national d’Histoire naturelle, Paris. Remaining paratype (dissected)

in the collection of the author.

Other specimen. — 1 2 from the ahermatypic coral Dendrophyllia sp., in 7 m, Nosy Tangam, near Nosy

Bé, 1 January 1964.

Etymology. — The specific name apiculus , Latin meaning “pointed”, alludes to the small sharp projection

on the first segment of leg 5.

Description of female

Body (Fig. 5a, b) moderately slender. Length 1.42 mm (1.36-1.50 mm) and greatest width

0.51 mm (0.48-0.54 mm), based on 3 specimens. Greatest dorsoventral thickness 0.33 mm. Somite

bearing leg 1 fused with cephalosome. Epimera of somites bearing legs 1-4 pointed posteriorly

in somites bearing legs 1 and 2, less pointed in somites bearing legs 3 and 4. Ratio of length

to width of prosome 1.58:1. Ratio of length of prosome to that of urosome 1.22:1.

Somite bearing leg 5 (Fig. 5c) 109 x 268 pm. Genital double-somite in dorsal view much

longer than wide, 273 pm long, 213 pm wide in expanded anterior third, and 122 pm wide in

posterior two-thirds with parallel sides. Posterolateral corners of anterior expanded portion bluntly

produced. Ratio of length to greatest width 1.28:1. Genital openings situated laterally (Fig. 5b)

and lacking setae but showing 2 minute knoblike projections. Three postgenital somites from

anterior to posterior 107 x 130, 78 x 122, and 65 x 120 pm.

Caudal ramus (Fig. 6a) subquadrate, 65 x 60 pm, only slightly longer than wide. Outer

lateral seta 44 pm, dorsal seta 88 pm, both smooth. Outermost terminal seta 96 pm, innermost

terminal seta 208 pm, and 2 long median terminal setae 360 pm (outer) and 580 pm (inner), all

with lateral setules, those on 2 median setae long and placed well apart. Postero-inner corner

of ramus with small spiniform projection 5 pm long.

Body surface without visible ornamentation.

Egg sac not seen.

Rostrum (Fig. 6b) triangular and bluntly pointed posteroventrally. Antennule (Fig. 6c)

385 pm long. Lengths of its 7 segments: 17 (52 pm along anterior margin), 47, 31, 90, 60, 44,

and 52 pm, respectively. Formula for armature: 4, 15, 6, 3, 4 + I aesthete, 2 + 1 aesthete, and

7 + 1 aesthete. All setae smooth.

Antenna (Fig. 6d) 4-segmented with formula 1, 1.3+ 1 spine, and 7. Second segment with

setules on both sides, third segment with outer setules, and fourth segment with inner setules.

Seta on segment 1 with bilateral setules. Seta on segment 2 with long unilateral setules. One

— 150 —

Fig. 5. — Hemicyclops apiculus, n. sp., female, a, dorsal (scale G); b, lateral (G); c, urosome, dorsal (B).

Hemicyclops apiculus, n. sp., femelle, a, vue dorsale (échelle G); b, vue latérale (G); c, urosome, vue dorsale (B).

seta on segment 3 with several unilateral spinules, other setae and spine smooth. One seta on

segment 4 with long spinules, other 6 setae smooth, 4 longest of these slightly geniculate.

Labrum (Fig. 6b) with row of short spinules anteriorly, followed by row of small blunt

spinules; another interrupted row of small setae posteriorly. Postoral area (Fig. 6e) with 6 sub¬

marginal spines and row of minute marginal spinules; corners of labium with prominent tooth.

— 151 —

Fig. 6. — Hemicyclops apiculus, n. sp., female, a, anal somite and caudal ramus, dorsal (scale D); b, rostrum and labrum, ventral

(E); c, antennule, ventral (E); d, antenna, anterior (E); e, postoral area, ventral (H); f, mandible, anterior (D); g, maxillule,

anterior (D); h, maxilla, anterior (D); i, second segment of maxilla, posterior (C). Ai = antennule, A 2 = antenna.

Hemicyclops apiculus, n. sp., femelle, a, somite anal et rames caudales, vue dorsale (échelle D) ; b, rostre et labre, vue

ventrale (E) ; c, antennule, face ventrale (E) ; d, antenne, face antérieure (E) ; e, région postorale, vue ventrale (H) ; f,

mandibule, vue antérieure (D) ; g, maxillule, vue antérieure (D); h, maxille, vue antérieure (D) ; i, deuxième segment de la

maxille, face postérieure (C). Ai = antennule, A 2 - antenne.

— 152 —

Mandible (Fig. 6f) terminally with 2 stout elements, one dentate, other spinulose, and 2

setae. Maxillule (Fig. 6g) bearing 8 setae, in groups of 5 + 3. Maxilla (Fig. 6h) with first segment

bearing 2 long setae. Second segment (Fig. 6i) bearing 1 smooth seta, clawlike spine bearing

1 inner spinule, and 1 spiniform seta with unilateral setules; segment with prolongation bearing

terminal row of 8 smooth spines. Maxilliped (Fig. 7a) with 2 setae on both first and second seg¬

ments. Small third segment unarmed. Minute fourth segment with 2 unequal clawlike spines

62 pm and 42 pm, and 3 slender setae.

Ventral area between maxillipeds and first pair of legs slightly protuberant (Fig. 5b) and

appearing as in Fig. 7b.

Legs 1-4 (Fig. 7c-f) with 3-segmented rami armed as follows:

coxa 0-1

basis 1-1

coxa 0-1

basis 1-0

P 3

coxa 0-1

basis 1-0

P*1

coxa 0-1

basis 1-0

exp 1-0;

1-1;

11,6

enp 0-1;

0-1;

1,5

exp 1-0;

1-1;

111,1,5

enp 0-1;

0-2;

UI,3

exp 1-0;

1-1;

111,1,5

enp 0-1;

0-2;

11,11,2

exp 1-0;

1-1;

11,1,5

enp 0-1 ;

0-2;

IIJI.l

Intercoxal plate of leg 1 with slender setules on ventral margin; intercoxal plates of legs

2-4 with marginal spines.

Basis of leg 1 with inner spine 50 pm. Exopod of leg 1 with 2 distal spines on third segment

setiform, with small spinules along outer margin and setules along inner margin; innermost seta

on third segment of endopod slender, short, and smooth. Distal spine on third segment of exopod

in legs 2-4 setiform.

Leg 5 (Fig. 7g) 2-segmented, with free segment in dorsal view 99 x 55 pm, in flat view

94 x 60 pm (as in Fig. 7h). Four terminal elements (3 spines and 1 seta) from outer to inner 39,

45, 65, and 58 pm. Both sides of free segment with prominent setules. First segment, fused with

body somite, having pointed outer projection and bearing dorsal seta 60 pm.

Leg 6 not identified but possibly represented by 2 minute knobs (Fig. 5b).

Color of living specimens in transmitted light slightly yellow, eye red, ovaries lemon yellow.

Male unknown.

Remarks

Among the many species of Hemicyclops, seven have an elongate genital double-somite

with rounded anterior expansions, as in the new species. Hemicyclops apiculus, new species,

may be distinguished from the female of all seven of these congeners in the following ways.

In H. kombensis Humes, 1965, and H. acanthosquillae Humes, 1965, the body is more than 2 mm

in length. In H. biflagellatus Humes, 1965, the body length is more than 2 mm and the somite

bearing leg 5 has a pair of setiform processes (setae?). In H. columnaris Humes, 1984, the body

is smaller (average length 1.13 mm) and the ratio of length to width of the caudal ramus is

— 153 —

FIG. 7. — Hemicyclops apiculus, n. sp., female, a, maxilliped, posterior (scale D); b, area between maxillipeds and first pair of

legs, ventral (E); c, leg 1 and intercoxal plate, anterior (B); d, leg 2, and intercoxal plate, anterior (B); e, leg 3 and intercoxal

plate, anterior (B); f, leg 4 and intercoxal plate, anterior (B); g, leg 5, dorsal (C); h, second segment of leg 5, flat ventral

view (C). MXPD = maxilliped, Pi = leg 1.

Hemicyclops apiculus, n. sp., femelle, a, maxillipède, vue postérieure ( échelle D) ; b, zone située entre les maxillipèdes et

la première paire de pattes, face ventrale (E) ; c, Pi et plaque intercoxale, face antérieure (B); d, P 2 et plaque intercoxale,

face antérieure (B) ; e, P 3 et plaque intercoxale, vue antérieure (B); f P 4 et plaque intercoxale, face antérieure (B); g, P 5 ,

vue dorsale (C) ; h, deuxième segment de P 5 , vue ventrale (C). MXPD = maxillipède. Pi - première paire de pattes.

— 154 —

1.7:1. In H. perinsignis Humes, 1973, the free segment of leg 5 is short and broad, ratio 1.5:1,

and the ratio of the caudal ramus is 1.72:1. In H. saxatilis Ho & Kim, 1991, the body length

is about 2:1, and. the genital double-somite has 2 pointed leaflike extensions laterally. Hemicy-

clops australis Nicholls, 1944, is poorly known, but may be distinguished by the posterolateral

projections on the swollen portion of the genital double-somite.

Hemicyclops vicinalis n. sp.

(Figs 8-11)

Type material. — 2 $ 9, 1 <? from one hermit crab, Dardanus guttatus (Olivier), in shell of Conus sp.,

1.5 m, Ankify, near Nosy Bé, Madagascar, 48°20’15”E, 13°30’30"S, 30 December 1963. Holotype $ (MNHN -

Cp 1057) and allotype â (dissected) (MNHN - Cp 1058) deposited in the Muséum national d’Histoire naturelle,

Paris. One paratype 9 (dissected) in the collection of the author.

Etymology. — The specific name vicinalis, Latin meaning “neighboring” or “near”, alludes to the close

similarity in external anatomy to H. columnaris and H. geminatus Stock, 1992.

Description of female

Body (Fig. 8a, b) with moderately broad prosome. Length 1.16 mm (1.08-1.23 mm) and

greatest width 0.44 mm (0.42-0.46 mm), based on 2 specimens. Greatest dorsoventral thickness

0.31 mm. Somite bearing leg 1 not separated dorsally from cephalosome. Epimera of metasomal

somites rounded except pointed on somite bearing leg 1. Ratio of length to width of prosome

1.26:1. Ratio of length of prosome to that of urosome 1.11:1.

Somite bearing leg 1 (Fig. 8c) 109 x 218 pm. Genital double-somite elongate, cylindrical,

in dorsal view 239 pm long, 185 pm wide at small lateral swellings in anterior third, and 156 pm

wide posteriorly. Ratio of length to greatest width 1.77:1. Genital openings, lacking setae or

spines, situated laterally at level of lateral swellings. Three postgenital somites from anterior to

posterior 88 x 133, 55 x 120, and 52 x 109 pm. Anal somite with posteroventral row of spines

on its distal edge (Fig. 8d).

Caudal ramus (Fig. 8d) subquadrate, little wider proximally than distally, length 61 pm, pro¬

ximal width 50 pm, distal width 44 pm. Ratio (taking width at middle) 1.17:1. Outer lateral seta

47 pm and dorsal seta 95 pm, both smooth. Outermost terminal seta 96 pm, innermost terminal

seta 170 pm, both with small lateral setules. Two median terminal setae in holotype: outer 350 pm

with lateral setules, inner broken at proximal joint. Inner distal corner of ramus with minute

setule 3 pm long.

Body surface lacking visible ornamentation.

Egg sac (Fig. 8e) elongate, 440 x 132 pm, with many eggs, each with diameter of approxi¬

mately 42 pm.

Rostrum (Fig. 8f) broadly rounded. Antennule (Fig. 9a) 320 pm long, lengths of its 7 seg¬

ments: 13 (44 pm along anterior margin), 37, 34, 55, 39, 44, and 42 pm, respectively. Armature:

4, 15, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete. All setae smooth. Antenna

(Fig. 8b) 4-segmented, with armature 1, 1,4, and 7. Seta on second segment relatively short

and weak. Otherwise similar to Hemicyclops acanthophorus, new species, except for minor seta-

tion.

— 155-

fig. 8. — Hemicyclops vicinalis, n. sp., female, a, dorsal (scale I); b, lateral (I); c, urosome, dorsal (B); d, anal somite and caudal

ramus, dorsal (D); e, egg sac, ventral (A); f, rostrum and edge of labrum, ventral (D). Ai = antennule, A 2 = antenna.

Hemicyclops vicinalis, n. sp., femelle, a, vue dorsale ( échelle I) ; b, vue latérale (I) ; c, urosorne, vue dorsale (B); d, somite

anal et rames caudales, vue dorsale (D) ; e, sac ovigère, vue ventrale (A); f, rostre et bord du labre, vue ventrale (D). Ai

= antennule, A 2 = antenne.

— 156 —

Fig. 9. — Hemicyclops vicinalis , n. sp., female, a, antennule, dorsal (scale E); b, antenna, anterior (D); c, labrum, ventral (C);

d, mandible, posteroventral (C); e, postoral area and paragnaths, ventral (C); f, maxilla, posterior (D); g, maxilliped, posterior

(D); h, area between maxillipeds and first pair of legs, ventral (B); i, leg 1 and intercoxal plate, anterior (E). MXPD =

maxilliped. Pi = leg 1.

Hemicyclops vicinalis, n. sp., femelle, a, antennule, face dorsale (échelle E) ; b, antenne, face antérieure (D) ; c, labre, face

ventrale (C); d, mandibule, vue posté ro-v entrale (C) ; e, région post-orale et paragnathes, vue ventrale (C) ; f, maxille, face

postérieure (D); g, maxillipède, vue postérieure (D) ; h, zone située entre les maxillipèdes et la première paire de pattes,

vue ventrale (B) ; i, Pi et plaque intercoxale, face antérieure (E). MXPD - maxillipède. Pi = première paire de pattes.

— 157 —

Labrum (Fig. 9c) with anterior row of long setae and subanterior row of blunt hyaline spines,

and having 2 rows of setules at each side. Postoral area (Fig. 9e) with row of hyaline spines

and median finely setulose area. Mandible (Fig. 9d) with stout elongate nonsetulose element,

spatulate element with marginal setules, and 2 slender setae, one spiniform with lateral setules.

Paragnath (Fig. 9e) attenuated lobe with distal hairlike setules. Maxillule as in Fig. 11c of male.

Maxilla (Fig. 9f) and maxilliped (Fig. 9g) as illustrated

Ventral area between maxillipeds and first pair of legs protruding slightly (Fig. 8b). Two

small sclerites in front of intercoxal plate of leg 1 (Fig. 9h).

Legs 1-4 (Figs 9i, lOa-c) with segmentation and armature as follows:

coxa 0-1

basis 1-1

coxa 0-1

basis 1-0

coxa 0-1

basis 1-0

coxa 0-1

basis 1-0

exp 1-0;

1-1;

11,6

enp 0-1;

0-1;

1,5

exp 1-0;

I-1 ;

111,1,5

enp 0-1;

0-2;

1,11,3

exp 1-0;

I-1 ;

111,1,5

enp 0-1;

0-2;

1,11,3

exp 1-0;

1-1;

11,11,5

enp 0-1;

0-2;

1,11,2

Intercoxal plate of leg 1 with slender setules along ventral edge; intercoxal plates of legs

2-4 with marginal spines.

In leg 1 inner spine on basis 49 pm long. In all 4 legs distal spine on third segment of

exopod setiform, and outer margin of coxa and basis with setules. Other setules and spinules

resembling those in other species, for example, Hemicyclops columnaris Humes, 1984.

Leg 5 (Fig. lOd) 2-segmented. First segment with dorsal seta 50 pm long. Second segment

elongate, narrow proximally, widened distally, 117 x 50 pm (greatest width). Ratio 2.34:1. Ter¬

minally with 3 spines and 1 seta, from outer to inner 44, 36, 62, and 52 pm. Innermost spine

with setules along inner margin. Seta reaching only to middle of genital double-somite. Both

outer and inner margins of second segment with strong setules.

Leg 6 not identified.

Living specimens in transmitted light nearly translucid, eye red, egg sacs dull orange brown.

Description of male

Body (Figs 10e, 11a) resembling in general form that of female. Length (of allotype)

1.25 mm and width 0.45 mm. Greatest dorsoventral thickness 0.26 mm. Ratio of length to width

of prosome 1.57:1. Ratio of length of prosome to that of urosome 1.24:1.

Somite bearing leg 5 (Fig. lib) 91 x 198 pm. Genital somite subquadrate, 146 x 156 pm,

slightly wider than long. Four postgenital somites from anterior to posterior 112 x 133, 88 x

122, 60 x 114, and 48 x 104 pm. Anal somite with posteroventral spinules as in female.

Caudal ramus like that of female.

Body surface lacking visible ornamentation.

— 158 —

Fig. 10. — Hemicyclops vicinalis, n. sp., female, a, leg 2 and intercoxal plate, anterior (scale E); b, leg 3 and intercoxal plate,

anterior (E); c, leg 4 and intercoxal plate, anterior (E); d, leg 5, ventral (D). Male, e, dorsal (I).

Hemicyclops vicinalis, n. sp., femelle, a, P 2 et plaque intereoxale, face antérieure ( échelle E) ; b, Pj et plaque intercoxale,

face antérieure (E); c, P 4 et plaque intercoxale, face antérieure (E); d, P 5 , vue ventrale (D). Mâle, e, vue dorsale (I).

— 159 —

Fig. II. — Hemicyclops vicinalis, n. sp., male, a, lateral (scale 1); b, urosome, dorsal (B); c, maxillule, antero-inner (D); d,

maxilla, anterior (D); e, second segment of maxilla, anterior (C); f, maxilliped, anterior (D); g, endopod of leg 1, anterior

(E); h, endopod of leg 2, anterior (E); i, leg 5, ventral (D); j, genital somite showing sixth pair of legs (E).

Hemicyclops vicinalis, n. sp., mâle. a. vue latérale ( échelle I) ; b, urosome, vue dorsale (B); c, maxillule, vue antéro-inteme

(D); d, maxille, face antérieure (D) ; e, deuxième segment de la maxille, face antérieure (C); f, maxillipède, vue antérieure

(D); g, endopodite de Pi, face antérieure (E) ; h, endopodite de P 2 , face antérieure (E); i, P 5 , vue ventrale (D) ; j, somite

génital montrant la sixième paire de pattes (E).

— 160 —

Rostrum as in female. Antennule similar to female but 1 seta added on segment 3 and

another seta on segment 4. Antenna like that of female. Labrum, postoral area, mandible, par-

agnath, and maxillule (Fig. lie) resembling those of female. Maxilla (Fig. lid) with 2 setae on

segment 1; segment 2 stoutly pointed and bearing 4 setae (Fig. 1 le). Maxilliped (Fig. Ilf) similar

to that of H. columnaris.

Legs 1-4 with segmentation and armature as in female, but basis of leg 1 lacking inner

spine seen in female (Fig. lg). Segments of endopod of leg 2 (Fig. 11 h) somewhat shorter than

in female.

Leg 5 (Fig. Hi) with second segment 127 x 49 pm. Ratio 2.59:1. Less tapered than in

female but otherwise similar.

Leg 6 (Fig. 1 lj) consisting of posteroventral flap on genital segment bearing 1 finely barbed

spinelike seta 36 pm.

Spermatophore not seen.

Color as in female.

Remarks

Three species of Hemicyclops, H. columnaris from Pacific Panama, H. geminatus from Cura¬

çao, and H. vicinalis from Madagascar, are at first glance remarkably similar. All have, in the

female, an elongate columnar genital double-somite with small rounded anterior lateral expansions

and a short caudal ramus with a length to width ratio less than 2:1. Flowever, there are subtle

differences that separate them and in particular distinguish the new species. In H. vicinalis, the

inner margin of the caudal ramus is smooth (in H. columnaris and H. geminatus this margin

bears setules); the length to width ratio of the caudal ramus in the female is 1.17:1 (1.7:1 in

H. columnaris, 1.27-1.52:1 in H. geminatus); the outer side of segment 2 of the antenna proximal

to the seta lacks setules (these setules present in H. columnaris and H. geminatus ); the second

segment of leg 5 is 117 x 50 pm (125 x 57 pm in H. columnaris, 85 x 40 pm in H. geminatus);

and the inner of the two stout elements on the mandible is tapered proximally and is expanded

and rounded distally (in H. columnaris this element is not tapered proximally and is rounded

distally, in H. geminatus it is not tapered proximally and has a truncate tip). In H. columnaris

the outermost terminal seta on the caudal ramus has a small thornlike process on its outer edge,

not found in the two congeners mentioned. In H. geminatus the seta on the second segment of

leg 5 in the female is much longer than in H. vicinalis, “reaching to 75-90% of length of genital

segment” (Stock, 1992).

In all three species discussed, there are ventrodistal spinules on the anal somite (these not

mentioned in the original description of H. columnaris).

Notes on Hemicyclops perinsignis Humes, 1973

This species, described from specimens associated with the sponge Agelas, has now been

found associated with the alcyonacean coral Tubipora musica L. as follows: 5 9 9 , 1 c? from 1

colony, Pointe Lokobe, Nosy Bé, Madagascar, 5 June 1967; 2 9 9, 1 <3 from 1 colony, same

locality, 12 June 1967.

— 161 —

Acknowledgements

The three new copepods described herein were collected during the International Indian Ocean Ex¬

pedition (1963-1964) while the author was chief scientist at Nosy Bé, Madagascar. The study of the material

was supported by a grant from the National Science Foundation of the United States (BSR 8821979).

REFERENCES

BOXSHALL, G. A., & A. G. Humes, 1987. — A new species of Hemicyclops (Copepoda: Poecilostomatoida)

associated with an echiuran worm in Hong Kong. Asian Mar. Biol.. 4 : 61-66.

HO, J.-S., & I.-H. KIM, 1990. — Hemicyclops ctenidis, a new poecilostomatoid copepod (Clausidiidae) associated

with a polychaete in Korea. Korean J. ZooL, 33: 231-237.

— 1991. — Two new species of the genus Hemicyclops (Copepoda, Poecilostomatoida, Clausidiidae) from

crab burrows in the Yellow Sea. Korean J. Zool., 34 : 289-299.

HUMES, A. G., 1965. — New species of Hemicyclops (Copepoda, Cyclopoida) from Madagascar. Bull. Mus.

Comp. Zool., 134:159-260.

— 1973. — Hemicyclops perinsignis, a new copepod from a sponge in Madagascar. Proc. Biol. Soc. Wash.,

86: 315-328.

— 1984. — Hemicyclops columnaris sp. n. (Copepoda, Poecilostomatoida, Clausidiidae) associated with a coral

in Panama (Pacific side). Zool. Scripta, 13 : 33-39.

HUMES, a. G., R. F., Cressey & R. U., Gooding, 1958. — A new cyclopoid copepod Hemicyclops visendus,

associated with Upogebia in Madagascar. J. Wash. Acad. Sci., 48 : 398-405.

ITOH, H., & S. NlSHIDA, 1993. — A new species of Hemicyclops (Copepoda, Poecilostomatoida) from a dredged

area in Tokyo Bay, Japan. Hydrobiologia, 254 : 149-157.

Kihara, T. C., & C. E. F. da ROCHA, 1993. — Two new species of Hemicyclops associated with mud shrimps

of the genus Callichirus from Brazil. Bijdr. Dierk., 63:243-254.

KIM, I.-H., & J.-S. Ho, 1992. — Copepodid stages of Hemicyclops ctenidis Ho and Kim, 1990 (Clausidiidae),

a poecilostomatoid copepod associated with a polychaete. J. Crust. Biol., 12:631-646.

NICHOLLS, A. G., 1944. — Littoral copepods from South Australia (II) Calanoida, Cyclopoida, Notodelphyoida,

Monstrilloida and Caligoida. Rec. So. Austral. Mus., 8:1-62.

STOCK, J. H., 1992. — A new species of Hemicyclops (Crustacea, Copepoda, Poecilostomatoida, Clausidiidae)

associated with hermit crabs in Curaçao. Stud. Nat. Hist. Carib. Region, 71 : 69-78.

VERVOORT, W., & F. Ramirez, 1966. — Hemicyclops thalassius nov. spec. (Copepoda, Cyclopoida) from Mar

del Plata, with revisionary notes on the family Clausidiidae. Zool. Med., 41 : 195-220.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 163-192.

Barnacles (Cirripedia: Thoracica)

of seas off the Straits of Gibraltar

by Brian A. FOSTER 1 and John S. BUCKERIDGE

Abstract. — The barnacles collected in dredges and grabs from stations in and off the Straits of Gibraltar

(BALGIM 1984 cruise) which were sorted at CENTOB in Brest, are systematically considered. A few other

deep-sea barnacles are also considered. Of the eighteen species illustrated, three are new to science. Particular

attention is given to species of Scalpellinae with eleven species (two new), and Verrucidae with four species

(one new). This paper determines only the species and their biogeography, leaving aside questions of generic

categorisation, phylogeny, and the particular ecological and hydrological circumstances of each cruise.

Keywords. — Gibraltar, Poecilasmatidae, Scalpellidae. Verrucidae, Pachylasmatidae, Balanidae, new taxa,

biogeography.

Cirripèdes (Cirripedia, Thoracica) récoltés dans les eaux du détroit de Gibraltar

Résumé. — Les Cirripèdes récoltés, principalement à l'aide de dragues et de bennes, au voisinage et dans

le détroit de Gibraltar lors de la campagne BALGIM, sont étudiés ici d’un point de vue systématique. Quelques

autres Cirripèdes de mer profonde sont également examinés. Tout le matériel étudié a été trié au CENTOB à

Brest. Au total dix-huit espèces sont recensées et figurées, trois d’entre elles sont nouvelles pour la science. Une

attention particulière est portée aux espèces des sous-familles Scalpellinae, au nombre de onze dont deux nouvelles,

et Verrucidae, au nombre de quatre dont une nouvelle. Dans ce travail, nous n'avons considéré que l’identification

des espèces et leur biogéographie. Les problèmes posés par les genres et la phylogénie, ainsi que les conditions

hydrologiques et les particularités écologiques propres à chaque campagne n’ont pas été abordés.

Mots-clés. — Gibraltar, Poecilasmatidae, Scalpellidae, Verrucidae, Pachylasmatidae, Balanidae, espèces nou¬

velles, biogéographie.

B. A. Foster & J. S. Buckeridge, Department of Civil and Environmental Engineering, UNITEC, Private Bag 92025, Auckland,

New Zealand

INTRODUCTION

This paper reports on a collection of barnacles the late Professor Brian FOSTER received

from the Centre national de tri d’océanographie biologique (CENTOB), Brest. They were col¬

lected by the BALGIM expedition, during May-June 1984, from either side of the Straits of

Gibraltar, particularly into the Ibero-Moroccan Bay of the Atlantic Ocean, by R. V. “Cryos” on

behalf of the Centre national de la recherche scientifique (PIROCEAN) under the direction of

Dr P. Bouchet. Eighteen different species of cirripedes, from 126 stations were collected. A

further ten samples from various deep-sea cruises in the Atlantic Ocean and three samples from

the CORINDON II cruise in Indonesia were also forwarded for determination and contained

specimens useful for present purposes. All stations and species data are listed in Appendix A.

1. Deceased, formerly of the Zoology Department, University of Auckland, Auckland, New Zealand.

— 164 —

The main collection results primarily from multiple deep-sea dredging in geographically

confined areas, and includes deep-sea material which has been known only from few specimens

and inadequate descriptions. The type localities for a number of early descriptions of Verruca

(sensu lato ) and Scalpellum (sensu lato) result from collections from “Talisman" cruises in the

Atlantic from the Bay of Biscay to the Cape Verde Islands, “Travailleur” cruises in Mediterranean

and the Bay of Biscay, and cruises by the Prince of Monaco in the Azores area (Aurivillius,

1898; Gruvel, 1900a, 1900b, 1902). The BALGIM material enables a better appreciation of

specific variation covered by a number of nominate species described from these early cruises.

This study deals solely with systematics and broad biogeography, leaving bathymetry, hy¬

drology and ecology for later consideration.

The paper is derived from a manuscript submitted to the Bulletin du Muséum national d'His-

toire naturelle , Paris, prior to Brian Foster’s untimely death. As the original manuscript incor¬

porated a study of both Indian and Atlantic faunas, it was recommended that it be split, with

each geographic area considered separately. The results for the Indian Ocean cruises has now

been published (Foster & Buckeridge, 1994). This manuscript incorporates any changes in

cirripede phylogeny that have occurred since the preparation of Foster’s manuscript, and further

drawings have been added where necessary, ensuring that all described taxa are illustrated.

MATERIALS AND METHODS

The cruise and station data relevant to the various samples are given in Appendix A. In

the following systematic treatment, the stations are referred to by the nomenclature on the labels

included in the bottles, namely collecting gear and station number. The gear used was as follows;

CP, beam trawl; DC, Charcot-Picard dredge; DR, rock dredge; DS, Sanders dredge; DW, Waren

dredge; FA, Faubert bottom net trawl; PI, submersible dive or SCUBA diver.

Specimens were sorted at CENTOB and preserved in alcohol. They were examined with

the aid of microscopy and dissection. Species regarded as new to science are given a full des¬

cription. Others are illustrated from specimens, and their anatomy commented on where existing

descriptions need amplification or to note points useful in identification. Illustrations of whole

animals and shells were drawn with the aid of a drawing tube. Appendages were mounted in

polyvinyl lactophenol and drawn with a camera lucida. Segments of cirri and caudal appendages

were counted per rami, and where a single average figure is given it has been rounded up to

an integer value for the complete rami.

All material is lodged in the Muséum national d’ Histoire naturelle (MNHN), Paris.

SYSTEMATIC PART

List of Species

Family Poecilasmatidae Annandale, 1910

Octolasmis nierstraszi (Hoek, 1907)

— 165 —

Poecilasma kaempferi Darwin, 1851

Family Scalpellidae Pilsbry, 1907

Smilium acutum (Hoek, 1883)

Trianguloscalpellum regium (W. Thomson, 1873)

Trianguloscalpellum gigas (Hoek, 1883)

Arcoscalpellum michelottianum (Seguenza, 1876)

Arcoscalpellum crenulatum sp. nov.

Amigdoscalpellum praeceps (Hoek, 1907)

Verum ?minutum (Hoek, 1883)

Meroscalpellum bifurcatum (Zevina, 1973)

Verum novaezelandiae (Hoek, 1883)

Verum frillosum sp. nov.

Ornatoscalpellum stroemii (Sars, 1859)

Scalpellum scalpellum (Linnaeus, 1767)

Family Verrucidae Darwin, 1854

Verruca trisulcata Gruvel, 1900

Altiverruca gibbosa Hoek, 1883

Altiverruca vertica sp. nov.

Metaverruca recta (Aurivillius, 1898)

Family Pachylasmatidae Buckeridge, 1983

Pachylasma giganteum (Philippi, 1836)

Family Bathylasmatidae Newman & Ross, 1971

Bathylasma hirsutum (Hoek, 1883)

Family Balanidae Leach, 1817

Megabalanus tulipiformes (Ellis, 1758).

Family POECILASMATIDAE Annandale, 1910

Octolasmis nierstraszi (Hoek, 1907)

(Fig. 1A)

Material. — CORINDON II: station 263, Strait of Makassar, 80 m (1 specimen).

Records. — Hoek, 1907a: 21, 19 stations East Indies, 16-120 m; Nilsson-Cantell, 1921: 268, Japan, 135 m;

Nilsson-Cantell, 1927: 762, Persian Gulf; Nilsson-Cantell, 1934a: 60, Malaysia, 73 m; Broch, 1931: 40, Kei

Is, 38-50 m; Hiro, 1937a: 414, Japan, 24 m (i.e., Persian Gulf, Malaysia to Japan, 16-135 m). Foster & Buck¬

eridge, 1994: 348, La Réunion, 75-92 m. On hydroids, etc.

Remarks

The bilobed tergum and narrowly split scutum clearly distinguish this species, but it is

apparent that there is variation in the shape of the basal arm of the scutum, in some cases

making the scutum tri-lobed.

— 166 —

Poecilasma kaempferi Darwin, 1851

(Fig. IB)

Material. — BALGIM: CP156, 1135 m (1 specimen); DW157, 1108 m (1).

Remarks

This is a frequently reported species (see Zevina, 1982: 98-100 for full records and sy¬

nonymies; also Rosell, 1989: 22 for a new Philippines record), as an epizooite on benthic

invertebrates, particularly decapod crustaceans, circumglobal, 126-1885 m. The subspecies auran-

tia Darwin, dubium Hoek, litum Pilsbry and novaeangliae Pilsbry are probably ecotype variants.

In specimens collected from the same lobster or crab in New Zealand waters, there is a wide

range of shell variations, seemingly associated with position on, or proximity to mechanical

disturbances from the various appendages of the host.

Family SCALPELLIDAE Pilsbry, 1907

Subfamily Calanticinae Zevina, 1978

Smilium acutum (Hoek, 1883)

(Fig. 1C)

Material. — BALGIM: CP65, 1805 m (1 specimen); CP68, 2035 m (3); CP69, 2028 m (23); DR82, 355 m

(4). BIOGAS V: CP07 (off Cape Finisterre), 2170 m (5). EPI I: CP38 (Gulf of Gascogne), 2100 m (1); CP39,

2175 m (1). BIOCYAN II (Bay of Biscay): PL18, 2000 m (3).

Recorded distribution. — Circum-global, 61-2480 m. See Zevina, 1981: 82 and Newman & Ross, 1971:

38 for synonymies and references; also Rosell, 1989: 19, Philippines, 882-770 m.

Remarks

This widely distributed, frequently encountered and distinctive barnacle was first described

by Hoek (1883) from material from both the North Atlantic and the South-West Pacific Oceans.

It occurs on a wide variety of substrata, including hydroids, crustaceans, spicules. It nicely

Fig. 1. — Poecilasmatidae and Scalpellidae: A,

Octolasmis nierstraszi CORINDON II

Stn 263. B, Poecilasma kaempferi BAL¬

GIM, CPI56. C, Smilium acutum BAL¬

GIM, CP69. Scales in mm. Capitular

plates: c, carina; s, scutum, t, tergum, ul,

upper latus, c.l., carinolatus, rl, rostrola-

tus, iml, inframedian latus, sc, subcarina.

Poecilasmatidae et Scalpellidae: A, Octo¬

lasmis nierstraszi CORINDON II Stn 263.

B, Poecilasma kaempferi BALGIM,

CP 156. C, Smilium acutum, BALGIM,

CP69. Échelles en mm. Plaques capitu¬

laires : c, carina ; s, scutum, t, tergum, ul,

latus supérieur, c.l., carinolatus, rl, rostro-

latus, iml, latus inframédian, sc, subcarina.

— 167 —

demonstrates the degree of variation in shell morphology (mostly ontogenetic change in capitulum

dimensions — the capitulum becoming more elongate and the tergum reflexed, the peduncle

with more numerous closely compact scales), and a circum-global distribution in moderately

deep water.

The collection of 41 specimens from 8 stations (355-2175 m) in the North Atlantic Ocean

of this report is comparable to the 199 specimens from 23 stations (391-1220 m) resulting from

New Zealand Oceanographic Institute sampling in New Zealand waters (FOSTER, 1978).

Subfamily Scalpellinae Pilsbry, 1907

Trianguloscalpellum regium (W. Thomson, 1873)

(Figs 2A-D, 3)

Material. — BIOGAS VII (Bay of Biscay): CP28, 3380 m (2 specimens). BIOGAS VIII (Bay of Biscay):

CP30, 3100 m (21).

FIG. 2. — Scalpellidae: A-D: Trianguloscalpellum regium. A, BIOGAS VII CP30 (47.0 mm c.l.). B. BIOGAS VII CP28, (36.0 mm

c.l.). C, BIOGAS VII CP30 (19.0 mm c.l.). D. BIOGAS VII CP30 (11.0 mm c.l.). E, Trianguloscalpellum gigas GASCOR,

CP42 (22.0 mm c.l.). F-I, Arcoscalpellum michelottianum BAGIM, CP95, F, ditto (23.0 mm c.l.). G, ditto , (14.5 mm c.l.).

H, ditto , (10.2 mm c.l.). I, ditto (3.6 mm c.l.). Scales in mm.

Scalpellidae: A-D: Trianguloscalpellum regium. A, BIOGAS VII CP30 (l.c. = 47,0 mm). B. BIOGAS VII CP28, (l.c. = 36,0 mm).

C, BIOGAS VII CP30 (l.c. = 19,0 mm). D, BIOGAS VII CP30 (l.c. = 11,0 mm). E, Trianguloscalpellum gigas GASCOR,

CP42 (l.c. = 22,0 mm). F-I, Arcoscalpellum michelottianum BAGIM, CP95, F, dito (l.c. = 23,0 mm). G, dito, (l.c. = 14,5 mm).

H, dito, (l.c. = 10,2 mm). I, dito (l.c. = 3,6 mm). Échelles en mm.

— 168 —

Records. — Thomson, 1873: 347, and Hoek, 1883: 106, North Atlantic, 5029-5212 m; Gruvel, 1920: 30,

off Cape Finisterre, 2779 m. Not Pilsbry, 1907a: 28, off Chesapeake Bay, 3740 m, = T. gigas, see below. As

molle Aurivillius, 1898: 191, Azores 845-1023 m; and Gruvel, 1920: 29, Azores, 4020 m. As var. ovale Hoek,

1883: 109, North Atlantic, 5212 m. As var. latidorsum Pilsbry, 1907a: 29, off New York, 1812-2940 m. (t'.e.,

North Atlantic, 845-5212 m). Foster & Buckeridge, 1994: 355, La Réunion, 4030 m.

Remarks

The illustrations of 4 specimens from CP30 show the change in capitulum form with growth.

The plates maintain close approximation, except for the carinolatera which become more widely

displaced from each other below the carina. There is a widening of the lateral aspect of the

capitulum to an oval form in larger specimens. The species can reach a large size (47 mm

capitulum length and 33 mm peduncle length in the present material). The plates are robust,

brownish, with relatively smooth surfaces and faint growth ridges. The carina has obvious lateral

sides. The integument between the plates is variously hirsute. The lower latera possess a low

profile, with the apex of the carinolatus characteristically curled across the base of the carina

to slightly overlap the upper latus at this position in smaller specimens.

The mandible is tridentoid. The segmentation of the posterior cirri increases with size of

the animal (Fig. 3); that of cirrus I and the caudal appendages, with lower numbers of segments,

Fig. 3. — Scalpellidae: Trianguloscalpellum regium BIOGAS VII CP30. Mean numbers of segments of rami as a function of

size of specimen. For anterior (a) and posterior (p) rami of cirrus I (average of 2); all rami of cirus 11-VI (average of 4);

both caudal appendages (ca), average of 2.

Scalpellidae: Trianguloscalpellum regium BIOGAS VII CP30. Nombres moyens de segments des rames en fonction de la

taille des spécimens. Rames antérieure (a) et postérieure (p) du cirrhe I (moyenne de 2 relevés) ; ensemble des rames des

cirrhes 7/-V7 (moyenne de 4 relevés) ; ensemble des appendices caudaux (ca), moyenne de 2 relevés.

— 169-

much more constant. The caudal appendage reaches about 0.5 times the length of the pedicle

of cirrus VI. Penis absent. Complemental males carried in interscutal pouches.

Trianguloscalpellum gigas (Hoek, 1883)

(Fig. 2E)

Material. — GASCOR (Bay of Biscay): CP42, 3815 m (3 specimens).

Record. — Hoek, 1883: 102, mid North Pacific, 3749 m.

Remarks

This species is similar to T. regium, but is distinguished by the less rounded profile of the

capitulum, with a proportionally longer upper latus and scutum. The plates are less glossy, and

the carina is rounded, without lateral sides. Most significantly, this species is a hermaphrodite;

it has a penis. The type specimen in the British Museum of Natural History conforms to this.

Hoek (1883) thought that there was a close relationship between T. gigas and T. darwini

(Hoek). This latter species was described on the basis of a single large specimen (48 mm

capitulum length, 31 mm peduncle length) from off Chile in the South-East Pacific. Examination

of this specimen in the British Museum shows it lacks a penis, and it is more likely that Hoek’s

Scalpellum darwini is T. regium, and that PiLSBRY’s (1907) and Zevina’s (1981: 309) specimens

are T. gigas. Both species would thus occur in deep seas around the world.

Arcoscalpellum michelottianum (Seguenza, 1876)

(Figs 2F-H, 4)

Material. — BALGIM: CP10, 1592 m (14 specimens); CP17, 1470 m (3); CP63.1510 m (27); CP65, 1805 m

(5); CP68, 2035 m (10); CP69, 2028 m (2); CP92, 1182 m (15); CP95, 1378 m (25); CP97, 1515 m (9); CP99,

1870 m (10). EPI VI: KG229 (off Brest), 2160 m (1).

Records. — Newman & Ross, 1971: 71, off Greenland, 3404-3422 m and off South Carolina, 3010 m;

Rao & Newman, 1972, Jonston Atoll, Mid-Pacific Ocean; Rosell, 1989: 17, Philippines, 682-770 m. As velutinum

Hoek, 1883: 96, off Cape St Vincent, 1656 m and off Tristan da Cunha, 2606 m; Gruvel, 1902: 56, off Canary

Is, 882-2000 m; Pilsbry, 1907a: 26, western North Atlantic, 64-1962 m; not Annandale, 1913: 229, India, 786 m,

= annandale Caiman, 1918: 109; Calman, 1918: 109, Java-Australia, 732-1280 m, and Aden-Zanzibar, 1097 m;

Gruvel, 1920: 27, Azores, 1331 m, off Nova Scotia, 1458 m; Barnard, 1925: 1, South Africa; Nilsson-Cantell,

1927: 743, Singapore; Broch, 1931: 18, off Hong Kong, 760 m; Stubbings, 1936: 28, Zanzibar, 690-762 m;

Zevina, 1972: 43, South-East Pacific, 660-700 m. As eximium Hoek, 1883: 100, off Tristan da Cunha, 1828 m.

As sordidum Aurivillius, 1898: 190, off New Foundland, 1267 m and off Portugal, 2028 m; Gruvel, 1920: 27,

Canary Is, 1098 m. As erectum Aurivillius, 1898: 192, Azores 1135-1165 m. As alatum Gruvel, 1900a: 192, 1902:

57. Distribution is, therefore, circumglobal, 64-3422 m. Long known as Arcocalpellum velutinum Hoek, 1883 but

synonymised with fossils by Withers (1953).

Remarks

This species is commonly encountered in the Atlantic Ocean (for full synonymy see Zevina,

1981: 343 and Newman & ROSS, 1971: 71). Newman & Ross (1971) gave much needed des¬

cription of young stages. To further augment the description, particularly with regard to changes

that occur during growth, 15 of the specimens from CP95 were dissected, and 4 are illustrated

in Figure 2.

— 170 —

This barnacle is hirsute, particularly in young stages, finely so in the larger specimens. The

plates are approximate, the capitulum elongate because of an elongate scutum and upper latus

(compared with T. gigas and T. regium). The lower latera are proportionately low, and the apeces

of the inframedian latus projects above the upper line of the rostro- and carinolatera. The apeces

of the carinolatera curve towards but do not overlap with the lower carinal border of the upper

latus.

Mandibles tridentoid except in one large specimen (21 mm capitulum length) in which a

small extra tooth occurs below the upper tooth. Penis absent. The numbers of segments in the

rami of the cirri and caudal appendage increase with size of the individuals (Fig. 4). It is in¬

teresting to compare the numbers and trends with those of T. regium (Fig. 3). Size for size it

is not possible to distinguish the two species on this sort of numerical data.

The material includes a solitary juvenile from CORINDON II Stn 248 at 170 m; it is clearly

scalpelline, and might be A. miche lottianum, but without co-occurring adults, identification is

uncertain. Isolated juvenile stages of any scalpelline are difficult to identify, because of the

similar arrangement of plates in juveniles.

FlG. 4. — Scalpellidae: Arcoscalpellum mi-

chelottianum BALGIM, CP95. Mean

numbers of segments of rami as a func¬

tion of size of specimen. Average of

rami for the numbers of specimens

shown above the abscissa.

Scalpellidae : Arcoscalpellum michelot-

tianum BALGIM . CP95. Nombres

moyens de segments des rames en fonc¬

tion de la taille des spécimens. Nombre

moyen de rames en fonction du nombre

de spécimens examinés indiqué au des¬

sus de l’abscisse.

Arcoscalpellum crenulatum n. sp.

(Fig. 5A-F)

Material. — BIOGAS VIII (Bay of Biscay): CP30, 3100 m (2 specimens: holotype MNHN Ci 2132;

paratype MHNH Ci 2133).

Description of holotype

15.2 mm capitulum length. Capitulum higher than wide. Tergum triangular, apex erect.

Carina simply bowed, flat roofed, with strong longitudinal ridges on each side. Scutum quad¬

rangular, apex acute and curved over the tergum, upper margin concave and with secondary

growth flange towards the tergum, upper latus margin hollowed out below the upper angle for

reception of apex of upper latus. Upper latus pentagonal, with concave margin towards scutum,

bearing a narrow rib along this margin, and lower margin mostly abutting the rostrolatus but

also touching the inframedian latus. Carinolatus pentagonal, carinal margin concave, and occlu-

dent margin projecting slightly beyond the carina below the carina. Inframedian latus triangular,

penetrating in these specimens, to the lowest corner of the upper latus. Rostrolatus rectangular,

with the lateral margin abutting both the inframedian latus and the upper latus, the margin against

the scutum crenulate to the extent of the abutment with the ridge of the upper latus.

Cirri I and II with anterior rami slightly shorter than posterior rami; segments of la 1.5

times as wide as those of posterior ramus. Caudal appendages as long as pedicle of cirrus VI.

Intermediate segments of posterior cirri with 3 pairs of setae on anterior edge; a single spine

on the posterior edge of posterior ramus only.

Segment counts, anterior ramus first:

III

8,9

12,17

17,19

21,17+

23,22

22,26

7,11

14,16

18,19

16+, 16+

21,21

22,25

Remarks

This species closely resembles Amigdoscalpellum aurivilli (Pilsbry) from off Cape Hatteras,

and also Arcoscalpellum pertosum Foster from New Zealand in the way the plates are apposed.

Fig. 5. — Scalpellidae: A-F, Arcoscalpellum crenulatum sp. nov. A, BIOGAS VIII, CP30 holotype (15.2 mm c.l.); B, ditto, paratype,

(13.0 mm c.l.); C, cirrus I; D, cirrus II; E, intermediate segments cirrus VI; F, pedicle cirrus VI and caudal appendage.

Scalpellidae : Arcoscalpellum crenulatum sp. nov. : A, BIOGAS VIII, CP30 holotype (15,2 mm l.c.); B, dito, paratype, (13,0 mm

l.c.) ; C, cirrhe I ; D, cirrhe II ; E, segments intermédiaires du cirrhe VI ; F, partie basale du cirrhe VI et appendice caudal.

— 172 —

However, the crenulate upper margin of the rostrolatus, the penetration of the inframedian latus

all the way to the upper latus, and the relatively short occludent margins of the carinolatera

below the carina, set the CP30 specimens apart. They are also from deeper water than the above

named species.

Amigdoscalpellum praeceps (Hoek, 1907)

(Fig. 6A)

Material. — BIOGAS VIII (Bay of Biscay): CP30, 3100 m (1 specimen). BIOGAS XI (Bay of Biscay):

CP35, 4720 m (1).

Record. — Hoek, 1907a: 114, East Indies, 411 m. Foster & Buckeridge, 1994: 356, La Réunion. 1660-

2970 m.

Remarks

This slender barnacle possesses lightly sculptured plates which show, through ontogeny, a

widening of the carinal tergal interspace and a reduction of the inframedian latus from full oc¬

cupation of the space between the rostra- and carinolatera to about 0.2 times of it and displaced

outwards in large specimens (Foster & Buckeridge, 1994).

Verum? minutum (Hoek, 1883)

(Fig. 6B-F)

Material. — BALGIM: CP68, 2077 m (13 specimens); CP69, 2028 m (3); CP99, 1870 m (3).

Record. — Hoek, 1883: 113, South-East Pacific, 2652 mm; Stubbings, 1936: 28, off Zanzibar, 802 m.

Foster & Buckeridge, 1994: 358, La Réunion, 1850-3520 m.

Fig. 6. — Scalpellidae: A, Amigdoscalpellum praeceps BIOGAS XI, CP35 (27.8 mm c.l.). B-F, Verum minutum'. B, BALGIM,

CP68 (5.8 mm c.l.). C, BALGIM CP69, (3.8 mm c.l.). D, BALGIM, CP99 (2.2 mm c.l.). E, BALGIM, CP68 (4.8 mm c.l.)

cirrus I. F, ditto, intermediate segment posterior ramus cirrus VI. Scales in mm.

Scalpellidae: A, Amigdoscalpellum praeceps BIOGAS XI, CP35 (l.c. = 27,8mm). B-F, Verum minutum.- B, BALGIM. CP68

(l.c. = 5,8 mm). C, BALGIM CP69, (l.c. = 3,8 mm). D, BALGIM, CP99 (l.c. = 2,2 mm). E, BALGIM, CP68 (l.c. = 4,8 mm)

cirrhe I. F, dito, segments intermédiaires de la rame postérieure du cirrhe VI. Échelles en mm.

— 173-

Remarks

Barnacle small, slender, non-hirsute, with smooth, non-striated plates, and faint growth

ridges. Scutum and upper latus elongate, without secondary growth flanges. Carina slightly bowed

in upper part. Tergum elongate, apex not curved. Carinolatus slightly higher than wide. Ros-

trolatus quadrangular, with slight convexity on occludent margin. Inframedian latus very narrow,

slightly wider near the upper extremity, umbo apical, and with very short scutal and upper latus

margins. The apex of the inframedian latus is club-shaped, tipped towards the rostral margin,

and displaced slightly from it in larger specimens (FOSTER & Buckeridge, 1994).

Meroscalpellum bifurcatum (Zevina, 1973)

(Fig. 7A, B)

Material. — BIOGAS XI: CP35 (off Tristan da Cunha), 4720 m (1 specimen). WALDA: CY09 (off Angola),

4613 m (1).

Record. — Zevina, 1973: 1003, Indian Ocean, 4151 m (in Zevina, 1981: 175).

Remarks

This is one of the scalpellines which show in the adult form variously reduced plates and

marked cuticular spaces between these plates. ZEVINA (1981) gives illustrations of 11 mm and

20 mm capitulum length specimens. Not only does the degree of capitular plate coverage decrease

in larger specimens, but the shapes of the plates themselves (except the carina) are increasingly

irregular.

Similar sequences of capitulum variations and changes have been documented for Neoscal-

pellum debile (Aurivillius) by ZEVINA (1981) and for Litoscalpellum intermedium (Hoek) by

Foster (1980). These instances serve as examples of the problems associated with a taxonomy

based on isolated specimens, and possible confusions among generic assignments.

Fig. 7. — Scalpellidae: A, Meroscalpellum bifurcatum WALDA, CY09 (16.2 mm c.l.). B, ditto , BIOGAS XI, CP35 (29.6 mm

c.l.). C-E ,Verum novaezelandiae: C, EPI I, CP39 (7.6 mm c.l.). D, ditto (6.0 mm c.l.). E, ditto (4.0 mm c.l.). Scales in mm.

Scalpellidae: A, Meroscalpellum bifurcatum WALDA, CY09 (l.c . = 16,2 mm). B, dito, BIOGAS XI, CP35 (l.c. = 29,6 mm).

C-E, Verum novaezelandiae; C, EPI I, CP39 (l.c. = 7,6 mm). D, dito (l.c. = 6,0 mm). E, dito (l.c. = 4,0 mm). Échelles en mm.

— 174 —

Verum novaezelandiae (Hoek, 1883)

(Fig. 7C-E)

Material. — BALGIM: CP10, 1592 m (1 specimen). EPI 1: CP38 (Bay of Biscay), 2100 m (14); CP39,

2100 m (7). CORINDON II (Makassar Strait): Stn 236, 1730 m (1).

Records. — Hoek, 1883: 124, New Zealand, 1280 m; Gruvel, 1902: 54, off Portugal, 2400-2500 m; An-

nandale, 1908 &19I3: 231, off Andaman Is, 896-2800 m; Calman, 1918: 123, Gulf of Aden, 2195 m; Weltner,

1922: 71 East-African coast, 1289-1644 m; Foster, 1978: 65 & 1980: 530, New Zealand, 455-2431 m.

Remarks

This species was well represented in collections from New Zealand (Foster, 1978, 1980).

It is clear that this deep water species is widespread. The present records confirm the Atlantic

Ocean records of Gruvel (1902).

V. novaezelandiae is recognised by the very slender capitulum in which the carinolatus is

markedly elongated to match a pronounced apical growth of the inframedian latus. This latter

plate remains narrow, with clear edges to the upper latus and scutum. With growth, the umbo

of the inframedian latus is placed closer to the basal margin. In some specimens the rostrolatus

is very narrow, particularly if growth of the peduncle has been bent to the rostral side.

Appendage counts for CP39 specimens are as follows:

cl(mm)

III

4.8

6.0

7.6

—

The appendage details conform to New Zealand specimens (Foster, 1978).

Verum frillosum nov. sp.

(Fig. 8)

Material. — BIOGAS VII: CP28 (Bay of Biscay), 3380 m (1 specimen, holotype MNHN Ci 2135).

Description of holotype

18.0 mm capitulum length. Peduncle only partially intact, scales wide. Tergum triangular,

apex recurved, dorsal margin notched for reception of carina. Carina simply bowed. Scutum

pentagonal, upper margin with secondary growth flange against tergum, lateral margin slightly

concave, lower margin sinuous in conjunction with rostro- and inframedian latus. Inframedian

latus hourglass-shaped, upper part wider than lower part, with sinuous margins to scutum and

upper latus. Upper latus pentagonal, upper margin with secondary growth flange towards apex,

margins to inframedian- and carinolatus sinuous. Carinolatus higher than wide, upper margin

sinuous, occludent margin beneath carina short. Rostrolatus broadly quadrangular.

Mandible tridentoid, lower angle acuminate. Maxillule with 2 stout spines at upper angle,

bunch of shorter spines at lower angle. Cirrus I with anterior ramus intermediate segments 2 times

— 175 —

width of those of posterior ramus. Mean numbers of segments in cirri as follows, anterior ramus

first:

III

8,12

16,19

20,20

24,24

25,25

Intermediate segments cirrus VI with 5 pairs of setae on anterior edge, no setae midway

on posterior edge. Caudal appendages 2-3 segments, just longer than pedicle of cirrus VI.

Remarks

This barnacle is similar to Verum constriction but lacks constricted rostrolatera, and the

growing edges of the upper latus, rostrolatus, carinolatus and inframedian latus are sinuous in

accord with the alternating and interlocking growth ridges of these plates.

Fig. 8. — Scalpellidae: Verum frillosum sp. nov. BIOGAS VII, CP28: A, capitulum holotype (18.0 mm cl). B, mandible. C,

maxillule. D, cirrus I. E, intermediate segments anterior and posterior rami cirrus VI. F, pedicle cirrus VI and caudal appendage.

Scales in mm.

Scalpellidae: Verum frillosum sp. nov. BIOGAS VII, CP28: A, capitulum holotype (Cl = 18.0 mm). B, mandible. C, maxillule.

D, cirrhe I. E, segments intermédiaires des rames antérieure et postérieure du cirrhe VI. F, partie basale du cirrhe VI et

appendice caudal. Échelles en mm.

Ornatoscalpellum stroemii (M. Sars, 1859)

(Fig. 9A)

Material. — BALGIM: DR82, 355 m (3 specimens).

Records. — Hoek, 1883: 73, northern Atlantic Ocean, 152-944 m; Pilsbry, 1907a: 21, North American

fishing banks, 143-214 m; and other references in Zevina, 1981: 111. As pressum Pilsbry, 1907a: 23, Le Havre

Bank, 549 m. As septentrionale Aurivillius, 1892: 128 and 1894: 51, North Sea, 590-890 m. As erosum Aurivillius,

1892: 119 and 1894: 52, North Atlantic, 1744 m. As obesum Aurivillius, 1892: 119 and 1894: 56, North Sea,

110 m. As luridum Aurivillius, 1892: 120 and 1894: 59, Baffin Bay, 309 m.

Remarks

The synonymy of 4 of Aurivillius’ (1892, 1894) and 3 of Pilsbry's (1907a) names within

O. stroemii (see Zevina, 1981: 111) illustrates the variation within at least some of the scalpelline

— 176-

species, and the tendency to name this variation by early taxonomists. Pilsbry (1907a: 21) was

troubled about Hoek’s (1883) illustration of the species, particularly the latter giving the im¬

pression that the scutum, inframedian latus and upper latus have subcentral umbos, and also

with the proportions of the upper latus. There may have been some over-artistry in HOEK’s

plates (not confined to this species), but the fact is that during ontogeny the position of the

umbo of the inframedian latus becomes progressively displaced from the apex, and both the

upper latus and carina also develop apical growth zones. The rectangularly bent carina, in its

upper part, results from displacement of the umbo, as it is in Scalpellum scalpellum.

This is a relatively shallow water scalpelline, often collected in fishing trawls. Stn DR82

is off the Morocco coast, and is a southernmost record of the species.

Scalpellum scalpellum (Linnaeus, 1767)

(Fig. 9B-D)

Material. — BALGIM: CP103, 347 m (21 specimens).

Records. — North-East Atlantic, Norway (68°N) to Congo (6°S), 50-366 m; Mediterranean (Reuni, 1980:

93, Italian seas, to 400 m). See Zevina, 1981: 95 for synonymies and references.

Remarks

Of all the 18 species considered in this collection, this is the only one in which the umbo

of the scutum is displaced from the apical position. The relatively square lateral shape of the

capitulum is produced by considerable subapical displacement of the umbos of the carina and

upper latus.

This is a relatively shallow occurring barnacle. It is frequently encountered on hydroids

and hydroid-like growths on trawling grounds, and is a well-known barnacle to European marine

naturalists. This is the only known species of the Scalpellinae from the Mediterranean Sea

(Relini, 1980).

Fig. 9. — Scalpellinae: A, Omatoscalpellum stroemii BALGIM, DR82, group, largest 5.5 mm c.l. B, Scalpellum scalpellum

BALGIM, CP103 (13.5 mm c.l.). C, ditto (6.2 mm c.l.). D, ditto (3.0 mm c.l.). Scales in mm.

Scalpellinae : A, Omatoscalpellum stroemii BALGIM, DR82, plus grand individu du groupe l.c. = 5,5 mm. B, Scalpellum

scalpellum BALGIM, CP103 (l.c. = 13,5 mm). C, dito (l.c. = 6,2 mm). D, dito (l.c. - 3,0 mm). Échelles en mm.

— 177 —

Family VERRUCIDAE Darwin, 1854

Discussion

There are 83 named species of verrucid (Buckeridge, 1994, Foster & Buckeridge, 1994).

They are asymmetrical sessile barnacles having a shell wall composed of carina, rostrum, fixed

scutum and fixed tergum, and an operculum of movable tergum and movable scutum. Nomen¬

clature of these plates is shown in Fig. 10A. A fuller discussion of the morphology and genera

is available in Buckeridge (1994).

The present material is particularly rich in verrucids, both in the number of determined

species (4) and individuals (420). Furthermore, the BALGIM collections are in the locality from

which Aurivillius (1898) and Gruvel (1900b, 1902, 1905, 1912) named and/or described many

species.

Verruca trisulcata Gruvel, 1900

(Figs 10, 11, 12)

Material. — BALGIM: CP90, 890 m (9 specimens); CP91. 948 m (4); CP92, 1182 m (70); CP95, 1378 m

(22); CP156, 1135 m (252); DW157, 1108 m (28); CP160, 1350 m (9).

Records. — Gruvel, 1900b: 243; 1902: 96; 1920: 44, Azores, 998 m. Gruvel, 1912: 348, Cape Spartel

(Straits of Gibraltar), 622 m. As imbricata Gruvel, 1900b: 244, 1902: 105, Azores, 441 m. As striata Gruvel,

1900b: 244; 1902: 98, Cape Verde I, 633 m. As radiata Gruvel, 1900b: 262, 1902: 94, Canaries, 912 m; Foster

& Buckeridge, 1994: 363, La Réunion, 980-1690 m.

Remarks

A range of shell forms is illustrated to show the range from low splayed (half as high as

wide) to quite upstanding (higher than wide) posture. Of 26 shells from CP92 (Fig. 11) the

height: width ratio ranges from 0.5 to 1.0, with no significant trend with size of specimen. Also,

with growth, the number of interlocking ribs between the movable tergum and scutum, and be¬

tween the carina and rostrum, increases from 1 in juveniles to 4 in specimens of about 6 mm

rostrocarinal length. A characteristic feature of the shell is the particularly wide superior rib on

the rostrum. The apeces of the rostrum and carina are never turned inwards. The “back” view

(i.e. of the fixed tergum and scutum) is highly variable; tall with easily discernable alae and

radial growth areas, or squat and contorted depending on irregularities of substratum. Movable

scutum apex curved over the superior articular groove, with a moderately deep apical pit inter¬

nally. Movable tergum broadly quadrangular, 1.5 times width of scutum, lower angle protruding

as apical ridge.

Mandible tridentoid, with a pectinate lower edge. Maxillule with an irregular cutting

edge, the lower angle protuberant (a feature common in verrucids). Cirrus I anterior ramus

just over 0.5 times length of posterior ramus. Cirrus II anterior ramus less than 0.5 times

length of posterior ramus. Cirrus III-VI with subequal rami. Caudal appendage about 4).66

times length of cirrus VI. Plots of the numbers of segments per rami (Figure 12) show in¬

creasing numbers of segments in all rami with increasing size of the specimen. Caudal ap¬

pendage 0.5 times length of cirrus VI.

— 178 —

Fig. 10. — Verrucidae: Verruca trisulcata. A, BALGIM, CP 160 (5.5 mm rcl). B, C, opercula, inner views, specimen A. D, E,

opercula, outer views, BALGIM, CP 160. F, G, “front” and “rear” views, BALGIM CP92 (5.0 mm rcl). H, I, “rear” views

of others BALGIM CP92; same scale. J, juvenile, BALGIM, CP 160 (0.8 mm rfcl). K, post-juvenile, BALGIM CP90 (1.8 mm

rcl). L, M, opercula, outer views BALGIM, CP90. N, cirrus I, BALGIM, CP90 (from 5.9 mm c.l.). O, cirrus II, ditto. P,

cirrus III, ditto. Q, intermediate segment cirrus VI, ditto. R, pedicle cirrus VI, penis and caudal appendage, ditto. N-R to

same scale. Scales in mm. Shell plates: c, carina; fs, fixed scutum; ft, fixed tergum; ms, movable scutum; mt, movable

tergum; r, rostrum, rcl = umbos of rostrum to carina length.

Verrucidae : Verruca trisulcata. A, BALGIM, CP 160 (Ire = 5,5 mm). B, C, opercule, vues internes, spécimen A. D, E, opercules,

vues externes, BALGIM, CP 160. F, G, vues antérieure et postérieure, BALGIM CP92 (Ire = 5,0 mm). H, I, vues postérieures

des autres spécimens BALGIM CP92 ; même échelle. J, juvénile, BALGIM, CP 160 (Ire = 0,8 mm). K, post-juvénile, BALGIM

CP90 (Ire = 1,8 mm). L, M, opercule, vues externes BALGIM, CP90. N, cirrhe I, BALGIM, CP90 (Ire = 5,9 mm). O, cirrhe

II, dito. P, cirrhe Ill, dito. Q, segment intermédiaire du cirrhe VI, dito. R, partie basale du cirrhe VI, pénis et appendice

caudal, dito. N-R dessinés à la même échelle. Échelles en mm. Plaques de la coquille : c, carène ; fs, scutum fixe ; ft, tergum

fixe; ms, scutum mobile; mt, tergum mobile ; r, rostre. Ire = distance séparant l’umbo du rostre de la carina.

The type locality for V. striata is also in the Atlantic Ocean. It seems unavoidable that

trisulcata , imbricata, striata and radiata are synonymous. Because the number of articular ribs

on the movable tergum is not constant, nor is the overall shell shape, it may be that V. grimaldi

number of segments

— 179 —

rostro-carinal length (mm)

Fig. 11. — Verrucidae: Verruca trisulcata, BALGIM, CP92: ratio of maximum shell height to maximum shell width plotted as

a function of rostrocarinal length. A ratio of 1.0 is a shell as high as wide; 0.5, half as high as wide. Regression: y = 0.49

+ 5.14x, n = 25, R = 0.028, no significant slope.

Verrucidae : Verruca trisulcata, BALGIM, CP92 : rapport de la hauteur maximale de la coquille à sa largeur maximale en

fonction de la longueur rostrocarinale. Un rapport de 1,0 correspond à une coquille aussi haute que large ; de 0,5, moitié

moins haute que large. Régression : y = 0,49 + 5,I4x, n = 25, R = 0,028, pas de pente significative.

rostro-carinal length (mm)

FiG. 12. — Verrucidae: Verruca trisulcata ,

BALGIM, CP92: Mean numbers of

segments of rami as a function of size

of specimen. For anterior (a) rami of

cirri I-III and posterior (p) rami of cirri

I and II, and all rami of cirri IV-VI,

and caudal appendages (ca).

Verrucidae: Verruca trisulcata, BAL¬

GIM, CP92: Nombres moyens de seg¬

ments des rames en fonction de la

taille des spécimens. Rame antérieure

(a) des cirrhes /-///, postérieure (p)

des cirrhes I et II, ensemble des rames

des cirrhes IV-VI, et appendices cau¬

daux (ca).

— 180 —

Gruvel. 1920 from off Madeira Is, V. entobapta Pilsbry, 1916 from off Florida, and V. macani

Stubbings, 1936 from off Zanzibar could be the same species.

Altiverruca gibbosa Hoek, 1883

(Fig. 13A, B)

Material. — BALGIM: CP108, 1527 m (1 empty shell).

Records. — Hoek, 1883: 134, Crozet Is, 1893 m; Nilsson-Cantell, 1928: 25, Celebes, 3056 m (see for

synonymy sulcata , bicornuta , mitra)\ Broch, 1931: 45, East Indies, 385 m; Zevina, 1971: 439, off Chile, 1230-

1700 m; Newman & Ross, 1971: 135, South Georgia, 1098-1153 m; Foster, 1981: 352, Kermadec Is and Norfolk

I, 590-949 m. As gibbosa somaliensis Nilsson-Cantell, 1929: 470, East-African coast, 1079-1289 m. As bicornuta

Pilsbry 1916: 43, off Nantucket, 3127 m. As sulcata Hoek, 1883: 139, off New Zealand, 950-1165 m. As mitra

Hoek, 1907b, Antarctic Peninsula, 555 m. As darwini Pilsbry, 1907a: 111, off Cape May, 2844 m. As rathburniana

Pilsbry, 1916: 41, off Cape Cod, 3235 m; Nilsson-Cantell, 1927: 776, Cape Verde Is, 1812 m.

Remarks

The single empty shell is a dirty yellow colour with clearly delimited growth lines. Basal

maximum length (rostrocarinal) 4.5 mm; apical rostrocarinal length (rcl) 5.2 mm; i.e. apex of

carina flared outward. Tergal height 5.0 mm. The operculum orifice is at an angle of 45 degrees

to the base.

A. gibbosa is distinguished from V. trisulcata by the ridges and shelf above the superior

articular ridge of the rostrum, and from V. sinuosa (above) by the very definite sculpturing of

the whole shell. A single specimen does not permit any examination of existing or further syn¬

onymy, but it does show how a widely distributed, recorded and considered barnacle can be

sparsely represented in intensive collecting.

Fig. 13. — Verrucidae: A, B, Altiverruca gibbosa BALGIM,

CP 108, “front” and “rear” views of empty shell. Scale in mm.

Verrucidae: A, B, Altiverruca gibbosa BALGIM, CP 108, vues

frontale et postérieure d'une coquille vide. Échelle en mm.

Altiverruca vertica n.sp.

(Fig. 14)

Material. — BALGIM: CP92, 1182 m (1 specimen: holotype MNHN Ci 2138).

Description of holotype

3.2 mm umbo rostrocarinal length (rcl): shell higher than wide, with operculum held nearly

perpendicular to base. Base calcareous. Carina higher than rostrum; upper margins to movable

— 181 —

opercula continuous and shallowly concave. Rostrum broadly quadrangular; suture with carina

without interlocking. All plates lightly sculptured with growth lines. Movable tergum square,

and with 2 broad articular ribs above a barely defined diagonal rib. Movable scutum 0.5 times

width of tergum, with diagonal rib placed close to articular edge, and internally with a deep

muscle pit apically.

Cirrus I with anterior ramus 1 segment longer than posterior ramus. Cirri II and III with

anterior rami shorter than posterior rami. Cirri IV-V damaged. Cirrus VI with 25 segments in

subequal rami, and intermediate segments with 2 pairs of setae on the anterior edge. Mean seg¬

ments per rami as follows, anterior ramus first:

I II III IV V VI ca

10,9 7,10 13,16 20,22 - 25,25 7

Caudal appendages 0.5 times length of basal podomere of pedicle of cirrus VI.

FlG. 14. — Verrucidae: Altiverruca vertica sp. nov. Holotype BALGIM, CP92: A, holotype (3.2 mm rcl). B, C, tergum and scutum,

inner views. D, cirrus I. E, cirrus II. F, cirrus III. G, pedicles of cirri VI and caudal appendages. H, intermediate segment

of cirrus VI. Scales in mm.

Verrucidae : Altiverruca vertica sp. nov. Holotype BALGIM, CP92 : A, holotype (Ire = 3,2 mm). B, C, tergum et scutum, vues

internes. D, cirrhe I. E, cirrhe II. F, cirrhe III. G, partie basale des cirrhes VI et appendices caudaux. H, segment intermédiaire

du cirrhe VI. Échelles en mm.

Remarks

This species is represented by a solitary specimen. It is similar to quadrangularis, being

lightly sculptured, smooth surfaced, the operculum held at a steep angle to the base, and the

— 182-

rostrum and carinal in “front” view broadly quadrangular, but there is no interlocking between

rostrum and carina; further, the scutum is much narrower. The specimen is larger than the largest

V. quadrangularis, and it is notable that the caudal appendages are more diminutive. The shell

resembles V. casula Hoek, 1913, of Malaysian seas; the appendages of V. casula have not been

described, and V. casula is notable for the broad rostral rib interlocking with the carina.

Metaverruca recta (Aurivillius, 1898)

(Fig. 15)

Material. — BALGIM: CP92, 1182 m (10 specimens); CP95, 1378 m (9); CP97, 1515 m (2); CP109,

1200 m (1); DW157, 1108 m (1). EPI I: CP38 (Celtic Sea), 2100 m (1).

Records. — Aurivillius, 1898: 195 and Gruvel, 1920: 46, Azores, 861-1385 m; Southward & Southward,

1958: 637. Bay of Biscay, 329-1774 m. As sculpta Aurivillius, 1898: 197 and Gruvel, 1920: 41, Azores, 454 m;

Nilsson-Cantell, 1929: 461, Scotland, 1326 m; Broch, 1931: 41, Kei Is, 345 m; Foster, 1981: 352, Kermadec

Is, 501-1180 m. As magna Gruvel, 1901: 261; 1902: 109, Gulf of Gascogne, 1480 m; Gruvel, 1920: 50. As

linearis Gruvel, 1900b: 243 and Gruvel, 1902: 107, Azores, 960-2018 m. As halotheca Pilsbry, 1907b: 188,

Hawaii, 1670 m; Pilsbry, 1916: 46, Hawaii, 417-430 m; Foster, 1978: 69, New Zealand, 252-896 m. As capsula

Hoek, 1913: 130, East Indies, 520-1301 m; Stubbings, 1936: 38, Zanzibar, 333 m. As coraliophila Pilsbry, 1916:

21, Florida — Bahamas, 506-794 m. As sculpta Buckeridge, 1983: 59; Miocene of New Zealand; Buckeridge,

1994: 116 (for full synoymy); Foster & Buckeridge, 1994: 368, La Réunion, 620-1230 m.

Fig. 15. — Verrucidae: Metaverruca recta. A, top view (4.5 mm rcl) BALGIM, CP95. B, oblique view (5.1 mm rcl) BALGIM,

CP92. C, D, outer views scutum and tergum (of 5.1 mm rcl specimen) BALGIM, CP92. E, F, inner views, ditto. G, cirrus

I; H, cirrus II; I, pedicle cirrus VI, penis and caudal appendages; J, intermediate segments cirrus VI; all of 5.0 mm rcl

specimen, BALGIM, CP95, and all to same scale. Scales in mm.

Verrucidae : Metaverruca recta. A, vue de dessus (Ire = 4,5 mm) BALGIM. CP95. B. vue oblique (Rcl - 5,1 mm) BALGIM, CP92.

C, D, vues externes du scutum et du tergum (spécimen ayant une Ire de 5,1 mm) BALGIM, CP92. E, F, vues internes, dito. G,

cirrhe I; H, cirrhe II; /, partie basale du cirrhe VI, pénis et appendices caudaux; J, segments intermédiaires du cirrhe VI;

provenant tous d’un spécimen ayant une Ire de 5,0mm, BALGIM, CP95, et tous à la même échelle. Échelles en mm.

— 183 —

Remarks

Accepting the synonymies, this is a widespread barnacle, from North Atlantic through Indian

to North and South Pacific Oceans. It is characterised by its particularly compact form, more

roundly sessile than narrowly erect like V. quadrangularis, with the apeces of the rostrum and

tergum set in from the basal margin (i.e. the shell is conical rather than splayed), with the upper

ribs between the rostrum and carina very wide, mostly 2, sometimes 3 of them; by the edge of

articulation of the movable plates forming a straight line between the apeces of the rostrum and

carina; by the general D-shape of the orifice; by the operculum being held at an angle 45-90

degrees to the perpendicular depending on the nature of the substratum; by the myophores; and

by the generally large size of grown specimens for the genus (8-10 mm basal length). The ap¬

pendages have been described by FOSTER (1978) and BUCKERIDGE (1994); notable is the shortness

of the caudal appendages.

Mean numbers of segments in cirri of 4 specimens (CP95) as follows, anterior ramus first:

rcl (mm)

III

1.0

6,5

5,6

8,10

15,17

16,14

14,17

1.3

7,6

5,7

9,12

15,16

19,19

21,21

2.9

12,9

8,11

18,21

22,26

26,28

30,31

4.3

12,10

8,12

19,20

20,21

23,22

20,21

Family PACHYLASMATIDAE Buckeridge, 1983

Pachylasma giganteum (Philippi, 1836)

(Fig. 16A, B)

Material. — BALGIM: DR37, 864 m (3 shells); DR40, 362 m (75 specimens, 14-34 mm basal diameter);

DR49, 521 m (1 specimen); DR111, 285 m (6 specimens); DRII5, 352 m (1 shell); DW116, 340 m, (1 shell);

DR152, 550 m (1 shell); DR153, 580 m (99 specimens, 6-47 mm basal diameter).

Records. — Darwin, 1854: 477, Mediterranean; Pilsbry, 1916: 329, Strait of Messina; Stubbings, 1967:

263, off West Africa; Weltner, 1897: 273, Mediterranean, on Errina aspera\ Reuni, 1980: 49, Italy, 150-200 m.

See Newman & Ross (1976: 40) for full synonymy; distribution quoted as “Mediterranean - West Coast Africa”.

Remarks

This barnacle is well described by Darwin (1854), who quoted a maximum size of 29.2 mm

basal rostrocarinal length. Very small specimens have no carinolatera. Few of the records of this

barnacle give depth data. The present records are precise, 285-864 m, just west of the Straits

of Gibraltar. Four of the stations yielded only shells (easily recognised by their thickness and

the irregular basal edges) and 2 of the stations yielded rock substratum with large numbers of

specimens growing on it and themselves.

— 184-

Family BATHYLASMATIDAE Newman & Ross, 1971

Bathylasma hirsutum (Hoek, 1883)

(Fig. 16C, D)

Material. — BALGIM: CP36, 990 m (1 shell only); DR37, 864 m (72 shells).

Records. —• See Newman & Ross (1976: 46) for full references, and distribution “Northeast Atlantic from

Faroe Is to Azores, 944-1829 m”. Southward & Southward (1958) give a useful list of collecting data, including

depth range of 384-1829 m.

Remarks

These 2 stations are a little further south than the stations listed by Southward & South¬

ward (1958) but do not constitute a significant addition to the known distribution. The large

number of shell plates at Stn DR37 (Bay of Cadiz) is quite reminiscent of similar collections

of plates of B. corolliforme from the Southern Ocean (Newman & Ross, 1971) and of B. alearum

from New Zealand (Foster, 1978). Evidently, the shell plates of deep water bathylasmatids

accumulate in-situ in proximity to appropriate attachment substrata, and in themselves provide

substrata for further recruitment and accumulation.

Family BALANIDAE Leach. 1817

Megabalanus tulipiformis (Ellis, 1758)

(Fig. 16E, F)

Material. — BALGIM: DR40, 362 m (4 shells); DR115, 352 m (1 shell plate); DR133, 195 m (1 specimen).

Distribution. — Mediterranean, east Atlantic of France to Cape Verde Is, 25-250 m (Newman & Ross,

1976: 69; Relini, 1980: 53).

Fig. 16. — Balanomorpha, inner views of opercula: A, B, Pachylasma giganteum, scutum and tergum, same individual, BALGIM,

DR40. C, D, Bathylasma hirsutum, ditto, different individuals, BALGIM, DR37. E, F, Megabalanus tulipiformis, ditto, same

individual, BALGIM, DR 133. Scales in mm.

Balanomorpha, opercule, vues internes : A, B, Pachylasma giganteum, scutum et tergum d’un même individu, BALGIM, DR40.

C, D, Bathylasma hirsutum, dito, provenant de divers individus, BALGIM, DR37. E, F, Megabalanus tulipiformis, dito, d’un

même individu, BALGIM, DR 133. Échelles en mm.

— 185 —

Remarks

Previous authors have noted the occurrence of M. tulipiformis with Pachylasma giganteum.

The single shell plate recovered from station DR 115 together with P. giganteum was readily

identified by its rose pink colour. In the BALGIM collections, M. tulipiformis is the only barnacle

recovered from any station in the Mediterranean (DR 133), and it was the only live material of

this species.

BIOGEOGRAPHY

Summary data are presented in Table 1, which gives the number of stations, number of

individuals and depth data from these collections and from the literature, for each of the species.

Full station data are given in Appendix A.

The frequency of recovery of barnacles in the deep sea grabs and dredges is 23 % (29 of

126 stations) for the BALGIM cruise, yielding 882 specimens (14 species).

The North Atlantic Ocean stations (29 BALGIM plus 9 others) provided 20 species, in¬

cluding 2 new to science ( Arcoscalpellum crenulatum and Verum frillosum).

There are 5 species common to New Zealand and the North Atlantic ( Poecilasma kaempferi,

Smilium acutum, Verum novaezelandiae, Altiverruca gibbosa and Metaverruca recta). These seem

to be part of a circumglobal, or at least North Atlantic to South-West Pacific barnacle fauna,

in moderately deep water (1000-2000 m).

Whether deeper sea (> 3000 m) species such as Trianguloscalpellum gigas, T. regium, Arcoscal¬

pellum michelottianum and Meroscalpellum bifurcatum are widespread is hard to ascertain from pub¬

lished records, but considering the more limited sampling from such depths they are likely to be so.

The more commonly collected species (i.e., 100 specimens) are: A. michelottianum, taken

at 11 stations (121 specimens), Verruca trisulcata, at 10 stations (400 specimens), and Pachylasma

giganteum, at 4 stations (181 specimens). The last of these, the shallowest occurring, is most

likely restricted to the northern Atlantic Ocean and the Mediterranean Sea. The predominance

of records from the North Atlantic, possibly reflects a longer history of collecting there. Bio¬

geographic restrictions are likely to apply to other shallower sea species, ( e.g., for the North

Atlantic Scalpellum scalpellum, Omatoscalpellum stroemii, Bathylasma hirsutum and Mega-

balanus tulipiformis), as they are to littoral species.

With respect to a principal aim of the BALGIM cruise, namely to relate Mediterranean

outflow water to its deflection to the northern side of the Ibero-Moroccan Bay, it is noteworthy

that stations that have Bathylasma hirsutum, Pachylasma giganteum or Megabalanus tulipiformis,

do not have verrucomorphs or lepadomorphs. Only one of these stations is in the Mediterranean

Sea (DR133, which has M. tulipiformis only, and is the only station there to provide barnacles),

two are on the sill of the Straits of Gibraltar (DR 152 and DR 153, with P. giganteum only), five

are grouped just due west of the sill (DR40, DR49, DR 111, DR115 and DW116, with both

M. tulipiformis and P. giganteum), and two further northwest (CP36 and DR37, both with P. gi¬

ganteum and B. hirsutum). The Mediterranean barnacle fauna does not have the deep ocean scal-

pellids and verrucids. The last 9 quoted stations are different from the other Atlantic stations,

and could represent a Mediterranean influence.

— 186 -

Table 1. — Summary data on numbers of stations, individuals and depths for each of the species.

Species

No of

Indiv. sample

Total no

Depth range (m)

stations

numbers

this paper

literature

Octolasmis niestraszi

16-135

Poecilasma kaempferi

1108

126-1885

Smilium acutum

1 3 23 45 1 1 3

355-2170

61-2480

Trianguloscalpellum regium

22 1

3100-3380

845-5212

Trianguloscalpellum gigas

3815

3749

Arcoscalpellum michelottianum

14 3 27 5 10 2 15

25 9 10 1

121

1182-2160

64-3422

Arcoscalpellum crenulatum n. sp.

3100

—

Amigdoscalpellum praeceps

1 1

3100-4720

411-2940

Verum ? minutum

13 3 3

1870-2077

802-3520

Meroscalpellum bifurcation

1 1

46613-4720

4151

Verum novaezelaniae

1 14 7 1

1592-2100

455-2500

Verum frillosum n. sp.

1 3

3380

—

Ornatoscalpellum stroemii

355

160-629

Scalpellum scalpellum

347

50-400

Verruca tricsulcata

9 4 70 252 28 9 22

394

890-1378

441-1690

Altiverruca gibbosa

1527

555-1289

Altiverruca vertica n. sp.

1182

—

Metaverruca recta

10 9 2 1 1 1

1108-2100

329-2110

Pachylasma giganteum

(3) 75 1 6 (1) (3) 75

1 6 (1) (1) (1) 99

187

285-864

150-250

Bathvslama hirsutum

(1) (72)

864-990

944-1829

Megabalanus tulipiformis

(4) (1) 1

135-195

Acknowledgements

The authors thank Michel Segonzac of Centre national de tri d’océanographie biologique, Brest, for

the invitation (to BAF) to study the material, and for sending them. Alain Crosnier of the Institut français

de recherche scientifique pour le développement en coopération (ORSTOM), Paris, provided extensive

support and advice on manuscript preparation, revision and publication.

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— 189-

Appendix A. — Cruise and station data, and barnacles and numbers

of barnacles collected at each station.

ATLANTIC OCEAN

BALGIM. — A study to correlate composition of benthic fauna and origin of water masses

between the Atlantic Ocean and Mediterranean Sea. This material was collected May-June 1984

aboard R.V. “Cryos” during cruise BALGIM, conducted for Centre national de la recherche scien¬

tifique (PIROCEAN). under the direction of Dr P. Bouchet (Museum). Dates: 29.5.84-18.6.84.

Stn

°N

°W

Species

CP10

1592

36.453

9.320

Arcoscalpellum michelottianum

Verum novaezelandiae

CP 17

1470

36.453

9.308

Arcoscalpellum michelottianum

CP36

990

36.166

7.137

Bathylasma hirsutum

(1)

DR37

864

36.178

7.154

Pachylasma giganteum

(3)

Bathylasma hirsutum

(72)

DR40

362

35.499

6.086

Pachylasma giganteum

Megabalanus tulipiformis

(4)

DR49

521

35.530

6.328

Pachylasma giganteum

CP63

1510

35.306

7.421

Arcoscalpellum michelottianum

CP65

1805

35.265

7.599

Smilium acutum

Arcoscalpellum michelottianum

CP68

2035

35.119

7.526

Smilium acutum

Arcoscalpellum michelottianum

Verum ? minutum

CP69

2028

35.114

7.505

Smilium acutum

Arcoscalpellum michelottianum

Verum ? minutum

DR82

355

33.455

8.320

Smilium acutum

Ornatoscalpellum stroemii

CP90

890

34.214

7.236

Verruca trisulcata

CP91

948

34.223

7.251

Verruca trisulcata

CP92

1182

34.243

7.303

Arcoscalpellum michelottianum

Verruca trisulcata

Altiverruca vertica

Metaverruca recta

CP95

1378

34.240

7.393

Arcoscalpellum michelottianum

Verruca trisulcata

Metaverruca recta

— 190 —

CP97

1515

34.254

7.411

Arcoscalpellum michelottianum

Metaverruca recta

CP99

1870

34.282

7.433

Arcoscalpellum michelottianum

Verum ? minutum

CP 103

347

34.107

7.298

Scalpellum scalpellum

CP108

1527

36.108

8.062

Altiverruca gibbosa

(1)

CP 109

1200

36.145

7.564

Metaverruca recta

DR111

285

35.569

6.221

Pachylasma giganteum

DR115

332

35.475

6.042

Pachylasma giganteum

Megabalanus tulipiformis

(1)

DW116

340

35.486

6.042

Pachylasma giganteum

(1)

DR133

195

35.258

4.174

Megabalanus tulipiformis

(1)

DR 152

550

35.567

5.347

Pachylasma giganteum

(1)

DR 153

580

35.558

5.353

Pachylasma giganteum

CP 156

1135

36.200

7.527

Poecilasma kaempferi

Verruca trisulcata

252

DW157

1108

36.210

7.558

Poecilasma kaempferi

Verruca trisulcata

Metaverruca recta

CP 160

1350

36.146

8.009

Verruca trisulcata

Bay of Biscay

. — This material

was collected during missions organised by IFREMER (Brest)

between 1974 and 1987.

BIOGAS V

CP07

2170

44.098

4.164

Smilium acutum

BIOGAS VII

CP28

3380

47.295

9.035

Trianguloscalpel 1 urn regium

Scalpellum frillosum

BIOGAS VIII

CP30

3100

47.296

9.061

Verum regium

Arcoscalpellum crenulatum

Amigdoscalpellum praeceps

BIOGAS XI

CP35

4720

46.343

10.230

Arcoscalpellum praeceps

Meroscalpellum bifurcation

EPI I

CP38

2100

47.338

8.422

Smilium acutum

Verum novaezelandiae

Metaverruca recta

CP39

2100

47.320

8.384

Verum novaezelandiae

Smilium acutum

— 191 -

EPI VI

KG229 2160 47.270 8.320 Arcoscalpellum michelottianum 1

GASCOR. — This material was collected during mission GASCOR, from deep-sea modules

placed by EPI (Environnement Profond: Impact), organised by IFREMER. Brest; chief, Gerard

AUFFREt. Date: 16.3.86.

CP42 3815 46.227 12.368 Trianguloscalpellum gigas 3

BIOCYAN. — This material was collected during mission BIOCYAN 2, with submersible

CYANA, organised by Centre océanologique de Bretagne (COB, Brest); chief, Myriam Sibuet.

Date: 26.6.83.

PLI8 2000 47.321 8.276 Smilium acutum 3

WALDA. — This material was collected during mission WALDA, organised by Centre océa¬

nologique de Bretagne (COB, Brest). Collected off Angola. Date: 23.6.71.

°S °E

CY09 4613 19.103 9.181 Meroscalpellum bifurcation 1

SOUTH-WEST PACIFIC

CORINDON II. — This material was collected during the cruise CORINDON II, organised by

ORSTOM: Responsible for biological data. Jacques Forest (Museum). Dates: 29.10.80-12.11.80.

Straits of Makassar, 15-2 400 m.

236

1730

0.067N

119.455

Verum novaezelandiae

248

170

0.542S

119.287

scalpellid juvenile indet.

263

1.568S

119.167

Octolasmis nierstraszi

Bull . Mus . natl . Hist , nat ., Paris, 4 e sér., 17, 1995

n° 1-2 : 193-200.

A new species of Didymochelia from New Caledonia

(Crustacea: Amphipoda: Didymocheliidae)

by Jim K. LOWRY & Helen E. Stoddart

Abstract. — Didymocheliid amphipods are extremely rare. Didymochelia ledoyeri sp. nov. is described

from New Caledonian material. It is only the fourth specimen collected and the third species described. This is

the first record of the family outside the subantarctic area.

Keywords. — Amphipoda, Didymocheliidae, taxonomy, new species, New Caledonia.

Une espèce nouvelle de Didymochelia de Nouvelle-Calédonie

(Crustacea : Amphipoda : Didymocheliidae)

Résumé. — Les amphipodes didymocheliides sont extrêmement rares. Didymochelia ledoyeri sp. nov. de

Nouvelle-Calédonie est seulement le quatrième spécimen capturé et la troisième espèce décrite. La famille est

signalée pour la première fois en dehors de la zone subantarctique.

Mots-clés. — Amphipoda, Didymocheliidae, taxonomie, espèces nouvelles, Nouvelle-Calédonie.

J. K. Lowry & H. E. Stoddart, Australian Museum, P. O. Box A285, Sydney South, NSW 2000, Australia.

INTRODUCTION

The didymocheliids are among the rarest amphipods known. K. H. Barnard (1931) first

described Didymochelia spongicola from one specimen living among sponges at South Georgia.

He was not able to place it in a family and the species has not been collected since. Bellan-

Santini & Ledoyer (1987) described the second species, D. edwardi , based on two specimens

from Prince Edward Island. They established a new family, Didymocheliidae, and compared it

to the Lysianassidae and the Acanthonotozomatidae. They considered the family to have closest

affinities with the Lysianassidae.

A fourth specimen of Didymochelia has been discovered among recent MUSORSTOM

collections from New Caledonia and is described here as Didymochelia ledoyeri sp. nov. The

unique specimen is lodged in the Muséum national d’Histoire naturelle, Paris (MNHN).

In this paper we change the terminology we have previously used to describe setae and

spines. This change is based primarily on arguments about the homology of setae and spines

presented by Oshel & Steele, 1988, and Watling, 1989. The terminology mainly follows

Watling, 1989, with a few modifications. What we have previously referred to as setae are

now referred to as slender setae and what we previously referred to as spines are now called

robust setae. What we previously referred to mainly as teeth (non-articulating extrusions of the

cuticle), are now referred to as spines.

The following abbreviations are used on the plates : A, antenna; G, gnathopod; MD,

mandible; MP, maxilliped; MX, maxilla; P, peraeopod; T, telson; U, uropod; 1, left; r, right.

— 194 —

Didymochelia K. H. Barnard, 1931

Didymochelia K. H. Barnard, 1931 : 429. - K. H. Barnard, 1932 : 247. - Bellan-Santini

& Ledoyer. 1987 : 367. - Barnard & Karaman, 1991 : 276.

Didymocheila J. L. Barnard, 1969 : 478 ( lapsus calami).

KEY TO the species OF Didymochelia

1. Pleonite 3 with dorsodistal boss; coxa 7 with produced acute posterodistal corner.2

— Pleonite 3 without dorsodistal boss; coxa 7 with non-produced subquadrate posterodistal

corner . D. ledoyeri sp. nov.

2. Pleonite 3 with rounded dorsodistal boss. D. edwardi Bellan-Santini & Ledoyer

— Pleonite 3 with recurved acute boss. D. spongicola K. H. Barnard

Didymochelia ledoyeri sp. nov.

(Figs 1-3)

Type material. — Loyalty Basin east of Thio, New Caledonia, 21° 28’ S 166° 21.5’ E, depth 700-1265

m, collected from deep-sea submersible, Cyana, 3 March 1989, H. Zibrowius, CALSUB : stn PL 12 : 1 S, holo-

type, 4.5 mm (MNHN-Am 4484).

Etymology. — This species is named in recognition of the contribution of Michel Ledoyer to amphipod

systematics.

Distribution. — Didymochelia ledoyeri is known only from New Caledonia in 700 to 1265 m depth.

Diagnosis

Peraeopod 4 : coxa without posteroventral lobe. Peraeopod 6 : coxa with rounded posteroven-

tral comer. Peraeopod 7 : coxa with subquadrate posteroventral corner. Pleonite 3 without dorsal

boss. Epimera 2-3 with tiny posteroventral tooth.

Description

Based on holotype male, female not known. Head and body : without setae, colour not

known. Head : exposed, deeper than long; lateral cephalic lobe large, broadly rounded; rostrum

absent; eyes apparently absent. Antenna 1 : medium length, 0.23 x body; peduncular article 1

short, length 1 x breadth, distal margin with small midmedial swelling; peduncular article 2

short, 0.4 x article 1; peduncular article 3 short, 0.13 x article 1; accessory flagellum long,

about 0.5 x primary flagellum, at least 3-articulate, article 1 long, 1.25 x article 2; flagellum

5-articulate; callynophore strong 2-field, without posterodistal setae or robust setae, without

flagellar robust setae, calceoli absent. Antenna 2 : slightly longer than antenna 1; peduncle without

brush setae; flagellum 7-articulate, calceoli absent.

Mouthpart bundle : conical. Epistome and upper lip : fused. Mandible : incisors serrate, each

with about 7 cusps; left and right laciniae mobilis present, left, a broad serrate blade, right, a

distally cuspidate peg; accessory setal row with simple robust setae distally and distally cuspidate

— 195 —

pegs proximally; with pappose intermediate setae; molar columnar with weakly triturating surface,

with large plumose seta on each molar; mandibular palp attached slightly proximally; article 1

short, length 1 x breadth, without setae; article 2 elongate, slightly broadened proximally, length

2.7 x breadth, 1.5 x article 3, with 7 posterodistal A2-setae; article 3 slender, blade-like, short,

length 2.5 x breadth, with 4 (left), 5 (right) distal D3-setae, and 2 apical E3-setae.

Fig. 1. — Didymochelia ledoyeri sp. nov., holotype male, 4.5 mm, MNHN-Am 4484, Loyalty Basin, New Caledonia.

Didymochelia ledoyeri sp. nov., holotype, mâle 4,5 mm, MNHN-Am 4484, bassin des Loyautés, Nouvelle-Calédonie.

Maxilla 1 : inner plate tapering distally, inner margin fully setose, 19 plumose setae; outer

plate narrow with 9 long, slender multicuspidate setal-teeth; palp large, 2-articulate, with 13

long, plumose terminal setae. Maxilla 2 : inner and outer plates broad, inner plate 0.9 x length

outer plate, with well developed oblique setal row. Maxilliped : inner plate large, subquadrate,

long plumose setae line apical and medial margins; outer plate small, subovate, with long, plu¬

mose apical, medial and submarginal setae; palp 4-articulate, article 2 slender, length 2.1 x

breadth, subequal in length to article 3, article 3 long, slender, length 2.7 x breadth; dactylus

well developed, unguis present (right dactylus damaged).

Peraeonites : 1 to 7 dorsally smooth. Gnathopod 1 : chelate; coxa large, as long as coxa

2, anterior and posterior margins straight; basis long, slender, length 3.2 x breadth, anterior

— 197 —

margin with 1 proximal simple seta; ischium short, length 1.3 x breadth; merus, posterior margin

with a few simple setae; carpus subtriangular, short, length 1.3 x breadth, shorter than (0.5 x)

propodus; propodus large, subrectangular, length 2.3 x breadth, margins subparallel, posterior margin

smooth, slightly concave, with a few plumose setae and very fine setae near posterior margin; palm

extremely obtuse, margin straight, smooth, posterior corner without robust setae; dactylus simple.

Gnathopod 2: chelate; coxa large, subequal in size to coxa 3; ischium short, length 1.7 x breadth;

carpus subrectangular, long, length 2.6 x breadth, subequal in length to propodus; propodus large,

subrectangular, length 2.7 x breadth, margins subparallel, posterior margin smooth, straight, with a

few plumose setae and very fine setae near posterior margin; palm extremely obtuse, margin straight,

smooth, posterior corner without robust setae; dactylus simple.

Peraeopods 3 and 4: coxa large, anterior margin straight with broadly rounded anteroventral

corner, posterior margin straight; merus weakly expanded anteriorly with short robust setae along

posterior margin; carpus short, broader than long, with short robust setae along posterior margin;

propodus twice as long as broad, with 3-4 short robust setae and 2 distal locking setae along

posterior margin; dactylus short, slender, with well developed unguis. Peraeopod 5: coxa bilobate,

anterior lobe slightly produced; basis slightly expanded posteriorly with slightly rounded, smooth

posterior margin; merus slightly expanded posterodistally with 2 distal robust setae; propodus

with 2 robust setae and 2 distal locking setae along anterior margin; dactylus short, slender,

with well developed unguis. Peraeopod 6: coxa small, slightly lobate posteriorly; basis slightly

expanded posteriorly with straight, smooth posterior margin; merus slightly expanded

posterodistally with 1 distal seta; propodus with 2 robust setae and 2 distal locking setae along

anterior margin; dactylus short, slender, with well developed unguis. Peraeopod 7: coxa small,

not lobate posteriorly; basis slightly expanded with sinusoidal, smooth posterior margin; merus

slightly expanded posterodistally with 1 marginal robust seta and 3 distal robust setae; propodus

with 2 robust setae and 2 distal locking setae along anterior margin; dactylus short, slender,

with well developed unguis.

Gills : from gnathopod 2 to peraeopod 7, simple, small; gill on peraeopod 7 very small.

Pleonites I to 3: dorsally smooth. Epimeron 1 : posteroventral corner rounded. Epimeron

2 : posteroventral corner with tiny tooth. Epimeron 3: posteroventral corner with tiny tooth. Uro-

somite 7: large with well developed, subacute dorsal boss. Urosomites 2 and 3: very small.

Uropod 1: peduncle with 4 dorsolateral, 1 apicolateral and 2 dorsomedial robust setae; rami

subequal in length; outer ramus with 4 lateral robust setae; inner ramus with 1 medial and 2

lateral robust setae. Uropod 2: peduncle with 1 dorsolateral and 1 apicolateral robust setae; rami

subequal in length; outer ramus with 2 lateral robust setae; inner ramus with 2 medial and 3

lateral robust setae. Uropod 3: peduncle short, with 1 slender seta; rami absent. Telson : length

0.75 x breadth, entire, distal margin truncated, with 2 pairs of subterminal penicillate setae and

2 short, terminal robust setae.

Remarks

Didymochelia spongicola K. H. Barnard, 1931 and D. edwardi Bellan-Santini & Ledoyer,

1987, are very similar. Didymochelia ledoyeri differs from both of these species in coxa 4 which

has no posteroventral lobe, coxae 6 and 7 which are not produced and epimera 2-3, both of

which have a tiny posteroventral tooth.

Didymochelia ledoyeri sp. nov.. holotype, male 4,5 mm, MNHN-Am 4484, bassin des Loyautés, Nouvelle-Calédonie. Échelles

= 0,1 mm.

— 199 —

Acknowledgements

We are particularly grateful to Alain Crosnier who originally encouraged us to study the amphipods

from the MUSORSTOM Expeditions and who arranged for one of us to come to Paris and sort the

collections. We thank the Australian Museum Trust who provided travel money for this project. We thank

Stephen Keable who illustrated the species and Roger Springthorpe who composed and inked the plates.

REFERENCES

Barnard, J. L., 1969. — The families and genera of marine gammaridean Amphipoda. U.S. natn Mus. Bull.,

271 : 1-535.

BARNARD, J. L., & G. S. KARAMAN, 1991. — The families and genera of marine gammaridean Amphipoda.

Rec. Aust. Mus. Suppl., 13 : 1-866.

Barnard, K. H., 1931. — Diagnosis of new genera and species of amphipod Crustacea collected during the

“Discovery” investigations, 1925-1927. Ann. Mag. nat. Hist., Ser. 10, 7 : 425-430.

— 1932. — Amphipoda. “ Discovery ” Rep., 5: 1-326.

Bellan-Santini, D., & M. Ledoyer, 1987. — Gammariens (Crustacea, Amphipoda) des Iles Marion et Prince

Edward. Boll. Mus. civ. Stor. nat. Verona, 13: 349-435.

OSHEL, P. E., & D. H. STEELE, 1988. — Comparative morphology of amphipod setae, and a proposed classification

of setal types. Crustaceana Suppl. , 13: 100-106.

WATLING, L., 1989. — A classification system for crustacean setae based on the homology concept. In: B. E.

FELGENHAUER, L. WATLING & A. B. THISTLE (eds). Functional Morphology of Feeding and Grooming in

Crustacea, Crustacean Issues, 6: 15-27.

Bull. Mus. natl. Hist, nat., Paris, 4 e sér., 17, 1995

n° 1-2 : 201-210.

Two new species of Nanocassiope from the Western Pacific

(Crustacea, Brachyura, Xanthidae)

by Peter J. F. Davie

Abstract. — Two new species of Nanocassiope Guinot. 1967, are described, N. oblonga from French Pol¬

ynesia, and N. tridentata from Indonesia. This brings to six the number of species attributed to the genus, with

four found in the Indo-West Pacific region.

Keywords. — Crustacea, Decapoda, Brachyura, Xanthidae, Nanocassiope , Pacific, Indonesia, French Po¬

lynesia, new species.

Deux nouvelles espèces de Nanocassiope de lTndo-Pacifîque

(Crustacea, Brachyura, Xanthidae)

Résumé. — Deux nouvelles espèces de Nanocassiope Guinot, 1967 sont décrites : N. oblonga de Polynésie

française et N. tridentata d'Indonésie. Six espèces sont donc attribuées à ce genre dont quatre ont été récoltées

dans T Indo-Pacifique occidental.

Mots-clés. — Crustacea, Decapoda, Brachyura, Xanthidae, Nanocassiope, Pacifique, Indonésie, Polynésie

française, espèce nouvelle.

P. J. F. Davie, Queensland Museum, P. O. Box, 3300, South Brisbane, Qld. 4101, Australia.

INTRODUCTION

The present study was initiated by the discovery of what appeared to be a new Nanocassiope

Guinot. 1967, species amongst xanthid crab collections made by J. POUPIN of the French Service

Mixte de Contrôle Biologique (SMCB). The collections were made from the F. R. V. Marara

which conducted biological surveying throughout French Polynesia. The present specimens of

this new species were collected by dredge on the outer slopes of the islands in depths ranging

from 100 to 200 m. Deep water xanthid species collected from this region by trapping have

been previously reported by Davie (1993).

While examining comparative specimens held at the MNHN, a single male specimen of a

further new species was discovered that had been collected by the Rumphius I Expedition from

the vicinity of Ambon, Indonesia (see Romimohtarto, 1974). The opportunity was therefore

taken to describe both species at the same time.

Abbreviations used in the text are: mm, millimetres; Gl, G2, first and second male gonopods;

MNHN, Muséum national d’Histoire naturelle, Paris; QM, Queensland Museum, Brisbane. The

— 202 —

abbreviated terminology used for carapace regions is that used by Serene (1984) following Dana

( 1852). Measurements given in the text are of carapace breadth (c.b.) followed by length.

Family Xanthidae MacLeay, 1838

Xanthinae MacLeay, 1838

Nanocassiope Guinot, 1967

Nanocassiope Guinot, 1967: 355-358; 1971: 1075. - Sakai, 1976: 433. - Serene, 1984:

209.

Diagnosis

Carapace broad, moderately convex, distinctly areolated with the protogastric, mesogastric

and hepatic regions, and epigastric lobes, separated by distinct grooves. Anterolateral border

usually with four teeth behind exorbital angle; second and third teeth prominent, third tooth in

particular most laterally pointed; fourth tooth more or less developed; anterolateral border granu¬

lar with first tooth connected to exorbital angle by crest, but also with clusters or rows of granules

passing ventrally. Front wide, straight or slightly convex, with long submarginal granular crest.

Anterior border of the buccal cavity with sharp, sinuous rim, with distinct lateral fissures. En-

dostomial crests incomplete; lacinie of first maxilliped very short transversely, such that distant

from median line, with anterior border incurved, and advanced little in front of prelabial space.

Chelipeds very unequal, large claw rather massive and with fingers short; small claw thinner

with cutting margins sharp, very elongated, pointed and with tips crossing. Ambulatory legs long

and narrow. Sternal plastron with epistemites having convex latero-extemal border. Male abdo¬

men short, broad. Male G1 stocky, incurved and twisted, with spiniform tubercles extending

over half length, and with apical bouquet of long and strong incurved setae. (Modified after

Guinot, 1967).

Remarks

Nanocassiope Guinot (1967) now contains six species: the type of the genus, N. melano-

dactylus (A. Milne Edwards, 1867) from the tropical and subtropical eastern Atlantic; N. polita

(Rathbun, 1893) from the Pacific coast of America; N. alcocki (Rathbun, 1902) from the western

Indian Ocean; N. granulipes (Sakai, 1939) from Japan and questionably from South Africa; N. ob-

longa sp. nov. from French Polynesia; and N. tridentata sp. nov. from Indonesia.

Nanocassiope oblonga sp. nov.

(Fig. 1)

Type material. — The single male (MNHN-B22782) is the holotype, the three females, paratypes.

Material examined. — French Polynesia. SMCB (J. Poupin): Marquises Islands: Eiao, Stn D74, 7 0 59.81’S,

140°45.23’W, dredged, 155 m, 19.01.1991: 1 S 4.7 x 3.0 mm, holotype (MNHN-B22782). Ibidem'. 1 2 4.3 x

— 203 —

2.7 mm, paratype (MNHN-B22783). Ibidem : 1 2 3.8 x 2.4 mm, paratype (QM). Nuku Hiva, Stn D83, 8°47.60’S,

140°05.00’W, dredged, 140 m, 25.01.1991: 1 ovig. 2 4.5 x 2.8 mm, paratype (MNHN-B22784).

Etymology. — Named in reference to the very wide carapace.

Distribution. — Only recorded from French Polynesia. Bathymetric range: 140-155 m.

Description

Carapace transversely ovoid, c. 1.57-1.61 times broader than long; conspicuously granular

over anterior half, receding posterolaterally; without setae; convex anteriorly, more or less flat

from side to side across postero-branchial regions, but depressed near lateral margin. Regions

poorly indicated, mostly marked by shallow smooth grooves; IF and 2F confluent, swollen,

coarsely granular; 1M marked anteriorly by row of coarse granules, confluent with inner branch

of 2M; 2M broad, divided anteriorly by short groove, outer branch also with raised granular

row; anterior prolongation of 3M clearly marked, narrow, posteriorly 3M less clearly marked;

4M not separated; 1L not defined; 2L, 3L, 4L and 5L confluent, with raised granular row ante¬

riorly; 6L not defined; IR, 2R confluent, with raised anterior granular row; IP and 2P separated

by short, indistinct, cardio-intestinal groove. Anterolateral border of carapace divided into four

granular teeth; first tooth clearly separated from exorbital angle, lower than second and third;

second slightly larger than third, anteriorly directed; third more laterally directed; fourth tooth

very small. Greatest carapace width across third teeth. Posterolateral border markedly convergent,

straight or slightly convex, longer than anterolateral. Posterior border costate. Front not produced,

divided into two slightly convex lobes by small median notch; laterally clearly separated from

supra-orbital angles. Supra-orbital border granular; median and lateral fissures closed, incon¬

spicuous. Infra-orbital border granular, with broad triangular tooth at inner end visible dorsally;

deep broad notch below exorbital angle. Eyestalk with 2-3 strong tubercles on edge of cornea,

and low tubercle on extension over cornea. Basal antennal segment minutely granular, in contact

with ventral prolongation of front; flagellum slightly longer than width of orbit. Basal segment

of antennule granular laterally, with moderately strong ridges on superior and lateral margins,

flagellum folding obliquely.

Third maxilliped: merus c. 0.6 times length of ischium, wider than long, anteroexternal

angle moderately produced, bluntly rounded, surface granular; ischium c. 1.6 times longer than

wide.

Chelipeds noticeably unequal, right largest and stoutest. Merus of right cheliped minutely

granular on outer face, upper border coarsely granular, without subterminal or terminal teeth.

Carpus rounded, with coarse granular striations, bearing a strong triangular tooth at inner angle.

Palm high, height c. 0.5 times length (including fixed finger); coarsely granulated, size diminish¬

ing ventrally; broad shallow depression before median longitudinal granular crest on upper mar¬

gin; inner proximal margin with 2-3 larger granular tubercles; immoveable finger relatively long,

ventral sub-marginal groove, cutting edge with largest tooth proximo-medially. Dactyl with clearly

defined, relatively sharp, longitudinal ridge forming entire superior margin; cutting margin with

low teeth increasing in size proximally, and larger, blunt, outwardly and backwardly projecting

molar basally. Left cheliped of same form but fingers thinner and cutting margins sharper; dactyl

lacking basal molar. Fingers of both chelae tan coloured on one female, not obviously coloured

on other specimens.

— 204 —

Fig. 1. — Nanocassiope oblonga sp. nov., Eiao, Marquises Islands, French Polynesia, holotype, 4.7 x 3.0 mm (MNHN-B22782):

A, dorsal carapace; B, third maxilliped; C, third walking leg; D, fourth walking leg; E, frontal view of epistome and infra-orbital

margin; F, abdomen; G, right chela; H, I, first gonopod. Scale lines: A-F = 0.5 mm, G = 1 mm, H = 0.1 mm.

Nanocassiope oblonga sp. nov., Eiao, îles Marquises, Polynésie française, holotype, mâle 4,7 x, 3,0 mm (MNHN-B22782) :

A, carapace, vue dorsale ; B, troisième maxillipède ; C, troisième patte ambulatoire ; D, quatrième patte ambulatoire ; E,

vue frontale de l’épistome et bord infra-orbitaire ; F, abdomen ; G, pince droite ; H, I, premier pléopode. Échelles : A-F =

0,5 mm, H = 0,1 mm.

— 205 —

Walking legs of medium length, second pair slightly the longest, c. 1.1 times carapace

width. Margins of meri, carpi, and propodi sharply, and distinctly granulated. Third leg: merus

c. 3.3 times longer than wide; carpus c. 2.3 times longer than wide; propodus 2.5 times longer

than wide; dactylus c. 1.1 times longer than propodus.

Sternum with groove separating fused sternites 3/4 strongly incised laterally, but not apparent

across most of width; long longitudinal fissure on sternite 4. Male abdomen smooth; segment

3-5 fused; telson broadly rounded, c. 1.5 times wider than long, deeply sunken into segment 6,

subequal in length to lateral border of sixth; sixth segment with lateral margins convergent

towards base, c. 1.6 times wider than long (measured at widest and longest points); third segment

the widest, laterally triangular. G1 as figured; three strong, broad, spines medially on inner mar¬

gin.

Remarks

Nanocassiope oblonga sp. nov. is most closely allied to N. alcocki (Rathbun, 1902) from

which it can be separated by the following characters:

1) the first anterolateral tooth is broader, lower, and less acute;

2) the fourth anterolateral tooth is much reduced and placed on the posterolateral margin,

whereas on N. alcocki it is prominent, broad, and only a little smaller than the third;

3) the anterolateral margins are generally less granular;

4) the outer face of the major chela is granular over most of its surface, whereas on N. alcocki

the ventral face is smooth;

5) the shape and pattern of setation on the male G1 is different, especially by N. oblonga

having a longer, less twisted, apex ( cf. fig.lH with Guinot, 1967: fig. 12a,b);

6) it is apparently a smaller species with one female being ovigerous at 4.5 mm c.b., and

the largest specimen, the male, being only 4.7 mm across; N. alcocki reaches 26 mm across the

carapace.

Nanocassiope tridentata sp. nov.

(Fig. 2)

Nanocassiope orientalis Serène, Romimohtarto & Moosa, 1974: 22 (non Microcassiope

orientalis Takeda and Miyake, 1969: 201).

Type material. — Holotype is the unique male available for study.

Material examined. — Indonesia, Ambon Bay, Rumphius Expedition I: Cruise 3, Stn D19, dredged, bottom

of coarse sand and stones, 25.01.1973: 1 â 4.2 x 2.9 mm, holotype (MNHN-B10015).

Etymology. — Name refers to the presence of only three obvious anterolateral teeth behind the exorbital

angle, which helps to distinguish the species.

Distribution. — Only known from the type locality, in the waters near Ambon, Indonesia.

206

Fig. 2. — Nanocassiope tridentata sp. nov. Ambon, Indonesia, holotype - 4.2 x 2.9 mm (MNHN-B10015): A, dorsal carapace;

B, third walking leg; C, fourth walking leg; D, frontal view of epistome and infra-orbital margin; E, third maxilliped; F,

right chela; G, left chela; H, male abdomen; 1, J, first gonopod. Scale lines: A-E, F = 0.5 mm; F, G = 1 mm; FI = 0.1 mm.

Nanocassiope tridentata sp. nov., Ambon, Indonésie, holotype, mâle, 4,2 mm x 2,9 mm (MNHN-BW015) : A, carapace, vue

dorsale ; B, troisième patte ambulatoire ; C, quatrième patte ambulatoire ; D, troisième maxillipède ; E, vue frontale de

l'épistome et bord infra-orbitaire ; F, abdomen ; G, pince droite : H, I, premier pléopode. Échelles : A-E = 0,5 mm, F, G =

/ mm. H = 0,1 mm.

— 207 —

Description

Carapace c. 1.45 times broader than long; conspicuously granular over anterior half, receding

posterolaterally; without marked setation; convex anteriorly, more or less flat from side to side

across postero-branchial regions. Regions relatively well marked by smooth grooves; IF and 2F

confluent, swollen, coarsely granular; 1M poorly but completely defined; 2M broad, undivided;

anterior prolongation of 3M clearly marked, narrow, posteriorly 3M less clearly marked; 4M

not separated; 1L not defined; 2L and 3L confluent; 4L and 5L shallowly separated; indistinct,

cardio-intestinal groove. Anterolateral border of carapace divided into three granular teeth; first

tooth largest, broad, triangular, broadly separated from exorbital angle; second slightly smaller

and narrower; third tooth very small. Greatest carapace width across third teeth. Posterolateral

border markedly convergent, straight or slightly convex, longer than anterolateral. Posterior

border costate. Front not produced, divided into two slightly convex lobes by small median

notch; laterally clearly separated from supra-orbital angles. Supra-orbital border granular; median

and lateral fissures small but distinct. Infra-orbital border granular, with broad, blunt, triangular

tooth at inner end; deep broad notch below exorbital angle. Eyestalk with 2-3 strong tubercles

on edge of cornea, and further tubercle on extension over cornea. Basal antennal segment minutely

granular, barely in contact with ventral prolongation of front; flagellum about as long as width

of orbit. Basal segment of antennule granular laterally, flagellum folding slightly obliquely. Ante¬

rior margins of epistome granular.

Third maxilliped: merus c. 0.6 times length of ischium, wider than long, anteroexternal

angle slightly produced, rounded, surface granular; ischium c. 1.3 times longer than wide.

Chelipeds noticeably unequal, right largest and stoutest. Merus of right cheliped granular

on outer face, upper border with row of large pointed tubercles separated from broad blunt

terminal lobe. Carpus with coarse granular elevations, bearing strong triangular granular tooth

at inner angle. Palm high, height c. 0.5 times length (including fixed finger); coarsely granulated,

size diminishing ventrally; broad shallow depression on upper outer margin before dorsal surface;

inner proximal margin with row of larger rounded granules; immoveable finger relatively long,

ventral submarginal groove, cutting edge with large teeth. Dactyl with clearly defined, relatively

sharp, granulated longitudinal ridge along entire superior margin; cutting margin with low teeth

increasing in size proximally, and larger, blunt, outwardly and backwardly projecting molar

basally. Left cheliped of same form but fingers thinner and cutting margins sharper; dactyl lacking

basal molar. Fingers of both chelae without trace of colouring in preserved specimen.

Walking legs of medium length, third pair c. 1.1 times carapace width. Anterior margins

of meri and carpi coarsely granulated, forming two broad lobes on each segment. Third leg;

merus c. 3.3 times longer than wide; carpus c. 2.3 times longer than wide; propodus c. 2.2 times

longer than wide; dactylus c. 1.1 times longer than propodus.

Sternum with groove separating fused sternites 3/4 incised laterally, but only broad shallow

groove across most of width; long longitudinal fissure on sternite 4. Male abdomen smooth;

segment 3-5 fused; telson broadly rounded, c. 1.3 times wider than long, moderately sunken

into segment 6, c. 1.2 times length of lateral border of sixth; sixth segment with lateral margins

slightly convergent towards base, c. 1.6 times wider than long (measured at widest and longest

points); third segment the widest, laterally triangular. G1 relatively narrow; four subapical re¬

flexed stout setae; otherwise as figured.

— 208 —

Remarks

This specimen was found in the collection of the MNHN with the label carrying the ques¬

tioned name “ Nanocassiope ? orientalis ?”, with the identifier also questioned as Serene. Indeed,

Serène et al. (1974) did list this species in the list of identifications of the collection made by

the Rumphius Expedition I. On close scrutiny however it is apparent that this preliminary iden¬

tification is not correct because there are many differences between the two taxa. Most impor¬

tantly Takeda and Miyake (1969) described their species in the genus Microcassiope Guinot,

1967, and there is no reason to doubt this placement. Microcassiope orientalis certainly does

not belong to Nanocassiope , as it is presently defined, because it lacks the long, stiff, recurved

bristles on the male G1 which are so characteristic of all the species of that genus.

Nanocassiope tridentata sp. nov. is most closely related to N. granulipes (Sakai, 1939). It

differs from that species, and all the other described species, by having only three anterolateral

teeth behind the exorbital angle instead of four, the first being vestigial and represented by only

one or two slightly raised granules. N. tridentata further differs from N. granulipes by having

the entire outer face of the cheliped palm covered in coarse granules; in the later species the

palm is “dorsally slightly carinate and granulated but its external surface near the inferior border

is smooth" (SAKAI, 1939: 547). The anterior extension of 3M is narrow, while on N. granulipes

it is relatively wide and more obviously tapering (cf. Sakai, 1939: fig. 59). Finally, the male

G1 is narrower and less stout than the other described species.

Acknowledgments

I am most grateful to Dr Alain Crosnier of ORSTOM for asking me to undertake this study. The

work was undertaken while at the Laboratoire de Zoologie (Arthropodes), Muséum national d’Histoire

naturelle, Paris, under a grant from the Institut français de Recherche Scientifique pour le Développement

en Coopération (ORSTOM). The Board of Trustees of the Queensland Museum very kindly helped with

travel funds. Allison Hill is thanked for her artwork used to illustrate N. ovalis. Danièle Guinot of the

MNHN, Paris, and Paul Clark of The Natural History Museum, London, both made valuable comments

on the manuscript, and to both I am very grateful.

REFERENCES

DANA, J. D„ 1852. — Crustacea Part 1. In: United States Exploring Expedition during the years 1838, 1839,

1840, 1841, 1842 under the command of Charles Wilkes, U.S.N., 13 : viii + 1-685. Philadelphia.

Davie, P. J. F., 1993. — Deepwater xanthid crabs from French Polynesia (Crustacea, Decapoda, Xanthoidea).

Bull. Mus. natn. Hist, nat., Paris, 4 e sér., 14, section A: 501-561, Figs 1-12, Pis 1-13.

GUINOT. D., 1967. — Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes

Brachyoures. II. Les anciens genres Micropanope Stimpson et Medaeus Dana. Bull. Mus. natn. Hist, nat.,

Paris, 2 e sér., 39 (2) : 345-374, fig. 1-42.

— 1971. — Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures.

VIII. Synthèse et bibliographie. Bull. Mus. natn. Hist, nat., Paris, 2 e sér., 42 (5), 1970 (1971): 1063-1090.

MACLEAY, W. S., 1838. — On the Brachyurous Decapod Cmstacea brought from the Cape by Dr SMITH. P.

63-72, pi. 3. In: Illustrations of the Annulosa of South Africa; being a portion of the objects of natural

history collected during an expedition into the interior of South Africa, under the direction of Dr Andrew

— 209 —

SMITH, in the years 1834, 1835, and 1836; fitted out by "The Cape of Good Hope Association for Exploring

Central Africa”, lllustr. Zool. S. Africa Invest., London: PI. 14, 75 p.

Milne Edwards, A., 1867. — Descriptions de quelques espèces nouvelles de Crustacés Brachyoures. Annls

Soc. Ent. Fr., 4 e sér„ (A2) 7 : 263-288.

RATHBUN, M. J., 1893. — Descriptions of new genera and species of Crabs from the West Coast of North

America and the Sandwich Islands. In: Scientific Results of Exploration by the U.S. Fish Commission

“Albatross”. No XXIX. Proc. U.S. natl. Mus., 16 (933): 223-260.

— 1902. — Crabs from the Maidive Islands. Bull. Mus. comp. Zool. Harv., 39 (5): 123-138, 1 PI.

ROMIMOHTARTO, K., 1974. — Report on the Rumphius Expedition 1. Part I. Description of the Expedition.

Oseanologi di Indonesia, 1: 1-11.

SAKAI, T., 1939. — Studies on the Crabs of Japan. IV. Brachygnatha, Brachyrhyncha. (Yokendo: Tokyo), p.

365-741, text-Figs 1-129, Pis 42-111, Tables 16.

— 1976. — Crabs of Japan and the Adjacent Seas. (Tokyo: Kodansha). 3 vols, p. i-xxix + 1-773, text Figs

1-379, maps 1-3 (English); p. 1-16, Pis 1-251 (Plates); p. 1-461, text Figs 1, 2 (Japanese).

Serene, R., 1984. — Crustacés Décapodes Brachyoures de l'Océan Indien occidental et de la Mer Rouge, Xan-

thoidea: Xanthidae et Trapeziidae. Avec un addendum par CROSNIER, A.: Carpiliidae et Menippidae. Faune

Tropicale. Office de la Recherche Scientifique et Technique Outre-Mer. Paris, 24 : 1-400, fig. A-C, 1-243,

pl. 148.

SerÈNE, R., K. ROMIMOHTARTO & M. K. MOOSA, 1974. — Report on the Rumphius Expedition I. Part II. 3.

The Hippidea and Brachyura collected by the Rumphius Expedition I. Oseanologi di Indonesia, 1: 17-26.

Takeda, M., & S. Miyake, 1969. — On two species of the family Xanthidae (Crustacea, Brachyura) from

southern Japan. OHMU 2 (9): 195-206, Figs 1-3.

Bull. Mus. nail. Hist. nal.. Paris, 4 e sér., 17, 1995

n° 1-2 : 211-218.

Une espèce nouvelle de Neocallichiriis aux îles Tuamotu,

Polynésie française

(Crustacea, Decapoda, Thalassinidea)

par Nguyen NGOC-HO

Résumé. — Une espèce nouvelle de la famille des Callianassidae récoltée en Polynésie française, Neocal-

lichirus taiaro, est décrite et illustrée. Elle est proche d’un groupe d’espèces dont l’endopodite des uropodes est

plus large que long et surtout de Neocallichirus lemaitrei Manning provenant de Colombie, à laquelle elle est

comparée.

Mots-clés. — Crustacea, Decapoda, Thalassinidea, Callianassidae, taxonomie, Neocallichirus, espèce nou¬

velle, Polynésie française.

A new species of Neocallichirus from the Tuamotu Islands, French Polynesia

(Crustacea, Decapoda, Thalassinidea)

Abstract. — A new species of the Callianassidae from French Polynesia, Neocallichirus taiaro , is described

and figured. It is related to a group of species whose uropodal endopod is broader than long, and especially to

Neocallichirus lemaitrei Manning from Colombia. Distinguishing characters between the two species are discussed.

Keyword. — Crustacea, Decapoda, Thalassinidea, Callianassidae, taxonomy, Neocallichirus, new species,

French Polynesia.

N. Ngoc-Ho, Laboratoire de Zoologie ( Arthropodes), Muséum national d'Histoire naturelle, 61, rue Buffon, F-75005 Paris.

INTRODUCTION

Dans le matériel de crustacés Décapodes récolté par Joseph Poupin lors de son voyage en

Polynésie française, en février 1994, se trouve un spécimen femelle de Callianassidae appartenant

à une espèce encore non décrite du genre Neocallichirus Sakai. Elle est présentée ici comme

une espèce nouvelle, Neocallichirus taiaro.

Le genre Neocallichirus a été établi en 1988 par Sakai qui y inclut les espèces australiennes

N. horneri Sakai 1988, espèce-type, N. darwinensis Sakai 1988, N. limosa (Poore, 1975), N. cae-

chabitator Sakai, 1988, ainsi que les espèces américaines N. grandimana (Gibbes, 1850) et N. ra-

thbunae (Schmitt, 1935), les espèces indonésiennes N. indica (de Man, 1905) et N. moluccensis

(de Man, 1905), l'espèce sud-africaine N. natalensis (Barnard, 1947) et l’espèce est-atlantique

N. sassendrensis (Le Lœuff & Intès, 1974).

À cette liste, Manning & Felder (1991) ajoutaient quatre espèces américaines ouest-at¬

lantiques N. guara (Rodrigues, 1971), N. guassutinga (Rodrigues, 1971), N. mirim (Rodrigues,

— 212

1971) et N. trilobata (Biffar, 1970). Ces espèces seront ultérieurement séparées par Manning

& Lemaître (1993) en un genre nouveau Sergio. En 1993, Manning signalait deux nouvelles

espèces de Neocallichirus en provenance de la mer des Caraïbes, N. nickellae et N. lemaitrei.

La première espèce de ce genre dans l’océan Indien a été décrite par Kazmi & Kazmi (1992),

et NGOC-HO (1994) décrit une autre espèce nouvelle d’Australie, N. denticulatus. N. taiaro, qui

fait l’objet de cette note, est la première espèce de Neocallichirus signalée en Polynésie française.

Les dimensions indiquées se rapportent à la longueur de la carapace (l.c.), mesurée de l’ex¬

trémité du rostre au bord postérieur de la carapace et à la longueur totale (l.t.), mesurée de

l'extrémité du rostre au bord postérieur du telson.

Genre NEOCALLICHIRUS Sakai, 1988

Neocallichirus taiaro sp. nov.

(Fig. 1-2)

Holotype. — Polynésie française : îles Tuamotu, atoll de Taiaro; 15°45’ S-144°40’ W; J. Poupin coll., à

la main dans terrier de sable fin corallien, 12-14 février 1994 / SMCB : $, l.c. 8,5 mm, 1. t. 32 mm (MNHN-Th

1291).

Etymologie. — L’espèce est nommée d’après sa localité-type.

Description

Carapace (fig. la, lb) avec ovale céphalo-thoracique bien délimité, sillon cervical profond,

placé au 4/5 e environ de la longueur de la carapace; linea thalassinica distincte. Rostre largement

triangulaire, à sommet obtus, dépassant à peine la base des pédoncules oculaires; saillies latérales,

de part et d’autre des pédoncules oculaires, très faibles.

Pédoncules oculaires comprimés dorso-ventralement, à extrémité obtuse, atteignant la base

du 2 e segment des antennules et portant vers le milieu des cornées latéro-dorsales globuleuses.

Pédoncules antennulaires plus courts que les pédoncules antennaires (fig. lb), munis de

longues soies, avant-dernier segment plus court que le dernier; longues soies denses sur les

flagelles.

Pédoncules antennaires munis de quelques longues soies, dépassant ceux des antennules de

la moitié du dernier segment, avant-dernier segment plus long que le dernier, écaille antennaire

en petit lobe ; flagelles avec peu de soies.

Premiers maxillipèdes (fig. ld) avec épipodite pourvu d’un lobe antérieur bien développé,

à sommet aigu ; endopodite rudimentaire, en un petit lobe arrondi.

Deuxièmes maxillipèdes (fig. le) munis d’un petit épipodite; exopodite d’aspect foliacé.

Troisièmes maxillipèdes (fig. If, lg) sans exopodite, ni épipodite; ischion et mérus subpé-

diformes, une rangée oblique d’épines à la face mésiale de l’ischion; propode dilaté ventralement,

plus de quatre fois plus large que le dactyle; dactyle effilé, environ quatre fois plus long que

large.

Grand péréiopode 1 (fig. 2a) à droite, puissant; ischion et mérus à peu près de même lon¬

gueur, ischion deux fois moins large, faiblement denté ventralement, mérus avec lobe ventral

saillant portant une douzaine d’épines; carpe plus court que le mérus, à peu près aussi large

— 213

Fig. 1. — Neocallichirus taiaro sp. nov., holotype : a, animal entier, vue dorsale; b, céphalothorax, vue dorsale; c, telson et

uropodes, vue dorsale; d, e, f, 1 er , 2' et 3 e maxillipède respectivement, vue ventrale; g, 3 e maxillipède, dénudé, vue mésiale.

Échelle : 2 mm.

Neocallichirus taiaro sp. nov., holotype: a, body in dorsal view; b, céphalothorax in dorsal view; c, telson and uropod in

dorsal view; d, e, f, maxilliped 1, 2 and 3 respectively in ventral view; g, maxilliped 3, setae omitted, in mesial view. Scale

bar: 2 mm.

— 214 —

que la portion palmaire de la main, muni de 3 épines ventrales distales ; paume environ 2,5 fois

plus longue que le carpe, portant 13 petites épines distribuées du 1/4 au 2/3 du bord ventral;

doigt fixe aussi long que le dactyle, avec 5 tubercules (un grand, quatre petits) à sommet arrondi

dans la région proximale du bord préhensile et une rangée de 3 autres tubercules proximaux,

visibles sur le côté externe ; dactyle à extrémité légèrement incurvée, bord préhensile avec 2

petites dents vers son milieu.

Petit péréiopode 1 (fig. lb) plus grêle, ischion avec une dizaine de denticules ventraux, à

peu près aussi long que le mérus; carpe 0,8 fois aussi long que le mérus et presque 2 fois plus

large; paume légèrement plus longue que le carpe; doigt fixe et dactyle de même longueur et

de même longueur que le carpe.

Péréiopodes 3 (fig. 2d) à propode dilaté ventralement mais ne dépassant pas le bord ventral

du carpe.

Péréiopodes 4 (fig. 2e) avec une saillie disto-ventrale au propode leur conférant un aspect

subchéliforme.

Péréiopodes 5 (fig. 20 à extrémité subchéliforme.

Abdomen (fig. la) dont le dernier segment est le plus long; rapports approximatifs de lon¬

gueur des autres segments par rapport à celui-ci (= 1) : 5 e et 2 e = 0,85; 4 e et 3 e = 0,65; 1 er =

0,75.

Pléopodes 1 (fig. 2g) biarticulés, grêles.

Pléopodes 2 (fig. 2h) munis d'un exopodite, un appendice interna placé au quart distal

environ du bord latéral interne de l’endopodite.

Pléopodes 3-5 (fig. 2i) à exopodite et endopodite larges, un petit appendice interna encastré

à peu près au milieu du bord interne de l’endopodite.

Telson (fig. le) légèrement plus large dans sa partie proximale que long, bord postérieur

faiblement arrondi.

Uropodes (fig. le) : exopodite à bord postérieur légèrement convexe, lobe antéro-dorsal

occupant presque la moitié de l’article, et s’écartant peu du bord distal; endopodite plus large

que long, à bord postérieur presque rectiligne.

Remarques

Le nouveau taxon présente les caractères de Neocallichirus signalés dans la diagnose du

genre par Sakai (1988), puis Manning & Felder (1991), à l’exception de ses cornées qui sont

dorso-latérales et globuleuses au lieu d’être dorsales et aplaties ( dorsal, disk-shaped ). Des cornées

semblables à celles de N. taiaro ont déjà été décrites chez certaines espèces ouest-atlantiques,

notamment chez N. grandimana (Gibbes), N. rathbunae (Schmitt), N. nickellae Manning et aussi

chez N. manningi (Kazmi & Kazmi) de Karachi. D'après Manning (1993), les trois espèces

ouest-atlantiques de Neocallichirus constituent, avec N. lemaitrei Manning de Colombie et N.

horneri Sakai d’Australie, un groupe d’espèces qui ont un endopodite des uropodes plus large

que long, aplati distalement, et un bord postérieur du telson arrondi ou faiblement concave. N.

taiaro est proche de ce groupe et en particulier de N. lemaitrei à laquelle elle ressemble aussi

par les caractères suivants :

1) rostre et projections frontales de la carapace très faibles;

2. — Neocallichirus taiaro sp. nov., holotype : a, grand péréiopode 1 ; b. petit péréiopode 1 : c, 3' stemite, vue ventrale ;

d, péréiopode 3 ; e, f, partie distale du péréiopode 4 et 5 respectivement; g, pléopode 1 ; h, pléopode 2; i, pléopode 3.

Échelle : 2 mm.

Neocallichirus taiaro sp. nov., holotype: a, large pereopod 1; b, small pereopod I: c, 3rd stemite in ventral view: d, pereopod

3; e, f distal part of pereopod 4 and 5 respectively: g, pleopod 1; h, pleopod 2 ; i, pleopod 3. Scale bar: 2 mm.

— 216

2) telson légèrement plus large que long ;

3) morphologie et spinulation du grand péréiopode 1 de la femelle ;

4) morphologie et spinulation des péréiopodes 3-5 ;

Les deux espèces peuvent être différenciées de la façon suivante :

1) cornées globuleuses chez N. taiaro, aplaties chez N. lemaitrei (voir Manning, 1993:

fig.la, lb, le);

2) propode du maxillipède 3 régulièrement arrondi et lisse au bord ventro-distal chez N.

taiaro, légèrement denticulé à ce bord chez N. lemaitrei ;

3) proportions différentes du petit péréiopode 1 de la femelle, en particulier le carpe est

plus court que la paume chez N. taiaro, plus long chez N. lemaitrei',

4) exopodite des uropodes approximativement aussi long que sa largeur au bord distal et

le lobe dorsal occupant moins de la moitié de l’appendice chez N. taiaro', exopodite des uropodes

environ 1,5 fois plus large que long et le lobe dorsal occupant à peu près la moitié de l’article

chez N. lemaitrei.

Remerciements

L’auteur remercie Joseph Poupin qui lui a confié ce matériel à étudier et Michèle de Saint-Laurent

pour ses commentaires sur le travail.

RÉFÉRENCES

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MANNING, R. B., & R. Lemaître, 1993. — Sergio, a new genus of ghost shrimp from the Americas (Crustacea:

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POORE, g. C. b., 1975. — Systematics and distribution of Callianassa (Crustacea, Decapoda, Macrura) from

Fort Phillip Bay, Australia, with description of two new species. Pacific Sci., 29 (2): 197-207.

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SAKAI, K., 1988. — A new genus and five new species of Callianassidae (Crustacea: Decapoda: Thalassinidea)

from Northern Australia. The Beagle, 5 (1): 51-69, Figs 1-10.

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1-21, Pis 1-5.

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