Éditions scientifiques du Muséum, Paris

BULLETIN DU MUSÉUM NATIONAL D’HISTOIRE NATURELLE

57, rue Cuvier, 75005 Paris

Section A : ZOOLOGIE

Rédacteur en chef : Christian Érard

Rédacteur adjoint : Danielle Defaye

Assistante de rédaction : Florence Kerdoncuff

S’adresser

pour les échanges :

Service des périodiques et des échanges de la Bibliothèque Centrale

du Muséum national d’Histoire naturelle

38, rue Geoffroy Saint-Hilaire, 75005 Paris

Tél. : [33] 01 40 79 36 41

pour les abonnements et achats au numéro :

Éditions Scientifiques du Muséum - Diffusion

57, rue Cuvier, 75005 Paris

Tél. : [33] 01 40 79 37 00

Fax: [33] 01 40 79 38 40

e-mail : dhenry-mnhn

pour l’envoi de manuscrits :

Éditions Scientifiques du Muséum - Rédaction du Bulletin

57, rue Cuvier, 75005 Paris

Tél. : [33] 01 40 79 34 38

Fax : [33] 01 40 79 38 58

e-mail : bulletin@mnhn.fr

Abonnements pour l'année 1996 (prix HT) :

Abonnement général : 1 800 FF

Section A : zoologie, biologie et écologie animales : 800 FF

Section B : botanique Adansonia : 500 FF

Section C : sciences de la Terre, paléontologie, géologie et minéralogie : 600 FF.

Numéro d'inscription à la Commission paritaire des publications et agences de presses : 1403 AlSource : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle, Paris

4' série, 18, 1996, Section A (Zoologie, Biologie et Écologie animales) n“ 3-4

SOMMAIRE

Alain Dubois & Annemarie Ohler. — Early scientific names of Amphibia Anura.

I. Introduction . 297-320

Alain Dubois & Annemarie OHLER. — Early scientific names of Amphibia

Anura. II. An exemplary case : Rana arborea Linnaeus, 1758 . 321-340

Edward B. Cutler & Norma J. CUTLER. — Sipuncula from the Indian Ocean and

New Caledonia . 341-365

Gabriel N’Zobadila, Johnny BOYER & Marie-Claude Durette-Desset. — Morpho¬

genèse d'Heligmosomoides polygyrus polygyrus (Dujardin, 1845) (Trichostrongy-

lina-Heligmosomoidea) chez Apodemus flavicollis en France. Comparaison avec

les espèces proches : Heligmosomoides laevis (Dujardin, 1845) et Heligmosomum

mixtum Schulz, 1954, parasites d’Arvicolidae . 367-385

Robert G. Moolenbeek & Dieter RÔCKEL. — Cones taken off Wallis and Futuna

Islands, South-West Pacific (Mollusca, Gastropoda, Conidae) . 387-400

Anatoly A. SCHILEYKO. — Guamampa n.g. (Gastropoda, Pulmonata) an “intermediate

link” . 401-408

David G. Herbert. — A critical review of the trochoidean types in the Muséum d'His-

toire naturelle, Bordeaux (Mollusca, Gastropoda) . 409-445

John S. Buckeridge. — A living fossil Waikalasma boucheti n.sp. (Cirripedia, Balano-

morpha) from Vanuatu (New Hebrides), Southwest Pacific . 447-457

Stefano Taiti and Franco FERRARA. — The terrestrial Isopoda of Corsica (Crustacea,

Oniscidea) . 459-545

Christopher C. Tudge. — Spermatophore morphology and spermatozoal ultrastructure

of the recently described hermit crab, Strigopagurus boreonotus Forest, 1995

(Decapoda, Anomura, Diogenidae) . 547-555

Alain CROSNIER. — Hypocolpus pararugosus, espèce nouvelle de l’Indo-Ouest Paci¬

fique (Crustacea, Decapoda, Brachyura, Xanthidae) . 557-564

Nguyen NGOC-HO. — Redescription des types de Naushonia perrieri (Nobili, 1904)

(Crustacea, Decapoda, Laomediidae) . 565-570

Ourida Abrous-Kherbouche & Jean-Paul Mauries. — Découverte du genre

Archipolydesmus en Algérie : description de trois espèces nouvelles (Diplopoda,

Polydesmida, Polydesmidae) . 571-587

Source : MNHN, Paris

Patrick Maréchal. — Psalistops gasci n.sp., première Barychelidae de Guyane fran¬

çaise (Araneae, Mygalomorphae) . 589-594

Claire VOISIN & Jean-François VOISIN. — Liste des types d’oiseaux des collections

du Muséum national d’Histoire naturelle de Paris. 4. Hérons (Ardeidae) . 595-609

Les articles du présent volume doivent être cités comme suit / The articles of the present

volume should be cited as follows:

Dubois A. & Ohler A. 1996. — Early scientific names of Amphibia Anura I. Introduc¬

tion. Bulletin du Muséum national d' Histoire naturelle , Paris, série 4, C 18 (3-4) : 297-320

ou / or

Dubois A. & Ohler A. 1996. — Early scientific names of Amphibia Anura I. Introduc¬

tion. Bull. Mus. natl Hist, nat., Paris, sér. 4, C 18 (3-4) : 297-320

Source : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle, Paris, 4 e série, 18, 1996

Section A, n“ 3-4 : 297-320

Early scientific names of Amphibia Anura

I. Introduction

by Alain DUBOIS & Annemarie OHLER

Abstract. — This paper opens a series of publications meant at ascertaining and stabilizing the status of

early scientific names of Amphibia Anura. Before starting the analysis of the status of these names, we present

a rather detailed discussion of several important rules and problems of zoological nomenclature, which will be

the basis for proposals and decisions presented in the forthcoming series of papers. We examine here the following

general questions: the status of nomina oblita and of nomina dubia ; the “suppression” of names by the International

Commission on Zoological Nomenclature; the nomenclatural role and importance of name-bearing types (ono-

matophores) and type-localities (type-populations); the availability of scientific names; the distinction between

different kinds of names and spellings; the distinction between three categories of syntypes; the criteria and

conditions for lectotype designations; the status of type-locality restrictions; and the relative priority between

names published simultaneously.

Key-words. —- Status of early scientific names, nomenclature, International Code, International Commission,

name-bearing types, type-localities. Amphibia, Anura.

Noms scientifiques anciens d'amphibiens anoures

I. Introduction

Résumé. — Cet article est le premier d’une série de publications qui auront pour but de confirmer et de

stabiliser le statut des noms scientifiques anciens d’amphibiens anoures. Avant de commencer l’analyse du statut

de ces noms, nous offrons une discussion assez détaillée de plusieurs règles importantes et de problèmes de la

nomenclature zoologique qui seront à la base des propositions et des décisions présentées dans la série d’articles

inaugurée par celui-ci. Nous y examinons les questions générales suivantes ; le statut des nomina oblita et des

nomina dubia ; la « suppression » de noms par la Commission internationale de Nomenclature zoologique ; le

rôle nomenclatural et l’importance des types porte-noms (onomatophores) et des localités-types (populations-

types) ; la disponibilité des noms scientifiques ; la distinction entre différentes catégories de noms et différentes

orthographes ; la distinction entre trois catégories de syntypes ; les critères et les conditions pour la désignation

de lectotypes; le statut des restrictions de localité-type; et la priorité relative de noms publiés simultanément.

Mots-clés. — Statut de noms scientifiques anciens, nomenclature. Code international. Commission inter¬

nationale, types porte-noms, localités-types, amphibiens. anoures.

A. Dubois & A. Ohler. Muséum national d'Histoire naturelle. Laboratoire des Reptiles et Amphibiens. 25 rue Cuvier, F-75231

Paris cedex 05.

PRELIMINARY NOTE

In this paper and in forthcoming papers of this series, the expression “the Commission”

refers to the International Commission on Zoological Nomenclature, and “the Code ” to the In¬

ternational Code on Zoological Nomenclature (ANONYMOUS 1985). The following expressions,

which are used in the Code or derived from expressions used therein, will always be written

with a dash, in order to show that they are well defined formulae with a precise technical meaning:

species-group, genus-group, family-group and class-group (names); type-specimen, type-series,

Source : MNHN, Paris

— 298 —

type-species, type-genus, type-locality, type-population, syntype-species, syntype-genera, name-

bearing type. Finally, Latin scientific names which are written between quotation marks are names

which are not nomenclaturally available according to the current Code.

INTRODUCTION

Since 1758, the year that was arbitrarily fixed for the beginning of availability of Latin

scientific names of animals within the frame of the Code , about 2 millions scientific names have

been proposed for animal taxa (Hammond 1992). The vast majority of these names were pub¬

lished during our century, after rules for zoological nomenclature had been devised and adopted

by the international community of zoologists. A much smaller number of names were published

between 1758 and the end of the 19th century, but these names are of particular importance,

since they are the oldest ones and have therefore a high probability of remaining the valid names

for the taxa they designate.

Most of these ancient names were published in books or papers that are now the classical

works of zoology, and one would expect these names to be well known, and their nomenclatural

status to be firmly established and stabilized. Strangely enough, careful examination of these

famous old works shows that this expectation is not borne out. As a matter of fact, many of

these classical works contain a number of names the status of which has not been properly

studied and stabilized by recent taxonomists.

While many of these old names are still currently in use to designate taxa considered valid

by present-day taxonomists, many others are not. Some names are not used because they are

considered junior synonyms or homonyms of valid names, or for more specific reasons, e.g.

because they have been expressly “suppressed” by the Commission. These names, although they

are not currently used, are not forgotten, and they are usually quoted in synonymies of names

now considered valid. Unless they have been “suppressed” by the Commission, they remain

therefore available and are not infrequently resurrected to designate new taxa, as the taxonomy

of an animal group evolves and more distinct taxa are recognized. But other names remain unused

without being properly allocated to any given taxon. These include both names that have been

“forgotten” by most or all members of the international scientific community (“ nomina oblita"),

and names which have not been completely forgotten, but the status of which is unclear, since

the information associated with the original publication of these names does not allow a clear

allocation of these names to biological taxa (“ nomina dubia").

“NOMINA OBLITA" AND “ NOMINA DUBIA"

Several attitudes can be and have been adopted by taxonomists towards nomina oblita and

nomina dubia.

Some taxonomists consider that these names should be definitively forgotten, and for ex¬

ample should not be quoted any more in synonymies, checklists and catalogues, “perhaps in the

hope that ‘they would go away’” (Duellman & Lynch 1981: 237). In other cases, some authors

ask for the “suppression” of these names by the Commission. For many taxonomists, “abandon-

Source : MNHN, Paris

— 299 —

ment” of these old names is justified by the fact that they were often poorly defined in the

original publications where they were created and should not be used for taxa recognized in

modem days through the use of more detailed methods and elaborate techniques, such as bio¬

acoustics, protein electrophoresis or other biochemical methods, etc. (see e.g. BÔHME & Wiedl

1994: 39). Finally, other taxonomists think that such names should be “suppressed” simply be¬

cause they have not been used for a long time, and that their “resurrection” would “threaten

the stability of nomenclature". However, this is true only in some cases, because, very often,

the biological taxon represented by such a “forgotten” name has had several successive names,

or has been very seldom mentioned in the scientific literature, so that one cannot seriously speak

of a long and “current” usage. The term of “stability of nomenclature” has a meaning only when

long periods of times and high numbers of publications and of independent authors are involved.

Understanding of these limitations is clearly the reason for the suppression, in the last edition

of the Code, of the so-called “nomen oblitum rule” which existed in Article 23b of the 1961

and 1964 editions (Anonymous 1961, 1964). In the current 1985 edition, there is no such rule,

and “suppression” of a nomen oblitum or of a nomen dubium can be obtained only through an

application to the Commission followed by the vote of an “Opinion” by the latter, but not by

any “automatic” rule.

The opposite attitude towards these old names, which we have been strongly advocating

for many years (see e.g.: DUBOIS 1977: 255, 1981, 1984, 1987; DUBOIS et al. 1988: Dubois &

Ohler 1995), is that every efforts should be made to allocate properly these ancient names,

through the use of the normal provisions of the Code, so that they stop being a potential cause

of doubt or of instability in zoological nomenclature. Such a proper allocation very often (al¬

though not always) requires the recourse to a lectotype or neotype designation. Once this is

done, the taxonomic status of the name is definitively fixed, and subsequent authors are free to

use it either as a valid name or as an invalid synonym, but not to ignore it. We believe that

the generalization of such an attitude towards ancient names would rather quickly lead to the

disappearance of most of the nomenclatural problems related to these ancient names, some of

which have already been the matter of very numerous, and sometimes “endless”, discussions

among zoologists. Some of these long discussions could have been avoided by a proper study,

understanding and use of the existing rules of the Code. The latter unfortunately remains too

little understood by many taxonomists, who tend more and more to appeal to the Commission

to solve nomenclatural problems which could be solved otherwise.

“SUPPRESSION” OF ZOOLOGICAL NAMES

We fully concur with Holynski (1994) that the recent tendency of the Commission to

“suppress” more and more zoological names has been extremely exaggerated and has done a

great deal of harm to the “image” of the Code. It is our feeling that the Commission has not

properly played its role of “Keeper of the Law” in front of the international community of

zoologists. Rules that are too often overlooked or circumvented are no more convincing to anyone.

Furthermore, in more and more cases, it is clear that recent zoologists are deliberately ignoring

Source : MNHN, Paris

— 300 —

the existing rules to follow “their own” rules (see e.g. the Mycetoglossini/Hemidactyliini case: Wake

1993; SMITH & Wake 1993, 1994, 1995; Dubois 1994, 1995b). This tendency, if continued and

supported by the Commission itself, is liable to lead soon to the ruin of the international consensus

on the Code , and to the disappearance of the universality and stability of names among zoologists.

After others, HOLYNSKI (1994) has dealt with these problems in a long and detailed dis¬

cussion, which we strongly urge our readers to study carefully (see Appendix I). Among his

proposals are that of changing the rules allowing the Commission to “suppress” names and to

place them on lists of “suppressed” names. We fully agree that the possibility to “suppress”

names should be limited to very specific cases, by making the conditions for a name to be

eligible for “suppression” by the Commission much more stringent (see HOLYNSKI 1994 and

Appendix II). We want to stress here that our attitude on this question is not new and has been

the attitude of many taxonomists in the past, including officers of the Commission, as shown

by this quotation from a letter sent on 21 February 1975 to one of us (A. D.) by Richard V. Mel¬

ville (London), then Secretary of the Commission : “Au fond, le ‘principe de conservation' est

justifié dans des cas de noms importants, employés par beaucoup de zoologistes et autres, et

surtout dans des domaines hors de la taxonomie - agriculture, eaux et forêts, médecine, bio¬

stratigraphie, etc. Mais lorsqu’il s’agit d’un groupe qui intéresse à l’heure actuelle surtout les

taxonomistes, et dont les noms ne sont pas très souvent cités, la Loi de Priorité peut être appliquée

normalement.” This remark points to an important difference between names widely used outside

the community of taxonomists, for which, for pragmatic reasons of communication among scien¬

tists, exceptions to the rules can be made in some cases, and names that have been used only

or mostly by taxonomists, for which exceptions should not be acceptable, even when the use of

these names has been rather important. In other words, among taxonomists themselves , changes

of names for nomenclatural reasons should be easily accepted, as taxonomists know the reasons

for such changes and should accept them with “fair play” when they are needed to correct a

mistake, even when the latter was made and perpetuated by themselves.

The recent tendency for the Commission to accept to “suppress” almost all names for which

“suppression” is requested by any individual zoologist (see HOLYNSKI 1994) is a strong en¬

couragement for zoologists to deal with nomenclatural problems without care. The rules tend

less and less to be considered as real rules, since they are more and more frequently set aside;

a rule that has to be followed only when this gives the same result as if there was no rule at

all (which becomes the case if all nomenclatural change due to a strict appliance of the rules

is liable to be cancelled by the Commission) is no more a rule. Many examples could be given

to support this claim, but, for sake of brevity, we will give only one, that of the recent “sup¬

pression" of the name Rana trilobata Mocquard, 1899.

The name Rana trilobata was given by MOCQUARD (1899: 158, fig. 1) to a frog from Mexico

which was accurately described and figured. BOULENGER (1920) placed this name in the syn¬

onymy of Rana pipiens Schreber, 1782. Later, the name had a rather complex history (see Hillis,

Frost & Frost 1983), but it always remained attached to a frog of the Rana pipiens complex.

On 10 March 1980, one of us (A. D.) wrote to Hobart M. Smith (Boulder) to suggest reexami¬

nation of the holotype of this species, still present in good condition in the Paris Museum, by

a specialist of this difficult group, in order to allow a proper identification of the species. This

was followed by two requests of loan of this specimen, first by John S. Frost (see Hillis,

Source : MNHN, Paris

— 301 —

Frost & Frost 1983), then by Robert G. Webb (see Webb 1991). The holotype being a young

specimen, its proper identification was not easy, but the final identification, as belonging to the same

species as the holotype of Rana megapoda Taylor, 1942 (Webb 1991), seems now clear and undi¬

sputed. The species until then known as Rana megapoda has been the matter of very few publications,

and replacement of its name by the senior synonym Rana trilobata Mocquard, 1899 would not have

caused any serious nomenclatural disturbance. However, Webb (1992) applied to the Commission

to ask “suppression” of the name Rana trilobata in order to “protect” the name Rana megapoda.

This request was followed by a single comment (Smith 1993), which clearly speaks against the

idea that “protection” of the recent and seldom used name Rana megapoda was a matter of high

concern for herpetologists. Despite this poor reaction, the Commission decided to follow the request,

and voted the “suppression” of the name Ram trilobata (ANONYMOUS 1994).

It is noteworthy that the name Rana trilobata cannot at all be considered to have ever been

a nomen oblitum, “since it has been used by several authors, although always in the wrong

sense, during the past 50 years” (HILLIS, FROST & Frost 1983: 74). Furthermore, the erroneous

allocation of this name did not come from its being based on a short, uncomplete or inadequate

description of a specimen now lost, as is the case for many old names, but to the carelessness

of all authors who have worked on Mexican frogs and on American leopard frogs, who failed

to examine the holotype which was publicly known (GUIBÉ 1950) to be kept in the collection

of one of the major natural history Museums in the world, until the curator of this collection

suggested its examination. The “well-known” name Rana megapoda cannot seriously be argued

to be much more known than the name Rana trilobata : to support his claim that “ Rana megapoda

is firmly entrenched in the primary literature”, Webb (1992: 211) only quoted two references

(Smith & Taylor 1948; Hillis, Frost & Wright 1983) to the use of this name for this poorly

known and little studied species of frogs, and in the first of these references, both names Rana

trilobata and Rana megapoda are mentioned. It would probably be impossible to find as many

references of publications where the name Rana megapoda appears as of papers mentioning the

name Rana trilobata, simply because the latter is almost 50 years older and was never “forgotten”

by systematists. Finally, it should be noted that, as stated by Webb (1991: 20) himself, Rana

trilobata and Rana megapoda are only subjective synonyms, based on different specimens from

different populations: the theoretical possibility exists that, despite their morphological similarity,

these populations could be later recognized as distinct species-group taxa.

As a matter of fact, in amphibians but also in many other zoological groups, recent biological

works using modern techniques such as protein electrophoresis or bioacoustics have shown that

species richness in these groups has long been underestimated by zoologists and is much higher

than had been believed in the past. A lot of old names which had once been considered as

synonyms can then be “resurrected” to designate the newly recognized species. This is much

more parsimonious than creating new names for these taxa, all the more that a synonym does

not “disappear” from scientific literature, but still has to be mentioned in synonymies, even if

it was “suppressed” by the Commission (see several examples in DUBOIS & Ohler 1995).

To summarize the discussion above, in 1994 the Commission voted for the “suppression”

of a well-known name, based on a well-known holotype kept in a major Museum, simply to

protect an invalid use introduced and maintained in a small number of publications, for less

than 50 years (1942-1991), by a small number of authors who had not done correctly their work.

Source : MNHN, Paris

— 302 —

This can hardly be argued to be an action taken for the sake of the stability and universality of

nomenclature. Unfortunately, although this example may be somewhat “extreme”, it is not unique.

A strong tendency exists, under the pretext of protecting “usage” rather than priority, to protect the

mistakes made by a few recent authors, especially those from some areas of the world. We strongly

disagree with this tendency and we support Holynski’s (1994) alternative approach.

Recent discussions, e.g. by Holynski (1994), have shown that it is impossible to give an

absolute definition of “current usage”, while “priority” is a simple, international and non-con-

troversial concept. “Current usage” depends on various factors and can, for a given name, change

very quickly when one factor changes (e.g., a group of colleagues publishing within a short

period several works using a name until then unused will make its usage become “current”).

On the other hand, the concept of “priority” relies upon a single uni-directional factor, i.e. time:

any publication date, once determined by careful bibliographic research, can unequivocally, and

independently of other information, be placed and recognized as prior or posterior to another

date. As in zoological nomenclature there is a starting date (1758) and stringent rules for the

availability of names, the number of names is finite and, to tell the truth, in any given zoological

group, rather small. It is fully misleading to claim that “bibliographic archaeologists” will “al¬

ways” uncover senior synonyms of currently used names: final nomenclatural stability will be

achieved when the bibliographic research work is completed, which is a finite and accessible

goal. Stabilization of nomenclature by this way is a true and final stabilization, while stabilization

by the bias of “Opinions” of the Commission is heavy and fragile. The same ideas can be ex¬

pressed in terms of the basic principles of thermodynamics: stabilization through the use of the

rules is definitive and costless, while “stabilization” through interventions of the Commission

is costly in terms of energy and liable to be changed again, for example “every 20 or 30 years”

(Holynski 1994: 14), as fashions and powers change within the small world of institutional

zoology.

As a matter of fact, in many cases, the so-called current “common usage” is simply the

use of a small group of zoologists, especially from “rich” countries which, having more funds

for research, tend to publish more papers than others. We predict that, if the Commission con¬

tinues to “suppress” well-known names, based on well-known holotypes kept in major Museums,

simply to please a small community of persons, it will be ruining the basis of its own existence.

If this tendency continues, the number of cases submitted to the Commission will grow each

year and some future zoologists will be tempted to overlook completely the nomenclatural aspect

of their taxonomic work: nothing is more easy for a “modern” taxonomist using “modern” tech¬

niques than to show that a new species can be recognized, e.g. by a study of biochemical

characters, and described on the basis of these “new characters” without care for the older scien¬

tific literature, and later, after a few years of use of the new name, if another serious taxonomist

points to the existence of an earlier synonym of this name, to ask for the “suppression” of the

latter. Such an extreme evolution of nomenclatural practices is likely to be rejected by a sig¬

nificant part of the international community of zoologists, especially from countries with an old

tradition of taxonomic research. We fear that, if the tendency continues for the Commission to

“suppress” names quickly, without proper discussion in the international community, just to “pro¬

tect the mistakes” of a few authors, this might lead to the end of the international consensus

upon a single system of zoological nomenclature.

Source : MNHN, Paris

— 303 —

NOMENCLATURAL ROLE AND IMPORTANCE OF NAME-BEARING TYPES

(ONOMATOPHORES) AND TYPE-LOCALITIES (TYPE-POPULATIONS)

It has become rather fashionable, in recent days, to express contempt for the early works

of the founders of zoology, for their “incomplete” or “inaccurate” descriptions, and this attitude

has been the basis for the neglect, rejection or “suppression” of many old names.

Of course, problems of allocation of names to biological taxa are more frequent and im¬

portant for ancient names than for names published recently. While some descriptions and il¬

lustrations from the very first days of zoology were “already" remarkably detailed and accurate

(see Dubois & Ohler 1995), it is true that early descriptions were often very brief and im¬

precise, and that type-specimens of many old names have been lost. However, this does not

mean that problems regarding the allocation of these names are of a nature different from

those concerning recent names. In all cases, the important point is the proper allocation to a

biological species of the name-bearing type(s), or, if this/these has/have been destroyed or lost,

of topotypical specimens. The Code is very clear about it: what allows the proper identification

of a nominal species is not its original description, but its original name-bearing type, which

in its turn refers to a type-locality (i.e. to a natural population of the animal species under

consideration).

Despite its misleading name, the role of the name-bearing type is not to provide a “typical”

specimen which should bear all “typical” characters of the species-group taxon it refers to: such

a conception of type-specimens clearly refers to “the typological thinking of essentialism” (Mayr

& ASHLOCK 1991: 11). It is unfortunate that the term “type” was used for a very long period

(more than 150 years) and retained in the current edition of the Code to designate the concept

of “name-bearing type”. It would have been much better to use for this concept another term,

such as Dennler’s (1939) term test or Simpson’s (1940, 1961) term onomatophore, which both

do not include the word “type” and therefore do not provide the “temptation” to view “name¬

bearing types” as “typical” specimens. However, due to the very long use of the term “type”

in nomenclature, it would seem difficult to introduce such a change now.

The major role of the name-bearing type (onomatophore) is not to provide characters (al¬

though it can “incidentally” also do this, especially if it is the only known specimen of the

species), but to provide an objective and non-ambiguous reference to a natural population of

animals. In this respect, it might be better to refer to type-populations rather than to type-lo¬

calities. In practice, it is true that generally the two terms are equivalent and that usually formal

designation of a type-locality is a clear and non-ambiguous designation of a natural population

as “name-bearing type” (or rather "onomatophore”). But there are exceptions to this general

rule. For example, changes may occur in the taxonomic composition of a fauna, which sometimes

can happen in short periods of time. A species may disappear from an area, or its natural range

may change: it may be replaced in the type-locality by another closely related species, but still

occur in other areas, etc. The “type-locality” may then refer to another animal populalion than

that used for the original description. For these reasons, and although this would be a rather

important change, we would favor the replacement, in the next edition of the Code, of the terms

“type-locality” by the terms “type-population”, or, even better although less palatable, “name¬

bearing population” or “onomatophoric population”.

Source : MNHN, Paris

— 304 —

World of language

(zoological nomenclature)

Name-bearing type

(onomatophore)

Real world of

natural populations

Non-type specimen

1 1

Zone of objective connection

between real world and

world of language

Type-population

Nominal taxa

Population not including type

Fig. 1. — The role of name-bearing types (onomatophores) as an objective connection between the real world of natural animal

populations and the world of language (zoological nomenclature). NFI, nominal family I. NGI and NG2. nominal genera

1 and 2. NS1 to NS3, nominal species-group taxa 1 to 3. PI to PS, natural animal populations I to 5. Tl to T3. name-bearing

types 1 to 3.

Source : MNHN, Paris

— 305 —

It is very important to stress that species-group name-bearing types (onomatophores) are

the only objective link between the real world of animals in nature and the world of language,

of scientific names (see Fig. 1). Descriptions, just as names, are part of the world of language,

not of the real world of animals, while name-bearing types are real specimens. A species-group

name is defined by an objective reference, through one or several type-specimens, to a natural

population of animals. In contrast, all other scientific names, i.e. names of higher taxa, rely on

name-bearing types which are names, not specimens. However, through the hierarchical structure

of zoological nomenclature, they ultimately refer also to the species-group type-specimens and

their type-populations (see Fig. 1).

Needless to say, in recent taxonomic works, it is important that description of species be

made with as much care and accuracy as possible: this will facilitate the work of other zoologists

to identify specimens and recognize species. But it should be clear that even very detailed des¬

criptions cannot be the basis for nomenclatural allocation of names. Even very recent descriptions,

as complete and accurate as they may be, may lack some characters which will be found only

later to be of importance for the actual identification of a specimen. Let us take just one example.

The North American tree-frog species Hyla versicolor Le Conte, 1825 and Hyla chrysoscelis

Cope, 1880 were described on the basis of morpho-anatomical characters of adult frogs only.

Later, subsequent studies showed that these species could be distinguished by other characters,

including their mating calls and their ploidy levels (see e.g. DUBOIS 1977). Since diploid and

tetraploid cells have different characteristics, it was possible to ascertain the ploidy of the name¬

bearing type of one of these nominal species, which had important nomenclatural consequences

(for more details, see: Fitzgerald et al. 1981; Smith et al. 1988; Anonymous 1993). The

same could well occur tomorrow concerning a species described today, but for which, at the

time of its description, the characteristics of the mating call or the ploidy level was not known

to be an important identification character.

The number of characters that can be studied and recognized in an animal is virtually

endless. As our knowledge progresses, new characters are regularly discovered. Information on

these characters will not be found in the descriptions, but can be obtained from a posteriori

study of the name-bearing type(s) or of topotypical specimens or populations. Therefore the

emphasis put by the Code on name-bearing types (and, through them, on the natural populations

from which they were collected), rather than on descriptions, is excellent indeed, and is the best

way to achieve the stability of nomenclature.

In conclusion, to ascertain the status of an ancient scientific name, the best and most efficient

way is to study its name-bearing type (onomatophore). This requires several steps:

1, to ascertain which specimens were the original name-bearing types (holotype or syntypes),

i.e. those on which the original description was based;

2, to ascertain if these specimens are still in existence, and where;

3, if they, or part of them, are still in existence, to study them;

4, to ascertain whether stabilization of the status of the name requires the designation of

a lectotype or, in case the holotype or all syntypes have disappeared, of a neotype, and, if this

is necessary, to designate and describe such a lectotype or neotype.

The purpose of the series of papers introduced here is to make this work for a number of

ancient scientific names of Amphibia Anura, in order to definitively stabilize their status. Before

Source : MNHN, Paris

— 306 —

doing this however, it may be useful to remind some of the rules and concepts on which such

a work is based, because some of them seem to be unknown or misunderstood by several recent

taxonomists.

AVAILABILITY OF SCIENTIFIC NAMES

Before discussing the status and allocation of any scientific name, the first question that

must be asked about it is whether this name is “available” in the very precise sense given to

this term by the Code. To be available, a name must, of course, have been published , in the

precise sense given also by the Code to this term, and accompanied by a description or indication.

But this is not enough. The name must also have been published by the author as a new name ,

a name nomenclaturally created in this publication, not a name already used by previous authors

and used again, sometimes under a slightly different meaning, in the publication under con¬

sideration. This may seem obvious but it is not. As a matter of fact, some recent authors tend

to deal with ancient taxonomic texts as if they were contemporary texts (see BOUR & DUBOIS

1984). Nowadays, when a taxonomist uses or quotes a scientific name, he/she usually refers to

the author and date of the original description of this nominal taxon. But in the old days this

was far from being the case. Many authors used a name without mentioning its author and date,

which were often considered to be “obvious”, well-known by all cultured readers. In many other

cases, they mentioned some secondary authors who had used this name but not the primary

author of the name. This was common practice then, and it would be completely inappropriate

to consider that, each time that a taxonomist has used a name without quoting its author, he

has created a new name (which would then be a junior primary homonym of the original name).

In order to know whether an author indeed “created” a name (i.e. a nominal taxon) in a

paper, we need some criteria. These may not be as simple to find as it would seem at first

glance. For example, many ancient authors used the formulae or letters “M/7?/” or “M.”, or

“ Nobis ”, “Nob." or “ N .”, to designate the new taxa that they were creating. But these formulae

cannot be used as non-ambiguous evidence of the creation of a new name, for several independent

reasons:

1, although these formulae were sometimes used to designate brand new taxa, they were

also used to designate new combinations, new spellings, or sometimes simply a modified concept

of the taxon (e.g. a more or less inclusive taxon, a newly defined taxon, etc.);

2, in the old days, it was frequent use for an author to describe as “new” a taxon in several

distinct publications; of course, with our current rules, the taxon must be understood as new

only in the first published work, and it would be completely irrelevant to consider any subsequent

publication of the same name by the same author, even if followed by mihi or nobis, as a new

name (and hence a junior homonym and synonym of the first published name).

Such a comment may seem to be unnecessary to many experienced taxonomists, who are

well aware of this situation, and this question may appear trivial, but it is not. Many taxonomists

in the past have recognized the existence of such unwarranted “junior homonyms”. It is incorrect

to consider that an author had created a new name when he had only modified the content or

Source : MNHN, Paris

— 307 —

definition of a taxon, while keeping for it the name proposed by an earlier author, even when

he did not quote this author, as was often the case then.

A very recent citation will allow to underline these statements: “The description of Hyla

rubra by Laurenti (1768) was based on an illustration [...] in Seba (1735). Daudin (1802)

also used the name Hyla rubra , referred to Seba’s illustration [...]. Daudin (1802; 1803) made

no reference to Laurenti ’s use of the name Hyla rubra, so it must be assumed that Daudin

considered his Hyla rubra to be a new name. Because of Daudin’s reference to the illustration

in Seba, it seems as though both descriptions of Hyla rubra were of the same animal; Laurenti’ s

description was based solely on Seba’s illustration, whereas Daudin’s description was based

on the specimen illustrated by Seba. Therefore, Hyla rubra Daudin, 1802, is a junior objective

homonym (sic) of Hyla rubra Laurenti, 1768.” (Duellman & Wiens 1993: 39-40). This short

citation contains an unusual amount of errors. First, it contains the curious creation of the category

of “junior objective homonym”, which clearly is the result of a lumping between “junior primary

homonym” and “junior objective synonym”. Second, if Daudin was indeed the author of a name

Hyla rubra, the correct citation for this authorship would be “Daudin, 1800a”, not “Daudin,

1802”: actually Daudin used the name Hyla rubra twice before his 1802 book, once in his

1800a booklet and once in SONNINI & LATREILLE’s 1801 book (for the respective publication

dates and authorship of these works, see Harper 1940). Third, Duméril & Bibron (1841: 595)

stated that the specimen illustrated and described by DAUDIN was not the specimen illustrated

by Seba, so that if it were true that Daudin had created a new nominal taxon, this would not

be an objective synonym of Laurenti’s name. But finally and more importantly, this citation

well exemplifies the mistake outlined above concerning junior primary homonyms. Of course,

Daudin (1800a, 1802, 1803; and in Sonnini & Latreille 1801) did not create a new name,

but used Laurenti’s name Hyla rubra, as shown by the fact that, in the synonymy of this name,

he mentioned not only Seba’s (1735) plate, but also Daubenton’s (1782: 668) and LacÉPÈDE’s

(1788: 327) works, who both clearly referred to Laurenti’s (1768) description. It is important

to underline such an error, because, if the same line of reasoning was to be generalized among

taxonomists, we would suddenly have to recognize hundreds, thousands or even millions of such

“junior objective homonyms”.

This does not mean that junior primary homonyms were not created in these old days. But

we need some criteria to recognize them. For reasons outlined above, absence of citation of the

original author and date of a name cannot be such a criterion, nor can the use of formulae like

mihi or nobis. Three major citeria can be pointed out:

Cl, when an author states in full words that he is creating a new name for a new taxon;

C2, when he states in full words that the name he is using is the same as the name used

by a previous author, but that he uses it with a different meaning, for another taxon;

C3, when the ancient name N is quoted in the synonymy of a name A, but, in the same

work, the same name N is applied as valid to another taxon.

Case Cl is very widespread and does not need to be exemplified. An example of case C2

is provided by Daudin (in Sonnini & Latreille 1801: 154), who wrote, in the text under the

name Rana pipiens: “Il ne faut pas rapporter à cette espèce le rana pipiens de Schreber, qui

est le synonyme de la grenouille criarde.” In other words, Daudin in this text considered the

name Rana pipiens Schreber, 1782 as a synonym of his Rana clamitans (“grenouille criarde”).

Source : MNHN, Paris

— 308 —

but he used the name Rana pipiens for another species, thus creating a junior primary homonym.

Finally, an example of case C3 is provided by Daudin (1802: 29-30) who included the name

Hyla lactea Laurenti, 1768 in the synonymy of his Hyla hypocondrialis, but used the name Hyla

lactea for another distinct species, which he had already described earlier (Daudin 1800b: 19):

in this case the name Hyla lactea Daudin, 1800b (not “1803” or “1802", as written in error

respectively by Duellman & LYNCH 1981 and by FROST 1985), is indeed a distinct nominal

species, which is a junior primary homonym of Hyla lactea Laurenti, 1768.

DISTINCTION BETWEEN DIFFERENT KINDS OF NAMES OR SPELLINGS

Once a scientific name has been published, it can be used again by other authors. As a

rule, ancient authors used names without changing them, but in some cases, and rather often in

older times of taxonomy, they modified their spelling, and sometimes they even replaced a name

by a different, new name, for the same taxon. In order to correctly deal with nomenclatural

problems, it is very important to distinguish between the following categories of new names

and spellings: new replacement names, justified and unjustified emendations, incorrect subsequent

spellings. One of us has already devoted a long and detailed discussion to this question (DUBOIS

1987: 31-48), to which we refer our readers: this paper contains in particular proposals of criteria

to distinguish the various categories of names and spellings, illustrated by many examples.

THREE CATEGORIES OF SYNTYPES

As already mentioned above, in many cases the best (or only) way to stabilize the status

of an ancient species-group name is through the study of its original name-bearing type: this

may be either a single specimen (the holotype ) or a series of specimens (the syntypes) 4 , in the

latter case, any subsequent author is entitled to designate among them a lectotype , which becomes

then the unique name-bearing type (the other syntypes then become paralectotypes and lose their

status of name-bearing types). In many cases, the name-bearing types of ancient nominal species

have been lost or destroyed and are not available for study. In such cases, the Code allows for

the possibility, if this is necessary to clarify and stabilize the status of a nominal taxon, to

designate a neotype for it. The latter should come from the original type-locality or very close

to it. Fig. 2 gives diagrammatic representations of the various kinds of name-bearing types and

the relationships they imply between natural populations and nominal species-group taxa.

Which specimens are the syntypes of a nominal taxon? In many ancient publications, the

author of a new name did not clearly state which specimens he considered to be the syntypes

of the new nominal taxon, and this must be reconstructed a posteriori through the help of external

evidence (Recommendation 72B of the Code). According to the Code, the original type-series

of a nominal species-group taxon can be composed:

Source : MNHN, Paris

309 —

FIG. 2. — Differenl kinds of name-bearing types (onomaiophorcs) in zoology. HT, holotype. LT, leclolype. NS. nominal species.

NT. neolype. PI lo P3. natural animal populalions I lo 3. PLI lo PL5, paralectotypes I to 5. ST1 to ST6. syntypes 1 to 6.

A. designation of a holotype (HT) in the original description of a species-group taxon gives an objective and non-ambiguous

reference to a single natural population of animals as "type-population". B, if the holotype is lost or destroyed, a neotype

(NT) may be described, but it should come from the same population as the holotype (or from a population as close as

a holotype. all these specimens are syntypes (ST) and they collectively bear the name. All the populations where these

specimens were collected share the status of type-populations. D, designation among the syntypes of a leclolype (LT) defini¬

tively suppresses the status of name-bearing types of all other syntypes, which become paralectotypes (PL) and lose per¬

manently their nomenclatural function regarding the species-group name concerned. This restriction also serves as a restriction

of the type-population to the population from which the lectotype was collected. E. if the lectotype is lost or destroyed, a

neotype may be chosen, but here also it must come from the same population as did the lectotype. The neotype may be

either one of the remaining paralectotypes or another specimen (even if some of the paralectotypes are still in existence).

Source : MNHN, Paris

— 310 —

1, either of specimens which had been examined, described and/or illustrated by the author

himself;

2, or of specimens which were not available to him but which had been examined, described

and/or illustrated by a previous author, in a work explicitly referred to in the original description;

3, or both.

In this respect, we think that matters will be made clearer by introducing here a distinction

between three categories of syntypes:

1, we call primary syntypes of a nominal species-group taxon T the specimens which had

been examined, described and/or illustrated by the author A of the original description D him/her¬

self;

2, we call secondary syntypes the specimens which had not been examined, described and/or

illustrated by the author A, but by a previous author B in an earlier work W quoted in the

original description D as a basis for the new taxon T;

3, we call tertiary syntypes specimens which had been examined by neither authors A and

B. but by a still earlier author C, quoted by author B in the work W quoted by A in the original

description D of the new taxon T.

To make these definitions clearer and to illustrate in detail these three categories, we have

devoted the second paper of this series (Dubois & Ohler 1996) to a thorough analysis of the

status of the syntypes of the species Rana arborea Linnaeus, 1758.

Finally, we want to stress that all specimens listed in the original description of a nominal

species are not always syntypes. Article 72(b) of the Code gives important particulars about the

criteria to be used to ascertain which specimens are and are not syntypes. Particularly important

is the following statement in Article 72(b)(i): “The type series of a nominal species-group taxon

consists of all the specimens eligible to be name-bearing types (...) included by the author in

the new nominal taxon, except any that the author expressly excludes from the type series (...),

or refers to as distinct variants (e.g., by name, letter, or number), or doubtfully attributes to the

taxon.” Therefore, specimens for which the author of a new nominal taxon expresses doubts

concerning their allocation to the taxon are not syntypes, and the same applies to specimens

referred in the original description to particular varieties (under indications such as: “var.",

“var. 8”, “var. B”, etc.). This rule has been overlooked by several recent authors, as we will see

in the following papers of this series.

LECTOTYPE DESIGNATIONS FOR SPECIES DESCRIBED

IN ANCIENT TAXONOMIC LITERATURE

A special, but not uncommon, situation exists when only some of the original syntypes of

a nominal species-group taxon are still in existence, the other ones having been lost. This is for

example the case of many of the species described by LINNAEUS (1758, 1761, 1766, 1767).

Should the subsequent authors feel then obliged to choose a lectotype among the remaining

syntypes? This is what a vast majority of authors until now have felt obliged to do, although

the Code does not at all require to do so. It is wrong, within the frame of the current Code, to

state that the fact that some (or all) of the original syntypes of a nominal species have been

lost or destroyed prevents from the possibility to designate one of them as lectotype. This is

Source : MNHN, Paris

— 311 —

underlined by Articles 72(c)(v) and 74(c) of the Code, which state that designation as lectotype

of a specimen described or illustrated in an old work is valid, even if this specimen cannot be

traced now. This designation must be understood as designation as name-bearing type of the

specimen illustrated or described, “not the illustration or the description”.

For old names which pose problems of allocation to biological species, this procedure has

very strong advantages, which have not been clearly pointed out by previous authors. Let us

make them clear by using a precise example, taken from our recent work on European green

frogs (Dubois & Ohler 1995).

Linnaeus (1758: 212) described Rana esculenta without stating the number and origin of

specimens he had before him. The type-locality he gave for this species was very vague: “Habitat

in Europae fontibus”. If we only had this information, the name esculenta could be available

for any of the European green frogs. Furthermore, according to LÔNNBERG (1896) and ANDERS-

SON (1900), the only original type-specimen of this species still in existence would appear to

belong to the brown frog species now known as Rana arvalis Nilsson, 1842 (see LÔNNBERG

1896: 35) ! Designation of the latter specimen as lectotype of Rana esculenta would have dramatic

nomenclatural consequences. Fortunately, the current Code does not oblige us to do this. In the

original description of Rana esculenta, LINNAEUS (1758: 212) made clear reference to Roesel

von Rosenhof’s (1758) description and figures of “Rana viridis aquaticae". Therefore, the speci¬

mens used by Roesel von ROSENHOF for his text and figures must also be considered (secondary)

syntypes of Rana esculenta. Designation of one of the figured or described specimens as lectotype

of this species (DUBOIS & Ohler 1995) is valid under the current Code. These rules have the

following nomenclatural consequences in this case:

1, by the designation as lectotype of Rana esculenta of one of the specimens figured by

Roesel von ROSENHOF (1758), all other syntypes of this nominal species (including the “ Rana

arvalis ” specimen) immediately and definitively lose their status of syntypes and cannot any

more threaten the stability of nomenclature;

2, the type-locality of Rana esculenta, which was “Europe” before this designation, becomes

definitively restricted to the locality where the figured specimen had been collected (Nürnberg,

Germany);

3, many external morphological and color characters of this specimen can be seen on this

figure, which in this case allows a non-ambiguous allocation of the name (furthermore, the text

gives detailed information on the developmental biology of these frogs which confirms the iden¬

tification).

Now, let us go further and suppose that the figure does not provide enough information

for an unequivocal allocation of the name, and that the illustrated specimen cannot be traced

now. It would then be possible to claim that we are in a case where the circumstances allow

the designation of a neotype (Article 75 of the Code), and this designation can be made in a

last step (4). But it cannot be made anyhow: the neotype must come from the restricted type-

locality or as close to it as practicable (Article 75(d)(5) of the Code).

This process can therefore be described more generally as follows. The designation as lec¬

totype of a figured or described but lost specimen can play the roles (1) and (2) above: “elimi¬

nation” of all other syntypes and fixation of the type-locality. In a step (3), close examination

of the figure and/or description may be enough to stabilize the status of the name. If not, in a

Source : MNHN, Paris

-312-

step (4), designation of a neotype from the restricted type-locality may be necessary to complete

the work of fixation of the status of a name.

We think that this process, which is described in detail here for the first time, is liable to

solve many, if not almost all, cases of old names without proper allocation to biological species.

We illustrate this process in full detail in our second paper of this series (Dubois & Ohler

1996), and, in subsequent papers of this series, we will use it more rapidly, as we think its

principles will have been clarified.

The three categories of syntypes recognized above have, in our view, different status re¬

garding their possible choice as lectotypes. Of course, still extant primary syntypes, which are

known to have been in the hands of the author when he described the new species, and which

are still kept in a Museum collection, should be preferred for lectotype designation when their

choice does not raise nomenclatural problems. But in some cases, as exemplified for the names

Rana esculenta (DUBOIS & OHLER 1995) or Rana arborea (DUBOIS & OHLER 1996), choice of

a still extant primary syntype as lectotype would lead to dramatic nomenclatural consequences.

In such cases, it may be better to choose a secondary syntype. which was not (or may not have

been) seen by the author of the name, but which was described or figured in a publication seen

by him/her. Only in extreme situations, when both primary and secondary syntypes do not belong

to the biological species currently known under the name under consideration, and when change

of this name would have disastrous consequences, should a tertiary syntype be chosen as lec¬

totype.

Finally, it should be stressed that the choice of a lectotype should be made after a careful

analysis of the nomenclatural situation of the name considered. As underlined above, not all

“old names” are “well-known”. Some, like Rana temporaria Linnaeus, 1758, are, while others,

sometimes appearing in the same ancient publication, such as Rana hyla Linnaeus, 1758 (see

Dubois & Ohler 1996). are not. The treatment of these two kinds of names should be different.

While for “well-known” names it is important to do the best to stabilize the use of the name

in its current, usual, or more common, sense, the same is not true for names which have been

almost or completely forgotten for dozens of years or for centuries. Allocation of the latter names

to biological taxa, through lectotype designation, should be done, if possible, in order not to

disturb the current nomenclature of the group where they belong. Therefore, if possible, in the

case of a “well-known” old name, designation of a lectotype (or, if necessary, of a neotype)

should be done in order to maintain the current use of the name, while in the case of an “obscure”

old name, it should be done in order to lead to the disappearance of this name as a junior

objective or subjective synonym. A junior objective synonym will remain permanently invalid,

but a junior subjective synonym is liable, in some cases, to be resurrected as the taxonomy of

a group progresses and more species-group taxa have to be recognized.

TYPE-LOCALITY RESTRICTIONS

According to Article 72(h) of the Code, the type-locality of a nominal species or subspecies

is the geographical place of capture or collection of its name-bearing type. When a species was

described from several syntypes from distinct localities, restriction of the type-locality to one

Source : MNHN, Paris

— 313 —

of these localities can be obtained only through the designation of a lectotype among them, or

of a neotype if all original syntypes have been destroyed or lost (see Fig. 2). So-called “restric¬

tions of type-localities”, not accompanied by formal designations of lectotypes or neotypes, are

invalid according to the Code. Many such invalid “restrictions” have been done by various authors

in the past. In many cases, it is possible to “make them valid a posteriori” through the designation

of a lectotype or of a neotype, and we think that, for the sake of stability, this should be done

as much as possible. But in some other cases, this is not possible, and type-localities will have

to be corrected in order to fit with the lectotype or neotype designation.

RELATIVE PRIORITY BETWEEN NAMES PUBLISHED SIMULTANEOUSLY

This will be the last comment of this introductory paper. Not infrequently, several scientific

names published simultaneously, either by the same or by different authors, are later considered

to apply to a single biological taxon. In such cases, according to the Code , the valid name

among them is determined by the action of the “first reviser”, i.e. the first author to have quoted

both names and chosen the order of priority between them. Such first reviser actions may be

difficult to trace, especially as concerns old names. In the subsequent papers of this series, we

will try to trace all such actions concerning the names discussed, and furthermore, in order to

avoid similar problems in the future, we will take such first reviser actions in all cases when

they have not been done yet.

REFERENCES

ANONYMOUS, 1961. — International Code of Zoological Nomenclature. First edition. London, International Trust

for zoological Nomenclature: i-xviii + 1-176.

— 1964. — International Code of Zoological Nomenclature. Second edition. London, International Trust for

zoological Nomenclature: i-xx + 1-176.

— 1985. — International Code of Zoological Nomenclature. Third edition. London, International Trust for

zoological Nomenclature: i-xx + 1-338.

— 1993. — Opinion 1716. Hyla chrysoscelis Cope, 1880 (Amphibia, Anura): specific name conserved by the

designation of a neotype. Bull. zoo!. Norn. 50 (1): 94-95.

— 1994. — Opinion 1776. Rana megapoda Taylor, 1942 (Amphibia, Anura): specific name conserved. Bull,

zool. Norn. 51 (2): 181.

ANDERSSON L. G., 1900. — Catalogue of Linnean type-specimens of Linnaeus's Reptilia in the Royal Museum

in Stockholm. Bihang K. Svenska Vet.-Akad. Handlingar 26 (4): 1-29.

BÔHME W. & WiEDL H., 1994. — Status and zoogeography of the herpetofauna of Cyprus, with taxonomic and

natural history notes on selected species (genera Rana, Coluber, Natrix, Vipera). Zool. Middle East 10:

31-52.

BOULENGER G. A., 1920. — A monograph of the American frogs of the genus Rana. Proc. amer. Acad. Arts

Sci. 55: 413-480.

BOUR R. & DUBOIS a., 1984. — Nomenclatural availability of Testudo coriacea Vandelli, 1761: a case against

a rigid application of the Rules to old, well-known zoological words. J. Herpet. 17: 356-361.

DAUBENTON M., 1782. — Les animaux quadrupèdes ovipares, et les serpens. In Encyclopédie méthodique -

Histoire naturelle des animaux. Tome premier, Paris, Panckoucke : 545-712.

Source : MNHN, Paris

— 314 —

Daudin F. M., 1800a. — Histoire naturelle des quadrupèdes ovipares. Livraison 1. Paris, Fuchs, Delalain, Treuttel

& Würtz : [1-11], pi. [I-Vl],

— 1800b. — Histoire naturelle des quadrupèdes ovipares. Livraison 2. Paris, Fuchs, Delalain, Treuttel &

Würtz: [13-24], pl. [VII-XII].

— 1802. — Histoire naturelle des rainettes, des grenouilles et des crapauds. Paris, Levrault : 1-108, pl. I-

XXXVIII.

— 1803. — Histoire naturelle, générale et particulière des Reptiles. Tome huitième. Paris, Dufart : 1-439, 8 pl.

Dennler J. G., 1939. — La importancia de la distribution geogrâfica en la sistemâtica de los vertebrados. Physis

16: 41-53, pl. I-Vm.

DUBOIS A., 1977. — Les problèmes de l’espèce chez les Amphibiens Anoures. Mém. Soc. zool. France 39 :

161-284.

— 1981. — Liste des genres et sous-genres nominaux de Ranoidea (Amphibiens Anoures) du monde, avec

identification de leurs espèces-types : conséquences nomenclaturales. Monit. zool. irai., (n. s.) 15, suppl. :

225-284.

— 1984. — La nomenclature supragénérique des Amphibiens Anoures. Mém. Mus. nail Hist, nat., (A) 131: 1-64.

— 1987. — Again on the nomenclature of frogs. Alytes 6: 27-55.

— 1994. — Comment on the proposed conservation of Hemidactyliini Hallowed, 1856 (Amphibia, Caudata).

Bull. zool. Norn. 51: 264-265.

— 1995a. — Discussion Draft of the Fourth Edition of the International Code of Zoological Nomenclature:

comments. (3). Bull. zool. Norn. 52: 299.

— 1995b. — Comments on the proposed conservation of Hemidactyliini Hallowed, 1856 (Amphibia, Caudata).

Bull. zool. Norn. 52: 337-338.

Dubois A., BOUR R., Brygoo E.-R., LESCURE J., Bouchet P. & Tillier S., 1988. — Comments on the proposed

suppression for nomenclature of three works by R.W. Wells and C.R. Wedington. (2). Bull. zool. Norn. 45:

146-149.

DUBOIS a. & OHLER a., 1995. — Frogs of the subgenus Pelophylax (Amphibia, Anura, genus Rana): a catalogue

of available and valid scientific names, with comments on name-bearing types, complete synonymies, pro¬

posed common names, and maps showing ad type localities. Zool. Polon. (1994) 39 (3-4): 139-204.

— 1996. — Early scientific names of Amphibia Anura. II. An exemplary case: Rana arborea Linnaeus, 1758.

Bull. Mus. not! Hist, nat., 4 e sér., A 18 (3-4): 321-340.

DUELLMAN W. E. & LYNCH J. D.. 1981. — Nomenclatural resolution of the identities of Hyla aurantiaca and

Hyla lactea. J. Herpet. 15: 237-239.

DUELLMAN W. E. & Wiens J. J„ 1993. — Hylid frogs of the genus Scinax Wagler, 1830, in Amazonian Ecuador

and Peru. Occ. Pap. Mus. nat. Hist. Univ. Kansas 153: 1-57.

DUMÉRIL A.-M.-C. & BlBRON G., 1841. — Erpétologie générale ou histoire naturelle complète des Reptiles.

Tome 8. Paris. Roret : i-vii + 1-792.

Fitzgerald K. V., Smith H. M. & Guillette L. J. Jr., 1981. — Nomenclature of the diploid species of the

diploid-tetraploid Hyla versicolor complex. J. Herpet. 15 (3): 356-360.

FROST D. R. (ed.), 1985. — Amphibian species of the world. Lawrence, Aden Press & Assoc. Syst. Coll.: [i-iv]

+ i-v + 1-732.

GUIBÉ J., 1950. — Catalogue des types d'amphibiens du Muséum national d'Histoire naturelle. Paris, Imprimerie

nationale: 1-71.

Hammond P., 1992. — Species inventory. In B. Groombridge (ed.). Global biodiversity - Status of the earth

living resources. London, Chapman & Had: 17-39.

Harper F., 1940. — Some works of Bartram, Daudin, Latreille, and Sonnini. and their bearing upon North

American herpetological nomenclature. Amer. Midi. Nat. 23 (3): 692-723.

HlLLlS D. M., FROST d. R. & FROST j. S., 1983. — Allocation and distribution of Rana trilobata Mocquard. J.

Herpet. 17 (1): 73-75.

HlLLlS D. M., FROST J. S. & Wright D. A., 1983. — Phylogeny and biogeography of the Rana pipiens complex:

a biochemical evaluation. Syst. Zool. 32 (2): 132-143.

HOLYNSKI R. B., 1994. — Structure and function or: what kind of nomenclatural regulations do we need? Crystal,

God, Hungary, (ser. Zool.) 2: 1-50.

Source : MNHN, Paris

— 315 —

LacÉPÈDE [B. G. E.] DE, 1788. — Histoire naturelle des quadrupèdes ovipares et des serpens. Vol. II, 16 cm.

Paris. Hôtel de Thou : [i-iv] + 1-464, pl. I-XV.

LAURENTI J. N., 1768. — Specimen medicum. exhibe ns synopsin Reptilium emendatam cum experimentis circa

venena et antidota Reptilium austriacorum. Viennae, Joan. Thom. Nob. de Trattnern: i-ii + 1-215, pl. I-V.

Linnaeus C. A., 1758. — Systema Naturae per régna tria naturae, secundum classes, ordines, genera, species,

cum characteribus, differentiis, synonymis, locus. Editio décima, reformata. Tomus I. Holmiae: i-iv + 1-824.

— 1761. — Fauna Suecica. Editio altera, auction Stocholmiae, Laurentii Salvii: [i-xl] + 1-579, pl. I-II.

— 1766. — Systema Naturae per régna tria naturae, secundum classes, ordines, genera, species, cum charac¬

teribus. differentiis. synonymis. locis. Editio duodecima, reformata. Tomus I, Pars I. Holmiae, Laurentii

Salvii: 1-532.

— 1767. — Systema Naturae per régna tria naturae, secundum classes, ordines, genera, species, cum charac¬

teribus. differentiis. synonymis, locis. Editio duodecima, reformata. Tomus I, Pars II. Holmiae, Laurentii

Salvii: 533-1327 + [i-xxxvii],

LÔNNBERG E., 1896. — Linnean type-specimens of birds, reptiles, batrachians and fishes in the Zoological

Museum of the R. University in Upsala. Bihang K. Svenska Vet.-Akad. Handlingar 22 (4): 1-45.

Mayr E. & ASHLOCK P. D., 1991. — Principles of systematic zoology. Second edition. New York, McGraw-Hill:

i-xx + 1-475.

MOCQUARD F. 1899. — Reptiles et batraciens recueillis au Mexique par M. Léon Diguet en 1896 et 1897. Bull.

Soc. philom. Paris (9) 1 : 154-169, pl. I.

ROESEL VON ROSENHOF A. I., 1758. — Historia naturalis ranarum nostratium. Nürnberg, Johann Joseph Fleisch-

mann: [i-viii] + i-viii + 1-116, pl. I-XX1V + frontispiece.

Seba A., 1735. — Locupletissimi rerum naturalium Thesauri accurata descriptio, et iconibus artificiosissimis

expressio. per universam physices historiam. Tomus II. Amstelaedami, Janssonio-Waesbergios, Wetstenium

& Smith: [i-xxxiv] + 1-154, pl. I-CXIV.

SIMPSON G. G., 1940. — Types in modem taxonomy. Amer. J. Sci. 238: 413-431.

— 1961. — Principles of animal taxonomy. New York, Columbia Univ. Press: i-xii + 1-247.

SMITH h. m., 1993. — Comment on the proposed conservation of the specific name of Rana megapoda Taylor,

1942 (Amphibia, Anura). Bull. zool. Norn. 50 (1): 57-58.

Smith H. M., Fitzgerald K. V. & Guillette L. J. Jr., 1988. — Case 2366. Hyla chrysocelis Johnson, 1961

(Amphibia, Anura): proposed conservation and designation of a neotype. Bull. zool. Norn. 45 (2): 138-140.

Smith H. M. & Taylor E. H., 1948. — An annotated checklist and key to the Amphibia of Mexico. Bull.

U. S. nat. Mus. 194: i-iv +1-118.

SMITH H. B. & Wake D. B., 1993. — Case 2869. Hemidactyliini Hallowell. 1856 (Amphibia, Caudata): proposed

conservation. Bull. zool. Norn. 50: 129-132.

— 1994. — Comment on the proposed conservation of Hemidactyliini Hallowell, 1856 (Amphibia, Caudata).

Bull. zool. Norn. 51: 341-342.

— 1995. — Comment on the proposed conservation of Hemidactyliini Hallowell, 1856 (Amphibia. Caudata).

Bull. zool. Norn. 53: 48-49.

SONNINI C. S. & LATREILLE P. A., 1801. — Histoire naturelle des reptiles. Tome II. Paris. Deterville : [i-iii] +

1-332, 19 pl.

WAKE D. B., 1993. — Phylogenetic and taxonomic issues relating to salamanders of the family Plethodontidae.

Herpetologica 49: 229-237.

Webb R. G., 1991. — The identity of Rana trilobata Mocquard (Anura: Ranidae). Herpetologica 47 (1): 13-21.

— 1992. — Case 2821. Rana megapoda Taylor, 1942 (Amphibia, Anura): proposed conservation of the specific

name. Bull. zool. Norn. 49 (3): 211-212.

Source : MNHN, Paris

— 316-

APPENDIX I

In the paper above, we made several references to a paper by HOLYNSKl (1994), which in

our view is a major contribution to the current international discussion on the Code , and in

particular on the problem of “usage versus priority”. Clearly the author of this paper does not

share the ideas which are accepted by most current members of the Commission, and, although

this paper was submitted in February 1991 to the Commission’s Secretary for publication in the

Bulletin of zoological Nomenclature, it was never published in this journal, without any answer

or explanation from the Commission’s Secretary. Finally this paper was published in the Hungar¬

ian journal Crystal. Unfortunately, this recent journal still does not have a large distribution

worldwide, and many colleagues may find difficult to trace a copy of it. We suggest that interested

colleagues order a copy of this paper by writing at the following address: Natural Science Foun¬

dation, God, Koros u. 15, 2131, Hungary. It would be useful if a major zoological journal with

large distribution worldwide decided to publish a complete reprint of this long paper, or, better

than nothing, important extracts from it. We provide below a copy of the Introduction of this

very stimulating paper:

“The goal of the International Code of Zoological Nomenclature is - among others - sta¬

bility of nomenclature. In my opinion, the Code acquits itself of this very difficult task surpris¬

ingly well: except for some taxonomically neglected groups the nomenclature is remarkably

stable, the rate of formal (i.e. not caused by taxonomical rearrangements) changes of names

remains within few percents in a century ! This splendid efficiency of the Code (together with

its comprehensively tested capability to make nomenclatural decisions easy and unequivocal; its

‘automatic action’, rendering any time-bureaucracy unnecessary; and - last not least - profoundly

fair, democratic character of its guiding principle, the Principle of Priority) seems to provide a

very strong argument for letting it work. Of course, nothing in this world is absolutely perfect,

so some modifications of the Code, as well as some exceptions from its rulings, are sometimes

warranted, but one would expect the modifications to be directed towards making the decisions

still more automatic, unequivocal and fair, and the exceptions limited to very rare cases of obvious

inevitability. To my great surprise, the opposite tendency dominates: many authors make repeated

efforts to upset the Code, or at least to obscure its regulations by introducing innumerable ab¬

rogations - setting aside validly designated types, changing dates of publication, ‘protecting’

some names, invalidating others, etc.; what is ‘in principle’ a perspicuous, logical system, be¬

comes impenetrable jungle of exceptions and special cases! On the following pages I wish to

present more detailed comments on some - in my opinion especially dangerous - recent pro¬

posals, discuss main general aspects of nomenclatural regulations, and, at last, suggest several

minor modifications which, in my intention, should further improve the above-mentioned merits

of the Code." (HOLYNSKl 1994: 4).

Source : MNHN, Paris

-317-

APPENDIX II

In the paper above, we suggest that some conditions be necessary for a name to be eligible

for “protection” by the Commission. We have already made specific suggestions in this respect,

first in a letter sent on 9 April 1991 by one of us (A. D.) to Otto Kraus (Hamburg), President

of the Commission, letter to which no answer on this point was given, and then in a paper sent

on 23 June 1994 to the secretariat of the Commission, of which only a brief abstract has been

published in the Bulletin of zoological Nomenclature (Dubois 1995b), and which is reproduced

below as it was submitted.

Proposals concerning the conditions needed for

a name being eligible for conservation

by Alain DUBOIS

Abstract. — New proposals are made concerning the conditions given in Article 79 for a name being

eligible for conservation: 1. the forgotten senior synonym recently rediscovered should not have been used by

a single author during a 50-year period before the date of publication of its “rediscovery”; 2, during the same

period, the junior synonym should have been used by at least 25 independent authors and in at least 100 publi¬

cations. A working definition of "independent authors” is provided. Other proposals are briefly discussed.

1. One of the basic principles of the Code is the rule of priority. Its consistent use for

roughly a century by almost all zoologists has had a considerable impact on zoological nomen¬

clature: in the overwhelming majority of cases, use of this rule has had the effect of stabilizing

nomenclature in the whole world. This basic rule therefore answers the major aims of the Code,

i.e. to ensure the universality and stability of zoological nomenclature. It is therefore beyond

all doubts that this rule should be maintained.

2. However, as pointed out, especially in the recent years, by various authors, in some

cases the “rediscovery” of a long “forgotten” name (simply forgotten, in most cases, because

subsequent authors have not done their work properly), may in some cases have a real disrupting

effect on zoological nomenclature. As was very aptly discussed recently by Holynski (1994),

the importance of this risk has been greatly overstressed by several recent authors, those who

are in favor of replacing the rule of priority by a “rule of usage”. My aim in this paper is not

to discuss in detail the idea of such a “rule of usage”, which has been done already by many

others (e.g. recently: NG 1991; Holynski 1994): in my opinion, such a rule would have so

many drawbacks that it cannot seriously challenge the rule of priority; if it were to replace the

latter in the next edition of the Code, no doubt this would cause major problems regarding the

current international consensus of zoologists to follow the Code and to recognize the competence

of the Commission in nomenclatural matters. My aim here is only to discuss the following ques-

Source : MNHN, Paris

— 318 —

tion: when can we seriously consider that the rediscovery of a “forgotten” name causes genuine

problems of stability in zoological nomenclature? In other words, what is the meaning of the

word “stability”?

3. In the Code currently in force, the following conditions are given (Art. 79) for a name

to be eligible for conservation by the Commission, through use of the Plenary Powers: the forgot¬

ten name must not have been used by a single author as a valid name during the 50 years

immediately preceding the request for action of the Commission; during the same period, the

junior synonym must have been used as a valid name by at least 5 different authors and in at

least 10 publications.

4. In my opinion, these rules are far too latitudinarian. In the recent years, on the basis of

these rules, the number of names submitted to the Commission for conservation has been regularly

progressing, and it will continue doing so in the future if these rules are maintained. In fact,

suppression of earlier synonyms and conservation of junior synonyms constitute a large proportion

of cases dealt with by the Commission and of pages published in the Bulletin of zoological

Nomenclature. As remarked by Holynski (1994), in the overwhelming majority of these cases,

once a name has been submitted for conservation to the Commission and the application put to

vote, the Commission follows this suggestion, and uses its Plenary Powers to suppress an older

synonym.

5. I fully agree that a name that has obtained general acceptance among zoologists, i.e.

that has been used by many of them in many publications and during a long period, should be

liable to be considered for protection, through use of the Commission’s Plenary Powers to

suppress a recently “rediscovered” “forgotten” name. Such a need is particularly clear for animal

species that have been the matter of very numerous scientific publications, including most species

of domestic animals, species widely used in experimental biology, often studied in population

biology, etc. However, I am strongly opposed to consider as widely used and of general accept¬

ance a name mentioned by a few dozens authors in a few dozens publications and during a few

dozens years, especially when these authors are taxonomists and when the name has never or

almost never appeared in non-taxonomic literature (publications in other fields of biology, popular

literature, etc.). Let us discuss separately these four aspects (numbers of authors and of publi¬

cations, kinds of publications, period).

6. Considering the number of authors, I feel that the number of authors having used the

junior synonym should be at least 25, if not 50. Furthermore, I suggest to add the following

qualification: “at least 25 independent authors”. This suggested rule is meant to address the fact

that authors who are working together, in a single research team or laboratory, or as collaborators

from different units, are likely to adopt a single nomenclature for the animals they study: if this

research team is numerous, prolific or financially healthy, it will tend to publish numerous papers

on these animals, and the local nomenclatural uses or habits of this team may tend to be presented

as the “common usage” of all zoologists, which may be wrong. Of course, there would be no

point in trying to track down research teams by administrative or police methods, and we need

an objective criterion to identify non-independent authors. I suggest to use co-authorship of pub¬

lished papers: for the purpose of this problem, any couple or team of authors who have co-signed

Source : MNHN, Paris

— 319 —

at least one paper on an animal group should be considered as a single independent unit regarding

the nomenclature of this group, and this unit should appear only once in the minimum list of

25 independent authors. Therefore, “independent authors” in this context will only refer to authors

who never published together on this group before the application to the Commission. Zoologists

submitting such an application should be invited to follow this rule; in case they ignored it, any

other zoologist would be entitled to point to publications that show that some of the authors of

the list submitted were not “independent” in the sense of this proposed ruling, which would

reduce accordingly the number of authors in support of the application.

7. The next criterion is the number of publications using the invalid junior synonym. I

suggest to raise this number to 100. In contrast to the preceding suggested rule, this allows to

take into account several publications of a single research team. But here again I suggest to

add an important qualification: this should include only publications that appeared before the

date of publication of the “ rediscovery” of the “forgotten" senior synonym. This is of course

to avoid that “current usage” be deliberately rigged or manipulated in order to support one’s

interpretation.

8. Finally, as concerns duration of the period during which: 1, the senior synonym has not

been used by a single author and 2, the junior synonym has been used by various authors, I

suggest to keep the number of 50 years, but, here also, to add the qualification before the date

of publication of the “rediscovery” of the “forgotten” senior synonym.

9. The numbers suggested above (25 independent authors, 100 publications, 50 years) seem

to me to be minimum values for the phrases “current usage” and “nomenclatural stability” to

have a real meaning. These figures might even have to be increased in some cases, if a distinction

was to be introduced between different kinds of names: for example, one might consider having

these figures for species-group names, but higher figures for genus-group names (e.g. 50 inde¬

pendent authors, 200 publications, 50 years) and still higher figures for family-group names (e.g.

100 independent authors, 500 publications, 50 years). Another distinction might be worth con¬

sidering: that between taxonomic and non-taxonomic publications. When a name has been used

only or almost only in taxonomic publications, the only biologists that are really acquainted

with this name are taxonomists. If discovery of a senior “overlooked” synonym occurs, all tax¬

onomists can understand the reason why the junior name must be replaced by the senior one,

and no real disturbance of their work follows this change. Of course, the disturbance may be

real when non-taxonomists are at stake. For this reason, it might be good to consider only non-

taxonomic publications among the 100 publications to be produced to demonstrate the existence

of a real current usage. This number will be easy to reach in many cases for species that have

a real economic value and importance, much less so for species that until now have interested

only taxonomists: but in the latter case “fair play” should be enough to accept the rare changes

which are from time to time necessary because of the “rediscovery” of “forgotten” names.

10 . At any rate, I suggest that names not meeting the above minimum requirements should

not even be considered by the Commission for protection, and therefore that applications con¬

cerning such cases should not be accepted for publication in the Bulletin. If such a rule had

been followed in the recent years, the Commission’s burden (in time, work and expenses) would

Source : MNHN, Paris

— 320 —

have been considerably reduced, and the credibility among zoologists of the Commission’s deci¬

sions (in particular as regards suppression of names or changes in priority between different

names) would have remained higher. I contend that below figures of a magnitude similar to

those suggested above, the claim that a name has had a “current usage” is simply ridiculous.

11 . In conclusion, I suggest to replace the conditions given in Art. 79 for a name being

eligible for conservation by the following conditions, which should all be met with together:

1, the “forgotten” senior synonym recently rediscovered should not have been used by a

single author during a 50-year period before the date of publication of its “rediscovery”;

2, during the same period, the junior synonym should have been used by at least 25 inde¬

pendent authors and in at least 100 publications. I also suggest that colleagues pay attention to

the other suggestions presented above in paragraph 9, and also to those provided by Holynski

(1994: 13).

REFERENCES

HOLYNSKI R. B., 1994. — Structure and function or: what kind of nomenclatural regulations do we need? Crystal,

God, Hungary, ser. Zool. 2: 1-50.

NG P. K. L., 1991. — How conservative should nomenclature be? Comments on the principle of priority. Bull,

zool. Norn. 48 : 87-91.

INDEX TO SCIENTIFIC NAMES OF AMPHIBIANS

“ aquaticae ”, "Rana viridis ” 311

arborea, Rana 310-312

arvalis, Rana 311

chrysoscelis, Hyla 305

clamitans, Rana 307

esculenta, Rana 311-312

Hemidactyliini 300

Hyla chrysoscelis 305

Hyla hypocondrialis 308

Hyla lactea 308

Hyla rubra 307

Hyla versicolor 305

hyla, Rana 312

hypocondrialis, Hyla 308

lactea, Hyla 308

megapoda, Rana 301

Mycetoglossini 300

pipiens, Rana 300-307-308

Rana arborea, 310-312

Rana arvalis 311

Rana clamitans 307

Rana esculenta 311-312

Rana hyla 312

Rana megapoda 301

Rana pipiens 300-307-308

Rana temporaria 312

Rana trilobata 300-301

"Rana viridis aquaticae” 311

rubra, Hyla 307

temporaria, Rana 312

trilobata, Rana 300-301

versicolor, Hyla 305

“viridis aquaticae'', "Rana'' 311

Source : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle, Paris, 4' série, 18. 1996

Section A, n“ 3-4 : 321-340

Early scientific names of Amphibia Anura

II. An exemplary case: Rana arborea Linnaeus, 1758

by Alain DUBOIS & Annemarie OHLER

Abslract. -— This paper provides a detailed analysis of the nomenclatural status of the scientific name

Rana arborea Linnaeus, 1758, with identification of its primary, secondary and tertiary syntypes, discussion of

the status of these specimens, and final designation of one of them (now lost) as lectotype of this nominal

species. During this analysis, the status of the following other names are also discussed: Rana hyla Linnaeus,

1758; Hyla Laurenti, 1768; Hyla ranaeformis Laurenti, 1768; Hyla rubra Laurenti, 1768; Hyla sceleton Laurenti,

1768; Hyla viridis Laurenti, 1768; Rana pentadactyla Laurenti, 1768; Hyla gibbosa Lacépède, 1788; Hyla auran-

liaca Daudin, 1802; Rana bilineata Shaw, 1802; Hyla arborea var. meridionalis Bouger, 1874; Sphaenorhynchus

eurhoslus Rivero, 1961. The lines of reasoning illustrated in detail in this case will be used again in further

papers of this series dealing with many other early scientific names of Amphibia Anura.

Key-words. — Nomenclature, Rana arborea Linnaeus, 1758, Hyla Laurenti, 1768, lectotype designation,

stabilization of name.

Noms scientifiques anciens d'amphibiens anoures

II. Un cas exemplaire : Rana arborea Linnaeus, 1758.

Résumé. — Cet article donne une analyse détaillée du statut nomenclatural du nom scientifique Rana

arborea Linnaeus, 1758, avec l’identification de ses syntypes primaires, secondaires et tertiaires, la discussion

du statut de ces spécimens, et finalement la désignation de l’un d’entre eux (actuellement perdu) comme lectotype

de l’espèce nominale. À l’occasion de cette analyse, le statut des noms suivants est également discuté : Rana

hyla Linnaeus, 1758; Hyla Laurenti. 1768; Hyla ranaeformis Laurenti, 1768; Hyla rubra Laurenti, 1768; Hyla

sceleton Laurenti, 1768; Hyla viridis Laurenti, 1768; Rana pentadactyla Laurenti, 1768; Hyla gibbosa Lacépède.

1788; Hyla aurantiaca Daudin, 1802; Rana bilineata Shaw, 1802; Hyla arborea var. meridionalis Bôttger, 1874;

Sphaenorhynchus eurhoslus Rivero, 1961. Les méthodes de travail et de raisonnement exposées de manière dé¬

taillée dans ce cas seront utilisées de nouveau dans les articles ultérieurs de cette série, qui traitera de nombreux

autres noms scientifiques anciens d’amphibiens anoures.

Mots-clés. — Nomenclature, Rana arborea Linnaeus, 1758, Hyla Laurenti, 1768, désignation d’un lectotype.

stabilisation du nom.

A. Dubois & A. Ohler, Muséum national dHistoire naturelle. Laboratoire des Reptiles et Amphihiens. 25 rue Cuvier. F-75231

Paris cedex 05.

INTRODUCTION

In the present series of papers, we intend to clarify and stabilize the status of a number

of ancient scientific names of Amphibia Anura, many of which have already been discussed by

and have posed problems to taxonomists. In the first of these papers (DUBOIS & Ohler 1996),

we exposed some general principles that will help us to deal with these problems, but such

Source : MNHN, Paris

— 322 —

questions always become clearer when they are illustrated by an example. The example we chose

to treat in full detail here is that of the name Rana arbore a Linnaeus, 1758. In further papers

of this series, we will not give as much detailed information, as the mode of reasoning will be

similar, and only factual details will differ.

The name Rana arborea Linnaeus, 1758 is a particularly exemplary case of the rather com¬

plicated situations with which one is sometimes confronted when dealing with very ancient scien¬

tific names. Despite several uses prior to this book, this name was first validly (in the sense of

the Code) used in the 1758 edition of Linnaeus’s Systema Naturae. As usual, Linnaeus did

not state how many specimens he had before him when naming this species, but he gave a

rather high number of “synonyms” of this name (Le. citations of descriptions that, according to

him, referred to the same species). This should not surprise us, because the Systema Naturae

was not an original work with primary descriptions, but a catalogue, similar to modern books

like Frost’s (1985) checklist, where he quoted and sometimes summarized previous detailed

descriptive papers. The characters Linnaeus (1758) gave for the species Rana arborea were so

few and so vague that they could well apply to several hundreds of tree-frog species from almost

all continents: in fact, LlNNAEUS’s concept of the species Rana arborea was closer to our current

concept of the family Hylidae (see e.g. Duellman & Trueb 1985) than to any current concept

of species within amphibians. In Linnaeus’s mind, this species occurred both in Europe and in

America. However, after a rather short period of uncertainty, it became clear that the European

species did not occur in America, and that Linnaeus’s concept of Rana arborea was a composite

concept applying to several distinct species. Since then, Linnaeus’s name has been consistently

applied by thousands of authors to the common European tree-frog, under the name Hyla arborea

(Linnaeus, 1758) (family Hylidae). The logical way to stabilize the status of this name would

then have appeared to be through the designation among Linnaeus’s syntypes of a lectotype

from Europe. However, nobody until now has dared to take such an action, clearly because all

the name-bearing types of this nominal species from Linnaeus’s collection which are still known

to be in existence (now in NHRM) appeared to belong to (several) American species of Hylidae

(LÔNNBERG 1896; Andersson 1900; Duellman 1977). Faced with this situation, all authors

have left this problem unsettled, and the species Hyla arborea still appears in checklists (e.g.

Duellman 1977; Frost 1985) as a species without name-bearing type and without type-locality.

Recently, after electrophoretic studies, NASCETTi, Lanza & BULLIN1 (1995) found that the

populations of Hyla from peninsular Italy show different allozymic patterns from those of central

Europe. These authors think that the Italian populations represent a species distinct from that

usually called Hyla arborea. They suggested to us (Lanza personal communication) that this

new situation made it necessary to fix the status of the latter name. Since all syntypes of Rana

arborea still in existence are from America, they suggested that it is urgent to ask the Commission

to suppress all these syntypes and to designate a neotype in agreement with the current usage

of this name. We note that the same question could also have been raised when the name Hyla

arborea var. meridionalis was created for the “meridional” tree-frog (BÔTTGER 1874: 66), and

a fortiori when it was raised to species level on the basis of bioacoustic studies (Paillette

1967; Schneider 1968), but at that time no one raised the problem. We agree that stabilization

of the status of the name Rana arborea Linnaeus, 1758 is necessary and requires the designation

of a lectotype or of a neotype and fixation of a precise type-locality, but we think that in this

Source : MNHN, Paris

— 323 —

case the regular provisions of the Code are enough to solve this nomenclatural problem, without

having to refer to the Commission. As a matter of fact, and as we already stressed it (DUBOIS

& Ohler 1995a, 1996), the Code does not at all make it compulsory to choose a lectotype

among the syntypes still available, which of course in this case would oblige us to designate

as lectotype a specimen belonging to an American species.

Fig. 1 gives a complete copy of the part of the text of Linnaeus (1758) that deals with

the name Rana arborea. As can be seen, beside the very short and little informative diagnosis

given by LINNAEUS for this species, this text contains additional information: LINNAEUS mentions

seven references to descriptions or figures which, according to him, refer to the species Rana

arborea. These references are clearly “indications” in the sense of the Code. We consider that

all the specimens mentioned in the publications listed in this synonymy (except that preceded

by B, see below) are syntypes of Rana arborea, as much as the specimens which were in

LlNNAEUS’s collections, a few of which only are known to be still in existence in the Stockholm

Museum.

Before designating a lectotype among these specimens, we will analyse in detail all the

information given in the seven references listed by Linnaeus (1758: 213), numbered R1 to R7

in the order of their appearance in Linnaeus’s text. This will allow us to build a list of (at

least) sixteen identified specimens, numbered SI to SI6 in chronological order of their descrip¬

tions, which, even if most of them are not yet in existence today, can be considered syntypes

of Rana arborea, and among which we can choose a lectotype.

Abbreviations

MNHN Muséum national d'Histoire naturelle, Paris. France

NHRM Naturhistoriska Rijkmuseet, Stockholm. Sweden

RMNH Rijksmuseum van Natuurlijke Historié. Leiden. Netherlands

COMMENTED LIST OF REFERENCES CITED IN THE SYNONYMY OF

THE ORIGINAL DESCRIPTION OF Rana arborea Linnaeus, 1758

Rl. “ Amoen. acad. 1. p. 135. Rana pedibus fissis; unguibus subrotundis, corpore laevi: postice

angustato.”

This citation refers to the description of the species numbered 20 in Hast’s thesis (1745:

29), as reprinted in the volume 1 of the Amoenitates Academicae (Linnaeus 1749: 135). This

description was based on a specimen from the “Donatio Caroli Gyllenborg”, about which we

give more information below under number S10. This specimen, which belongs to an American

species of tree-frog, is a secondary (or possibly primary) syntype of Rana arborea Linnaeus,

1758.

This species described by Hast (1745: 29) under number 20 was the only one referred by

him to the genus Rana among the specimens of the GYLLENBORG collection. In the original

edition of his thesis, Hast (1745: 29) included in the synonymy of this species two frogs from

Seba’s (1734, 1735) books, i.e. S3 and S6 below, and one from the Saint Petersburg Museum,

Source : MNHN, Paris

— 324 —

AMPHIBIA REPTILIA. Rana. 213

Hyla. 15. R. dorfo angulato tranfverfe gibbo, abdomine fafcia

replicata inguinali intercepto.

Ge/n. pifc. 809. Rana gibbofa 4. 5.

Habitat in Europa.

Sonus campanarum boatum c longinquo imitatur.

arborea. 16. R. corpore lævi : fubtus punclis contiguis tuberculato,

pedibus fiffis, unguibus orbiculato-dilatatis.

Am am. acad. 1. p. 135. Rana pedibus fiffis; unguibus

fubrotundis, corpore lævi: poftice anguftato.

Muf. Ad. Fr. 1. p. 47. Rana eadem.

Gron. muf. 2. p. 84. n. 63. Rana.

Seb. muf. 1. t. 73. f. 3. Rana brafilienfis gracilis.

Seb. muf. 2. t. 78. /. 5. Rana americana rubra.

Gefn.pifc. 808. Ranunculus viridis.

$ Amain, acad. 1. p. 285. Rana pedibus fiffis. palmis

tetrada&ylis, plantis pentadactylis : geniculis fubtus

tuberofis.

Habitat fub foliis arborum Europæ, Americas, Mufcas

in fauces revocans.

boans. 17. R. corpore lævi : fubtus pun£tis contiguis, pedibus pal-

matis : plantis pentadactylis, palmis tetradafiylis, ungui¬

bus orbiculato-dilatatis.

Amam.acad. 1. p. 285. Rana palmis tetradaftylis, plan¬

tis pentadaflylis palmatis, digitorum apicibus fub¬

rotundis.

Muf. Ad. Fr. \. p. 47. Rana eadem.

Seb. muf. 1. /. 71. /. 3, 4. Rana furinamenfis.

Habitat in America.

Simillima R. arborea:. fed pedes omnes palmati cv corpus

album magnum, maculis etiam laclcis, modo hœc fuffi-

ciant pro fpecie diflingucnda.

o 3 II. SER-

Rana aquatics ovipara fubcunt metamorphojin.

Fig. 1. — Reproduction of page 213 of ihe 1758 edition of Linnaeus's Systema Naturae, including the original descriptions of

Rana hyla and Rana arborea. (Bibliothèque du Laboratoire des Reptiles et Amphibiens, MNHN).

Source : MNHN, Paris

— 325 —

i.e. S7 below. In the reprinted edition of his thesis. Hast (1749: 135) added a fourth synonym

from the Saint Petersburg Museum, i.e. S8 below. All four synonyms can be considered also

syntypes of Rana arborea Linnaeus, 1758. As will be shown in detail below, all four specimens

belong to American tree-frogs.

R2. "Mus. Ad. Fr. 1. p. 47. Rana eadem.”

This citation refers to the diagnosis of the species called Rana arborea in LINNAEUS’S (1754)

book. Three American specimens (SI3-15) are known to correspond to this reference (see below).

Three synonyms are mentioned under this name: the description of HAST (1749: 135) just dis¬

cussed above, based on the specimen S10, and the two species of Seba’s (1734, 1735) frogs

quoted in its synonymy, i.e. the specimens S3 and S6 below.

R3. “ Gron. mus. 2. p. 84. n. 63. Rana.”

This citation refers to the description of the species numbered 63 in Gronovius’s (1756)

book. This specimen is discussed below under SI6. This frog species is said to inhabit Suriname.

Five names are included in the synonymy of this species: Hast's (1745, 1749) species discussed

above, i.e. specimen SIO, and the two Seba’s (1734, 1735) species included in its synonymy,

i.e. S3 and S6; Catesby’s (1743) description of "Rana viridis arborea", a species from Northern

America (S9); and a third species of Seba (1735) from Lemnos (S5).

R4. "Seb. mus. 1. t. 73. /. 3. Rana brasiliensis gracilis.”

This citation is a direct reference to Seba’s (1734) species “Rana brasiliensis gracilis",

about which more information is given below under S3.

R5. "Seb. mus. 2. t. 78. /. 5. Rana americana rubra.”

This citation is a direct reference to Seba’s (1735) species "Ranula americana rubra”, about

which more information is given below under S6.

R6. “Gesn. pise. 808. Ranunculus viridis.”

Conrad Gessner (or Conradus GESNERUS; see Adler 1989: 8) wrote several books, which

furthermore had several editions, but Linnaeus’s reference to the name “Ranunculus viridis ” in

a page 808 suggests that the edition he had in his hands when he wrote his 1758 book was the

second or the third edition of the liber IV of the Historia Animalium, subtitled De Piscium et

Aquatilium Animantium Natura (GESNERUS 1604, 1620; contrary to the statement by ADLER 1989:

7, the 1620 edition of this book is the third, not the second). Three “species” of frogs are dealt

with in page 808 of this book, but LiNNAEUS’s reference is clearly to the first one, “ Calamite ”,

for which GESNERUS mentions two other names (“synonyms”): "Muta" of PLINIUS, and “Ranun¬

culus viridis ” of his own book De Quadrupedibus Oviparis (GESNERUS 1554, reprinted 1586

and 1617). This paragraph in page 808 of GESNERUS’s (1604, 1620) book contains two different

pieces of information, which we regard as evidence of reference to two distinct specimens (at

least): 1, a Latin text about this frog: this text is exactly copied from Rondeletius’s (1555:

224) text entitled “De Calamite ”, which must be assumed to have been based on at least one

specimen (SI); 2, a figure which is an exact reproduction of the figure of “Ranunculus viridis”

that appears in GESNERUS (1554: 55, 1586: 60, 1617: 60). and which we regard as based on a

distinct specimen (S2). Both specimens SI and S2 are secondary syntypes of Rana arborea

Linnaeus, 1758.

Source : MNHN, Paris

— 326 —

R7. “B Amoen. acad. 1. p. 285. Rana pedibus fissis, palmis tetradactylis, plantis pentadactylis:

geniculis subtus tuberosis.”

This citation refers to the description of the species numbered 9 in Balk’s (1746: 8) thesis,

as reprinted in the volume 1 of the Amoenitates Academicae (LINNAEUS 1749: 285-286). This

description was based on two specimens from the “Donatio Adolphi Friderici”, about which we

give more information below under numbers SI 1-12. However, it is important to stress that

these two specimens cannot be considered syntypes of the nominal species Rana arborea, ac¬

cording to Article 72(b)(i) of the Code (see Dubois & Ohler 1996): the presence of the Greek

letter B before the reference clearly shows that LINNAEUS considered that these specimens rep¬

resented a “distinct variant”, which excludes these specimens from the type-series.

Finally, in the text concerning this species. Balk (1746, 1749) also refers to another Seba’s

( 1734) specimen, discussed in more detail below under S4: of course, this specimen also is

excluded by the letter B from the syntypes of Rana arborea.

COMMENTED LIST OF IDENTIFIED SYNTYPES

OF Rana arborea Linnaeus, 1758

51. Specimen(s) referred to in p. 224 of RONDELETIUS (1555) as "Calamite". Origin: region of

Montpellier, Hérault, Languedoc, France. Secondary syntype(s) of Rana arborea Linnaeus, 1758.

Rondeletius’s (1555) “description” does not refer precisely to particular specimens. It was

composed in part of information taken from the works of PLINIUS and Nicander, but this frog

was apparently also known personally to the author, who mentioned several of its medicinal

properties and who wrote: "Ranette nostri nominant” (RONDELETIUS 1555: 224); “Nous l’appel¬

ions en Languedoc Rainette” (Rondelet 1558: 167). Since Guilaume (sic) Rondelet (or Guliel-

mus Rondeletius) was professor at the Montpellier University (as printed on the front pages

of his books), it is logical to consider that his text was in part based on specimen(s) from

Montpellier’s region (Southern France). However, it would be very ill-inspired to designate for¬

mally as (lost) lectotype of Rana arborea one of these specimens, since the only tree-frog that

lives in this region is the species currently known as Hyla meridionalis Bottger, 1874 (see e.g.

Arnold & Burton 1978; Parent 1981; Castanet & Guyétant 1989), a name that deserves

protection since it has already been used in a high number of general publications for many

years and by many distinct authors.

52. Specimen shown in figure of p. 55 of GESNERUS (1554), and reproduced in Gesnerus (1558:

950; 1586: 60; 1604: 808; 1617: 60; 1620: 808), as “Ranunculus viridis". Origin: region of

Zürich, Switzerland. Secondary syntype of Rana arborea Linnaeus, 1758.

This figure (reproduced here in Fig. 2A) shows a specimen of frog on a tree leaf (indicating

an arboreal mode of life). The drawing was apparently made from a living specimen (in contrast,

some other drawings in the same book are easily recognizable as made from dead collection

specimens). It is of a rather poor quality, however it shows a character, the presence of a spot

Source : MNHN, Paris

C %àJDC.

J.J.OUJUjcjt'Mr. Q

Source : MNHN, Paris

— 328 —

near the groin, that can be interpreted as the end of the dark lateral stripe on the flank of the

species Hyla arborea, which allows its distinction from the closely related species Hyla meri-

dionalis (see e.g. Arnold & BURTON 1978: pi. 9; Matz & WEBER 1983: pi. IX; DlESENER &

REICHHOLF 1986: 76-79; GÜNTHER 1986: 154-157). In the front page of GESNERUS’s (1558)

volume, it is expressly stated that the drawings in this work are new, i.e. that they were made

especially for these books. According to Petit (1965: 203), most of the drawings in these books

were done by GESSNER himself, others by other Zurich’s artists. As, at the time of writing this

book, Gessner was living in Zurich (see e.g. Salzmann 1965; Adler 1989), we can assume

that this drawing was made from a specimen of tree-frog collected in the region of Zürich.

Since the drawing was apparently made from a live specimen, since it shows a character that

fits with Hyla arborea. and since the latter is the only species of tree-frog known to occur in

Zurich’s region (see e.g. GROSSENBACHER 1988), the specimen which was used to prepare this

figure, although now lost, is a good candidate for the designation as lectotype of Rana arborea

Linnaeus, 1758.

S3. Specimen shown in fig. 3 of pi. LXXIII and described in p. 117 of SEBA (1734) as “Rana.

Brasiliensis, gracilis”. Origin: Brazil. Secondary syntype of Rana arborea Linnaeus, 1758.

This figure and description were used as the basis of the name Hyla sceleton by Laurenti

( 1768: 35), and Seba’s figure was reproduced by Bonnaterre (1789: pi. 7, fig. 4). To the best

of our knowledge, this name has never been allocated to any biological species, although it

seems clear to us that the drawing and short description could well be applied to one (or several)

Brazilian species of hylids. The specimen shown in fig. 3 of pi. LXXIII of Seba (1734), being

the only specimen on which Hyla sceleton Laurenti, 1768 was based, is the holotype of this

species. This specimen was still in the collections of the Paris Museum at the beginning of the

19th century (Daudin 1802: 28, 1803: 58), but has disappeared from these collections since

then (Guibé 1950; personal observation). We suggest that designation of a neotype from Brazil,

that would fit with the characters given by Seba (1734), would allow the status of this name

to be fixed.

Despite the numerous discussions already devoted to the name Hyla aurantiaca Daudin,

1802 (see e.g. Rivero 1969; Duellman & Lynch 1981; Lynch & Duellman 1984), all authors

until now have ignored the fact that this name, which is a junior homonym of Hyla aurantiaca

Laurenti, 1768, is nothing but a strict replacement name for Hyla sceleton Laurenti, 1768. In

both his texts referring to this species, Daudin (1802: 28, 1803: 58) stated in full words that

he changed the name of this species because of its inadequacy, and that he only with doubt

referred to this species a second specimen in the collections of the Paris Museum. Therefore,

both nominal species Hyla sceleton Laurenti, 1768 and Hyla aurantiaca Daudin, 1802 have the

same holotype, the specimen shown in Seba’s figure mentioned above. The second specimen

mentioned by Daudin (1802, 1803) is still in the Paris Museum collections, under the number

MNHN 4871, but contrary to the statement of Guibé (1950: 18), followed by Rivero (1969:

701), DUELLMAN & LYNCH (1981: 238) and Lynch & Duellman (1984: 122), it is not the

holotype of Hyla aurantiaca Daudin, 1802. Therefore, Rivero’s (1969) proposal of the “new

name” (nomen novum) Sphaenorhynchus eurhostus for Hyla aurantiaca Daudin, 1802 (preoc-

Source : MNHN, Paris

— 329 —

cupied) is incorrect, because, strictly speaking, being a new name for Hyla aurantiaca Daudin,

1802, and hence an objective synonym of it, this name is also a new name for and an objective

synonym of the name Hyla sceleton Laurenti, 1768. FROST (1985: 175) decided to follow the

suggestion of Duellman & LYNCH (1981) and LYNCH & DUELLMAN (1984) to apply the name

Sphaenorhynchus lacteus (Daudin, 1800b) (not “1802” or “1803”, as written by mistake in the

above quoted works) to the species which Rivero (1969) meant to designate under the name

Sphaenorhynchus eurhostus. Let us note however that, to be valid, this action will have to be

confirmed by a vote of the International Commission on Zoological Nomenclature. We will come

back to the problems raised by these names in a forthcoming paper of this series.

Wagler (1833: 890) referred the specimen S3 in Seba (1734) to the species “Discodactylus

ruber mihi”. Despite appearances, this is not a new name for this species, but only a new com¬

bination (hence the term “mihi”) of the specific name Hyla rubra Laurenti, 1768 with the new

generic name Discodactylus , proposed in the same paper by Wagler (1833: 888) as a nomen

novum for the generic name Hyla Laurenti, 1768 (this objective synonym of Hyla was overlooked

by Duellman 1977: 24).

In conclusion, the specimen S3, a frog from Brazil, is the holotype of the following three

nominal species: Hyla sceleton Laurenti, 1768; Hyla aurantiaca Daudin, 1802 (nec Hyla auran¬

tiaca Laurenti, 1768); and Sphaenorhynchus eurhostus Rivero, 1961. It cannot be chosen as

lectotype of Rana arborea Linnaeus, 1758 if stability of the use of this name is to be maintained.

S4. Specimen shown in fig. 1 of pi. LXXV and described in p. 119 of Seba (1734) as “Rana,

maxima, Virginiana, eximia, rara; foemina”. Origin: “Pennsylvania”, no doubt in error (see

Heyer 1979: 13); emended to “Indiis” by Laurenti (1768: 32). Specimen expressly excluded

(as variety 6) from the syntypes of Rana arborea by LINNAEUS (1758).

This specimen was the one on which was based the name Rana pentadactyla Laurenti,

1768. The second specimen, stated by Laurenti (1768: 32) to be kept in the “Museo Illustrissimi

Comitis Turriani”, was expressly excluded from the name-bearing types of the latter nominal

species, according to Article 72(b)(i) of the Code, by its clear inclusion in a “var. 6”. It is

therefore in error that HEYER (1979: 13) considered both specimens as syntypes of this nominal

species, but fortunately this author treated the specimen illustrated by Seba (1734) as “name

bearer”, i.e. lectotype, of Rana pentadactyla, so that the nomenclatural status of the name remains

unchanged. Müller (1927: 277) restricted the type-locality of this species to “Surinam”, but,

as this action was not accompanied by a neotype designation, it is not valid. Heyer (1979: 13)

ignored MÜLLER’s designation and refrained from restricting the type-locality, as this action

“would involve arbitrary decisions”. However, it is clear that final stabilization of the status of

this name will require such a restriction, which can be validly obtained only through the des¬

ignation of a neotype from a known population. In order not to upset MÜLLER’s (1927) action,

we suggest it would be better to choose a neotype from Suriname. Pending such a designation,

SEBA’s specimen, now lost, remains the holotype of the species Rana pentadactyla Laurenti,

1768. As mentioned above, according to Article 72(b)(i) of the Code, this specimen cannot be

considered syntype of Rana arborea Linnaeus, 1758.

Source : MNHN, Paris

— 330 —

S5. Specimen shown in fig. 2 of pi. XIII and described in p. 16 of Seba (1735) as “Rana lemnia".

Origin: Limnos, Greece. Tertiary syntype of Rana arborea Linnaeus, 1758.

This specimen is the holotype of the nominal species Hyla ranaeformis Laurenti, 1768 (orig¬

inal description shown here in Fig. 3) and Hyla gibbosa Lacépède, 1788, and Seba’s figure was

reproduced under the latter name by Bonnaterre (1789: pi. 5, fig. 1). We discussed elsewhere

(DUBOIS & Ohler 1995b) the status of these two objective synonyms, which we think apply

to a European green frog of the subgenus Rana (Pelophylax). This subgenus has a very com-

R E P T I L I ü M. 33

crura poftica longilfima : hinc faltus in¬

gens ; digiti fcandentes apicibus in àrbi-

cula vifcida plana, inftar oris hirudinisdi-

latabilibus. Catesby. 2. pag. 71. bint

J'edet fiib folio , digitis adbarcns, m Jupra

incumbens,

XXV. Hyla ransformis. Seba II. 13. 2.

Diagn.

Capite rotundo piano ; oculis promineriti-

bus ; pedibus fafciatis ; Iateribus iaturatius

tin&is; dorfo diftindtiflimo gibbo.

Var. ( 3 . ( Seba II. 70. 4. ) fuperne maculis af-

perl'a.

Habitat prior in Lmno. /?. Surin.mii.

XXVI. Hyla viridis. Rtfel. Tab. IX. X. XL

fiontifptc.

Diagn.

Supra virens, infra albens, utroque latere

linea flava.

Var. f 3 . (Catesby Carol. 2. 71. )' corpora té-

reti, linea u'rinqne flava, led re&a; diftin.

guitur ciimove tfchic, tfchit, tfchit, dam

noftra clamat ra, ra, ra.

c Ha-

Quam ob rem hxc l’ujfi Hylx f.LCeiJü! aomeu ejus>

dem m.-rn- vit.

34 TABULA

Habitat prior in Europœ arboribus ; p. in

America.

XXVII Hyla fufca.

Diagn.

Corpora fufco, pedibus fubtus ad talos, &

ad iingulos digitorum articulos tuberofo-la-

ciniatis.

Hofpitatnr in Mttf.o Academico Upfalienjt ,

fij 5 Vienna in Turriano tbefauro.

XXVIII. Hyla ladtea.

Diagn.

Corpora niveo, maculis Iadteis, femoribus.

tibiisque Tuhlividis ; hypochondriis obfolete

cinereo-fafciatis; ore amplifiimo.

Var. |3- coloris fupra cærulefcentis fubplumbei.

Habitat prior in America ; bojpitatur in mu-

feo Academico Upfalieti/i. p in mufeo Pe¬

tr opolitano.

XXIX. Hyla viridi - fufca. Merian Surinam.

Tab. 56.

Diagn.

Corpora fufco, maculis viridibus emargina-

tis; pedibus viridi-fafciatis; collo utrinque

facco conico viridi ocellato.

Habitat Surinam’.

XXX. Hyla tibiatrix. Seba 1. 71 . Fig. i.& 2 .

Diagn.

Fig. 3. — Reproduction of pages 33 and 34 of LAURENTl's (1768) Specimen Medicum, including Ihe original descriptions of Hyla

ranaeformis and Hyla viridis. (Bibliothèque du Laboratoire des Reptiles et Amphibiens, MNHN).

Source : MNHN, Paris

— 331 —

plicated taxonomy and nomenclature (see DUBOIS & Ohler 1995a) and we refrain from dis¬

cussing further the status of this name here. At any rate, it is clear that this specimen cannot

be chosen as lectotype of Rana arborea Linnaeus, 1758.

S6. Specimen shown in fig. 5 of pi. LXVIII (not LXXVIII, as printed by error in Linnaeus,

1758: 213) and described in p. 70 of SEBA (1735) as “ Ranula, Americana, rubra”. Origin: Ame¬

rica. Secondary syntype of Rana arborea Linnaeus, 1758.

This figure and description were used as the basis of the name Hyla rubra by Laurenti

( 1768: 35). The same name was used again, with reference to the same figure in Seba, by many

ancient authors, including LacÉPÈDE (1788: 327, 459), Bonnaterre (1789: 10), Daubenton

(1782: 668), Daudin (1800a: 11; 1802: 26; 1803: 53) and Daudin in Sonnini & Latreille

( 1801: 176). According to Daudin (1800a: 11; 1802: 27; 1803: 54), the specimen shown in figs

1-2 of pi. VI of Daudin (1800a), in fig. 1 of the plate facing p. 176 of Sonnini & Latreille

( 1801), and in figs 1-2 of pi. IX of Daudin (1802) was kept in the Paris Museum collections,

and originated from Seba’s collection; however, Duméril & BlBRON (1841: 595) stated that

this specimen did not come from Seba’s collection, so that it cannot be the specimen shown in

fig. 5 of pi. LXVIII of Seba (1735). Despite this discrepancy, it is clear that Daudin did not

propose a new name, but simply used Laurenti’s name Hyla rubra , even if he did not expressly

quote the latter, as was often the case in his time (see the detailed discussion of this case in

Dubois & Ohler 1996). Therefore we disagree with Duellman & Wiens (1993) who recognized

a nominal species Hyla rubra Daudin, 1802, distinct from Hyla rubra Laurenti, 1768.

The holotype of Hyla rubra Laurenti, 1768, from Seba’s collection, is not in the Paris

Museum collections (Duméril & Bibron 1841; GUIBÉ 1950; personal observation) and must

be considered lost. Therefore, Duellman & Wiens’s (1993) designation of a neotype (RMNH

25883, from Paramaribo, Suriname) for this species is valid, and stabilizes the status of this

name.

Given all this information, the specimen S6 would of course be a very bad choice for the

designation of a lectotype for Rana arborea Linnaeus, 1758.

S7. Specimen from the Saint Petersburg collection described under Nr. 47 in p. 427 of ANONY¬

MOUS (1742) as “Rana surinamensis prone coerulescentis supine albi coloris, ad latera utrimque

maculis nigris notata cum foetibus exclusis Pipae”. Origin: Suriname. Tertiary syntype(s) of Rana

arborea Linnaeus, 1758.

We are not aware of any publication that would elucidate the status of the specimen, kept

in alcohol, on which was based the above reference, but since this frog is said to be from

Suriname, this specimen would not appear to be a good candidate for designation of a lectotype

for Rana arborea Linnaeus, 1758.

Source : MNHN, Paris

— 332 —

S8. Specimen from the Saint Petersburg collection described under Nr. 55 in p. 428 of ANONY¬

MOUS (1742) as “Rana americana parva ventre albido; dorso plumbei coloris, lateribus ex albo

et nigro varietatis". Origin: America. Tertiary syntype(s) of Rana arborea Linnaeus, 1758.

The case of this name is similar to the preceding one. As this specimen is said to be from

America, it would not be a reasonable choice for lectotype designation for Rana arborea

Linnaeus, 1758.

S9. Specimens shown in pi. 71 and described in p. 71 of Catesby (1743) as “ Rana viridis

arborea". Origin: “Virginia and Carolina", U.S.A. Tertiary syntypes of Rana arborea Linnaeus,

1758.

These specimens (one shown on the plate, additional ones suggested by the text), which

belong to a North American tree-frog, were the basis of “var. B” of Laurenti’s (1768) Hyla

viridis. Holbrook (1842: 121-122) restricted the use of the latter name to Catesby’s (1743)

species, and so doing he “almost” designated a lectotype, but this action was incorrect because,

according to Article 72(b)(i) of the Code, mention of “var. B” excludes these specimens from

the syntypes. The same specimens were later the basis of the description by SHAW (1802: 136)

of Rana bilineata, of which they are therefore the syntypes. According to Duellman (1977:

46), they belong to the species currently known as Hyla cinerea (Schneider, 1799), which inhabits

the southeastern U.S.A. Therefore these specimens would be a very bad choice for lectotype

designation for Rana arborea Linnaeus, 1758.

S10. Specimen from the “Donatio Caroli Gyllenborg” described under Nr. 20 in p. 29 of Hast

( 1745) and in pp. 135-136 of Hast (1749) as “ Rana pedibus fissis, unguibus subrotundis, corpore

laevi, pone angustato". Origin: unknown. Secondary (or possibly primary) syntype of Rana ar¬

borea Linnaeus, 1758.

According to Lônnberg (1896: 11), a single specimen is kept in the Linnaeus collection

(now in NHRM) with the label “ Rana arborea, Mus. Gyllenb.”, but this specimen, which ap¬

parently belongs to the American species Hyla leucophyllata Beireis, 1783, was probably not

part of the original Gyllenborg collection described by Hast (1745, 1749). The identity of

the original specimen(s) described by Hast (1745, 1749) remains therefore in doubt, but, since

this author listed only American species in the synonymy of this species, it seems appropriate

to admit that the latter was probably an American species. This specimen also would therefore

be a very bad choice for the lectotype of Rana arborea Linnaeus, 1758.

Sll-12. Specimens from the “Donatio Adolphi Friderici” described under Nr. 9 in pp. 8-9 of

Balk (1746) and in pp. 285-286 of Balk (1749) as “ Rana pedibus fissis, palmis tetradactylis,

plantis pentadactylis; geniculis subtus tuberosis". Origin: unknown. Specimens expressly ex¬

cluded (as variety B) from the syntypes of Rana arborea by LINNAEUS (1758).

Source : MNHN, Paris

— 333 —

According to LÔNNBERG (1896: 13), two specimens from the “Donatio Adolphi Friderici”

were kept in the LINNAEUS collection (now in NHRM). LÔNNBERG (1896: 13) writes: “The two

discoloured specimens can hardly with certainty be classified, but certainly they do not belong

to the Hyla arborea of recent authors. I think, it is some American species with better developed

vomerine teeth.” Duellman (1977: 31) did not mention these two specimens in his discussion

of the Linnaean syntypes of Rana arborea still in existence. As a matter of fact, as mentioned

above, these two specimens cannot be candidates for the lectotype designation of Rana arborea

Linnaeus, 1758 because, according to article 72(b)(i) of the Code, they are not to be considered

syntypes of this nominal species.

S13-15. Specimens from the “Museum Drottningholmense" collection referred to in p. 47 of

LINNAEUS (1754) as “Rana arborea". Origin: America. Primary syntypes of Rana arborea Lin¬

naeus, 1758.

According to Andersson (1900: 17), three specimens corresponding to this reference were

kept in the Linnaeus collection (now in NHRM). Two of these specimens were identified by

ANDERSSON (1900: 17) as Hyla punctata (Schneider, 1799), a species from South America.

Duellman (1974: 10, 1977: 31, 89) further stated that these two specimens (NHRM 155) are

part of the syntypes of Calamita punctata Schneider, 1799. The third specimen was tentatively

referred by Andersson (1900: 18) to the species Hyla inframaculata Boulenger, 1882 from

Brazil, “or some other species, belonging to the same American group of the genus Hyla". Ac¬

cording to DUELLMAN (1977: 31), this third specimen is not to be found now in the NHRM

collection. At any rate, these three American specimens are not good candidates for the lectotype

designation of Rana arborea Linnaeus, 1758.

S16. Specimen(s) from Gronovius’s “Museum Ichthyologicum” described under Nr. 63 in p. 84

of GRONOVIUS (1756) as "Rana palmis tetradactylis fissis, plantis pentadactylis semipalmatis.

unguibus digitorum subrotundis, corpore laevi, pone angustato". Origin: Suriname. Secondary

syntype(s) of Rana arborea Linnaeus, 1758.

We are not aware of any publication that would elucidate the status of the specimen(s)

which GRONOVIUS (1756) had before him when he wrote the rather detailed description of this

species, but he clearly stated that this frog was from Suriname, so that this/these specimen(s)

would not appear to be good candidate(s) for designation of a lectotype for Rana arborea

Linnaeus, 1758.

LECTOTYPE DESIGNATION FOR Rana arborea Linnaeus, 1758

The detailed survey above has allowed to identify at least sixteen specimens that could be

considered as possible syntypes of Rana arborea Linnaeus, 1758. The number of sixteen is a

number by default, because in some cases the exact number of specimens which had been used

to prepare the original description cannot now be ascertained.

Source : MNHN, Paris

— 334 —

A detailed analysis of the data concerning these specimens shows that the latter can be

referred to four categories, as follows.

1. Three specimens (SI3-15) are primary syntypes of Rana arborea. All three refer to Amer¬

ican species of Hylidae, and would therefore be very bad choices for lectotype designation for

Rana arborea. Although two of these specimens (NHRM 155) are apparently the only original

syntypes of Rana arborea to be still in existence, we propose to discard them as name-bearing

types, in order to maintain the stability of nomenclature. Designation below of another, not pri¬

mary, syntype, as lectotype makes these two specimens become paralectotypes of Rana arborea ,

i.e. specimens which do not play any more role for the clarification of the nomenclatural status

of this name.

2. Three specimens (S7-9) are tertiary syntypes of Rana arborea. As we already underlined

it (Dubois & Ohler 1996), tertiary syntypes should as much as possible be avoided for lectotype

choice. In the present case, this is all the more justified that these three specimens originated

from America.

3. Three specimens (S4, SI 1-12) were expressly excluded (as variety 6) from the syntypes

of Rana arborea by Linnaeus (1758) himself. Furthermore, one of these specimens (S4) clearly

belonged to an American species, and the other two probably also.

4. Finally, seven specimens were identified as secondary syntypes of Rana arborea. Four

of these specimens (S3, S6, S10, S16) can immediately be discarded for possible lectotype des¬

ignation, as they originated from America. The three remaining specimens (SI, S2, S5) were of

European origin. The detailed analysis presented above shows that these (at least) three specimens

belonged to three distinct biological species. Specimen(s) SI belong(s) to the species currently

known as Hyla meridionalis Bbttger, 1874. Specimen S5 belongs to a green frog of the Rana

(Pelophylax) subgenus, not to the genus Hyla. The only syntype which clearly belongs to the

species currently known as Hyla arborea Linnaeus, 1758 is the specimen S2. Although this

specimen is now lost, it is clearly the best choice for lectotype designation for Rana arborea:

this choice will allow this name to remain objectively and definitively attached to the central

European tree-frog which has been called Hyla arborea or Hyla arborea arborea in thousands

of publications already.

In conclusion, we hereby formally designate as lectotype of the nominal species Rana ar¬

borea Linnaeus, 1758 the specimen shown in the figure of page 55 of Gesnerus’s (1554) book,

here reproduced in Fig. 2A. In agreement with this designation, we hereby restrict the type-

locality of this nominal species to: “region of Zürich (47°23’N, 8°32’E), canton of Zurich, Swit¬

zerland”.

If in the future it appeared essential, in order to stabilize further the status of the name

Rana arborea Linnaeus, 1758, to have a type-specimen to which some biological information

(such as bioacoustic, biochemical or cytogenetic data) be attached, it would be possible to des¬

ignate a neotype for this taxon. The only requirement that would then follow from our present

action is that this specimen should have been collected in the region of Zurich. Such a work

should be carried out paying attention to the conservation status of this species, which is now

rare in the Zürich region (Grossenbacher 1988: 109-111).

Source : MNHN, Paris

— 335 —

STATUS OF THE NAME Rana hyla Linnaeus, 1758

The name Rana hyla Linnaeus, 1758 has been traditionally considered a synonym of Rana

arborea Linnaeus, 1758 (see e.g. MERTENS & Wermuth 1960: 49; Gorham 1974: 94;

Duellman 1977: 31). Probably this synonymy was initially proposed because of the identity

between Linnaeus’s (1758) specific name hyla and LAURENTl’s (1768) generic name Hyla , the

valid name of the genus including the European tree-frogs. However, this is incorrect.

This statement is not based on the examination of original syntypes of this nominal species,

since no such syntypes are known to exist (LOnnberg 1896; Andersson 1900; Duellman

1977). The original text of LINNAEUS (1758: 213), which is here reproduced in Fig. 1, gives a

short diagnosis of Rana hyla and refers to a single synonym for this name: “Rana gibbosa ”

in GESNERUS (1604: 809, 1620: 809). This name is in a situation similar to that of the name

“Ranunculus viridis ” discussed above under S2: here also the descriptive notes and drawing

in liber IV of the Historia Animalium refer to the earlier description and drawing in liber

II. The figure (reproduced here in Fig. 2B) is of a good quality and clearly shows an European

brown frog of the group of Rana (Rana) temporaria Linnaeus, 1758 (see DUBOIS 1992). In

the 1558, 1604 and 1620 editions of liber IV, GESNERUS did not provide a detailed description,

but referred to the description of “Rana gibbosa ” that he had published earlier in liber II

(GESNERUS 1554: 58, 1586: 63, 1617: 63). The latter description also clearly fits with Rana

temporaria. Besides, in both books, GESNERUS provided comments on the mating calls of

these frogs (with a discussion about so-called “voiceless” populations), notes on their habitats,

and discussions of previous authors who had already written about them. All this information

is quite clear evidence that the species meant by this author under the name “Rana gibbosa”

is our current Rana temporaria.

In order to definitively stabilize the status of the name Rana hyla Linnaeus, 1758, we hereby

formally designate as lectotype of this nominal species the specimen shown in the figure of

page 63 of GESNERUS’s (1554) book, here reproduced in Fig. 2B. In agreement with this des¬

ignation, and following the conclusions of the discussion above under S2, we hereby restrict

the type-locality of this nominal species to “region of Zurich (47°23’N, 8°32’E), canton of Zürich,

Switzerland”.

The name Rana hyla Linnaeus, 1758 should therefore now be treated as a subjective syn¬

onym of Rana temporaria Linnaeus, 1758. The latter name is the name of the type-species of

the genus Rana Linnaeus, 1758, and it has been used in thousands of publications by thousands

of authors. It should therefore be protected. In order to avoid any possible threatening of this

name, we hereby take a first reviser action and we afford priority to the name Rana temporaria

Linnaeus, 1758 over Rana hyla Linnaeus, 1758.

Source : MNHN, Paris

— 336 —

STATUS OF THE NAME Hyla viridis Laurenti, 1768

The name Hyla viridis Laurenti, 1768 is traditionally referred to the synonymy of Hyla

arborea (Linnaeus, 1758) (see e.g. Mertens & Wermuth 1960; Gorham 1974; Duellman

1977). This name is of particular nomenclatural importance, because, following STEJNEGER’s

(1907: 75) designation, it is the type-species of the nominal genus Hyla Laurenti, 1768 (which

had been created with nine originally included species, some of which are now placed in other

genera). So its assignation to a biological species must be clear and definitive, and not liable

to raise problems in the future.

Fig. 3 provides a copy of the original description of the nominal species Hyla viridis

Laurenti, 1768. It contains three parts. The first part starts with reference to plates IX, X, XI

and frontispiece of ROESEL VON ROSENHOF (1758), followed by a short diagnosis; this clearly

refers to the common European tree-frog, studied and figured in many details by ROESEL von

ROSENHOF (1758) under the name “ Rana arborea". The second part concerns the “var. 6” of

this species: it consists in a reference to Catesby’s (1743: 71) text, followed by descriptive

notes, including notes on the calling behavior of the frog; this clearly refers to the American

tree-frog discussed above under S9. Finally, the third part concerns the distribution of this frog,

which is said to inhabit “first” the trees of Europe, and America for variety 6. We already pointed

out above that Holbrook’s (1842) “restriction” of the use of the name Hyla viridis to the Amer¬

ican species is invalid, since mention of "var. B" excludes Catesby’s specimens from the syn-

types. Therefore the only remaining syntypes are those shown in ROESEL VON ROSENHOF’s (1758)

plates quoted in Laurenti ( 1768: 33). All these specimens clearly belong to the species currently

known as Hyla arborea (Linnaeus, 1758).

We hereby formally designate as lectotype of the nominal species Hyla viridis Laurenti,

1768 the calling male shown in middle left position in plate IX of ROESEL von ROSENHOF

(1758), here reproduced in Fig. 2C. Since Roesel von Rosenhof lived and worked in Nürnberg

(see e.g. Adler 1989: 10), and since his text and illustrations were clearly prepared from living

specimens which he must have collected close to his working place, we hereby restrict the type-

locality of this nominal species to: “region of Nürnberg (49°27’N, 11°04’E), Bayern, Germany”.

This nominal species should currently stand as a junior subjective synonym of Hyla arborea

(Linnaeus, 1758).

Acknowledgements

We are very grateful to Leo J. Borkin (Saint-Petersburg) for providing a partial photocopy of a very

rare volume (Anonymous 1742), to Roger Bour (Paris) for bibliographical advice, and to Monique

Ducreux (Paris) for the facilities provided for our work on the ancient books of the Central Library of

the Paris Museum.

Source : MNHN, Paris

— 337 —

REFERENCES

ANONYMOUS, 1742. — Musei Imperialis Petropolitani. Vol. I, Pars prima qua continentur res naturales ex regno

animali. Petropolitanae, Typis Academiae Scientiarum: i-viii + 1-332.

ADLER K.. 1989. — Herpetologists of the past. In K. ADLER (ed.). Contributions to the history of herpetology,

Oxford, Ohio. S.S.A.R.: 5-141.

ANDERSSON L. G., 1900. — Catalogue of Linnean type-specimens of Linnaeus’s Reptilia in the Royal Museum

in Stockholm. Bihang K. Svenska Vet.-Akad. Handlingar 26 (4): 1-29.

ARNOLD E. N. & BURTON J. a., 1978. — A field guide to the reptiles and amphibians of Britain and Europe.

London, Collins: 1-272, pi. 1-40.

Balk L., 1746. — Museum Adolpho-Fridericianum. Holmiae, Laurentii Salvii: [i-viii] + 1-50, pi. I-II.

— 1749. — Museum Adolpho-Fridericianum. In C. LINNAEUS (ed.). Amoenitates Academicae, 1, Holmiae &

Lipsiae, Godofredum Kiesewetter: 277-326, pi. XIII-XIV.

BÔTTGER 0., 1874. — Reptilien von Marocco und von den canarischen Inseln. AM. Senckenb. naturf. Ges. 9:

121-191. 1 pi.

BONNATERRE Abbé, 1789. — Tableau encyclopédique et méthodique des trois règnes de la nature. Erpétologie.

Paris, Panckoucke : i-xxviii + 1-70, pl. 1-7 + 1-6 + 1-12.

Castanet J. & GUYÉTANT R. (eds.), 1989. — Atlas de répartition des amphibiens et reptiles de France. Paris,

Société Herpétologique de France : 1-191.

CATESBY M., 1743. — The natural history of Carolina. Florida and the Bahama Islands. Vol. II. London: [i-iv]

+ 1-100 + i-xliv + [i-vi], pl. 1-100.

DAUBENTON M., 1782. — Les animaux quadrupèdes ovipares, et les serpens. In Encyclopédie méthodique -

Histoire naturelle des animaux. Tome premier, Paris, Panckoucke : 545-712.

DAUDIN F. M., 1800a. — Histoire naturelle des quadrupèdes ovipares. Livraison 1. Paris, Fuchs, Delalain, Treuttel

& Würtz : [1-11], pl. [I-VI],

— 1800b. — Histoire naturelle des quadrupèdes ovipares. Livraison 2. Paris, Fuchs. Delalain. Treuttel &

Würtz : [13-24], pl. [VII-XI1],

— 1802. — Histoire naturelle des rainettes, des grenouilles et des crapauds. Paris. Levrault : 1-108, pl. I-

XXXVIII.

— 1803. — Histoire naturelle, générale et particulière des Reptiles. Tome huitième. Paris, Dufart : 1-439,

8 pl.

DlESENER G. & REICHHOLF J., 1986. — Lurche und Kriechtiere. München, Mosaik Verlag: 1-288.

DUBOIS a., 1992. — Notes sur la classification des Ranidae (Amphibiens Anoures). Bull. Soc. linn. Lyon 61

(10) : 305-352.

DUBOIS a. & OHLER a., 1995a. — Frogs of the subgenus Pelophylax (Amphibia, Anura, genus Rana): a catalogue

of available and valid scientific names, with comments on name-bearing types, complete synonymies, pro¬

posed common names, and maps showing all type localities. Zool. Polon., (1994) 39 (3-4): 139-204.

— 1995b. — Catalogue of names of frogs of the subgenus Pelophylax (Amphibia, Anura, genus Rana): a few

additions and corrections. Zool. Polon., (1994) 39 (3-4): 205-208.

— 1996. — Early scientific names of Amphibia Anura. I. Introduction. Bull. Mus. natl Hist, nat., 4 e ser.,

section A 18 (3-4): 297-320.

DUELLMAN W. E., 1974. — A reassessment of the taxonomic status of some Neotropical hylid frogs. Occ. Pap.

Mus. nat. Hist. Univ. Kansas 27: 1-27.

— 1977. — Liste der rezenten Amphibien und Reptilien. Hylidae, Centrolenidae, Pseudidae. Das Tierreich 95:

i-xix + 1-225.

DUELLMAN W. E. & LYNCH J. D., 1981. — Nomenclatural resolution of the identities of Hyla aurantiaca and

Hyla lactea. J. Herpet. 15: 237-239.

DUELLMAN W. E. & TRUEB L., 1985. — Biology of amphibians. New York, McGraw-Hill: i-xix + 1-670.

DUELLMAN W. E. & Wiens J. J., 1993. — Hylid frogs of the genus Scinax Wagler, 1830, in Amazonian Ecuador

and Peru. Occ. Pap. Mus. nat. Hist. Univ. Kansas 153: 1-57.

Source : MNHN, Paris

— 338 —

DUMÉRIL A.-M.-C. & BlBRON G., 1841. — Erpétologie générale ou histoire naturelle complète des Reptiles.

Tome 8. Paris, Roret : i-vii + 1-792.

FROST D. R. (ed.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst. Coll.: [i-iv]

+ i-v + 1-732.

GESNERUS C., 1554. — Historia Animalium. Liber II. De Quadrupedibus Oviparis. Tiguri [Zürich],

C. Froschoverus: [i-vi] + 1-110 + 1-27.

— 1558. — Historia Animalium. Liber IV. De Piscium et Aquatilium Animantium Natura. Tiguri [Zürich],

Christoph Froschoverus: [i-xxxix] + 1-1297.

— 1586. — Historia Animalium. Liber II. De Quadrupedibus Oviparis. Francofurdi, Roberti Cambieri: [i-vi]

+ 1-119.

— 1604. — Historia Animalium. Liber IV. De Piscium et Aquatilium Animantium Natura. Editio secunda.

Francofurti, Andreae Cambieri: [i-xl] + 1-1052 + 1-38.

— 1617. — Historia Animalium. Liber IL De Quadrupedibus Oviparis. Francofurti, Henrici Laurentii: [i-vi]

+ 1-119.

— 1620. — Historia Animalium. Liber IV. De Piscium et Aquatilium Animantium Natura. Editio secunda [sic].

Francofurti, Henrici Laurentii: [i-xl] + 1-1052 + 1-30.

GORHAM S. W„ 1974. — Checklist of world amphibians up to January I. 1970. Saint-John, The New Brunswick

Museum: 1-173.

GRONOVIUS L. T., 1756. — Musei Ichthyologici. Tomus secundus. Lugduni Batavorum, Theodorum Haak: [i-viii]

+ 1-88, pi. V-VII.

GROSSENBACHER K., 1988. — Atlas de distribution des amphibiens de Suisse. Documenta faunistica Helvetiae

8 : 1-208.

GuiBÉ J., 1950. — Catalogue des types d'amphibiens du Muséum national d’Histoire naturelle. Paris, Imprimerie

nationale: 1-71.

GÜNTHER R., 1986. — Ordnung Anura, Froschlurche. In W.-E. ENGELMANN, J. FRITZSCHE, R. GÜNTHER & F.

J. Obst, Lurche und Kriechtiere Europas, Stuttgart & München, Enke & DTV: 113-184.

Hast B. R., 1745. — Amphibia Gyllenborgiana. Dissertatione Academica, Upsaliae: [i-viii] + 1-36.

— 1749. — Amphibia Gyllenborgiana. In C. LINNAEUS (ed.). Amoenitates Academicae, 1, Holmiae & Lipsiae,

Godofredum Kiesewetter: 107-140.

HEYER W. R., 1979. — Systematics of the pentadactylus species group of the frog genus Leptodactylus (Amphibia:

Leptodactylidae). Smithsonian Contrib. Zool. 301: i-iii + 1-43.

HOLBROOK J. E., 1842. — North American herpetology; or, a description of the reptiles inhabiting the United

States. Volume IV. Philadelphia, J. Dobson: i-vi + 7-138, pi. 1-35.

LaCÉPÈDE [B.G. E.] DE, 1788. — Histoire naturelle des quadrupèdes ovipares et des serpens. Vol. II, 16 cm.

Paris, Hôtel de Thou : [i-iv] + 1-464, pl. I-XV.

LAURENTI J. N., 1768. — Specimen Medicum, exhibens synopsin Reptilium emendatam cum experimentis circa

venena et antidota Reptilium austriacorum. Viennae, Joan. Thom. Nob. de Trattnern: i-ii + 1-215, pl. I-V.

LINNAEUS C. (ed.), 1749. — Amoenitates Academicae, 1, Holmiae & Lipsiae, Godofredum Kiesewetter: [i-iv]

+ 1-563, pl. I-XVII.

— 1754. — Museum Adolphi Friderici Regis Svecorum, Gothorum, Vandalorumque. Holmiae, Typographia

Regia: [i-iii] + i-xxx + 1-96 + [i-vii], pl. I-XXXIII.

— 1758. — Systema Naturae per régna tria naturae, secundum classes, ordines, genera, species, cum charac-

teribus, differentiis, synonymis, locus. Editio décima, reformata. Tomus I. Holmiae: i-iv + 1-824.

LÔNNBERG E., 1896. — Linnean type-specimens of birds, reptiles, batrachians and fishes in the Zoological

Museum of the R. University in Upsala. Bihang K. Svenska Vet.-Akad. Handlingar 22 (4): 1-45.

LYNCH j. D. & DUELLMAN W. E., 1984. — Hyla lactea Daudin, 1803 (Amphibia): request for conservation under

the plenary powers. Bull. zool. Norn. 41: 122-124.

MATZ G. & Weber D., 1983. — Guide des amphibiens et reptiles d'Europe. Neuchâtel & Paris, Delachaux &

Niestlé : 1-292, pl. I-XLIV.

MERTENS R. & Wermuth H., I960. — Die Amphibien und Reptilien Europas. (Dritte Liste, nach dem Stand

vom 1. Januar 1960). Frankfurt am Main. Waldemar Kramer: i-xi + 1-264.

Source : MNHN, Paris

— 339 —

MÜLLER L., 1927. — Amphibien und Reptilien der Ausbeute Prof. Bresslau’s in Brasilien 1913-1914. Abh. Senk-

kenb. nalurf. Ges. 40 (3): 259-304.

Nascetti G., Lanza B. & BULLINI L., 1995. — Genetic data support the specific status of the Italian treefrog

(Amphibia: Anura: Hylidae). Amphibia-Reptilia 16: 215-227.

Paillette M., 1967. — Valeur taxinomique des émissions sonores chez les Hyla (Amphibiens, Anoures) de la

faune française. C. r. Acad. Sci. (D) 264 : 1626-1628.

Parent G. H., 1981. — Matériaux pour une herpétofaune de l'Europe occidentale. Contribution à la révision

chorologique de l'herpétofaune de la France et du Benelux. Bull. Soc. linn. Lyon 50 (3) : 86-111.

PETIT G., 1965. — Conrad Gesner, zoologiste. Gesnerus 22: 195-204.

RIVERO J. A., 1969. — A new name for Sphaenorhynchus aurantiacus (Daudin) (Amphibia, Salientia). Copeia

1969: 700-703.

ROESEL VON ROSENHOF A. I., 1758. — Historia Naluralis Ranarum Nostratium. Nürnberg, Johann Joseph Fleisch-

mann: [i-viii] + i-viii + 1-116, pl. I-XXIV + frontispiece.

RONDELET G., 1558. — Histoire entière des poissons. Seconde partie. Lion, Mace Monhome : [i-iv] + 1-181 +

[i-ix],

RONDELETIUS G., 1555. — Universae Aquatilium Historia. Pars altera. Lugduni, Matthiam Bonhomme: [i-xii]

+ 1-242 + [i-ix],

SALZMANN C., 1965. — Conrad GeBners Persônlichkeit. Gesnerus 22: 115-133.

Schneider H., 1968. — Bio-akustiche Untersuchungen am Mittelmeerlaubfrosch. Z. vergl. Physiol. 61: 369-385.

SEBA A., 1734. — Locupletissimi rerum naturalium Thesauri accurata descriptio, et iconibus artificiosissimis

expressio. per universam physices historiam. Tomus I. Amstelaedami, Janssonio-Waesbergios. Wetstenium

& Smith: [i-xxxiv] + 1-178, pl. I-CXI + 2 unnumbered pl.

— 1735. — Locupletissimi rerum naturalium Thesauri accurata descriptio, et iconibus artificiosissimis expres¬

sio, per universam physices historiam. Tomus II. Amstelaedami, Janssonio-Waesbergios, Wetstenium &

Smith: [i-xxxiv] + 1-154, pl. I-CXIV.

SHAW G., 1802. — General zoology, or systematic natural history. Vol. Ill, part I. Amphibia. London, Kearsley:

i-viii + 1-312, pl. 1-86.

SONNINI C. S. & LATREILLE P. A., 1801. — Histoire naturelle des reptiles. Tome II. Paris, Deterville : [i-iii] +

1-332, 19 pl.

STEJNEGER L., 1907. — Herpetology of Japan and adjacent territory. Bull. J. S. nat. Mus. 58: i-xx + 1-577, pl.

I-XXXV.

WAGLER J., 1833. — Deutung der in Seba’s Thesauro rerum naturalium T. 1 et 2 enthaltenen Abbildungen von

Lurchen, mit critischen Bemerkungen. Isis von Oken 1833: 885-905.

Source : MNHN, Paris

— 340 —

INDEX TO SCIENTIFIC NAMES OF AMPHIBIANS

“americana parva", “ Rana ” 331

“americana rubra", "Rana" 324-325, 331

“americana rubra”, “Ranula" 325, 331

arborea arborea, Hyla 334

arborea var. meridionaiis, Hyla 321-322

arborea, Hyla 322, 328, 332, 334-336

arborea, Hyla arborea 334

“arborea", “Rana" 327, 333, 336

arborea, Rana 321-326, 328-329, 331-334

“arborea", “Rana viridis” 325, 332

auraniiaca, Hyla 321, 328-329

bilineala, Rana 321, 332

“brasiliensis gracilis", “Rana" 324-325, 328

Calamila punctata 333

“ Calamite ” 325-326

cinerea, Hyla 332

Discodactylus 329

Discodactylus ruber 329

eurhostus, Sphaenorhynchus 321, 328-329

“fissis”, “Rana palmis tetradactylis" 333

“fissis", “Rana pedibus" 324, 326, 332

gibbosa. Hyla 321, 330

“ gibbosa ”, “Rana" 324, 327, 335

“gracilis", “Rana brasiliensis” 324-325, 328

Hyla 321-322, 329, 333-336

Hyla arborea 322, 328, 332, 334-336

Hyla arborea arborea 334

Hyla arborea var. meridionaiis 321-322

Hyla aurantiaca 321, 328-329

Hyla cinerea 332

Hyla gibbosa 321, 330

Hyla inframaculata 333

Hyla leucophyllata 332

Hyla meridionaiis 326, 328, 334

Hyla punctata 333

Hyla ranaeformis 321, 330

Hyla rubra 321, 329, 331

Hyla sceleton 321, 328-329

Hyla viridis 321, 330, 332, 336

hyla, Rana 321, 324, 334-335

Hylidae 322, 333

inframaculata, Hyla 333

lacteus, Sphaenorhynchus 329

“ lemnia ”, “Rana" 329

leucophyllata, Hyla 332

“maxima virginiana", “Rana" 329

meridionaiis, Hyla 326, 328, 334

meridionaiis, Hyla arborea var. 321-322

“Muta" 325

“palmis tetradactylis fissis", “Rana" 333

“parva", “Rana americana" 331

“pedibus fissis”, “Rana” 324, 326, 332

(Pelophylax), Rana 330, 334

pentadactyla. Rana 321, 329

punctata, Calamila 333

punctata, Hyla 333

“Rana" 325

Rana 323, 335

“Rana americana parva” 332

“Rana americana rubra " 324-325, 331

“Rana arborea" 327, 333, 336

Rana arborea 321-326, 328-336

Rana bilineata 321, 332

“Rana brasiliensis gracilis" 324-325, 328

“Rana gibbosa" 324, 327, 335

Rana hyla 321, 324, 335

“Rana lemnia" 330

“Rana maxima virginiana ” 329

“Rana palmis tetradactylis fissis" 333

“Rana pedibus fissis" 324, 326, 332

Rana pentadactyla 321, 329

“Rana susinamensis ” 331

Rana temporaria 335

"Rana viridis arborea" 325, 332

Rana (Pelophylax) 330, 334

Rana IRana) temporaria 335

(Rana) temporaria, Rana 335

ranaeformis, Hyla 321, 330

“Ranula americana rubra" 325, 331

“Ranunculus viridis ” 324-327, 335

ruber, Discodactylus 329

rubra, Hyla 321, 329, 331

“rubra", Rana americana 324-325, 331

“rubra", “Ranula americana" 325, 331

sceleton, Hyla 321, 328-329

Sphaenorhynchus eurhostus 321, 328-329

Sphaenorhynchus lacteus 329

“surinamensis”, “Rana" 331

temporaria, Rana 335

temporaria, Rana (Rana) 335

“tetradactylis fissis”, “Rana palmis ” 333

“virginiana", “Rana maxima" 329

“viridis arborea", “Rana” 325, 332

viridis. Hyla 321, 330, 332, 335-336

“viridis", “Ranunculus" 324-327, 335

Source : MNHN, Paris

Bulletin du Musé h

um national d Histoire naturelle, Paris, 4' série, 18, 1996

Section A, n M 3-4 : 341-365

Sipuncula from the Indian Ocean and New Caledonia

by Edward B. CUTLER & Norma J. CUTLER

Abstract. — Collections of over 4200 sipunculans made by six different teams of French biologists from

depths of 1-5340 m are assigned to thirty-four species. The diverse Indian Ocean locations were mostly around

the Comoran Islands and the sub-Antarctic southern islands, plus a few from off Sri Lanka and French Polynesia.

No new taxa are described, but eight have not been previously reported from the Indian Ocean ( Golfingia mar-

garitacea, Nepliasoma culleri, Phascolion lucifugax, P. hibridum , P. tuberculosum, Onchnesoma magnibathum,

O. squantatum, Apionsoma mttrinae bilobatae). Most of these, as well as nine other range extensions, are based

on deep or cold-water material.

Key-words. — Taxonomy. New Caledonia, Indian Ocean, Sipuncula.

Les Sipunculiens de l'océan Indien et de Nouvelle-Calédonie

Résumé. — Les récoltes de plus de 4 200 Sipunculiens, réalisées par six équipes de biologistes français

à des profondeurs de 1 à 5 340 m, ont porté sur trente-quatre espèces. Les divers sites de l’océan Indien étaient

le plus souvent situés autour des Comores et des îles du sud de la région sub-antarctique, et quelques-uns au

large du Sri Lanka et de la Polynésie française. Aucun taxon nouveau n’a été décrit, mais huit étaient déjà

connus de l’océan Indien.

Mots-clés. — Systématique, Nouvelle-Calédonie, océan Indien, Sipunculiens.

E. B. Cutler, Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138 USA & N. J. Cutler, Biology

Department, Hamilton College. Clinton, NY 13323.

INTRODUCTION

While much has been written about the Indian Ocean sipunculan fauna, a good contemporary

starting place is the comprehensive review presented at the 1970 International Symposium on

the Biology of Sipuncula and Echiura (Haldar 1975) in which 135 species were listed (not all

currently considered valid). Since 1975, the report covering the widest geographic range (CUTLER

& CUTLER 1979) was based on 4000 worms (fifty-four species) most collected during the In¬

ternational Indian Ocean Expedition of 1963-1964, plus others by B. Thomassin and associates,

most coming from along the western part of the ocean. The three subsequent reports are more

restricted to shallow water material. One on the Nicobar and Andaman islands (nine species)

was offered by Haldar (1976), while Haldar’s major work (1991) focused on the Indian coastal

fauna (1900 specimens and 37 species). Most recently, Saiz (1993) reported on an interesting

collection, from the “MD-32 La Réunion” project, consisting of seventeen species, four new to

the Indian Ocean.

Source : MNHN, Paris

— 342 —

Between 1972 and 1986, six groups of French biologists made collections of benthic in¬

vertebrates, including more than 4200 sipunculans, from some part of the Indian Ocean. Many

of these expeditions were conducted on board the research vessel Marion Dufresne. Three col¬

lections were processed by the Centre National de Tri d’Océanographie Biologique (CENTOB)

and made available by M. Segonzac. These were: Benthedi, from the northern Mozambique Chan¬

nel (most near the Comoran Islands of Mayotte and Glorieuses) by the vessel Suroit in 1977;

Jasus MD-50, from St Paul and Amsterdam Islands in 1986 under the direction of M. Arnaud,

Marseille; and Safari I & II from south of Madagascar in 1979 and south-west of Sri Lanka in

1981. A series of three collections made by A. Guille, Paris, from the far southern islands came

to us directly in part (Kerguelen Islands, 1972-1974) and partly via CENTOB. The latter were

from MD-04 in 1975 from Kerguelen, MD-08 in 1976 from Marion (Prince Edward Islands),

and Crozet Island. A collection from coral around Reunion was made by P. Delacroix, île de la

Réunion, in 1980. The sixth contribution was made by B. Gout, Montpellier, from Mayotte Island

in the Comorans in 1983 and 1986, and from New Caledonia, French Polynesia in 1985.

This new material covers a very wide variety of Indian Ocean habitats, at latitudes of 2-50°S,

and longitudes of 38-87°E, from warm tropical sand and coral to cold sub-Antarctic mud, and

from depths of 1-5340 m. The inclusion of the New Caledonia material from outside the Indian

Ocean (166.5°E) was to avoid producing another small report on a set of nine species, when

only two of these are not already in this report ( Aspidosiphon spiralis and A. laevis).

None of the material represents new species but seventeen are zoogeographically interesting,

eight new to the Indian Ocean (Golfingm margaritacea, Nephasoma cutleri, Phascolion lucifugax,

P. hibridum, P. tuberculosum, Onchnesoma magnibathum, O. squamatum, Apionsoma murinae

bilobatae), only one coming from shallow water ( P ; lucifugax). Nine species show significant

range extensions ( Nephasoma constrictum , Phascolion convestitum, P. lutense, Onchnesoma

steenstrupii , Phascolosoma noduliferum , Apionsoma murinae murinae , Aspidosiphon muelleri,

A. spiralis, A. zinni). All but one of these latter extensions were into deep or cold water in the

Indian Ocean. The only warm shallow exception being A. spiralis to New Caledonia in the

Pacific.

It is difficult to discuss a limited Indian Ocean fauna since the eastern boundary is so

ill-defined. More commonly one considers an Indo-West Pacific fauna (CUTLER 1994). Also,

this area from Southeast Asia through Indonesia, the Philippines, to Australia is so rich in species

that inclusion of this boundary region would greatly inflate any species list. This brief con¬

sideration of the sipunculan fauna is primarily limited to that area west of the Ninetyeast Ridge

(see Table 1). The exception to this is the fauna from New Caledonia included here for reasons

mentioned above.

One common feature of these collections is the large number of small examples, i.e. trunks

less than 7 mm long. This can represent adult size for a few of these taxa but not most. It seems

clear that the methods used to collect, sort, and process the material have improved so that

smaller representatives are not lost to science. This does make the task of identification more

difficult and in several cases we could not, with certainty, apply specific names. Included in

this report are thirty-four species from ten of the seventeen genera. Eight of the species are

represented by more than 100 specimens, sixteen have from 10-100, while ten species have fewer

than ten representatives in these collections.

Source : MNHN, Paris

— 343 —

In the following section the station data for each species will be summarized, but more

precise data can be found in the Appendix. For a general treatment of this phylum including

synonymies and illustrated keys to all taxa, see CUTLER 1994.

SYSTEMATIC SECTION

Family Sipunculidae Rafinesque, 1814

Genus SIPUNCULUS Linnaeus, 1766

Sipunculus nudus Linnaeus, 1766

The 134 worms were collected around the Comoran Islands (11.5-13°S, 45-47.5°E) at depths

between 5-38 m (one station at 275 m).

Many are less than 10 mm long, and only a few reach a trunk length of 25 mm, none

approaching the more common length of 50-150 mm. All are white, transparent, and with only

twenty-three or twenty-four longitudinal muscle bands which is at the low end of the range for

this species. Some of the very smallest appear to be recently settled larvae. Given the small

size of this material it is possible that the distinctions between S. nudus and S. robustus are not

yet developed ( e.g . nephridial attachment and brain processes) so this identification is somewhat

tenuous.

Distribution

This has been collected from temperate, sub-tropical and tropical waters in all oceans. Most

are from intertidal to 30 m, but a few records are from 100-900 m. Therefore, this material is

not outside the known range.

Family Golfingiidae Stephen & Edmonds', 1972

Genus GOLFINGIA Lankester, 1885

Subgenus GOLFINGIA Lankester, 1885

Golfingia (Golfingia) margaritacea (Sars, 1851)

Sipunculus margaritaceus Sars, 1851. Phascolosoma margaritaceum Danielssen & Koren,

1877. Golfingia margaritacea; Fisher 1952; Saiz 1995. Golfingia margaritacea margaritacea ;

Cutler & Cutler 1987; Cutler 1994.

I. Dr. Stanley J. Edmonds, who contributed much to our knowledge about the biology of Sipuncula, died at age 86 in

July, 1995.

Source : MNHN, Paris

— 344 —

Phascolosoma ohlini Theel, 1911. Golfingia ohlini; WESENBERG-LUND 1955. Golfingia

margaritacea ohlini; Cutler & Cutler 1987; Cutler 1994.

All 617 worms were collected from sub-Antarctic waters (46-50°S), 254 came from Marion

(38°E, 95-204 m), thirty-six were collected near the Crozet Islands (49.5-50.5°E, 192-275 m),

the remaining 327 came from around the Kerguelen Islands (68.5-70.5°E) at 17-230 m.

Nomenclatural note

When Cutler & Cutler (1987) reviewed the genus Golfingia they retained two subspecies,

but acknowledged the ill-defined nature of the boundary between the two taxa. This boundary

was based partially on a non-overlapping range, but largely on two morphological differences:

the presence/absence of introvert hooks and the shape of the posterior end of the trunk. Saiz

( 1995) examined a large collection from the Weddell Sea ranging in length from 1.3-110 mm.

He confirmed the deciduous nature of the hooks as those less than 24 mm carried hooks, but

larger ones are hookless, and the end of the trunks graded from pointed to round. Saiz concluded

that the retention of subspecific rank can not be upheld and we here concur with his judgement.

The trunk length of this material ranged from 2-12 mm, but most are in the 3-7 mm range,

quite small for this species. While both ends are opaque the body wall in the middle of the

trunk is transparent. The introverts are 65-135% of the trunk length and may or may not bear

small (30-45 pm) clear hooks. The pattern of presence/absence was less consistent than that

reported above by Saiz (1995) since even in these worms smaller than 10 mm, hooks are not

always present. Most worm have eight to twelve small tentacles but one 3 mm worms exhibits

sixteen. The posterior ends vary from round to pointed, a very small tail occasionally present.

Sexual maturity seems to come early in this population since even in worms as small as 4 mm,

gametes are present, eggs in some, sperm in others.

Distribution

Widely distributed in the Atlantic, Arctic, and Antarctic oceans (80°N to 78°S), and less

commonly in the northern and southern Pacific. They have been found at depths of 1-5300 m,

but most come from less than 300 m. The deeper records are mostly at lower latitudes making

this an example of tropical submergence, not bipolar distribution. The species was unknown

from the Mediterranean Sea, and the Indian Ocean per se.

While this species has been reported from Indian Ocean longitudes, the southern latitudes

have been in the 66-67° range (STEPHEN 1948 as G. mawsoni). Therefore, while the border

between the Antarctic and the Indian Oceans is not precise, the Southwest Indian Ridge can

serve as that limit. These current records are the first in this part of the world, from north of

the 50th parallel, i.e. the Southwest Ridge. Thus, this is a new record for the Indian Ocean.

Golfingia (Golfingia) muricaudata (Southern, 1913)

These eighty worms came from five areas: eighteen worms near Mayotte (11.5-13°S,

45-47.5°E) at depths of 325-3716 m, two worms from south of Sri Lanka (5.5°S, 78.5°E) at

Source : MNHN, Paris

— 345 —

3660 m, and nine worms near Amsterdam and St Paul Islands (38-39°S, 77-78°E) at 315-1685 m.

From further south, there are eleven worms from Marion Island (47°S, 38°E) at 204 m, and forty

worms off the Kerguelens (49°S, 67-69.5°E) at 230-315 m.

The trunks are 2-35 mm long (most under 15 mm) and exhibit the distinctive posterior

papillae and tail (0.5-4 mm) characteristic for this species. The small deciduous hooks may be

present (in smaller) or absent (in larger worms). The body walls are transparent to translucent

in the under 15 mm animals, becoming opaque and coarse in larger worms.

Distribution

Common in deep water from the North Atlantic, and it occurs off South Africa and Bouvet

Island. It is known from the northern and southern Pacific. In the Indian Ocean it has been

reported from the east coast of Africa at bathyal depths, but previously unknown from the rest

of the Indian Ocean. Therefore, the Sri Lanka and sub-Antarctic specimens represent a significant

extension of this taxon and suggest a low density, cosmopolitan, cold water distribution.

Subgenus SPINATA Cutler & Cutler, 1987

Golfingia (Spinata) pectinatoides Cutler & Cutler, 1979

The three worms came from off Glorieuses (11.5°S, 47.5°E), at 24 m.

These worms having rings of unidentate hooks with basal spinelets, bilobed nephridia, but

lacking a posteriorly attached spindle muscle and longitudinal muscle bands, clearly belong to

this taxon.

Distribution

Previously known from southern Madagascar and French Polynesia at intertidal depths. This

new material represents only a small northward extension of its western Indian Ocean range.

Genus NEPHASOMA Pergament, 1940

Nephasoma constrictum (Southern, 1913)

Eight of these worms came from near Mayotte (13°N, 45°E), at 350 m. The other nine were

collected around Sri Lanka (2-8°S, 79-87°E) between 1095-4350 m.

The plump spindle-shaped trunks range from 3-40 mm long with introverts about 50-75%

of the trunk length. The peculiar neck-like constriction at the base of the introvert and large,

dark, dome-shaped papillae characterize this species. The hooks appear to be deciduous so that

smaller worms tend to have more than larger worms. Internally the nephridia open a little behind

the anus and the retractor muscles originate in the middle of the trunk. Two worms had been

living in polychaete tubes and several had remnants of a grey mud coating.

Source : MNHN, Paris

— 346 —

Distribution

In the northeastern Atlantic Ocean and western Mediterranean (34-51°N) at 150-4000 m. A

recent find by Saiz (1993) from Réunion at 75-750 m (plus a few to 3500 m) was the first for

the Indian Ocean. Therefore, these records reinforce and extend its range northward and eastward

in the colder water of the Indian Ocean.

Nephasoma cutieri (Murina, 1975)

Most of these thirty-nine worms were collected off Marion Island (47°S, 38°E) at 95-204 m.

The remaining four worms lived further northeast near Sri Lanka (5.5°S, 78.5°E) at 3660 m.

They are thread-like animals, the largest being 6 mm long, with introverts 50-75% of the

trunk length. This introvert length helps distinguish it from the similar N. constricticervix that

has a much shorter introvert. The pair of very short nephridia and medium-large hooks (50-

150 pm) are typical for this species.

Distribution

Scattered Pacific Ocean reports from 80°S to 11°N, at depths between 2600-4600 m. There¬

fore, this report is the first for the Indian Ocean and a significant extension of its longitudinal

and bathymetric range.

Nephasoma diaphanes diaphanes (Gerould, 1913)

Of these 561 worms, 382 came from around the Comoran Islands (11.5-13°N, 45-47.5°E)

at 250-1390 m. Eight worms came from off Sri Lanka (1.5-6°S, 78.5-87°E) at depths of 3660-

5175 m (the only species from over 4500 m in these collections), and thirty-five were collected

near St Paul and Amsterdam Islands (37.5-39°S, 77.5°E) between 262-2200 m. From the sub-

Antarctic islands there are seventy-six worms from Marion Island (47°S, 38°E) at 31-192 m,

forty-nine worms from the Crozet Islands (46-47°S, 49.5-51.5°E) at 143-1230 m, and eleven

from the Kerguelen Islands (49-50°S, 68.5-70.5°E) at 43-195 m.

The present material may contain some animals that might be called N. abyssorum (with

smooth white skin) or N. diaphanes corrugatum (with rough brown skin), if they are in better

condition. A few of the worms from the far southern waters had been living in arenaceous

foraminiferan tests and exhibited larger dome-shaped dark papillae. These have slender, straight

trunks 2-12 mm long (most less than 5 mm) that are often transparent. The introvert bears scat¬

tered, small, bluntly pointed hooks and a few tentacular lobes.

Distribution

Cosmopolitan in cold water, most from bathyal and abyssal depths (down to 5300 m). These

new data reinforce its presence in the Indian and sub-Antarctic oceans.

Source : MNHN, Paris

— 347 —

Family Phascolionidae Cutler & Gibbs, 1985

Genus PHASCOLION Théel, 1875

Subgenus ISOMYA Cutler & Cutler, 1985

Phascolion (Isomya) con vest itum Sluiter, 1902

Seven of these eight worms were collected off New Caledonia (22.5°S, 166.5°E) at 10-28 m,

while a single worm came from Mayotte Island (12.5°S, 45°E) at 27 m. All had been living in

a shelter such as an empty gastropod shell.

Since some of these are badly damaged and the trunks are only 1.5-4 mm long, we have

some reservations about this identification. Nevertheless, we place these here given the equal

size of the retractor muscles, the broad-based, recurved, 40-60 pm. Type II Phascolion hooks

(fig. 30E in Cutler 1994), and the weakly developed holdfast papillae. The papillae around the

anterior end of the trunk are large and crowded, thus, except for the smaller hook size, they

strongly resembles P. tuberculosum.

Distribution

Mediterranean, Red Sea, Gulf of Aden and Indonesia, from 25-275 m. New reports by Saiz

from the Weddell Sea (74.5°S, 30°W) at 600-700 m indicate the presence of this species in the

southern seas. Therefore, this material extends the known range further south in the Indian, and

further east into the Pacific Ocean.

Phascolion (Isomya) lucifugax Selenka & de Man, 1883

All four worms came from Mayotte Island (12.5°S, 45-46.5°E) at 5-25 m, and had been

living in empty gastropod shells.

This uncommon species has blunt Type III hooks (fig. 301 in CUTLER 1994) and no obvious

holdfast papillae. Another diagnostic feature is the numerous tentacles, usually more than thirty.

The anterior trunk papillae are dome-shaped and crowded, but otherwise the trunk appears

smooth. The two equal sized retractor muscles originate very close to the posterior end. The

intestine does not form coils but has irregular loops only.

Distribution

Philippines and northern Japan at unknown depths, probably less than 100 m. This material

is the first record from the Indian Ocean thus a significant range extension.

Phascolion (Isomya) tuberculosum Théel, 1875

The forty-six specimens were collected from three regions: thirty-nine came from Glorieuses

(11.5-13°S, 45-47.5°E) at 23-440 m; two from St Paul and Amsterdam Islands (38-39°S, 77.5°E)

at 262-315 m; five worms from New Caledonia (22.5°S, 166.5°E) at 22 m.

Source : MNHN, Paris

— 348 —

The two equal sized retractor muscles place these worms in this subgenus. Despite their

very small size (1-2 mm) most do show the large, recurved, dark. Type II hooks (Fig. 30F in

Cutler 1994). Pale, round holdfast papillae resembling thick crepes that characterize this species

are present on many, but others show only large bulbous papillae, some are almost smooth, and

a few of the smallest still carry the larval array of numerous, very small papillae. When tentacles

are visible there are 10-12 short ones serving to differentiate this from P. lucifugax. Most trunks

are white to transparent and the gut is without regular coils.

Distribution

Common in the northeastern Atlantic at depths of 25-2700 m. The few specimens from

Japan and New Zealand from 93-300 m suggest a low-density population in the Pacific, a pattern

reinforced by this New Caledonia record. This new material also represents the first records

from the Indian Ocean, a significant extension.

Subgenus MONTUGA, Gibbs, 1985

Phascolion (Montuga) lutense Selenka, 1885

While one of these twenty-eight worms was found southeast of Madagascar (32°S, 48.5°E)

at 3825 m, most came from the area around St Paul and Amsterdam Islands (37.5-38.5°S, 77.5°E)

at 975-2200 m.

The smooth, almost transparent skin and the distinctive grey “cap” at the anterior end of

the trunk is typical for this species. They create soft cocoon-like mud tubes and, this collection,

are 2-9 mm long. Internally the esophagus leaves the retractor column prior to its subdivision,

and the gut does form into coils.

Distribution

Generally a cold-water species (1800-6860 m) that appears to be absent in the lower lati¬

tudes. In the southern Hemisphere it has been found in the Pacific Ocean between 36-66°S, and

from 20-32°S in the Atlantic. In the Indian Ocean the only prior record is from the southern

end of the Mozambique Channel (32.5°S, 13°E). In the northern Hemisphere it is known from

the northwestern Pacific and the northeastern Atlantic, 47-56°N. Therefore, this material rein¬

forces its presence in the southern Indian Ocean with a significant eastward extension.

Subgenus PHASCOLION Théel, 1875

Phascolion (Phascolion) hibridum Murina, 1981

These forty-five worms were collected near St Paul and Amsterdam Islands (38-39°S, 77.5°E)

at 262-975 m.

Source : MNHN, Paris

— 349 —

These worms, that had been living in empty gastropod and scaphopod shells, are up to

10 mm long. They have distinct dark hooks, but lack holdfasts and regular gut coils. The small

tentacles are well-formed and the anterior end of the trunk is surrounded by an aggregation of

tall, pointed, dark papillae giving the impression of a shield. The single (fused) large dorsal

retractor muscle originates near the posterior end of the trunk.

Distribution

Malaysia and Samoa at 1500-2380 m. These worms constitute not only a significant west¬

ward extension of the known range, but also the first Indian Ocean record. The water depth is

somewhat shallower but, due to the latitude, the temperature is still cold.

Phascolion (Phascolion) strombus strombus (Montagu, 1804)

326 of these 332 worms were living around the Comoran Islands (mostly from Mayotte,

11.5-13°S, 46-47.5°E) at 250-1390 m, in discarded gastropod shells. The remaining six were col¬

lected off the Kerguelen Islands (47-49.5°S, 70-71.5°E) at 31-1390 m.

While the holdfast papillae of this most common Phascolion are variable, most are U-shaped

with pointed, thin, hardened borders. Some are rounded with the hardened protein distributed

over much of its dome-like surface, reminding one of P. widens but with only a single point.

Since these are small worms their body walls are largely transparent. Around the anterior trunk

are large, mammiform, dark reddish papillae. Type I hooks (fig. 30A in Cutler 1994) and fewer

than 20 tentacles are present. The two retractor muscles are of very different dimensions and

the gut is arranged in loops.

Distribution

Very common and eurytopic in the North Atlantic and Arctic Oceans. There are scattered

records from the Caribbean, Mediterranean, and Red Sea, the Gulf of Aden, southwest Indian

Ocean, Argentina, and Chile. Other records exist from the far South Pacific, and off Japan. It

is known from depths of 1-4030 m, most commonly between 50-2000 m. These new worms con¬

firm the presence of this taxon in the western and far southern Indian Ocean.

Genus ONCHNESOMA Koren & Danielssen, 1875

Onchnesoma magnibathum Cutler, 1969

Both worms came from very deep water, one from east of Madagascar (32°S, 48.5°E) at

3825 m, and the other from southwest of Sri Lanka (1.5°S, 87°E) at 4420 m.

The characteristic posterior radiating folds of skin or “keels”, and a trunk lacking papillae

that tapers into the introvert are distinctive in these specimens.

Source : MNHN, Paris

— 350 —

Distribution

Widespread in the Atlantic Ocean with one record from the southeastern Pacific. Generally

from depths between 3000-5500 m. This report constitutes the first from the Indian Ocean.

Onchnesoma squamatum (Koren & Danielssen, 1875)

These twelve worms came from one station off Mayotte (12.5°S, 45°E) at 325 m.

The spherical trunks are 2-5 mm long and mostly covered with closely packed large gray

papillae. A few worms have smooth patches where these papillae were no longer present.

Distribution

Known from the northern Atlantic (commonly 150-1400 m, a few down to 2300 m), and

Mediterranean Sea (shallow water, 10-55 m). Thus, these worms are another first record from

the Indian Ocean.

Onchnesoma steenstrupii Koren & Danielssen, 1875

The 145 worms were collected near Mayotte (12.5-13°S, 45°E) at 235-815 m.

The spherical trunks are 2-5 mm long and have a series of keels radiating out from the

posterior tip composed of tiny plate-like papillae. The introvert is several times the length of

the trunk and internally it appears to have a single retractor muscle originating very near the

posterior end.

Distribution

Found in much of the Atlantic Ocean, (23°S to 57°N), and in the higher latitudes of the

western Pacific, and the southwestern Indian Oceans (35°S, 22.5°E). In general it inhabits waters

at bathyal depths on continental slopes (100-1600 m, rarely 40-3000 m). So this new material

extends the range further up into the western Indian Ocean.

Family Phascolosomatidae Stephen & Edmonds, 1972

Genus PHASCOLOSOMA Leuckart, 1828

Phascolosoma nigrescens (Keferstein, 1865)

The fifteen worms were present in three collections at depths of 1-30 m: nine came from

Réunion (21.5°S, 56°E), five from the Comoran Islands (12.5-13°S, 45.5-46.5°E), and one from

New Caledonia (22.5°S, 166.5°E).

These 4-30 mm worms have body walls that vary from almost transparent in the smallest,

to a uniform pale brown color. One significant attribute is the many rings of hooks (more than

fifty) that have a distinctive internal structure. Often the rings extend to near the base of the

Source : MNHN, Paris

— 351 —

introvert, which is longer than the trunk. The unremarkable trunk papillae are of uniform size

and of the same color as the skin.

Distribution

Very widespread circumtropical species; generally between 30°N and 30°S, in shallow waters

of the world’s oceans. Thus, this material fills in within the known range.

Phascolosoma noduliferum Stimpson, 1855

These eleven worms came from near the St Paul and Amsterdam Islands (38°S, 77.5°E) at

depths of 75-90 m.

The trunks are only 3-5 mm long making them difficult to identify with confidence. They

do have large papillae with the distinctive platelets, and the hooks are in many rings varying

somewhat in detail. They are referred to this species with some hesitation.

Distribution

Intertidal from southern Australia and Tasmania, plus deeper water off the Philippines, New

Guinea and New Zealand. This record is, therefore, a major westward extension into the southern

Indian Ocean.

Phascolosoma scolops (Selenka & de Man, 1883)

Most of these worms (24/29) were collected on Réunion (21.5°S, 56°E) from intertidal

depths, while the remaining five came from the Comoran Islands (11.5-12.5°S, 46-47.5°E) at

25 m.

The trunk lengths are from 3-21 mm, most less than 10 mm and most are white with darker

red dome to cone-shaped papillae that are larger and more concentrated at each end. A few of

the very smallest appear to be recently settled juveniles. The long introvert has dark patches

along its dorsal surface and fewer than twenty-five rings of hooks.

Distribution

Common throughout the Indo-West Pacific, including northern Japan, south to northern

Australia, and east to Hawaii. Usually found at intertidal and shallow depths (< 30 m). Recently

reported from Reunion at 1-10 m (Saiz 1993). Some reports from off western Africa exist but

these may be misidentified P. stephensoni. This material fits into the known range.

Phascolosoma stephensoni (Stephen, 1942)

The two worms came from the intertidal zone of Réunion (21.5°S, 56°E).

Source : MNHN, Paris

— 352 —

Neither of these worms were complete being represented by the anterior portion only. The

identification is based on the unique hooks with both a clear streak and crescent, and the cone-

shaped preanal papillae that are smooth and red.

Distribution

In the Mediterranean Sea, eastern Atlantic from the Azores south, western and northwestern

Indian Ocean (Durban, Mozambique, Somalia, southwestern India), and western Pacific. These

Reunion collections are a small extension of the known range into the central Indian Ocean.

Genus ANTILLESOMA (Stephen & Edmonds, 1972)

Antillesoma antillarum (Grube & Oersted, 1858)

Five of these six worms lived on Réunion (21.5°S, 56°E) in intertidal water, and the sixth

was collected off Mayotte (12.5°S, 45.5°E) at 8 m.

These worms are small for this species being only 4-6 mm long. Nevertheless, they do

exhibit the large dark papillae over most of the body, a short introvert with no hooks, bushy

tentacles, and many contractile vessel villi that characterize this species.

Distribution

This is a cosmopolitan species found in tropical and subtropical, intertidal and shallow

waters, including many locations in the Indian Ocean. The most recent report is also from Réunion

(Saiz 1993), so this new material only reinforces our prior knowledge.

Genus APIONSOMA Sluiter, 1902

Apionsoma misakianum (Ikeda, 1904)

Most of these 192 worms were collected off New Caledonia (22.5°S, 166.5°E) at 8-28 m,

fifty-four came from the Comoran Islands (11.5-13°S, 45-47.5°E) at 15-38 m, and four had been

living on Réunion (21.5°S, 56°E) between 1-15 m.

The slender, 2-10 mm, spindle-shaped worms have very long thin introverts bearing rings

of hooks near the tip. The round posterior papillae help distinguish this from the hookless but

similar A. trichocephala, both having four slender retractor muscles and bilobed nephridia. The

small hooks have the comb-like array of basal spinelets.

Distribution

Widespread but not common in shallow tropical and subtropical waters of the Indian (East

Africa, Madagascar, western India, western Australia), Pacific, and western Atlantic oceans. Most

recently Saiz (1993) has reported this from Réunion at 58-77 m, so these new specimens merely

confirm its presences in the Indo-West Pacific.

Source : MNHN, Paris

— 353 —

Apionsoma murinae murinae (Cutler, 1969)

The waters around the Comoran Islands yielded twenty-six of these forty-three worms (11.5-

13°S, 45-47.5°E) at 250-1125 m. Another sixteen came from near St Paul and Amsterdam Islands

(38-38.5°S, 77.5°E) at 1430-2200 m, and the deepest specimen was collected south of Sri Lanka

(5.5°S, 78.5°E) at 3600 m.

The large mammiform papillae on the posterior quarter of these plump, spindle-shaped,

2-10 mm worms with single lobed nephridia, are characteristic. Internally the four very thin

retractor muscles originate close to the ventral nerve cord.

Distribution

Widespread in the northern Atlantic, down to 15°S on the eastern side at 1000-5200 m. In

the Pacific it has been reported in the Bering Sea and in deep waters of the southern Pacific.

It is also known from both sides of the Indian Ocean (Kenya, Philippines, Indonesia, at 300-

600 m).

These Indian Ocean records are from Cutler (1977), but have been mistakenly credited

to A. murinae bilobatae in subsequent works including Cutler 1994. So while this new material

nicely fills in large gaps within the Indian Ocean, it is not new for the ocean. This does complete

the circle showing this to be well established around the globe.

Apionsoma murinae bilobatae (Cutler, 1969)

Two samples yielded sixty-eight worms from Mayotte (13°S, 45.5°E) at 350-625 m.

The presence of a short second lobe on the nephridia (10-40% of the primary lobe), and

a flask-shaped trunk with pale thin walls and rounded posterior end differentiate this taxon from

the nominate form.

Distribution

A northern Atlantic subspecies that is more restricted than the nominate form (300-1900 m).

In the Mediterranean it has been found at 160-1230 m. Earlier attributions of this taxon from

the Indian Ocean were errors, see A. murinae murinae. Therefore, this is the first record from

the Indian Ocean and a significant extension of its known range.

Apionsoma trichocephalus Sluiter, 1902

Most of the 286 worms came from the Comoran Islands (12.5-13°S, 45-46.5°E) at 2-32 m.

The remaining forty-three were collected off New Caledonia (22.5°S, 166.5°E) at 8-22 m.

This small (2-10 mm) slender, spindle-shaped, sand-dwelling worm is the only member of

this genus lacking hooks on the thin introvert that is 6-8 times the length of the trunk. Tentacles

also appear to be lacking. The trunk is smoother than the similar hook-bearing A. misakianum

Source : MNHN, Paris

— 354 —

with inconspicuous papillae. Several of the worms from Mayotte, even those with trunks only

5 mm long, are carrying large eggs.

Distribution

Southeastern United States and Gulf of Mexico, West and South Africa, around the western

and northern margins of the Indian Ocean, Vietnam, Japan, Indonesia, Australia, New Zealand,

and Costa Rica (unknown from the central Pacific). Found at intertidal depths to 100 m. These

new collections reaffirms its presence in the western Indian Ocean while the New Caledonia

records complement other Pacific Ocean reports.

Family ASPIDOSIPHONIDAE Baird, 1868

Genus ASPIDOSIPHON Diesing, 1851

Sub-genus ASPIDOSIPHON s. str.

Aspidosiphon (Aspidosiphon) elegans (Chamisso & Eysenhardt, 1821)

All twenty-seven specimens were collected from intertidal water on Réunion (21.5°S, 56°E).

While several of the worms are damaged or incomplete, trunk lengths are from 2.5-20 mm,

but most are less than 10 mm long with almost transparent skin. The ungrooved, uniformly granu¬

lar anal shields are golden in color. Hooks of two types are present. The Type A, compressed,

recurved, bidentate hooks in rings, and the Type C, conical, scattered hooks on the dorsal side

of the introvert that are less dark in the smaller worms (figs 55A & 62A in Cutler 1994).

Distribution

Widespread and common in the Indian and western Pacific oceans to Hawaii, and in the

Caribbean. Lives in dead coral and soft rock in shallow waters. These new records reinforce

their presence in the central Indian Ocean.

Aspidosiphon (Aspidosiphon) gracilis gracilis (Baird, 1868)

Three of these six worms were taken from Mayotte (12.5°S, 45°E) at 2-25 m, and the others

came from New Caledonia (22.5°S, 166.5°E) at 16 m.

Partially due to their small size (2-8 mm trunks) these worms are referred here with some

reservation. The distinguishing attributes are the weakly developed ungrooved anal shield and

the presence of only unidentate hooks. One 6 mm specimen died with the introvert fully extended

showing six short lobes for nuchal tentacles.

Distribution

The northern and eastern Indian Ocean to the Philippines, generally in intertidal coral rock.

Therefore, this questionable material extends the range southward on the western side and a

little further into the western Pacific Ocean.

Source : MNHN, Paris

— 355 —

Aspidosiphon (Aspidosiphon) muelleri Diesing, 1851

This is by far the most commonly encountered species in these collections coming from

four areas: the Comoran Islands yielded 1346 worms (11.5-13°S, 45-47.5°E) at 5-705 m, four

worms were collected near the Kerguelens (49.5°S, 70°E) at 31 m, seven worms were found off

St Paul and Amsterdam Islands (38.5°S, 77.5°E) at 135-575 m, and forty-four came from New

Caledonia (22.5°S, 166.5°E) at 10-28 m. About 1300 of the 1400 specimens were collected from

depths greater than 250 m. Most had been living in gastropod and some in scaphopod shells

while at least a few had been living in coral rock.

Most of these worms are transparent or white and 1-4 mm long. Some are 5-10 mm long,

only a few are in the 10-25 mm range. The smallest worms, around 1 mm long, still exhibit

larval papillae and are clearly recently settled juveniles. Since most of these worms are less

than 5 mm they have just the beginning of grooves and patches of hardened protein on their

shields. The smallest have only light brown dots of denser material, with intermediate forms,

from pale to the typical heavy dark shields in those over 15 mm long. The small array of cone-

shaped spines along the ventral margin of the anal shield was present in some of the larger

worms but not evident on most of those under 5 mm long. Those with undeveloped shields can

look very similar to some Phascolion species. Only unidentate compressed hooks are present,

and in a few specimens scattered hooks existed along the whole introvert. Eggs were present

in worms as small as 4 and 6 mm.

Distribution

Common in the northeastern Atlantic (48-10°N), through the Mediterranean and Red seas,

and along the east coast of Africa to Madagascar and South Africa. Sri Lanka and Indonesia

are the other Indian Ocean records. There are reports from the western margin of the Pacific

from Japan through southeast Asia to Australia and the Kermadec Islands. It seems to be absent

from most of the Pacific Ocean; one record near Chile. In the western Atlantic only one record

from southern Brazil is known. Often found at depths of 5-400 m, occasionally as deep as 1000 m

and rarely as deep as 2900 m. Most often found in discarded gastropod or scaphopod shells.

Some from shallow, warm water live in the bases of solitary corals (previously known as A.

corallicola or A. jukesii). Thus, while this Comoran material is not surprising, the few from the

far southern islands constitute an extension into that part of this ocean and the New Caledonia

worms constitute a small extension into the Pacific.

Aspidosiphon (Aspidosiphon) spiralis Sluiter, 1902

These twenty-five worms all came from New Caledonia (22.5°S, 166.5°E) at 13-22 m, and

had been living in gastropod shells.

This rare species was originally based on only four worms and the 4-7 mm animals we

have are much smaller than the 40 mm type specimen. Ours differ in a few ways. The main

factor pointing to this species (and differentiating it from the similar A. muelleri ) is the ungrooved

anal shield that is made up of irregular squares and dark patches that spread past the anus and

over the anterior end of the trunk. The anterior ventral edge shows a few small cone-shaped

Source : MNHN, Paris

— 356 —

spines on some worms. Rings of hooks, present in the type, could not be located in this material.

We found only scattered Type B, pyramidal hooks (15-30 (im) that are dispersed over most of

the introvert.

Distribution

Indonesia, in gastropod shells, at 10-90 m. Thus, this new material, if real, extends its range

a small distance into the western Pacific Ocean.

Sub-genus PARASPIDOSIPHON (Stephen, 1964)

Aspidosiphon (Paraspidosiphon) laevis de Quatrefages, 1865

The four specimens were collected off New Caledonia (22.5°S, 166.5°E) at 11-12 m.

Distinct longitudinal muscle bands can be seen through the body wall in these 8-20 mm

worms. Unidentate Type A hooks are arranged in rings, and the dark anal shield is grooved.

The anterior end of the spindle muscle does branch with the larger strand going to the body

wall posterior to the anus, and the other part continuing anteriorly as in most species. The retractor

muscles originate in the third quarter of the trunk, not near the posterior end as in the similar

A. coyi.

Distribution

Widespread, but low density, in warm waters in the Indo-West Pacific east to Hawaii. Also

present in warm water on both sides of the Atlantic. Inhabits shallow-water coral rock. This

new material fits into the known range.

Aspidosiphon (Paraspidosiphon) steenstrupii Diesing, 1859

Twelve of these thirteen worms were collected from Réunion (21.5°S, 56°E) at 1-20 m,

while a single specimen was found off Mayotte (12.5°S, 45.5°E) at 20 m.

Many of these have trunks measuring 20-27 mm, but a few are in the 5-10 mm range. Some

worms have been damaged and are without a posterior end. The longitudinal muscle bands can

be seen through the body wall, better developed in the larger worms, and the ungrooved, granular

anal shields are of the creamy white variety. The skin at the anterior end of the larger worms

has a rough “corky” appearance.

Distribution

Throughout the western and northern Indian Ocean and the western Pacific islands to Hawaii.

Also collected from the Caribbean, Cape Verde Islands, and the Gulf of Guinea. It lives in

shallow-water coral rocks. This new material does not expand its known range.

Source : MNHN, Paris

— 357 —

Sub-genus AKRIKOS Cutler & Cutler, 1989

Aspidosiphon (Akrikos) thomassini Cutler & Cutler, 1979

All but three of these seventeen worms came from Mayotte (12.5°S, 45°E) at 2-3 m (one

station at the unusual depth of 340 m), the other three were taken at New Caledonia (22.5°S,

166.5°E) at 19 m.

The external morphology of this species is one of the least “aspidosiphonid” in this genus,

sometimes looking more like a Nephasoma species. It has a very weakly developed anal shield,

no caudal shield, and an introvert that is 3-4 times the trunk length coming off at much less

than a right angle to the trunk. Hooks are not present, and one worm has its introvert fully

extended showing previously unseen tentacular lobes. The anterior of these 2-12 mm trunks do

have dispersed papillae and a rough or reticulated appearance. The two thin retractor muscles

originate at, or very near, the posterior end.

Distribution

Madagascar and French Polynesia, in intertidal coral sands. This new material is from the

same or very close to previously known locations. The deeper records are of interest.

Aspidosiphon (Akrikos) zinni 1 Cutler, 1969

The four worms came from one station near St Paul and Amsterdam Islands (34.5°S, 77.5°E)

at 2200 m.

While this species rarely exceeds 5 mm, the largest worm in this collection is 4 mm long.

They are transparent, very thin, and have the characteristic golden-brown, fine-grained, flat anal

shield with distinct borders.

Distribution

Common in the northern Atlantic Ocean at depths of 1100-4400 m, plus two records from

off the Congo River and two from the Indian Ocean (Mozambique Channel and Reunion at

45-1425 m). Often lives in arenaceous foraminiferan tests. This material does extend its range

further into the southern Indian Ocean.

Genus CLOEOSIPHON Grube, 1868

Cloeosiphon aspergiilus (de Quatrefages, 1865)

All six worms were collected on Réunion (21.5°S, 56°E) at 1-30 m.

1. Prof. Donald J. Zinn, for whom this species was named, died at age 85 in September. 1996.

Source : MNHN, Paris

— 358 —

Table 1. — Sipunculan species in this report with Indian Ocean literature records. Regions: We - western (Africa. Madagascar.

Source : MNHN, Paris

— 359 —

REFERENCES

6. 22, 24. 29, 30. 56

2, 3, 8, 33, 41, 46, 54, 56

5. 8, 23, 24, 29, 37, 45, 49, 54

3, 8, 12, 13, 24, 30, 40, 41, 45, 46, 54

A. murinae murinae

I. murinae bilobalae

Family Aspidosiphonidae

Genus ASPIDOSIPHON

i. (Akrikos) Ihomassini #

A. (Paraspidosiphon) la

i. (Paraspidosiphon) steenstrupii

Genus CLOEOSIPHON

This is probably the easiest sipunculan to identify with its distinctive white cone-shaped

anal shield made up of closely set diamond-shaped units with the introvert protruding through

its center along the main trunk axis.

Distribution

Widespread in the Indo-West Pacific stopping west of Hawaii. Lives in coral rock. The

most recent record is also from Réunion (Saiz 1993) so this material does not add to the known

range.

REFERENCES

Numbers relate to numbers in Table 1.

1. AUGENER H., 1903. — Beitrage zur Kenntnis der Gephyreen nach Untersuchung der im Gottinger zoologischen

Museum befindlichen Sipunculiden und Echiuriden. Arch, fiir Naiurg. 69 : 297-371.

2. BOUVIER E. L„ 1894. — Un nouveau cas de commensalisme: Association de vers du genre Aspidosiphon

avec des polypes madréporaires et un mollusque bivalve. C. R. Acad. Sri., Paris 119: 96-98.

3. CUTLER E. B„ 1965. — Sipunculids of Madagascar, cah. ORSTOM, Océanogr. 3 (4): 51-63.

4. — 1969. — New species of Sipuncula from the western North Atlantic. Proc. Biol. Soc. Wash. 82 : 209-218.

5. — 1977a. — The bathyal and abyssal Sipuncula. Galaihea Rep. 14 : 135-156.

Source : MNHN, Paris

— 360 —

6. — 1977b. — Shallow water sipunculans collected by the Galathea Expedition (Sipuncula). Steenstrupia 4:

151-155.

7. — 1994. — The Sipuncula, their systematics. biology, and evolution. Cornell University Press. Ithaca, NY,

437 p.

8. CUTLER E. B. & CUTLER N. J., 1979. — Madagascar and Indian Ocean Sipuncula. Bull. Mus. nail. Hist.

not.. Paris, 4' série, A I (4): 941-990.

9. — 1980. — Sipuncula collected by the R/V Verna. J. Zoo!., 190: 193-209.

10. CUTLER E. B. & GIBBS P. E., 1985. — A phylogenetic analysis of higher taxa in the phylum Sipuncula. Svst.

Zool. 34: 162-173.

11. CUTLER e. B. & Kirsteuer E., 1968. — Additional notes on some Sipuncula from Madagascar. Results of

the Australian Indo-West Pacific Expedition, 1959-1960, part 12. Zool. An:. 180: 352-356.

12. FISCHER W., 1892. — Übersicht der von Herm Dr. F. Stuhlmann auf Sanzibar und an der gegeniiberliegenden

Festlandskiiste gesammelten Gephyreen. Jahrb. Hamb. Wiss. Anst. 9 (2): 79-89.

13. — 1895. — Die Gephyreen des Naturhistorischen Museums zu Hamburg. Abh. Geb. Naturwiss., Hamburg

13: 1-24.

14. —- 1913. — Über einige Sipunculiden des Naturhistorischen Museums zu Hamburg. Jahrb. Hamb. Wiss.

Anst. 30 (2): 93-101.

15. — 1914. —- Weitere Mitteilungen iiber die Gephyreen des Naturhistorischen (Zoologischen) Museums zu

Hamburg. Jahrb. Hamb. Wiss. Anst. 31: 1-28.

16. — 1920. — Gephyreen der antarktischen und subantarktischen Meere. Deutsche Siidpolar Expedition 16.

Zoologie 8: 409-430.

17. — 1922. — Gephyreen der deutschen Tiefsee Expedition, auf des Dampfer “Valdivia" 1898-1899. Wiss.

Ergebn. Deulsch. Tiefsee Exped.. Valdivia 22 (1): 1-26.

18. — 1923a. — Zwei Sipunculiden des Kapgebietes nebst einer Zustammenstellung der bekannten Arten dieses

Gebietes, und ihrer faunistischen Beziehung zur Ost- und Westkuste. Gôteborgs Kungl. Vetensk. Vitterhets-

Samhalles Hand!. 25 (4): 1-8.

19. — 1923b. — Gephyreen des Golfes von Siam. Vidensk. medd. Dansk nat. hist. Foren. 76: 21-27.

20. GEROULD J. H., 1913. — The sipunculids of the eastern coast of North America. Proc. U.S. Nat. Mus. 44:

373-437.

21. GIBBS P. E., 1985. — On the genus Phascolion with particular reference to the northeast Atlantic species.

J. Mar. Biol. Ass. U. K. 65: 311-323.

22. GRUBE E., 1868b. — Beschreibungen einiger von Georg Ritter von Frauenfeld gesammelter Anneliden und

Gephyreen des rothen Meeres. Verh. Raise Konigli. Zool. Bot. Ges. Wien 18: 629-650.

23. HALDAR B. P., 1976. — Sipuncula from the Andaman and Nicobar islands. Rec. Zool. Surv. of India 70:

109.

24. — 1991. — Sipunculans of the Indian coast. Mem. Zool. Surv. of India 17 (4): 1-169.

25. HAMMARSTEN O., 1915. — Gephryeen von Madagaskar gesammelt van W. Kaudern 1911-1912. Ark. Zool.

9 (10) 26: 1-3.

26. HÉRUBEL M. A., 1904. — Liste des sipunculides et des échiurides rapportés par M. Ch. Gravier du Golfe

de Tadjourah (Mer Rouge). Bull. Mus. nail Hist, nat., Paris 10 : 562-565.

27. — 1907. — Recherches sur les sipunculides. Mém. Soc. zool. Fr. 20 : 107-418.

28. IKEDA I., 1904. — The Gephyrea of Japan. J. Coll. Sci., lmp. Univ. Tokyo 20 (4): 1-87.

29. JOHNSON P., 1964. — Two new species of Aspidosiphon (Sipunculoidea). Ann. Mag. nat. Hist., ser 13 7:

331-335.

30. LANCHESTER W. F., 1905b. — Gephyrea in the marine fauna of Zanzibar and British East Africa from the

collections made by C. Crossland in 1901-1902. Proc. Zool. Soc. Lond. 1: 28-35.

31. LEROY P., 1936. — Les sipunculiens du Muséum National d'Histoire Naturelle de Paris. Bull. Mus. nail

Hist, nat., Paris, ser. 2 8 : 423-426.

32. MURINA V. V., 1967. — New facts about the distribution of marine sipunculid worms in the Red Sea and

the Gulf of Aden. In Some results of Expedition III to the Red Sea Inbuma. Express information. Academy

of Sciences UkSSR: 52-56.

33. — 1970. — Contributions to the fauna of sipunculid worms from the Aden Gulf. Vest. Zool. 2: 65-71.

Source : MNHN, Paris

— 361 —

34. — 1972. — Contribution of the sipunculid fauna of the Southern Hemisphere. Zool. Inst. akad. Nauk SSSR

11 (19): 294-314.

35. — 1975. — New taxa of the genus Golfingia. Zool. Zh. 54: 1085-1089.

36. — 1981a. — First report of the sea worms Sipuncula and Echiura from littoral Somali. Vest. Zool. 1: 11-18.

37. — 1981. — New species of the genus Phaseolion (Sipuncula) from the shells of Scaphopoda. Zool. Zh. 60:

348-352.

38. PERGAMENT T. S., 1940. — On a new genus Nephasoma from the Arctic Ocean. Results of cruising expedition

on icebreaker G. Sedov 1937-1940. Arctic Scientific Institute of Arctic Sea Route. Council of Ministry

USSR: 1-3. [Reprinted 1946, in Trudy ekspedition Sedov Moskva-Leningrad 3: 189-193.)

39. PRASHAD B„ 1936. — On the Sipuncula from Indian water in Indian Museum, Calcutta. Rec. Ind. Mus..

Zool. Surv. India, Calcutta: 231-238.

40. QUATREFAGES M.A. DE, 1865b. — Gephy riens Inermes. Histoire Naturelle des Annelés Marins et d’Eau

Douce, Paris 2 : 599-632.

41. SAIZ SAUNAS J. I., 1993. — Sipuncula from Réunion Island (Indian Ocean). J. Nat. Hist. 27: 535-555.

42. — 1995. — Sipuncula of the southeastern Weddell Sea (Antarctica). Polar Biol. 15: 307-317.

43. SELENKA E., 1885. — Report on the Gephyrea collected by H.M.S. Challenger during 1873-76. Report of

scientific results of the Voyage of Challenger, Zoology 13 (36): 1-25.

44. SELENKA E„ MAN J. G. de & BÜLOW C., 1883. — Die Sipunculiden, eine systematische Monographie.

Reisen in Archipel Phillippinen von Dr. C. Semper, Weisbaden 2 4: 1-131.

45. SHIPLEY A. E., 1902. — Sipunculoidea, with an account of a new genus Lithacrosiphon. In J. S. GARDINER

(ed.). Fauna and geography of the Maidive and Laccadive archipelagoes. Cambridge, Cambridge University

Press: 131-140.

46. — 1903. — Report on the Gephyrea collected by Prof. Herdman at Ceylon in 1902. Ceylon Pearl Oyster

Fish. Suppl. Rep. 3: 169-176.

47. SLUITER C. P, 1902. — Die Sipunculiden und Echiuriden der Siboga Expedition, nebst Zusammenstellung

der Ueberdies aus den indischen Archipel bekannten Arten. Siboga Expeditie, Monographie 25: 1-53.

48. SOUTHERN R„ 1913. — Gephyrea of the coasts of Ireland. Scientific Investigations of the Fisheries Branch,

Department of Agriculture for Ireland 3: 1-46.

49. STEPHEN A. C., 1942. — The South African intertidal zone and its relation to ocean currents. Notes on the

intertidal sipunculids of Cape Province and Natal. Ann. Natal Mus. 10 (2): 245-256.

50. — 1948. — Sipunculids. British, Australian, New Zealand Antarctic Research Expedition Reports, ser. B 5

(4): 213-220.

51. — 1968. — Sipuncula and Echiura from Nouvelle-Caledonie. Cahiers du Pacifique 10: 44-50.

52. Stephen A. C. & CUTLER E. B., 1969. — On a collection of Sipuncula, Echiura, and Priapulida from South

African waters. Trans. Roy. Soc. South Africa 38: 111-121.

53. Stephen A. C. & Edmonds S. J., 1972. — The phyla Sipuncula and Echiura. London, Trustees of the British

Museum ( Natural History ), 528 p.

54. Stephen A. C. & Robertson J. D„ 1952. — A preliminary report on the Echiuridae and Sipunculidae of

Zanzibar. Proc. Roy. Soc. Edinb. 64 (4): 426-444.

55. THÉEL H., 1911. — Priapulids and sipunculids dredged by the Swedish Antarctic expedition 1901-1903 and

the phenomenon of bipolarity. K. Svenska Vet. Akad. Handl. 47 (1): 3-36.

56. WESENBERG-LUND E., 1959a. — Sipunculoidea and Echiuroidea from Mauritius. Vidensk. Medd. Dansk nat.

hist. Foren. Knbenh. 121: 53-73.

57. — 1959b. — Sipunculoidea and Echiuroidea from Tropical West Africa. Atlantide Report 5: 177-210.

58. — 1963. — South African sipunculids and echiuroids from coastal waters. Vidensk. Medd. Dan. nat. hist.

Foren. Knbenh. 125: 101-146.

Source : MNHN, Paris

— 362 —

APPENDIX — STATION DATA

Genera and species arranged alphabetically. Within species, stations arranged by longitude

from West to East. Source code: 42 = Guille, 50 = Dellacroix, 52 = SAFARI, 55 = BENTHEDI,

75 = JASUS, 77 = Gout.

LATIT LONGIT DEPTH NUMB SORC YR STATION

Antillesoma antillarum

Apionsoma misakianum

Apionsoma murinae murinae

Apionsoma murinae bilobatae

Apionsoma trichocephalus

Aspidosiphon elegans

Aspidosiphon gracilis

Aspidosiphon laevis

Source : MNHN, Paris

— 363

Aspidosiphon muelleri

Aspidosiphon spiralis

Aspidosiphon steenstrupii

Aspidosiphon thomassini

Aspidosiphon zinni

Cloeosiphon aspergillus

Golfingia margaritacea

310SMK

Source : MNHN, Paris

— 364 —

Nephasoma constrictum

BB219/20- F

85SMK

Nephasoma cutleri

Nephasoma diaphanes

4926 07018

Golfingia pectinatoides

Onchnesoma^ magnibathum

Onchnesoma souamatum

Source : MNHN, Paris

— 365 —

Onchnesoma

steenstrupii

1255 04516 0350 1 55 77

Phascolion

convestitum

2220 16630 (

Phascolion

hibridum

Phascolion

lucifuaax

Phascolion

lutense

3152 04829 •

«0 l 75 86

Phascolion

strombus

;!« jjjjjj j

||j 'I | ”

Phascolion

tuberculosum

Phascolosoma nigrescens

Phascolosoma noduliferum

3747 07734 0075 7 75 66 3

Phascolosoma scolops

Phascolosoma stephensoni

Sipunculus nudus

Source : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle. Paris, 4' série, 18, 1996

Section A, n“ 3-4: 367-385

Morphogenèse d 'Heligmosomoides polygyrus polygyrus

(Dujardin, 1845) (Trichostrongylina, Heligmosomoidea) chez

Apodemus flavicollis en France. Comparaison avec les espèces

proches : Heligmosomoides laevis (Dujardin, 1845) et

Heligmosomum mixtum Schulz, 1954, parasites d’Arvicolidae

par Gabriel N’ZOBADILA, Johnny BOYER & Marie-Claude DURETTE-DESSET

Résumé. — Seulement trois espèces d'Heligmosomidae sont communes dans la faune de France : Helig¬

mosomoides laevis parasite de Microtus arvalis, Heligmosomoides polygyrus polygyrus parasite d’Apodemus spp.

et de Mus musculus, Heligmosomum mixtum parasite de Clethrionomys glareolus. La morphogenèse de ces trois

espèces est décrite chez leurs hôtes naturels respectifs. La morphogenèse larvaire ne confirme pas la position

phylétique des adultes puisque les synlophes de la Lt d'Heligmosomoides laevis et d'Heligmosomum mixtum

sont comparables et s’opposent à ceux d'Heligmosomoides polygyrus polygyrus et d'Heligmosomoides yamagutii.

Les deux genres Heligmosomoides (crêtes longitudinales) et Heligmosomum (crêtes obliques sur au moins une

des quatre faces du corps) sont donc extrêmement proches. Nous pensons donc que la nouvelle subdivision du

genre Heligmosomum en deux sous-genres, Heligmosomum et Pseudoheligmosomum, ne peut être acceptée, d’au¬

tant plus que l’espèce-type du sous-genre Pseudoheligmosomum a été mal identifiée : en effet, Heligmosomum

mixtum sensu Asakawa et Satoh, 1987, n’est pas l’espèce de Schulz (1954). Nous proposons de la nommer

Heligmosomum asakawai n.sp.

Mots-clés. — Heligmosomoides, Heligmosomum. Nematoda, Trichostrongylina, Rongeurs, zone paléarctique,

morphogenèse.

Morphogenesis of Heligmosomoides polygyrus polygyrus (Dujardin, 1845)

(Trichostrongylina, Heligmosomoidea), a parasite of Apodemus flavicollis in France.

Comparison with related species: Heligmosomoides laevis (Dujardin, 1845) and

Heligmosomum mixtum Schulz, 1954, parasites of Arvicolidae

Abstract. — Only three species of the Heligmosomidae are common in France. Heligmosomoides laevis,

a parasite of Microtus arvalis, Heligmosomoides polygyrus polygyrus, a parasite of Apodemus spp. and Mus

musculus, and Heligmosomum mixtum, a parasite of Clethrionomys glareolus. The morphogenesis of these three

species is studied in their respective natural hosts. The larval morphogenesis does not confirm the current phyletic

position of the adults, since the synlophes of Heligmosomoides laevis and Heligmosomum mixtum fourth stage

larvae are similar and differ from those of Heligmosomoides polygyrus polygyrus and Heligmosomum yamagutii.

Therefore, the two genera, Heligmosomoides (longitudinal ridges) and Heligmosomum (oblique ridges on at least

one quarter of the body), are very closely related. The subdivision of the genus Heligmosomum into two subgenera

cannot be accepted since the type species of Pseudoheligmosomum was not identified correctly: in fact, Helig¬

mosomum mixtum sensu Asakawa and Satoh, 1987 is not the species as described by Schulz (1954). We propose

to name it Heligmosomum asakawai n.sp.

Key-words. — Heligmosomoides, Heligmosomum, Nematoda, Trichostrongylina, rodents, Palaearctic region,

morphogenesis.

G. N’zobadila, J. Boyer & M.-C. Durette-Desset, Muséum national d'Histoire naturelle, Laboratoire de Biologie parasitaire.

Protistologie. Helminthologie, associé au CNRS (URA 114). 61 rue de Buffon. F-7S23I Paris Cedex 05.

Source : MNHN, Paris

— 368 —

INTRODUCTION

Sur les quarante-six cycles réalisés chez les Trichostrongylina, seulement sept sont connus

chez les Heligmosomoidea (c/. Durette-Desset & Chabaud 1993), qui représentent la sous-

famille morphologiquement la plus évoluée de tous les Trichostrongles. La plupart de ces cycles

concernent des Heligmonellidae, sous-famille des Nippostrongylinae parasites de Rongeurs Myo-

morphes. Actuellement seuls deux cycles biologiques sont connus chez les Heligmosomidae :

celui d’Heligmosomoides polygyrus bakeri Durette-Desset et al., 1972, parasite de Mus musculus

et de Cricetidae en zone néarctique, décrit brièvement pour la première fois par Spurlock (1943),

et celui d 'Heligmosomoides kurilensis kobayashii, parasite d 'Apodemus speciosus au Japon, décrit

par Asakawa (1987). Dans les deux cas, les cycles ont été réalisés chez des animaux d’origine

expérimentale, souris de laboratoire (Mus musculus) pour le premier, souris de laboratoire et

mérions ( Meriones unguiculatus) pour les seconds, ces derniers étant des hôtes inhabituels.

Heligmosomoides polygyrus bakeri est d’après Durette-Desset et al. (1972), une des deux

sous-espèces américaines d'Heligmosomoides polygyrus polygyrus (Dujardin, 1845), parasite -

à'Apodemus spp. et plus exceptionnellement de la Mus musculus en Europe. Dès 1971,

FORRESTER avait noté qu’il existait une différence entre les deux populations, européenne

et américaine. Ce n’est qu’en 1983 qu’HASEGAWA et al. ont clairement exprimé que les très

nombreux travaux réalisés sur Heligmosomoides polygyrus (= Nematospiroides dubius Baylis,

1926) concernaient, non pas la sous-espèce européenne Heligmosomoides polygyrus polygyrus,

mais la sous-espèce américaine Heligmosomoides polygyrus bakeri. La souche, nommée « souche

50» par FORRESTER (1971) a été isolée par Ehrenford en 1950 à partir d’œufs trouvés dans

les fèces de Peromyscus maniculatus en Californie (voir Ehrenford 1954). Depuis 1950, elle

a été entretenue sur souris de laboratoire et distribuée dans le monde entier.

C’est probablement une des raisons pour lesquelles les travaux concernant Heligmosomoides

polygyrus polygyrus sont restés très fragmentaires. Seule la phase libre du cycle a été étudiée

en détail par ROMAN (1951) puis Fahmy (1956). Rappelons cependant que ce sont Elton et al.

(1931) qui ont découvert les premiers, chez Apodemus sylvaticus en Angleterre, la présence de

pseudokystes contenant des Heligmosomoides polygyrus polygyrus. Or la présence de ces pseu¬

dokystes paraît caractéristique du cycle des Heligmosomidae.

Enfin, très récemment, Brown et al. (1994) ont étudié le rythme de production des œufs

d'Heligmosomoides polygyrus polygyrus chez Apodemus sylvaticus.

Il nous a donc paru intéressant d’étudier de façon approfondie le cycle biologique

d'Heligmosomoides polygyrus polygyrus avec un double objectif :

— Heligmosomoides polygyrus polygyrus est une des rares espèces d’Heligmosomidae qui

parasite des Muridae et non des Arvicolidae . Nous avons donc cherché à situer cette espèce

systématiquement et biologiquement par rapport aux deux autres espèces d’Heligmosomidae les

plus communes en France, Heligmosomoides laevis (Dujardin, 1845) et Heligmosomum mixtum

(Schulz, 1954), toutes deux parasites d’Arvicolidae.

1. L’espèce a été signalée à différentes reprises chez des Arvicolidae en Europe occidentale mais il pourrait s'agir d’erreurs

d'identification comme le suggèrent QUINNEL et at. (1991).

Source : MNHN, Paris

— 369 —

— Récemment des études comparatives d’immunité croisée et de variabilité génétique entre

Heligmosomoides polygyrus polygyrus et sa sous-espèce américaine Heligmosomoides polygyrus

bakeri ont été réalisées (Quinell et al. 1991 ; Abu-Madi et al. 1994). Nous avons donc cherché

à savoir s’il existait des différences entre la biologie A'Heligmosomoides polygyrus polygyrus

et celle à'Heligmosomoides polygyrus bakeri.

Pour réaliser ces deux objectifs, nous avons choisi de travailler avec les hôtes naturels de

ces différentes espèces. L'objet du présent travail concerne l’étude de la morphogenèse

d 'Heligmosomoides polygyrus polygyrus comparée à celles d’ Heligmosomoides laevis et

d 'Heligmosomum mixtum.

MATÉRIEL ET MÉTHODES

Les hôtes

Les cycles biologiques des trois espèces de la faune de France ont été établis chez des

hôtes naturels sains, en élevage au laboratoire. Les hôtes infestés naturellement ont fourni les

fèces contenant les œufs donnant les larves de Trichostrongles : Apodemus flavicollis Melchior,

1834 pour le cycle d’Heligmosomoides polygyrus polygyrus, Microtus arvalis Pallas, 1779 pour

le cycle d'Heligmosomoides laevis et Clethrionomys glareolus Schreber, 1780 pour celui

d'Heligmosomum mixtum.

Les coprocultures

Les coprocultures sont faites à partir des œufs se trouvant dans les fèces d’animaux rapportés

de Septfontaines (Doubs-France). Deux méthodes de coproculture ont été utilisées pour obtenir

les stades larvaires libres : la méthode de Brumpt (1922), modifiée par Durette-Desset &

Cassone (1987), qui permet d’obtenir directement des larves infestantes, récoltées une à une à

la pipette, comptées et conservées à l’étuve à 22°C dans de l’eau à pH neutre et la méthode

d’HUBERT & Kerbœuf (1984), qui permet d’étudier la morphogenèse des stades libres.

Modes d’infestation

Tous les rongeurs sont infestés par voie buccale en dose unique avec des larves 3 engainées.

Les larves sont concentrées dans un petit volume d’eau et aspirées à l’aide d'une seringue munie

d’une aiguille à embout recourbé, constituant une sonde au moyen de laquelle elles sont injectées

dans l’estomac du rongeur.

Les Apodemus sont infestés avec 150 larves d’Heligmosomoides polygyrus polygyrus âgées

de une à trois semaines, les Microtus avec 100 larves d'Heligmosomoides laevis âgées de quatre

à douze jours, et les Clethrionomys avec 100 larves d'Heligmosomum mixtum, âgées de six à

seize jours. La disparité du nombre et de l’âge des larves infestantes inoculées est due au fait

que ces deux paramètres sont fonction du matériel disponible (hôtes sains et larves infestantes).

Source : MNHN, Paris

— 370 —

Recherche et récolte des parasites

Les stades libres proviennent du développement des œufs mis en culture. Ils sont recueillis

à midi, puis tous les jours de J| à Jô. Les larves obtenues à ces dates sont mises entre lame et

lamelle, légèrement chauffées à la flamme pour permettre leur extension et leur immobilisation.

Elles sont ensuite dessinées à frais.

Les stades parasites sont recueillis par autopsie des rongeurs infestés expérimentalement.

Lors de l’autopsie, les différents organes, foie, poumons, cœur, estomac et intestin grêle sont

prélevés et dilacérés dans de l’eau physiologique. L’intestin est coupé en quatre parties de lon¬

gueurs équivalentes (notées Int], Int 2 , Int? et InD du duodénum jusqu’au cæcum. Les vers sont

récoltés sous la loupe, comptés et fixés à l’éthanol 10" bouillant dans lequel ils sont conservés.

L’étude du synlophe est faite selon la méthode de Durette-Desset (1985).

Symboles utilisés

Li larve du premier slade;

Lj larve du deuxième slade:

(L3) larve 3 engainée dans la cuticule de la L2:

L3 larve du troisième stade:

(I 4 ) larve 4 engainée dans la cuticule de la L 3 :

La larve du quatrième stade :

(Im) ver immature ou juvénile, enfermé dans la cuticule de la La ;

Im ver immature ou juvénile :

Ad adulte.

RÉSULTATS

Description et identification des stades larvaires libres

Chez les trois espèces, les deux premiers stades sont très proches morphologiquement et

sont différenciés principalement par leur taille. De plus entre la L| et la L 2 , chez Heligmosomoides

laevis, l’ébauche génitale grandit notablement (Fig. 1), chez Heligmosomoides polygyrus poly-

gyrus, les renflements cuticulaires latéraux ont une forme différente (Fig. 2F, G) et, chez Helig-

mosomum mixtum, la queue de la L 2 est beaucoup plus longue (Fig. 3A, B). Le troisième stade

se reconnaît par sa double cuticule puisqu’il reste enfermé dans la cuticule du deuxième stade.

Les sexes des stades libres ne peuvent pas être déterminés.

Les trois stades larvaires ont un pharynx bien différencié. L’œsophage est rhabditoïde chez

la L| et la L 2 , et strongyloïde chez la (L 3 ). L’anneau nerveux est situé au milieu de la longueur

de l’œsophage. Les deirides ne sont visibles que chez les larves d ’Heligmosomoides polygyrus

polygyrus (Fig. 2E), le pore excréteur n'apparaît généralement que chez la larve 3. L’ébauche

génitale est située légèrement en arrière du milieu du corps.

Les principales mensurations des stades libres sont données dans le tableau 1 pour Helig¬

mosomoides laevis , le tableau 2 pour Heligmosomoides polygyrus polygyrus et le tableau 3 pour

Heligmosomum mixtum.

Source : MNHN, Paris

— 371 —

3 . I. — Heligmosomoides laevis (Dujardin, 1845). A-C

C, (Lj) à Jj. D-H : stades larvaires parasites et immatures, coupes Iran

mâle à Ja, le synlophe de la La n'est pas encore formé. F. La mâle à

les coupes de corps sont orientées comme la figure D. d, dos :

la cuticule de la L? ; (La), larve 4 enfermée dans la cuticule

La. Echelles : A-C : 100 pm, D-H : 25 pm.

Ja. G, (1m) femelle à Js. H, 1m. femelle à J 7 . Toutes

g. gauche ; dr, droite ; (Lj), larve 3 enfermée dans

enfermé dans la cuticule de la

Source : MNHN, Paris

— 372 —

Fig. 2. — HeUgmosomoides polygyrus polygyrus (Dujardin. 1845). A, La mâle à J 4 , tête en vue apicale. B. Im. mâle à Je. id.

C-E : stades larvaires libres. C, Li à J2, vue latérale gauche. D, La à J4. vue latérale droite. E, (Lj) à Je. vue latérale droite.

F-L : coupes transversales au milieu du corps. F-G : stades larvaires libres. F, L| à J2. G, L2 à J 4 . H-L : stades larvaires

parasites et immatures. H. Lj femelle à Jj. I. (La) femelle à Jj. J. La mâle à J5. K, (Im) femelle à Je. L. Im mâle à Je. Toutes

les coupes de corps sont orientées comme la figure F. d. dos; v. ventre; g, gauche; dr. droite. (L.i), Larve 3 enfermée dans

la cuticule de la L2; (La), Larve 4 enfermée dans la cuticule de la L3; (Im), ver immature enfermé dans la cuticule de la

La. Échelles : A. B. G-L : 25 pm, C-E : 50 pm. F : 10 pm.

Source : MNHN, Paris

— 373 —

TABLEAU 1. — Principales mensuralions exprimées en pm des stades libres Li, L2, (L3) d 'Hetigmosomoides laevis.

Jours après mise en culture

des œufs

Stades larvaires

(L3)

Nombre de larves mesurées

Longueur du corps

366,5

305 - 406

386

360 - 412

433,5

397 - 475

480

402 - 500

Largeur du corps

24,5

22 - 28

25 - 28

25 - 30

26,5

21 - 32

Hauteur du pharynx

11,5

9 - 13

11,5

8 - 14

13 - 15

10 - 17

Position anneau nerveux

par rapport à la tête

80,5

68 - 108

76 - 85

70 - 90

79,5

75 - 82

Position pore excréteur

par rapport à la tête

79,5

75-82

Longueur de l'œsophage

122,5

108 - 148

121,5

115 - 125

135

122 - 148

133

125 - 138

Longueur de l'ébauche

qénitale

10,5

9 - 14

1 2

10 - 14

13,5

12 - 16

14,5

13 - 16

Dist. ébauche gén. / queue

sur lonqueur du corps

44,5

39 - 50,5

48,5

47 - 49,5

47,5

45,5 - 52

47 - 54

Longueur de la queue

76,5

60 - 87

86,5

85 - 87

90-110

86,5

82 - 93

Identification et Morphogenèse des stades larvaires parasites et des vers immatures

Identification du sexe

À partir de J 2 après l’infestation et très nettement à J 3 , la morphologie et la migration de

l’ébauche génitale de la larve 3 deviennent caractéristiques du sexe, l’ébauche migrant plus pos¬

térieurement chez la femelle que chez le mâle (Fig. 4A, B ; Fig. 5A, B ; Fig. 6 A, F).

Identification des différents stades

Les L 4 se différencient des L 3 par la disparition du pharynx, l’apparition d’un synlophe

probablement liée à l’enroulement du corps, l'apparition d’une épine caudale dans les deux

sexes et, de plus, chez le mâle, par le gonflement de la queue.

Les immatures se différencient des L 4 par l’apparition d’une vésicule céphalique, d'un nou¬

veau synlophe et par la différenciation complète des organes génitaux.

Les mues 3 et 4 se font de la même façon dans les deux sexes : la cuticule de la L 4 présente

des ondulations ventrales, tandis que celle de l’immature présente d’abord des ondulations dor¬

sales, puis ventrales. Ces ondulations correspondent à la formation des crêtes cuticulaires. Puis,

la cuticule, aussi bien chez la L 4 que chez l’immature, se décolle d’abord sur la face ventrale,

puis dans la région caudale, la face dorsale et enfin la région apicale par où sortira le ver.

Source : MNHN, Paris

— 374 —

TABLEAU 2. — Principales mensurations exprimées en pm des stades libres Lt, L;, (Lj) d' Heligmosomoides polygyrus polygyrus.

Jours après mise en culture

des œufs

Stades larvaires

(L3)

Nombre de larves mesurées

Longueur du corps

494

443 - 546

514

440 -548

522,5‘

494 - 633

Largeur du corps

28,5

23 - 34

25 - 30

24 - 36

Hauteur du pharynx

18,5

16 - 20

15 - 19

Position anneau nerveux

par rapport à la tête

91,5

75 - 100

81 - 108

104

85 - 130

Position pore excréteur

par rapport à la tête

115,5

95 - 140

Longueur de l'œsophage

147,5

140 - 158

150

144 - 158

179,5

142 - 230

Longueur de l'ébauche

qénitale

13,5

11-20

10 - 22

12 - 25

Dist. ébauche gén. / queue

sur lonqueur du corps

222,5

167 - 257

229

203 - 250

244

222 - 295

Longueur de la queue

94,5

72 - 102

90,5

87 - 99

84 - 100

Morphogenèse

La morphogenèse des stades larvaires parasites et des immatures apparaît sur la Fig. 4 pour

Heligmosomoides laevis, la Fig. 5 pour Heligmosomoides polygyrus polygyrus et la Fig. 6 pour

Heligmosomum mixtum. Nous ne décrivons ici que le synlophe, les autres caractères étant iden¬

tiques à ceux décrits chez les Nippostrongylinae (voir DURETTE-DESSET & Cassone 1987).

Chez la larve 3, il n’y a pas de véritable synlophe, mais la cuticule chez les trois espèces

porte deux renflements latéraux bifides qui s’étendent tout le long du corps (Figs ID, 2H, 3D).

Chez la larve 4, le synlophe est formé de crêtes cuticulaires longitudinales, naissant en

arrière de la vésicule céphalique et disparaissant au niveau de l’anus ou à l’emplacement de la

vulve chez la femelle et juste en avant de la future bourse caudale chez le mâle. Chez Helig¬

mosomoides laevis (Fig. 1F) et Heligmosomum mixtum (Fig. 3E), le synlophe larvaire est iden¬

tique : il est formé de cinq crêtes cuticulaires dont trois ventrales bien développées et deux

petites crêtes latérales, droite et gauche. La crête latérale droite étant dirigée en sens inverse

des autres crêtes, il existe un axe d’orientation incliné à environ 45° sur l’axe sagittal. Chez

Heligmosomoides polygyrus polygyrus (Fig. 2J), le synlophe est formé de six crêtes cuticulaires

ventrales, dirigées de la droite vers la gauche. Il n’existe pas de crêtes en face de chaque champ

latéral, ni d’axe d’orientation.

Source : MNHN, Paris

— 375 —

Chez l’immature, le synlophe formé est celui de l’adulte. Chez Heligmosomoides laevis, il

comprend dix-sept à vingt-quatre crêtes cuticulaires, dont huit à quinze crêtes dorsales et huit

à dix crêtes ventrales (Fig. 1H). Celui d 'Heligmosomoides poly gyrus poly gyrus est formé de vingt-

quatre à trente-deux crêtes, au nombre de treize à seize sur la face dorsale et de onze à seize

sur la face ventrale (Fig. 2L). Celui d’ Heligmosomum mixtum est formé de vingt et une à vingt-six

crêtes, dont neuf à douze crêtes dorsales obliques et douze à quinze crêtes ventrales longitudinales

(Fig. 3G).

Chez les trois espèces, les crêtes sont orientées de la droite vers la gauche sur les deux

faces, selon un axe d’orientation sub-frontal.

FIG. 3. — Heligmosomum mixtum (Schulz. 1954). A-C : slades larvaires libres en vue latérale gauche. A. L, à h. B. U à h. C,

Lj à Jj. D-G : stades larvaires parasites et immatures, coupes transversales au milieu du corps. D. L3 mâle à J3. E. La femelle

à J7. F, (Im) mâle à J9. le synlophe de l'immature n'est pas encore formé. G, (Im) femelle à J9, les synlophes de la La et

de l'immature sont visibles. Toutes les coupes de corps sont orientées comme la figure E. d. dos: v, ventre; g. gauche; dr,

droite; (L3), Larve 3 enfermée dans la cuticule de la L2; (La). Larve 4 enfermée dans la cuticule de la L3; (Im). ver immature

enfermé dans la cuticule de la La. Échelles : A-B : 100 gm, C : SO gm, D : 20 gm. E-G : 25 gm.

Source : MNHN, Paris

— 376 —

Jours après mise en culture

des œufs

Stades larvaires

(L3)

Nombre de larves mesurées

1 0

Longueur du corps

356

304 - 405

375

332 - 414

393

360 - 457

Largeur du corps

19 - 26

21 - 27

20 - 27

Hauteur du pharynx

13,5

12 - 15

13,5

12 - 17

13,5

11 - 16

Position anneau nerveux

par rapport à la tête

50 - 77

61 - 81

107

Position pore excréteur

par rapport à la tête

76 - 94

75 - 110

Longueur de l'œsophage

105

90-119

106

101 - 120

113

102 - 129

Longueur de l'ébauche

qénitale

6-10

6-11

6-10

Dist. ébauche gén. / queue

186

194

199

sur lonqueur du corps

167 - 219

169 - 234

170

250

Longueur de la queue

56 - 67

67 - 83

65 - 86

Chez les deux Heligmosomoides, les trois crêtes ventrales gauches sont plus développées

que les autres crêtes et le nombre de crêtes est généralement légèrement plus élevé sur la face

dorsale. Chez Heligmosomum mixtion , les crêtes sont de taille équivalente et leur nombre est

plus important sur la face ventrale que sur la face dorsale.

Notons par ailleurs que, comme chez les autres Trichostrongles, les glandes excrétrices de

la L 4 sont très développées et que les papilles labiales externo-latérales n’apparaissent que chez

l’adulte. De plus, ce n’est qu’au stade immature que la queue de la femelle subit une torsion

de 45° par rapport à la vulve, soit vers la gauche chez Heligmosomoides laevis, soit vers la

droite chez Heligmosomoides polygyrus polygyrus et chez Heligmosomum mixtum.

Mensurations

Les principales mensurations des stades libres, des stades larvaires et des immatures sont

données dans les tableaux 1 et 4 pour Heligmosomoides laevis, les tableaux 2 et 5 pour Helig¬

mosomoides polygyrus polygyrus et les tableaux 3 et 6 pour Heligmosomum mixtum.

Source : MNHN, Paris

— 377 —

FIG. 4. — Heligmosomoides laevis (Dujardin. 1845), stades larvaires parasites et immatures. A, Lj mâle à h, vue latérale droite.

B. Lj femelle à h. vue latérale gauche. C, (L4) mâle à Ja. vue latérale gauche. D-G : parties postérieures. D. La femelle à

Js, en vue latérale gauche. E. La mâle à Js, vue latérale droite. F. (Im) femelle à J6. vue latérale droite. G, (lm) mâle à Js,

vue latérale droite. (La), larve 4 enfermée dans la cuticule de la L3; (Im), ver immature enfermé dans la cuticule de la La:

g.e., glande excrétrice : c.d., canal déférent ; test., testicule ; vs., vésicule séminale ; sp„ spicule : vest., vestibule ; sph., sphinc¬

ter; tr„ trompe. Sur la figure D, les flèches indiquent les différentes parties de l'appareil génital femelle : ovaire (ov.), utérus

(ut.), ovéjccteur (ovéj.). Échelles : A-C, E : 100 pm, D : 25 pm, F-G : 50 pm.

Source : MNHN, Paris

— 379 —

FIG. 5. — Heligmosomoides polygyrus polygyrus (Dujardin. 1845), stades larvaires parasites et immatures. A. L3 mâle à J3, vue

latérale droite. B. L_i femelle à J3. vue latérale droite. C, (La) femelle à J4. vue latérale gauche. D. (La) mâle à J4, vue latérale

droite. E. La femelle à J5, vue latérale gauche. F-H : parties postérieures. F. La mâle à J4, vue latérale gauche. G, (Im) mâle

à Js, vue latérale gauche. H. (Im) femelle à J„, vue latérale gauche. (La). Larve 4 enfermée dans la cuticule de la L,. (Im).

ver immature enfermé dans la cuticule de la La: sp., spiculé; ov„ ovaire: ut., utérus; vest., vestibule; sph., sphincter: tr..

trompe; g.e., glande excrétrice. Sur la figure F, les flèches indiquent les différentes parties de l'appareil génital mâle : testicule,

(test.), vésicule séminale (v.s.), canal déférent (c.d.). Échelles : A. E. G-H : SO pm, B-C : 100 pm, D. F : 75 pÆ urce : MNHN ‘ Farls

TABLEAU 5. — Principales mensurations exprimées en pm des stades larvaires parasites et des vers immatures i'Heligmosomoides polygyrus polygyrus de Ji

à J7 après l'infestation d’Apodemus flavicollis. M : mâle, F : femelle.

Source : MNHN, Paris

— 381

FlO. 6 . — Heligmosomum mixtum (Schulz, 1954), siades larvaires parasiles el immatures. A, L3 mâle à J2, vue latérale droite.

B-E : extrémités antérieures, vues latérales droites. L.i femelle à J3. C. La femelle à Js. D, (Im.) mâle à J6. E, 1m mâle à J*.

F-I : extrémités postérieures femelles, vues latérales droites. F, L3 à J3. G, La à J3. H. La à J5. I. Im à J9. J, L3 mâle à J2,

ébauche génitale. K-L : mâle, parties postérieures, vues latérales droites. K, La à J5. L, (Im) à J 6 . (La), larve 4 enfermée

dans la cuticule de la L 3 ; (Im), ver immature enfermé dans la cuticule de la La. sp.. spicule: ovéj., ovéjecleur; ov„ ovaire:

ut., utérus: vest., vestibule; sph., sphincter; tr., trompe; g.e., glande excrétrice. Échelles: A : 50 pm, B : 25 pm, C : 75 pm,

D : 200 pm, E : 150 pm, F-H, K : 100 pm, 1 : 200 pm, J : 30 pm, L : 50 pm.

Source : MNHN, Paris

Tableau 6 . — Principales mensurations exprimées en pm des stades larvaires parasites et des vers immatures d 'Heligmosomum mixtum de J2 à J12 après

l'infestation de Clelhrionomys glareolus. M : mâle, F : femelle.

Temps après l'infestation

J10

Localisation

intestin

Stades parasites

Im.

Nombre de specimens

mesurés

1 3

Longueur du corps

380

684

1020

1300

4350

4490

4450

7700

373 - 387

657 - 711

985 - 1075

1187 - 1475

2800 - 5230

3810 - 5070

3100 - 6580

4100 - 9200

1140

1350

4860

5110

5100

8840

1100 - 1300

1235 - 1420

3700 - 5650

4700 - 5720

4090 - 5970

7700 - 10300

Position anneau nerveux

110

125

148

215

212

245

par rapport à la tête

86 - 88

100 - 120

98 - 152

67 - 119

127 - 182

178 - 239

170 - 350

205 - 309

140

103

173

167

225

230

102 - 162

97 -112

150 - 212

160 - 189

160 - 277

175 - 270

Position pore excréteur

89,5

134

139

105

293

328

320

390

par rapport à la tête

89 - 90

129 - 139

131 - 158

85 - 122

282 - 310

314 - 352

246 - 396

285 - 440

152

110

315

200

385

395

140 - 166

97 - 119

295 - 335

163 - 242

285 - 470

370 - 435

Longueur de l'œsophage

142

219

229

383

419

522

540

557

138 - 146

211 - 226

209 - 254

280 - 459

400 - 447

518 - 534

487 - 594

514 - 594

324

281

450

550

602

556

297 - 345

246 - 320

440 - 470

477 - 590

495 - 677

525 - 595

Longueur de l'ébauche

177

183

1210

génitale

18 - 22

44 - 47

143 - 212

167 - 205

995 - 1360

170

195

2100

127 - 211

95 - 324

1960 - 2230

Longueur de la queue

50 - 52

52 - 57

55 - 59

61 - 66

54 - 66

55 - 69

62 - 71

64 -73

Source : MNHN, Paris

— 383 —

DISCUSSION

La morphogenèse larvaire des genres Heligmosomoides et Heligmosomum révèle leurs très

grandes affinités. Contrairement aux Nippostrongylinae (Heligmosomoidea) déjà connus, la

larve 4 ne possède pas de vésicule céphalique et présente, dans les deux sexes, une épine caudale

que l’on retrouvera chez la femelle adulte. La torsion de la queue ne se fait qu’au stade immature,

de 45°, soit à gauche chez Heligmosomoides laevis, soit à droite chez Heligmosomoides polygyrus

polygyrus et Heligmosomum mixtum. Les synlophes larvaires sont très proches, comme l’avait

déjà observé Asakawa (1990), en décrivant le synlophe de la L 4 A'Heligmosomum yamagutii

Chabaud et al., 1963. Cependant, ce synlophe n’a pas de crête latérale droite, mais trois crêtes

ventrales et une crête latérale gauche. Contrairement à ce que l’auteur a écrit, il n’existe donc

pas d’axe d’orientation, puisque toutes les crêtes sont orientées dans la même direction.

Il est remarquable de constater que les synlophes larvaires d 'Heligmosomoides polygyrus

polygyrus et A'Heligmosomum yamagutii ne possèdent pas d’axe d’orientation, alors qu’il en

existe un chez Heligmosomoides laevis et Heligmosomum mixtum. Du point de vue de l’onto¬

genèse larvaire, rien n’implique donc qu’ Heligmosomum soit plus évolué qu 'Heligmosomoides.

Ceci nous conforte dans l’idée que, du point de vue systématique, la division du genre

Heligmosomum en deux sous-genres, proposée par Asakawa & Satoh (1987), est inutile. De

plus, dans leur article, les auteurs choisissent Heligmosomum mixtum Schulz, 1954 comme

espèce-type du sous-genre Paraheligmosomum. Dans la définition de ce sous-genre, ils s’appuient

sur les caractères du synlophe de spécimens récoltés chez Clethrionomys rutilus mikado au Japon

et qu’ils ont identifiés, par erreur, à Heligmosomum mixtum décrit pour la première fois par

Schulz (1954) chez Clethrionomys rutilus en Russie. En fait, il s’agit d’une autre espèce A'He¬

ligmosomum, puisque le synlophe A'Heligmosomum mixtum décrit par TRAVASSOS & Darriba

(1929) (sous le nom A'Heligmosomum costellatum) chez Clethrionomys rutilus de Russie, puis

par Durette-Desset (1968) et par Genov & Jancev (1981), chez le même hôte et dans la

même région, est caractérisé par des crêtes obliques sur toute la face dorsale et par des crêtes

longitudinales sur toute la face ventrale, alors que les spécimens japonais possèdent des crêtes

longitudinales uniquement sur la face ventrale gauche. Nous proposons de nommer l’espèce

d'Asakawa & Satoh, 1987, Heligmosomum asakawai n.sp. [ = Heligmosomum ( Paraheligmoso¬

mum ) mixtum sensu Asakawa & Satoh, 1987, nec Schulz, 1954]; en effet, la description des

auteurs est suffisamment précise pour en déduire que l’identification comme Heligmosomum mix¬

tum était erronée. Si le sous-genre était utilisé, il serait logique de considérer Heligmosomum

asakawai comme l’espèce-type. Nous pensons personnellement qu’il est plus simple de ne pas

utiliser ce sous-genre.

Dans la faune de France, Heligmosomoides polygyrus polygyrus fait partie du contingent

arrivé en Europe occidentale à partir du centre de dispersion d’Europe orientale (voir DURETTE-

DESSET 1967), avec quelques autres espèces dont les deux seules communes sont Heligmoso¬

moides laevis et Heligmosomum mixtum. Heligmosomoides laevis est bien la forme la plus

primitive avec une bourse caudale symétrique et des crêtes cuticulaires longitudinales. Heligmo¬

somoides polygyrus polygyrus est plus évoluée que la précédente, puisque la bourse caudale est

fortement asymétrique et que le nombre de crêtes est plus élevé que chez Heligmosomoides

laevis. Elle reste cependant très proche de cette espèce, ce qui semblerait indiquer que la capture

Source : MNHN, Paris

— 384 —

par les Muridés s’est effectuée précocément à partir d’espèces proches des formes primitives

comme Heligmosomoides laevis. L’espèce la plus évoluée est Heligmosomum mixtum, puisque

les crêtes dorsales sont obliques et non longitudinales.

La morphogenèse larvaire ne permet donc pas de confirmer la position phylétique des adultes

puisque les synlophes de la L 4 d 'Heligmosomoides laevis et d’ Heligmosomum mixtum sont compa¬

rables et s’opposent à ceux d'Heligmosomoides polygyrus polygyrus et A'Heligmosomoides

yamagutii. Par contre, elle met en évidence les relations étroites existant entre Heligmosomoides

(crêtes longitudinales) et Heligmosomum (crêtes obliques sur au moins une des quatre faces du

corps).

Remerciements

Nous remercions le Dr P. Delattre et Mr. J. P. Quéré du Laboratoire d’éco-éthologie de l’Institut

national de recherches agronomiques de Montpellier, ainsi que le Dr P. Giraudoux, du Laboratoire d’éco¬

logie de l’université de Bourgogne, qui nous ont facilité la capture des rongeurs.

RÉFÉRENCES

Abu-Madi M. A., Reid A. P., LEWIS J. W. & Hominick W. M., 1994. — Genomic variability within laboratory

and wild subspecies of Heligmosomoides polygyrus. J. Helminthol. 68: 93-96.

ASAKAWA M., 1987. — Genus Heligmosomoides Hall, 1916 (Heligmosomidae: Nematoda) from the Japanese

wood mice, Apodemus spp. III. The life cycle of Heligmosomoides kurilensis kobayashii (Nadtochii, 1966)

in ICR mice and preliminary experimental infection to jirds. J. College Dairying 12: 131-140.

— 1990. — Heligmosomum (Paraheligmosomum) yamagutii Chabaud et al.. 1963 (Heligmosomidae: Nematoda),

from red-backed vole, Clethrionomys rex (Microtidae; Rodentia) on Mt. Petegari, Japan. J. Rakuno Gakuen

Univ. 15 : 159-165.

Asakawa M. & Satoh R., 1987. — Discovery of the genus Heligmosomum Raillet et Henry, 1909 (Heligmo¬

somidae: Nematoda) from japanese Clethrionomys and establishment of Paraheligmosomum n. subgen.

J. College Dairying 12: 111-119.

BAYLIS H. A., 1926. — On a trichostrongylid nematode from the wood-mouse ( Apodemus sylvaticus). Ann. Mag.

Nat. Hist. 18: 455-464.

Brown, E. D., Macdonald D. W., Tew T. E. & TODD I. A.. 1994. — Rhythmicity of egg production by Helig¬

mosomoides polygyrus in wild wood mice, Apodemus sylvaticus. J. Helminthol. 68: 105-108.

BRUMPT E., 1922. — Enseignement de parasitologie appliquée au diagnostic, à l'hygiène et à l’épidémiologie.

Masson, Paris, série 3, 7 e leçon 16 : 4-24.

Chabaud A. G., Rausch R. L. & DESSET M.-C., 1963. — Nématodes parasites de Rongeurs et d’insectivores

japonais. Bull. Soc. Zool. France 88 : 489-512.

DUJARDIN F., 1845. — Histoire naturelle des Helminthes ou Vers intestinaux. Paris, XVI, 645 + 15 p.

DURETTE-DESSET M.-C., 1967. — Évolution des Nématodes Héligmosomes en rapport avec celle de leurs hôtes

fondamentaux, les Microtidae. C. R. Acad. Sci., Paris 265 : 1500-1503.

— 1968. — Identification des Strangles de Mulots et de Campagnols décrits par Dujardin. Ann. Parasit. Hum.

Comp. 43 : 387-404.

— 1985. — Trichostrongyloid nematodes and their vertebrate hosts: reconstruction of the phylogeny of a

parasitic group. Adv. Parasitol. 24: 239-306.

DURETTE-DESSET M.-C. & CASSONE J., 1987. — Sur deux Nématodes Trichostrongyioïdes parasites d'un Muridé

africain. III. - Dualité physiologique des larves infestantes, liée aux rythmes saisonniers. Ann. Parasit.

Hum. Comp. 62 : 577-589.

Source : MNHN, Paris

— 385 —

DURETTE-DESSET M.-C., KlNSELLA J. M. & FORRESTER D. J., 1972. — Arguments en faveur de la double origine

des Nématodes néarctiques du genre Heligmosomoides Hall, 1916. Ann. Parasit. Hum. Comp. 62 : 365-382.

Durette-Desset M.-C. & Chabaud A. G., 1993. — Nomenclature des Strongylida au-dessus du groupe-famille.

Ann. Parasit. Hum. Comp. 68 : 111-112.

ELTON C., ford E. B., Baker J. R. & Gardner A. d., 1931. — The health and parasites of a wild mouse

population. Proc. Zool. Soc. London 2: 657-721.

EHRENFORD F. A.. 1954. — The life cycle of Nematospiroides dubius Baylis (Nematoda: Heligmosomidae).

J. Parasitol. 40: 480-481.

Fahmy M. A. M., 1956. — An investigation on the life cycle of Nematospiroides dubius (Nematoda: Heligmo¬

somidae) with special reference to the free-living stages. Z. Parasitenk. 17: 394-399.

FORRESTER D. j., 1971. — Heligmosomoides polygyrus (= Nematospiroides dubius ) from wild rodents of northern

California: natural infections, host specificity and strain characteristics. J. Parasitol. 57: 498-503.

GENOV T. & Yancev Y., 1981. — The morphology and taxonomy of nematodes of the genera Heligmosomoides

Hall, 1916 and Heligmosomum Raillet et Henry. 1909 (Heligmosomidae Cram, 1927) in Bulgaria. Khel-

minthologiya 12: 8-30.

HASEGAWA H., OTSURU M. & TOB1TA K., 1983. — First record of Heligmosomoides polygyrus from Japan

(Nematoda : Heligmosomidae). Jap. J. Parasitol. 32: 419-423.

Hubert J. & Kerbœuf D., 1984. — A new method for culture of larvae used in diagnosis of ruminant gastroin¬

testinal strongylosis: comparison with fecal cultures. Can. J. Comp. Med. 48: 63-71.

QUINNELL R. J., BEHNKE J. M. & KEYMER A. E„ 1991. — Host specificity of and a cross-immunity between

two strains of Heligmosomoides polygyrus. Parasitology 102: 419-427.

ROMAN E., 1951. — Étude écologique et morphologique sur les Acanthocéphales et les Nématodes parasites des

Rats de la région lyonnaise. Mém. Mus. nain. Hist, nat., Paris, sér. À, Zool. 2 : 249-270.

SCHULZ R. S., 1954. — Dictyocaulides et Heligmosomatides, In Skrjabin K. I., SCHIKHOBALOVA N. P. & SCHULZ

R. S-, Osnovy Nematodologii , Moscou 4: 1-123.

SPURLOCK G. M., 1943. — Observations on host -parasite relations between laboratory mice and Nematospiroides

dubius Baylis. J. Parasitol. 25: 303-311.

TRAVASSOS L. & DARRIBA A. R., 1929. — Notas sobre Heligmosominae. Sciencia Medica 7: 432-438.

Source : MNHN, Paris

Bulletin du Muséum national d’Histoire naturelle, Paris, 4 e série, 18 , 1996

Section A, n°‘ 3-4 : 387-400

Cones taken off Wallis and Futuna Islands, South-West Pacific

(Mollusca, Gastropoda, Conidae)

by Robert G. MOOLENBEEK & Dieter RÔCKEL

Abstract, — Deep-water benthos off Wallis and Futuna Islands was sampled by dredging and trawling

during the MUSORSTOM 7 expedition in 1992. Twenty eight species of the genus Conus were obtained, some

of which had been translocated from shallow into deeper water. Four species remain unidentified and eighteen

represent new regional records, the total number of Conus species recorded from this archipelago now standing

at fifty-three. Conus paciftcus n.sp., from 295-600 m, and C. neptunus futunaensis n.ssp., from 370-455 m, are

described.

Key-words. — Gastropoda, Conidae, Pacific, Wallis & Futuna Islands, systematics, new records, new spe¬

cies, new subspecies.

Gastéropodes Conidae du bathyal de Wallis et Futuna, Pacifique Sud

Résumé. — Le benthos profond de la zone économique des îles Wallis et Futuna a été échantillonné par

dragages et chalutages lors de la campagne MUSORSTOM 7 en 1992. Vingt-huit espèces de Conus sont reconnues

dans le matériel récolté, dont quelques-unes du domaine littoral. Quatre espèces ne sont pas identifiées avec

certitude et dix-huit n'étaient pas connues de la région, ce qui porte à cinquante-trois espèces l’inventaire des

Conidae de ce petit archipel. Deux taxons du benthos profond sont décrits comme nouveaux : Conus pacificus

n.sp., récolté entre 295 et 600 m, et C. neptunus futunaensis n.ssp., récolté entre 370 et 455 m.

Mots-clés. — Gastropoda. Conidae, Pacifique, Iles Wallis & Futuna, systématique, nouvelle espèce, nouvelle

sous-espèce.

R. G. Moolenbeek, Zoôlogisch Museum, University of Amsterdam, P.O. Box 94766, GT Amsterdam, The Netherlands.

D. ROckel, Neckaranlage 6, 69412 Eberbaeh am Neckar. Germany.

INTRODUCTION

Our knowledge on the composition and distribution of the deep-water benthos in the Indo-

Pacific is still fragmentary, since very few regions have been adequately sampled. In the present

paper, we record material of the gastropod family Conidae from the slopes of the islands of

Futuna, Alofi (together constituting the Hoorn or Horne Islands) and Wallis, and of the seamounts

lying within the 200 mile economic zone of these islands. The material was obtained by Philippe

Bouchet, Bernard Métivier and Bertrand Richer de Forges on R.V. “ Alis" during the MUS¬

ORSTOM 7 expedition carried out in 1992 by the Muséum national d’Histoire naturelle, Paris,

and the Institut Français de Recherche Scientifique pour le Développement en Coopération (ORS¬

TOM). A total of 142 dredgings and trawlings were carried out in the upper bathyal zone, down

to 1300 m, but the collection also contains some material from less than 100 m depth. A narrative

Source : MNHN, Paris

— 388 —

of the cruise, together with a review of the tectonic history of the area and a station list were

presented by Richer de Forges & Menou (1993).

Abbreviations and text conventions

MNHN Muséum national d'Histoire naturelle. Paris;

ZMA Zoôlogisch Museum Amsterdam;

spm specimen(s), doubtful if alive or dead collected ;

sp species ;

Iv live collected specimen(s);

dd dead collected specimen(s);

stn station ;

RD relative diameter (maximum diameter of last whorl relative to aperture height);

RSH relative spire height (shell length minus aperture height/aperture height).

All material is deposited in the Muséum national d’Histoire naturelle (MNHN, Paris) unless

otherwise stated. We refer to Rockel et al. (1995a) for references to the original descriptions

and other relevant literature concerning the taxa discussed here.

SYSTEMATICS

Family Conidae Fleming, 1822

Genus CONUS Linnaeus, 1758

Conus acutangulus Lamarck, 1810

Material examined. — Waterwitch Bank: stn DW 538, 12°30.8’S - 176°40.3’W, 275-295 m, 1 spm.

Futuna Island: stn DW 497, 14°I9.6’S - 178°04.8’W, 355-369 m, 1 dd.

Remarks

The species has a vast Indo-West Pacific distribution and is known so far from depths of

0.5 m to 100 m. It is uncertain whether the present specimens have been translocated to the

depths of 295-355 m, or represent a deep-water population of the species. The specimen from

Waterwitch Bank is subadult (length 10.4 mm). The specimen from Futuna is adult (length

25.9 mm) and has strong, beaded spiral ribs on the last whorl.

Conus aphrodite Petuch, 1979

Material examined. — Futuna Island: stn DW 511, 14°14.0’S - 178°I1.5’W. 400-450 m. 1 dd.

Remarks

Conus aphrodite is a true deep-water species, known from the Ryukyu Islands (Japan), the

Philippines, and New Caledonia where empty shells are found in 105-370 m. The single specimen

Source : MNHN, Paris

— 389 —

from Futuna (length 15.1 mm), which may have been translocated to 400-450 m depth, represents

the first record from the area.

Conus bullatus Linnaeus, 1758

Conus bullatus — Richard 1983: 19.

Material examined. — Field Bank: stn DW 588, 12°17.3'S - 174°44.6'W, 490-500 m, 1 dd.

Remarks

Conus bullatus is known from East Africa to the Marquesas and Hawaii and is most common

from shallow sublittoral to about 50 m depth, although in the Philippines there are records down

to 240 m. The subadult and dead specimen from Field Bank was probably translocated to

490-500 m.

Conus chiangi (Azuma, 1972)

Material examined. — Futuna Island: stn DW 514. 14°13.3 , S - 178°10.7 - W, 349-355 m, 1 dd.

Remarks

First record from this area. Conus chiangi is a deep-water species found so far in Japan

to the Philippines and New Caledonia, in 200-400 m depth.

Conus corallinus Kicner. 1845

Material examined. — Field Bank: stn DW 597, 12°31.4’S - 174°18.6’W, 469-475 m, 1 dd.

Remarks

New record for this area. Conus corallinus was known from Okinawa (Japan) to Papua

New Guinea and New Caledonia, mainly in subtidal depths but also in deep-water down to

240 m.

Conus crocatus Lamarck, 1810

Material examined. Field Bank: stn DW 596, 12°31.8’S - 174°18.9’W, 32 m, 1 spm.

Remarks

This is an Indo-West Pacific species living in depths of 1-80 m. In the Pacific, recorded

from Japan to the Marshall Islands, Samoa, Papua New Guinea and the Solomon Islands. The

single shell from Field Bank is subadult and probably dead collected.

Source : MNHN, Paris

— 390 —

Conus cylindraceus Broderip & Sowerby I, 1830

Material examined. — Bayonnaise Bank: stn CP 629, 11°53.7’S - 179°32.3’W, 400-420 m, 1 fragment.

Remarks

Conus cylindraceus is known from the Indian Ocean to Polynesia in depths down to about

25 m, indicating that the fragment from Bayonnaise Bank must have been translocated downslope.

Conus dusaveli (H. Adams, 1872)

Material examined. — Field Bank: stn DW 597, 12°31.4’S - 174°18.6’W, 469-475 m, 1 dd.

Wallis Island: stn DW 529, 12°31.4'S - 176°39.6'W. 500 m, 2 dd.

Remarks

First record for the area. Conus dusaveli was earlier known from Ryukyu Islands (Japan)

to the Philippines, and New Caledonia in depths of 50-290 m. One of the specimens from Wallis

Island is a juvenile (length 15.9 mm) that probably represents this species.

Conus eburneus Hwass in Bruguière, 1792

Conus eburneus — Richard 1983: 19.

Material examined. — Wallis Island: stn Lagon sud, 13°22.3’S - 176°11.2’W, 52-55 m, 1 spm.

Remarks

A species common in the Indo-Pacific from East Africa to Polynesia, intertidal to about

70 m, usually in depths of 1-15 m.

Conus eugrammatus Bartsch & Rehder, 1943

Material examined. — Tuscarora Bank: stn DW 556, 11°48.7’S - 178°18.0 , W, 440 m, 1 dd.

Remarks

The single specimen from Tuscarora Bank is in bad condition and its identity is somewhat

equivocal. The presence in this region of Conus eugrammatus , currently known from Japan to

the Philippines, Queensland and Hawaii, would not be unexpected.

Conus floccatus Sowerby II, 1841

Conus floccatus — Richard 1983: 19.

Source : MNHN, Paris

— 391 —

Material examined. — Bayonnaise Bank: sin CP 629, 11°53.7’S - 179°32.3’W, 400-420 m, 1 dd.

Remarks

Conus floccatus is a species with a Western Pacific distribution living in depths of less

than 100 m. The specimen from Bayonnaise Bank has probably been translocated.

Conus generalis Linnaeus, 1767

Material examined. — Bayonnaise Bank: stn DW 625, 11°52.4’S - 179°33.8’W, 425-430 m, 1 juv spm.

Remarks

An Indo-West Pacific species usually living in depths of 1-50 m, in the Philippines recorded

down to 240 m.

Conus imperialis Linnaeus, 1758

Conus imperialis — RICHARD 1983: 19.

Material examined. — Wallis Island: stn DW 529, 12°31.4’S - 176°39.6’W, 500 m, 2 dd.

Field Bank: stn DW 596, 12°31.8’S - 174°18.9’W, 32 m, 1 dd.

Remarks

An Indo-West Pacific species, living intertidally to about 75 m. The specimens found at

500 m depth must have been translocated.

Conus kanakinus Richard, 1983

(Fig. 10)

Conus kanakinus — RÔCKEL et al. 1995b: 575-576.

Material examined. — Wallis Island: stn DW 522, 13°10.7’S - 176°15.0’W, 650-765 m, 2 dd; stn DW

526, 13°13.4’S - 176°15.5’W, 355-360 m, 1 spm.

Waterwitch Bank: stn DW 538, 12°30.8’S - 176°40.3’W, 275-295 m, 1 spm.

Remarks

We provisionally identify the four specimens from Wallis Island and Waterwitch Bank as

Conus kanakinus , so far known only from the New Caledonia region, although doubts remain.

The present specimens differ from the New Caledonian ones in having a less elongate last whorl

and a colour pattern of dashes and lines instead of only spiral lines. On the other hand, the

general facies are similar and the broad, paucispiral protoconch and the spire sculpture are iden-

Source : MNHN, Paris

— 392 —

tical. Two of the specimens are juveniles. The larger ones are 13.7 and 14.2 mm long, and may

be adult.

Conus kimioi (Habe, 1965)

Material examined. — Field Bank: stn DW 589, 12°16.2’S - 174°41.4'W, 400 m, 1 dd.

Futuna Island: sin DW 499, 14°19.6’S - 178°04.6’W, 290-395 m, 1 dd; stn DW 514, 14°13.3’S - 178°10.7’W,

349-355 m, 1 dd.

Remarks

A new regional record of this deep-water species, earlier known from Japan to the Philippines

and New Caledonia, in 120-250 m.

Conus memiae (Habe & Kosuge, 1970)

Material examined. — Futuna Island: stn DW 499, 14°19.6’S - I78°04.6’W, 290-395 m, 1 dd; stn DW

509, 14°14.8’S - 178°11.5’W, 200-240 m, 1 dd.

Wallis Island; stn DW 526, 13°13.4’S - 176°15.5’W, 355-360 m, 1 dd.

Remarks

A new record for this species, earlier known from Japan to the Philippines and Indonesia,

the Solomon Islands and Fiji, generally in depths of 50-250 m.

Conus neptunus futunaensis n.ssp.

(Figs 1-3)

Type material. — Holotype and 4 paratypes MNHN, 1 paratype ZMA 3.95.014, 1 paratype collection

Rôckel.

Type locality. — Futuna Island, R.V. Mis, MUSORSTOM 7, stn DW 497, 14°19.6’S - 178°04.8'W, 355-

369 m, 10 May 1992, P. Bouchet, B. Métivier & B. Richer de Forges coll.

Material examined. Futuna Island: stn DW 497, 14°19.6'S - 178°04.8’W, 355-369 m (2 lv, holotype and

paratype MNHN 44.7 x 19.0 mm); stn DW 499, 14°19.6’S - 178°04.6’W, 290-395 m (2 dd, paratypes MNHN

44.9x21 mm, 31.1 x 12.8 mm); stn DW 504, 14°19.6’S - m^.S’W, 300-390 m (3 dd, paratype ZMA

50.8 x 21.2 mm, coll Rôckel 46.3 x 20.4 mm, MNHN 47.8 x 20.3 mm).

Wallis Island: stn DW 523, 13°12.0 , S - 176°15.6’W, 455-515 m, 1 dd.

Etymology. — Named after its type locality.

Description (holotype)

Shell medium-sized and moderately solid. Last whorl ovate (RD0.52), outline convex,

slightly constricted at the base. Shoulder angulate. Spire of moderate height (RSH 0.17), outline

deeply concave. Protoconch broken, with probably more than two whorls, maximum diameter

0.8 mm. Teleoconch with 10.5 whorls, slightly stepped. Sutural ramp concave, with 2 strong and

Source : MNHN, Paris

— 393 —

1 weak spiral grooves, crossed by close-set axial ribs. First 8 whorls tuberculate. Upper part of

last whorl smooth, basal part with 8-10 strong spiral ribs.

Ground colour cream with irregularly scattered dark brown, curved or angular axial dashes

and triangular spots, concentrated and underlain light brown or light violet flecks on both sides

of centre. Protoconch and base white. Aperture light violet.

Dimensions: shell length 45.5 mm, maximum diameter 19.5 mm, aperture height 37.6 mm.

Remarks

Dimensions of paratypes range from 44.7 to 50.8 mm, RD 0.51-0.53, RSH 0.14-0.19, relative

weight 0.11-0.22 g/mm. Number of protoconch whorls about 3, maximum diameter 0.75-0.8 mm.

Number of teleoconch whorls 10.5-11.5, sutural ramps with 3-5 spiral grooves. Colour pattern

generally sparser, light brown and paler than in the holotype. A light brown band may be present

just below shoulder.

Conus neptunus futunaensis n.ssp. differs from C. neptunus neptunus Reeve, 1843 mainly

in its ovate shape and smaller size. The last whorl of the nominal subspecies is elongately conical

with almost straight sides, while the new subspecies has an ovate shape with convex sides.

C. neptunus neptunus, known so far only from the Philippines and the South China Sea, attains

a length of 80 mm, whereas the largest Futuna specimen is 50.8 mm long.

Conus neptunus futunaensis also resembles C. laterculatus Sowerby III, 1870, C. australis

Holten, 1802, and C. lienardi Bernardi & Crosse, 1861. Conus laterculatus differs in its narrower

and more cylindrical last whorl (RD 0.42-0.51), which is strongly sculptured with grooves and

ribbons from base to shoulder. The spire of Conus laterculatus is generally lower (RHS0.10-

0.16), the colour pattern consists of more regularly arranged spots and dots instead of scattered

lines and triangles. Conus australis can be distinguished by its larger size (shell length 60-

105 mm), its generally narrower last whorl (RD 0.44-0.52) and its heavily sculptured last whorl.

Finally, Conus lienardi, from New Caledonia, is similar in size and shape, occasionally also in

its pattern, but differs in its less pronounced tubercles on the spire and the spiral grooves on

the teleoconch sutural ramp. It also has a more rounded shoulder and an almost smooth last

whorl with a white ground colour. The protoconch of C. neptunus futunaensis has three or more

whorls whereas C. lienardi has only 2 to 2.5 whorls.

<ï •

Conus orbignyi Audouin, 1831

Material examined. — Futuna Island: stn DW 499, 14°19.6’S - 178°04.6'W, 290-395 m, 1 dd; stn DW

500, 14°19.5’S - 178*04.1*W, 350-394 m, 2 dd; stn DW 511, 14°14.0’S - 178°11.5’W, 400-450 m, 1 dd; stn

DW 516, 14°13.5’S - 178*11.6’W. 441-550 m, 1 dd.

Remarks

3 Conus orbignyi is a deep-water species known from depths of 50-400 m. The Futuna Island

population belongs to the nominal subspecies, whereas Conus orbignyi coriolisi Moolenbeek &

Richard, 1995 occurs off New Caledonia and in the Coral Sea. This is a new record for the

area.

Source : MNHN, Paris

— 394 —

Figs 1-7. — New taxa of Conus from Wallis and Fuluna Islands. 1-3, Conus neplunus futunaensis n.ssp., Futuna Island; 1-2,

holotype, length 45.5 mm, dorsal and ventral views; 3, paratype ZMA, length 50.8 mm, ventral view. 4-7, Conus pacificus

n.sp., Bayonnaise Bank; 4-5, holotype, length 20.2 mm, dorsal and ventral views; 6-7, paratype ZMA, Waterwitch Bank,

length 23.8 mm, dorsal and ventral views.

Source : MNHN, Paris

— 395 —

Conus pacificus n.sp.

(Figs 4-7)

Type material. — Holotype and 1 paratype MNHN, 1 paratype ZMA 3.95.013.

Type locality. — Bayonnaise Bank, R.V. “ Alis ", MUSORSTOM 7, sin DW 626, 11°53.6'S - 179°32.0’W,

597-600 m, 29 May 1992. P. Bouchet, B. Métivier & B. Richer de Forges coll.

Material examined. — Waterwitch Bank: sin DW 538, 12°30.8’S - 176°40.3’W, 275-295 m (1 dd, paratype

ZMA, 23.8 mm x 8.7 mm. aperture height 16.5 mm, RD 0.53, RSH 0.31).

Field Bank: stn DW 588, 12°17.3’S - 174°44.6’W, 490-500 m (1 dd, paratype MNHN, 14.4x5.1 mm,

aperture height 10.5 mm, RD 0.49, RSH 0.27).

Etymology. — Named after the Pacific Ocean, where in the deep-sea many unknown animals still await

to be discovered.

Description (holotype)

Shell small, cylindrical, fusiform, smooth, and glossy. Relative diameter (RD) of last whorl

0.49, relative spire height (RSH) 0.27. Protoconch partly broken, maximum diameter 0.8 mm.

Teleoconch with 7.25 whorls, the first three to four whorls with small nodules and two spiral

grooves. On the last whorl only one spiral groove. Shoulder slightly angulate, spire a little convex,

sutural ramp almost flat. A few indistinct basal grooves.

Ground colour white, with light brown axial streaks and very fine spiral lines consisting

of very fine white spots. On the last whorl four continuous axial brown streaks from base to

suture.

Dimensions: shell length 20.2 mm, diameter 7.3 mm, aperture height 14.7 mm.

Remarks

The paratypes agree in shape and colour with the holotype, one paratype (Figs 6-7) has

repaired the severely broken base. None has a well preserved protoconch.

Superficially Conus pacificus n.sp. appears to be most similar to the sympatric species Conus

cylindraceus Broderip & Sowerby 1, 1830, especially in its shape. It differs by the colour which

is a redder brown in C. cylindraceus, and the presence of spiral white spotted lines. The pro¬

toconch of Conus cylindraceus is narrower (diameter 0.7 mm), its spire outline is sigmoid (upper

part concave, following part convex) instead of convex, and the tuberculation of the spire whorls

is less pronounced. Conus auratinus Da Motta, 1982, although attaining a larger size and having

a lower spire, is also similar in shape and pattern and might be a closely related species. Juveniles

of Conus episcopatus Da Motta, 1882 may have a similar colour pattern but always have a more

flat-sided spire.

Conus saecularis Melvill, 1898

Material examined. — Bayonnaise Bank: stn CP 629, 11°53.7’S - 179°32.3’W, 400-420 m, 1 dd; stn DW

625, 11°52.4’S - 179°33.8’W, 425-430 m, 1 dd.

Source : /MNHN, Paris

— 396 —

Remarks

Conus saecularis is a deep-water species, living in depths of 85-400 m in the northwestern

Indian Ocean and the western Pacific from the Ryukyu Islands (Japan) to the Solomon Islands

and Papua New Guinea. This is the first record from the region.

Conus samiae Da Motta, 1982

Material examined. — Futuna Island: stn DW 499, 14°I9.6’S - 178°04.6’W, 290-395 m, 1 dd.

Remarks

The validity of Conus samiae is questionable. In the Philippines, it is found in deep water,

where it may intergrade with C. sulcatus Hwass, 1792, variety bocki Sowerby III, 1881. Therefore

C. samiae may be a form of C. sulcatus. The single specimen from Futuna Island is a strongly

sculptured juvenile (length 26.5 mm) and is only tentatively identified as Conus samiae. This is

the first record from the region of this species otherwise known from the Philippines and the

Solomon Islands.

Conus sponsalis Hwass in Bruguière, 1792

Conus sponsalis — RICHARD 1983: 19.

Material examined. — Combe Bank: stn DW 543, 12°25.6'S - 177°28.2’W, 27-30 m, 1 spm.

Remarks

Conus sponsalis is a wide-spread Indo-Pacific shallow-water species.

Conus sulcocastaneus Kosuge, 1981

Material examined. — Field Bank: stn DW 594, I2°31.0’S - 174°19.9’W, 495-505 m, 1 dd.

Remarks

Conus sulcocastaneus is a deep-water species, found in depths of 120-240 m in the Phi¬

lippines and the Marshall Islands. This is the first record from the area.

Conus tessulatus Born, 1778

Conus tessulatus — Richard 1983: 19.

Material examined. — Combe Bank: stn DW 542, 12°26.4’S - 177°28.2’W, 370 m, I dd.

Field Bank: stn DW 596, 12°31.8’S - 174°18.9’W, 32 m, 1 lv.

Waterwitch Bank: stn DW 538, 12°30.8’S - 176°40.3’W, 275-295 m, 2 dd.

Source : MNHN, Paris

— 397 —

Remarks

An Indo-Pacific species, living intertidally and subtidally to depths of 40 m. The empty

shells from Combe and Waterwitch Banks have most probably been translocated.

Conus sp. 1

(Figs 8-9)

Material examined. — Futuna Island: stn DW 499, 14°19.6’S - 178°04.6’W, 290-395 m, 1 dd; stn DW

512, 14°13.5’S - 178°10.3’W, 210-245 m, 1 dd.

Remarks

The identity of these two shells from Futuna Island is unclear. They superficially resemble

Conus praecellens A. Adams, 1854, but differ in their less pyriform last whorl and particularly

in protoconch characters. While typical Conus praecellens has a paucispiral protoconch with

about 2 whorls, the protoconch of the smaller shell (length 22.3 mm) from Futuna Island is

multispiral with three or more whorls. The protoconch of the larger specimen (length 35.9 mm)

is broken. These two specimens apparently represent a distinct species.

Conus sp. 2

(Fig. 11)

Material examined. — Wallis Island: stn DW 522, 13°10.7’S - 176°15.0’W, 650-765 m, 1 dd.

Remarks

The single specimen is in very bad condition and measures 24.4 mm in length (RD 0.50,

RSH is 0.25). It may prove to be a juvenile of an already described species.

Conus sp. 3

(Fig. 12)

Material examined. — Futuna Island: stn DW 514, 14°13.3’S - 178°10.7’W, 349-355 m, 1 dd.

Remarks

The 16.4 mm high specimen, which may be a juvenile, cannot at present be identified with

any known species.

Conus sp. 4

(Figs 13-14)

Material examined. — Combe Bank: stn DW 539, 12°27.3'S - 177°27.3'W, 700 m, 2 dd; stn CP 552,

12°15.7’S - 177°27.8’W, 786-800 m, 1 dd.

Source : MNHN, Paris

— 398 —

FIGS 8-14. — Deep-water species of Conus from Wallis and Futuna Islands. 8-9. Conus sp. I, Fuluna Island. MUSORSTOM 7,

stn DW 499, length 35.8 mm. ventral and dorsal views. 10, Conus kanakinus, Wallis Island, MUSORSTOM 7, stn DW 522,

length 14.2 mm, dorsal view. 11, Conus sp. 2, Wallis Island, MUSORSTOM 7, stn DW 522, length 24.4 mm. 12, Conus

sp. 3, Futuna Island, MUSORSTOM 7, stn DW 514, length 16.4 mm, dorsal view. 13-14, Conus sp. 4, Combe Bank,

MUSORSTOM 7, stn DW 539, length 23.6 mm, ventral and dorsal views.

Source : MNHN, Paris

— 399 —

Remarks

The three specimens belong to the Conus profundorum complex, which includes Conus

vaubani Rôckel & Moolenbeek, 1995 from the New Caledonia area. They differ from the latter

species in having a brown protoconch and indistinct spiral striae on the spire. They may represent

a local form of Conus vaubani or possibly an undescribed species.

DISCUSSION

Of the twenty-eight species of Conus collected during the MUSORSTOM 7 expedition, thir¬

teen are truly deep-water species, with a main distribution below 100 m, and eleven are shal¬

low-water species represented by translocated shells. In addition, four species, which are not

identifiable with certainty, may belong to the deep-water fauna. This cone fauna appears to be

considerably less diverse than elsewhere in the tropical western Pacific at similar depths, though

it is by no means depauperate. Thirty-nine species are recorded from the New Caledonia region

from depths greater than 100 m (Rôckel et al. 1995b), but this is the result of a much more

intensive sampling effort. The cone fauna of the nearby Fiji Islands is known mainly through

the work of CERNOHORSKY (1964), but the deep-water species have not been sampled.

Richard et al. (1981, 1982) and Richard (1983) have already recorded thirty-seven shal¬

low-water species of Conus (actually thirty-five as we regard two of them as synonyms) from

Wallis and Futuna. There had been no earlier records of deep-water Conidae from there. All

thirteen deep-water species, as well as five shallow-water species collected during the expedition,

are new records for the archipelago, thus bringing the total inventory to fifty-three species of

Conus.

Acknowledgements

The material and information on the MUSORSTOM 7 cruise on which this paper is based was gener¬

ously placed at our disposal by Dr P. Bouchet and Ms V. Héros (both MNHN). We are greatly indebted

to Dr P. Bouchet and Mr B. Marshall (New Zealand) for critically reading the manuscript and for their

comments. Mr M. Filmer (Chobham, U.K.) gave valuable suggestions concerning the English text.

REFERENCES

Cernorhorsky W. O., 1964. — The Conidae of Fiji (Mollusca: Gastropoda). Veliger 7: 61-94.

Richard G„ 1983. — Wallis et Futuna. Ses îles, ses lagons, ses coquillages. Xenophora 18: 9-20.

Richard G., Galzin R., Salvat B., Bagnis R., Bennett J., Denizot M. & Ricard M., 1981. — Geomor-

phology, ecology and socio-economy of the Futuna marine ecosystem (Horn archipelago-Polynesia). Proc.

4th Int. Coral Reef Symp.. Manila 1: 269-274.

Richard G., Bagnis R., Bennett J., Denizot M., Galzin R., Ricard M. & Salvat b., 1982. — Wallis et

Futuna. Etude de l’environnement lagunaire et récifal des îles Wallis et Futuna (Polynésie occidentale).

Rapport définitif. Rapport Ecole Pratique des Hautes Etudes RL9: 1-101.

Source : MNHN, Paris

— 400 —

Richer de Forges B. & MENOU J.-L., 1993. — La campagne MUSORSTOM 7 dans la zone économique des

îles Wallis et Futuna. Compte rendu et liste des stations. In A. CROSNIER (ed.). Résultats des campagnes

MUSORSTOM, volume 10. Mém. Mus. nail Hist. nai.. Paris 156: 9-25.

RÜCKEL D., KORN W. & Kohn A. J., 1995a. — Manual of the living Conidae. Volume I: Indo-Pacific region.

Christa Hemmen, Wiesbaden, 517 p.

RÔCKEL D., Richard G. & Moolenbeek R. G., 1995b. — Deep-water Cones (Gastropoda: Conidae) from the

New Caledonian region. In P. BOUCHET (ed.). Résultats des campagnes MUSORSTOM, volume 14. Mém.

Mus. nail Hist. not.. Paris 167 : 557-594.

Source : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle. Paris. 4 e série, 18. 1996

Section A, n“ 3-4 : 401-408

Guamampa n.g. (Gastropoda, Pulmonata),

a bradybaenid land snail with monadeniid characters

by Anatoly A. SCHILEYKO

Abstract. — Guamampa n.g. is a genus of Aegistinae (Bradybaenidae) characterized by the presence of a

single mucus gland entering the stylophore; the distal portion of vas deferens much enlarged: a flagellum of

peculiar appearance; the penial verge grooved; and the vagina enlarged and coated with dense white cover. The

type species is Helix tuba Albers, 1854 from Sulawesi, Indonesia. Certain characters of Guamampa and of

Tricheulota, from the Philippines, are shared by the American genus Monadenia. Monadeniinae is elevated to

family rank within the Xanthonychoidea.

Key-words. — Gastropoda, Pulmonata, Xanthonychoidea, Monadeniidae, Guamampa, Indonesia, systema-

Guamampa n.g. (Gastropoda, Pulmonata),

un Bradybaenidae avec des caractères de Monadeniidae

Résumé. — Guamampa n.g. est un nouveau genre de Bradybaenidae Aegistinae défini par un ensemble

de caractères de l’appareil génital : une seule glande muqueuse implantée sur le sac du dard; la partie distale

du vas deferens très développée; un flagelle d’un type particulier, ressemblant à un poing; la verge pénienne

ouverte, présentant un profond sillon; le vagin grand, couvert d’une enveloppe externe résistante. L'espèce-type

est Helix tuba Albers, 1854, de Sulawesi (= Célèbes) en Indonésie. Guamampa et Tricheulota, des Philippines,

partagent un certain nombre de caractères anatomiques avec le genre Nord-américain Monadenia. Le taxon

Monadeniinae est élevé au rang de famille à l'intérieur de la superfamille Xanthonychoidea.

Mots-clés. — Gastéropodes, Pulmonés, Xanthonychoidea, Monadeniidae, Guamampa, Indonésie, systéma-

A. A. Schileyko, Institute of Problems of Evolution and Ecology. Leninski Prospect 33. 117071 Moscow. Russia.

INTRODUCTION

The problem of relationships of Eurasian and American helicoid land molluscs has been

the subject of number recent discussions (Miller & Naranjo-Garcia 1989, 1991; NORDSIECK

1987; SCHILEYKO 1991). The author united Asian Bradybaenidae and American Xanthonychidae

under a superfamily Xanthonychoidea (SCHILEYKO 1991). In this group, the condition of the

sexual apparatus found in the subfamily Aegistinae is viewed as plesiomorphic when compared

with that found in the Bradybaeninae, on one hand, and to Xanthonychinae (Xanthonychidae),

on the other hand (Schileyko 1991: 196, fig. 5). The condition observed in the subfamily

Monadeniinae, from northwest America, is thus seen as immediately derived from the condition

in Xanthonychinae. Conversely, Miller & Naranjo-Garcia (1989, 1991) include Monadeniinae

Source : MNHN, Paris

— 402 —

directly into the Asian Bradybaenidae. In this connection, it should be mentioned that more than

100 years ago Pilsbry (1894) pointed out the similarity of the mucus glands of the bradybaenid

Tricheulota, from the Philippines, and of the American Monadenia.

In April 1995 I had an opportunity to dissect the species originally described as Helix tuba

Albers, 1854 from Sulawesi (= Celebes), Indonesia. Peculiar features in the organisation of the

reproductive apparatus lead me to establish a new genus for this species, and to discuss its

systematic position and possible phylogenetic relationships.

Abbreviations

ag albumen gland;

e epiphallus;

f flagellum;

mg mucus gland:

p penis:

pr proslate;

ps penis shealh;

r penial retractor muscle;

s stylophore;

st spermatheca;

vd vas deferens;

vdl slender proximal part of vas deferens:

vd2 distal part of vas deferens;

ve verge.

Family Bradybaenidae Pilsbry, 1939

Subfamily Aegistinae Kuroda & Habe, 1949

GUAMAMPA n.g.

Type species. — Helix tuba Albers, 1854.

Material examined. — About fifteen specimens (four dissected) from limestone outcrop named Gua Mampu,

near Uloe (Kampung Luppang, Desa Cabbeng). 30 km NNE of Watampone. South-West Sulawesi; altitude c.50 m.

Collected by P. Bouchet, MNHN, 10 September 1991. under leaves of aroids.

Etymology. — This genus is named after the locality where the present material was collected.

Description (Fig. 1)

Shell helicoid, depressed, rather thin but solid, somewhat translucent, of about 4.5 whorls;

last whorl evenly rounded at periphery and slightly descending in front. Basic colouration con¬

sisting of yellow background with two brown or reddish bands above and below periphery; umbi¬

licus encircled by area of same dark colour; in addition, two ill-defined bands, darker than

background, one between adapical band and suture, the other between abapical band and circu-

mumbilical area. Band width varies from reduced to hypertrophied. Initial part (0.3-0.4 whorl)

of embryonic whorls smooth and polished, subsequently regularly radially wrinkled. Postnuclear

surface nearly smooth, densely covered with very short golden hairs arranged in oblique series.

Aperture wide, with broadly expanded and reflected whitish or pinkish lip. Umbilicus open,

rather narrow. Height 17-20 mm, diameter 30-36 mm.

Talon, a small vesicle on a long slender duct, lying on surface of albumen gland. Vas deferens

consisting of two parts: a long slender duct arising from prostate and an enlarged club-shaped

portion entering epiphallus apically. Boundary between vas deferens and epiphallus marked by

Source : MNHN, Paris

— 403 —

flagellum of peculiar appearance, somewhat resembling a human fist with extended forefinger.

Flagellum containing internally a series of narrow cavities entering the principal lumen of the

duct at right angle, or nearly so. Epiphallus cylindrical, penial retractor attached to its middle

portion. Apical part of the penis more or less bulbous, containing a verge in form of longitudinally

folded fleshy plate, i.e. it is not closed, but grooved. Inner surface of distal part of penial tube

covered by rows of prismatic tubercles. Stylophore voluminous, the only tubercular-alveolar

mucus gland entering its lower part via thin duct; accessory sac absent. Vagina enlarged, with

white dense external layer and thick walls filled with loose fibrous tissue. Spermathecal duct

nearly cylindrical, spermathecal head lying in situ on surface of spermoviduct.

Discussion

Besides the type species, Guamampa probably also include the following nominal species,

as described and illustrated by Sarasin & Sarasin (1899): Helix zonalis Férussac, 1821;

H. exceptinucula Beck, 1837, H. expansa L. Pfeiffer, 1861, H. halmaherica Kobelt, 1892, and

H. surrecta Kobelt, 1894. All of these, and H. tuba , are usually placed in Planispira

(Camaenidae), see e.g. Richardson (1985). However, the occurrence of a stylophore, in asso¬

ciation with other morphological genital characters, clearly excludes Guamampa from the

Source : MNHN, Paris

— 404 —

Camaenidae and allows its inclusion in the Bradydaenidae. In this family the subfamilies Aegis-

tinae and Bradybaeninae differ by the presence (Aegistinae) or absence (Bradybaeninae) of the

flagellum and by the occurrence in the Aegistinae of a small, but closed, tubular verge in the

penis. The unusual structure of the flagellum described above and the latter character clearly

connects Guamampa to the Aegistinae, a subfamily with taxa distributed throughout southern

Asia including China, Japan, the Philippines and Indonesia.

From my dissections of Aegista subchinensis (Moellendorff, 1884), which is obviously very

close to A. chinensis (Philippi, 1845), the type species of Aegista, the unusual structure of the

flagellum described above in Guamampa is shared with Aegista. However other genital characters

allow distinction between the two genera (Figs 2D, 3).

Genital morphology in Guamampa also resembles that observed in Tricheulota Pilsbry, 1895,

type species (OD) Chloritis spinosissima Semper, 1880. The history of the type species is a

little complicated. Semper (1873, pi. 14, figs9a-b) first illustrated the reproductive anatomy of

a species identified by him as Helix sanziana Hombron & Jacquinot, 1841. Later (Semper 1874,

pi. 9, fig. 10) he published an illustration of a shell identified as Helix sanziana var. When the

relevant text, with reference to the two aforementioned illustrations, was issued much later

(Semper 1880: 235), Semper had changed the identification to Chloritis spinosissima n.sp.

Whatever the real identity of the type species is, this would not change the present discussion

because Helix sanziana and Tricheulota spinosissima are congeneric and most probably closely

related. Conchological and genital characters are very similar in Guamampa and Tricheulota

(Fig. 2A, B), but the flagellum exhibits the usual conical shape and the enlarged portion of the

vas deferens is very short in Tricheulota. judging from SEMPER’s illustration. (In addition, SEMPER

did not show a penial retractor but this was probably not intentional).

Source : MNHN, Paris

— 405

These aegistine genera can be compared to Monadenia, the type genus of the subfamily

Monadeniinae which belongs to the family Xanthonychidae (Figs 2C, 4). Monadenia occurs in

northwestern North America, roughly from Alaska to northern California (PlLSBRY 1939). I have

had the opportunity to dissect M. infumata (Gould, 1855) (material from Point Reyes National

Seashore Park, Marin Co., California, 19 April 1989, coll. W. B. Miller) and M. (Shastelix)

troglodytes Hanna & Smith, 1933 (material from Shasta Co., California, 16 August 1969, coll.

B. Roth). Genital morphology in these species did not differ significantly from that in published

illustrations of M. fidelis (Gray, 1834), the type species of the genus (e.g. PlLSBRY 1939).

Monadenia exhibits a mosaic of characters and/or character states shared with Aegista, Tricheulota

and Guamampa :

- penial sheath and closed verge as in Aegista ;

- a single club-shaped mucus gland as in Tricheulota and Guamampa;

- a simple conical flagellum like in Tricheulota.

However each taxon shows characters not shared with others:

Source : MNHN, Paris

— 406 —

- no penial sheath, grooved verge and enlarged distal region of the vas deferens in

Guamampa and Tricheulotcr,

- occurrence of several mucous glands exhibiting a globular shape and alveolar structure

in Aegista ;

- mucous gland internally lamellar (Fig. 4) and clearly divided into gland proper and muscu¬

lar duct in Monadenia.

FIG. 4. — Monadenia fidelis. Middle Sisters Rock. Oregon, Field Museum of Natural History, Chicago, April 27, 1960. A, shell.

B, sexual apparatus (albumen gland omitted). C, distal parts opened.

Guamampa has a single gland with alveolar structure as in Aegista , but this gland opens

into a single duct as in Monadenia , although the duct is very short, slender and hardly muscu-

larized. This duct could represent the first step of a morphocline leading to the condition observed

in Monadenia , which would imply that the heavily muscular duct observed in Monadenia is

derived from the basal region of the gland, irrespective of its initial structure. The unusual and

complex structure of the lower vagina and atrium observed in Monadenia could also easily be

derived from, or lead to, the very thick wall and thick external envelope of the homologous

regions in Guamampa.

By comparison with other helicoid taxa, I propose the following conditions as plesiomor-

phic / apomorphic in the group of taxa discussed above:

la. Occurrence of paired mucus glands with alveolar structure, opening into the accessory

sac in (Aegista)! lb. Reduction of one mucous gland associated with differentiation of the other

Source : MNHN, Paris

— 407 —

into a basal duct and an upper tubular glandular portion ( Tricheulota, Guamampa and still more

in Monadenia );

2a. Accessory sac well developed ( Aegista ) / 2b. Reduction of the accessory sac (all other

genera discussed above);

3a. Penial verge grooved ( Tricheulota and Guamampa ) / 3b. Penial verge closed (tubular)

(Aegista and Monadenia );

4a. Penial sheath present ( Aegista and Monadenia) / 4b. Penial sheath reduced ( Tricheulota

and Guamampa );

5a. Flagellum simple, tapering ( Tricheulota and Monadenia ) / 5b. Complex structure of

flagellum described above ( Aegista and Guamampa)-,

6a. Vas deferens slender, evenly cylindrical (in Monadenia and Aegista) / 6b. Vas deferens

differentiated into two regions (Tricheulota and Guamampa)-,

7a. Walls of the adatrial region with simple structure (Aegista and Tricheulota) / 7b. Walls

of adatrial region thickened and complex in structure (Guamampa and still more in Monadenia).

When submitted to a procedure of parsimony (Hennig86), this data set resolves monophyly

of Tricheulota and Guamampa (penial sheath reduced and distal vas deferens differentiated), but

does not allow resolution of the trichotomy formed by this group, Monadenia and Aegista. How¬

ever, similarities and mosaic pattern of character states shown and discussed above suggest close

relationships of the Bradybaenidae (Guamampa, Tricheulota, Aegista) and Xanthonychidae

(Monadenia). This interpretation is reinforced by the occurrence in some North American

xanthonychid taxa (Xanthonyx, Metostracon, Trichodiscina, Miraverellia) of a pair of mucus

gland similar to those observed in Aegista. However, in these taxa the glands are not alveolar

but exhibit internal anastomosing folds as in Monadenia. In consequence, I propose to include

the Bradybaenidae in the Xanthonychoidea to which Monadenia belongs. As we have come to

the conclusion that Monadenia was derived independently from ancestors other than those of

the rest of american helicoids, its taxonomic rank should be elevated up to family (Monadeniidae).

Further investigation should allow reanalysis of this vast set of taxa, and in particular allow

exploration of the various biogeographical interpretations of the apparent vicariance of the Brady¬

baenidae and Xanthonychidae on each side of the Behring strait.

Aknowledgements

I thank Dr P. Bouchet who collected the material of Guamampa tuba and invited me to work as

visiting professor in MNHN, and Dr S. Tillier for his views on the classification discussed in the present

paper. I am very much indebted to Dr Walter B. Miller and Dr Barry Roth (Santa Barbara Museum of

Natural History) and Dr Clifton C. CoNEYt (Los Angeles County Museum of Natural History) who gave

me a free hand on their material of Monadenia and Aegista.

Source : MNHN, Paris

REFERENCES

MILLER W. b. & Naranjo-Garcia E., 1989. — Familial relationships and biogeography of the Western American

and Caribbean Helicoidea (Gastropoda: Pulmonata). Abstracts of Meeting. Los Angeles, 25-30 June 1989,

40 p.

— 1991. — Familial relationships and biogeography of the Western American and Caribbean Helicoidea

(Gastropoda: Pulmonata). Amer. Mal. Bull. 8 (2): 147-153.

NORDSIECK H„ 1987. — Revision des Systems der Helicoidea (Gastropoda: Stylommatophora). Arch. Moll. 118

(1/3): 9-50.

PlLSBRY H. A., 1890. — Manual of Conchology, vol. 6. Philadelphia, 324 p.

— 1894-1895. — Manual of Conchology, vol. 9. Philadelphia, 366 + 126 p.

— 1939. — Land Motlusca of North America (North of Mexico), vol. 1, part 1. Philadelphia, xvii + 573 p.

Richardson L„ 1985. — Camaenidae: Catalog of species. Tryonia 12: 1-479.

Sarasin P. & Sarasin F., 1899. — Die Land-Mollusken von Celebes. Materialen zur Naturgeschichte der Insel

Celebes , vol. 2. Wiesbaden, 248 p.

SCHILEYKO A. A., 1991. — Taxonomie status, phylogenetic relations and system of the Helicoidea sensu lato

(Pulmonata). Arch. Moll. 120 (4/6): 187-236.

SEMPER C., 1873-1880. — Reisen im Archipel der Philippinen. Wissenschaftliche Resultate. Band 3 (Landmol-

lusken); Heft 2 (1873): 81-128, pis 11-14; Heft 3 (1874): 129-168, pis 9, 15, 17; Heft 5 (1880): 225-264,

pis 1920, 22-23. Wiesbaden.

Source : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle, Paris, 4' série, 18, 1996

Section A, n“ 3-4 : 409-445

A critical review of the trochoidean types in the Muséum

d’Histoire naturelle, Bordeaux (Mollusca, Gastropoda)

by David G. HERBERT

Abstract. — The type material of twenty-two nominal species of New Caledonian trochoidean gastropods,

housed in the Muséum d'Histoire naturelle, Bordeaux, is discussed. The species were described by Fischer.

Lambert, Montrouzier and Souverbie in the Journal de Conchyliologie from 1858-1879. Lectotypes are desig¬

nated where more than one specimen is present and the primary types of all are illustrated. In the majority of

cases the relationships of the taxa to other tropical western Pacific species require further investigation. Few of

the names can be used as the earliest available names with any degree of certainty. New synonyms: Tectaria

montrouzieri Fischer, 1878 = Monodonta angulifera A. Adams, 1853; Trochus gilberti Montrouzier in Fischer,

1878 = Ziziphimts polychromus A. Adams, 1853; Trochus (Euchelus) fossulatulus Souverbie in Souverbie &

Montrouzier, 1875 = Stomatella cancellata Krauss. 1848.

Key-words. — Type material, Mollusca, Trochidae, Bordeaux Museum, New Caledonia.

Révision critique des types de Trochoidea néo-calédoniens du Muséum d'Histoire naturelle,

Bordeaux (Mollusca, Gastropoda)

Résumé. — Le statut de vingt-deux espèces nominales de Trochoidea indo-pacifiques est révisé sur la base

du matériel-type conservé au Muséum d’Histoire naturelle de Bordeaux. Ces taxons, publiés dans le Journal de

Conchyliologie de 1858 à 1879, ont été décrits de Nouvelle-Calédonie par Fischer, Lambert, Montrouzier et

Souverbie. Des lectotypes sont désignés chaque fois que le matériel-type est représenté par plusieurs syntypes,

et tous les échantillons pertinents sont illustrés. Dans la plupart des cas, l'identité de ces taxons ne peut encore

être établie de façon définitive et requiert des comparaisons supplémentaires avec d’autres espèces du Pacifique

occidental. Seuls quelques noms paraissent pouvoir être utilisés avec certitude comme les noms valides d’espèces

néo-calédoniennes ou à plus large répartition indo-pacifique. Synonymes nouveaux : Tectaria montrouzieri Fischer,

1878 = Monodonta angulifera A. Adams, 1853; Trochus gilberti Montrouzier in Fischer, 1878 = Ziziphinus poly¬

chromus A. Adams, 1853; Trochus (Euchelus) fossulatulus Souverbie in Souverbie & Montrouzier, 1875 = Sto¬

mate l/a cancellata Krauss, 1848.

Mots-clés. — Matériel-type. Mollusca, Trochidae, Muséum Bordeaux, Nouvelle-Calédonie.

D.G. Herbert. Natal Museum. Private Bag 9070. Pietermaritzburg. 3200, South Africa.

INTRODUCTION

The Muséum d’Histoire naturelle, Bordeaux (MHNB) [in latin Museum Burdigalense ],

houses an extensive collection of molluscs from the New Caledonian archipelago. The bulk of

this material was collected by French Marist missionaries such as R. P. Xavier Montrouzier

(1820-1897) (Crosse 1898; O'Reilly 1931) and R. P. Pierre Lambert (1823-1903) (H. Fischer

1904). The majority of new taxa were described in the Journal de Conchyliologie by S.-M. Sou-

Source : MNHN, Paris

— 410 —

VERBIE, J.-B. Gassies, P. FISCHER and H. CROSSE, and also by the missionaries themselves.

Souverbie, director of the MHNB, collaborated extensively with MONTROUZIER, particularly in

respect to marine molluscs, and frequently co-authored publications with him.

The material in this collection is historically important since it contains the type material

of many New Caledonian species. In some instances, particularly where the specimens were

numerous, it is evident that additional type material was sent to the Muséum national d'Histoire

naturelle, Paris (MNHN), and it is still present in the typothèque there. Otherwise it seems that

all the original material was retained in Bordeaux. Not infrequently, however, specimens acquired

after the publication of the original descriptions were sent to the MNHN and some of these “ex

auctore" or "ex auteur ” specimens have since been accorded type status (FlSCHER-PlETTE 1950).

Clearly this is not justified.

The MHNB collection seems to have been largely overlooked by subsequent workers, par¬

ticularly those concerned with marine taxa, and the types have rarely if ever been consulted for

verification of identifications. This situation is exacerbated by the museum’s present policy of

not sending type material out on loan. My personal interest in the MHNB New Caledonian types

concerns trochoidean vetigastropods for which, in many cases, the most recent illustrations are

those given by Fischer in Kiener’s “ Spécies général'' (FISCHER 1875-1880) and PlLSBRY in

TRYON’s “Manual of Conchology" (PlLSBRY 1888, 1889, 1890). More recent literature makes

occasional reference to some Souverbie and Montrouzier names, but as the taxa involved

are by and large very poorly known, the identifications are to some extent suspect.

In an attempt to resolve a number of uncertainties regarding Indian Ocean trochids and to

obtain photographs of types for comparative purposes, I visited the MHNB. Whilst there I pho¬

tographed type material of all the trochoidean taxa present, with the intention of publishing the

following review.

Since New Caledonia lies near the centre of a very large zoogeographic province, it is

likely that endemicity in shallow water habitats is low and that most species are widely dis¬

tributed. Although the MHNB collections were amongst the earliest from the New Caledonian

archipelago (a region which is still being studied extensively and which has an astonishingly

rich marine molluscan fauna, BOUCHET in lit.), molluscan material was already available from

neighbouring areas of the central Indo-West Pacific (e.g. in the collection of Hugh Cuming). As

a result, it is likely that a significant proportion of the taxa named by the French authors had

already been studied and described by earlier workers. Establishing whether or not such is the

case, however, is complicated by the paucity of material available and the poor descriptions

provided by relatively prolific authors such as A. Adams, often with neither locality data nor

illustrations. In a number of cases I have been unable come to definite conclusions regarding

the validity of the taxa under consideration and resolution of these uncertainties will require

comparison of topotypic series and perhaps examination of soft parts. At present few of the

names can be used as the earliest available names for distinct taxa with any degree of certainty.

Difficulty has also been experienced in trying to assign the taxa to genera, largely because the

genera themselves have not yet been adequately defined.

Source : MNHN, Paris

— 411 —

Abbreviations

ams

ANSP

BMNH

ICZN

MHNB

MNHN

ZSIC

Australian Museum, Sydney:

Academy of Natural Sciences, Philadelphia:

The Natural History Museum, London:

International Code of Zoological Nomenclature;

Muséum d'Histoire naturelle, Bordeaux;

Muséum national d’Histoire naturelle, Paris;

Zoological Survey of India, Calcutta.

REVIEW

Taxa are listed using their original names and in alphabetical order, by genus, subgenus

and species. No other trochoidean types are present in the MHNB and I know of no further

trochoidean taxa for which type material should be housed there.

Monodonta fischeri Montrouzier in Souverbie & Montrouzier, 1866

(Figs 1-2)

Monodonta fischeri Montrouzier in Souverbie & Montrouzier, 1866: 142, pi. 6, fig. 7;

FISCHER 1878 in 1875-1880: 246, pi. 84, fig. 1. Type loc.: “ins. Art” (New Caledonia).

Euchelus fischeri ; PiLSBRY 1889: 443, pi. 38, figs 18, 19; KAICHER 1990: 5704.

Type material. — Two specimens, labelled “types décrits Joum. de Conch, t. 14, p. 4 [sic]. No. 1 celui

figuré pi. 6, fig. 7. île Art, don. de l’auteur", are present in the MHNB. A further three specimens labelled "syn-

types probables" and donated by Montrouzier are present in the MNHN. The original description, however, men¬

tions only four specimens. In order to resolve this discrepancy I designate the figured specimen (No. 1 in the

MHNB) as lectotype (Figs 1-2) (dimensions: diameter 3.3 mm, length 4.0 mm). It is whitish with scattered maroon-

chestnut spots.

Remarks

This species clearly belongs in the tribe Chilodontini of the trochoidean subfamily Eucy-

clinae. It is close to members of Herpetopoma Pilsbry, 1889, type species Euchelus scabriusculus

Adams & Angas, in Angas, 1867, but has a more strongly protruding columella pillar with a

strong basal tooth and a weaker upper one. In this respect it resembles a number of Indo-West

Pacific taxa such as Trochus gemmatus Gould, 1845, T. instrictus Gould, 1849, Monodonta

exasperata A. Adams, 1853, Euchelus seychellarum G. & H. Nevill, 1869, and Clanculus

crassilabrum Sowerby, 1905, which seem to combine shell characters of several chilodontine

genera, including Euchelus, Herpetopoma and “ Agathodonta ”. Although it will not be possible

to reliably assign these species to genera until the genera themselves have been adequately

studied, at present fischeri seems best referred to Herpetopoma.

Compared with T. instrictus, this taxon is much smaller, has a more obviously beaded sculp¬

ture and more extensive deposition of callus in the parietal and columella regions, such that the

umbilicus is occluded to a greater degree. C. crassilabrum is larger and has more numerous

Source : MNHN, Paris

— 412-

spiral cords with finer granules. M. exasperala and E. seychellarum are more depressed and

have a wider umbilicus.

Jansen (1994) listed fischeri in the synonymy of Trochus gemmalus (under Euchelus s.l.),

but did not provide discussion of the matter. Compared with the present species, Hawaiian (topo-

typic) material of T. gemmalus has a relatively strong cord at the umbilical margin, retains a

wider umbilicus at maturity and has more evenly sized spiral cords. In fact, the lectotype of

T. fischeri resembles more closely the specimens referred by Jansen to Euchelus cf. gemmatus

which are smaller and have a narrower umbilicus than more typical ones. She did not, however,

consider these differences consistent enough to warrant regarding them as a separate species. A

more definite conclusion must await further study.

Figs 1-2. — Monodonta fischeri Montrouzier in Souverbie & Montrouzier, 1866, leclolype (diameter 3.3 mm. length 4.0 mm).

Conclusion

Provisionally referred to Herpetopoma (Eucyclinae, Chilodontini); a possible junior synonym

of H. gemmatus (Gould, 1845).

Rotella montrouzieri Souverbie, 1858

(Figs 3-5)

Rotella montrouzieri Souverbie, 1858: 376; 1860a: 123, pi. 2, fig. 11; FISCHER 1878b: 207;

1879 in 1875-1880: 379. Type loc.: “Insula Art” (New Caledonia).

Ethalia guamensis var. montrouzieri ; Pilsbry 1889: 459, pi. 59, fig. 37.

Ethalia guamensis montrouzieri ; Kaicher 1990: 5700.

Type material. — One specimen, labelled “type décrit et figuré Joum. de Conch, t. 6 [sic], p. 376 et t.

8, p. 123, pl. 2, f. 11. Art, don. de l'auteur”, is present in the MHNB. This is the holotype (Figs 3-5) (dimensions:

diameter 15 mm, length 13 mm). Two further ‘Va- auteur" specimens are in the MNHN, but they have no type

status since the original description stated that there was only one specimen.

Source : MNHN, Paris

-413-

Figs 3-5. — Role lia montrouzieri Souverbie, 1858, holotype (diameter 15 mm, lenglh 13 mm).

Remarks

Fischer (1878b) considered R. montrouzieri to be no more than a colour variety of Trochus

callosus Koch in Philippi, 1844 ( non Gmelin, 1791, nec Wood, 1828) and at the same time

referred the species to a new supraspecific taxon, Liotrochus. Koch’s T. callosus, however, is

itself now regarded as a synonym of Rotella guamensis Quoy & Gaimard, 1834, type species

of Ethalia H. & A. Adams, 1854. Pilsbry (1889) followed Fischer’s proposal and named (1905)

two further varieties, sanguinea and selenomphala , treating these and montrouzieri as subspecies

of guamensis. More recently Hickman & McLean (1990) figured E. guamensis and E. mont¬

rouzieri as separate species and Hickman (in lit.) has indicated that the guamensis complex of

taxa (as per Pilsbry) is a heterogeneous group, including both umboniine and non-umboniine

taxa. Further work is needed to clarify this issue, but the accompanying figures will help, at

least, in defining montrouzieri.

Conclusion

Part of the Ethalia guamensis species complex, but in need of further study.

Source : MNHN, Paris

— 414 —

Stomatella (Gena?) crassa Montrouzier in Souverbie & Montrouzier, 1870

(Figs 6-8)

Stomatella (Gena?) crassa Montrouzier in Souverbie & Montrouzier, 1870: 74, pi. 9, fig. 6.

Type loc.: “ins. Art” (New Caledonia).

Stomatella (Synaptocochlea) crassa; PlLSBRY 1890: 26, pi. 55, figs 22, 23.

FIGS 6-11. — Stomatella (Gena ?) crassa Montrouzier in Souverbie & Montrouzier. 1870, and Stomatella granosa Lambert. 1874.

6-8, S. crassa, lectotype (length 10.1 mm, width 6.6 mm); 9-11, S. granosa, lectotype (length 5.1 mm, width 3.8 mm).

Source : MNHN, Paris

— 415-

Type material. — The original description mentioned two specimens and both remain in the MHNB. The

lot is labelled "type décrit Journ. de Conch, t. 18, p. 40 [sic]. No. 1 celui figuré, pi. 9, f. 6. lie Art, ex auciore’’.

The specimen labelled No. 1 is here refigured and designated lectotype (Figs 6-8) (dimensions: length 10.1 mm,

width 6.6 mm, height 4.4 mm).

Remarks

This specimen has very much the appearance of a Stomatella. It may well prove to be

nothing more than a large, somewhat worn specimen of Stomatella stellata Souverbie in Souverbie

& Montrouzier, 1863, see below. Comparison should also be made with Gena caledonica Crosse,

1871.

Conclusion

One of many nominal Stomatella species (Stomatellinae); its validity requires further study.

Stomatella granosa Lambert, 1874

(Figs 9-11)

Stomatella granosa Lambert, 1874: 374; Souverbie & MONTROUZIER 1875: 35, pi. 4, fig. 2;

Pilsbry 1890: 27. Type loc.: “ins. Lifou” (Loyalty Islands, New Caledonia).

Type material. — The original description stated that there were two specimens, but Souverbie & Mont¬

rouzier (1875) mentioned only a single example and this is the only one now_ present in the MHNB. It is labelled

“type décrit Journ. de Conch, t. 22, p. 374; et t. 23, p. 35, pi. 4, f. 2. île Lifou, ex auctore". There is no

additional type material in the MNHN and the second specimen must be considered lost. I here figure and

designate the remaining one as lectotype (Figs 9-11) (dimensions: length 5.1 mm, width 3.8 mm, height 2.3 mm).

Remarks

This is a characteristic species with coarse spiral cords rendered irregularly granular where

they are crossed by well-developed growth-lines. Its overall facies is that of Synaptocochlea

rather than Stomatella. Shell coloration is similar to that of Synaptocochlea caliginosa (H. &

A. Adams, 1864) (of unknown provenance; holotype BMNH 1968: 138, with operculum. Figs 59-

61 herein), but S. granosa has much coarser spiral sculpture.

Conclusion

Evidently referable to Synaptocochlea (?Eucyclinae: Chilodontini) and probably a valid spe-

Stomatella picta Montrouzier in Souverbie & Montrouzier, 1862

(Figs 12-14)

Stomatella picta (non Stomatia picta d’Orbigny, 1842) Montrouzier in Souverbie & Mon¬

trouzier, 1862: 239, pi. 9, fig. 7. Type loc.: “ins. Art” (New Caledonia).

Source : MNHN, Paris

— 416 —

Stomatella montrouzieri PlLSBRY 1890: 27, pi. 53, figs 74-75; nom. nov.

Type material. — One specimen, labelled "type décrit et figuré Joum. de Conch, t. 18, p. 139, pi. 8, f. 7,

île Art, don. de l'auteur", is present in MHNB (dimensions: length 4.7 mm, width 3.0 mm, height 2.2 mm).

This specimen fits the dimensions given and matches the figure well, and, since the description stated that there

was only one specimen, I consider this to be the holotype (Figs 12-14). There are additional “ex auctore" specimens

in the MNHN, four in the typothèque and two (including operculum) in the Journal de Conchyliologie collection

(cited by Fischer-Piette 1950), but these have no type status.

Figs 12-17. — Siomalella picta Montrouzier in Souverbie & Montrouzier, 1862, and Stomatella stellata Souverbie in Souverbie

& Montrouzier, 1863. 12-14, S. picta, holotype (length 4.7 mm, width 3.0 mm): 15-17, S. stellata, holotype (length 7.5 mm,

width 4.8 mm).

Source : MNHN, Paris

-417-

Remarks

Pilsbry (1890) considered Stomatia picta d’Orbigny, 1842, from the tropical western Atlan¬

tic and Stomatella picta Montrouzier in Souverbie & Montrouzier, 1862, to be congeneric and

thus secondarily homonymous, and proposed the replacement name Stomatella montrouzieri for

the junior name. Regardless of whether or not this secondary homonymy is real (1 believe it

is), Montrouzier’ s picta is permanently invalid (ICZN Art. 59b) and the replacement name is

the valid name for the taxon. At the same time, PlLSBRY proposed a new supraspecific taxon,

Synaptocochlea, for small, spirally Urate species otherwise similar to Stomatella, and cited

S. montrouzieri as the type species. The MHNB specimen is thus the holotype of the type species

of Synaptocochlea Pilsbry, 1890.

Most authors have accepted Synaptocochlea as a generically distinct taxon and have referred

it, together with Stomatella, to the Stomatellinae/Stomatellidae (cf KEEN 1960). Synaptocochlea

species are smaller than those of Stomatella, lack an obvious interior nacreous layer, have fewer

whorls, a coarser spiral sculpture that is rendered somewhat granose by growth-lines and retain

an operculum. More recently, Hickman & McLean (1990) have indicated that the genus is not

in fact stomatelline, and should be referred to the Eucyclinae (tribe Chilodontini).

The relationships of S. montrouzieri to other species of Synaptocochlea, particularly the

widespread Indo-West Pacific S. concinna (Gould, 1845), need to be investigated. At first glance

S. montrouzieri appears to have a finer, less obviously beaded sculpture and (the type at least)

has a bold colour pattern that lacks the red spiral lines/flecks so common in S. concinna. However,

the range of variation shown in S. concinna is extensive, particularly in respect of coloration,

and S. montrouzieri may well prove to fall within this when studied in detail. S. picta d’Orbigny,

1842, also merits comparison with S. concinna as it too is of very similar appearance. Abbott

( 1958) has suggested that concinna should be treated as no more than a subspecies of picta.

Conclusion

Type species of Synaptocochlea Pilsbry, 1890 (?Eucyclinae, Chilodontini), and probably a

synonym of S. concinna (Gould, 1845).

Stomatella stellata Souverbie in Souverbie & Montrouzier, 1863

(Figs 15-17)

Stomatella stellata Souverbie in Souverbie & Montrouzier, 1863a: 169, pi. 5, fig. 10;

Pilsbry 1890: 25, pi. 53, figs 76, 77, pi. 2, figs 35-37 (var. ornatissima); Hedley 1901: 128;

1909: 353. Type loc.: “ins. Art” (New Caledonia).

Gena stellata; Melvill & Standen 1895 in 1895-1897: 126.

Synaptocochlea stellata ; CERNOHORSKY 1978: 37, text-fig. 4; Wilson 1993: 69.

Type material. — One specimen, labelled “type décrit et figuré en Journ. de Conch, t. 11, p. 6, pl. 5, fig.

l[ii'c]. île Art, don. de l'auteur'’, is present in the MHNB (dimensions: length 7.5 mm, width 4.8 mm, height

3.2 mm). A further two specimens, “ex auctore ”, are present in the MNHN. Since the original description stated

that only a single specimen had been seen and the MHNB specimen (Figs 15-17) matches both the figure and

dimensions given reasonably well, I regard it as the holotype and consider the MNHN specimens to have no

type status.

Source : MNHN, Paris

-418-

Remarks

Cernohorsky (1978) referred this species to Synaptocochlea , but it is difficult to be certain

whether this was justified. There are features of the shell, in particular its finer sculpture, which

suggest that the original referral to Stomatella may in fact be correct. The spiral lirae, although

crossed by growth-lines, are not rendered granose by them. There is, in addition, some trace of

nacre on the interior. A more conclusive appraisal must await examination of the soft parts of

topotypic material.

Pilsbry (1890) placed Stomatella ornata Brazier, 1877, described from north-eastern

Australia, in synonymy with this species. However, operculae glued inside the apertures of the

four syntypes of S. ornata in the AMS (one here figured, Figs 62-64), ally that taxon with

Synaptocochlea. See also remarks above regarding Stomatella (Gena ?) crassa Montrouzier, 1870.

Conclusion

Probably a species of Stomatella (Stomatellinae); its validity requires further study.

Tectaria montrouzieri Fischer, 1878

(Figs 18-19)

Tectaria montrouzieri Fischer, 1878c: 212; Souverbie & Montrouzier 1879: 31, pi. 3,

fig. 6. Type loc.: “île Art” (New Caledonia).

Turcica montrouzieri ; HEDLEY 1909: 354.

Type material. — Fischer indicated the original material to be in the “Mus. Burdigalense”, but did not

specify the number of specimens. Souverbie & Montrouzier (1879) later stated that there was only one. This

remains in the MHNB and must be regarded as holotype (Figs 18-19): its dimensions correspond with those

given by Fischer in the original description (diameter 7.4 mm, length 10.2 mm). It is labelled “type décrit Journ.

de Conch, t. 26. p. 212 et t. 27. p. 31, pi. 3, f. 6. Souverbie - lie Art, ex auctore". A further specimen “ex

auteur” is present in the MNHN, but it has no type status.

Remarks

This species was not mentioned by Rosewater (1972) in his revision of the Indo-Pacific

Tectariinae, even as a dubious or excluded taxon. Its original placement in Tectaria[us] was

erroneous and it should instead be referred to the Trochidae (cf. HEDLEY 1909). The holotype

is very similar to the type material of Monodonta angulifera A. Adams, 1853, from the Philippines

(three syntypes, BMNH 1968215) and is clearly conspecific therewith (Herbert in prep.). M. an¬

gulifera is the type species of the chilodontine genus Perrinia H. & A. Adams, 1854 (s.d. Pilsbry

1889).

Conclusion

A junior synonym of Monodonta angulifera A. Adams, 1853, type species of Perrinia H. &

A. Adams, 1854 (Eucyclinae, Chilodontini).

Source : MNHN, Paris

— 419 —

Ta montrouzieri Fischer, 1878, and Trochus artensis Fischer. 1878. 18-19, Teclaria montrouzieri,

l, length 10.2 mm); 20-21. Trochus artensis, holotype (diameter 7.9 mm, length 10.2 mm).

holotype

Trochus artensis Fischer, 1878

(Figs 20-21)

Trochus artensis Fischer, 1878b: 208. Type loc.: ‘Tile Art” (New Caledonia).

Cantharidus artensis ; PlLSBRY 1889: 129.

Type material. — The single “Musée de Bordeaux" specimen cited in the original description remains in

the MHNB (Figs 20-21). It must be considered the holotype (dimensions: diameter 7.9 mm, length 10.2 mm). It

is labelled “type décrit Joum. de Conch, t. 26, p. 208, non figuré. île Art, ex auctore ”,

Remarks

There is no previous illustration of this taxon. The holotype is a badly worn specimen

belonging within the Thalotia-Calthalotia-Prothalotia complex. It has a weak bulge at the base

Source : MNHN, Paris

— 420 —

of the columella and there is a fine granulation on the spiral cords. It is patterned with opistho-

cline, maroon, axial flames on a near white ground. PlLSBRY (1889) merely translated Fischer’s

original diagnosis and remarks, adding nothing to our knowledge of the taxon. HEDLEY (1908)

suggested that it might be a synonym of Thalotia crenellifera A. Adams, 1853, from northern

Australia; however, his figure of the BMNH type of that species indicates it [crenellifera] to

be more elevated and to have a stronger columella tooth (see remarks under Trochus gilberti).

Other similar taxa include Trochus (Ziziphinus) arruensis Watson, 1880, from the Arrou

[Aru] Islands (Arafura Sea), Thalotia marginata Tenison-Woods, 1880, and Trochus (Thalotia)

torresi Smith, 1884, both from northern Australia. These appear to differ somewhat from artensis,

judging from the types and original descriptions, but without good series of topotypic material

by which to assess intraspecific variability it is impossible to meaningfully evaluate these diffe¬

rences. Before the true identity and relationships of this species can be established, fresh material

will need to be studied and compared with a range of cantharidine species from the central

Indo-West Pacific.

Conclusion

Fresh, topotypic material requires comparison with other tropical western Pacific can¬

tharidine taxa (Trochinae, Cantharidini).

Trochus constellatus Souverbie in Souverbie & Montrouzier, 1863

(Figs 22-25)

Trochus constellatus Souverbie in Souverbie & Montrouzier, 1863b: 279, pi. 12, fig. 3;

Fischer 1878 in 1875-1880: 271, pi. 90, fig. 1. Type loc.: “Balade et ins. Art” (New Caledonia).

Monodonta constellata ; PlLSBRY 1889: 108, pi. 35, figs 9, 10, pi. 62, figs 69-71.

Type material. — The original description cited “Mus. Burdigalense” and indicated that there were eleven

specimens, two of which remain in the MHNB, labelled “types décrits Journ. de Conch, t. 11, p. 279, No. 1

celui figuré, pi. 12, fig. 3, No. 2 sujet de la note, don de l'auteur. île Art et Balade”. An additional twelve

specimens labelled as types are present in the MNHN (five cited by Fischer-Piette 1950), but since this exceeds

the number cited originally, some must be specimens sent to Paris subsequent to the publication of the description.

Unfortunately, the status of all these MNHN specimens as types is thus compromised. The MHNB specimen

labelled No. 1 is here refigured (Figs 22-25) and designated lectotype (dimensions: diameter 7.9 mm, length

Remarks

This species clearly belongs to the gibbuline genus Diloma Philippi, 1845, and was in fact

referred there by FISCHER (1879 in 1875-1880). Its small size, globose-conical profile, smooth

columella and concave base suggest a relationship with the subgenus Cavodiloma Finlay, 1926,

type species Diloma coracina (Philippi, 1851) from New Zealand. The latter has a well-developed

ridge on the base extending from the columella-basal lip junction, around the margin of the

basal concavity; this is scarcely evident in T. constellatus.

Source : MNHN, Paris

— 421 —

Conclusion

A species of Diloma Philippi, 1845 (Trochinae, Gibbulini), probably belonging to subgenus

Cavodiloma Finlay, 1926. The question of validity must await comparison with other members

of the genus.

Figs 22-25. — Trochus constellants Souverbie, in Souverbie & Montrouzier, 1863, lectotype (diameter 7.9 mm, length. 7.1 mm).

Trochus gilberti Montrouzier in Fischer, 1878

(Figs 26-27)

Trochus gilherti Montrouzier in Fischer, 1878b: 207. Type loc.: “File Art” (New Caledonia).

Trochus giliberti (unjustified emendation) Souverbie & Montrouzier, 1879: 33, pi. 3, fig. 7;

Fischer 1879 in 1875-1880: 401, pi. 119, fig. 2.

Cantharidus giliberti; PlLSBRY 1889: 128, pi. 45, figs 37, 38; SCHEPMAN 1908: 41, pi. 9,

fig. 4 (radula); Cernohorsky 1978: 34, pi. 8, fig. 10.

Cantharidus (Cantharidus) gilberti ; Adam & Leloup 1938: 19, pi. 2, fig. 7.

Cantharidus (Jujubinus) gilberti; SPRINGSTEEN & LEOBRERA 1986: 34, pi. 5, fig. 4.

Jujubinus gilberti; Wilson 1993: 80.

Type material. — Two specimens, labelled “types décrits Joum. de Conch. (Fischer) t. 26, p. 207 et t. 27

(Montrouzier) t. 27 [sic], p. 33, f. 7. lie Art, ex auctore", are present in the MHNB (Figs 26-27). No indication

of the number of specimens originally available was given in Fischer (1878b), but Souverbie & Montrouzier

Source : MNHN, Paris

— 422 —

(1879) indicated that there were only two and figured both. The larger specimen, which is in better condition

(Fig. 26). is designated lectotype (length 12.8 mm). A further six specimens, “ex auteur ", are present in the

MNHN, but they have no type status.

Flos 26-27. — Trochus gilberti Montrouzicr in

Fischer. 1878; 26, lectolype (lenglh 12.8 mm);

27. paraleclotype (length 11.7 mm).

Remarks

Souverbie & MONTROUZIER (1879) slated that the species was named in honour of the

Reverend Father Gilibert and that the original spelling given (Fischer 1878b) was incorrect.

However, there was nothing in the original publication to indicate that such patronymy was

intended and therefore, in terms of ICZN Art 32c(ii), it cannot be classed as an incorrect original

spelling and the name must thus be used in its original form. Trochus giliberti, following ICZN

Art 33b(iii), should be regarded as an unjustified emendation with its own author and date (i.e.

Souverbie & Montrouzier 1879), and is a junior objective synonym of T. gilberti. This si¬

tuation is unfortunate, but compliance with ICZN (1985) offers no defensible alternative.

T. gilberti is synonymous with Ziziphinus picturatus and Z. polychromus, both described

from the Philippines by A. Adams (1853). The synonymy of these two simultaneously published

taxa was noted by Ponder (1978) who afforded Ziziphinus polychromus precedence. The syntypes

of both are in the BMNH (Z. polychromus BMNH 1968 111, four specimens; Z. picturatus BMNH

196844, two specimens), those of Z. polychromus are in better condition; lectotypes of both are

here designated and figured (Figs 65, 66). Cantharidus (Jujubinus) tristis Thiele, 1930, described

from north-western Australia, is a further synonym (Ponder 1978) and so too may be the northern

Australian Thalotia crenellifera A. Adams, 1853 (c/. Wilson 1993).

The shell is elevated conical, with a strong peripheral spiral cord (usually shallowly bifid)

and fine incised spiral striae on the adapical surface; the base is rather more coarsely lirate, the

umbilicus narrow or closed and there is a denticle of rather variable prominence at the base of

the columella. The ground colour is usually red or green and is very variably patterned with

Source : MNHN, Paris

— 423 —

white axial flames, zig-zags and/or flecks. Comparison of Figs 26-27 with Figs 65-66 shows the

spiral striation of the gilberti types to be more close-set than that of polychromus or picturatus,

but this is evidently a variable character within the species.

Trochus fournieri Crosse, 1863, also from New Caledonia, is similar but, judging from the

holotype (BMNH 1896.12.1.9), is smaller in relation to the number of whorls, has a more evenly

rounded periphery, lacks a basal columella denticle and has a bright green internal iridescence

(Fig. 67). Komaitrochus pulcher Kuroda & Taki, 1958, from southern Japan, is broader, lacks

an enlarged peripheral spiral cord and has much finer, almost obsolete spiral sculpture above

the periphery.

The generic affinities of this species are problematic. It clearly belongs within the

Cantharidus-Jujubinus-Thalotia-Komaitrochus complex, but supraspecific taxa within this group

are not yet sufficiently well defined as to permit a definite statement. The species was recorded

from a number of localities in Indonesia by SCHEPMAN (1908) and it will probably prove to be

distributed throughout the central western Pacific.

Conclusion

A junior synonym of Ziziphinus polychromus A. Adams, 1853 (Trochinae, Cantharidini);

generic affinity uncertain.

Trochus reevei Montrouzier in Souverbie & Montrouzier, 1866

(Figs 28-31)

Trochus reevei Montrouzier in Souverbie & Montrouzier, 1866: 141, pi. 6, fig. 8; Fischer

1875: 49: 1878 in 1875-1880: 327, pi. 102, fig. 2. Type loc.: “ins. Art” (New Caledonia).

Gibbula reevei , PlLSBRY 1889: 229, pi. 32, figs 59, 60.

Type material. — The original description stated that there were eight examples; two of these remain in

the MHNB labelled “types décrits Journ. de Conch, t. 14, p. 141, No. 1 celui figuré pi. 6, f. 8. île Art, don. de

l'auteur". The other six were sent to Paris by Montrouzier and are in the MNHN. All may be considered syntypes.

The two MHNB specimens have become detached from the original board and it is now impossible to establish

which was No. 1. None the less, they are very similar and both in good condition, obviously live taken. The

one with the more well-developed apertural dentition is here illustrated and designated lectotype (Figs 28-31)

(dimensions: diameter 7.5 mm, length 6.0 mm).

Remarks

This species superficially resembles members of the genus Clanculus Montfort, 1810, but

lacks both a columella disjunction and strong columella teeth. In sculpture and apertural dentition

it resembles Clanculus danieli Crosse, 1862, the type species of Eurytrochus Fischer, 1879, and

was in fact listed thereunder by Fischer (1879 in 1875-1880: 417). Eurytrochus comprises a

small, but seemingly well defined group of trochids from the central Indo-West Pacific (Japan

to New South Wales and India to Samoa) and is probably worthy of recognition at generic level.

Trochus (Clanculus) bathyraphe Smith, 1876, described from the nearby Solomon Islands,

is extremely similar to T. reevei, even to the extent of having a turquoise-green tinted apex.

Source : MNHN, Paris

— 424 —

It differs, however, in having much more obviously beaded spiral cords (two syntypes BMNH

76.1.10.62, Figs 68-69). More material is needed in order to establish whether these forms differ

consistently in this respect or whether they simply represent opposite ends of a graded series

including specimens with intermediate sculpture.

Conclusion

A member of the genus Eurytrochus Fischer, 1879 (Trochinae,?Gibbulini), and almost cer¬

tainly a valid name; perhaps an earlier name for Trochus bathyraphe Smith, 1876.

Trochus scrobiculatus Souverbie in Souverbie & Montrouzier, 1866

(Figs 32-33)

Trochus scrobiculatus Souverbie in Souverbie & Montrouzier, 1866: 140, pi. 6, fig. 9;

Fischer 1878 in 1875-1880: 248, pi. 84, fig. 2. Type loc.: “ins. Art” (New Caledonia).

Euchelus scrobiculatus; PlLSBRY 1889: 437, pi. 38, figs 2, 3; Hidalgo 1904-1905: 256;

Dautzenberg & BOUGE 1933: 406; Franc 1956: 23; Mastaller 1979: 31.

Source : MNHN, Paris

— 425 —

Type material. — Two examples (Mus. Burdigalense) were mentioned in the original description and two

remain in the MHNB, labelled “types décrits Journ. de Conch, t. 14. p. 140, No. 1 celui figuré pi. 6, f. 9. île

Art, ex auctore". The one identified as the figured specimen is here refigured and designated lectotype (Figs 32-33)

(dimensions: diameter 5.4 mm, length 6.5 mm). No type material is present in the MNHN.

Remarks

This name is one of a number given to a group of small, white, cancellate, chilodontine

species from a range of localities in the Indo-West Pacific. The group appears to be divisible

into at least two subgroups, depending upon whether or not there are ridge-like denticles inside

the outer lip when mature, but at present it is not clear whether each of the described taxa

represent good species or whether a smaller number of somewhat variable species is involved.

T. scrobiculatus belongs with those in which such denticles are present, and is thus allied to

Monodonta foveolata A. Adams, 1853, from Lord Hood’s Island (Marutéa Atoll, Tuamotu

Archipelago), and Turbo semilugubris Deshayes, 1863, from La Réunion. Species which do not

seem to develop these apertural denticles include Monodonta clathrata A. Adams, 1853, from

the Philippines, and Euchelus cavernosus Sowerby, 1905, from Sri Lanka. Euchelus favosus

Melvill & Standen, 1896, from the Loyalty Islands, also probably belongs in this group, but the

figured syntype (Manchester Museum) is too juvenile to assess.

Turbo semilugubris differs from Trochus scrobiculatus in being smaller, in possessing an

umbilicus and in having a bold colour pattern (MNHN syntype. Figs 70-71). Monodonta foveolata

is more similar in size, but, judging from the lectotype (Figs 72-73, BMNH 1968071/1, designated

Marshall 1979), has an open (albeit narrow) umbilicus, is more depressed and has less deeply

cancellate sculpture. The significance of the apparent differences between T. scrobiculatus and

M. foveolata, however, needs to be assessed in terms of intraspecific variation.

An element of confusion has surrounded the generic affinity of these taxa. The overall shell

facies is similar to that of Vaceuchelus Iredale, 1929, the type species of which is Euchelus

FIGS 32-33. — Trochus scrobiculatus Souverbie in Souverbie & Montrouzier. 1866, specimen identified as figured specimen, here

designated lectolype (diameter 5.4 mm, length 6.5 mm).

Source : MNHN, Paris

426 —

angulatus Pease, 1868, from “ins. Annaa” (= Anaa Atoll, Tuamotu Archipelago). PlLSBRY (1889),

in contrast to IREDALE (1929), believed E. angulatus to be no more than a variety of M. Fove-

olatus and Melvill & Standen (1901) followed suit. The two were regarded as distinct species

by MARSHALL (1979) on the grounds that E. angulatus lacked denticles inside the aperture. The

figure of the lectotype recently provided by JOHNSON (1994, pi. 7, fig. 13) is too small to permit

this observation to be confirmed, but personal examination of the lectotype (ANSP 40671), in

fact, reveals such denticles to be present inside the outer lip (Figs 74, 75). There can be little

doubt therefore, that T. scrobiculatus is referable to Vaceuchelus. Members of this genus differ

from those of Herpetopoma Pilsbry, 1889, in lacking a deep notch between the denticles at the

junction of the basal and columellar lips, and in having a generally coarser sculpture.

The lectotype of Euchelus angulatus has a relatively narrow supra-peripheral spiral cord

which lies closer to the peripheral cord than does the sub-peripheral one, and has a steeply

sloping shoulder (Figs 74, 75); in this respect it differs from the types of T. scrobiculatus and

M. foveolata. However, until such time as a good topotypic series can be compared, the question

of the validity of these nominal taxa remains unresolved.

Conclusion

A species of Vaceuchelus Iredale, 1929 (Eucyclinae, Chilodontini), but specific validity

requires further study.

Trochus (Euchelus) fossulatulus Souverbie in Souverbie & Montrouzier, 1875

(Figs 34-35)

Trochus (Euchelus) fossulatulus Souverbie in Souverbie & Montrouzier, 1875: 39, pi. 4,

fig. 5; G. & H. Nevill 1875: 103; FISCHER 1876: 151; 1879 in 1875-1880: 391, pi. 117, fig. 2.

Type loc.: “ins. Art” (New Caledonia).

Euchelus fossulatus [sic]; PlLSBRY 1889: 444, pi. 38, figs 15, 16; Hedley 1915: 710.

Type material. — The two specimens mentioned in the original description remain in the MHNB. They

are labelled “types décrits Journ. de Conch, t. 23, p. 39, No. 1 celui figuré, pi. 4, f. 5. île Art. ex auctore".

Specimen No. 1 is here refigured (Figs 34-35) and designated lectotype (dimensions: diameter 9.4 mm, length

8.3 mm). There is no material in the MNHN.

Remarks

Pilsbry (1889) correctly referred this species to his new taxon Hybochelus, noting that it

differed from the type species, Hybochelus cancellatus (Krauss, 1848), in having a slightly more

prominent spire. I can find nothing to indicate that more than one species is involved and thus

regard T. fossulatulus as a junior synonym of H. cancellatus , as suggested by Hedley (1915).

PlLSBRY’s description of his H. cancellatus orientalis (Pilsbry, 1904), from Japan, fits the types

of fossulatulus extremely well, but I think it unlikely that such subspecific distinction is justified.

The taxon is evidently distributed over a wide section of the central Indo-West Pacific, from

Japan to the Philippines, Andaman Islands and New Caledonia, and probably further. It differs

Source : MNHN, Paris

— 427 —

from H. mysticus (Pilsbry, 1889) and H. sagamiensis Kuroda & Habe, 1971, in having a relatively

wide umbilicus.

The South African locality (Table Bay) given in Krauss’s original description of H. can-

cellatus is erroneous; no tropical Indo-West Pacific trochids are known to occur there. In fact,

there have been no subsequent reports of the species in southern or eastern Africa as a whole.

FIGS 34-35. — Trochus (Euchelus) fossulatulus Souverbie in Souverbie & Montrouzier, 1875. specimen identified as figured

specimen, here designated lectotype (diameter 9.4 mm, length 8.3 mm).

Conclusion

A synonym of Siomatella cancellata Krauss, 1848, type species of Hybochelus Pilsbry, 1889

(Eucyclinae, Chilodontini).

Trochus (Euchelus) lamberti Souverbie in Souverbie & Montrouzier, 1875

(Figs 36-37)

Trochus (Euchelus) lamberti Souverbie in Souverbie & Montrouzier, 1875: 37, pi. 4, fig. 4;

G. & H. NEVlLL 1875: 102; Fischer 1876: 151; 1878b: 210; 1879 in 1875-1880: 385, pi. 116,

fig. 2. Type loc.: “Insula Nou” (New Caledonia).

Euchelus lamberti ; Hedley 1907: 479; 1915: 710.

Type material. — The single specimen mentioned in the original description remains in the MHNB (Figs

36-37); it must be considered the holotype (dimensions: diameter 9.0 mm, length 8.7 mm). It is labelled “type

décrit Journ. de Conch, t. 13, p. 37, pi. 4, f. 4, et figuré dans Kiener [pi. 116, fig. 2]. lie Nou, ex auctore".

There is no material in the MNHN.

Remarks

G. & H. Nevill (1875) placed this name in the synonymy of Tallorbis roseola G. &

H. Nevill, 1869, described from Ceylon, a proposal that was accepted by Fischer (1879 in

1875-1880) and Hedley (1915). There are two syntypes of T. roseola in the ZSIC (M2258/1,

R. N. Kilbum, pers. comm.), the larger of which is here illustrated (Fig. 76) and designated

Source : MNHN, Paris

— 428 —

lectotype. This specimen has a bolder colour pattern and more pronounced cancellation than the

holotype of T. lamberti, but is in other respects very similar. Although it is perhaps unwise to

assess synonymy with so little material available, 1 believe the apparent differences result pri¬

marily from the fresher condition of the T. roseola lectotype, and concur with earlier authors

in considering there to be only one species involved.

The generic affinity of this species is unclear. The Nevill brothers assigned the taxon to

their new genus Tallorbis G. & H. Nevill, 1869, but this has subsequently been synonymised

with Euchelus Philippi, 1847 (e.g. Keen 1960). Certainly the taxon appears referable to the

Chilodontini, but its relationships within this group require further study. Hedley (1915) regarded

Tallorbis as an earlier name for Hybochelus Pilsbry, 1889.

FIGS 36-37. — Trochus (Euchelus) lamberti Souverbie in Souverbie & Montrouzier, 1875, holotype (diameter 9.0 mm, length

8.7 mm).

Conclusion

A synonym of Tallorbis roseola G. & H. Nevill, 1869 (Eucyclinae, Chilodontini), but generic

affinity requires further study.

Trochus (Monilea) lifuanus Fischer, 1878

(Figs 38-40)

Trochus (Monilea) lifuanus Fischer, 1878a: 63; Cernohorsky 1978: 36. Type loc.: “ins.

Lifu” [Lifou](Loyalty Islands, New Caledonia).

Monilea lifuana; SOUVERBIE & MONTROUZIER 1879: 30, pi. 3, Fig. 5; FISCHER 1879 in

1875-1880: 388, pi. 116, fig. 4; Pilsbry 1889: 252, pi. 41, figs 6, 7, pi. 59, figs 64, 65; HEDLEY

1899: 405, 1909: 353.

Trochus (Monilea) lifuana ; SMITH 1884: 73, pi. 6, figs B, Bl.

Minolia lifouana [sic]; Melvill & Standen 1895 in 1895-1897: 125.

Monilea (Monilea) lifuana ; Ladd 1966: 40, pi. 5, figs 13, 14.

Source : MNHN, Paris

— 429 —

Type material. — A single specimen labelled "type décrit Journ. de Conch, t. 26, p. 63, et Souverbie t. 27.

Monilea lifuana p. 30, pi. 3, f. 5. île Lifou, ex auctore" is present in the MHNB (Figs 38-40). It may be regarded

as the holotype (dimensions: diameter 13.6 mm, length 10.7 mm). There is no material in the MNHN.

Remarks

Iredale (1929) rightly compared his new species Talopena gloriola, from New South Wales,

with Trochus lifuanus , and at the same time proposed that lifuanus be referred to Talopena.

There can be little doubt that lifuanus and gloriola are congeneric since their conchological

similarities are considerable. Whether in fact they are referable to Talopena Iredale, 1918, how¬

ever, is less clear, since its type species. Monilea incerta Iredale. 1912, has strong spiral sculpture

and is clearly nothing more than a small species of Monilea s. str. (c/. MARSHALL 1979). Greater

similarity is shown with a cluster of species conchologically intermediate between Ethalia

H. & A. Adams, 1854, and Ethminolia Iredale, 1924 (Herbert 1992, figs 131-136). The present

species differs from Ethminolia s. str. in being larger and in possessing a distinct umbilical

funicle, and from Ethalia s. str. in that the shell is thin and the terminal funicular callus largely

separate from the parietal region. It is impossible to reliably assign these intermediate taxa to

genus using conchological characters alone. Resolution of the dilemma must await studies of

the radula and anatomy.

FIGS 38-40. — Trochus ( Monilea) lifuanus Fischer, 1878, holotype (diameter 13.6 mm, length 10.7 mm).

Source : MNHN, Paris

— 430 —

Whether or not T. lifuanus and T. gloriola represent distinct species also requires further

study. Comparison of the type material (cf Herbert 1992, fig. 131) indicates gloriola to be

somewhat taller and to have a slightly less well-developed callus (that is brownish rather than

green) at the end of the umbilical funicle, but these differences are not great and might easily

be encompassed within the variability of a single species. CERNOHORSKY (1978) and Wilson

( 1993) placed T. lifuanus in synonymy with Monilea vernicosa Gould, 1861, described from the

Ryukyu Islands, but I consider this unlikely. The holotype of M. vernicosa (JOHNSON 1964,

pi. 20, fig. 2; HERBERT 1992, fig. 136), though sculpturally indistinguishable from T. lifuanus,

is slightly more depressed and has a much weaker umbilical funicle. It could be suggested that,

with a diameter of only 5.3 mm, the vernicosa holotype is juvenile and thus could be expected

to have a more weakly developed funicle, but specimens of T. lifuanus of a similar size already

have a much more strongly developed funicle and, furthermore, have an altogether narrower

umbilicus.

Conclusion

Probably a valid species and perhaps an earlier name for Talopena gloriola Iredale, 1929;

somewhat intermediate between Ethminolia and Ethalia in shell characters (Umboniinae).

Trochus (Monilea) rhodomphalus Souverbie in Souverbie & Montrouzier, 1875

(Figs 41-43)

Trochus (Monilea) rhodomphalus Souverbie in Souverbie & Montrouzier, 1875: 36, pi. 4,

fig. 3. Type loc.: “ins. Lifou” (Loyalty Is).

Trochus rhodomphalus; FISCHER 1878b: 210; 1879 in 1875-1880: 392, pi. 117, fig. 3.

Monilea rhodomphala', PlLSBRY 1889: 262, pi. 41, figs 22-24.

Minolia rhodomphala', MELVILL & STANDEN 1897 in 1895-1897: 414.

Type material. — The original description cites “Mus. Burdigalense" and mentions three specimens. Only

two, bearing the label “types décrits Journ. de Conch, t. 23, p. 36, No. 1 celui figuré pi. 4, f. 3, No. 2 celui

figuré dans Kiener et Fischer [= Fischer 1879 in 1875-1880, pi. 117, fig. 3]. île Lifou, ex auctore", are now

present in the MHNB. The first of these is here refigured (Figs 41-43) and designated lectotype (dimensions:

diameter 7.6 mm, length 5.0 mm). There is no material in the MNHN.

Remarks

Similar to a number of taxa described from the central Indo-West Pacific. Ethalia rhodom¬

phala Smith, 1903, from the Maidive and Laccadive archipelagos (three syntypes BMNH

1903.9.17.57-59, one here figured and designated lectotype. Figs 77-79), differs in having non-

shouldered whorls and thus a more flat-sided spire; it also has a well-developed, linguiform

callus deposit at the junction of the columella and parietal region, strong plicae at the umbilical

margin and is more glossy. Ethalia floccata Sowerby, 1903, from Japan (holotype BMNH

1903.12.7.15), is perhaps the most similar species, but besides differences in coloration, the

holotype of that taxon has more or less obsolete spiral sculpture and has a wider umbilicus with

a less strongly thickened margin (Figs 80-82). Isanda pulchella A. Adams, 1855, from Mindoro,

Source : MNHN, Paris

— 431 —

Philippines (holotype BMNH 1968350) and Ethalia capillata Gould, 1862, from China, are doser

to E. rhodomphala Smith, 1903, but /. pulchella has a more funnel-shaped umbilicus with weaker

marginal plicae (Figs 83-85), and E. capillata a larger linguiform callus that almost totally oc¬

cludes the umbilicus (lectotype figured by JOHNSON 1964, pi. 5, fig. 14).

Pilsbry (1905) united these taxa (save for £. capillata ) in his new genus Ethaliella, citing

£. floccata as the type species. If conchological criteria alone are considered, there is perhaps

justification for this, but additional data from the external anatomy and radula may prove other¬

wise. Whether these five nominal taxa represent distinct species is also open to question.

T. rhodomphalus and £. floccata could easily represent variations of a single species, perhaps

belonging in Ethminolia Iredale, 1924 or Ethalia H. & A. Adams, 1854. Similarly, £. rhodom¬

phala, E. capillata and /. pulchella may be another single species. However, in the absence of

adequate series of specimens and data on external anatomy and radula, it is worthless to speculate

further on this. Three additional taxa, Minolia ceraunia, M. edithae and M. malcolmia, described

by Melvill (1891) from the Philippines also merit comparison with this group of species.

Fischer (1878b) indicated that he believed Trochus rotellaeformis Philippi, 1849, of un¬

known provenance, to be a synonym of T. rhodomphalus. Philippi stated that the original material

was in the collection of Silvanus Hanley, but it could not be traced at the Leeds Museum (NORRIS

in lit.) and thus I cannot confirm the synonymy.

Conclusion

A member of the Ethaliella group of species (Umboniinae) and probably an earlier name

for Ethalia floccata Sowerby, 1903.

Figs 41-43. — Trochus (Monilea) rhodomphalus Souverbie in Souverbie & Montrouzier, 1875. specimen figured wilh original

description and here designaled leclotype (diameter 7.6 mm, length 5.0 mm).

Source : MNHN, Paris

— 432 —

Trochus (Polydonta) calcaratus Souverbie in Souverbie & Montrouzier, 1875

(Figs 44-46)

Trochus (Polydonta) calcaratus Souverbie in Souverbie & Montrouzier, 1875: 41, pi. 4,

figs 7, 7a. Type loc.: “ins. Art” (New Caledonia).

Trochus calcaratus ; Fischer 1875: 48; 1879 in 1875-1880: 347, pi. 109, fig. 2; Pilsbry

1889: 30, pi. 2, fig. 15, pi. 8, figs 83, 84; HIDALGO 1904-1905: 246; Hedley 1909: 353; Kaicher

1979: 2174.

Infundibulum (Lamprostoma) calcaratum ; Dautzenberg & Bouge 1933 : 405.

Type material. — The original description stated that twelve examples had been seen; only two, labelled

“types! décrits Joum. de Conch, t. 23, p. 41, et figurés pi. 4, f. 7 & 7a. île Art, ex auctore 1 .", are present in

the MHNB. There are none in the MNHN. Although one of the specimens is labelled 7a, the original plate did

not distinguish fig. 7 from fig. 7a. Because specimen 7a (Figs 44-45) has more mature apertural and umbilical

features, I designate it lectotype (dimensions: diameter 23 mm, length 28 mm).

FIGS 44-46. — Trochus (Polydonta) calcaratus Souverbie in Souverbie & Montrouzier, 1875. 44-45, lectotype (diameter 23 mm,

length 28 mm); 46, paralectotype (diameter 21 mm, length 23 mm).

Source : MNHN, Paris

— 433 —

Remarks

Pilsbry (1889), with some reservations, relegated a number of taxa to the synonymy of

T. calcaratus , some of which in fact pre-date it. More recently, CERNOHORSKY (1978) and WILSON

(1993) placed T. calcaratus in synonymy with T. histrio Reeve, 1861 (apparent syntype in BMNH

illustrated by Kaicher 1979, under T. sacellum, card No. 2169). Our current understanding of

the taxonomy of the genus Trochus, however, is woefully inadequate. Numerous species of Tro-

chus were described during the last century, many of which simply represent individual variants

of intraspecifically variable species. The situation is complicated further by the fact that descrip¬

tions and illustrations were often poor and that the type material of many is now lost. Although

the synonymy proposed by CERNOHORSKY may be sound, it needs to be investigated in detail.

It remains quite possible that there is a still earlier name for the species (for example Trochus

sacellum Philippi, 1855).

Both remaining specimens in the type lot of T. calcaratus are here illustrated (Figs 44-46)

to show the variation in the development of the peripheral angle and projections. The species

is relatively small for the genus, showing mature apertural characters at length 30 mm; one of

its most notable features is the series of hollow triangular projections at the periphery, but this

is not a unique character (cf. Trochus aemulans (A. Adams, 1855) from China and T. tubiferus

Kiener, 1850, from New Caledonia). The specimens are white to pale buff, with broad reddish

axial markings.

Conclusion

One of the many dubiously valid species of Trochus s. str. (Trochinae, Trochini); in need

of detailed comparison.

Trochus (Tectus) fabrei Montrouzier in Fischer, 1878

(Figs 47-48)

Trochus (Tectus) fabrei Montrouzier in Fischer, 1878a: 64; Fischer 1879 in 1875-1880:

384, pi. 116, figs 1, la. Type loc.: “ins. Lifu” [Lifou] (Loyalty Islands, New Caledonia).

Trochus fabrei ; PlLSBRY 1889: 21, pi. 3, figs 21, 22.

Type material. — Two specimens are present in the MHNB; one (leg. Montrouzier) from "Lifu" [Lifou]

and a second (leg. Lambert), a fossil from raised beach deposits on the île des Pins (both localities in the New

Caledonian archipelago). They bear the label “J. Conch, t. 26: 64 & 206”. No figure was provided with the

original description (Fischer 1878a), the first illustrations being those given in the "Spécies général" (Fischer

1879 in 1875-1880). Both specimens were illustrated, the Lifou one as fig. la, and the larger, fossil one as fig. 1.

The Lifou specimen is here refigured (Figs 47-48) and designated lectotype (dimensions: diameter 26.5 mm,

length 34.3 mm).

Remarks

This material is indeed referable to Tectus Montfort, 1810, a taxon currently afforded full

generic rank. The strong columella pleat suggests referral to Tectus s. str.

Source : MNHN, Paris

— 434 —

Fischer’s remarks on the species are pertinent (Fischer 1879 in 1875-1880). There is con¬

siderable similarity with Tectus triserialis (Lamarck, 1822), but T. fabrei would seem to be less

elevated and to have a sculpture of granular spiral cords rather than the spirally aligned rows

of nodules typical of T. triserialis. However, the extent to which T. triserialis varies in length

and sculpture needs to be further investigated. T. pyramis (Born, 1778) is less elevated, generally

more coeloconoid and, with the exception of strong peripheral granules on the spire whorls, is

smoother.

FlGS 47-48. — Trochus (Tectus) fabrei Montrouzier in Fischer, 1878, leclotype (diameler 26.5 mm, length 34.3 mm).

Conclusion

A species of Tectus s. str. (Trochinae, Trochini); further comparison with T. triserialis

(Lamarck, 1822) is needed.

Trochus (Zizyphinus) poupineli Montrouzier in Souverbie & Montrouzier, 1875

(Figs 49-50)

Trochus (Zizyphinus) poupineli Montrouzier in Souverbie & Montrouzier, 1875: 40, pi. 4,

fig. 6; Fischer 1878b: 210; 1879 in 1875-1880: 387, pi. 116, fig. 3. Type loc.: “ins. Art” (New

Caledonia).

Calliostoma poupineli ; PlLSBRY 1889: 350, pi. 17, fig. 4L

Dactylastele poupineli ; Marshall 1995, figs 79-82, 135, 155.

Source : MNHN, Paris

— 435 —

Type material. — The single specimen mentioned in the original description remains in the MHNB, it

must be regarded as the holotype (Figs 49-50) (dimensions: diameter 8.4 mm, length 11.7 mm). It is labelled

"type décrit Journ. de Conch, et figuré dans Kien. t. 23, p. 40, pi. 4, f. 6 [= figure in J. Conch. Paris], île Art,

ex auctore". There are two “ex auteur" specimens in the MNHN.

Remarks

Frequently regarded a synonym of Ziziphinus complus A. Adams, 1854, and used in place

of that name to avoid confusion with Calliostoma comtus (Philippi, 1855) (see for example

PlLSBRY 1889). The species has recently been discussed in detail by MARSHALL (1995) who

recognised it as distinct from Calliostoma comptum, and referred it to the new genus Dactylastele.

Conclusion

A valid species (Calliostomatidae).

Turbo artensis Montrouzier in Souverbie, 1860

(Fig. 51)

Turbo artensis Montrouzier in Souverbie, 1860b: 370; SOUVERBIE 1861: 274, pi. 11, fig. 5;

Fischer 1873: 58, pi. 37, fig. 1, pi. 38, fig. 1; PlLSBRY 1888: 196, pi. 45, figs 96-97; FlSCHER-

PlETTE 1950: 19; Kaicher 1988: 5291. Type loc.: “ins. Art” (New Caledonia).

Turbo (Senectus) artensis', Melvill & Standen 1895 in 1895-1897: 124.

Turbo (Marmorastoma) artensis ; CERNOHORSKY 1978: 39, pi. 10, fig. 3.

Type material. — The original description did not specify the number of specimens available, but Souverbie

(1861) stated that numerous examples had been seen. Two specimens are present in the MHNB, labelled “No. 1

et 2, types décrits Journ. de Conch, t. 8, p. 370 et t. 9, p. 274, No. 1 type figuré pi. 11, fig. 5. île Art, don de

l'auteur”. Two further lots, labelled as types, are present in the MNHN. One of these, in the typothèque, contains

four adult specimens and three juveniles; one of the adults bears a note stating “un des exemplaires ayant servi

Source : MNHN, Paris

— 436 —

à la diagnose. Ex auctore !”. The second lot, in the Journal de Conchyliologie collection, has two specimens

which were listed as types by Fischer-Piette (1950: 19), the larger of these he also cited as the figured specimen.

There is, however, nothing associated with this specimen to indicate that this was the case (normally in the

Journal de Conchyliologie collection there would be, V. Héros in lit.) and in view of the fact that the MHNB

specimen No.l is specifically annotated as being the figured one I designate it as lectotype (here refigured.

Fig. 51) (dimensions: diameter 44.5 mm, length 47 mm). This is also the course of action advised by Recom¬

mendation 74D of the ICZN, since the majority of the author’s types are in Bordeaux. The other MHNB specimen

and the MNHN specimen cited as being one of those upon which the diagnosis was based should be considered

paralectotypes, but the type status of the remainder is dubious.

Remarks

This would seem to be a fairly distinct species characterised by shape and sculpture. It has

relatively strong spiral cords, the intervals between which have a single finer spiral thread which

is crossed by >-shaped axial pliculae producing a close-set, herring-bone sculpture. There is no

umbilicus and the peristome is markedly drawn out and flaring where the columella and outer

lip meet. The ground colour is orange-brown (perhaps somewhat faded) with a few darker and

lighter axial stripes. The operculum is for the most part smooth, but possesses some rippling

on the outer lip side; the markings shown in Fig. 51 are a result of discoloration.

Conclusion

A valid species of Turbo (Turbinidae, Turbininae).

Turbo laetus Montrouzier in Souverbie & Montrouzier, 1863

(Figs 52-54)

Turbo laetus Montrouzier in Souverbie & Montrouzier, 1863b: 277, pi. 12, fig. 2. Type

loc.: “Balade et ins. Art” (New Caledonia).

Source : MNHN, Paris

— 437 —

Leptothyra beta; PlLSBRY 1888: 258, pi. 63, figs 29, 30; Hedley 1899: 408, 1907: 479,

1909: 355; Shopland 1902: 176; HIDALGO 1904-1905: 245; Melvill 1918: 152; Iredale 1929:

273; Viader 1937: 55.

Type material. — There are four syntypes in the MHNB. labelled “types décrits Joum. de Conch. No. 1

celui figuré, pi. 4 [sic], f. 2. île Art. don. de l'auteur". No. 1 is here refigured (Figs 52-54) and designated

lectotype (diameter 5.0 mm). There are a further eleven specimens labelled as syntypes in the MNHN and four

specimens “ex auteur". The original description stated that there were twenty-five specimens.

Remarks

This name is a junior primary homonym of Turbo laetus Philippi, 1849. SOWERBY (1886) pro¬

posed the replacement name costulosus, an adaptation of “ Turbo costulatus Gould” (ms ?) (non Wood,

1828). This is one of numerous species of Collonista Iredale, 1918, described from the Indo-West

Pacific. J. H. McLean, who is currently revising the Colloniinae, has indicated (McLean in lit.)

that the earliest non-homonymous name for this taxon is Collonia granulosa Pease, 1868, based on

material from the Caroline Islands. A lectotype for the latter was designated and figured by Johnson

( 1994), but the figure number was inadvertently transposed with that of C. picta Pease, 1868. John¬

son’s fig. 21 is in fact that of the lectotype of C. granulosa (error noted by McLean).

Conclusion

A junior primary homonym, the earliest available name for which is Collonia granulosa

Pease, 1868 (fide McLean in lit.); should be referred to Collonista Iredale, 1918 (Turbinidae,

Colloniinae).

Figs 52-54. — Turbo laetus Montrouzier in Souverbie & Montrouzier. 1863. specimen figured with original description and here

designated lectotype (diameter 5.0 mm).

Source : MNHN, Paris

— 438 —

Turbo naninus Souverbie in Souverbie & Montrouzier, 1864

(Figs 55-58)

Turbo naninus Souverbie, in Souverbie & Montrouzier, 1864: 263, pl. 10, fig. 6; SOUVERBIE

1875: 293, pl. 13, fig. 9. Type loc.: “ins. Art” (New Caledonia).

Leptothyra nanina-, Pilsbry 1888: 259, pl. 58, figs 55, 56; Hedley 1907: 479; 1909: 355;

Cernohorsky 1978: 39, pl. 10, fig. 5.

Type material. — There are two separate type lots of this species in the MHNB, each with one specimen.

The first is labelled "types décrits et figurés Journ. de Conch, t. 12, p. 235 [s/c], pl. II [sic], fig. 6 (Mala.). lie

Art, don de l’auteur”, whilst the label attached to the second states "Journ. de Conch, t. 12, p. 243 [s/c], pl. 10.

fig. 6 (Mala.) in ibid., t. 23, p. 293, pl. 13, fig. 9. Type characleribus emendatis. ex auteur." The original

description stated that only a single specimen was available and thus the specimen in the first lot, upon which

the original description and figure were based, must be regarded as the holotype (Fig. 55. diameter 3.0 mm),

despite the fact that Souverbie (1875) considered it “un peu imparfait". The specimen in the second lot (cf. Figs

56-58) is simply a fresher example that Souverbie (1875) used to augment the description, but it has no type

status. There are two further specimens labelled syntypes in the MNHN; these are ex auctore specimens, but

they likewise have no status as types.

Remarks

Resembles the preceding species, but the last adult whorl is somewhat biangular with a

distinct angle at the shoulder and a weaker one marking the periphery of the base. The sculpture

Figs 55-58. — Turbo naninus Souverbie in Souverbie & Montrouzier. 1864. 55. holotype (diameter 3.0 mm): 56-58. fresher.

characleribus emendatis specimen of Souverbie (1875) (diameter 3.0 mm).

Source : MNHN, Paris

— 439 —

is finer and, judging from the material available, the species would seem to be smaller. This

taxon also belongs within Collonista and is being studied by J. H. McLean, who indicated (in

lit.) that it may prove to be a New Caledonian endemic.

Acknowledgements

1 would like to thank the authorities of the MHNB for permission to examine the New Caledonian

types in their custody, and in particular Mme Nathalie MEMOIRE without whose assistance this study would

not have been possible. 1 thank also Bernard MÉtivier (MNHN) and Kathie Way (BMNH) for their help

whilst examining types under their care, Ian LOCH (AMS) for the loan of Brazier types, Gary Rosenberg

(ANSP) for Pease types and Dick KlLBURN for photographs of types in the Zoological Survey of India.

Philippe Bouchet, Dick Kilburn, Bruce Marshall and Jim McLean provided valuable comments on

the manuscript and Virginie Héros checked the accuracy of my statements regarding the MNHN types.

The study was funded through my personal Foundation for Research Development grant.

Source : MNHN, Paris

— 440 —

FIGS 65-69. — Ziziphinus polycliromus A. Adams, 1853, Ziziphinus picturatus A. Adams, 1853, Trochus fournieri Crosse, 1863

and Trochus (Clanculus) bathyraphe Smith, 1876. 65, Z. polycliromus, lectotype (BMNH), length 12.3 mm: 66, Z. picturatus,

lectotype (BMNH), length 11.5 mm; 67, T. fournieri, holotype (BMNH), length 7.9 mm; 68-69. T. hathyraphe, one of two

syntypes (BMNH), diameter 8.3 mm.

Source : MNHN, Paris

— 441 —

FIGS 70-76. — Turbo semilugubris Deshayes, 1863, Monodonta foveolala A. Adams, 1853, Euchelus angulatus Pease, 1868, and

Tallorbis roseola G. & H. Nevill, 1869. 70-71, T. semilugubris, one of two syntypes (MNHN), diameter 2.66 mm; 72-73,

M. foveolala, lectotype (BMNH), diameter 5.2 mm; 74-75, E. angulatus Pease, 1868, lectotype (ANSP 40671), diameter

4.6 mm, denticles inside outer lip present, but not obvious (arrows); 76, T. roseola, lectotype (ZSIC), diameter 9.1 mm

(photograph courtesy of R. N. Kilburn).

Source : MNHN, Paris

-442-

Figs 77-85. — Ethalia rhodomphala Smith. 1903, Elhalia floccala Sowerby. 1903 and Isanda pulchella A. Adams, 1855. 77-79.

E. rhodomphala, lectotype (BMNH). diameter 6.9 mm; 80-82, E. floccala, holotype (BMNH). diameter 7.9 mm; 83-85,

I. pulchella, holotype (BMNH), diameter 7.5 mm.

Source : MNHN, Paris

— 443 —

REFERENCES

Abbott R. T., 1958. — The marine mollusks of Grand Cayman Island, British West Indies. Monogr. Acad. nat.

Sci. Philad. 11, 138 p., plus plates and index.

Adam W. & Leloup E., 1938. — Résultats scientifiques du voyage aux Indes Orientales Néederlandaises. Mém.

Mus. r. Hist. nat. Belg. 2 (19) : 1-209.

ANONYMOUS, 1985. — International Code of Zoological Nomenclature, third edition. London, International Trust

for zoological Nomenclature, 338 p.

CERNOHORSKY W. O., 1978. — Tropical Pacific marine shells. Sydney: Pacific Publications.

CROSSE h., 1898. — Nécrologie - le R. P. Xavier Montrouzier. J. Conch., Paris 46: 42-44.

DaUTZENBERG P. & BOUGE J-L., 1933. — Les mollusques testacés marins des établissements français de

l’Océanie. J. Conch., Paris 33 : 41-108, 145-326, 351-469.

FISCHER H., 1904. — Nécrologie - le R. P. Lambert. J. Conch., Paris 52 : 99-100.

FISCHER P., 1873. — Genre Turbo. In L. C. KlENER (ed.). Spécies général et iconographie des coquilles vivantes.

Paris. J.-B. Baillère et fils 10 : 1-128, pis 1-42.

— 1875. — Catalogue des mollusques appartenant aux genres Turbo, Calcar et Trochus, recueillis dans les

mers de l’archipel calédonien. J. Conch., Paris 23 : 44-51.

— 1875-1880. — Genres Calcar, Trochus, Xenophora. Tectarius et Risella. In L. C. KlENER (ed.). Spécies

général et iconographie des coquilles vivantes. Paris, J.-B. Baillère et fils 11.

— 1876. — Remarques sur la synonymie et l'habitat de quelques espèces de mollusques de la Nouvelle-Calé¬

donie. J. Conch., Paris 24 : 148-151.

— 1878a. — Diagnoses trochorum novorum. J. Conch., Paris 26: 62-67.

— 1878b. — Catalogue des mollusques appartenant aux genres Turbo, Calcar et Trochus, recueillis dans les

mers de l'archipel calédonien (Supplément), suivi de la liste des espèces des genres Delphinula, Liotia et

Phasianella. J. Conch., Paris 26 : 205-211.

— 1878c. — Diagnoses molluscorum novorum. J. Conclu, Paris 26: 211-213.

FlSCHER-PlETTE E., 1950. — Liste des types décrits dans le Journal de Conchyliologie et conservés dans la

collection de ce journal. J. Conch., Paris 90 : 8-22, 65-81, 149-179.

Franc a., 1956. — Mollusques marins. In Campagne de la Calypso en Mer Rouge. Annls Inst, océanogr.,

Monaco 32: 19-60.

HEDLEY C., 1899. — The Mollusca of Funafuti. Part 1. - Gasteropoda. Mem. Aust. Mus. 3 (7): 397-488.

— 1901. — A revision of the types of the marine shells of the Chevert Expedition. Rec. Aust. Mus. 4: 121-130.

— 1907. — The Mollusca of Mast Head Reef, Capricorn group, Queensland. Part 2. Proc. Linn. Soc. N.S.W.

32: 476-513.

— 1908. — Studies on Australian Mollusca. Part X. Proc. Linn. Soc. N.S.W. 33: 456-489.

— 1909. — Catalogue of the marine Mollusca of Queensland. Rep. Australas. Ass. Advmt Sci. 12: 343-371.

— 1915. — Studies on Australian Mollusca. Part XII. Proc. Linn. Soc. N.S.W. 39: 695-755, dated 1914.

Herbert D. G., 1992. — Revision of the Umboniinae in southern Africa and Mozambique (Mollusca: Gastropoda:

Trochidae). Ann. Natal Mus. 33 (2): 379-459.

— (in prep). — A revision of the Eucyclinae, tribe Chilodontini (Gastropoda: Trochidae) of southern Africa.

HIDALGO J. G., 1904-1905. — Catalogo de los moluscos testâceos de las islas Filipinas, Joiô y Marianas. I.

Moluscos mari nos. Madrid: Real Academia de Ciencias Exactas, Fiscias y Naturales.

HICKMAN C. S. & McLean J. H., 1990. — Systematic revision and suprageneric classification of trochacean

gastropods. Science series. Natural History Museum of Los Angeles County 35: 1-169.

IREDALE T., 1929. — Queensland molluscan notes I. Mem. Qd Mus. 9: 261-297.

JANSEN R, 1994. — Notes on Australian species of Euchelus and Herpetopoma (Gastropoda: Trochidae) with

description of five new species. Molluscan Research 15: 55-66.

JOHNSON R. I., 1964. — The Recent Mollusca of Augustus Addison Gould. Washington, Smithsonian Institution.

— 1994. — Types of shelled Indo-Pacific mollusks described by W. H. Pease. Bull. Mus. comp. Zool. Harv.

154 (1): 1-61.

Source : MNHN, Paris

— 444 —

KAICHER S. D„ 1979. — Card catalogue of world shells. Pack No. 21, Trochidae I. St Petersburg. Florida,

Kaicher.

— 1988. — Card catalogue of world shells. Pack No. 52, Turbinidae 1. St Petersburg, Florida, Kaicher.

— 1990. — Card Catalogue of world shells. Pack No. 56, Trochidae V. St Petersburg, Florida, Kaicher.

KEEN a. M., 1960. — In R. C. MOORE (ed.). Treatise on Invertebrate Paleontology. Part I. Mollusca 1. Boulder,

Colorado, Geological Society of America and University of Kansas.

Ladd h. S., 1966. — Chitons and gastropods (Haliotidae through Adeorbidae) from the western Pacific islands.

Prof. Pap. U. S. Geol. Surv. 531: 1-98, pis 1-16.

LAMBERT. P., 1874. — Description d'un Stomatella provenant de la Nouvelle-Calédonie. J. Conch.. Paris 22 :

374.

Marshall B. A., 1979. — The Trochidae and Turbinidae of the Kermadec Ridge (Mollusca: Gastropoda).

N. Z. J. Zool. 6 (4): 521-552.

— 1995. — Calliostomatidae (Mollusca: Gastropoda: Trochoidea) from New Caledonia, the Loyalty Islands

and the northern Lord Howe Rise. In P. BOUCHET (ed.). Résultats des Campagnes MUSORSTOM. Vol. 14.

Mém. Mus. nain. Hist. not. 167: 381-458.

MASTALLER M., 1979. — Beitràge zur faunistik und ôkologie der Mollusken und Echinodermen in der Koral-

lenriffen bei Aquaba. Rotes Meer. Ph. D. Thesis. Munich.

Melvill J. C., 1891. — Descriptions of eleven new species belonging to the genera Columbarium. Pisania,

Minolia, Liotia. and Solarium. J. Conch.. London 6: 405-411.

— 1918. — Descriptions of thirty-four new species of marine Mollusca from the Persian Gulf, Gulf of Oman,

and Arabian Sea, collected by Mr F. W. Townsend. Ann. Mag. nat. Hist. ser. 9 1: 137-158.

Melvill J. C. & Standen R„ 1895-1897. — Notes on a collection of shells from Lifu and Uvea, Loyalty

Islands, formed by the Rev. James and Mrs Hadfield, with a list of species. J. Conch., London 8: part I,

1895-1896: 84-132; part 2, 1896: 273-315, 1897: 379-381; part 3, 1897: 396-421.

— 1901. — The Mollusca of the Persian Gulf, Gulf of Oman and Arabian Sea, as evidenced mainly through

the collections of Mr F. W. Townsend, 1893-1900; with descriptions of new species. Proc. zool. Soc. Lond..

1901 2: 327-460.

NEVILL G. & NEVILL H., 1875. — Descriptions of new marine Mollusca from the Indian Ocean. J. Asiatic Soc.

Beng. 44: 83-104, pis vii-viii.

O'REILLY R, 1931. — Un missionnaire naturaliste - Xavier Montrouzier (1820-1897) [including bibliography].

Rev. Hist. Missions. March 1931: 1-23.

PlLSBRY H. A., 1888. — Turbinidae. In G. W. TRYON (ed.). Manual of Conchology, Philadelphia, Academy of

Natural Sciences. 10: 161-290, pis 37-64.

— 1889. — Trochidae. In G. W. TRYON (ed.). Manual of Conchology. Philadelphia, Academy of Natural

Sciences. 11: 1-519, pis 167.

— 1890. — Stomatellidae, Scissurellidae, Pleurotomariidae, Haliotidae, Scutellinidae, Addisoniidae, Cocculi-

nidae, Fissurellidae. In G. W. TRYON (ed.). Manual of Conchology, Philadelphia, Academy of Natural

Sciences. 12: 1-323, pis 164.

— 1904. — New Japanese Mollusca: Gastropoda. Proc. Acad. nat. Sci. Philad. 56: 3-37.

— 1905. — New Japanese Mollusca. Proc. Acad. nat. Sci. Philad. 57: 101-122.

PONDER W. F., 1978. — The unfigured Mollusca of J. Thiele, 1930 published in Die Fauna Südwest-Australiens.

Rec. W. Aust. Mus. 6 (4): 423-441.

ROSEWATER J., 1972. — The family Littorinidae in the Indo-Pacific. Part II. The subfamilies Tectariinae and

Echininae. Indo-Pacific Mollusca 2 (12): 507-533.

SCHEPMAN M. M., 1908. — The Prosobranchia of the Siboga Expedition. Part 1. Rhipidoglossa and Docoglossa,

with an appendix by Prof. R. Bergh. Siboga Expeditie 1899-1900. Leiden, E. J. Brill 49a: 1-107, pis 1-9.

SHOPLAND E. R., 1902. — List of marine shells collected in the neighbourhood of Aden between 1892 and 1901.

Proc. malac. Soc. Lond. 5: 171-179.

SMITH E. A., 1884. — Mollusca. In Report of the zoological collections made in the Indian Ocean during the

voyage of HMS “Alert”. 1881-1882. London, British Museum: 34-116.

SOUVERBIE S.-M., 1858. — Descriptions d’espèces nouvelles de l'archipel calédonien. J. Conch., Paris 7: 376.

— 1860a. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 8 : 123-126.

Source : MNHN, Paris

— 445 —

— 1860b. — Descriptions d'espèces nouvelles. J. Conch., Paris 8 : 369-371.

— 1861. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 9 : 271-284.

— 1875. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 23: 282-295.

SOUVERBIE S.-M. & MONTROUZIER X., 1862. — Descriptions d’espèces nouvelles de l’archipel calédonien.

J. Conch.. Paris 10 : 231-248.

— 1863a. — Descriptions d’espèces nouvelles de l’archipel calédonien et des îles Salomon et Woodlark.

J. Conch., Paris 11: 161-176.

— 1863b. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 11 : 276-287.

— 1864. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 12: 261-275.

— 1866. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 14: 138-151.

— 1870. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 18 : 71-83.

— 1875. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 23: 33-44.

— 1879. — Descriptions d’espèces nouvelles de l’archipel calédonien. J. Conch., Paris 27: 25-34.

Sowerby G. B.. 1886. — Turbo. In Thesaurus conchytiorum. London, Sowerby 5: 189-234, pis 1-14 (493-506).

Springsteen F. J. & Leobrera F. M„ 1986. — Shells of the Philippines. Manila, Carfell Seashell Museum.

VlADER R., 1937. — Revised catalogue of the testaceous Mollusca of Mauritius and its dependencies. Mauritius

Inst. Bull. 1 (2): i-xiii, 1-111, plus map.

WILSON B., 1993. — Australian Marine Shells, Prosobranch Gastropods. 1, Kallaroo, Odyssey.

Source : MNHN, Paris

Bulletin du Muséum national d'Histoire naturelle, Paris, 4 e série. 18. 1996

Section A, n“ 3-4 : 447-457

A living fossil Waikalasma boucheti n.sp.

(Cirripedia, Balanomorpha) from Vanuatu (New Hebrides),

Southwest Pacific

by John S. BUCKERIDGE

Abstract. — This paper describes Waikalasma boucheti n.sp., the first known living representative of the

Eolasmatinae, a cirripede subfamily previously known only from the Palaeœcene-Miocene of Australasia. The

present material, recovered from the bathyal environment off Vanuatu, strengthens the case for Waikalasma being

considered as the outgroup of all modern acom barnacles, and provides further evidence to confirm the Eolas¬

matinae as one of the most primitive groups of the Balanomorpha.

Key-words. — Waikalasma boucheti, Eolasmatinae, Balanomorpha, Cirripedia, Vanuatu, Southwest Pacific.

Découverte d’une balane, fossile vivant, Waikalasma boucheti n.sp. (Cirripedia, Balanomorpha)

aux Vanuatu (Nouvelles-Hébrides), sud-ouest Pacifique.

Résumé. — Cet article décrit Waikalasma boucheti n.sp., première espèce vivante des Eolasmatinae, une

sous-famille de cirripèdes seulement connue du Paléocène-Miocène d’Australasie. Ce matériel, découvert dans

un faciès bathyal au large de Vanuatu, confirme le fait que Waikalasma est un groupe extérieur des cirripèdes

balanes et fournit une autre évidence que les Eolasmatinae sont l’un des groupes les plus primitifs des Balano-

Mots-clés. — Waikalasma boucheti, Eolasmatinae, Balanomorpha, Cirripedia, Vanuatu, sud-ouest Pacifique.

J. S. Buckeridge, Department of Civil and Environmental Engineering, UNITEC, Private Bag 92025, Auckland, New Zealand.

INTRODUCTION

The first known species of Waikalasma was described as a fossil by BUCKERIDGE (1983),

from the early Miocene Waikawau cast beds of Port Waikato, New Zealand. The genus is signi¬

ficant in that it possesses eight plates in the shell wall and is surrounded by two or more rows

of smaller, imbricating plates. These characters are considered primitive (BUCKERIDGE 1983),

and place Waikalasma not only early, but central in the phylogeny of the Balanomorpha (BUCK¬

ERIDGE & Newman 1992; Newman & Yamaguchi 1995; Buckeridge 1996). The specimens

described in this paper were collected either by a Waren dredge (DW), or beam trawl (CP),

from waters near Vanuatu as part of the MUSORSTOM 8 (1994) expedition. Both specimens

were recovered from what appears to have been a rocky substrate.

The barnacles are preserved in alcohol, and have been examined with the aid of light mi¬

croscopy and dissection. In addition to photographs of the exterior of both holotype and paratype,

Source : MNHN, Paris

— 448 —

illustrations of the whole animal, shell and appendages, have been drawn with the aid of a

camera lucida.

The holotype MNHN-G2428 and the paratype MNHN-Ci2506, have been deposited in the

Muséum national d’Histoire naturelle (MNHN), Paris, France.

SYSTEMATIC PART

Family PACHYLASMATIDAE Utinomi, 1968

Subfamily EOLASMATINAE Buckeridge, 1983

Diagnosis (emend.). — Shell of eight distinct compartmental plates, rostrolatera not entering sheath, second

carinolatera barely entering sheath; basis membranous.

Distribution (emend.). — Upper Palæocene to Lower Miocene (New Zealand), Lower Miocene (Victoria,

Australia); Recent, 700-850 m (off Vanuatu).

Discussion

When originally erected, this subfamily comprised two monotypic genera, represented by

Eolasma maxwelli Buckeridge, 1983 (Lower Palæocene, New Zealand) and Waikalasma juneae

Buckeridge, 1983 (Lower Miocene, New Zealand). Since then, the geographic distribution has

been extended to the Miocene of Victoria, Australia, by Eolasma rugosa Buckeridge, 1985 and

is now known from the Recent in seas off Vanuatu (previously known as the New Hebrides).

Initially it was thought that there were no imbricating plates present, but close examination of

the holotype of Waikalasma juneae demonstrated the remains of at least two whorls of small

imbricating plates (Buckeridge & Newman 1992). In the same paper, following Yamaguchi

& Newman (1990), a new interpretation of capitular plate arrangement in Waikalasma was pro¬

vided. It is this interpretation, of rostrum-rostrolatus-carinolatus-second carinolatus-carina

(RRL-CL-CL 2 -C), that is adopted here.

Genus WAIKALASMA Buckeridge, 1983

Diagnosis (emend.). — Shell bilaterally symmetrical with eight solid, calcareous, weakly articulated com¬

partmental plates, including very wide carina (C), moderately narrow rostrum (R), and narrower, paired rostrolatera

(RL), carinolatera (CL) and second carinolatera (CL 2 ); alae well developed, particularly on carina, but almost

confluent with paries; radii absent; with two or more whorls of imbricating plates; basis membranous; caudal

appendages absent.

Distribution (emend.). — Lower Miocene to Recent. Oceania.

Type species. — Waikalasma juneae Buckeridge, 1983. Miocene (Aquitanian), Port Waikato, New Zealand.

Source : MNHN, Paris

-449-

Discussion

The origin of the Balanomorpha is within the Brachylepadomorpha, and a clear lineage can be

deduced with Waikalasma and Eochionelasmus placed at the earliest stages of balanomorph radiation

(Buckeridge & Newman 1992; Newman & Yamaguchi 1995). However, the hard parts of Wai¬

kalasma differ primarily from a Neobrachylepas form in only two ways: the loss of median latera,

and the adoption of a configuration whereby imbricating plates are added to the shell from outside

the preceding whorl. The soft parts of Waikalasma are quite distinct from those of Neobrachylepas ,

which are rather unusual and have been interpreted as adaptations to feeding (Newman & YAMAGU-

CHi 1995). The structural arrangement of Waikalasma represents a grade of organisation between

balanomorphs with three pairs of partially and fully integrated latera, and also between those with

numerous whorls of basal imbricating plates and those with none.

Waikalasma boucheti n.sp.

(Figs la-f; 2a-g; 3a-h; 4a, b)

Material examined. — Vanuatu, MUSORSTOM 8: stn CP 1080, 15°57’S - 167°27’E, 799-850 m, 5.X.1994.

1 specimen (holotype); stn DW 1113, 14°53'S - 167°06’E, 700-736 m, 8.X.1994. 1 specimen (paratype).

Etymology. — The new species is named for Philippe Bouchet, Muséum national d’Histoire naturelle,

Paris. Dr Bouchet was one of the principal scientists on the 1994 MUSORSTOM expedition.

Habitat. — Station CP 1080 was situated eleven kilometres off the north east coast of île Malekula. The

barnacle was attached to a brown, weathered, tuffaceous sandstone, the broken surface of which suggests that

this may have been an outcrop rather than a large pebble. Associated fauna at this site was rich, including

teleosts, scaphopods, gastropods and other crustaceans. The specimen from stn DW 1113 was recovered from

nine kilometres off the north east cape of île Santo, attached to a large pebble of fossiliferous sandy tuff. Both

areas are characterised by expanses of both rocky (predominantly basaltic) and muddy bottoms. A full account

of substrate and conditions is provided in Richer de Forges et al. (1996).

Diagnosis

Waikalasma with two or more whorls of large imbricating plates, inner whorl with total of

eight plates; scutum triangular and very elongate with long, low articular ridge, adductor muscle

scar weak, central; tergum narrow, inverted “V-shaped”, with deeply excavated basal margin;

spur at basiscutal angle.

Description

Holotype (MNHN-CÎ2428): rostro-carinal diameter 30.7 mm; width 32.5 mm; height

24.7 mm. Base membranous.

Paratype (MNHN-Ci2506): rostro-carinal diameter 26.2 mm; width 23.2 mm; height

12.8 mm.

Carina well developed, semi-conic, with extended alae, approximately 1.5 times height of

rostrum. Lateral plates approximate, separated by very narrow alar zones, RL and CL 2 clearly

separated from paries of rostrum and carina respectively by broad exposed alar areas on latter

plates. Internally, RL not entering the sheath, CL 2 only slightly, extending into sheath as very

Source : MNHN, Paris

— 450 —

narrow zone of less than 1 mm wide; RL exposed for approximately 70% of distance between

basal margin and sheath, being otherwise overlapped by CL. All plates transversely sculptured

with fine growth lines, each lateral plate with central, very weakly developed longitudinal rib.

On most plates, growth lines with slight basal deflection approaching rib. Carina, with three

similar, but less clearly defined ribs. Basally ribs approximately aligned with edges of first whorl

of imbricating plates. Alae almost confluent with paries, possessing fine apico-basal striae trans¬

versely cutting well formed growth lines, latter slightly inflected at alar margin, welting absent.

Holotype with imbricating plates arranged in two whorls with eight plates in each; inner plates

larger, up to 6.3 mm high and 9.4 mm wide, and placed to overlap area of abutment of com-

partmental plates, inner whorl of imbricating plates in turn overlapped by outer whorl of plates

of up to 1.7 mm height (Fig. If); imbricating plates develop “alar extensions” about 1 mm wide

on overlapped margins; both whorls of imbricating plates closely approximate basally, forming

3.2 mm thick skirt of laminae, angling inwards to outer edge of the compartmental plates. Skirt,

and base of compartmental plates in contact with substrate. Imbricating plates grow by addition

of laminae from base, with outermost plates newest; laminae of both first and second whorls

confluent, although in holotype, plates in outer whorl (= earliest) lack laminae.

Opercula

Tergum narrow, inverted “V-shaped”, basal margin deeply excavated; apico-basal ridge

broadly concave towards carina, basally becoming full width of projected basal angle; articular

ridge low, quadrangular; interior lacking adductor muscle scars or furrows; exterior with well

developed transverse growth lines on apico-basal and articular ridges. Scutum elongate, triangular,

occludent margin 2.6 times length of basal margin; internally with very weakly developed, cen¬

trally placed, adductor muscle pit; articular ridge low, about half length of gently concave articular

margin; exterior with well developed transverse growth lines, crossed by weak longitudinal striae.

Soft parts

Mandible tridentate, lower angle accuminate with two groups of short spines; first maxilla

with two large and three small upper spines, notch poorly developed, centrally with group of

five large spines, lower angle with three medium spines, relatively hirsute overall. Penis rudi¬

mentary, without basidorsal point. Labrum gently curved, with two or three small teeth on each

side near palps, and very finely denticulated surfaces lower on either side. Palps well separated,

rounded, with setae on both inner and outer sides. No eggs or branchiae were noted.

Cirri

Cirri I and II with anterior rami with one segment fewer than posterior rami. Cirrus II

more like cirrus III than cirrus I. Anterior rami of cirri III, V and VI with two or three more

segments than posterior rami. Cirrus IV with anterior rami longer than posterior, but appendage

may have been damaged. Cirrus VI with four pairs of large setae on anterior margin of inter¬

mediate segments. Segment count per ramus as follows (anterior, posterior):

24, 25 29, 32

Source : MNHN, Paris

— 451 —

Source : MNHN, Paris

— 452 —

Colour (in alcohol)

The holotype shell is creamy-white internally and externally stained a dark brownish yellow.

The paratype is cream both internally and externally. Soft tissue in the holotype has a strong

purple tint, particularly on the cirri.

Remarks

The paratype, although smaller, possesses a total of twenty-seven imbricating plates. These

may be recognised as an inner whorl of eight plates, each overlapping a suture in the com-

partmental wall, a second whorl of twelve plates (variously interleaved and overlapping), a third

whorl of six plates (generally overlapping) and a fourth whorl of one plate (Fig. 2f). The plates

are smaller than in the holotype, with only one being over 5 mm in height. From the structure

of the paratype it is easy to see how new plates are added to the outer imbricating whorl(s):

addition is not regular, being accomplished either by interleaving with, or overlapping of, adjacent

plates. At the base of the carina, (the region of greatest compartmental growth), plates are first

interleaved, then placed to overlap (Fig. 4b). This pattern is essentially the same as that observed

in shell compartmental growth: a primary wall of four plates (R-CL-C-CL) is enlarged by re¬

plication of the CL to produce the CL 2 , which is interleaved between the C and CL; this is

followed by addition of the RL, which overlaps both R and CL (see Fig. 4).

In the holotype, both opercula show significant wear in the apical regions, with the tergum

this “smoothed area” extends for more than half the length of the apico-basal ridge; with the

scutum, wear is confined to the upper third of the occludent margin.

The absence of caudal appendages in W. boucheti is intriguing, as these would have

strengthened the status of Waikalasma as an excellent outgroup for the Pachylasmatinae. But

caudal appendages, as are found in Pachylasma, are considered to be plesiomorphic. Natural

processes however, are notably irregular, and this is certainly observed in sessilian phylogeny,

where clear linear relationships, confirmed by sequential loss of pleisiomorphies, are rarely found

(Buckeridge 1996). The presence of a rudimentary penis may indicate the possibility of this

species possessing complemental males, none were observed.

The discovery of two specimens of an extant Waikalasma species provides an opportunity

to confirm a number of unusual characteristics that previously defined a unique, intact, but fos¬

silized, specimen of Waikalasma juneae. In particular, the tight grouping of the lateral plates,

with very little space between RL, CL and CL 2 is confirmed. Further, it is clear that the RL

does not enter the sheath. Although the latter character was suspected in the fossil material, the

fragility of that specimen prevented excavation of sediment from the interior to confirm this.

Unfortunately both specimens of W. boucheti are adults, so do not provide conclusive evidence

of the manner in which the lateral plates are added, i.e. during ontogeny, is the CL 2 intercalated

into the wall as in higher balanomorphs ? This is certainly something additional specimens may

be able to clarify.

Waikalasma boucheti n.sp. (Fig. la-f), may be distinguished from W. juneae (Fig. lg), by

possessing proportionately shorter and clearly rostrally directed lateral plates. Newman (pers.

comm.) now considers that W. juneae probably had about thirty imbricating plates, possibly

arranged in two whorls. His conclusion is based on the ribs or groves in the compartmental

Source : MNHN, Paris

— 453 —

Fig. 2. — Waikalasma boucheli n.sp. a-e, holotype, MNHN-CÎ2428: a. lateral view of whole specimen (left side); b. lateral view

of whole specimen (right side); c, dorsal view of whole specimen; d. carinal view of whole specimen; e. rostral view of

whole specimen, f-g. paratype. MNHN-Ci2506: f. dorsal view of whole specimen; g. lateral view of whole specimen (right

side), (scale: a-e x 1.74; f-g x 2.17). Photography by P. Lozouet, MNHN, Paris (BIMM).

Source : MNHN, Paris

— 454 —

Source : MNHN, Paris

— 455 —

plates, and is accepted here. It necessitates considerably fewer imbricating plates than in the

reconstruction given in BUCKERIDGE & Newman (1992), and is now much closer to the ar¬

rangement seen in W. boucheti. Although a very weak medial rib was observed on lateral plates

of IV. juneae, these appear more clearly defined on the living material. The scuta of W. boucheti

are quite unlike that figured and tentatively attributed to W. juneae by BUCKERIDGE (1983: 64).

I am now confident that his figure 48 (showing a rather broad valve, with a well formed adductor

muscle scar) is not of IV. juneae, rather, it probably represents another bathylasmatid or a worn

balanid scutum. Based on the present reconstruction of W. juneae (Fig. lg), it is clear that the

scuta of this species would have been proportionately much shorter than those of W. boucheti.

FIG. 4. — Waikalasma boucheti n.sp. Schematic plaie arrangement (dorsal plans): a, holotype, MNHN-Ci2428, showing inner

(stippled) and outer (hatched) whorls, both of eight plates, each overlapping a suture in the inner whorl adjacent: b. paratypc,

MNHN-Ci2506, showing inner whorl (stippled), of eight plates, each overlapping a suture in the compartmentai wail, a

second whorl (black), of twelve plates (variously interleaved and overlapping); a third whorl (hatched) of six plates, generally

overlapping: a fourth? whorl of one plate, (clear), overlapping plates on second and third whorls. (Note: this diagram de¬

monstrates relationships between plates. It is neither to scale, nor in proportion.). Drawning by J. Rebière. MNHN. Paris

(Lab. Zoologie Arthropodes).

Affinities

The number of both compartmentai plates and imbricating whorls are sufficient to distinguish

this material from Chionelasmus, which has a rostrum, carina and four latera in the primary

wall, and one whorl of imbricating plates in adults; Pachylasma and Bathylasma may be diffe¬

rentiated from Waikalasma by a lack imbricating whorls. The comparatively larger carina and

the intercalation of six lateral plates into the compartmentai wall distinguish this genus from

Source : /MNHN, Paris

— 456 —

Eochionelasmus. Of the plate structure, the lack of welting in Waikalasma places it closer to

the Chionelasmatinae than the Pachylasmatinae. As previously noted, the soft parts are quite

distinct from Eochionelasmus , but as with Neohrachylepas, this is likely to be a special trophic

adaptation in the latter (Yamaguchi & Newman 1990). In general however, the soft parts of

Waikalasma houcheti are not particularly noteworthy, the maxilla, mandible and labrum are sim¬

ilar to many species within the Pachylasmatinae and Bathylasmatinae, although the absence of

caudal appendages distinguishes this genus from Eochionelasmus, Chionelasmus and Pachylasma.

Using the absence of characters like caudal appendages in establishing “antiquity” for a group

must be approached with caution. If Waikalasma is indeed a good candidate as an outgroup for

the Pachylasmatinae, and I believe this is so, then it is quite likely that caudal appendages may

have been lost some time between the Middle Miocene and the Present, long after the first

pachylasmatines appear in the fossil record.

The discovery of Waikalasma boucheti n.sp. from the living fauna was quite unexpected.

The Waikalasma body plan was initially considered as a brief interlude in balanomorph phylogeny

(Buckeridge 1983). In this sense IV. boucheti may truly be viewed as a living fossil. Unlike

many other recent and significant discoveries in cirripede phylogeny (e.g. Newman 1979, 1985),

this material is not associated with a hydrothermal vent environment.

Biogeography

The Southwest Pacific is confirmed to be of particular significance in sessilian evolution,

as 73% of primitive balanomorphs with a generic age older than the Miocene have their earliest

records there (Buckeridge 1996). Local endemicity at high taxonomic levels is attributed by

NEWMAN (1985) to be either the result of isolation, or association with severe environmental

gradients or ecotomes. Although this species is not part of a hydrothermal vent community, such

as described by Newman ( loc. cit.), it is both insular and bathyal, like so many other pachy-

lasmatine and bathylasmatine réfugiai taxa.

Acknowledgements

The author wishes to sincerely thank Alain Crosnier, ORSTOM (Institut français de recherche scien¬

tifique pour le développement en coopération), for the invitation to work on the MUSORSTOM collections,

for providing access to the material, and for providing a stimulating working environment. Patsy

McLaughlin (visiting scientist at the Muséum national d'Histoire naturelle, Paris), provided thoughtful

and invaluable encouragement, and reviewed the manuscript, as did William Newman (Scripps Institution

of Oceanography, California, United States).

REFERENCES

BUCKERIDGE J. S., 1983. — Fossil barnacles (Cirripedia: Thoracica) of New Zealand and Australia. N. Z. Geol.

Surv. Paleontol. Bull. 50: 1-151.

— 1985. — Fossil barnacles (Cirripedia: Thoracica) from the lower Miocene Limestone. Batesford. Victoria.

Proc. Roy. Soc. Victoria 97 (3): 139-150.

Source : MNHN, Paris

— 457 —

— 1996. — Phylogeny and Biogeography of the Primitive Sessilia and a consideration of a Tethyan origin for

the group. Crustacean Issues , A. A. Balkema Publishers, Rotterdam, 1995 (1996) 10: 255-267.

BUCKERIDGE J. S. & Newman W. A, 1992. — A re-examination of genus Waikalasma (Cirripedia: Thoracica)

and its significance in balanomorph phylogeny. Jo. Pal. 66 (2): 341-345.

Newman W. A.. 1979. — A new scalpellid (Cirripedia); a Mesozoic relic living near an abyssal hydrothermal

spring. Trans. San Diego Soc. Nat. Hi stn. 19 (11):

— 1985. — The abyssal hydrothermal vent invertebrate fauna: a glimpse of antiquity? Biol. Soci. Wash. Bull.

6: 231-242.

Newman W. A. & Yamaguchi T., 1995. — A new sessile barnacle (Cirripedia, Brachylepadomorpha) from the

Lau Back-Arc Basin, Tonga; first record of a living representative since the Miocene. Bull. Mus. nail

Hist. not.. Paris, 4 e sér., 17 (3-4): 221-243.

Richer de Forges B., Faliex E. & Menou J.-L., 1996. — La campagne MUSORSTOM 8 dans l'archipel de

Vanuatu. Compte-rendu et liste des stations. In A. CROSNIER (ed.) Résultats des Campagnes MUSORSTOM

Vol. 15. Mém. Mus. natl Hist. nat. 168 : 9-32.

Yamaguchi T. & Newman W. A., 1990. — A new and primitive Barnacle (Cirripedia: Balanomorpha) from

the North Fiji Basin Abyssal Hydrothermal Field, and its evolutionary implications. Pacific Sci. 44: 135-155.

Note added in proof

Since the acceptance of this manuscript for publication, a further incomplete specimen, probably

attributable to Waikalasma houcheti has been identified by Diana Jones (Western Australian Museum, Perth,

Australia), from Loyalty Ridge, Bathus 3, stn DW 778, 24°43’S - 170°07’E, 750-760 m.

Source : MNHN, Paris

Bulletin du Muséum national d Histoire naturelle, Paris, 4' série. 18. 1996

Section A. n“ 3-4 : 459-545

The terrestrial Isopoda of Corsica (Crustacea, Oniscidea)

by Stefano TAITI & Franco FERRARA

Abstract. — Seventy-six species of terrestrial isopods are recorded from Corsica. Eight species ( Oritoniscus

punctatus, Tiroloscia ntontana, Platyarthrus corsicus, Cylisticus uncinatus, Alloschizidium campaneltii, Armadil-

lidium lanzai, A. littorale and A. torchiai ) are described as new and twelve are newly recorded from the island.

The subspecies Oritoniscus paganus pagamts Racovitza and 0. paganus ocellalus Vandel are raised to species

rank. Phalloniscus pygmaeus (Budde-Lund) is transferred to the genus Sardoiiiscus Arcangeli, and Paraschizidium

rentyi Vandel and Typhloschizidium cottarellii Argano & Pesce to the genus Alloschizidium Verhoeff, of which

Typhloschizidium Arcangeli and Nesolidium Verhoeff are considered to be junior synonyms. About half of the

species have a West-Mediterranean distribution, the majority being strictly Tyrrhenian (38%). About 25% of the

species are endemic, which demonstrates the importance of Corsica as a spéciation area. The oniscidean fauna

of Corsica is strictly related to that of the Tuscan Archipelago and, to a lesser extent, to that of Sardinia. Some

important faunistic affinities exist with southern France (massifs of Maures and Estérel, and the Grasse region)

and the Pyrénées. No species of southern origin (Sicilian or North African) has been collected in Corsica, which

seems to exclude a zoogeographic relationship with those regions. The distribution of the species within Corsica

is discussed. A key to all the species is given.

Key-words. — Crustacea, Isopoda, Oniscidea, taxonomy, new species, zoogeography, Corsica.

Les isopodes terrestres de Corse (Crustacea, Oniscidea)

Résumé. — Soixante-seize espèces d’isopodes terrestres sont signalées de Corse. Huit espèces nouvelles

(Oritoniscus punctatus, Tiroloscia montana, Platyarthrus corsicus, Cylisticus uncinatus, Alloschizidium campa-

nellii, Armadillidium lanzai, A. littorale et A. torchiai) sont décrites et douze sont nouvellement citées de l’île.

Les sous-espèces Oritoniscus paganus paganus Racovitza et O. paganus ocellatus Vandel sont élevées au rang

d'espèce. Phalloniscus pygmaeus (Budde-Lund) est transféré dans le genre Sardoniscus Arcangeli, Paraschizidium

remyi Vandel et Typhloschizidium cottarellii Argano & Pesce dans le genre Alloschizidium Verhoeff, dont

Typhloschizidium Arcangeli et Nesolidium Verhoeff sont des synonymes plus récents. La moitié environ des espèces

a une distribution Ouest-méditerranéenne, la plupart étant strictement tyrrhénienne. A peu près 25 % des espèces

sont endémiques, ce qui prouve l’importance de la Corse en tant que zone de spéciation. Les caractères de la

faune des isopodes terrestres de Corse montrent une stricte corrélation avec les caractères de la faune de l’archipel

toscan et, à un degré moindre, avec ceux de la Sardaigne. Des affinités faunistiques importantes existent entre

la Corse et la France méridionale (massifs des Maures et de l’Estérel et région de Grasse) et aussi avec les

Pyrénées. Aucune espèce d'origine plus méridionale (sicilienne ou nord-africaine) n’a été recueillie en Corse, ce

qui semble exclure toute relation zoogéographique avec ces territoires. La distribution des espèces en Corse est

aussi discutée. Une clé des espèces est proposée.

Mots-clés. — Crustacea, Isopoda, Oniscidea, taxonomie, espèces nouvelles, zoogéographie, Corse.

S. Taiti & F. Ferrara. Centro di Studio per la Faunistica ed Ecologia Tropicali del Consiglio Nazionale dette Ricerche. Via

Romana 17, 50125 Firenze, Italy.

Source : MNHN, Paris

— 460 —

INTRODUCTION

The terrestrial isopods from Corsica, as well as from many islands in the western Medi¬

terranean, are apparently well known, thanks to the work of Vandel who provided an important

picture of the oniscidean population of the island in several contributions (1944a, 1944b, 1945,

1953a, 1954a, 1954b, 1954c, 1960, 1962, 1968a). Moreover, the Oniscidea from many Corsican

caves have been thoroughly investigated by Beron (1972a, 1972b). However, the systematics

of this group of crustaceans has changed markedly in recent years, becoming more precise and

reliable, and the study of limited collections has already provided interesting results, with the

description of a new genus, two new species and the report of a new form for the island (Taiti

& Ferrara 1980; Ferrara & Taiti 1983). Therefore, we believed it appropriate to carry out

the most complete study possible of the terrestrial isopod fauna of Corsica.

Most of the material studied was collected during six expeditions of the Centro di Studio

per la Faunistica ed Ecologia Tropicali, C.N.R., Florence, which covered most of the island and

its characteristic biotopes. Material deposited in the following institutions has also been examined:

Muséum national d’ Histoire naturelle, Paris (MNHN); Staatliches Museum fiir Naturkunde, Stutt¬

gart (SMNS); National Natural History Museum, Sofia; Museo Civico di Storia Naturale, Verona

(MV), and Dipartimento di Biologia Evolutiva dell’Università, Siena. Specimens collected during

some expeditions of the Museo Zoologico “La Specola" dell’Università, Florence (MZUF), on

many of the islets surrounding Corsica (Lanza & Poggesi 1986) have also been studied.

Family Tylidae

G enus TYLOS Audouin, 1826

Tylos europaeus Arcangeli, 1938

(Fig. 1)

ITylos latreillei; Vandel 1954a: 73.

Tylos europaeus; Giordani Soika 1954: 75.

Tylos latreillei europaeus; VANDEL 1960: 108, fig. 48A-C.

Material examined. — 13 66, 6 9 9, 1 juv., Calzarello (near Ghisonaccia), beach, leg. S. Taiti and

S. Campanelli, 18.X.1982.

Previous Records. — Porto-Vecchio (Giordani Soika 1954); Corsica (Vandel 1954a).

Distribution. — This littoral species is known with certainly from the coasts of the whole Mediterranean

Sea and the Atlantic coasts of Europe as far north as Brittany (Giordani Soika 1954; Vandel 1960).

Remarks

Two taxa of Tylos occur in Corsica, as well as in Sardinia and some islands of the Tuscan

Archipelago. According to Vandel (1960) they represent two subspecies of Tylos latreillei

Audouin, 1826 (T. latreillii europaeus Arcangeli, 1938 and T. latreillii sardous Arcangeli, 1938),

while Giordani Soika (1954, 1972) and Kussakin (1982) consider these two taxa as distinct

species since they are well characterised from both morphological and ecological points of view

Source : MNHN, Paris

— 461 —

Source : MNHN, Paris

— 462

(VANDEL 1960). The main problem is the correct identification of the true T. latreillii, to which

one of these two forms most probably corresponds. GlORDANl Soika (1972) suggests the fol¬

lowing synonyms: T. sardous Arcangeli, 1938 = T. ponticus Grebnitzky, 1874, and, tentatively,

T. europaeus Arcangeli, 1938 = T. latreillii Audouin, 1826. In contrast, KUSSAKIN (1982) con¬

siders T. europaeus and T. ponticus as valid species, and T. latreillii a taxon incertae sedis. In

our opinion T. ponticus , rather than T. europaeus, is a more probable synonym of T. latreillii.

In fact, in the original figures of T. latreillii (AUDOUiN 1826, pi. 13, fig. 1.3) the ventral plates

of pleonite 4 are arched and apically pointed, and those of pleonite 5 have apices obliquely

truncate; these features are certainly more similar to those of T. ponticus than T. europaeus (see

also Vandel 1960). Since we are unable to solve with certainty the real synonymy of T. latreillii,

for the moment we prefer to follow Kussakin’s opinion and maintain the names T. europaeus

and T. ponticus for the two taxa.

It is difficult to define the exact distribution of the two species T. europaeus and T. ponticus,

since in the literature both taxa have often been recorded with the name T. latreillii. Re-exami-

nation of the material studied by the different authors is necessary in order to determine to

which of the two species those records belong.

Tylos europaeus is readily distinguished from T. ponticus by: the more developed groove

on the lateral margin of pereonite 1 and the wider epimera of pereonites 2 to 7 (compare Fig. IA

and Fig. 2A); the ventral plates of pleonite 4 shorter and stouter, and those of pleonite 5 having

rounded, instead of truncate, apices (compare Fig. IB and Fig. 2B); the basis of pereopod 1

without a prominent triangular process on the tergal margin (compare Fig. 1C and Fig. 2C); and

the male pleopod 2 endopod with a narrower and more curved apical part (compare Fig. ID and

Fig. 2D).

Tylos ponticus Grebnitzky, 1874

(Fig. 2)

Tylos sardous; GlORDANl Soika 1954: 73.

Tylos latreillei sardous; VANDEL 1960: 109, fig. 48D-E.

Tylos ponticus; Lanza & POGGESI 1986: 121, 180.

Material examined. — 8 S3, Golfo di Sogno, leg. S. Taiti, 13.IV.1981; 6 33, 8 9 9, rocher d'Acciaju

Nord (N of golfe de Sta Giulia), leg. B. Lanza, 8.VIII. 1972.

Previous Records. — Porto- Vecchio (Giordani Soika 1954); Corsica (Vandel 1960); rocher d'Acciaju Nord

(Lanza & Poggesi 1986).

Distribution. — This species occurs on the coasts of the Mediterranean Sea, the Atlantic coast of north¬

western Africa (Giordani Soika 1954), and probably the Red Sea (Taiti & Ferrara 1991).

Source : MNHN, Paris

— 463 —

FIG. 2. — Tylos ponlicus: A. pereonal epimera, dorsal; B. pleon and uropods. ventral; C. pereopod 1; D. S pleopod 2 endopod.

Source : MNHN, Paris

464 —

Genus HELLERIA Ebner, 1868

Helleria brevicornis Ebner, 1868

Helleria brevicornis Ebner, 1868: 95, pi. I; SCHARFF 1894: 163; Dollfus 1899: 208; Arcan-

GELI 1914: 481; 1925: 55; 1947: 373; 1950: 146, 149; COLLINGE 1941: 522; VANDEL 1954a:

73; 1960: 110, figs 28, 47B, 49; Beron 1972a: 9; Lanza & POGGESI 1986: 121, 180.

Syspastus brevicornis ; Budde-Lund 1885: 280; Verhoeff 1926: 263.

Material examined. — Many 6 â and 9 9 collected from the following localities: Camera (near Centuri);

2 km SE of col de Ste-Lucie (between Pino and Luri); marine de Giottani; Selmacci (SW of Pietracorbara);

St-Léonard (near marine de Pietracorbara); marine de Pietracorbara; Sta Catterina de Sisco; along the Sisco river,

near Crosciano; marine de Sisco; Guado Grande (N of Nonza); Sta Maria-di-Lota (W of Miomo); Serra di Pigno

(Bastia); between Bastia and St-Florent; défilé de Lancone, between Oletta and Casatorra; near Capo a u Cavallo

(SW of Calvi); Francardo; Campana (N of Piedicroce); near Stazzona (S of Piedicroce); 2 km S of Carticasi;

forêt de Valdo-Niello, Albertacce; Gorges de la Restonica; Elbo (NW of Girolata); plage de Caspio (NW of

Porto); Porto and environs; Vecchio valley (S of Venaco); Sagone river valley. 6 km SW of Vico; U Castagno

(Ghisoni); Ghisoni; col de Vizzavona; Cascades des Anglais (Vizzavona); 1 km W of Tolla (Prunelli river valley);

near Suarella (N of Cauro); Zicavo; near Solenzara; 3 km N of col de la Vaccia (N of Aullène); between Alza

and Argiavara (col de Bavella); col de Bavella; N of Aullène; Serra-di-Scopamène (S of Aullène); Zonza; Fiu-

micicoli river valley; S of Orone (W of l’Ospedale); forêt de l'Ospedale; Golfo di Sogno (N of Porto- Vecchio);

Tizzano (SW of Sartène); rocher d'Acciaju nord (N of golfe de Sta Giulia).

Previous Records. — Ajaccio (Ebner 1868; Budde-Lund 1885; Scharff 1894); St-Florent (Budde-Lund

1885; Scharff 1894); Bastia (Scharff 1894; Dollfus 1899); Orezza. Vico, Vizzavona (Dollfus 1899); Ghisernia,

Iatemi (Arcangeli 1914); between Bastia and Cardo, Sta Lucia, San Martino, near grotte de Brando, forêt d'Aitone

(Evisa), Vizzavona, Punta di Borgo (Ajaccio) (Verhoeff 1926); Corsica, very common (Vandel 1960); grotte

de Corte (Beron 1972a); rocher d’Acciaju nord (Lanza & Poggesi 1986).

Distribution. — This species has a North-Tyrrhenian distribution. It occurs in southern Provence, Corsica,

Sardinia, some islands of the Tuscan Archipelago (Capraia, Elba, Pianosa) and Monte Massoncello (Livorno).

Budde-Lund (1885) recorded this species at Genoa and Verhoeff (1910) at Ospedaletti (Imperia), but these

records need confirmation; according to Vandel (1960) they represent occasional introductions.

Family LIGI1DAE

Genus LIGIA Fabricius, 1798

Ligia italica Fabricius, 1798

Ligia italica; Vandel 1954a: 73; 1960: 122, figs 54-56; BEAUCOURNU 1967: 562; Beron

1972a: 9.

Material examined. — Many <5 6 and 9 9 collected from the following localities: Cap Corse; Capo

Pertusato (SE of Bonifacio); île de la Giraglia; île Pietricaggiosa (Cerbicale); îlot Toro Piccolo; îlot La Folaca

(near Palombaggia); îlot de la Roscana (golfe de Pinarello); îlot Sperduto Grande and îlot Sperduto Piccolo (E

of île Cavallo).

Previous Records. — Corsica (Vandel 1960); grotte des Pigeons (Sagone) (Beaucournu 1967; Beron

1972a).

Distribution. — This species is common on the coasts of the whole Mediterranean, the Atlantic coast of

the Iberian Peninsula and Atlantic islands (Azores, Madeira and Canaries).

Source : MNHN, Paris

— 465 —

Family Trichoniscidae

Genus FINALONISCUS Brian, 1951

Finaloniscus briani Vandel, 1953

(Fig. 9)

Finaloniscus Briani Vandel, 1953a: 154, figs 1-4; 1954a: 73.

Finaloniscus briani ; Vandel 1960: 143, figs 61-64, 1968a: 356; Beron 1972a: 10, 1972b:

Material examined. — IS, grotte de Sisco (N of Bastia), leg. W. Schawaller, 8.VI11.1980.

Previous records. — Grotte de Corte (Pietracorbara) (Vandel, 1953a); alluvial plains of lower Gravona

river, E of Ajaccio (Vandel, 1960; 1968a; Beron, 1972a).

Distribution. — Known only from the above localities.

Finaloniscus franciscoloi (Brian, 1951)

(Figs 3, 9)

Material examined. — 7 8 9, 3 juvs, marine de Giottani (between Nonza and Centuri-Port), leg. S. Taiti

and A. Poggesi, 24.V1.1984; 3 9 9, Ogliastro (N of Nonza), leg. S. Taiti and A. Poggesi, 24.VI.1984; 4 â <S,

1 9, plage de Caspio (NW of Porto), leg. S. Taiti, 16.IV. 1981.

Distribution. — Finaloniscus franciscoloi was previously recorded from some caves of Savona Province

(Liguria) (Bologna & Vigna Taglianti 1985), from Tuscany (Taiti & Ferrara 1989a, 1995b) and Sicily (Caruso

et al. 1987).

Remarks

These specimens correspond in all details to the descriptions of F. franciscoloi by Brian

( 1951) and Vandel (1953b). Besides the two species present in Corsica, the genus Finaloniscus

includes also F. berberensis Vandel, 1959 from Algeria and Morocco, which, according to its

description (Vandel 1959), is probably synonymous with F. franciscoloi.

Genus NESIOTONISCUS Racovitza, 1908

Nesiotoniscus corsicus Racovitza, 1908

(Figs 4, 9)

Trichoniscus (Nesiotoniscus) corsicus Racovitza, 1908: 360, pis XVI-XVIII, figs 226-257;

Verhoeff 1943: 159.

Trichoniscus corsicus ; Jeannel & RACOVITZA 1908: 389.

Trichoniscus (Phymatoniscus) corsicus ; WOLF 1934: 73.

Nesiotoniscus corsicus; Vandel 1947: 36; Remy 1950: 25; Brian 1953: 33; Vandel 1953a:

160; Taiti & Ferrara 1995a: 313, fig. 4.

Source : MNHN, Paris

— 466 —

1 exopod; G, pleopod 2 exopod; H, pleopod 2 endopod.

Source : MNHN, Paris

— 467 —

Nesiotoniscus corsicus corsicus; Vandel 1954b: 153; 1960: 179, fig. 80; 1968a: 357; Beron

1972a: 10; 1972b: 809; DALENS 1977: 181.

Nesiotoniscus paganus; Vandel 1954a: 73.

Nesiotoniscus corsicus Racovitzai Vandel, 1954b: 154, fig. 2E ( partim: Francardo, ?

Propriano,? Sartène).

Nesiotoniscus corsicus racovitzai-, Vandel 1960: 181, fig. 81C (partim: Francardo,?

Propriano,? Sartène).

Material examined. — 4 <5S. 1 2. grotte de Pietralbello (Moltifao), leg. S. Taiti and S. Campanelli,

16.X.1982; 2 9 2, same locality, leg. P. Beron, 24.XI.1967; 1 6 , same locality, leg. S. Taiti and S. Vanni,

7.III.1994; 4 dd, 10 2 2, 3 juvs, 1.6 km from Stazzona, near road to Eaux d'Orezza, leg. S. Taiti and A.

Poggesi, 20.VI.1984; 3 dd, 2 2 2, 8 juvs, Francardo, right bank of Golo river, leg. S. Taiti and A. Poggesi.

22.V1.1984; 1 d, 3 2 2, Francardo, rive droite du Golo, à 100 m en aval du pont de la route nationale 193,

ait. 266 m, leg. P. Rémy, 22.VIII.1942 (syntypes of N. corsicus racovitzai , MNHN); 1 d, 1 2, Monte Rosso

(SE of Sartène), meadow near ilex grove, leg. S. Taiti and S. Campanelli, 20.X.1982;? 1 2, Propriano, au nord

de Sartène, sur le bord de la mer près du château, ait. 15 m, leg. P. Rémy, 18.IX.1948 (syntype of N. corsicus

racovitzai, MNHN);? I 2, Sartène, jardins à l'ouest du couvent de San Damiano, ait. 300 m, leg. P. Rémy,

16.IX.1948 (syntype of N. corsicus racovitzai, MNHN).

Previous records. — Grotte de Pietralbello (Racovitza 1908; Jeannel & Racovitza 1908; Wolf 1934;

Remy 1950; Brian 1953; Vandel 1953a, 1954b, 1960, 1968a; Beron 1972a; Dalens 1977); Francardo, right bank

of Golo river (Vandel 1947, 1953a, 1954b, I960); ? Propriano, ? Sartène (Vandel 1954b, 1960); grotte de Grottone

(Lama), grotte Roumandella (Caporalino), grotte de Cabanuli (Omessa) (Vandel 1968a; Beron 1972a).

Distribution. — Known only from Corsica.

Remarks

Vandel (1960) considers the taxa of Nesiotoniscus with a hook-like process on the male

pereopod 7 merus as subspecies of N. corsicus. According to Tabacaru (1993) and TAITI &

Ferrara (1995a) they belong to distinct species: N. corsicus and N. racovitzai Vandel, 1955

from Corsica, N. ribensis Vandel, 1948 from the Grasse region (Provence) and N. bernardi

(Vandel, 1943) from the massif of Estérel. These four species together with N. harpagonifer

Taiti & Ferrara, 1995 from Capraia Island (Tuscan Archipelago) constitute a homogeneous group

within the genus (cors/cus-group).

Nesiotoniscus corsicus is widely distributed in the granitic part of Corsica. Some populations

show small differences in the shape of the male pleopod 1 exopod (compare Fig. 4B, specimen

from “grotte de Pietralbello”, and Fig. 4E, specimen from Monte Rosso, SE of Sartène), but in

our opinion these differences are within the variability of the species.

Re-examination of the type material of N. racovitzai showed that only the specimen from

Solenzara (lectotype, see below) fits the description of the species (Vandel 1954b, fig. 2D),

while those from Francardo belong to N. corsicus. Most probably also the two syntypes females

from Sartène and Propriano should be ascribed to N. corsicus, as suggested by the presence of

this species in the same area (Monte Rosso).

Source : MNHN, Paris

— 468 —

Source : MNHN, Paris

— 469 —

Nesiotoniscus racovitzai Vandel, 1954

(Figs 5, 9)

Nesiotoniscus corsicus Racovitzai Vandel, 1954b: 153, figs 2D, 3 (partira: Solenzara).

Nesiotoniscus corsicus racovitzai; Vandel 1960: 180, fig. 81 A, B, D (partira : Solenzara).

Nesiotoniscus racovitzai; Taiti & Ferrara, 1995a: 313, fig. 4.

Material examined. — 1 6 lectotype (specimen in micropreparation, MNHN-Is4105), Solenzara, Commune

de Sari-di-Porto- Vecchio, ravin de Fontanaccia, entre la mer et la route nationale, alt. 2 m, leg. P. Remy, 3.IX. 1948.

Previous records. — Solenzara (Vandel 1954b, 1960).

Distribution. — Known only from Corsica.

Source : MNHN, Paris

— 470 —

Remarks

Nesiotoniscus racovitzai belongs to the corsicus-group on account of the hook-like process

on the male pereopod 7 merus. It is readily distinguished from all other species in the group

by the structure of the male pleopod 1 exopod which has a long narrow posterior point and a

large subrectangular lobe on the external margin.

Genus ORITONISCUS Racovitza, 1908

Oritoniscus paganus Racovitza, 1908

(Figs 6A-D, 9)

Trichoniscus (Oritoniscus) paganus Racovitza, 1908: 233, pis XV-XVI, figs 181-211; WOLF

1934: 72.

Trichoniscus paganus-, Jeannel & Racovitza 1908: 389.

Oritoniscus paganus; Remy 1950: 7, 9, 25; VANDEL 1953a: 159 ( partim : nec grotte des

Tobi Pinnuti); 1954a: 73; Beron 1972b: 809.

Oritoniscus paganus paganus; Vandel 1960: 213, fig. 100B (partim : nec grotte des Tobi

Pinnuti); 1968a: 357 (partim : nec grotte des Tobi Pinnuti); BERON 1972a: 10 (partim : nec grotte

des Tobi Pinnuti).

Material examined. — 2 6 6 , 1 9, grotte de Pietralbello (Moltifao), leg. P. Beron, 24.XI.1967; 3 9 9,

same locality, leg. S. Taiti and S. Campanelli, I6.X.1982; 1 6, 1 9, grotte de Roumandella (Caporalino), leg.

S. Taiti, 12.V.1982; 14 6 6 , 12 9 9, grotte de Sahara (Castiglione). leg. S. Taiti and S. Campanelli. 17.X.1982:

2 <5i, 1 juv., same locality, leg. S. Taiti and S. Vanni, 10.III. 1994; 1 6. grotte de Cherpinede (Lano), leg. A.

Torchia and S. Zoia, 26.V. 1982; 2 9 9, same locality, leg. S. Taiti and A. Poggesi, 22.VI. 1984.

Previous Records. — Grotte de Pietralbello (Racovitza 1908; Remy 1950; Vandel 1953a, 1960, 1968a;

Beron 1972a); grotte de Sisco, grotte de Brando (Remy 1950; Vandel 1968a; Beron 1972a); grotte de Corte

(Pietracorbara) (Vandel 1968a; Beron 1972a); grotte de Cherpinede, grotte de Sulane (Furiani), grotte de Cabanuli

(Omessa), grotte de Sahara, grotte de Valletto (Santo-Pietro-di-Venaco), grotte de Leccia-Torta (Castiglione), grotte

de Gudrone (Sorio), grotte des Paladini (Solaro) (Vandel 1953a, 1960, 1968a; Beron 1972a); grotte Tavona di

Nuaia No. 2 (Conca) (Vandel 1960, 1968a; Beron 1972a); grotte Manuel-Ange (Lozzi), grotte Chevalier (Lano)

(Vandel 1968a; Beron 1972a); Canella bridge (S of Solenzara), banks of lower Gravona river (E of Ajaccio)

(Vandel 1960, 1968a).

Distribution. — Oritoniscus paganus is endemic to Corsica.

Remarks

On the basis of the presence or absence of the eye, Vandel (1953a, 1960) divides the

species into two subspecies: the nominal one, blind, spread throughout Corsica, and the subspecies

ocellatus, limited to the northern peninsula. In the caves of Brando, Sisco and Corte, both sub¬

species seemed to be present (Vandel 1968a).

The material examined here revealed the following important facts:

1. In the caves of Brando and Sisco (we have not studied material from the grotte de Corte)

only the form ocellatus has been collected, and most probably this is the only species of the

genus present in the whole Cap Corse peninsula;

Source : MNHN, Paris

— 471 —

Source : MNHN, Paris

— 472 —

2. Vandel (1960) considers the specimens described by RACOVITZA (1908) to be immature

stages of O. paganus paganus. Vandel’s opinion is certainly incorrect, since all the male speci¬

mens examined from the caves of Pietralbello (type locality), Sahara and Roumandella, up to

4 mm long and certainly adult, have pleopod 1 (Fig. 6A) as illustrated by Racovitza (1908)

and Vandel (1960, fig. 100B);

3. Vandel (1953a) identifies O. paganus from the grotte des Tobi Pinnuti on the basis of

a female and two juveniles. Some specimens with no trace of an eye, collected from the same

cave, possess male characters (<J 5.5 mm long) which differ from those of both paganus and

ocellatus (see below).

Due to the consistent differences in male characters between paganus and ocellatus, we

consider these two taxa to be distinct species. This conclusion has also been confirmed by a

genetic analysis (COBOLLI SBORDONI et al. 1995). Most probably also the specimens from the

grotte des Tobi Pinnuti belong to another distinct species, but we prefer not to nominate it because

of the small amount of material examined.

The exact distribution within Corsica of O. paganus is not clear: it is certainly present in

the four caves from which we examined specimens, while the records by Vandel need confir¬

mation after re-examination of the material studied by the French author.

Oritoniscus ocellatus Vandel, 1953

(Figs 6E, F, 7A, B, 9)

Oritoniscus paganus var. ocellata Vandel, 1953a: 159.

Oritoniscus paganus ocellatus; Vandel 1960: 216, figs 99-100A, C; 1968a: 360; Beron

1972a: 10.

Material examined. — 3 <J<J, 6 2 9, grotte de Sisco, leg. B. Lanza, 11.1V.1977; 8 â <?, 13 2 2, same

locality, leg. W. Schawaller, 8.VIII. 1980; 6 ââ, 19 9 2, same locality, leg. S. Taiti, 19.IV.1981; 3 d<J, 8 2 9,

same locality, leg. S. Taiti, 21.V.1982; 14 âô, 11 2 9, same locality, leg. S. Taiti and S. Campanelli, 13.X.1982:

4 66, 11 99, same locality, leg. S. Taiti and A. Poggesi, 27.VI.1984; 1 6, 1 2, same locality, leg. S. Taiti

and S. Vanni, 6.III.1994; 2 6 6, 3 22, grotte de Brando, leg. S. Taiti and S. Campanelli, 13.X.1982; 2 22,

same locality, leg. P. Magrini, 1.XI.1983; 1 6, 3 juvs. Camera (near Centuri), leg. S. Taiti, 18.IV.1981; 2 6 6,

1 2, near Camera, ilex grove, leg. S. Taiti and A. Poggesi, 19.VI.1984.

Previous records. — Grotte de Corte, grotte de Sisco, grotte de Brando (Vandel 1953a, 1960, 1968a;

Beron 1972a); Pino (Vandel 1960, 1968a).

Distribution. — This species occurs in the Cap Corse peninsula. A closely related form is present on the

islands of Montecristo and Giannutri in the Tuscan Archipelago (Taiti & Ferrara 1995b).

Remarks

Oritoniscus ocellatus differs from O. paganus in the presence of the eye, consisting of a

single ommatidium, and the male modifications. In particular, the pereopod 5 merus shows a

distinct setose swelling at the base (absent in O. paganus ); the pereopod 7 basis is more swollen;

the pleopod 1 exopod is clearly longer than the endopod (vice versa in O. paganus), with the

basal lobe narrow and rounded (conspicuous and rectangular in O. paganus), the medial margin

straight with many small spines in the distal part (it shows a step and no spines in O. paganus),

and the terminal spine stouter with no distal setae.

Source : MNHN, Paris

— 473 —

Fig. 7. — Oritoniscus ocellaius. 6 from grotte de Sisco: A, pereopod 6; B, pereopod 7. Oritoniscus sp., 6 from grotte des Tobi

Pinnuti: C, pleopod 1; D, pereopod 5; E. pereopod 6; F, pereopod 7.

Source : MNHN, Paris

— 474 —

Oritoniscus sp.

(Figs 7C-F, 9)

Oritoniscus paganus; Vandel 1953a: 159 ( partira : grotte des Tobi Pinnuti).

Oritoniscus paganus paganus; VANDEL 1960: 216 (partira : grotte des Tobi Pinnuti); 1968a:

359 (partim: grotte des Tobi Pinnuti); BERON 1972a: 10 ( partira : grotte des Tobi Pinnuti).

Material examined. — 1 <J, 3 2 2,5 juvs, grotte des Tobi Pinnuti (Sorio), leg. S. Taiti, 15.X.1982.

Previous records. — Grotte des Tobi Pinnuti (Vandel 1953a, 1960, 1968a; Beron 1972a).

Remarks

These specimens show clear differences in the male characters, particularly the shape of

pleopod 1 (Fig. 7C), from both 0. paganus and, to a lesser extent, 0. ocellatus. They probably

belong to a distinct species but more material is needed for a certain identification.

Oritoniscus punctatus n.sp.

(Figs 8, 9)

Oritoniscus punctatus (nomen nudum); Ferrara & Taiti 1984: 417.

Material examined. — 1 <î holotype (MZUF), 1 $ juv., 1 2 paratype (MNHN-Is4103), 1 2 paratype

(MV), 1 2 juv. paratype (MZUF), Punta de la Parata (W of Ajaccio), under big stones on rocky cliff near sea,

leg. S. Taiti, 14.V. 1982.

Etymology. — L. punctatus = having a point. The name refers to the distinct triangular point on the medial

margin of the basal part of the male pleopod I exopod.

Description

Maximum length: 66 and $9,4 mm.

Colourless body. Eye absent. Dorsum with distinct granulations, each bearing a large scale-

spine on top. Antennule of three articles with 6-7 aesthetascs at the apex. Antenna with flagellum

of five articles, the second of which with a row of 5-6 aesthetascs.

Male

Pereopods 1-6 without distinct modifications. Pereopod 7 basis with a deep depression on

the distal part of the sternal margin. Pleopod 1 exopod with a basal part having a large concavity

and a rounded lobe on the external margin, a distinct triangular point on the medial margin,

and ending with a very long and strong distal spine; endopod with a rectangular basal part on

which a long glabrous flagellum is inserted, not surpassing the tip of the exopod. Pleopod 2

exopod about three times as wide as long, with short posterior point apically rounded; endopod

of two articles, the distal one pointed and slightly longer than the basal one.

Source : MNHN, Paris

— 475 —

Source : MNHN, Paris

— 476 —

Remarks

The new species shows close affinities with O. paganus and O. ocellatus. It is easily dis¬

tinguishable from both in having dorsal granulations and in the modifications of the male

pereopod 7 and pleopod 1, particularly the shape of the exopod with a distinct acute point on

the medial margin. This last character is absent in all the species of the paganus- group.

Genus TRICHONISCUS Brandt, 1833

Trichoniscus pusillus provisorius Racovitza, 1908

Trichoniscus pusillus provisorius ; Vandel 1954a: 73; 1960: 321, figs 30, 150.2.

Material examined. — Many 6 6 and 2 2 collected from the following localities: Barcaggio (Cap Corse);

Minervio (S of Pino); Carbonacce (S of Luri); marine de Giottani (between Nonza and Centuri-Port); Ponticello

(near Pietracorbara); marine de Pietracorbara; Nonza; near grotte de Brando; Lavasina (S of Brando); col de

Teghime (W of Bastia); Oletta; Bevinco river valley, below col de S. Stefano; Urtaca; 2 km SW of Ponte Novu

(Golo river valley); Asco river valley, S of Moltifao; 3 km S of Ponte Leccia, left bank of Golo river; forêt de

Bonifato (S of Calvi); Punta di Chiarsgioli (Monte San Petrone); Francardo; near Lano (NW of Corte); near

Stazzona; Corte; near Albertacce (forêt de Valdo-Niello); col de Vergio (NE of Evisa); Porto; SSE of Evisa;

Tavignano river valley, between Volta and Scandulaie (NW of Aléria); col de Vizzavona; col de Verde (S of

Ghisoni); 1 km E of Tolla (Prunelli river valley); between Cauro and Bocca San Giorgio (E of Ajaccio); 3 km

N of col de la Vaccia (N of Aullène); col de Bavella; 2 km NW of Aullène; Casalabriva (N of Propriano); below

chapelle Pianelli, between Casalabriva and Olmeto; Golfo di Sogno (N of Porto- Vecchio); Monte Rosso (SE of

Sartène); Orasi (S of Sartène); Tizzano (SW of Sartène).

Previous records. — All Corsica (Vandel 1954a, 1960).

Distribution. — Trichoniscus pusillus provisorius is recorded from France, Great Britain, Ireland. Denmark,

Poland, Spain, Switzerland, Italy, Turkey. Lebanon and Algeria. It has been introduced also to the Azores and

Newfoundland.

Trichoniscus pygmaeus Sars, 1899

Trichoniscus pygmaeus; VANDEL 1954a: 73; 1960: 325, figs 153-154.

Material examined. — Id, between Pietrosella and Fogolina, SE of Porticcio (Ajaccio), ilex and arbutus

wood, leg. S. Taiti and S. Campanelli, 21.X.1982.

Previous Records. — Corsica (Vandel 1954a, 1960).

Distribution. — This is a widespread species known from Europe (except Portugal and the Balkans).

Morocco, Newfoundland and the USA.

Trichoniscus fragilis Racovitza, 1908

(Fig. 9)

Trichoniscus fragilis ; Vandel 1954a: 73; 1960: 333, fig. 158; 1968a: 356; Beron

1972a: 11.

Source : MNHN, Paris

— All —

Fig. 9. — Distribution in Corsica of Trichoniscidae species (except Trichoniscus pusillus provisorius, T. pygmaeus and Haplo-

phihalmus danicus ): 1, Finaloniscus brianr, 2, F. franciscoloi; 3, Nesioloniscus corsicus; 4, N. racovitzai; 5, Oritoniscus

paganus', 6, O. ocellatus', 7, Oritoniscus sp.; 8, O. punciatus; 9, Trichoniscus fragilis ; 10, T. halophilus ; 11, T. pedronensis;

12, Cyrnoniscus remyi\ 13, Carloniscus dol/usv, 14, Buddelundiella cataractae.

Source : MNHN, Paris

— 478 —

Material examined. — 8 6 6, 8 9 9, entrance of the grotte de Sisco, leg. S. Taiti, 19.IV.1981; 4 6 6,

8 9 9, same locality, leg. S. Taiti and S. Vanni, 8.III.1994; 3 6 6, 3 9 9, Phare de Fomali (W of St-Florent),

leg. S. Taiti, 17.VII.1978.

Previous records. — Sisco, Solenzara, Porto-Vecchio, Bonifacio (Vandel 1960); grotte de St-Barthélémy

(Bonifacio) (Vandel 1968a; Beron 1972a); grotte de Sisco (Beron 1972a).

Distribution. — Atlantic and Mediterranean coasts of France, Corsica, Tuscany, southern Italy, Crete and

Algeria.

Trichoniscus halophilus Vandel, 1951

(Fig. 9)

Material examined. — 1 6, Barcaggio, Cap Corse, leg. S. Taiti and S. Campanelli, 14.X.1982; 2 6 6,

7 9 9, same locality, leg. S. Taiti and A. Poggesi, 19.V1.1984; 2 9 9, Puma de la Parata (W of Ajaccio), leg.

S. Taiti and S. Vanni, 8.III. 1994; 1 6,6 9 9, Punta di e Botte di Tizzano (SW of Sartène), small stream near

sea, leg. S. Taiti, 15.V.1982.

Distribution. — This species is known from some French Mediterranean islands, Corsica, Sardinia, Tuscany,

Sicily, Panteileria Island, Lampedusa Island and Malta. In Morocco it has been recorded from a cave near Taza

(Vandel 1955).

Trichoniscus pedronensis Vandel, 1947

(Fig. 9)

Trichoniscus pedronensis Vandel, 1947: 49, figs 13-14; 1954a: 73; 1960: 332, fig. 157.

Material examined. — 2 66,2 9 9 (syntypes), Monte San Pedrone [= San Petrone], 1600 m, versant

est, dans la hêtraie sous les pierres, leg. P. Remy, 26.VIII. 1942 (MNHN).

Previous records. — Monte San Petrone (Vandel 1947, 1960).

Distribution. — Known only from the type locality.

Remarks

Investigations carried out in the beech woods of Monte San Petrone revealed no specimens

that could be referred to this species. We re-examined the type material studied by Vandel,

but unfortunately pereopods 7 and pleopods 1 and 2 of both males are preserved in micropre¬

parations which have deteriorated with age, so that we are not able to provide new illustrations

of the sexual characters of this species.

Genus HAPLOPHTHALMUS Schoebl, 1861

Haplophthalmus danicus Budde-Lund, 1885

Haplophthalmus danicus; VANDEL 1954a: 73; 1960: 362, fig. 172.

Material examined. — Many 6 6 and 9 9 collected from the following localities: near marine de Giottani

and near grotte de Brando (Cap Corse peninsula); Folelli, along Fium Alto; between Stazzona and the Eaux

Source : MNHN, Paris

— 479 —

d’Orezza; 3 km S of Ponte Leccia, along left bank of Golo river; Francardo, right bank of Golo river, near a

spring.

Previous records. — Corsica, very common (Vandel 1960).

Distribution. — This species is known from all Europe, northern Africa, Asia Minor, St Helena Island,

most of North America, Hawaii and Japan.

Genus CYRNONISCUS Vandel, 1953

Cyrnoniscus remyi Vandel, 1953

(Fig. 9)

Cyrnoniscus remyi Vandel, 1953a: 163, figs 5-6; 1954a: 73; 1960: 394, figs 190-191; 1968a:

360; Beron 1972a: 11; 1972b: 809.

Material examined. — 4 6 6,6 9 9, grotte de Corte (Pietracorbara), leg. P. Beron, 27.XI.1967; 1 9,

grotte de Sisco, leg. S. Taiti and S. Campanelli, 13.X.1982; 2 9 9, same locality, leg. W. Schawaller, 8.VIII. 1980;

1 9, same locality, leg. S. Taiti and A. Poggesi, 27.VI. 1984; 2 66, same locality, leg. S. Taiti and S. Vanni,

6.III.1994; 2 9 9, same locality, leg. S. Taiti, S. Vanni and A. M. Nistri, 3.IV.1995; 1 9, Nonza, near cemetery,

leg. S. Taiti and A. Poggesi, 24.VI.1984; 1 6, 1 9, grotte des Tobi Pinnuti (Sorio), leg. S. Taiti, 15.X.1982; 14

9 9,1 juv., 1.6 km from Stazzona, near road to the Eaux d’Orezza, leg. S. Taiti and A. Poggesi, 20.VI. 1984.

Previous records. — Grotte de Corte (Pietracorbara), grotte de Gudrone (Sorio) (Vandel 1953a, 1960,

1968a; Beron 1972a); grotte de Sisco, grotte de Ostriago (Pietracorbara) (Vandel 1968a; Beron 1972a).

Distribution. — Cyrnoniscus remyi, the sole species in the genus, is endemic to Corsica.

Genus CARLONISCUS Verhoeff, 1936

Carloniscus dollfusi (Carl, 1908)

(Fig. 9)

Carloniscus Dollfusi ; Vandel 1954a: 73.

Carloniscus dollfusi; Vandel 1960: 388, figs 187A, 189.

Records. — Francardo, right bank of Golo river (Vandel 1960).

Distribution. — This species occurs in the French Alps and Corsica.

Remarks

Vandel’ s record of this species on the right bank of the Golo river at Francardo needs to

be confirmed. It was based on a single female specimen and several recent investigations in the

same locality aimed at collecting this species have not been successful. It could be a matter of

an occasional introduction: in fact, another species {Haplophthalmus danicus), certainly intro¬

duced, is abundant in that area.

Source : MNHN, Paris

— 480 —

Genus BUDDELUNDIELLA Silvestri, 1897

Buddelundiella cataractae Verhoeff, 1930

(Fig. 9)

Buddelundiella borgensis; Vandel 1954a: 73.

Buddelundiella cataractae ; Vandel 1960: 410, figs 198-201.

Records. — Bonifacio, Gurgazo (Vandel 1960).

Distribution. — This species has a widespread distribution in Europe.

Undetermined family

Genus BUCHNERILLO Verhoeff, 1942

Buchnerillo litoralis Verhoeff, 1942

Lereboulletia littoralis; Vandel 1945: 104, figs V-XVI.

Buchnerillo littoralis ; Vandel 1954a: 73; 1960: 400, figs 195-197.

Material examined. — 1 6, 2 9 9, 1 juv., Barcaggio (Cap Corse), leg. S. Taiti and S. Campanelli,

14.X.1982; 1 rf, 9 9 9, same locality, leg. S. Taiti and A. Poggesi, 19.VI.1984; 1 6, 7 9 9, 2 juvs, l’île-Rousse,

leg. S. Taiti, 16.VII.1978; 1 6, Punta de la Parata (W of Ajaccio), leg. S. Taiti, 14.V.1982.

Previous records. — Marine de Sisco (Vandel 1945, 1960).

Distribution. — It is known from the coasts of Madeira Island, Provence, Corsica, Sardinia, Tuscany,

Ischia Island, Aegadean islands, Sicily and Malta.

Family Stenoniscidae

Genus STENONISCUS Aubert & Dollfus, 1890

Stenoniscus pleonalis Aubert & Dollfus, 1890

Stenoniscus pleonalis', Vandel 1944b: 24; 1954a: 73.

Stenoniscus pleonalis pleonalis', Vandel 1962: 425, figs 206, 208-210; BERON 1972a: 11.

Records. — Lavasina (S of Brando), marine de Sisco (Vandel 1944b, 1962); grotte de Sdragonato (Boni¬

facio) (Beron 1972a).

Distribution. — This species occurs on the northern coasts of the Mediterranean, from France to Greece,

and Madeira Island. Vandel (1968b) reported this species in the Galapagos, but this record needs confirmation.

Remarks

Vandel (1962) considers S. carinatus Silvestri, 1897 to be a subspecies of S. pleonalis.

However, it is certain that the two taxa are distinct species (Caruso 1976; Taiti & Ferrara

1980). Without re-examination, it is not possible to define to which of the two species belong

Source : MNHN, Paris

— 481 —

the specimens from Corsica recorded in the literature as S. pleonalis. However, it is very probable

that both species are present in Corsica, as in the Tuscan Archipelago.

Stenoniscus carinatus Silvestri, 1897

Stenoniscus carinatus ; Lanza & POGGESI 1986: 121, 178.

Material examined. — Many 6 6 and 9 9, Puma de la Parata (W of Ajaccio), leg. S. Taiti and S. Vanni,

8.III.1994; 3 <3 <3, 17 9 9, Capo Pertusato (SE of Bonifacio), leg. S. Taiti and S. Campanelli, 19.X.1982; 2 6 6,

île Pietricaggiosa (Cerbicale), leg. S. Taiti and S. Campanelli, 17.V. 1982.

Previous Records. — Phare de Pertusato and île Pietricaggiosa (Lanza & Poggesi 1986).

Distribution. — This species is known from Tenerife (Canaries), Portugal, Corsica, Sardinia, the Tuscan

Archipelago, Sicily and surrounding islands (Ustica, Pantelleria, Aegadean and Pelagean islands), Malta and the

coasts of the northern Adriatic. Most probably the records of S. pleonalis by Schultz (1972) from Bermuda

islands, and Paoletti & Stinner (1989) from Florida, belong to S. carinatus ; this species is certainly present in

Florida (many 6 6 and 9 9, Key West and Long Key, leg. S. Taiti, 15-16.XII.1984).

Family SCYPHACIDAE

Genus ARMADILLONISCUS Uljanin, 1875

Armadilloniscus candidus Budde-Lund, 1885

(Fig. 10A, B)

Armadilloniscus candidus; VANDEL 1954a: 74; 1962: 472, figs 234-235.

Material examined. — 1 <3,2 9 9, Tollare (Cap Corse), leg. S. Taiti, 18.IV.1981; 1 <3, 1 9, l’île-Rousse,

leg. S. Taiti, 16.VII.1978; 7 <3<3, 32 9 9, Puma de la Parata (W of Ajaccio), leg. S. Taiti, 14.V.1982; 1 3,1 9,

Fautea (between Solenzara and Porto- Vecchio), leg. S. Taiti, 17.V.1982.

Previous records. — Sisco (Vandel 1962).

Distribution. — This species occurs on the coasts of southern France, Corsica, Sardinia, Tuscany, Algeria,

Tunisia, Malta and the Azores.

Armadilloniscus ellipticus (Harger, 1878)

(Fig. IOC, D)

Material examined. — 24 6 6, 37 9 9, Barcaggio (Cap Corse), leg. S. Taiti and S. Campanelli, 14.X.1982;

3 66,2 9 9, same locality, leg. S. Taiti and A. Poggesi, 19.VI.1984.

Distribution. — Armadilloniscus ellipticus is known from the Atlantic coasts of North America, the coasts

of the Mediterranean, the Azores, Madeira Island, Madagascar, Malaysia, Hong Kong, Korea and Hawaiian islands

(Taiti & Ferrara 1989b; Garthwaite et al. 1992).

Source : MNHN, Paris

— 482 —

Fig. 10. — Armadilloniscus candidus, 6 : A. pleopod 1; B, pleopod 2. ArmadiHoniscus ellipticus , <5 : C, pleopod 1; D, pleopod 2.

Source : MNHN, Paris

— 483 —

Remarks

The synonymy of Armadilloniscus litoralis Budde-Lund, 1885 with A. elliplicus has been

proved recently by Garthwaite et al. (1992) on the basis of both morphological and genetic

data.

Family Halophilosciidae

Genus HALOPHILOSCIA Verhoeff, 1908

Halophiloscia couchii (Kinahan, 1858)

Philoscia Couchii; Dollfus 1897: 72; Arcangeli 1925: 50.

Philoscia Couchi ; DOLLFUS 1899: 207.

Halophiloscia Couchi ; Vandel 1954a: 74.

Halophiloscia (Halophiloscia) couchi ; VANDEL 1962: 477, figs 237-238.

Material examined. — Many 6 6 and 2 2 collected from the following localities: Barcaggio and Tollare

(Cap Corse); marine de Sisco: Phare de Fornali (W of St-Florent); anse de Faggiola (désert des Agriates); Punta

de la Parata (W of Ajaccio); Golfo di Sogno (N of Porto- Vecchio).

Previous records. — Bastia and Porto-Vecchio (Dollfus 1897, 1899); Etang de Biguglia (Dollfus 1899).

Distribution. — This widespread species populates all the coasts of the Mediterranean, the Atlantic coasts

of Europe and Africa as far south as Dakar, the Azores, Canaries, Madeira and Cape Verde. It has also been

introduced to Virginia, Bermuda, Argentina and Western Australia.

Halophiloscia hirsuta Verhoeff, 1928

Halophiloscia hirsuta; Vandel 1954a: 74; Lanza & Poggesi 1986: 120, 176, 178, 179.

Halophiloscia (Halophiloscia) hirsuta; Vandel 1962: 483, figs 241-242; Beron 1972a: 12.

Halophiloscia sp.; Beaucoumu, 1967: 562.

Material examined. — Many 6 <$ and 2 2 collected from the following localities: Tollare (Cap Corse);

marine de Giottani (between Nonza and Centuri-Port); entrance of the grotte de Sisco; anse de Faggiola (désert

des Agriates); Punta de la Parata (W of Ajaccio); Tizzano and environs (SW of Sartène); Capo Pertusato (SE

of Bonifacio); île de la Giraglia; îlot Intermediaire (Finocchiarola, E of Cap Corse); île Pjetricaggiosa (Cerbicale);

îlot La Folaca (near Palombaggia, SE of Porto-Vecchio); îlot Sperduto Piccolo (E of île Cavallo); îlot Cala di

u Ghiunco (île Lavezzi).

Previous records. — Grotte des Pigeons (Sagone), grotte de Saragonato (Bonifacio) (Beron 1972a); Corsica

(Vandel 1954a, 1962); îlot Intermediaire (Finocchiarola), île Pietricaggiosa (Cerbicale), îlot Toro Grande, îlot La

Folaca (Lanza & Poggesi 1986).

Distribution. — This species is known from the north-central coasts of the Mediterranean, from France

to Greece.

Halophiloscia ischiana Verhoeff, 1933

Material examined. — 3 ââ, 6 2 2, Golfo di Sogno (N of Porto-Vecchio), leg. S. Taiti, 13.IV.1981.

Source : MNHN, Paris

— 484 —

Distribution. — This species was previously recorded from southern France, Sardinia, Tuscany (Giglio

Island and Monte Argentario promontory), Campania (Ischia Island and Sorrento), and the Balearic islands

(Menorca Island).

Halophiloscia tyrrhena Verhoeff, 1928

Material examined. — 12, Barcaggio (Cap Corse), leg. S. Taiti and S. Campanelli, 14.X.1982; 1 d,

3 2 2, marine de Sisco, leg. S. Taiti and S. Campanelli, 12.X.1982; 3 <3<J, 3 2 2, Puma de la Parata (W of

Ajaccio), leg. S. Taiti and S. Vanni, 8.III.1994.

Distribution. — It is known from the Mediterranean coasts of France, Corsica, Sardinia, Liguria, Tuscany

(Gorgona and Elba islands, promontories of Monte Massoncello, Uccellina and Monte Argentario).

Genus STENOPHILOSCIA Verhoeff, 1908

Stenophiloscia zosterae Verhoeff, 1928

Material examined. — Many â <5 and 2 2 collected from the following localities: Barcaggio (Cap Corse);

marine de Pietracorbara; marine de Sisco; l’île-Rousse; Calzarello and environs (E of Ghisonaccia).

Distribution. — Known from southern France, Corsica, Sardinia, Liguria, Tuscany, Sicily and surrounding

islands (Ustica, Aeolian islands, Pantelleria, Pelagean islands), Malta, Dalmatia and Greece.

Family Philosciidae

Genus CHAETOPHILOSCIA Verhoeff, 1908

Chaetophiloscia elongata (Dollfus, 1884)

Philoscia pulchella Budde-Lund, 1885: 215.

Philoscia elongata; Scharff 1894: 163; Dollfus 1897: 95; 1899: 198; Arcangeli, 1925: 48.

Chaetophiloscia elongata ; Vandel 1954a: 74.

Material examined. — Many 6 â and 2 2 collected from the following localities: Barcaggio and Tollare

(Cap Corse); Centuri-Port; near Meria; marine de Pietracorbara; marine de Giottani (between Nonza and

Centuri-Port); Sta Catterina de Sisco; between Ogliastro and Lainosa (N of Nonza); Nonza; near Crosciano, along

Sisco river; near Erbalunga; near grotte de Brando; col de Teghime (W of Bastia); désert des Agriates; plage

d'Ostriconi (NE of Lozari); between Casatorra and Oletta; Asco river valley; Folelli, along Fium Alto; Prunete

(E of Cervione); plage de Caspio (NW of Porto); Sollacaro (N of Propriano); near Propriano; Fautea (between

Solenzara and Porto Vecchio); Lovo Santo, swamp at mouth of Cavo river; source de Caldane (NE of Sartène);

Golfo di Sogno (N of Porto-Vecchio); Monte Rosso (SE of Sartène); Tizzano and baie d’Avena (SW of Sartène);

golfe de Sant’Amanza (NE of Bonifacio).

Previous records. — Ajaccio, Bastia (Scharff 1894, Dollfus 1899); Sartène, Bonifacio (Dollfus 1899).

Distribution. — This species is widespread in the lands encompassing the Mediterranean Sea.

Chaetophiloscia sicula Verhoeff, 1908

Chaetophiloscia sicula; Vandel 1954a: 74; 1962: 496, fig. 247.

Source : MNHN, Paris

— 485 —

Material examined. — Many d d and 9 9 collected from the following localities: Barcaggio (Cap Corse);

Centuri-Port; between Macinaggio and Meria; Guado Grande (N of Nonza); near grotte de Brando; Cardo (W

of Bastia); near San Martino-di-Lota; Bastia; défilé de Lancone (W of Casatorra); Asco river valley; Francardo

(N of Corte); near Propriano; île de la Giraglia.

Previous records. — Corsica (Vandel 1954a, 1962).

Distribution. — Known from Menorca Island, southern France, Corsica, all the Italian peninsula, the Tuscan

Archipelago (islands of Gorgona, Elba, Palmaiola, Giglio and Giannutri), Ischia Island, Sicily and Greece.

Chaetophiloscia cellaria (Dollfus, 1884)

Philoscia cellaria ; DOLLFUS 1897: 91.

Chaetophiloscia cellaria ; REMY 1950: 9; Vandel 1953a: 163; 1954a: 74; 1962: 499,

fig. 248; Beron 1972a: 12; SCHMALFUSS 1990: 170.

Material examined. — Many d d and $ 9 collected from the following localities: between Macinaggio

and Meria; marine de Giottani (between Nonza and Centuri-Port); grotte de Sisco; grotte de Brando and environs;

Miomo; Cardo (W of Bastia); île de la Giraglia.

Previous records. — Bonifacio (caves) (Dollfus 1897, Vandel 1962); grotte de Brando (Remy 1950;

Vandel 1953a, 1962; Beron 1972a); grotte de St François (Bonifacio) (Vandel 1953a; Beron 1972a); grotte de

Sisco (Vandel 1962; Beron 1972a); cave N of Bastia (Schmalfuss 1990).

Distribution. — Widespread in the northern Mediterranean region, from Spain to Lebanon (Schmalfuss

1991).

Genus CTENOSCIA Verhoeff, 1928

Ctenoscia dorsalis Verhoeff, 1928

Material examined. — Many dd and 9 9, Tollare (Cap Corse), leg. S. Taiti, 18.IV.1981; 1 d, 2 9 9,

same locality, leg. S. Taiti and A. Poggesi, 19.VI.1984.

Distribution. — This species is known from western Spain, Corsica, Sardinia, Liguria, Sicily and sur¬

rounding islands (Aeolian and Aegadean islands, Pantelleria and Pelagean islands), and Malta.

Ctenoscia minima (Dollfus, 1892)

Ctenoscia minima; Beron, 1972a: 11.

Records. — Grotte de Sisco (Beron 1972a).

Distribution. — Portugal, Spain, Menorca Island, La Galite Archipelago and Corsica (?).

Remarks

Ctenoscia minima differs from C. dorsalis in having the male pleopod 1 endopod bent

outwards instead of straight. The record of this species by Beron (1972a) is doubtful, since it

is based on a single specimen with no indication of the sex. Most probably it refers to C. dorsalis.

Source : MNHN, Paris

— 486 —

Genus PHILOSCIA Latreille, 1804

Philoscia affinis Verhoeff, 1908

Philoscia muscorum; Scharff 1894: 163; Dollfus 1899: 190; Arcangeli 1925: 47; VER¬

HOEFF 1926: 263.

Philoscia affinis ; Vandel 1954a: 74; 1962: 511, figs 254C, 255; Beron 1972a: 12.

Material examined. — Many <5 6 and $ 9 collected from the following localities: Barcaggio and Tollare

(Cap Corse): Centuri-Port; Camera (near Centuri); Rogliano; between Macinaggio and Meria; Morsiglia; 2 km

SE of col de Ste-Lucie (between Pino and Luri); 2 km E of Luri; Carbonacce (S of Luri); marine de Pietracorbara;

Ponticello (near Pietracorbara); Selmacci (SW of Pietracorbara); St-Léonard (W of marine de Pietracorbara);

Ogliastro (N of Nonza); Guado Grande (N of Nonza); near Crosciano along Sisco river; marine de Sisco; Nonza;

near Erbalunga; near grotte de Brando; Sta Maria-di-Lota; Cardo (W of Bastia); désert des Agriates; Casta (SW

of St-Florent); Olmeta-di-Tuda; col de S. Stefano (S of Olmeta-di-Tuda); défilé de Lancone; 1 km N of San-

Gavino-di-Tenda; plage d’Ostriconi (NE of Lozari); Urtaca; near Murato; 1 km W of Altiani (SE of Speloncato);

near grotte de Pietralbello (Moltifao); Asco river valley; 2 km SW of Ponte Nuovo along Golo river; 3 km S

of Ponte Leccia; forêt de Bonifato (SE of Calvi); Francardo (N of Corte); Lano (NW of Corte); Punta di Chiarsgioli

(Monte San Petrone); near the Eaux d’Orezza; near Stazzona; 2 km S of Carticasi; Prunete (E of Cervione);

2 km S of Corte; SE of Porto; near Cargèse; col de Vizzavona; 7 km SSW of Ghisoni; 1 km E of Tolla (Prunelli

river valley); between Alza and Argiavara (col de Bavella); col de Bavella: between col de Bavella and Punta

Velaco; 2 km N of Aullène; between col de St-Eustache and col de Tana (W of Aullène); near Zonza; below

chapelle Pianelli, between Casalabriva and Olmeto; Taglio Rosso (Ste-Lucie-de-Porto- Vecchio); Lovo Santo,

swamp at mouth of Cavo river; near Propriano; 2 km W of Orone (W de l'Ospedale); source de Caldane (NE

of Sartène); Golfo di Sogno (N of Porto- Vecchio); Tizzano and baie d'Avena (SW of Sartène); golfe de

Sant'Amanza (NE of Bonifacio); Bonifacio; Capo Pertusato (SE of Bonifacio).

Previous records. — Ajaccio (Scharff 1894); Bastia (Scharff 1894, Dollfus 1899, Arcangeli 1925);

St Florent (Scharff 1894); Corsica, very common (Vandel 1962); grotte de Grottone (Lama) (Beron 1972a).

Distribution. — This widespread species has been recorded from Italy, Slovenia, Croatia, Germany, France,

northern Spain, Algeria, and perhaps Tunisia (Vandel 1962). It occurs on all the Tyrrhenian islands and Malta.

Remarks

According to Vandel (1954a, 1962) the records from Corsica of P. muscorum (Scopoli,

1763) most probably refer to P. affinis. Also in our opinion this is the only species of the genus

present in Corsica, as shown by the abundant material examined. Thus, P. muscorum is considered

not to belong to the Corsican fauna.

Genus TIROLOSCIA Verhoeff, 1926

Tiroloscia Corsica Corsica (Dollfus, 1888)

Philoscia Corsica Dollfus, 1888: 10, figs 1-14; SCHARFF 1894: 163; DOLLFUS 1897: 94;

1899: 207; ARCANGELI 1925: 47.

Philoscia (Tiroloscia) Corsica; ARCANGELI 1950: 130; Vandel 1954a: 74.

Tiroloscia Corsica; Vandel 1962: 518, fig. 256.

Material examined. — Many 6 6 and 9 9 collected from the following localities: Barcaggio and Tollare

(Cap Corse); marine de Giottani (between Nonza and Centuri-Port); Sta Catterina, N of marine de Sisco; col de

Source : MNHN, Paris

— 487 —

S. Stefano (S of Olmeta-di-Tuda); entrance of the grotte de Grottone, near Lama; 1 km W of Altiani (SE of

Speloncato); between Barchetta and Casamozza on the left bank of Golo river; 2 km SW of Ponte Nuovo (Golo

river valley); Asco river valley, NW of Ponte Leccia; S of Moltifao (Asco river valley); forêt de Bonifato; near

grotte de Sahara (near Castiglione); Punta di Chiarsgioli (Monte San Petrone); Francardo; Albertacce (Golo river

valley); col de Vergio (NE of Evisa); forêt de Valdo-Niello; forêt de la Restonica; near Porto; SSE of Evisa,

along Porto river; near col de Sévi (SE of Evisa); Venaco; Lac de Melo (Restonica river valley); between Volta

and Scandulaie along a tributary stream of Tavignano river (NW of Aléria); 6 km SW of Vico, Sagone river

valley; near Ghisoni; col de Vizzavona; Capannelle (SE of Vizzavona); Bocognano; col de Verde (S of Ghisoni);

1 km E of Tolla (Prunelli river valley); Palneca (Taravo river valley); S of col de Marcuccio (between Cauro

and Bastelica); Monte Incudine; between Pietrosella and Fogolina (SE of Porticcio); 3 km N of col de la Vaccia

(N of Aullène); col de Bavella; Punta Velaco (Bavella); between col de St-Eustache and col de Tana (W of

Aullène); forêt de Valle Mala, near col de St-Eustache; Chiovone river valley (Aullène); near Aullène; near

Serra-di-Scopamène (S of Aullène); 4 km E of Zonza; between Zonza and Quenza; Punta di u Carapono (Coti-

Chiavari); between Casalabriva and Olmeto; S of col d'llarata, between Zonza and TOspedale; between l’Ospedale

and Zonza, E of Punta di u Diamante; I'Ospedale and surroundings; near Propriano; Monte Rosso (SE of Sartène);

Tizzano and environs (SW of Sartène); La Trinité (N of Porto-Vecchio).

Previous records. — L'Ospedale, Vizzavona, Vico (Dollfus 1888, 1897, 1899; Arcangeli 1925); Porto,

Sartène, Porto-Vecchio, Melo (Dollfus 1888, 1897; Arcangeli 1925); Ajaccio (Scharff 1894; Dollfus 1897);

Corte (Sharff 1894); Gravona (Dollfus 1897).

Distribution. — The nominal subspecies is present in Corsica and Sardinia. A closely related subspecies,

T. Corsica giuslii Taiti & Ferrara, 1980, is known from the islands of Capraia, Gorgona and Montecristo

(unpublished datum), in the Tuscan Archipelago.

Tiroloscia macchiae Verhoeff, 1931

Philoscia (Tiroloscia) macchiae Verhoeff, 1931: 545.

Material examined. — Many 6 6 and 2 2 collected from the following localities: Ersa; Rogliano; Camera

and environs (near Centuri); between marine de Meria and Macinaggio; Morsiglia; between Morsiglia and Pastina;

near Meria; near Carbonacce (S of Luri); marine de Giottani (between Nonza and Centuri-Port); St-Léonard (W

of marine de Pietracorbara); marine de Pietracorbara; Sta Catterina (N of marine de Sisco); near Crosciano along

Sisco river; marine de Sisco; Nonza; near grotte de Brando; Erbalunga; Cardo (W of Bastia).

Previous records. — Corsica (Verhoeff 1931).

Distribution. — Tiroloscia macchiae is known from the Cap Corse peninsula, the south-western part of

the Tuscan mainland and the Tuscan Archipelago (Elba and Pianosa islands) (Taiti & Ferrara 1989a, fig. 83).

Tiroloscia montana n.sp.

(Figs 11, 12)

Material examined. — 1 6 holotype (MZUF), 1 S juv., 1 2 paratypes (MNHN-Is4102), 6 2 2,6 juvs

paratypes (MZUF), Haut-Asco, leg. S. Taiti and A. Poggesi, 21.VI. 1984; 1 2 paratype (MZUF), Haut-Asco.

1600 m, leg. S. Taiti and S. Campanelli, 16.X.1982; 9 22 paratypes (SMNS), Haut-Asco, 1500-1600 m, leg.

W. Schawaller, 27.VIII.1980; 1 6 juv., 9 2 2 paratypes (MZUF), forêt de Carozzica, ca. 2 km NE of Haut-Asco,

leg. S. Taiti and A. Poggesi, 21.VI.1984.

Etymology. — L. montanus = montane. The name refers to the collecting sites of the specimens, at an

altitude of over 1500 m.

Description

Maximum dimensions: <J, 6.5 x 2.5 mm; 9, 8.5 x 4 mm.

Source : MNHN, Paris

— 488 —

Source : MNHN, Paris

— 489 —

Brown colour with the usual yellowish muscle spots. Dorsal cuticle covered with numerous

tiny semicircular scales and many lanceolate scale-spines. Marginal line of pereonites slightly

distinct, more distant from the lateral margin in the anterior half where the numerous gland

pores (> 50) are situated. Co-ordinates of the noduli latérales as in Fig. 1 IB. Eye with 15-17

ommatidia. Cephalon with median lobe at obtuse angle, slightly protruding; lateral lobes small,

rounded; frontal line slightly bent down in the middle; suprantennal line straight. Pereonite 1

with posterior margin broadly rounded; pereonite 2 with subright posterior corners; pereonites

3-7 with posterior comers progressively more acute. Pleonites 3-5 with large falciform epimera,

directed backwards. Telson triangular with broken sides and acute apex. Antenna with fifth article

of peduncle slightly longer than flagellum; ratio of flagellar articles 5:3:5. Mandible with molar

penicil dichotomized. Maxillular outer branch bearing ten (five cleft) teeth; inner branch with

two penicils and a short posterior point. Maxillipedal endite with three strong spines at apex

and no penicil. Pleopodal exopods with no respiratory areas.

Male

Pereopods 1-4 with a brush of short spines on merus and carpus. Pereopod 7 without distinct

modifications. Pleopod 1 exopod ovoidal, with a slight indication of a posterior point; endopod

with straight distal part bearing a line of short spines, apex with two spines and a tuft of setae.

Pleopod 2 endopod slightly longer than exopod.

Remarks

Tiroloscia montana is readily distinguished from T. Corsica Corsica and T. macchiae, the

other two species in the genus present in Corsica, by the cephalon with more developed lateral

lobes, the different shape of the dorsal scale-spines, the telson with broken sides, and the shape

of the male pleopod 1 exopod with a slight indication of a posterior point.

In the shape of the dorsal scale-spines the new species resembles T. esterelana (Verhoeff,

1918), from which it differs in the broader body shape, larger pleonal epimera, the shape of the

cephalon, telson and male pleopod 1.

It differs from T. exigua (Budde-Lund, 1885) and T. pyrenaica (Dollfus, 1897) in the shape

of the scale-spines, telson and male pleopod 1 endopod; from T. apenninorum Verhoeff, 1908

essentially in the lack of a hook-shaped process on the male pereopod 7 merus.

Family ONISCIDAE

Genus SARDONISCUS Arcangeli, 1939

Sardoniscus pygmaeus (Budde-Lund, 1885)

Philoscia pygmaea Budde-Lund, 1885: 212; DOLLFUS 1887: 11; 1897: 91; 1899: 207.

Phalloniscus pygmaeus; Vandel 1954a: 74; 1954c: 226, figs A-D; 1962: 532, figs 262-265;

Arcangeli 1954a: 123.

Material examined. — Many <5 S and 9 9 collected from the following localities: Barcaggio (Cap Corse):

Ersa; Camera (near Centuri); Rogliano; between Macinaggio and marine de Meria; between Morsiglia and Pastina;

Source : MNHN, Paris

— 490 —

2 km SE of col de Ste-Lucie (between Pino and Luri); Minervio (S of Pino); Carbonacce (S of Luri); marine

de Giottani (between Nonza and Centuri-Port); Ponticello (near Pietracorbara); St-Léonard (W of marine de Pietra-

corbara); near Crosciano along Sisco river; Sta Catterina, N of marine de Sisco; between Ogliastro and Lainosa

(N of Nonza); near Erbalunga; near grotte de Brando; Lavasina (S of Brando); Sta Maria-di-Lota; San Martino-

di-Lota; Cardo (W of Bastia); col de Teghime (W of Bastia); near Murato (St Florent); Bevinco river valley,

below col de S. Stefano; 1 km W of Altiani (SE of Speloncato); near Ponte Nuovo along Golo river; near grotte

de Pietralbello (Moltifao); near Ponte Leccia; left bank of Golo river, 3 km S of Ponte Leccia; forêt de Carozzica

FIG. 12. — Tiroloscia montana n.sp., 6: A, pereopod 7; B, pleopod 1 exopod; C, pleopod 1 endopod; D, pleopod 2.

Source : MNHN, Paris

— 491 —

(near Haut-Asco); Francardo (N of Corte); Castirla (N of Corte); Lano (NW of Corte); near the Eaux d’Orezza

(NE of Piedicroce); 2 km S of Carticasi (NE of Corte); Cervione; forêt de Valdo-Niello (Golo river valley);

Punta Cervello (SE of Cauro); Zicavo; between Pietrosella and Fogolina (SE of Porticcio); between Alza and

Argiavara (col de Bavella); col de Bavella; between col de St-Eustache and col de Tana (SW of Aullène); Chiovone

river valley, near Aullène; Serra-di-Scopamène (S of Aullène); 4 km E of Zonza; valley below chapelle Pianelli,

between Casalabriva and Olmeto; Golfo di Sogno (N of Porto- Vecchio); Monte Rosso, SE of Sartène; Orasi (S

of Sartène); near Tizzano, baie d'Avena (SW of Sartène): Viagenti (near Pianotolli, NW of Bonifacio); near

Ermitage de la Trinité (W of Bonifacio).

Previous records. — La Solenzara (Dollfus 1897, 1899; Vandel 1962); Bastia, Francardo, Lano, Solaro,

Sari-di-Porto- Vecchio, Ste-Lucie-de-Porto-Vecchio, Propriano, Bonifacio (Vandel 1954c, 1962).

Distribution. — Sardoniscus pygmaeus is known from Corsica, Sardinia, Elba Island and some hills of

central-western Tuscany (Taiti & Ferrara 1989a, fig. 84).

Remarks

The genus Sardoniscus was erected by Arcangeli (1939) to include the new species

S. pusillus from Sardinia, described on specimens that he had previously identified as Agabi-

formius hirtus (Aubert & Dollfus, 1890) (Arcangeli 1925). Vandel (1954a) states that S. pusil¬

lus corresponds to “ Philoscia ” pygmaea Budde-Lund, 1885 and it belongs to Phalloniscus

Budde-Lund, 1908, a genus instituted to accommodate some species from New Zealand. Ac¬

cording to the French author the genus is also present in the Mediterranean and neotropical

regions. Thus, according to Vandel the correct taxonomic status of the species is Phalloniscus

pygmaeus (Budde-Lund, 1885), of which Sardoniscus pusillus is a junior synonym. The ascription

of this species to Phalloniscus was questioned by Arcangeli (1954a) and Ferrara & Taiti

( 1978), but this status has been maintained in anticipation of comparison of species of Phal¬

loniscus from New Zealand. However, we have to point out that, in one of his latest papers,

Vandel himself (1977) states that the genus Phalloniscus is endemic to New Zealand.

We have had the possibility recently to examine two specimens (1 <î, 1 $) collected at

Wellington, New Zealand, belonging to Phalloniscus armatus Bowley, 1935. Even if this is not

the type-species of Phalloniscus ', it certainly belongs to this genus. The Corsican species differs

from P. armatus in some important generic characters and cannot be ascribed to Phalloniscus.

In fact, it has a single nodulus lateralis (two in P. armatus) per side on the pereonites (see

Vandel 1962: 35 and Ferrara & Taiti 1978: 31, pi. 3, fig. XI, 1) and some cleft teeth (all

teeth entire in P. armatus ) on the outer branch of the maxillule. Thus, we consider valid the

genus Sardoniscus , to which the species pygmaeus must be ascribed. All the other species from

the Mediterranean area presently in Phalloniscus must be re-examined to define their correct

generic position.

In a recent paper, SCHULTZ (1995) transferred the species of Phalloniscus from the neotropi¬

cal region (some only tentatively) to the new genus Novamumdoniscus.

1. The type species of Phalloniscus is Oniscus punctatus Thomson, 1879, which, due to its vague description, has never

been encountered again. In fact VANDEL (1977: 12) propose to suppress this name.

Source : MNHN, Paris

— 492 —

Family Platyarthridae

G enus PLATYARTHRUS Brandt, 1833

Platyarthrus costulatus Verhoeff, 1908

(Figs 13 A, 15)

Platyarthrus Nabeleki Frankenberger, 1939: 115, figs 1-2.

Platyarthrus costulatus ; Arcangeli 1950: 126; Vandel 1953a: 163; 1954a: 73; Lanza &

POGGESI 1986: 120, 178, 179.

Platyarthrus costulatus costulatus ; Vandel 1962: 461, figs 230, 231 A; Beron 1972a: 11.

Material examined. — Many 6 6 and 2 $ collected from the following localities: Capo Grosso, Tollare

and Barcaggio (Cap Corse); moulin Mattéi (NE of Centuri-Port); Ersa; Rogliano; Centuri-Port; Camera (near

Centuri); between Macinaggio and marine de Meria; Morsiglia; near Meria; marine de Giottani (between Nonza

and Centuri-Port); Sta Catterina, N of marine de Sisco; Nonza; anse de Faggiola (désert des Agriates); col de

S. Stefano (S of Olmeta-di-Tuda); near Sorio; col de S. Colombano (E of Belgodère); left bank of Golo river

between Barchetta and Casamozza; near grotte de Pietralbello (Moltifao); Asco river valley; near Folelli, along

Fium Alto; Francardo (N of Cone); Prunete (E of Cervione); plage de Caspio (NW of Porto); Casanova (S of

Corte); between Volta and Scandulaie, along a tributary stream of Tavignano river (NW of Aléria); Aléria; Punta

de la Parata (W of Ajaccio); Porticcio; between Pietrosella and Fogolina (SE of Porticcio); Punta di u Carapono

(Coti-Chiavari); valley below chapelle Pianelli, between Casalabriva and Olmeto; Lovo Santo, swamp at mouth

of Cavo river; Bocca Albitrina (SW of Sartène); la Trinité (N of Porto-Vecchio); Viagenti (near Pianotolli, NW

of Bonifacio); Ermitage de la Trinité (NW of Bonifacio); Capo Pertusato (SE of Bonifacio); île de la Giraglia;

île Pietricaggiosa (Cerbicale); Mot Toro Grande; Mot du Torello.

Previous records. — Ajaccio (Frankenberger 1939); grotte des Paladini (Solaro) (Vandel 1953a; Beron

1972a); all Corsica (Vandel 1962); grotte de Sisco II (Beron 1972a); Me Pietricaggiosa, Mot du Torello, Mot Toro

Grande (Lanza & Poggesi 1986).

Distribution. — This species occurs in northern Spain, the Balearic islands, Atlantic and Mediterranean

part of France, Morocco and Algeria. In Italy it is recorded from Liguria, Tuscany, including the Tuscan

Archipelago (Capraia, Gorgona and Elba islands), Sardinia, Ischia Island, Tremiti islands, Sicily and surrounding

islands (Aeolian, Aegadean and Pantelleria).

Platyarthrus corsicus n.sp.

(Figs 13B, C, 14, 15)

Material examined. — 1 6 holotype (MZUF), 2 66, 3 9 2 paratypes (MNHN-Is4098), many 66 and

2 2 paratypes (MZUF), between Cauro and col de St-Georges (E of Ajaccio), 650-680 m, leg. S. Taiti, 14.V. 1982;

1 6, 8 2 2 paratypes (MZUF), Punta de la Parata (W of Ajaccio), leg. S. Taiti, 14.V. 1982; many <5 6 and 9 9

paratypes (MZUF), between Pietrosella and Fogolina (SE of Porticcio), ilex and arbutus wood, leg. S. Taiti and

S. Campanelli, 21.X.1982; 2 66, 1 2 paratypes (MZUF), between Alza and Argiavara (col de Bavella), along

the first left tributary stream of S. Pietro river, ilex grove, leg. S. Taiti, 11.VII.1978; 2 66,3 9 2 paratypes

(MNHN-Is4099), many 6 6 and 9 2 paratypes (MZUF). col de Bavella, leg. S. Taiti, 14.IV. 1981; 4 6 6, 6 2 9

paratypes (MZUF), between col de St Eustache and col de Tana (W of Aullène), coniferous forest, leg. S. Taiti

and S. Campanelli, 22.X.1982; 2 6 6, 3 2 2 paratypes (MZUF), forêt de Valle Mala (S of col de St-Eustache),

leg. S. Taiti and S. Campanelli, 22.X.1982; many 6 6 and 2 2 paratypes (MZUF), Punta di u Carapono (Coti-

Chiavari), leg. S. Taiti and S. Campanelli, 21.X.1982; many 6 6 and 2 2 paratypes (MZUF), valley below

chapelle Pianelli, between Casalabriva and Olmeto, leg. S. Taiti and S. Campanelli, 22.X.1982; 1 9 paratype

(MZUF), Fiumicicoli river valley, 700 m, leg. W. Schawaller, 1.VIII. 1980; many 6 6 and 9 2 paratypes (MZUF),

Monte Rosso (SE of Sartène), ilex grove and meadow, leg. S. Taiti and S. Campanelli, 20.X.1982; 2 6 6,3 2 2

paratypes (MNHN-Is4096), many 6 6 and 9 2 paratypes (MZUF), Tizzano and surroundings (SW of Sartène),

ilex grove, leg. S. Taiti and S. Campanelli, 15.V.1982.

Source : MNHN, Paris

— 493 —

Source : MNHN, Paris

— 494 —

Description

Maximum dimensions: â, 2.5 x 1.1 mm; 9, 3.3 x 1.3 mm.

Colourless body, eye absent. Dorsal ornamentation: cephalon with 2 + 2 large tubercles on

vertex; pereonite 1 with 3 + 3 ridges and a small tubercle at the base of epimera; pereonites 2-6

with 3 + 3 ridges, very protruding except the second one on each side which progressively gets

Source : MNHN, Paris

— 495 —

smaller from front to back; pereonite 7 with 2 + 2 ridges, well developed and protruding back¬

wards. Dorsum with many rounded and flattened scale-spines. Body very convex, ovoidal.

Cephalon with median lobe rounded, protruding frontwards and slightly hollow dorsally; lateral

lobes subquadrangular, tranversely directed frontwards. Pereonite 1 with posterior corners right-

angled, becoming progressively more acute in the following pereonites. Pleonites 3-5 with

epimera falciform, narrow and long, directed backwards. Telson with a short basal part and a

narrow triangular distal part, clearly protruding backwards compared with the uropodal protopods.

Antenna with thickset peduncular articles; second flagellar article about three times as long as

first. Uropods with thickset exopod, slightly shorter than protopod.

Male

Pleopod 1 exopod small and ovoidal; endopod with apical part slightly swollen and pointed.

Pleopod 2 endopod much longer than exopod.

Remarks

Platyarthrus corsicus is morphologically close to P. costulatus, from which it differs in

having more developed lateral and paramedian ridges on pereonites 1-6, and 2 + 2 instead of

3 + 3 ridges on pereonite 7. These differences are constant and no specimens with intermediate

characters have been observed, even from localities where both forms have been collected. In

our opinion these two forms belong to distinct, even if closely related, species.

In the past P. corsicus has certainly been confused with P. costulatus and some records

from Corsica of the latter refer to the new species, which is present only in the southern part

of the island.

Platyarthrus caudatus Aubert & Dollfus, 1890

(Fig. 15)

Platyarthrus caudatus ; VANDEL 1954a: 73; 1962: 457, figs 228-229.

Material examined. — 1 9, Cardo (W of Bastia), leg. F. Bernini, 19.IV. 1980; 1 juv., same locality, leg.

S. Taiti and S. Campanelli, 12.X.1982; 5 9 9, Ermitage de la Trinité (W of Bonifacio), leg. S. Taiti, 12.IV.1981.

Previous records. — Brando, Sisco, Bonifacio (Vandel 1962).

Distribution. — Known from most of the lands of the western Mediterranean.

Platyarthrus aiasensis Legrand, 1953

(Fig. 15)

Material examined. — 8 9 9, Ermitage de la Trinité (W of Bonifacio), leg. S. Taiti, 12.IV.1981.

Distribution. — This species has a West-Mediterranean-Atlantic distribution. It is known from Aix Island,

the Canaries. Madeira Island, Algeria, Corsica, Sardinia, south-western Tuscany and the Tuscan Archipelago (Elba,

Gorgona and Giannutri islands), Sicily and surrounding islands (Aeolian, Ustica, Aegadean, Pantelleria, Pelagean),

and Malta. It has been introduced to South Africa (Cape Town), the USA (California and Texas) and St Barthélémy

Island in the Caribbean (Garthwaite & Taiti 1989).

Source : MNHN, Paris

— 496 —

Source : MNHN, Paris

— 497 —

Platyarthrus schoeblii schoeblii Budde-Lund, 1883

(= P. schoeblii intermedius Vandel, 1946)

(Fig. 15)

Platyarthrus schoeblii ; SCHARFF 1894: 163; DOLLFUS 1899: 190; ARCANGELI 1925 : 446;

Vandel 1954a: 73.

Platyarthrus schôbli schôbli', Vandel 1962: 456, fig. 227B.

Material examined. — Many 6 6 and 2 2 collected from the following localities: moulin Mattéi (NE of

Centuri-Port); Rogliano; Morsiglia; Carbonacce (S of Luri); Ponticello (near Pietracorbara); Sta Catterina, N of

marine de Sisco; near Crosciano along Sisco river; between Ogliastro and Lainosa (N of Nonza); near grotte de

Brando; Cardo (W of Bastia); col de S. Stefano (S of Olmeta-di-Tuda); left bank of Golo river between Barchetta

and Casamozza; between Volta and Scandulaie along tributary stream of Tavignano river (NW of Aléria).

Previous records. — St-Florent (Scharff 1894); Vico, Ajaccio (Dollfus 1899; Arcangeli 1925); Corsica

(Vandel 1962).

Distribution. — Due to the uncertainties in the taxonomy of the schoeblii-gtoup (see also Caruso &

Lombardo 1982; Caruso et al. 1987), it is difficult to define the distributions of the various subspecies. Besides

Corsica, P. schoeblii schoeblii Budde-Lund (= P. schoeblii intermedius Vandel) is present in southern France,

Sardinia, the Tuscan mainland and the Tuscan Archipelago, Sicily, Aeolian islands, Malta and Algeria.

Remarks

According to Vandel (1962) the subspecies P. schoeblii schoeblii is present and widely

distributed in Corsica. The numerous specimens examined by us belong instead to the subspecies

P. schoeblii intermedius Vandel, which, according to the French author, is distinguished from

the nominal taxon by the presence of a residual ridge 4 on the pereonites (absent in the subspecies

schoeblii, see Vandel 1962, figs 227A and 227B). We have been able to re-examine the syntypes

of P. schoeblii described by BUDDE-LUND (1885) from Bona, Algeria, deposited in the Natural

History Museum, London and the Zoologisk Museum, Copenhagen. All the specimens have the

dorsal ridges as in P. schoeblii intermedius Vandel, which therefore is a junior synonym of the

nominal subspecies.

Most probably P. schoeblii schoeblii sensu Vandel is a distinct subspecies from P. schoeblii

schoeblii sensu Budde-Lund but it cannot take this name. However, in Corsica only P. schoeblii

schoeblii sensu Budde-Lund nec Vandel (= P. schoeblii intermedius Vandel) is present.

Platvarthrus hoffmannseggii Brandt, 1833

(Fig. 15)

Material examined. — 12, Cardo (W of Bastia), leg. F. Bernini, 19.IV.1980; 1 6, 1 2, Aléria, leg.

F. Giusti, 30.XI.1983.

Distribution. — This species is distributed in central-southern Europe.

Source : MNHN, Paris

— 498 —

Family Cylisticidae

Genus CYLISTICUS Schnitzler, 1853

Cylisticus convexus (De Geer, 1778)

(Fig. 17)

Cylisticus convexus; Vandel 1954a: 74; 1962: 559, figs 275, 276.

Records. — Bastia (Vandel 1962).

Distribution. — Cylisticus convexus is widely distributed in Europe and Asia Minor, and it has been in¬

troduced with human activities to northern Africa, St Helena Island and America. Also the record of this species

in Corsica, based on a single specimen, is certainly due to an occasional introduction.

Cylisticus vandeli Taiti & Ferrara, 1980

(Fig. 17)

Cylisticus gracilipennis; Dollfus 1887: 10 (partim: Corsica); 1899: 187 (partim : grotte

d’Erbalunga); ARCANGELI 1925: 47 (partim: grotte d’Erbalunga).

Cylisticus esterelanus ; ARCANGELI 1950: 176; CASSOLA 1982: 654.

Cylisticus nasutus; VANDEL 1953a: 164; 1954a: 74; 1962: 573, figs 286, 287; Beron 1972a:

12; Taiti & Manicastri 1980: 256, fig. 9 (partim : Corsica).

Cylisticus vandeli Taiti & Ferrara, 1980: 270, fig. VI.

Material examined. — Many â 6 and 9 9 collected from the following localities: Tollare (Cap Corse);

Ersa; near Rogliano; Camera (near Centuri); between Macinaggio and marine de Meria; between Morsiglia and

Pastina; near Carbonacce (S of Luri); marine de Giottani (between Nonza and Centuri-Port); Ponticello (near

Pietracorbara); St-Léonard (W of marine de Pietracorbara); Sta Catterina, N of marine de Sisco; near Crosciano

along Sisco river; between Ogliastro and Lainosa (N of Nonza); Nonza; near grotte de Brando; Erbalunga; near

Bastia; Cardo (W of Bastia); Olmeta-di-Tuda; Urtaca; between Barchetta and Casamozza along left bank of Golo

river; near grotte de Pietralbello (Moltifao); grotte de Pietralbello; 3 km S of Ponte Leccia; Francardo; grotte

de Sahara; Fium Alto valley, loc. Casette; near Lano (E of Omessa); Casanova (S of Corte); Fontaine de Padula

(between Vivario and Vezzani); col de Vizzavona; near Ghisoni; Bocognano; Rocchio Pinzuto, lungo il ruscello

Fiumicelle, affluente del Fiume Solenzara: between Cauro and col de St-Georges (W of Ajaccio); between

Pietrosella and Fogolina (SE of Porticcio); Punta di u Carapono (Coti-Chiavari); between col de St-Eustache and

col de Tana (W of Aullène); forêt de Valle Mala, near col de St-Eustache; valley below chapelle Pianelli, between

Casalabriva and Olmeto; Ermitage de la Trinité (W of Bonifacio).

Previous records. — Grotte d'Erbalunga (Dollfus 1899; Arcangeli 1925; Vandel 1962); grotte des Maures

(Poggio di Venaco) (Vandel 1953a, 1962; Beron 1972a); forêt de Valdo-Niello (Vandel 1962); grotte de Grottone

(Lama) (Beron 1972a); Ghisoni (Taiti & Ferrara 1980).

Distribution. — This species is endemic to Corsica, where it is widely distributed.

Cylisticus uncinatus n.sp.

(Figs 16, 17)

Material examined. — 1 6 holotype, 3 <3<3, 4 9 9 paratypes (MZUF), 1 <3, 2 9 9 paratypes (MNHN-

Is4104), 1 <3 juv., 1.6 km from Stazzona, near road to the Eaux d'Orezza, leg. S. Taiti and A. Poggesi, 20.VI.1984;

2 <3 <5 paratypes (MZUF), col de Teghime. humus, leg. F. Bernini, 20.IV. 1980; 1 â paratype (SMNS), Bevinco

river valley, below col de S. Stefano, leg. W. Schawaller, 27.VII.1980; I (3,2 9 9 paratypes, near Sorio, near

Viacale bridge on Chiaraggio stream, leg. S. Taiti and S. Vanni, 9.III.1994 (MZUF); 2 <3<3, 5 9 9 paratypes

Source : MNHN, Paris

— 499 —

Source : MNHN, Paris

— 500 —

FIG. 17. — Distribution in Corsica of Cylisticidae species: 1, Cylislicus convexus : 2. C. vandelr, 3, C. uncinatus ; 4. Troglocylisticus

cyrnensis.

Source : MNHN, Paris

— 501 —

(MZUF), Monte S. Petrone slopes, 1000-1500 m, leg. A. Torchia and S. Zoia, 25.V.1982; 3 S<S, 6 2 9 paratypes

(MZUF), Punta di Chiarsgioli, Monte S. Petrone, NW slope, 1100 m, beech wood, leg. S. Taiti and A. Poggesi,

21.VI.1984; ? 1 d, 8 2$ (MZUF), col de Bavella, between Alza and Argiavara, along first left stream of

S. Pietro river, ilex grove, leg. S. Taiti. 11.VII.1978; ? 3 6 6, 6 2 2 (MZUF), col de Bavella, coniferous forest,

leg. S. Taiti, 14.IV. 1981.

Etymology. — L. uncinatus = having a hook. The name refers to the characteristic triangular process on

the male pereopod 7 merus.

Description

Maximum length: 6, 11.5 mm; $, 13 mm.

Body very pale brown. Dorsum smooth, with tiny triangular scale-spines. Noduli latérales

almost at the same distance from lateral margin; d/c co-ordinates with a maximum on pereonite

5. Eye small, with 5 to 10 ommatidia, according to the animal’s size. Cephalon with median

lobe triangular, very obtuse, slightly protruding above the vertex and frontwards; lateral lobes

small, quadrangular. Pereonite 1 with posterior margin slightly concave at the sides. Telson with

distal part much narrower than basal, triangular, subacute apex distinctly protruding backwards

in comparison with uropodal protopods. Antenna with second flagellar article 1.5 times as long

as the first.

Male

Pereopods 1-4 with a brush of spines on carpus and, more sparsely, on merus. Pereopod 7

ischium with rostral surface transversely depressed and covered with short setae, sternal margin

slightly concave with long setae; merus with a protruding setose triangular process on tergal

margin. Pleopod 1 exopod triangular with a short rounded posterior point bent outwards, outer

margin sinuous; endopod with straight apical part and no particular modifications.

Remarks

Cylisticus uncinatus , as well as C. vandeli, belongs to the nasuft/s-group of Cylisticus (see

Ferrara & Taiti 1978, 1985; Taiti & Manicastri 1980; Taiti & Ferrara 1980), which

includes fifteen more species and subspecies distributed in southern France, central Italy and

Sardinia. The new species is readily distinguishable from all others by the distinct process on

the male pereopod 7 merus. In the shape of the cephalon, telson and male pleopod 1 exopod,

C. uncinatus is close to the other Corsican species, C. vandeli, from which it differs, besides

the above-mentioned character, in its larger dimensions, presence of a feeble pigmentation, more

developed eye, and the male pleopod 1 endopod with apical part straight instead of bent outwards.

The specimens from col de Bavella show some differences from the type specimens in the

scale-spines longer and more pointed (Fig. 161), which give a setose look to the specimens, and

in the shape of the male pleopod 1 exopod (Fig. 16J). All the other characters, particularly the

shape of the male pereopod 7, are identical to those of the type specimens. They might belong

to a distinct species but more material is needed for a certain identification.

Source : MNHN, Paris

— 502 —

Genus TROGLOCYLISTICUS Ferrara & Taiti, 1983

Troglocylisticus cyrnensis Ferrara & Taiti, 1983

(Fig. 17)

Troglocylisticus cyrnensis Ferrara & Taiti, 1983: 485, figs I-II.

Records. — Grotte de Brando (Ferrara & Taiti 1983).

Distribution. — Known only from the type locality.

Family Porcellionidae

G enus PROTRACHEONISCUS Verhoeff, 1917

Protracheoniscus babori Frankenberger, 1938

Protracheoniscus occidentalism Vandel 1954a: 74; 1962: 578, figs 288-290.

Protracheoniscus babori ; Taiti & Ferrara 1980: 278.

Material examined. — Many 6 6 and 9 9 collected from the following localities: between Barcaggio and

Punta di Agnello (Cap Corse); marine de Pietracorbara; Lovo Santo, swamp at mouth of Cavo river.

Previous records. — Marine de Pietracorbara (Vandel 1962); mouth of Cavo river (Taiti & Ferrara

1980).

Distribution. — This species has a wide distribution: besides Corsica, it is recorded from southern France,

Catalonia, Italy (Tuscany and Sicily), Croatia, Albania (cited as Protracheoniscus albanicus Arcangeli, 1952; see

Schmalfuss 1983), Greece and probably southern Russia.

Remarks

Examination of the abundant material from Corsica shows that these specimens belong to

P. babori as redescribed by Taiti & Ferrara (1980). In fact, the male pereopod 7 ischium has

a deep concavity covered with setae, the merus bears a process at the base, and the carpus is

distinctly arched on the tergal margin. However, these characters are clearly visible only in fully

adult specimens.

Thanks to the courtesy of Dr. H. Dalens (Toulouse), we could examine some specimems

of Protracheoniscus occidentalis Vandel, 1939 collected near Toulouse, which proved this species

to be synonymous with P. babori.

Genus PORCELLIONIDES Miers, 1877

Porcellionides pruinosus (Brandt, 1833)

Metoponorthus (Metoponorthus) pruinosus ; Vandel, 1954a: 74.

Material examined. — Many 6 6 and 9 9 collected from the following localities: Rogliano: near moulin

Mattéi, N of Centuri-Port; Centuri-Port; between Bastia and St-Florent; défilé de Lancone; entrance of grotte de

Grottone, near Lama; Asco river valley; near Ermitage de la Trinité (W of Bonifacio); île de la Giraglia.

Source : MNHN, Paris

— 503 —

Previous records. — Corsica (Vandel 1954a).

Distribution. — Cosmopolitan species.

Porcellionides sexfasciatus sexfasciatus (Budde-Lund, 1885)

Metoponorthus sexfasciatus; Scharff 1894: 163; DOLLFUS 1899: 189.

Metoponorthus (Polytretus) sexfasciatus; VANDEL 1954a: 74.

Metoponorthus (Polytretus) sexfasciatus sexfasciatus; VANDEL 1962: 608, figs 299-302.

Material examined. — Many â 6 and 9 9 collected from the following localities: near Bastia; Casta (SW

of St-Florent); entrance of grotte de Grottone, near Lama; Calzarello, SE of Ghisonaccia; Punta de la Parata (W

of Ajaccio).

Previous records. — Corte (Scharff 1894); Vizzavona, Porto-Vecchio (Dollfus 1899); littoral regions of

Corsica (Vandel 1962).

Distribution. — It is known from the Atlantic islands, Morocco, Algeria, Tunisia, Spain, France, Italy and

Malta.

Genus ACAEROPLASTES Verhoeff, 1918

Acaeroplastes melanurus sardous Verhoeff, 1918

(Fig. 18)

Metoponorthus melanurus; BUDDE-LUND 1885: 181 (partim; Corsica); SCHARFF 1894: 163;

Dollfus 1899: 189.

Metoponorthus (Acaeroplastes) melanurus; ARCANGELI 1950: 115.

Metoponorthus (Acaeroplastes) melanurus sardous; Vandel 1954a: 74.

Acaeroplastes melanurus sardous; VANDEL 1962: 633, fig. 313; Taiti & Ferrara 1980:

280.

Aceroplastes [sic!] melanurus sardous; Lanza & Poggesi 1986: 121, 176, 180, 188.

Material examined. — Many $ 6 and 9 9 collected from the following localities: Tollare (Cap Corse);

Centuri-Port; marine de Giottani (between Nonza and Centuri-Port): marine de Pietracorbara; anse de Faggiola,

désert des Agriates; Casta (SW of St-Florent); Pas du Diable (W of Casta); col de S. Stefano (S of Olmeta-di-

Tuda); col de S. Colombano (E of Belgodère); between Barchetta and Casamozza along left bank of Golo river;

forêt de Tartagine; Ghisonaccia; Porticcio; below Punta di Boccarona, along Solenzara river; between col de

Bavella and Punta Velaco; Sollacaro (N of Olmeto); Fautea (between Solenzara and Porto-Vecchio); Lovo Santo,

swamp at mouth of Cavo river; mouth of Rizzanèse river (SW of Propriano); Golfo di Sogno (NE of Porto- Vec¬

chio); Bocca Albitrina (SW of Sartène); Monte Rosso (SE of Sartène); dolmen de Fontanaccia (S of Sartène);

Tizzano and environs, baie d'Avena (SW of Sartène); Viagenti (near Pianotolli, NW of Bonifacio); Capo Pertusato

(SE of Bonifacio); île Spano (NE of Calvi); îlot Fautea; îlot sud de la Tonnara (golfe de Ventilegne); rocher sud

de Ratino (W of île Cavallo); îlot Sperduto Grande (E of île Cavallo).

Previous records. — Corsica (Budde-Lund 1885); Ajaccio (Scharff 1894; Dollfus 1899); étang de Biguglia

(Dollfus 1899); all Corsica, Ponte Leccia, Sta-Maria-Figaniella and Sartène (Vandel 1962); mouth of Cavo river

(Taiti & Ferrara 1980); îlot Fautea, îlot Sperduto Grande, rocher sud de Ratino, île Spano (Lanza & Poggesi

1986).

Distribution. — Known from Corsica, Sardinia and Tuscany (Montecristo and Elba Islands, La Scarpa

islet near Pianosa Island. Monte Massoncello, Uccellina and Monte Argentario).

Source : MNHN, Paris

— 504 —

Source : MNHN, Paris

— 505 —

Remarks

The abundant material examined revealed some differences in the male uropodal exopod

in the various populations. Southern populations (e.g. Capo Pertusato) (Fig. 18A) have a much

more flattened and widened exopod than northern ones ( e.g. Cap Corse peninsula) (Fig. 18E),

even in male specimens of the same length (9 mm). In some populations (e.g. Cavo river mouth)

this appendage has an intermediate development, while the shape of the male pereopod 7 and

pleopod 1 is identical in all populations. In our opinion these differences must be considered

within the variability of this subspecies.

Genus LEPTOTRICHUS Budde-Lund, 1885

Leptotrichus panzerii (Audouin, 1826)

Leptolrichus Panzerii ; Budde-Lund 1885: 193, 194; Arcangeli 1914: 477; 1950: 121.

Leptotrichus Panzeri ; DOLLFUS 1887: 10; ARCANGELI 1925: 17; VANDEL 1954a: 74.

Leptotrichus panzeri ; VANDEL 1962: 645, figs 317-319.

Material examined. — Many 6 6 and 2 2 collected from the following localities: Centuri-Port; Bonifacio;

Capo Pertusato (SE of Bonifacio).

Previous records. — Corsica (Budde-Lund 1885; Dollfus 1887; Vandel 1954a); Bonifacio and Propriano

(Vandel 1962).

Distribution. — This species occurs in all the lands encompassing the Mediterranean Sea, Madeira Island,

the Canaries, Cape Verde and the Bermudas.

Genus AGABIFORMIUS Verhoeff, 1908

Agabiformius lentus (Budde-Lund, 1885)

Agabiformius lentus ; Vandel 1954a: 74; 1962: 640, figs 315-316.

Material examined. — 12, Cardo (W of Bastia), leg. S. Taiti and A. Poggesi, 27.VI.1984.

Previous records. — Corsica (Vandel 1954a); Sisco (Vandel 1962).

Distribution. — This species is very common in all the lands of the Mediterranean basin. It has been

introduced by man to Madeira Island, the Canaries, Senegal, Oman, Seychelles, China, Mexico, Haiti, Venezuela,

Bermudas and Hawaiian islands. It has also been recorded from greenhouses in England.

Genus LUCASIUS Kinahan, 1859

Lucasius pallidus (Budde-Lund, 1885)

Lucasius pallidus ; VANDEL 1954a: 74.

Lucasius pallidus pallidus ; Vandel 1962: 651, figs 320-321.

Material examined. — Many â 6 and 2 2 collected from the following localities: valley below chapelle

Pianelli, between Casalabriva and Olmeto; mouth of Rizzanèse river, S of Propriano; Monte Rosso. SE of Sartène.

Previous records. — Corsica (Vandel 1954a).

Distribution. — Portugal, Spain, southern France, Morocco, Corsica, Sardinia and Tuscany.

Source : MNHN, Paris

— 506 —

Genus PORCELLIO Latreille, 1804

Porcellio spatulatus Costa, 1882

(Fig. 19)

Porcellio latissimus ; Budde-Lund 1885: 95, 302; DOLLFUS 1887: 10; 1899: 188; 7SCHARFF

1894: 163.

Porcellio spatulatus; ARCANGELI 1925: 15, pi. 4, figs 1-2; Vandel 1954a: 73; 1962: 660;

Lanza 1979: 49; CASSOLA 1982: 654; Lanza & POGGESi 1986: 66, 71, 121, 177, 178, 179,

180, 182, 184, 185.

Source : MNHN, Paris

— 507 —

Porcellio (Polyplatus) spatulatus ; Arcangeli 1950: 90.

Material examined. — Many 66 and 2 2 collected from the following localities: near Tizzano (SW of

Sartène); Capo Pertusato (SE of Bonifacio); île de Comuta (golfe de San Cipriano); îlot La Folaca (near Palombag-

gia, SE of Porto-Vecchio); rocher de Vacca (Cerbicale); île Pietricaggiosa (Cerbicale); îlot Toro Grande; îlot Toro

Piccolo; îlot Porraggia Piccola and îlot Porraggia Grande (N of île Cavallo); grand îlot des Bruzzi (W of golfe

de Figari); îlot Fazzuolo Piccolo (NW of Bonifacio); rocher sud de Ratino (W of île Cavallo); île Saint-Antoine

(Capo Pertusato, SE of Bonifacio).

Previous records. — Corsica and île Lavezzi (Budde-Lund 1885; Arcangeli 1925, 1950); ?Corte (Scharff

1894); île Saint-Antoine (Lanza 1979; Lanza & Poggesi 1986); île de Comuta, rocher de Vacca, île Pietricaggiosa,

îlot Toro Grande, îlot Toro Piccolo, îlot La Folaca, îlot Porraggia Piccola, îlot Porraggia Grande, rocher sud de

Ratino, île Lavezzi, îlot Fazzuolo Piccolo, grand îlot des Bruzzi (Lanza & Poggesi 1986).

Distribution. — Southern Corsica, Sardinia and northern Tunisia (ca 28 km N of Sousse, unpublished

datum).

Remarks

In Corsica this species is strictly littoral. Thus, the record from Corte (Sharff 1894) most

probably is due to a misidentification or a mislabelling of the locality.

The male characters of this species are illustrated in Fig. 19.

Porcellio scaber Latreille, 1804

Material examined. — 2 6 6, 2 2 2, near Tizzano, SW of Sartène, leg. S. Taiti, 15.V.1982.

Distribution. — Cosmopolitan species.

Porcellio diiatatus dilatatus Brandt, 1833

Porcellio dilatatus ; Scharff 1894: 163; Remy 1950: 6, 7, 9; Vandel 1953a: 164; 1954a:

74; BERON 1972a: 12.

Material examined. — Many 6 6 and 2 2 collected from the following localities: near Meria; marine de

Giottani (between Nonza and Centuri-Port); near Ponticello (near Pietracorbara); St-Léonard (W of marine de

Pietracorbara); Sta Catterina, N of marine de Sisco; grotte de Sisco; near Crosciano along Sisco river; grotte de

Brando and environs; Sto Pietro-di-Tenda; grotte de Sahara (Castiglione).

Previous records. — Ajaccio (Scharff 1894); grottes de Sisco and Brando (Remy 1950; Vandel 1953a;

Beron 1972a); grottes de Sahara, Cabanuli (Omessa), Sisco 11, Ostriago (Pietracorbara) and Manuel-Ange (Lozzi)

(Beron 1972a).

Distribution. — Known from all Europe but rare at the east of the Italian peninsula. It has been introduced

to North and South America, Sri Lanka and Hawaii.

Porcellio laevis Latreille, 1804

Porcellio laevis ; Scharff 1894: 163; Verhoeff 1926: 263; Vandel 1953a: 164; 1954a:

74; Beron 1972a: 13; Lanza & Poggesi 1986: 121, 182.

? Porcellio ragusae\ Verhoeff 1926: 263; Vandel 1954a: 73; 1962: 660.

Source : MNHN, Paris

— 508 —

Material examined. — Many 6 â and 2 2 collected from the following localities: Tollare (Cap Corse);

marine de Pietracorbara; marine de Sisco; Casta (SW of St Florent); défilé de Lancone; Folelli, along Fium Alto;

Ghisonaccia; Porticcio; valley below chapelle Pianelli, between Casalabriva and Olmeto; mouth of Rizzanèse

river, SW of Propriano; Bocca Albitrina, W of Sartène; Monte Rosso, SE of Sartène; Tizzano, SW of Sartène;

dolmen de Fontanaccia and menhir de Renaggiu, S of Sartène; Bonifacio; île de la Giraglia; île Lavezzi.

Previous records. — Bastia and St-Florent (Scharff 1894); road from Biguglia to St-Florent, Sagone

(Verhoeff 1926); grotte de St-Francois (Bonifacio) (Vandel 1953a; Beron 1972a); île Lavezzi (Lanza & Poggesi

1986).

Distribution. — Cosmopolitan species.

Remarks

On the basis of a re-examination of the types of Porcellio ragusae Dollfus, 1896, CARUSO

& Lombardo (1982: 26) demonstrated this species to be a junior synonym of P. laevis. Most

probably also the record of P. ragusae from Corsica refers to P. laevis.

Porcellio orarum vizzavonensis Verhoeff, 1928

(Fig. 20)

Porcellio variabilis (= P. transmutatus); DOLLFUS 1899: 188.

Porcellio lugubris vizzavonensis; Verhoeff 1926: 263 (nomen nudum); 1928: 123.

Porcellio (Porcellio) Verhoeffi; ARCANGELI 1950: 87.

Porcellio orarum vizzavonensis; Vandel 1951: 159, fig. 32; 1954a: 74; 1962: 731, figs

350, 352D-E; Taiti & Ferrara 1980: 282; Lanza & Poggesi 1986: 121, 170, 175, 176, 177,

178, 179, 181, 183, 185, 186, 187, 188.

Material examined. — Many 6 6 and 2 2 collected from the following localities: Capo Grosso, Tollare,

Barcaggio and pond between Barcaggio and Punta di Agnello (Cap Corse); marine de Giottani (between Nonza

and Centuri-Port); phare de Fornali, W of St-Florent; near Capo a u Cavallo (SW of Calvi); near grotte de

Pietralbello, S of Moltifao; Folelli, along Fium Alto; Haut-Asco; forêt de Carozzica (near Haut-Asco); near Asco;

Francardo; near Caporalino; N slope of Monte Cinto; W slope of Monte Cinto near refuge Altare; Cala Maiora,

anse de Gattoia (W of Girolata); Prunete (E of Cervione); plage de Caspio (NW of Porto); col de Vergio (NE

of Evisa); Porto; lac de Melo and lac de Capitello (Restonica river valley); Restonica river valley; Vecchio river

valley (S of Venaco); near Ghisoni; col de Vizzavona; Capannelle (SE of Vizzavona); lac de Vitelaca (upper

valley of Prunelli river); Punta de la Parata, W of Ajaccio; below Punta di Boccarona, along Solenzara river;

3 km N of col de la Vaccia (N of Aullène); col de Bavella and environs; foot of Punta Velaco (S of Bavella);

Fautea (between Solenzara and Porto- Vecchio); Lovo Santo, swamp at mouth of Cavo river; valley below chapelle

Pianelli between Casalabriva and Olmeto; S of Orone (W of I'Ospedale); Barrage de l’Ospedale; Taglio Rosso,

W of Ste-Lucie-de-Porto- Vecchio; baie d'Avena, SW of Sartène; golfe de Sant'Amanza; near Ermitage de la

Trinité (NW of Bonifacio); old fortress near Capo Pertusato (SE of Bonifacio); Capo Pertusato; île de la Giraglia;

îlot Terre and îlot Finocchiarola (N of Macinaggio); îlot Nord de Morsetta (N of Galéria); îlot Porri and rochers

d’Elpa Nera (W of Stolli, golfe de Girolata); îlot Palazzinu and îlot Palazzu (N of Punta Palazzo); îlot Garganellu

(near île Gargalu, S of Punta Palazzo); Zeccu d’a Furmicula (golfe d’Elbo); îlot Guardiola (SW of Porto); île

de Cala d'Alga (Sanguinaires); île Mezzu Mare (Sanguinaires); île Piana de Portigliolo (golfe d'Ajaccio); îlot

Fautea; îlot de la Roscana (golfe de Pinarello); île de Pinarello; îlot La Folaca (near Palombaggia, SE of Porto-

Vecchio); rocher de Vacca and île Pietricaggiosa (Cerbicale); îlot Toro Grande; îlot Toro Piccolo; grand îlot des

Bruzzi (NW of golfe de Figari); île Cavallo; îlot Luigi Giafferri (N of île Lavezzi); îlot Silene (NW of île

Lavezzi); île Lavezzi.

Previous records. — Vizzavona (Verhoeff 1926, 1928); Sargone (Verhoeff 1928); Corsica (Vandel 1962);

mouth of Cavo river, Fautea, Velaco, Diamante (Taiti & Ferrara 1980); île de la Giraglia, îlot Terre, îlot Finocchi¬

arola, îlot Fautea, îlot de la Roscana, île de Pinarello, rocher de Vacca, île Pietricaggiosa, îlot Toro Grande, îlot

Source : MNHN, Paris

— 509 —

Source : MNHN, Paris

— 510 —

Toro Piccolo, îlot La Folaca, île Cavallo, îlot Luigi Giafferri, îlot Gian Pietro Gaffori, îlot Silene, grand îlot des

Bruzzi, île Piana de Portigliolo, île Mezzu Mare, île de Cala d'Alga, îlot de Cala Maiora. îlot Garganellu, îlot

Palazzinu, îlot Porri, îlot Nord de Morsetta (Lanza & Poggesi 1986).

Distribution. — Known from Corsica and some islets around Sardinia (Argano & Manicastri 1991).

Remarks

The large number of specimens examined permits some morphological observations on this

taxon:

a, the maximum length is 15 mm for the male and 21 mm for the female (13 mm according

to Vandel 1951);

b, the shape of both cephalon and telson is very variable, also within the same population;

c, dorsal granulations are in general feeble, but in some specimens they are well developed;

d, the male pereopod 7 ischium has a setose area, the tergal margin of the carpus is usually

not or only slightly humped, also in very large specimens.

The variability found in the Corsican populations shows that the systematics of the orarum-

group is still unclear and a re-examination of the different subspecies (see Vandel 1962) is

necessary. Moreover, in central-southern Italy there is a form of this group which was considered

to be a subspecies of P. orarum Verhoeff, 1910 (P. orarum verhoeffi Dahl, 1916) (FERRARA &

Taiti 1978; Taiti & Ferrara 1980), but which certainly refers to Porcellio pumicatus Budde-

Lund, 1885 (Taiti & Ferrara 1989a). Thus, P. orarum should be a junior synonym of P. pumi¬

catus. Since we have not examined any specimen referable to the typical P. orarum , while

awaiting a complete analysis of all the forms in this group, we prefer to reserve the name

pumicatus only for the Italian taxon.

Porcellio lamellatus sphinx (Verhoeff, 1931)

Porcellio lamellatus ; Dollfus 1899: 188; Vandel 1954a: 74.

Porcellio lamellatus sphinx; Vandel 1962: 744, figs 357-358A, D-F; Lanza & POGGESI

1986: 121, 176, 185.

Material examined. — Many 6 â and 2 2 collected from the following localities: Barcaggio (Cap Corse);

anse de Faggiola (désert des Agriates); Calzarello (SE of Ghisonaccia); baie d'Avena (SW of Sartène); Punta di

Fautea (between Solenzara and Porto- Vecchio); Golfo di Sogno (NE of Porto- Vecchio); Capo Pertusato (SE of

Bonifacio); îlot terre and îlot intermédiaire (near îlot Finocchiarola, N of Macinaggio); île Piana de Portigliolo

(golfe d’Ajaccio).

Previous records. — Etang de Biguglia (Dollfus 1899); îlot terre, îlot intermédiaire, île Piana de Portigliolo

(Lanza & Poggesi 1986).

Distribution. — This littoral subspecies occurs in the eastern Iberian Peninsula, southern France. Corsica,

Sardinia, Italy and Dalmatia.

Remarks

Schmalfuss (1992: 4, 12) suggests to transfer P. lamellatus to the genus Proporcellio

Verhoeff, 1907, on the basis of the clear likeness of this species with Proporcellio quadriseriatus

Verhoeff, 1917. On the contrary, in an electrophoretic analysis of some species of Porcellio

Source : MNHN, Paris

— 511 —

including P. lamellatus, VIGLIANISI et al. (1992) show that the genetic distance of this species

falls within the range of the genus. Thus, we prefer to keep this taxon in Porcellio.

Family Armadillidiidae

G enus ALLOSCHIZIDIUM Verhoeff, 1919

Alloschizidium was erected by Verhoeff (1919) as a subgenus of Armadillidium Brandt,

1833 to include Armadillidium pruvoti Racovitza, 1907, from a cave in the French Maritime

Alps. It was later considered by the same author (VERHOEFF 1933a) to be a subgenus of

Troglarmadillidium Verhoeff, 1900, and finally by VERHOEFF (1933b) and ARCANGELI (1948) to

be a full genus.

Vandel (1944a, 1954d, 1962) divided the family Armadillidiidae into two subfamilies on

the basis of cephalic structure: Eluminae, with eleven genera, caracterized by a cephalon with

no postscutellar line 1 , and Armadillidiinae, including the sole genus Armadillidium, with a

cephalon bearing such a line. According to Vandel (1954d), since the postscutellar line is present

in A. pruvoti, this species has to be in Armadillidium, of which consequently Alloschizidium

was considered to be a junior synonym. The presence of a schisma at the posterior comers of

pereonite 1 justified only the institution of a group of species (the pruvoti- group), which included

A. pruvoti (with the typical form and a Corsican form), A. mateui Vandel, 1953 and A. racovitzai

Vandel, 1954.

Examination of many specimens belonging to the Corsican form of A. pruvoti shows that

this taxon represents a distinct species, even if morphologically very close to A. pruvoti, which

cannot be placed in Armadillidium. Moreover, it seems difficult to include this species in a

subfamily (Armadillidiinae) different from the other species ascribed by Vandel himself to the

Eluminae (e.g. considering only the Corsican fauna, Paraschizidium remyi Vandel, 1944).

Some questions arise from the above considerations:

— Is A. pruvoti a real Armadillidium and therefore Alloschizidium is synonymous with

Armadillidium ?

— Does Paraschizidium remyi, apparently so similar to A. pruvoti, belong to a different

genus and subfamily?

— In general, what is the validity of morphologically very close genera such as Para¬

schizidium Verhoeff, 1919, Typhloschizidium Arcangeli, 1933 and Nesolidium Verhoeff, 1941?

In order to clarify the taxonomic status of the Corsican species, we examined the following

species, also using the scanning electron microscope: Armadillidium pruvoti “forme de Corse”

(sensu Vandel); Paraschizidium coeculum (Silvestri, 1897), of which the type species P. olearum

Verhoeff, 1919, is a junior synonym (Manicastri & Taiti 1994); P. olearum sensu Vandel,

1962 nec Verhoeff, 1919; P. remyi; Typhloschizidium sardoum Arcangeli, 1933, type species of

the genus; T. igiliense Ferrara & Taiti, 1978; T. cottarellii Argano & Pesce, 1974; and Nesolidium

buchnerorum Verhoeff, 1941, type species of the genus (type specimens from Ischia Island).

1. According to VANDEL (1944a, 1954d, 1962) the postscutellar line is a neoformation. On the contrary, SCHMALFUSS

(1989: 209) states that this line is probably homologous to the frontal line in other families. Though the latter opinion

is certainly more likely, we prefer to keep using VANDEL's terminology, awaiting a specific study on this matter.

Source : MNHN, Paris

— 512 —

The following results emerged:

— P. coeculum and P. olearum sensu Vandel have a very simple cephalon, i.e. with a scutel-

lum laterally defined only by the antennary depressions and superiorly with no distinct margin,

and with a rudimentary postscutellar line. Moreover the antennule consists of two articles and

the posterior corners of pereonite 1 are not cleft. In our opinion Paraschizidium is a valid genus

and includes with certainty these two species. All the other species presently in Paraschizidium

must be re-examined to define their correct taxonomic status. Certainly the Greek species

described in this genus by Schmalfuss (1981) and Sfenthourakis (1992; 1995) do not belong

to Paraschizidium since they lack a postscutellar line, the scutellum is well defined on all its

sides and the posterior comers of pereonite 1 are distinctly cleft.

— Armadillidium pruvoti “forme de Corse”, Paraschizidium remyi, Typhloschizidium

igiliense, T. cottarellii and N. buchnerorum possess a cephalon with distinct postscutellar and

frontal lines and a well-defined scutellum (in T. cottarellii and N. buchnerorum the superior

margin is interrupted in the middle), pereonite 1 with a schisma at the posterior comers, and

the antennule with three articles. In our opinion, these species constitute a homogeneous group,

clearly different from Armadillidium, characterized by the ability to roll up in the form of a

lengthened ovoid (as in Paraschizidium), the cephalon with antennary lobes directed frontwards

and distinct postscutellar and frontal lines, antennule of three articles and pereonite 1 with

schisma. For all these species we propose to revalidate the genus Alloschizidium, of which Neso-

lidium is a junior synonym.

— Typhloschizidium sardoum apparently differs from the above-mentioned species in lack¬

ing the frontal line, while it shares all the other characters. The lack of the frontal line is certainly

a secondary feature caused by the very deep concavity of the antennary depressions. In fact, a

species from a cave in southern Tuscany (Alloschizidium cavernicolum Taiti & Ferrara, 1995),

morphologically very close to T. sardoum, shows the antennary depressions with concavities

slightly less marked and both lines (postscutellar and frontal) still present and very close to

each other (Taiti & FERRARA 1995b). In conclusion, also T. sardoum must be included in

Alloschizidium, of which Typhloschizidium is a junior synonym.

At present, Alloschizidium includes the following species (Taiti & Ferrara 1995b): A. pru¬

voti, A. campanellii n.sp. (= A. pruvoti “forme de Corse” sensu Vandel), A. racovitzai, A. igiliense,

A. remyi, A. cottarellii, A. buchnerorum, A. sardoum, A. eeae (ARGANO & UTZERI 1973) and

A. cavernicolum.

Alloschizidium campanellii n.sp.

(Figs 21, 22, 27)

Armadillidium Pruvoti; Vandel 1954a: 74; 1954d: 54, fig. 4 (partim: specimens from

Corsica).

Armadillidium pruvoti “forme de Corse”; Vandel 1962: 785, fig. 379.

Material examined. — 1 6 holotype (MZUF), 14 <3 6, 19 9 9 paratypes (MZUF), 2 6 6, 2 9 2 paratypes

(MNHN-Is4097), Tizzano (SW of Sartène), ilex grove, leg. S. Taiti and S. Campanelli, 15.V.1982; 2 6 6,

5 2 9 paratypes (MZUF), valley below chapelle Pianelli, between Casalabriva and Olmeto, leg. S. Taiti and

Source : MNHN, Paris

— 513 —

Source : MNHN, Paris

— 514 —

S. Campanelli, 22.X.1982; 6 S â, 21 9 9 paraiypes (MZUF), Monte Rosso (SE of Sartène), meadow near ilex

grove, leg. S. Taiti and S. Campanelli, 20.X. 1982; 1 6 paratype (MZUF), same data, ilex grove.

Previous records. — Ste-Lucie-de-Porto- Vecchio, Bonifacio, Sartène and environs, Ajaccio and environs

(Vandel 1954d, 1962).

Etymology. — The new species is named after our friend S. Campanelli for his invaluable help in collecting

the specimens.

Fig. 22. — AUoschizidium campanellii n.sp., 6 : A, antennule: B, uropod; C, pleopod 1 exopod; D, pleopod 1 endopod; E, pleopod 2.

Source : MNHN, Paris

-515-

Description

Maximum diameter of the animal when rolled up: <î, 3 mm; ?, 3.5 mm.

Colourless body, eye absent. Dorsum smooth, covered with many tiny triangular scale-spines.

Body strongly convex with vertical epimera, able to roll up into a perfect ball. Cephalon with

triangular scutellum, very wide superiorly, bent over vertex but clearly separated and protruding

from it; postscutellar line semicircular on both sides; antennary lobes quadrangular, directed front¬

wards. Pereonite 1 with posterior margin slightly sinuous; posterior corners bearing a schisma

with external lobe distinctly protruding backwards compared with the inner one; a light depression

along the whole lateral margin. Telson trapezoidal, slightly broader than long. Antennule of three

articles with a tuft of superimposed aesthetascs at apex. Antenna with second flagellar article

three times as long as first. Uropodal exopod longer than wide.

Male

Pleopod 1 exopod subrectangular, with no distinct posterior point; endopod with thickset

distal part, rounded apex. Pleopod 2 as in Fig. 22E.

Remarks

These specimens undoubtedly correspond to those from Corsica identified by Vandel

( 1954a, 1954d, 1962) as Armadillidium pruvoti. According to the French author, the morpho¬

logical differences between the Corsican population and the typical one are due to the subadult

stage of the specimens observed. However, the abundant material examined by us, including

male specimens of similar size to that of the typical A. pruvoti , shows that the differences noticed

by Vandel are constant and do not depend on the age of the specimens. Thus, the Corsican

population belongs to a distinct species of the genus Alloschizidium together with the closely

related A. pruvoti.

Alloschizidium campanellii differs from A. pruvoti in the shape of the telson with a broader

basal part, uropodal exopod narrower and longer, male pleopod 1 exopod with no distinct posterior

point. In cephalic structure A. campanellii resembles A. remyi, from which it is readily distin¬

guished by the triangular instead of piliform dorsal scale-spines, the longer telson and uropodal

exopod.

Alloschizidium remyi (Vandel, 1944) n. comb.

(Figs 23, 24, 27)

Paraschizidium Remyi Vandel, 1944a: 84, figs I-IV; 1954a: 74; Arcangeli, 1948: 258.

Paraschizidium remyi ; Vandel 1962: 766, figs 367-368.

Material examined. — 19, grotte de Sisco, leg. S. Taiti, S. Vanni and A.M. Nistri, 3.IV.1995; 1 9, Cardo

(W of Bastia), humus under ilex tree, leg. F. Bernini, 19.IV.1980; 2 63, 3 9 9, same locality, leg. S. Taiti and

A. Poggesi, 25.VI. 1984.

Previous records. — Convent of Ste-Catherine-de-Sisco (Vandel 1944a).

Distribution. — Species endemic to Corsica.

Source : MNHN, Paris

— 516 —

Source : MNHN, Paris

-517-

Remarks

On the basis of the presence of a postscutellar line, schisma at the posterior comers of

pereonite 1 and antennule with three articles, this species belongs to the genus Alloschizidium

and not to Paraschizidium. The main diagnostic features of this species, including the previously

unknown male characters, are illustrated in Figs 23, 24.

Alloschizidium cottarellii (Argano & Pesce, 1974) n. comb.

(Figs 25, 27)

Material examined. — 1 9, Punta de la Parala (W of Ajaccio), leg. S. Taiti, 14.V.1982; 2 9 9, 1 juv.,

same locality, leg. S. Taiti and S. Vanni, 8.III. 1994; 5 9 9,1 juv., between Cauro and Bocca di San Giorgio (E

of Ajaccio), ilex grove, leg. S. Taiti, 14.V.1982; 10 S <?, 12 9 9, between Pietrosella and Fogolina (SE of

Porticcio), ilex and arbutus grove, leg. S. Taiti and S. Campanelli, 21.X.1982; 7 9 9, forêt de Valle Mala, near

col de St-Eustache (between Petreto-Bicchisano and Aullène), leg. S. Taiti and S. Campanelli, 22.X. 1982; 1 9,

near Punta di u Carapono (Coti-Chiavari), leg. S. Taiti and S. Campanelli, 21.X.1982; 3 â â , 5 9 9, Capo

Source : MNHN, Paris

— 518 —

Source : MNHN, Paris

-519-

Fig. 26. — Alloschizidium buchnerorum : A, dorsal scale-spine; B. cephalon, frontal; C, left epimeron of pereonite 1, dorsal;

D. telson and uropods; E. antennule; F. antenna.

Pertusato (SE of Bonifacio), leg. S. Taiti, I8.V.1982; 1 9, same locality, leg. S. Taiti and S. Campanelli, 19.X.1982;

1 â, 1 9, same locality, leg. S. Taiti, 29.VI. 1991.

Distribution. — Southern Corsica and northern Sardinia.

Remarks

These specimens correspond well to the description by Argano & Pesce (1974) of

Typhloschizidium cottarellii from Sardinia (Ardara, Sassari).

Due to the presence of a postscutellar line on the cephalon, this species is ascribed to the

genus Alloschizidium. The main diagnostic characters of this species are illustrated in Fig. 25.

A. cottarellii is morphologically similar to A. buchnerorum (Verhoeff, 1941) from Ischia

Island, from which it differs in the longer scale-spines on the dorsum, more broadly rounded

Source : MNHN, Paris

— 520 —

Fig. 27. — Distribution in Corsica of Alloschizidium species: 1. A. campanellii; 2, A. remyr, 3, A. cottarellii.

Source : MNHN, Paris

— 521 —

posterior comers of pereonite 1, and longer telson and uropodal exopod. Re-examination of three

syntypes (9$) of A. buchnerorum , in micropreparations deposited at the Zoologischen Staats-

sammlung, Munich, allowed us to illustrate the main characters of this species (Fig. 26).

Genus ARMADILLIDIUM Brandt, 1833

Armadillidium nasatum nasatum Budde-Lund, 1885

(Fig. 31)

Armadillidium nasatum; Vandel 1954a: 74; 1962: 787, fig. 380.

Material examined. — 1 d, 1 9,1 juv., Folelli, along Fium Alto, leg. P. Magrini, 5.XI.1983; many d d

and 9 9, same locality, leg. S. Taiti and A. Poggesi, 20.V1.1984.

Previous records. — Bonifacio (Vandel 1962).

Distribution. — Central-northern Italy, France, northern Spain, Netherlands, Great Britain and Ireland.

It has also been introduced to the USA and recorded from many hothouses in Europe.

Remarks

These specimens correspond perfectly to the nominal subspecies since they lack dorsal

granulations, the cephalic scutellum is very protruding above the vertex with parallel sides and

upper margin slightly concave, and the indentation at the base of the epimera on the posterior

margin of pereonite 1 is slightly marked.

Armadillidium nasatum sardoum Arcangeli, 1950

(Fig. 31)

Material examined. — 1 <5, Asco river valley, loc. Le Cabanelle, leg. S. Taiti and A. Poggesi, 20.VI. 1984;

13 dd, 6 9 9, Lovo Santo, swamp at mouth of Cavo river, leg. B. Lanza, VIII.1971; 15 dd, 12 9 9, same

locality, leg. S. Taiti, 10.VII.1978.

Distribution. — Corsica and Sardinia.

Remarks

These specimens differ from those of the preceding subspecies in the smaller size (ovigerous

9 13 mm long vs. 17 mm), the uniformly grey colour, the presence of tiny but clearly visible

granulations on the dorsum, the cephalic scutellum less protruding with convergent sides and

upper margin convex. On the basis of those characters they correspond to A. nasatum sardoum

described by ARCANGELI (1950) from several localities in Sardinia.

Armadillidium granulatum Brandt, 1833

Armadillidium granulatum; DOLLFUS 1899: 186; ARCANGELI 1914: 467; Vandel 1954a:

74; 1962: 796, fig. 383; Lanza & Poggesi 1986: 120, 177.

Source : MNHN, Paris

— 522 —

Material examined. — Many â 6 and 9 9 collected from the following localities: Centuri-Port; marine

de Pietracorbara; Miomo (N of Bastia); Fautea (between Solenzara and Porto-Vecchio); between Bonifacio and

col d’Arbia; Bonifacio; Capo Pertusato (SE of Bonifacio); île de Pinarello.

Previous records. — Bonifacio, Bastia (Dollfus 1899); “Ghisernia laterni” (Arcangeli 1914); Corsica

(Vandel 1954a, 1962); île de Pinarello (Lanza & Poggesi 1986).

Distribution. — Littoral species distributed along all the coasts of the Mediterranean, the Atlantic coasts

of Morocco, Portugal, France, Madeira and the Azores.

Armadillidium sordidum Dollfus, 1887

(Fig. 31)

Armadillidium sordidum', DOLLFUS 1892: 176-177; 1896: 358; 1899: 187; ARCANGELI

1925: 8; Vandel 1954a: 74; 1962: 816, figs 392-393.

Armadillidium (Armadillidium) sordidum sordidum-, ARCANGELI 1954b: 155, pis 15-16.

Material examined. — Many ââ and 9 9 collected from the following localities: Camera (near Centuri);

near Meria; between Morsiglia and Pastina; 2 km SE of col de Ste-Lucie (between Pino and Luri); Carbonacce

(S of Luri); Selmacci (SW of Pietracorbara); St-Léonard (W of marine de Pietracorbara); between Bastia and

St-FIorent; Bevinco river valley, below col de S. Stefano; Urtaca; near Murato; lower valley of Asco river; 3

km S of Ponte Leccia, along left bank of Golo river; Folelli, along Fium Alto; Francardo; 1.6 km down from

Stazzona (near Piedicroce).

Previous records. — Bastia and Porto-Vecchio (?) (Dollfus 1892, 1899; Arcangeli 1925); Orezza (Dollfus

1899).

Distribution. — This species has a typical Tyrrhenian distribution. It is found in southern France (Maures

massif), Corsica, Sardinia, Liguria, Tuscany and Umbria.

Remarks

In Corsica this species is common on the north-eastern part of the island (in the so-called

“Alpine Corsica”), while it seems to be absent in the granitic part. Thus, the record from Porto-

Vecchio by Dollfus (1892) is doubtful and most probably is due to a misidentification.

Armadillidium assimile Budde-Lund, 1885

Armadillidium Zenckeri ; DOLLFUS 1887: 9; ARCANGELI 1914: 461.

Armadillidium esterelanum; DOLLFUS 1892: 139; 1896: 358; SCHARFF 1894: 163.

Armadillidium vizzavonense Verhoeff, 1926: 263; 1928: 121; ARCANGELI 1950: 80; Vandel

1953a: 164.

Armadillidium vizzavonensis; Vandel 1954a: 74.

Armadillidium assimile', Vandel 1962: 803, fig. 386; Beron 1972a: 13.

Material examined. — Many 6 S and 9 9 collected from the following localities: Barcaggio (Cap Corse);

near Rogliano; Centuri-Port; between Macinaggio and marine de Meria; near Meria; near Carbonacce (S of Luri);

marine de Giottani (between Nonza and Centuri-Port); marine de Pietracorbara; near convent of Sta Catterina

de Sisco; near Crosciano along Sisco river; Cardo (W of Bastia); anse de Faggiola (désert des Agriates); St-

FIorent; Casta (SW of St-Florent); Olmeta-di-Tuda; col de S. Stefano (S of Olmeta-di-Tuda); défilé de Lancone,

between Casatorra and Oletta; left bank of Golo river, between Barchetta and Casamozza; near grotte de Pietral-

bello (Moltifao); Asco river valley; plage de Caspio (NW of Porto); 4 km E of Zonza; Fautea (between Solenzara

Source : MNHN, Paris

— 523 —

and Porio- Vecchio); Lovo Samo, swamp ai mouth of Cavo river; Taglio Rosso (W of Sta-Lucia-di-Porto- Vecchio);

Golfo di Sogno; near Porto-Vecchio; Monte Rosso (SE of Sartène); dolmen de Fontanaccia (S of Sartène); Tizzano

and environs (SW of Sartène); near Ermitage de la Trinité (NW of Bonifacio).

Previous records. — Bastia (Dollfus 1892; Scharff 1894; Verhoeff 1928); Corte, Vico. La Solenzara,

I'Ospedale, Porto-Vecchio (Dollfus 1892); Vizzavona (Dollfus 1892; Verhoeff 1928; Arcangeli 1950); between

Bastia and S. Martino (Verhoeff 1926; Arcangeli 1950); Île-Rousse (Verhoeff 1928; Arcangeli 1950); grotte

de Carpiniccia (Pietroso) (Vandel 1953a; Beron 1972a); all Corsica (Vandel 1962).

Distribution. — Beside Corsica, it is known from Sardinia, Tuscany, central-southern France, eastern and

southern Spain. Portugal and the Azores.

Armadillidium lanzai n.sp.

(Figs 28, 31)

Armadillidium lanzai (nomen nudum); Lanza & POGGESI 1986: 42, 55, 120, 175, 177, 179,

183.

Material examined. — 1 6 holotype (MZUF), 38 <5 <5, 52 9 9 paratypes (MZUF), 2 6 6,2 9 9 paratypes

(MNHN-Is4101), îlot Toro Grande, leg. S. Taiti, 17.V. 1982; 1 9 paratype (MZUF), Fautea (between Solenzara

and Porto-Vecchio), leg. B. Conti and B. Lanza, VII-VIII.1977; 2 6 6, 2 9 9 paratypes (MZUF), same locality,

leg. S. Taiti, 17.V.1982; 1 6 paratype (MZUF), same locality, leg. S. Taiti and S. Campanelli, 19.X.1982; 3 6 6,

1 9 paratypes (MZUF), île de la Giraglia, leg. R. Brizzi, R. Innocenti and S. Turillazzi, 30.IX.1972; 3 9 9

paratypes (MZUF), same locality, leg. R. Brizzi and R. Pirozzi, 1.VI. 1972; many 6 6 and 9 9 paratypes (MZUF),

same locality, leg. S. Taiti and A. Poggesi, 26.VI.1984: 1 6 paratype (MZUF), île de Pinarello. leg. B. Conti

and N. Lanza. I0.VIII.1973; 2 66, 3 9 9 paratypes (MZUF), îlot Toro Piccolo, leg. S. Campanelli, 17.V.1982;

13 6 6 , 23 9 9 paratypes (MZUF), îlot Luigi Giafferri (Lavezzi), leg. B. Lanza, 6.VIII. 1974.

Etymology. — The new species is named after our friend Prof. B. Lanza, Florence, in recognition of his

valuable contribution to the knowledge of the Corsican fauna.

Description

Maximum dimensions: <3, 11.5 x 4.4 mm; 9, 13 x 5.5 mm.

Colour variable according to the population: usually grey-brown with large pale spots,

irregularly arranged; the specimens from îlot Toro Grande and îlot Toro Piccolo have a yellowish

ground colour which blends with the pale spots. Dorsum with light granulations. Animals able

to roll up into a ball, mesosphaeric type, with anterior comers of pereonite 1 slightly bent up.

Eye with 18-20 ommatidia. Cephalon with wide scutellum, slightly protruding and not bent over

the vertex, anteriorly convex and posteriorly strongly concave, so that a deep fossette is visible

between the scutellum and the anterior margin of the vertex; upper margin of the scutellum

slightly concave; antennary lobes quadrangular, protruding frontwards, with antennary grooves

shallow. Pereonite 1 with posterior margin regularly sinuous; lateral margin slightly impressed

in the posterior third. Telson somewhat broader than long, trapezoidal, with more or less narrow

apex and posterior comers rounded.

Male

Pereopod 7 ischium with sternal margin almost straight with some small verrucae, rostral

surface with an area of short setae; merus with a small tuft of short setae on the basal part of

the rostral surface. Pleopod 1 exopod with a long posterior point, medial margin regularly curved

with short spines.

Source : MNHN, Paris

— 524 —

FIG. 28. — Armadillidium lanzai n.sp., specimens from îlot Toro Grande: A. cephalon, frontal; B, cephalon, dorsal; C. cephalon,

from back; D, cephalon and pereonite 1, lateral; E, telson and uropods; F. <5 pereopod 7; G, 6 pleopod 1.

Source : MNHN, Paris

— 525 —

Remarks

The new species shows morphological affinities with Armadillidium quinquepustulatum

Budde-Lund, 1885, originally described from Algeria and later recorded from the Maures Massif

in southern France, and with A. maculatum Risso, 1816, from the Maritime Alps. It is distin¬

guished from the former by the different colour pattern, the more developed scutellum, and the

male pereopod 7 ischium more thickset, with no spines on tergal margin and sternal margin less

concave with no brushes of setae. It differs from the latter in the colour pattern, the scutellum

shorter and not bent over the vertex, telson with wider apex, uropodal exopod shorter and the

male pleopod 1 exopod with much more developed posterior point.

The record of this species from an islet near Sardinia (Argano & Manicastri 1991) is

due to a misidentification (Manicastri pers. com.).

Armadillidium vulgare (Latreille, 1804)

Armadillidium vulgare ; DOLLFUS 1899: 187; VERHOEFF 1926: 263; Vandel 1954a: 74;

Beron 1972a: 13; Lanza & Poggesi 1986: 120, 183.

Material examined. — Many 6 6 and 9 9 collected from the following localities: Tollare and Barcaggio

(Cap Corse); Centuri-Port; near Meria; St-Léonard (W of marine de Pietracorbara); marine de Pietracorbara; Sta

Catterina de Sisco; Bastia; anse de Faggiola and désert des Agriates; Casta (SW of St-Florent); défilé de Lancone;

col de S. Stefano (S of Olmeta-di-Tuda); Urtaca; near Murato; col de S. Colombano (E of Belgodère); 1 km W

of Altiani (SE of Speloncato); Asco river valley, S of Moltifao; Folelli, along Fium Alto; near grotte de Sahara

(Castiglione); Francardo; Prunete (E of Cervione); plage de Caspio (NW of Porto); 2 km S of Corte; Ghisoni;

col de Verde (S of Ghisoni); Ghisonaccia; 1 km W of Tolla (Prunelli river valley); Porticcio; Monte Incudine;

col de Bavella; Taravo river valley; Conca (N of Ste-Lucie-de-Porto-Vecchio); Fautea (between Solenzara and

Porto- Vecchio); Lovo Santo, swamp at mouth of Cavo river; valley below chapelle Pianelli, between Casalabriva

and Olmeto; mouth of Rizzanèse river, S of Propriano; Monte Rosso, SE of Sartène; la Trinité (N of Porto-

Vecchio); Porto-Vecchio; between Bonifacio and col d’Arbia; near ermitage de la Trinité, W of Bonifacio; Boni¬

facio; Capo Pertusato (SE of Bonifacio); îlot Sperduto Piccolo (E of île Cavallo); île Lavezzi; îlot Cala di u

Ghiuncu (île Lavezzi).

Previous records. — Between Bastia and Santa Lucia, Santa Reparata, Gravona (Ajaccio) (Verhoeff 1926);

grotte Roumandella (Caporalino) (Beron 1972a); îlot Cala di u Ghiuncu (Lanza & Poggesi 1986).

Distribution. — Cosmopolitan species.

Armadillidium album Dollfus, 1887

(Fig. 31)

Armadillidium album ; BIGOT, 1977: 7, 12.

Material examined. — 8 Ô d, 2 9 9, Calzarello, near Ghisonaccia, beach, leg. S. Taiti and S. Campanelli,

18.X. 1982.

Source : MNHN, Paris

— 526 —

Previous records. — Eastern coast of Corsica (Bigot 1977).

Distribution. — This halophilic species is widespread along the coasts of the Mediterranean and the Atlantic

coasts of Europe.

Armadillidium littorale n.sp.

(Figs 29, 31)

Material examined. — 1 6 holotype (MZUF), many <5 <5 and 9 9 paratypes (MZUF), 2 6 6,2 9 9

paratypes (MNHN-Is4100), Capo Pertusato (SE of Bonifacio), under stones near beach, leg. S. Taiti and

S. Campanelli, 19.X.1982; 5 9 9 paratypes (MZUF), same locality, leg. S. Taiti, 18.V.1982; 1 9 paratype

(MZUF), same locality, leg. S. Taiti, 29.VI. 1991.

Etymology. — L. litioralis = littoral. The name of the species refers to the habitat where the specimens

were collected, i.e. under stones beyond the upper limit of the beach.

Description

Maximum dimensions: S and $ 4 x 1.5 mm.

Brown colour with epimera and muscle spots pale. Dorsum rough, with numerous tiny triangular

scale-spines. Animals able to roll up into a perfect ball, eusphaeric type. Eye with 10-12 om-

matidia. Cephalon with postscutellar and frontal lines, the former clearly visible only in

the medial part; scutellum slightly protruding over the vertex, with upper margin con¬

tinuing the frontal line, and anterior surface distinctly hollow; antennary lobes oblique,

neither thickened nor bent backwards. Pereonite 1 with posterior margin regularly con¬

cave; lateral margin slightly swollen and grooved. Telson trapezoidal, much wider than

long, with posterior corners broadly rounded. Antenna short and thickset, with second

flagellar article about three times as long as the first. Pleopods 1-2 exopod with an in¬

dentation on outer margin of the tracheal field. Uropodal exopod much wider than long.

Male

Pereopods with no particular modifications. Pleopod 1 exopod with rounded posterior margin

and no posterior point; endopod with stout distal part, apex slightly bent outwards. Pleopod 2

as in Fig. 29H.

Remarks

Armadillidium littorale is morphologically close to A. album, from which it is readily dis¬

tinguished by the different shape of the scale-spines, the lateral margin of pereonite 1 slightly

swollen and grooved, no modifications on the male pereopod 7 and the male pleopod 1 exopod

with no posterior point.

Source : MNHN, Paris

— 527 —

Source : MNHN, Paris

— 528 —

Source : MNHN, Paris

— 529 —

Source : MNHN, Paris

— 530 —

Armadillidium torchiai n.sp.

(Figs 30, 31)

Material examined. — 1 6 holotype (MZUF). 1 3 paratype (MZUF), forêt de Vizzavona, beech wood,

1000-1400 m, leg. A. Torchia and S. Zoia, 27-28.V.1982; 1 3 paratype (MV), Vizzavona, leg. M. Meregalli,

25.VIII.1982.

Etymology. — The new species is named after one of its collectors, Mr A. Torchia, Genoa.

Description

Male

Maximum dimensions: 3.5 x 1.2 mm.

Brown colour with pale muscle spots. Dorsum rough with many small triangular scale-spines.

Animals able to roll up into a perfect ball, eusphaeric type. Eye small, consisting of about eight

ommatidia. Cephalon with postscutellar and frontal lines, the former visible for almost its entire

length; scutellum slightly protruding above vertex, with upper margin continuing the frontal line

and anterior surface distinctly concave; antennary lobes oblique, neither thickened nor bent back¬

wards. Pereonite 1 with posterior margin regularly concave, lateral margin very slightly swollen.

Telson wider than long, triangular with very broadly rounded apex. Antenna short and stout,

with second flagellar article about three times longer than first. Pereopods without particular

modifications. Pleopod 1-2 exopods with an indentation on the outer margin of the tracheal

field. Pleopod 1 exopod with a short posterior point; endopod with thickset distal part, straight.

Pleopod 2 as in Fig. 30G. Uropodal exopod trapezoidal, wider than long.

Remarks

Armadillidium torchiai is morphologically very close to A. littorale , from which it differs

in having the lateral margin of pereonite 1 less swollen, the shape of the telson and the male

pleopod 1 exopod with a short posterior point.

DISCUSSION

With the data from the present analysis, seventy-six species of terrestrial isopods are known

from Corsica, of which eight are described as new and twelve are newly recorded (marked with

* in the following list):

Source : MNHN, Paris

— 531 —

1) Tylos europaeus Arcangeli, 1938 38)

2) Tylos ponticus Grebnitzky, 1874 *39)

3) Helleria brevicornis Ebner, 1868 40)

4) Ligia iialica Fabricius, 1798 41)

5) Finaloniscus briani Vandel, 1953 *42)

*6) Finaloniscus franciscoloi (Brian, 1951) 43)

7) Nesioioniscus corsicus Racovitza, 1908 *44)

8) Nesioioniscus racovitzai Vandel, 1954 45)

9) Oritoniscus paganus Racovitza, 1908 *46)

10) Oritoniscus ocellatus Vandel, 1953 47)

*11) Oritoniscus punctatus n.sp. 48)

12) Trichoniscus pusillus provisorius *49)

Racovitza, 1908 50)

13) Trichoniscus pygmaeus Sars, 1899 51)

14) Trichoniscus fragiiis Racovitza, 1908 52)

*15) Trichoniscus halophilus Vandel, 1951 53)

16) Trichoniscus pedronensis Vandel, 1947 54)

17) Haplophthalmus danicus Budde-Lund, 1885 55)

18) Cyrnoniscus remyi Vandel, 1953 56)

19) Carioniscus dollfusi (Carl, 1908) 57)

20) Buddelundiella cataractae Verhoeff. 1930 58)

21) Buchneriilo litoralis Verhoeff, 1942 *59)

22) Sienoniscus pleonalis Aubert & Doll- 60)

fus, 1890 61)

23) Sienoniscus carinatus Silvestri, 1897 62)

24) Arniadilloniscus Candidas Budde-Lund, 1885 63)

*25) Arniadilloniscus elliplicus (Harger, 1878) *64)

26) Halophiloscia couchii (Kinahan, 1858) 65)

27) Halophiloscia hirsuta Verhoeff, 1928 *66)

*28) Halophiloscia ischiana Verhoeff, 1933 67)

*29) Halophiloscia lyrrhena Verhoeff, 1928 *68)

*30) Sienophiloscia zosterae Verhoeff, 1928 69)

31) Chaetophiloscia elongata (Dollfus, 1884) 70)

32) Chaetophiloscia sicula Verhoeff, 1908 71)

33) Chaetophiloscia cellaria (Dollfus, 1884) *72)

*34) Ctenoscia dorsalis Verhoeff, 1928 73)

35) Ctenoscia minima (Dollfus, 1892) 74)

36) Philoscia affinis Verhoeff, 1908 *75)

37) Tiroloscia Corsica Corsica (Dollfus, 1888) *76)

Tiroloscia macchiae Verhoeff, 1931

Tiroloscia montana n.sp.

Sardoniscus pygmaeus (Budde-Lund. 1885)

Platyarthrus costulatus Verhoeff, 1908

Platyarthrus corsicus n.sp.

Platyarthrus caudatus Aubert & Dollfus, 1890

Platyarthrus aiasensis Legrand, 1953

Platyarthrus schoeblii schoeblii Budde-Lund, 1885

Platyarthrus hoffmannseggii Brandt, 1833

Cylisticus convexus (De Geer, 1778)

Cylisticus vandeli Taiti & Ferrara, 1980

Cylisticus uncinatus n.sp.

Troglocylisticus cyrnensis Ferrara & Taiti, 1983

Protracheoniscus babori Frankenberger, 1938

Porcellionides pruinosus (Brandt, 1833)

Porcellionides sexfasciatus sexfasciatus (Budde-Lund, 1885)

Acaeroplastes melanurus sardous Verhoeff, 1918

Leptotrichus panzerii (Audouin, 1826)

Agabiformius lentus (Budde-Lund, 1885)

Lucasius pallidus (Budde-Lund, 1885)

Porcellio spatulatus Costa, 1882

Porcellio scaber Latreille, 1804

Porcellio dilatatus dilatatus Brandt. 1833

Porcellio laevis Latreille, 1804

Porcellio or arum vizzavonensis Verhoeff, 1928

Porcellio lamellatus sphinx Verhoeff, 1931

Alloschizidium campanellii n.sp.

Alloschizidium remyi (Vandel, 1944)

Alloschizidium cottarellii (Argano & Pesce, 1974)

Armadillidium nasatum nasal urn Budde-Lund, 1885

Armadillidium nasatum sardoum Arcangeli, 1950

Armadillidium granulatum Brandt, 1833

Armadillidium sordidum Dollfus, 1887

Armadillidium assimile Budde-Lund. 1885

Armadillidium lanzai n.sp.

Armadillidium vulgare (Latreille, 1804)

Armadillidium album Dollfus, 1887

Armadillidium littorale n.sp.

Armadillidium torchiai n.sp.

According to their present distributions, the species can be grouped in the following

categories:

1. Cosmopolitan species (5)

Porcellionides pruinosus, Agabiformius lentus, Porcellio scaber, P. laevis and Armadillidium

vulgare. All the species are of Mediterranean origin, with the sole exception of P. scaber, of

West-European origin and most probably introduced to Corsica.

Source : MNHN, Paris

— 532 —

2. European species (8)

Trichoniscus pusillus provisorius, T. pygmaeus, Haplophthalmus danicus, Buddelundiella

cataractae, Platyarthrus hoffmannseggii, Cylisticus convexus, Porcellio dilatants dilatants and

Armadillidium nasatum nasatum.

3. Mediterranean-Atlantic species (9)

Tylos europaeus, T. ponticus, Ligia italica, Trichoniscus fragilis, Armadilloniscus ellipticus,

Halophiloscia couchii, Stenophiloscia zosterae, Armadillidium granulatum and A. album.

4. Holomediterranean species (2)

Chaetophiloscia elongata and Leptotrichus panzerii.

5. West-Mediterranean-Atlantic species (12)

Buchnerillo litoralis, Stenoniscus pleonalis , S. carinatus, Armadilloniscus candidus, Ctenos-

cia dorsalis, C. minima, Platyarthrus costulatus, P. aiasensis, Porcellionides sexfasciatus

sexfasciatus, Lucasius pallidus, Porcellio lamellatus sphinx and Armadillidium assimile.

6. West-Mediterranean species (8)

Trichoniscus halophilus, Halophiloscia hirsuta, H. ischiana, H. tyrrhena, Chaetophiloscia

cellaria, Philoscia affinis, Platyarthrus caudatus and P. schoeblii schoeblii.

7. North-Mediterranean species (2)

Chaetophiloscia sicula and Protracheoniscus babori.

8. Alpine species (1)

Carloniscus dollfusi.

9. Tyrrhenian species (29)

Helleria brevicornis, Finaloniscus briani, F. franciscoloi, Nesiotoniscus corsicus, N. racovit-

zai, Oritoniscus paganus, O. ocellatus, O. punctatus, Trichoniscus pedronensis, Cyrnoniscus

remyi, Tiroloscia Corsica Corsica, T. macchiae, T. montana, Sardoniscus pygmaeus, Platyarthrus

corsicus, Cylisticus vandeli, C. uncinatus, Troglocylisticus cyrnensis, Acaeroplastes melanurus

sardous, Porcellio spatulatus, P. orarum vizzavonensis, Alloschizidium campanellii, A. remyi,

A. cottarellii, Armadillidium nasatum sardoum, A. sordidum, A. lanzai, A. littorale and A. torchiai.

This last group of species can be further divided according to their distributions:

9a. Corsican endemic species (18)

Finaloniscus briani, Nesiotoniscus corsicus, N. racovitzai, Oritoniscus paganus, O. ocellatus,

O. punctatus, Trichoniscus pedronensis, Cyrnoniscus remyi, Tiroloscia montana, Platyarthrus

corsicus, Cylisticus vandeli, C. uncinatus, Troglocylisticus cyrnensis, Alloschizidium campanellii,

A. remyi, Armadillidium lanzai, A. littorale and A. torchiai. Almost one quarter of the species

Source : MNHN, Paris

— 533 —

recorded from Corsica are exclusive to the island. However, the endemism is generally weak,

as most of the species are closely related to taxa present in the surrounding area (Mediterranean

France, the Pyrénées, the Tuscan Archipelago and Tuscan coastal hills, and Sardinia). Two mo¬

nospecific genera ( Cyrnoniscus and Troglocylisticus) appear to be exclusive to Corsica. They

are very specialised endogean and/or cavemicolous taxa. The former is related to taxa present

in the northern Mediterranean area, and the latter to the nasutus- group of the genus Cylisticus

widely distributed in the Tyrrhenian area.

9b. Corsican-Sardinian endemic species (4)

Tiroloscia Corsica Corsica, Porcellio orarum vizzavonensis, Alloschizidium cottarellii and

Armadillidium nasatum sardoum.

9c. North-Tyrrhenian species (5)

Helleria brevicornis, Tiroloscia macchiae, Sardoniscus pygmaeus, Acaeroplastes melanurus

sardous and Armadillidium sordidum.

9d. Tyrrhenian ( sensu lato) species (2)

Finaloniscus franciscoloi and Porcellio spatulatus.

About 60% of the species have wide distributions (groups 1 to 7), almost all of Mediter¬

ranean origin. Seventeen of these species are halophilic (species 1, 2, 4, 14, 15, 21-30, 63 and

74 on the list), and can be considered as real expansive species due to their great facility for

dispersal. All are widespread along the coast of the Mediterranean Sea and some also occur on

the Atlantic coasts of Africa as far south as the river Senegal.

The number of species with a Tyrrhenian distribution is particularly high ( ca 38%) and this

group is the most interesting from a zoogeographic point of view. In fact, it consists of endemic

endogean, cavemicolous or montane taxa, with a low capacity for dispersal, which provide valu¬

able information about the faunistic affinities of Corsica. Indeed a comparison of the oniscidean

fauna of Corsica with that of the surrounding regions [Sardinia (Argano et al. 1995); Tuscan

Archipelago and coastal hills (Taiti & Ferrara 1989a and unpublished data), Sicily and

circum-Sicilian islands (CARUSO et al. 1987), mainland Italy (ARGANO et al. 1995), southern

France (Vandel 1960, 1962), Iberian Peninsula (SCHMÔLZER 1971) and North Africa (Vandel

1955, 1958, 1959)] leads to the following considerations.

In Sardinia the presence of many significant species (Corsican-Sardinian and North-

Tyrrhenian elements, vicarious taxa of the genera Oritoniscus, Cylisticus of the nasutus- group,

and Nesiotoniscus ) demonstrates the close faunistic relationship of this island with Corsica. Be¬

sides the presence of characteristic taxa, a low number of species of the genera Porcellio and

Armadillidium in both Sardinia and Corsica must be stressed. Only five species of Porcellio are

known from these islands, mainly widespread or cosmopolitan in distribution, with the exception

of P. orarum vizzavonensis and P. spatulatus. The latter has probably been recently introduced

to Corsica if we consider its limited distribution in the southern coastal area of the island. The

genus Armadillidium is present in the two islands with ten species that are mainly widespread,

except A. sordidum and the three new species; however the characteristic forms of the Italian

Source : MNHN, Paris

— 534 —

peninsula and Sicily are completely missing. It is important to note that in Italy this genus has

very many species, with over sixty forms known at present. Thus, Corsica, like Sardinia, seems

to have been excluded from the spéciation phenomena and colonizations of these two genera

which occurred in Italy. Besides the clear affinities between the two islands, some important

differences must also be pointed out. In fact, several characteristic taxa of the Sardinian fauna

are missing in Corsica, i.e. species of the genera Scotoniscus Racovitza, 1908, Catalauniscus

Vandel, 1953, Alpioniscus Racovitza, 1908, Cordioniscus Graeve, 1914, Trichophiloscia Arcan-

geli, 1950, Tritracheoniscus Taiti & Manicastri, 1985, and the species Armadillo officinalis

Duméril, 1816; on the contrary, not present in Sardinia are species of the genera Cyrnoniscus,

Carloniscus and Troglocylisticus.

The Tuscan Archipelago and coastal hills have numerous species of great zoogeographic

value in common with Corsica: the North-Tyrrhenian elements, Finaloniscus franciscoloi,

Armadillidium assimile , and closely related species of the genera Nesiotoniscus, Oritoniscus,

Tiroloscia, Cylisticus and Alloschizidium. Likewise, zoogeographically important species present

in Sardinia but not in Corsica are also missing in this area.

As far as Sicily is concerned, there are no significant examples of faunistic affinities with

Corsica, except for the species with a wide distribution, Finaloniscus franciscoloi, and a species

of Nesiotoniscus (A/. helenae Brisolese & Caruso, 1974), which, however, does not belong to

the corsicus-group.

With regard to a comparison with mainland Italy, all the typical Italian Alpine-Apennine

elements are missing in Corsica, e.g. species of the genera Spelaeonethes Verhoeff, 1932,

Androniscus Verhoeff, 1908, Alpioniscus, Haplophthalmus and Buddelundiella (with the exception

of H. danicus and B. cataractae, certainly introduced), Orthometopon Verhoeff, 1917, Trachelipus

Budde-Lund, 1908, Armadillidium of the depressum-, carniolense-, alassiense-, furcatum- and

vulgare- groups. Moreover, species of Helleria, Oritoniscus, Nesiotoniscus and Alloschizidium pre¬

sent in Corsica are absent along the Alpine and Apennine chains.

In southern France (Maures and Estérel massifs, and the Grasse region) there are several

zoogeographically important species which are in common with or closely related to forms present

in Corsica: Helleria brevicornis, Porcellio orarum, Armadillidium sordidum, species of Orito¬

niscus, Nesiotoniscus, Tiroloscia and Cylisticus.

Oritoniscus paganus, O. ocellatus and O. punctatus from Corsica show close relationships

with species of the same genus from the Pyrénées, while the genus Tiroloscia is present in

Corsica with T. Corsica Corsica, T. macchiae and T. montana and in the Pyrénées with T. pyrenaica

(Dollfus, 1897).

Even though the terrestrial isopods from North Africa are poorly known, it is interesting

to point out the presence there of species of the genera Finaloniscus ( F '. berberensis) and

Nesiotoniscus (N. delamarei Vandel, 1954 and N. sebaouensis Vandel, 1955).

With regard to the distribution of the species within Corsica, ten species populate exclusively

the north-eastem part of the island, i.e. the so-called “Alpine Corsica” ( Oritoniscus ocellatus,

Trichoniscus pedronensis, Cyrnoniscus remyi, Ctenoscia dorsalis, C. minima, Tiroloscia mac¬

chiae, Cylisticus convexus, Troglocylisticus cyrnensis, Alloschizidium remyi and Armadillidium

sordidum), while twelve species occur only in the granitic part ( Nesiotoniscus corsicus, N. raco-

vitzai, Oritoniscus punctatus, Tiroloscia montana, Platyarthrus corsicus, P. aiasensis, L. pallidus,

Source : MNHN, Paris

— 535 —

Table 1. — Distribution of terrestrial isopods in circum-Corsican islands.

{

j !

| |

Ile de la Giraglia

•

• •

•

Hot Terre

•

Ilot Intermediaire

•

Hot Finocchiarola

Hot Fautca

•

Hot dc la Roscana

•

lie de Pinarello

•

He de Comuta

•

Rocher de Vacca

•

He Pietricaggiosa

•

Hot du Torello

•

Hot Toro Grande

•

Hot Toro Piccolo

•

Hot La Folaca

•

Rocher d'Acciaju Nord

•

Ilot Porraggia Piccola

Hot Porraggia Grande

•

Hot Sperduto Grande

•

Hot Sperduto Piccolo

•

Rocher Sud dc Ratino

•

He Cavallo

•

Hot Luigi Giafferri

•

Hot Gian Pietro GalTori

•

Hot Silcnc

•

Hot Cala di u Ghiuncu

•

lie Lavezzi

•

lie Saint Antoine

•

Hot Fazzuolo Piccolo

•

Ilot Sud dc la Tonnara

•

Grand îlot des Bruzzi

•

Ile Piana de Portigliolo

•

He Mezzu Marc

Ile de Cala d‘ Alga

Ilot de Cala Maiora

Ilot Guardiola

——

Ilot Garganellu

Hot Palazzu

Hot Palazzinu

Zcccu d’a Furmicula

Hot Porri

Rochers d'Elpa Nera

Ilot Nord de Morsetta

Ile Spano

•

Source : MNHN, Paris

— 536 —

P. spatula tus, P. scaber, Alloschizidium campanellii, A. cottarellii and Armadillidium torchiai).

One species ( Armadillidium littorale ) has been collected only in the Miocenic molasse of Capo

Pertusato, near Bonifacio, and three species are exclusive to montane areas above 1000 m altitude

(Trichoniscus pedronensis in Monte San Petrone, Tiroloscia montana in the Haut-Asco area and

Armadillidium torchiai in the Vizzavona forest).

As concerns the oniscidean fauna of the islands surrounding Corsica (Lanza & POGGESI

1986), seventeen species are presently known from forty-three islets (Table 1). However, we

must point out that, except for the île de la Giraglia, the île Pietricaggiosa and the îlots Toro

Grande and Toro Piccolo which have been thoroughly investigated, most of the data derive from

occasional collectings. Most of the species are littoral or widespread eurytopic ones and occur

also in Corsica, with the exception of Porcellio spatulatus, which populates only the southernmost

islets and the southern coast of the main island, and Armadillidium lanzai, recorded from the

île de la Giraglia in the North and the île de Pinarello, the îlots Toro Grande, Toro Piccolo,

Luigi Giafferri (Lavezzi) and the coast near Fautea in the South.

Twenty-four species are presently known from Corsican caves, most of which must be con¬

sidered subcavernicolous (RUFFO 1955), i.e. trogloxenous, which live near the cave entrances

(twelve species: Helleria brevicornis, Ligia italica, Stenoniscus pleonalis, Halophiloscia hirsuta,

Ctenoscia minima, Philoscia affinis, Platyarthrus costulatus, Porcellionides pruinosus, P. sex-

fasciatus, Porcellio laevis, Armadillidium assimile and A. vulgare) or subtroglophilic, which nor¬

mally live in caves but do not show any peculiar adaptation to cave life (five species: Trichoniscus

fragilis, Chaetophiloscia cellaria, Cylisticus vandeli, Porcellio dilatatus dilatatus and Alloschizidium

remyi). The remaining seven species are eucavemicolous (Ruffo 1955), five of them being eutro-

glophilic ( Finaloniscus briani, Nesiotoniscus corsicus, Oritoniscus paganus, O. ocellatus and

Oritoniscus sp.) and two troglobitic ( Cyrnoniscus remyi and Troglocylisticus cyrnensis). It is

important to point out that all the eucavemicolous species are endemic to Corsica.

GENERAL CONCLUSIONS

In synthesis, with the present study of the terrestrial isopods from Corsica, the following

conclusions can be drawn:

Seventy-six species are known from the island, including eight new species and twelve

taxa recorded for the first time. This number of species is certainly conspicuous (for comparison,

sixty-six species are recorded from Sardinia, seventy-two from the Tuscan Archipelago and about

eighty from Sicily).

About half of the species are West-Mediterranean in distribution, the majority being strictly

Tyrrhenian (38%).

Many endemic species are present (about 25% of the total), which demonstrates the

importance of Corsica as a spéciation area.

The oniscidean fauna of Corsica is closely related to that of the Tuscan Archipelago and,

to a lesser extent, to that of Sardinia.

Source : MNHN, Paris

— 537 —

Some important faunistic affinities exist with southern France (massifs of Maures and

Estérel, and the Grasse region) and the Pyrénées.

No species of southern origin (Sicilian or North African) has been collected in Corsica,

which seems to exclude a zoogeographic relationship with those regions.

Key to species of terrestrial Isopods from Corsica

1. Able to roll up into a ball. 2

— Unable to roll up into a ball. 23

2. Epimera of pereonites 2-7 separated from tergites . 3

— Epimera of pereonites fused with tergites. 5

3. Pleonal tergites fused with each other. Helleria brevicornis

— Pleonal tergites separated. 4

4. Ventral plates of pleonite 5 apically rounded . Tylos europaeus

— Ventral plates of pleonite 5 apically truncate. Tylos ponticus

5. Exoantennal rolling up ability. 6

— Endoantennal rolling up ability. 9

6. Eye absent; maxillular inner branch with 6-8 penicils. Troglocylisticus cyrnensis

— Eye present; maxillular inner branch with 2 penicils. 7

7. Pigmented body; eye with 20-25 ommatidia . Cylisticus convexus

— Colourless body; eye reduced with maximum 10 ommatidia . 8

8. Male pereopod 7 merus with a distinct triangular process . Cylisticus uncinatus

— Male pereopod 7 merus without any process. Cylisticus vandeli

9. Antennal flagellum of 3 articles not clearly distinct; uropodal exopod cylindrical ... 10

— Antennal flagellum of 2 distinct articles; uropodal exopod lamellar. 11

10. Uropods not completely covered by the telson, clearly visible in dorsal view.

. Buddelundiella cataractae

— Uropods completely covered by the telson, not visible in dorsal view .. Buchnerillo litoralis

11. Pereonite 1 with a notch (schisma) at the posterior corner. 12

— Pereonite 1 without a notch at the posterior comer. 14

12. Cephalic scutellum with no upper margin, fused with vertex- Alloschizidium cottarellii

— Cephalic scutellum clearly separated from vertex. 13

13. Dorsum covered with long piliform scale-spines. Alloschizidium remyi

— Dorsum with inconspicuous triangular scale-spines. Alloschizidium campanellii

14. Cephalon with postscutellar line and no frontal line . 15

— Cephalon with both postscutellar and frontal lines. 21

15. Telson triangular. 16

— Telson trapezoidal. 19

16. Posterior margin of pereonite 1 with a deep incision at the base of epimeron. 17

— Posterior margin of pereonite 1 regularly concave, not incised- Armadillidium assimile

17. Telson with pointed apex; male pereopod 7 ischium with distinct verrucae on sternal

margin. Armadillidium granulatum

— Telson with rounded apex; male pereopod 7 ischium without verrucae on sternal margin. . 18

Source : MNHN, Paris

— 538 —

18. Dorsum smooth; cephalic scutellum protruding above vertex about as much as its width

. Armadillidium nasatum nasatum

— Dorsum granulated; scutellum protruding above vertex about half as much as its width

. Armadillidium nasatum sardoum

19. Male pereopod 7 ischium distinctly enlarged distally. Armadillidium sordidum

— Male pereopod 7 ischium not enlarged distally . 20

20. Cephalic scutellum bent over vertex, not protruding; no fossette between scutellum and vertex

. Armadillidium vulgare

— Cephalic scutellum straight, distinctly protruding above vertex; fossette between scutellum

and vertex clearly visible. Armadillidium lanzai

21. Telson triangular with broadly rounded apex . Armadillidium torchiai

— Telson trapezoidal. 22

22. Male pereopod 7 basis with a distinct conical process in the distal part Armadillidium album

— Male pereopod 7 basis without any process. Armadillidium littorale

23. Antennal flagellum consisting of indistinct articles, so that it appears as a single conicle

article. 24

— Antennal flagellum consisting of distinct articles. 38

24. Pleonite 3 with 1 or 2 tubercles. 25

— Pleonite 3 without tubercles. 26

25. Pleonite 3 with a distinct median tubercle . Cyrnoniscus remyi

— Pleonite 3 with 2 median tubercles fused at the base. Carloniscus dollfusi

26. Pereonites with 6+6 longitudinal ridges. Haplophthalmus danicus

— Pereonites smooth, granulated or tuberculated, but with no longitudinal ridges. 27

27. Male pereopod 7 merus with a distinct hook-like process. 28

— Male pereopod 7 merus without any process . 29

28. Male pleopod 1 exopod with a protruding lobe on outer margin . Nesiotoniscus racovitzai

— Male pleopod 1 exopod without any lobe on outer margin. Nesiotoniscus corsicus

29. Male pleopod 1 exopod with a long posterior spine. 30

— Male pleopod 1 exopod without a posterior spine. 32

30. Dorsum smooth. 31

— Dorsum granulated. Oritoniscus punctatus

31. Eye absent; male pleopod 1 endopod longer than exopod. Oritoniscus paganus

— Eye consisting of a single ommatidium; male pleopod 1 endopod shorter than exopod

. Oritoniscus ocellatus

32. Pereonites with 2 paramedian tubercles. Finaloniscus briani

— Pereonites with no tubercles. 33

33. Male pleopod 2 exopod much longer than wide, as long as endopod Finaloniscus franciscoloi

— Male pleopod 2 exopod wider than long, much shorter than endopod. 34

34. Eye consisting of 3 ommatidia . 35

— Eye reduced, with 1 or 2 ommatidia . Trichoniscus halophilus

35. Male pleopod 1 endopod apically swollen . Trichoniscus pedronensis

— Male pleopod 1 endopod apically pointed. 36

36. Antennal flagellum of 3 articles; body with few traces of pigment. 37

Source : MNHN, Paris

— 539 —

Antennal flagellum of 4-5 articles; body well pigmented Trichoniscus pusillus provisorius

Male pleopod 1 exopod with an incision and some setae on outer margin .

. Trichoniscus pygmaeus

Male pleopod 1 exopod without incision and setae on outer margin . Trichoniscus fragilis

Antennal flagellum of more than 10 articles. Ligia italica

Antennal flagellum of 2-4 articles. 39

Antennal flagellum of 4 articles. 40

Antennal flagellum of 2 or 3 articles . 41

Uropodal endopod reaching half of exopod. Armadilloniscus candidus

Uropodal endopod protruding backwards compared with exopod Armadilloniscus ellipticus

1. Antennal flagellum of 3 articles .

- Antennal flagellum of 2 articles.

’. Male genital papilla distally bilobed ...

- Male genital papilla entire.

1. Dorsum granulated .

- Dorsum smooth.

1. Male pleopod 1 endopod apically with a

conspicuous spine directed outwards .

. Halophiloscia tyrrhena

— Male pleopod 1 endopod apically with a conspicuous transverse lobe and a triangular process

. Stenophiloscia zosterae

45. Male pleopod 1 endopod apically with a conspicuous spine . 46

— Male pleopod 1 endopod apically without any spine. Halophiloscia ischiana

46. Male pleopod 1 endopod apically with a spine on outer margin .... Halophiloscia couchii

— Male pleopod 1 endopod apically with a spine on inner margin .... Halophiloscia hirsuta

47. Cephalon with lateral lobes absent or weakly developed; pleon distinctly narrower than

pereon. 48

— Cephalon with prominent lateral lobes; body outline continuous ... Sardoniscus pygmaeus

48. Eye consisting of a single ommatidium . 49

— Eye consisting of several ommatidia. 50

49. Male pleopod 1 endopod with distal part bent outwards. Ctenoscia minima

— Male pleopod 1 endopod with distal part straight. Ctenoscia dorsalis

50. Frontal line absent . 51

— Frontal line present. 53

51. Telson rounded . Chaetophiloscia cellaria

— Telson triangular . 52

52. Pereonal epimera with an inner dark and an outer pale stripe ... Chaetophiloscia elongata

— Pereonal epimera with an outer dark and an inner pale stripe. Chaetophiloscia sicula

53. Male pereopod 7 merus with a hook-like process on sternal margin . Philoscia affinis

— Male pereopod 7 merus with no distinct modifications. 54

54. Cephalon with lateral lobes absent . 55

— Cephalon with small but distinct lateral lobes. Tiroloscia montana

55. Male pleopod 1 exopod with outer margin indented and equipped with spines; length 10-

13 mm . Tiroloscia Corsica Corsica

Source : MNHN, Paris

— 540 —

— Male pleopod 1 exopod with outer margin not indented and without spines; length 5-8 mm

. Tiroloscia macchiae

56. Uropods entirely covered by the telson, not visible in dorsal view. 57

— Uropods clearly visible in dorsal view. 58

57. Dorsum with very feeble tubercles, distinctly pubescent . Stenoniscus pleonalis

— Dorsum with distinct tubercles, not pubescent . Stenoniscus carinatus

58. Body colourless; eye absent. 59

— Body pigmented; eye present. 64

59. Telson triangular, much shorter than uropodal protopod. 60

— Telson with a long distal part, distinctly protruding compared to uropodal protopod . 62

60. Dorsum with longitudinal ridges. 61

— Dorsum without ridges . Platyarthrus hoffmannseggii

61. Cephalon with median lobe rounded . Platyarthrus schoeblii schoeblii

— Cephalon with median lobe apically indented. Platyarthrus aiasensis

62. Dorsum with no ridges. Platyarthrus caudatus

— Dorsum with longitudinal ridges. 63

63. Pereonite 7 with 3+3 feeble ridges. Platyarthrus costulatus

— Pereonite 7 with 2+2 prominent ridges . Platyarthrus corsicus

64. Pleopods 1-2 exopods with lungs . 65

— Pleopods 1-5 exopods with lungs . Protracheoniscus babori

65. Pereonite 1 with posterior margin straight. 66

— Pereonite 1 with posterior margin sinuous. 70

66. Body bordered with long setae . Leptotrichus panzerii

— Body not bordered with long setae. 67

67. Suprantennal line present, V-shaped. 68

— Suprantennal line absent. 69

68. Male pleopod 1 exopod wider than long with rounded distal part. .. Porcellionides pruinosus

— Male pleopod 1 exopod longer than wide with triangular distal part.

. Porcellionides sexfasciatus sexfasciatus

69. Tip of telson not surpassing uropodal protopod; male pleopod 1 exopod with acute apex

. Acaeroplastes melanurus sardous

— Tip of telson distinctly surpassing uropodal protopod; male pleopod 1 exopod with truncate

apex . Agabiformius lentus

70. Cephalic median lobe spatuliform, projecting well above vertex . Porcellio lamellatus sphinx

— Cephalic median lobe triangular or rounded. 71

71. Tip of telson rounded or truncate. 72

— Tip of telson pointed. 73

72. Tip of telson rounded . Porcellio dilatatus dilatatus

— Tip of telson truncate . Porcellio spatulatus

73. Male pleopod 1 exopod with truncate or rounded apex. 74

— Male pleopod 1 exopod with acute apex. 75

74. Male pleopod 1 exopod with truncate apex . Porcellio scaber

— Male pleopod 1 exopod with rounded apex . Porcellio orarum vizzavonensis

Source : MNHN, Paris

— 541 —

75. Telson with a triangular distal part, much narrower than basal; eye with more than 20

ommatidia; length up to 20 mm . Porcellio laevis

— Telson triangular with slightly and regularly concave sides; eye with 10-12 ommatidia; length

up to 8 mm . Lucasius pallidus

Acknowledgements

Our sincerest thanks are due to the following colleagues for the loan of materials: Dr P. Beron,

Sofia; Dr H. Schmalfuss, Stuttgart; Dr J. C. Weulersse, Paris; Dr L. Sorbini, Verona; Prof. F. Giusti,

Siena. Dr H. Dalens, Toulouse, is particularly acknowledged for the loan of part of the material studied

by Prof. A. Vandel and specimens of Protracheoniscus occidentalis. We are very grateful to the Mayor

of Moltifao (Corsica) for permission to visit the grotte de Pietralbello.

REFERENCES

ARCANGEU A., 1914. —- La collezione di Isopodi terrestri del R. Museo di Zoologia degli Invertebrati di Firenze.

Atti Soc. ital. Sci. nat. 52: 455-486.

— 1925. — Gli Isopodi terrestri della Sardegna. Boll. Musei Zool. Anal. comp. R. Univ. Torino 39: 1-75, pis

4-5.

— 1939. — Sardoniscus pusillus. Nuovo genere e nuova specie di Isopodo terrestre di Sardegna. Boll. Musei

Zool. Anal. comp. R. Univ. Torino 47: 125-129, pis 1-2.

— 1947. — La posizione sistematica del genere Helleria Ebn. (Crostacei Isopodi terrestri). Archo zool. ital.

31: 373-389.

— 1948. — Schizidiinae sottofamiglia di Armadillidiidae (Crostacei Isopodi Terrestri). Boll. 1st. Mus. Zool.

Univ. Torino 1 (1943-1948): 213-272.

— 1950. — Gli Isopodi terrestri della Sardegna. 2a Edizione. Boll. 1st. Mus. Zool. Univ. Torino 2 (1949-1950):

77-191, pis I-VII.

— 1954a. — Phalloniscus pygmaeus (B.L.) è proprio un Phalloniscus ? (Crostacei Isopodi terrestri). Boll. 1st.

Mus. Zool. Univ. Torino 4 (1953-1954): 123-126.

— 1954b. — Revisione di alcune specie italiane del genere Armadillidium Br. (Crostacei Isopodi terrestri).

Boll. 1st. Mus. Zool. Univ. Torino 4 (1953-1954): 135-165, pis I-XVI.

ARGANO R., Ferrara F., Guglielmo L., RIGGIO S. & Ruffo S., 1995. — Crustacea Malacostraca II (Tanaidacea,

Isopoda, Amphipoda, Euphausiacea). In A. MINELLI, S. RUFFO & S. La Posta (eds). Checklist delle specie

della fauna italiana, 30, Calderini, Bologna, 52 p.

ARGANO R. & Manicastri C, 1990. — II genere Spelaeonelhes Verhoeff, 1932 in Sardegna (Crustacea, Onis-

cidea, Trichoniscidae). Fragm. ent. 22: 1-10.

— 1991. — A preliminary report on Oniscidean fauna (Crustacea, Isopoda) from central Tyrrhenian islands:

3-8. In P. JUCHAULT & J. P. MOCQUARD (eds). The Biology of terrestrial Isopods. III. Proceedings of the

Third International Symposium on the Biology of Terrestrial Isopods, Poitiers (France), July 10-12, 1990:

VI + 221.

ARGANO R. & PESCE G. L., 1974. — Un Elumino mirmecofilo di Sardegna: Typhloschizidium cottarellii n.sp.

(Isopoda, Oniscoidea, Armadillidiidae). Fragm. ent. 9: 283-291.

AUDOUIN J. V., 1826. — Explication sommaire des planches de Crustacés de l'Égypte et de la Syrie, publiées

par Jules-César Savigny, Membre de l'Institut, offrant un exposé des caractères naturels des genres, avec

la distinction des espèces. In Description de l’Égypte, ou recueil des observations et des recherches qui

ont été faites en Égypte pendant l’expédition de l'Armée française, publié par les ordres de sa Majesté

l’Empereur Napoléon le Grand. Histoire Naturelle I (4 e partie) : 77-98.

Source : MNHN, Paris

— 542 —

BEAUCOURNU J. C.. 1967. — Contribution à la connaissance de la biologie à' Ixodes <Eschatocephalus) vesperti-

lionis Koch, 1844 et d’ixodes (Pomerantzevella) simplex Neumann, 1906 (Acarina, Ixodoidea), parasites

des Chiroptères. Ann. Spéléol. 22 : 543-580.

BERON P., 1972a. — Aperçu sur la faune cavernicole de la Corse. Série documents. Laboratoire souterrain du

C.NJI.S., Moulis 3 : 1-55.

— 1972b. — Aperçu sur la faune cavernicole de la Corse. Annls Spéléol. 27 : 807-810.

BIGOT L., 1977. — Contribution à la connaissance des zoocoenoses ripicoles de surface en Corse. Ecol. méditer.

3 : 5-12.

Bologna M. A. & Vigna Taglianti A., 1985. — Res Ligusticae CCXIV. Fauna cavernicola delle Alpi Liguri.

Annali Mus. civ. Stor. nat. Giacomo Doria 84-bis (1984): 1-388.

BRIAN A., 1951. — Descrizione di un nuovo genere di Trichoniscidae raccolto in una grotta ligure da Mario

Franciscolo. Boll. Soc. ent. ital. 81: 22-25.

— 1953. — Di alcuni Trichoniscidi nuovi della fauna endogea italiana (Isopodi terrestri). Boll. Soc. ent. ital.

83: 26-35.

BUDDE-LUND G., 1885. — Crustacea Isopoda terrestria per familias et genera et species descripta. Hauniae,

320 p.

Caruso D., 1976. — Isopodi terrestri dell’Isola di Pantelleria. Animalia 3: 105-124.

Caruso D., Baglieri C., di Maio M. C. & Lombardo B. M., 1987. — Isopodi terrestri di Sicilia ed isole

circumsiciliane (Crustacea, Isopoda Oniscoidea). Animalia Suppl. 14: 5-211.

CARUSO D. & LOMBARDO B. M., 1982. — Isopodi terrestri delle isole maltesi. Animalia 9: 5-52.

CASSOLA F., 1982. — Il popolamento cavemicolo della Sardegna. Lav. Soc. ital. biogeogr, (N.S.) 7 (1978):

615-755, pis I-m.

COBOLLI SBORDONI M., Taiti S., de Matthaeis E. & Ketmaier V., 1995. — Genetic relationships among

populations of the Oritoniscus paganus group (Crustacea, Isopoda, Oniscidea) from Corsica and Giannutri

(Tuscan Archipelago). Mém. Biospéol. 22: 13-16.

COLLINGE W. E., 1941. — Some remarks on the genus Helleria v. Ebner (Terrestrial Isopoda). Ann. Mag. nat.

Hist. 8 (11): 522-526.

DALENS H., 1977. — Comportement constructeur chez l'lsopode Nesiotoniscus corsicus Racovitza. Bull. Soc.

Hist. nat. Toulouse 113: 181-182.

DOLLFUS A., 1887. — Catalogue provisoire des espèces françaises d'Isopodes terrestres. Bull. Soc. étud. scient.

Paris 9:8-11.

— 1888. — Description d'une espèce nouvelle du genre Philoscia. Bull. Soc. étud. scient. Paris 11: 10-11,

1 pl.

— 1892. — Tableaux synoptiques de la faune française. Le genre Armadillidium (Crustacés Isopodes terrestres).

Feuille jeun. Nat. 22 : 15-19, 39-42, 135-141, 175-179.

— 1896. — Sur la distribution géographique des Armadilliens en Europe. In P. P. C. HOEK (ed.). Compte-rendu

des Séances du troisième Congrès international de Zoologie, Leyde, E.J. Brill : 356-358.

— 1897. — Tableau iconographique des Philoscia d’Europe. Crustacés Isopodes terrestres. Feuille jeun. Nat.

27 : 70-74, 91-95.

— 1899. — Catalogue des Crustacés Isopodes terrestres (Cloportides) de France. Feuille jeun. Nat. 29 : 186-190,

207-208.

EBNER V. VON, 1868. — Helleria, eine neue Isopoden-Gattung aus der Familie der Oniscoiden. Verh. zool.-bot.

Ges. Wien 18: 95-114, pl. I.

FERRARA F. & Taiti S., 1878. — Gli Isopodi terrestri dell'Arcipelago toscano. Studio sistematico e biogeografico.

Redia 61: 1-106.

— 1983. — Troglocylisticus cyrnensis n. gen. n.sp. di isopodo terrestre della Corsica (Cylisticidae). Redia 66:

485-590.

— 1984. — Oritoniscus beroni nuova specie di isopodo terrestre della Sardegna (Trichoniscidae). Redia 67:

417-420.

— 1985. — Su una nuova specie del genere Cylisticus Schnitzler, 1853, della Liguria orientale (Crustacea,

Oniscidea). Fragm. ent. 18: 33-37.

Source : MNHN, Paris

— 543 —

FRANKENBEROER Z., 1939. — Über eine neue Art der Gattung Platyarthrus Brdt. (Isopoda-Oniscoidea). Eut.

Listy 2: 115-117.

GARTHWA1TE R. L., LAWSON R. & Taiti S., 1992. — Morphological and genetic relationships among four species

of Armadilloniscus Uljanin. 1875 (Isopoda: Oniscidea: Scyphacidae). J. nat. Hist. 26: 327-338.

GARTHWAITE R. L. & TAITI S., 1989. — Platyarthrus aiasensis Legrand (Isopoda: Oniscidea: Platyarthridae) in

the Americas. Bull. Sth. Calif. Acad. Sci. 88: 42-43.

GlORDANl SOIKA A., 1954. — Studi di ecologia e biogeografia - XII. Ecologia, sistematica, biogeografia ed

evoluzione del Tylos Latreillei Auct. (Isop. Tylidae). Boll. Mus. civ. Stor. nat. Venezia 7: 63-83, pis I-XI.

— 1972. — Nuove ricerche sul genere Tylos (Crust. Isop.). Boll. Mus. civ. Stor. nat. Venezia 22-23: 193-210,

pis I-VI.

Jeannel R. & Racovitza E. G., 1908. — Biospéologica VI. Enumération des grottes visitées, 1906-1907

(2 e Série). Archs Zool. exp. gén. 8 (4) : 327-414.

KUSSAKIN O. G., 1982. — Marine and salt water isopod crustaceans (Isopoda) of cold and temperate waters of

the Northern Hemisphere. Opred. Faune SSSR , 131: 1-456 [in Russian].

LANZA B., 1979. — Universo in miniatura attomo alia Corsica. Geos. Milano 1 (2): 34-51.

Lanza B. & Poggesi M., 1986. — Storia naturale delle isole satelliti della Corsica. L’Universo. Firenze 66

(1): 1-198.

MANICASTRI C. & TAITI S., 1994. — Gli Isopodi terrestri dell’Appennino umbro-marchigiano (Crustacea, Onis¬

cidea). Biogeographia 17 (1993): 125-150.

PAOLETTI M. G. & STINNER B. R., 1988. — Two new terrestrial Isopoda (Oniscidea) from coralline cays of

Venezuela’s Caribbean coast. Proc. ent. Soc. Wash. 91 (1989): 71-80.

Racovitza E. G„ 1908. — Biospéologica IX. Isopodes terrestres (Seconde Série). Archs Zool. exp. gén. 9 (4) :

239-415, pis IV-XXIII.

Rem Y R, 1950. — Grottes de Corse. Ann/s Spéléol. 5 : 3-47.

RUFFO S., 1955. — Le attuali conoscenze sulla fauna cavernicola della regione pugliese. Memorie Biogeogr.

adriat. 3: 1-143.

SCHARFF R. F., 1894. — Einige Bemerkungen iiber eine Reise in Corsica. Ber. senckenh. naturf. Ges.: 153-167.

SCHMALFUSS H., 1981. — Die Isopoden der Nôrdlichen Sporaden (Àgais). Stuttg. Beitr. Naturk., A 343: 1-24.

— 1983. — Die Landisopoden (Oniscoidea) Griechenlands 4. Beitrag: Gattung Protracheoniscus , Teil I

(Trachelipidae). Stuttg. Beitr. Naturk., A 367: 1-9.

— 1989. — Die Land-Isopoden (Oniscidea) Griechenlands. 10. Beitrag: Gattung Trichodillidium n.g. (Armadil-

lidiidae). Sher. ost. Akad. Wiss. (Matliem.-naturw. Kl., Abt. I ) 197: 207-214.

— 1990. — Die Landisopoden (Oniscidea) Griechenlands. 11. Beitrag: Gattung Chaetophiloscia (Philosciidae).

Revue suisse Zool. 97: 169-193.

— 1991. — The terrestrial isopod genus Chaetophiloscia in western Asia (Oniscidea: Philosciidae). Stuttg.

Beitr. Naturk., A 463: 1-9.

— 1992. — The terrestrial isopod genus Porcellio in western Asia (Oniscidea: Porcellionidae). Stuttg. Beitr.

Naturk., A 475: 1-45.

SCHMOLZER K., 1971. — Die Landisopoden der Iberischen Halbinsel. Monogr. Cien. moder. 80: I-XI, 1-161,

10 maps.

SCHULTZ G. A., 1972. — Ecology and systematics of terrestrial isopod crustaceans from Bermuda (Oniscoidea).

Crustaceana Suppl. 3: 79-99.

— 1995. — Terrestrial isopod crustaceans (Oniscidea) from Paraguay with definition of a new family. Revue

suisse Zool. 102: 387-424.

SFENTHOURAKIS S., 1992. — New species of terrestrial isopods (Isopoda, Oniscidea) from Greece. Crustaceana

63: 199-209.

— 1995. — New species of terrestrial isopods (Oniscidea) from central Aegean islands. Stuttg. Beitr. Naturk.,

A 519: 1-21.

Tabacaru I., 1993. — Sur la classification des Trichoniscidae et la position systématique de Thaumatoniscellus

orghidani Tabacaru, 1973 (Crustacea, Isopoda, Oniscidea). Trav. Inst. Spéol. Emile Racovitza 32 : 43-85.

Source : MNHN, Paris

— 544 —

Taiti S. & FERRARA F., 1980. — Nuovi studi sugli Isopodi terrestri dell'Arcipelago toscano. Redia 63: 249-300.

—- 1989a. — Biogeography and ecology of terrestrial isopods from Tuscany. Moniiore zoo!, iial. (N.S.) Monogr.

4: 75-101.

— 1989b. — New species and records of Armadilloniscus Uljanin. 1875 (Crustacea Isopoda Oniscidea) from

the coasts of the Indian and Pacific oceans. Trop. Zool. 2: 59-88.

— 1991. — New species and records of terrestrial isopods (Crustacea) from the Arabian Peninsula. Fauna

Saudi Arabia 12: 209-224.

— 1995a. — Su una nuova specie di Nesioioniscus (Crustacea. Isopoda, Oniscidea) dell'Isola di Capraia

(Arcipelago Toscano) con considerazioni sul genere. Boll. Mus. re g. Sci. nai. Torino 13: 313-321.

— 1995b. — Isopodi terrestri (Crustacea. Oniscidea) delle grotte della Toscana (Italia centrale). Mém. Biospéol.

22: 169-196.

Taiti S. & Manicastri C., 1980. — Cylisticus apruiianus, nuova specie di Isopode terrestre dell’Abruzzo

(Crustacea, Oniscoidea). Fragm. eni. 15: 253-257.

Vandel A., 1944a. — Isopodes terrestres récoltés par M. Remy, au cours de son voyage en Corse (juillet-

septembre 1942). I. - Description de Paraschizidium remyi, n.sp., suivie de considérations sur l'histoire et

la répartition de la famille des Armadillidiidae. Archs Zool. exp. gén. (Notes et Revue) 83 : 83-98.

— 1944b. — Isopodes terrestres récoltés par M. Remy au cours de son voyage en Corse (juillet-septembre

1942). II. - La famille des Stenoniscidae B.-L. Archs Zool. exp. gén. (Notes et Revue) 84 : 23-47.

— 1945. — Isopodes terrestres récoltés par M. Remy, au cours de son voyage en Corse (juillet-septembre

1942). III. - La famille des Buddelundiellidae. Archs Zool. exp. gén. (Notes et Revue) 84: 100-113.

— 1947. — Espèces françaises, nouvelles ou peu connues, de Trichoniscidae (Isopodes terrestres) (Troisième

Partie). Bull. Soc. zool. Fr. 72: 35-53.

— 1951. — Le genre “Porcellio" (Crustacés; Isopodes: Oniscoidea). Évolution et systématique. Mém. Mus.

nail Hist, nat ., N. S., A 3 : 1-192.

— 1953a. — Isopodes terrestres récoltés dans les grottes de la Corse, par le Professeur P.-A. Remy, en 1942

et 1948. Notes biospéol. 8 : 153-166.

— 1953b. — Remarques systématiques, morphologiques et biogéographiques sur un groupe de Trichoniscidae

Nord-Atlantiques. (Crustacés; Isopodes terrestres). Bull. Mus. natl Hist, nat., Paris 25 : 368-375.

— 1954a. — Le peuplement isopodique de la Corse ; étude biogéographique. Revue fr. Ent. 21 : 72-84.

— 1954b. — Les Trichoniscides endogés et cavernicoles appartenant au genre Nesiotoniscus Racovitza

(Crustacés; Isopodes terrestres). Notes biospél. 9: 149-156.

— 1954c. — Sur une espèce mal connue d'Isopode terrestre, Phalloniscus pygmaeus (B.-L.). (Philoscia pygmaea

B.-L.). (Crustacé). Bull. Mus. natl Hist, nat., Paris, 26 : 226-230.

— 1954d. — Note sur deux Armadillidiidae endogés et cavernicoles de la région de Grasse (Crustacés; Isopodes

terrestres). Notes biospél. 9 : 49-60.

— 1955. — La faune isopodique cavernicole de l’Afrique du Nord (Berbérie). Notes biospél. 10 : 63-80.

— 1958. — Les Isopodes terrestres du Rif. Tamuda 6 : 127-134.

— 1959. — Les Styloniscidae et les Trichoniscidae de l'Afrique du Nord (Crustacés; Isopodes terrestres).

Bull. Mus. natl Hist, nat., Paris 31 : 159-167.

— 1960. — Isopodes terrestres (Première Partie). Faune Fr. 64 : 1-416.

— 1962. — Isopodes terrestres (Deuxième Partie). Faune Fr. 66 : 417-931.

— 1968a. — Les Isopodes terrestres et cavernicoles de la Corse. Annls Spéléol. 23 : 355-362.

— 1968b. — /. Isopodes terrestres. In Mission Zoologique Belge aux îles Galapagos et en Ecuador (N. et J.

Leleup. 1964-1965). Résultats Scientifiques. Première Partie. Vol. 1, Bruxelles, L’Imprimerie des Sciences :

37-168.

— 1977. — Les Oniscoïdes (Isopodes terrestres) de la Nouvelle-Zélande et de l’Archipel Kermadec. Mém.

Mus. natl Hist, nat., Paris, N.S., A 102 : 1-56.

VERHOEFF K. W„ 1910. — Ueber Isopoden, 16. Aufsatz, Armadillidium und Porcellio an der Riviera. Jh. Ver.

vaterl. Naturk. Württ.: 115-143.

— 1919. — Ueber augenlose Armadillidien und kritische Prüfung der Familie Armadillidiidae. 25. Isopoden-

Aufsatz. Arch. Naturgesch., A 83 (1917): 160-170.

Source : MNHN, Paris

— 545 —

— 1926. — 7. Isopoda. In Beitràge zur Kenntnis der Land- und Siisswasserfauna Korsikas. I. Ergebnisse der

Dr. Paul Schottlander-Lehrexpedition des Jahres 1914. 1m Auftrag des Kuraloriums der Dr. P. Schottlander-

Stiftung herausgegeben von W. Amdl, Berlin. Mill. zool. Mus. Berl. 12 (2): 263.

— 1928. — Über alpenlandische und italienische Isopoden. 37. Isopoden-Aufsatz. Zool. Jb. ( Sysl .) 56: 93-172.

— 1931. — Über Isopoda terrestria aus Italien. 45. Isopoden-Aufsatz. Zool. Jb. ( Sysl .) 60: 489-572, pis 6-8.

— 1933a. — Arthropoden aus südostalpinen Hôhlen gesammelt von Karl Strasser, Triest. 7. Aufsatz. Mill.

Hôhl.- u. Karstforsch .: 1-21.

— 1933b. — Zur Systematic Géographie und Ôkologie der Isopoda ten-estria Italiens und über einige

Balkan-Isopoden. 49. Isopoden-Aufsatz. Zool. Jb. (Sysl.) 65: 1-64, pis 1-2.

— 1943. — Cavernicole Diplopoden, Chilopoden und Land-Isopoden der Insel Capri, gesammelt von Dr. G.

Buchner, Ischia. Z. Karst- u. Hôhlenk.: 153-165.

VIGLIANISI R, LOMBARDO B. M. & CARUSO D., 1992. — Differenziamento genetico e morfologico in alcune

specie siciliane di Isopodi terrestri del genere Porcellio e descrizione di tre nuove specie (Crustacea, Isopoda,

Oniscoidea). Animalia 19: 235-273.

WOLF B., 1934. — Animalium Cavernarum Catalogus. Pars 2. 3: 65-144.

Source : MNHN, Paris

Bulletin du Muséum national d’Histoire naturelle, Paris. 4' série. 18, 1996

Section A. n os 3-4: 547-555

Spermatophore morphology and spermatozoal ultrastructure of

the recently described hermit crab, Strigopagurus boreonotus

Forest, 1995 (Decapoda, Anomura, Diogenidae)

by Christopher C. TUDGE

Abstract. — The spermatophore morphology and spermatozoal ultrastructure of the diogenid hermit crab,

Strigopagurus boreonotus, is described and compared with that of previously investigated diogenid genera. The

spermatophores show similarities with those described for the genera Calcinus and Dardanus. The spermatozoa

have an overall morphology which is reminiscent of representatives in the genus Clibanarius, above all, the

genus Calcinus, while still retaining a particular suite of spermatozoal characters so far unique to Strigopagurus.

Key-words. — Spermatozoa, spermatophores, ultrastructure, Strigopagurus, Diogenidae.

Morphologie du spermatophore et ultrastructure du spermatozoïde du bernard-l'hermite

récemment décrit, Strigopagurus boreonotus Forest, 1995 (Decapoda, Anomura, Diogenidae)

Résumé. — La morphologie du spermatophore et Tultrastructure du spermatozoïde du bernard-l'hermite

Diogenidae, Strigopagurus boreonotus, sont décrites et comparées avec celles des autres genres de Diogenidae

précédemment étudiés. Le spermatophore montre des similarités avec ceux décrits dans les genres Calcinus et

Dardanus. Le spermatozoïde a une morphologie générale qui rappelle les espèces du genre Clibanarius et surtout

du genre Calcinus, tout en montrant un ensemble original de caractères spermatologiques trouvés jusqu'ici seu¬

lement chez Strigopagurus.

Mots-clés. — Spermatozoïde, spermatophores, ultrastructure, Strigopagurus, Diogenidae.

C. C. Tudge, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington D.C.

20560, USA ; current address: Crustacean Laboratory, Museum of Victoria, 71 Victoria Crescent. Abbotsford, Vic. 5067,

Australia.

INTRODUCTION

The anomuran family Diogenidae is a morphologically diverse taxon currently encompassing

eighteen genera. Although comprised of a heterogeneous assemblage of genera the family is

considered to be an ancient monophyletic group (FOREST 1995). Genera are often characterised

by a combination of morphological characters but each character can vary among the other mem¬

bers of the family. Single generic apomorphies are rare. This lack of good generic apomorphies

has made elucidating the phylogenetic relationships between the different genera in the Dioge¬

nidae very difficult (Forest 1984, 1995) and has led Forest to state “...qu’il est difficile

d'éclaircir la phylogénie des Diogenidae et de préciser les liens de parenté entre des genres si

disparates.” (FOREST 1995: 28).

Source : MNHN, Paris

— 548 —

This diversity of adult morphological form in the Diogenidae is reflected in the equally

diverse morphology of reproductive components such as spermatophores (Mouchet 1930, 1931;

Hamon 1939; Matthews 1953, 1956, 1957; Tuzet & Manier 1961; Uma & Subramoniam

1984; TUDGE 1991, 1995a) and spermatozoa (KOLTZOFF 1906; Nath 1942; Dhillon 1964, 1968;

Jamieson 1991; Tudge 1992; Tudge & Justine 1994; Tudge 1995a, b). With the increasing

use of transmission electron microscopy to study the ultrastructural characters of spermatophores

and spermatozoa within the Diogenidae (and the Anomura in general) some doubt has been cast

on the monophyly of the family (TUDGE 1991, 1992, 1995a, b).

The present study describes and illustrates the spermatophore and spermatozoal ultrastructure

from the holotype specimen of the diogenid Strigopagurus boreonotus and compares it with the

ultrastructure of previously studied species in other diogenid genera.

MATERIAL AND METHODS

The specimen of Strigopagurus boreonotus Forest, 1995 was collected by Dr B. Richer de

Forges during the Bathus 2 cruise off the west coast of New Caledonia (22°46’S - 167°14’E),

south-west Pacific in May 1993. It was collected at station number 718 at a depth of 430-436 m.

This specimen is now the designated holotype (MNHN-Pg 5181).

The male reproductive material (testes and the ducts of the vasa deferentia) was removed

from the single male specimen and fixed in cold glutaraldehyde for a minimum of two hours

at 4°C then posted to Brisbane (Queensland, Australia) at ambient temperature where the

remainder of the fixation and embedding process was carried out.

For light microscopy, glutaraldehyde-fixed spermatophores were viewed under an Olympus

BH2 Nomarski interference contrast microscope. Micrographs were taken on an attached Olympus

OM-2 camera.

Transmission electron microscopy

The gonad tissue of Strigopagurus boreonotus was processed in the Zoology Department,

The University of Queensland, by the standard fixation procedure (outlined below) for transmis¬

sion electron microscopy. This was carried out in a Lynx-el. Microscopy Tissue Processor, after

the initial glutaraldehyde fixation and first phosphate buffer wash.

Portions of the testis (approximately 1 mm 3 ) were fixed in 3% glutaraldehyde in 0.2 M

phosphate buffer (pH 7.2), with 1-3% sucrose added, for a minimum of one hour at 4°C. They

were washed in phosphate buffer (3 washes in 15 min), postfixed in phosphate buffered 1%

osmium tetroxide for 80 min; similarly washed in buffer and dehydrated through ascending con¬

centrations of ethanol (40-100%). After being infiltrated and embedded in Spurr’s epoxy resin,

thin sections (50-80 nm thick) were cut on a LKB 2128 UM IV microtome with a diamond

knife. Sections were placed on carbon-stabilized colloidin-coated 200 p.m mesh copper grids and

stained (according to Daddow 1986) in Reynold’s lead citrate for 30 s, rinsed in distilled water,

then 6% aqueous uranyl acetate for 1 min, Reynold’s lead citrate again for 30 s and a final rinse

in distilled water. Micrographs were taken on an Hitachi H-300 transmission electron microscope

at 80 kV.

Source : MNHN, Paris

— 549 —

RESULTS

Spermatophore morphology

The pedunculate spermatophores of Strigopagurus boreonotus are composed of an ovoid to

spherical, sperm-filled ampulla (220 urn long x 170 pm wide) attached to a long, relatively thick

stalk (Fig. 1). The spermatophores are large, being approximately 850 pm in length.

Spermatozoal morphology

The spermatozoa are composed of a spherical acrosome vesicle capped by a convexly domed

operculum and penetrated posteriorly by an extensive perforatorial chamber. The acrosome vesicle

has a length of 5.3 pm and is 4.5 pm wide. Posterior to the acrosome vesicle is the cytoplasmic

region with three microtubular arms and, more posteriad, the nucleus. The entire sperm cell is

approximately 10 pm in length (refer to Figs 2, 3 throughout).

Acrosome

The apical or anterior pole of the acrosome vesicle of Strigopagurus boreonotus is covered

by a high domed, electron-dense operculum (Fig. 3A, D). Subjacent to, and filling the inside

of, the domed operculum is a coarsely granular, homogeneous subopercular zone. The subopercu-

lar zone extends posteriorly to meet the perforatorial chamber and laterally to abut on the inner

and outer acrosome zones (Fig. 3A). Along this boundary a thin, electron-lucent area containing

small electron-dense granules occurs. Thinly enveloping the anterior end of the perforatorial

chamber is the inner acrosome zone. This homogeneous, finely granular zone forms an

Source : MNHN, Paris

— 550 —

subopercular zone

foratorial chamber

peripheral acrosome zone

opercul

microvil

projections

FIG. 2. — Strigopagurus boreonolus (Diogenidae). Semidiagrammatic longitudinal section of a spermatozoon, based on a tracing

of a micrograph. Scale bar = 1 pm.

electron-dense ring which extends no further posteriad than the widest point of the perforatorial

chamber (Fig. 3A, E). External to the inner acrosome zone, and comprising the bulk of the acro¬

some vesicle, is the outer acrosome zone. This zone is similar in form to the inner acrosome

zone but less electron-dense. The outer acrosome zone is prevented from reaching the exterior

acrosome membrane by the intervention of a thin, moderately electron-dense zone which lies

beneath the acrosome membrane, the peripheral acrosome zone (Fig. 3A, E, F). This peripheral

zone has a slightly irregular appearance (pale lacunae in a darker matrix) and extends from the

operculum to the basal opening of the perforatorial chamber.

The invaginated perforatorial chamber is spherical, about 3 |xm at its widest point and

occupies a large portion of the centre of the sperm cell (Fig. 3A, E, F). It has a constricted

basal opening and the anteriormost (apical) region can vary in form, giving the appearance of

Source : MNHN, Paris

— 551 —

some asymmetry. Posteriorly, the walls of the perforatorial chamber produce short, microvillar

projections which extend laterally into the chamber (Fig. 3A, E). The contents of the perforatorial

chamber is divisible into two areas of differing form and the more posterior region appears

continuous with the cytoplasm in the region of the constricted opening. The posteriormost portion

is coarsely granular, heterogeneous and can appear almost reticulate. This region changes ante¬

riorly to give a more homogeneous, finely granular zone. The boundary between the two areas

is approximately at the midpoint of the perforatorial chamber (Fig. 3A).

Cytoplasmic region

The cytoplasm forms a thick collar around the posterior part of the acrosome vesicle, al¬

though a thin layer also occurs beneath the vesicle. Abundant membranes and membrane systems

associated with numerous cristate mitochondria are a conspicuous part of the cytoplasm (Fig. 3A,

C, F). A single, concentrically arranged, membraneous whorl or lamellar scroll is apparent in

many of the spermatozoa (Fig. 3A, C). Bundles of microtubules representing the bases of the

three microtubular arms pass through the cytoplasm and a pair of centrioles is seen in the cy¬

toplasm directly below the constricted opening of the perforatorial chamber (Fig. 3B). A disrupted

nuclear membrane forms a discontinuous partition between the two regions.

Nuclear material

The nucleus is amorphous, but maintains an approximately globular shape with a crenulated

external surface, and is surrounded by a thickened nucleo-plasma membrane. The contents of

the nucleus are finely granular, relatively homogeneous and electron-pale (Fig. 3A).

DISCUSSION

The spermatophore morphology described for Strigopagurus boreonotus is consistent with

the pedunculate spermatophore type common in, and perhaps diagnostic of, the Anomura (Tudge

1995a). This pedunculate spermatophore morphology is a tripartite arrangement with a sperm-

filled ampulla connected to a pedestal or basal plate via a stalk.

Within the Diogenidae the spermatophore morphology has been recorded from twenty-two

species in eight genera (see list of references in the Introduction) and from these representatives

the morphology of the spermatophore of Strigopagurus boreonotus (Fig. 1) appears most similar

to the genera Calcinus and Dardanus , and especially that described for Dardanus megistos

(TUDGE 1991, 1995a); except that the spermatophore of S. boreonotus is nearly six times larger.

The importance of size of spermatophores as a reliable distinguishing character in the Anomura

is undermined by the fact that spermatophore size is proportional to the diameter of the vas

deferens of the reproductive tract which is in turn proportional to the size of the individual

specimen. Despite this fact, the size of the spermatophore in S. boreonotus (850 (im) more closely

approximates the range of sizes recorded for representatives of the genus Dardanus (160-

3500 (im) than those recorded for the genus Calcinus (150-180 pm) (TUDGE 1991, 1995a). The

Source : MNHN, Paris

— 553 —

spermatophore morphology of S. boreonotus is unlike any morphology recorded for other

paguroids (outside the Diogenidae) or even other anomurans (TUDGE 1991, 1995a).

The spermatozoa of Strigopagurus boreonotus conform to a general paguroid sperm type

characterised by a concentrically zoned acrosome vesicle, apically capped by an electron-dense

operculum; the acrosome vesicle shape may vary from spherical, through ovoid, to more elongate

and cylindrical; the acrosome vesicle is penetrated from its posterior end by a perforatorial cham¬

ber, which may terminate pre-equatorially or extend to a subterminal position immediately

beneath the operculum; the acrosome vesicle is embedded in the cytoplasm and/or nucleus and,

most importantly, there are three microtubular arms (of cytoplasmic origin) which emerge from

the cytoplasm below the acrosome vesicle (Pochon-Masson 1968a, b; Chevaillier 1970;

Hinsch 1980; Jamieson 1991; Tudge & Jamieson, 1991; Tudge 1992, 1995a, b).

In size and shape the spermatozoa of Strigopagurus boreonotus are similar to investigated

representatives in the genera Calcinus and Clibanarius; although many ultrastructural differences

are apparent (Jamieson 1991; Tudge 1992, 1995a, b). The position of the thin inner acrosome

zone on the anterior region of the perforatorial chamber in S. boreonotus (Figs 2, 3A, E) is similar

in Calcinus minutus and the bulbous, almost spherical perforatorial chamber with small micro-

villar projections (Figs 2, 3A, E, F) is approximated by all three investigated species in the

genus Calcinus (TUDGE 1995a, b). The tendency for the anterior wall of the perforatorial chamber

in S. boreonotus to be irregular (Figs 2, 3A) may indicate links to the investigated Calcinus

species in which this region of the perforatorial chamber divaricates to form two or more distinct

fingers or lobes (TUDGE 1995a, b). This latter character’s presence is an autapomorphy for the

genus Calcinus. There is no dense perforatorial ring, an autapomorphy of the investigated

Clibanarius species, seen in the spermatozoa of S. boreonotus. Some (apomorphic?) structures

of the acrosome vesicle present in the sperm of S. boreonotus which have not been seen in other

investigated diogenids are the extreme width and bulbous shape of the perforatorial chamber,

the electron-lucent, granular region forming the boundary between the perforatorial chamber and

subopercular zone and the loculated appearance of the peripheral acrosome zone (Figs 2, 3A,

E, F).

The spermatozoa of Strigopagurus boreonotus have a distinct ultrastructural morphology

which distinguishes this species from the other investigated genera in the Diogenidae. Some

overall similarities to investigated members in the genus Clibanarius and also representatives

in the genus Calcinus are apparent. A preliminary phylogenetic analysis based on spermatozoal

and spermatophore characters (analysis in progress, results available upon request) consistently

placed Strigopagurus boreonotus in the same clade with, and between, the genera Calcinus and

Clibanarius (TUDGE 1995a). At present, only six of the eighteen genera in the Diogenidae have

been investigated for spermatophore and spermatozoal morphology. With future research into

the remaining genera, spermatophore and spermatozoal ultrastructural morphology may further

assist in the elucidation of inter-generic relationships within the family Diogenidae.

Acknowledgements

I wish to thank Dr Bertrand Richer de Forges (ORSTOM, Nouméa) for the collection and initial

fixation of the specimen of Strigopagurus boreonotus. Mrs Lina Daddow (Zoology Department, The Uni¬

versity of Queensland) is heartily thanked for expert assistance with all aspects of electron microscopy.

Source : MNHN, Paris

— 554 —

I am also grateful to Prof. Jacques Forest (MNHN, Paris) for identification of the specimen. The use of

facilities at the Natural History Museum, Washington D.C., is gratefully acknowledged.

REFERENCES

CHEVAILL1ER P., 1970. — Recherche sur la structure et les constituants chimiques des cellules germinales mâles

des Crustacés Décapodes. Doctor of Science Thesis, University of Rennes, Rennes, France : 1-323.

Daddow L. Y. M., 1986. — An abbreviated method of the double lead stain technique. J. Submicrosc. Cytol.

18: 221-224.

DHILLON B., 1964. — Sperm nucleus of Clibanarius longitarsus. Experientia 20: 505-506.

— 1968. — Radial processes of decapod sperm. Microscope 76: 365-368.

FOREST j., 1984. — Révision du genre Aniculus (Decapoda, Diogenidae). Crustaceana (Suppl. 8) : 1-91.

— 1995. — Crustacea Decapoda Anomura: révision du genre Trizopagurus Forest. 1952 (Diogenidae), avec

l’établissement de deux genres nouveaux. In A. CROSNIER (ed.). Résultats des Campagnes MUSORSTOM,

Vol. 13. Mém. Mus. natl Hist. nat. 163 : 9-149.

Hamon M., 1939. — La constitution chimique des spermatophores de Crustacés supérieurs du groupe des

Pagurides. C. R. Soc. Biol. 130: 1312-1314.

HlNSCH G. W„ 1980. — Spermiogenesis in Coenobita clypeatus, 1. Sperm structure. Int. J. Invert. Reprod. 2:

189-198.

JAMIESON B. G. M., 1991. — Ultrastructure and phylogeny of crustacean spermatozoa. Mem. Qld. Mus. 31: 109-

142.

KOLTZOFF N. K., 1906. — Studien iiber die gestalt der zelle. I. Untersuchungen iiber die spermien der decapoden,

als einleitung in das problem der zellengestalt. Arch. Mikroskop. Anat. 67: 364-572.

MATTHEWS D. C., 1953. — The development of the pedunculate spermatophore of a hermit crab, Dardanus asper

(De Haan). Pac. Sci. 7: 255-266.

— 1956. — The probable method of fertilization in terrestrial hermit crabs based on a comparative study of

spermatophores. Pac. Sci. 10: 303-309.

— 1957. — Further evidences of anomuran non-pedunculate spermatophores. Pac. Sci. 11: 380-385.

MOUCHET S., 1930. — Mode de formation des spermatophores chez quelques Pagures. C. R. Acad. Sci. Paris

190:691-692.

— 1931. — Spermatophores des Crustacés Décapodes Anomoures et Brachyoures et castration parasitaire chez

quelques Pagures. Stat. Océanogr. Salammbô. Ann. 6 : 1-203.

NATH V., 1942. — The decapod sperm. Trans. Nat. Inst. Sci. India 2: 87-119.

POCHON-MASSON J., 1968a. — L’ultrastructure des spermatozoïdes vésiculaires chez les Crustacés Décapodes

avant et au cours de leur dévagination expérimentale. I. Brachyoures et Anomoures. Ann. Sci. Nat., Zool.

Biol. 10: 1-100.

— 1968b. — L'ultrastructure des spermatozoïdes vésiculaires chez les Crustacés Décapodes avant et au cours

de leur dévagination expérimentale. II. Macroures. Discussion et conclusions. Ann. Sci. Nat., Zool. Biol.

10 : 367-454.

TUDGE C. C., 1991. — Spermatophore diversity within and among the hermit crab families, Coenobitidae, Dio¬

genidae and Paguridae (Paguroidea, Anomura, Decapoda). Biol. Bull. 181: 238-247.

— 1992. — Comparative ultrastructure of hermit crab spermatozoa (Decapoda: Anomura: Paguroidea). J. Crust.

Biol. 12: 397-409.

— 1995a. — The Ultrastructure and Phylogeny of Anomuran Crab Spermatozoa. PhD. Thesis. Zoology De¬

partment, The University of Queensland, Australia: 1-346.

— 1995b. — Ultrastructure and Phylogeny of the Spermatozoa of the Infraorders Thalassinidea and Anomura

(Decapoda, Crustacea). In B. G. M. JAMIESON, J. AUSIO & J.-L. JUSTINE (eds). Advances in Spermatozoal

Phylogeny and Taxonomy. Mém. Mus. natl Hist. nat. 166 : 251-263.

Source : MNHN, Paris

— 555 —

TUDGE C. C. & Jamieson b. g. M., 1991. — Ultrastructure of the mature spermatozoon of the coconut crab

Birgus latro (L.) (Coenobitidae, Paguroidea, Decapoda). Mar. Biol. 108: 395-402.

TUDGE C. C. & Justine J.-L., 1994. — The cytoskeletal proteins actin and tubulin in the spermatozoa of four

decapod crabs (Crustacea. Decapoda). Ada Zool. (Stockholm) 75: 277-285.

Tuzet O. & Manier J.-F., 1961. — Les constituants cytoplasmiques au cours de la spermatogenèse de six

pagures. Bull. Soc. zool. Fr. 86 : 305-315.

Uma K. & SUBRAMONIAM T., 1984. — A comparative study of the spermatophore in Scylla serrata (Forskal)

(Decapoda: Brachyura) and Clibanarius longitarsus (De Haan) (Decapoda: Anomura). J. Mar Biol. Assoc.

India 26: 103-108.

Source : MNHN, Paris

Seclic

/ d' Histoire naturelle, Paris, 4 e série, 18. 1996

on A, n“ 3-4 : 557-564

Hypocolpus pararugosus, espèce nouvelle de l’Indo-Ouest

Pacifique (Crustacea, Decapoda, Brachyura, Xanthidae)

par Alain CROSNIER

Résumé. — Une espèce nouvelle d 'Hypocolpus, H. pararugosus, est décrite d’après des spécimens récoltés

en Nouvelle-Calédonie et au Sri Lanka. Elle avait, jusqu’à maintenant, été confondue avec H. rugosus (Henderson,

1893). Sa comparaison avec les formes voisines amène à élever au rang d’espèce la sous-espèce stenocoelus

Guinot-Dumortier, 1960, de H. rugosus.

Mots-clés. — Crustacea, Decapoda, Brachyura, Xanthoidea, Xanthidae, eau profonde, océan Pacifique,

Nouvelle-Calédonie, espèce nouvelle.

Hypocolpus pararugosus , new species from the Indo-west Pacific

(Crustacea, Decapoda, Brachyura, Xanthidae)

Abstract. — A new species belonging to the genus Hypocolpus, H. pararugosus, is described after specimens

from New Caledonia and Sri Lanka. It had, until now, been confused with H. rugosus (Henderson, 1893). The

comparison of this species with closely related ones leads to the elevation of the subspecies rugosus stenocoelus

Guinot-Dumortier, 1960, to full species rank.

Key-words. — Crustacea, Decapoda, Brachyura, Xanthoidea, Xanthidae, deep water. Pacific Ocean,

New Caledonia, new species.

A. Cbosnif-r, Océanographe biologiste ORSTOM, Muséum national d'Histoire naturelle. Laboratoire de Biologie des Invertébrés

Marins et Malacologie. 55 rue de Buffon, F-75231 Paris cedex 05.

Lors de la campagne BATHUS 1, effectuée avec le navire de l’ORSTOM Alis au large de

la Nouvelle-Calédonie, un spécimen A'Hypocolpus a été capturé par 140-150 m de profondeur.

Il s’est révélé appartenir à une espèce nouvelle, dont un exemplaire avait déjà été capturé au

Sri Lanka au début de ce siècle, mais avait alors été identifié, à tort, par Laurie (1906), puis

par Guinot-Dumortier (1960), à Hypocolpus rugosus (Henderson, 1893).

Cette espèce est décrite ci-après et comparée aux espèces qui lui sont les plus proches.

Hypocolpus pararugosus n.sp.

(Figs 1, 2A-C, 3A-B)

Hypocolpus rugosus rugosus Guinot-Dumortier, 1960, en partie : 197 (seulement la femelle

identifiée par Laurie et mentionnée ci-après dans le matériel examiné). Non Henderson, 1893.

Matériel examiné. — Nouvelle-Calédonie, BATHUS 1 : stn DW 692, 20°35,43’S - 164°58,88’E, 140-150 m,

17.III.1993 : 1 ô 15,7 x 23,8 mm (MNHN-B 22937).

Source : MNHN, Paris

— 558 —

Sri Lanka, Golfe de Manaar, Hermann coll., Laurie det., 1906: 1 $ ovigère 12,4 x 18,0 mm (BM

1907.5.22.215).

Types. — Le mâle récolté à la station DW 692 de BATHUS 1 est l’holotype. La femelle provenant du Sri

Lanka est un paratype.

Étymologie. — L’adjonction du mot grec para, près, à rugosus rappelle combien cette espèce nouvelle est

proche de celle décrite par Henderson, en 1893, sous ce dernier nom.

Description (faite d’après l’holotype).

La carapace, pratiquement glabre, est 1,5 fois plus large que longue et présente des lobules

proéminents, ornés de granules assez gros, pas très serrés et séparés par des espaces lisses ou

garnis de minuscules ponctuations. Les sillons séparant les lobules sont lisses. La disposition

de ces lobules correspond à celle proposée par Dana en 1852 (Fig. 3A); on notera toutefois

que le lobule 2M est divisé en deux par un sillon longitudinal et que l’aire cardiaque (IP) est

divisée en trois : deux gros lobules antérieurs, disposés côte à côte et un petit lobule postérieur

impair.

Le front présente deux lobes peu proéminents, reliés aux orbites par une faible concavité.

Le bord antérolatéral de la carapace est fortement saillant, lisse à l’œil nu (avec de minus¬

cules granules à un assez fort grossissement), et légèrement convexe ; il s’étend, en arrière, jus¬

qu’au niveau de la pointe externe du lobule 3L. Le reste du bord latéral de la carapace porte

ensuite des granules dont deux sont plus gros que les autres, puis est excavé pour permettre

aux pattes ambulatoires 4 et 5 de venir s’appliquer étroitement contre la carapace lorsqu’elles

se replient. Les deux gros granules se trouvent, l’un légèrement en arrière de l’extrémité pos¬

térieure du bord antérolatéral lisse, l’autre un peu en avant de la partie excavée.

Source : MNHN, Paris

— 559 —

Le bord postérieur de la carapace, légèrement sinueux, est bordé par deux rangées de gra¬

nules; ceux de la rangée postérieure sont nettement plus petits que ceux de la rangée antérieure.

La cavité sous-hépatique, qui demeure bien visible lorsque le chélipède correspondant est

replié, est de grande taille et s’étend transversalement. Elle est quatre fois plus longue que large.

Ses bords antérieur et postérieur, à peine arqués, sont subparallèles sur presque toute leur longueur

et la cavité ne présente pas de rétrécissement marqué dans sa partie externe; son bord interne,

régulièrement arrondi, dépasse l’aplomb du bord externe de l’orbite; sa partie externe forme

une surface plate, non bordée. L’intérieur de la cavité, lisse, est plus creusée dans sa partie

postérieure que dans sa partie antérieure et présente, sur presque toute sa longueur, un très léger

renflement. Le bord postérieur de la cavité est longé extérieurement, sur ses deux tiers internes

environ, par une dépression bien marquée, bordée postérieurement par des granules.

FlG. 2. — A-C, Hypocolpus pararugosus n.sp. A, <5 holotype, 15,7 x 23,8 mm, Nouvelle-Calédonie (MNHN-B 22937) : vue vemrale

de l'animal enlier; B. carpe el propode du chélipède droit, vue externe. C, 5 ovigère paratype, 12,4 x 18,0 mm, Sri Lanka

(BM 1907.5.22.215) : vue dorsale de l'animal entier. D, Hypocolpus rugosus (Henderson. 1893). 2 lectotype, 12,6 x 18,5 mm,

Inde : carpe et propode du chélipède droit, vue externe. E, Hypocolpus stenocoelus Guinot, 1960, 6 lectotype, 15,0 x 22,9 mm,

île Maurice (BM 1889.3.27.22) : carpe et propode du chélipède droit, vue externe.

Source : MNHN, Paris

— 560 —

Les parties visibles des stemites sont creusées de dépressions peu granuleuses, séparant des

protubérances fortement granuleuses. L’ensemble présente ainsi un aspect fortement érodé.

Les troisièmes maxillipèdes sont granuleux, avec un mérus présentant deux dépressions lisses

et un ischion avec un sillon longitudinal médian également lisse.

Les parties visibles des chélipèdes et des pattes ambulatoires repliés (à l’exception des dac¬

tyles) présentent de gros nodules et des granules plus ou moins gros. On notera, en particulier,

une forte protubérance arrondie sur la partie antérieure de la face externe du carpe des chélipèdes.

L’abdomen, dont les segments 3-5 sont soudés, présente sur les segments 2-5 un fort ren¬

flement transversal, qui, à peine divisé transversalement par un sillon sur les segments 2 et 3,

présente une division bien marquée sur les segments 4 et 5. Sur le segment 6, on observe deux

renflements largement séparés.

Les pléopodes 1 et 2 présentent l’aspect habituel observé chez les espèces du genre. On

notera seulement que les grandes soies distales des pléopodes 1 sont au nombre de deux (Fig. la-b).

Le paratype (Fig. 2C) est bien conforme au type. On notera seulement que le rapport

longueur/largeur de la cavité sous-hépatique est de 3,8 et que la partie antérieure de l’aire

cardiaque (IP) n’est pas divisée en deux.

Remarques

Guinot-Dumortier (I960: 197) a bien analysé la plupart des différences séparant la

femelle du Sri Lanka, identifiée à H. rugosus par Laurie, des types d’HENDERSON, mais elle

n’a pas conclu à son appartenance à une espèce différente.

Onze espèces d 'Hypocolpus ont été décrites jusqu’à présent: H. abbotti Rathbun, 1894;

H. diverticulatus (Strahl, 1861); H. guinotae Vannini, 1982; H. haanii Rathbun, 1909 [=//. gra¬

nular us de Haan, 1837, cf. Yamaguchi & Baba 1993: 443]; H. kurodai Takeda, 1980;

H. mararae Crosnier, 1991; H. pardii Galil & Vannini, 1960; H. perfectus Guinot-Dumortier,

1960; H. punctatus (Miers, 1884); H. rugosus rugosus (Henderson, 1893); H. rugosus steno-

coelus Guinot-Dumortier, 1960.

H. pararugosus, par la forme de sa cavité sous-hépatique, est proche de H. haanii Rathbun,

1909, H. rugosus rugosus (Henderson, 1893) et H. rugosus stenocoelus Guinot-Dumortier, 1960.

Nous avons pu examiner le matériel suivant de ces dernières espèces :

H. haanii : Japon, spécimen acheté au revendeur Frank par le Muséum de Paris et provenant

de la collection faite par Von Siebold et Bürger (T. YAMAGUCHI in litt.). 1 S, 26,6 x 37,9 mm

(MNHN-B 8588).

H. rugosus : Inde (côte sud-est), golfe de Manaar (Tuticorin), Thurston coll., syntypes :

2 9 9, 9,1 x 13,3 mm et 12,6 x 18,5 mm (BM 1939.9.20.1-2). Nous sélectionnons la femelle

de 12,6 x 18,5 mm comme lectotype.

Indonésie, Iles Kai, expédition danoise de 1922, stn 61, 50m, 14.V.1922: 1 9,

8,9 x 14,1 mm (MNHN-B 22294). Ce spécimen, qui n’était pas encore identifié, est ici men¬

tionné pour la première fois. Il est malheureusement en très mauvais état.

H. rugosus stenocoelus : Ile Maurice, syntypes : 2 â â 14,1 x 21,2 et 15,0 x 22,9 mm (BM

1889.3.27.22). C’est par erreur que GUINOT-DUMORTIER (1960 : 198) mentionne, pour les syn-

Source : MNHN, Paris

— 561 —

FIG. 3. — Vue dorsale de l'animal entier et cavité sous-hépatique gauche. A-B, Hypocotpus pararugosus n.sp., Nouvelle-Calédonie :

<5 holotype. 15,7x23.8 mm (MNHN-B 22937). C-D. Hypocotpus rugosus (Henderson, 1893), Inde, golfe de Manaar : 9

lectotype, 12,6x18,5mm (BM 1939.9.20.1-2). E-F, Hypocotpus stenocoelus Guinot, I960, île Maurice: â lectotype,

15,0x22,9 mm (BM 1889.3.27.22).

Source : MNHN, Paris

— 562 —

types, une femelle et un mâle. Par ailleurs, cet auteur n’ayant pas désigné d’holotype pour cette

sous-espèce, nous sélectionnons le mâle de 15,0 x 22,9 mm comme lectotype.

H. haanii se sépare d'H. pararugosus par, entre autres :

— le bord postérieur de la cavité sous-hépatique qui est fortement convexe, de sorte que

la largeur de la cavité sous-hépatique est très variable, son maximum se situant vers le milieu

de sa longueur (c/. Guinot-Dumortier 1960, pl. Il, fig. 12). Le rapport longueur/largeur de la

cavité est voisin de 2,75. Par ailleurs, aucune dépression ne borde, extérieurement, le bord

postérieur de la cavité et aucun renflement longitudinal ne s’observe chez celle-ci;

— les granules couvrant le corps qui sont beaucoup plus nombreux (plus serrés) et nettement

mieux marqués. Cette différence est très nette, tant sur la face dorsale de la carapace que sur

la face ventrale du corps (sternites et abdomen en particulier);

— le pléopode 1 qui porte cinq longues soies distales (au lieu de deux).

H. rugosus rugosus (Fig. 3C-D) se distingue par :

— la cavité sous-hépatique proportionnellement plus courte et dont la largeur décroît

régulièrement de sa partie interne à sa partie externe (Fig. 3D et Guinot-Dumortier 1960, pl. Il,

fig. 13). Le rapport longueur/largeur de la cavité est compris entre 2,5 et 2,7. Par ailleurs la

dépression qui borde extérieurement le bord postérieur de la cavité est beaucoup moins marquée ;

— la partie saillante du bord antérolatéral de la carapace plus courte (en liaison avec le

moindre développement en longueur de la cavité sous-hépatique) et se terminant par une dent

granuleuse bien marquée (dent qui est totalement absente chez H. pararugosus ) ;

— les granules couvrant le corps beaucoup mieux marqués, le relief apparaissant ainsi

nettement plus accentué; le bord antérolatéral saillant de la carapace, notamment, est granuleux

(au lieu de paraître lisse à l’œil nu). Les dents latérales de la carapace sont nettement plus

fortes. Sur la face dorsale de la carapace, entre les amas de granules, on note des petites

dépressions nettes et profondes qui manquent chez H. pararugosus.

H. rugosus stenocoelus (Fig. 3E-F) se différencie par :

— la cavité sous-hépatique proportionnellement plus courte mais surtout beaucoup plus

étroite et dont la largeur décroît régulièrement de sa partie interne à sa partie externe (Fig. 3F

et Guinot-Dumortier 1960, pl. II, fig. 15). Le rapport longueur/largeur de la cavité est très

voisin de 5. Chez cette espèce, on observe, comme chez H. pararugosus, une dépression bordant

extérieurement le bord postérieur de la cavité, dépression encore mieux marquée que chez

H. pararugosus;

— la partie saillante du bord antérolatéral de la carapace plus courte (en liaison avec le

moindre développement en longueur de la cavité sous-hépatique);

— les granules semblables à ceux de H. rugosus rugosus; cependant, l’ensemble de la face

dorsale de la carapace semble avoir un relief un peu moins accentué, ceci semblant essentiel¬

lement dû à un meilleur développement de la pilosité de la carapace qui amortit les reliefs. On

notera également que, chez les spécimens de H. rugosus stenocoelus que nous avons examinés,

les soies sont brunes (au lieu d’être blanchâtres chez H. rugosus rugosus); de ce fait les granules,

entourés par ces soies, sont plus visibles.

Source : MNHN, Paris

— 563 —

Si les comparaisons ci-dessus montrent que H. pararugosus se distingue somme toute

facilement des espèces qui lui sont le plus proches, elles montrent également, à notre avis, qu’il

n’est pas justifié de considérer H. stenocoelus comme étant une sous-espèce de H. rugosus, la

forme particulière de la cavité sous-hépatique suffisant, à elle seule, à justifier l’élévation au

rang d’espèce de ce taxon. D’autres différences existent également entre H. rugosus et H. ste¬

nocoelus et ont bien été mises en relief par GUINOT-DUMORTIER (1960 : 201-202) que l’on aura

intérêt à relire. En particulier, comme cet auteur le fait remarquer, le bord latéral de la carapace,

en arrière de la partie saillante correspondant à la cavité sous-hépatique, «forme une crête dé¬

primée et surmontée de granules petits, assez réguliers, nombreux [chez H. stenocelus] tandis

que chez H. rugosus le bord est formé par une succession de dents granuleuses peu nombreuses

dont l’irrégularité masque la dépression marginale». Ces caractères s’observent bien sur les pho¬

tos C et E de la Fig. 3. Par ailleurs la granulation des chélipèdes est plus fine chez H. stenocoelus

que chez H. rugosus (comparer les photos D et E de la Fig. 2).

Comme nous l’avions mentionné lors de la description de H. mararae (CROSNIER 1991 :

427), seule cette espèce, parmi les Hypocolpus, avait été capturée à des profondeurs assez grandes

(130-150 m), toutes les autres profondeurs mentionnées dans la littérature pour les captures d'Hy¬

pocolpus ne dépassant pas, jusqu’alors, 30 m. La capture d’une seconde espèce vivant aux alen¬

tours de 150 m de profondeur est donc intéressante. Ceci dit, le spécimen de Laurie, d’après

les indications fournies par cet auteur, a été capturé dans un récif de corail du golfe de Manaar ;

aucune profondeur n’est indiquée, mais a priori, compte-tenu des autres récoltes de crustacés

étudiées par Laurie, elle ne devrait pas être bien grande ; ceci semblerait donc indiquer que

H. pararugosus aurait une répartition bathymétrique assez large.

Remerciements

Bertrand Richer de Forges (ORSTOM) nous a envoyé les récoltes parmi lesquelles le spécimen

étudié ici a été trouvé. Paul Clark (The Natural History Museum, Londres) nous a envoyé en prêt la

plupart des spécimens nécessaires à cette étude. Maurice Gaillard et Jacques Rebière sont les auteurs

respectivement des dessins et des photos qui illustrent cette note. Le Prof. Danièle Guinot et Peter Davie

ont bien voulu critiquer notre manuscrit. À tous nous adressons nos remerciements.

RÉFÉRENCES

CROSNIER A., 1991. — Hypocolpus mararae, espèce nouvelle de Polynésie française (Crustacea Decapoda Bra-

chyura Xanthidae). Bull. Mus. nail Hist, nat., Paris, 4 e sér., A 13 (3-4) : 423-428, fig. 1, pi. I.

Dana J. D., 1851. — On the Classification of the Cancroidea. Am. J. Sci. Arts, sér. 2 12 (34): 121-131.

Galil B. & Vannini M., 1990. — Research on the coast of Somalia. Xanthidae. Trapeziidae, Carpiliidae. Menip-

pidae (Crustacea Brachyura). Tropical Zoology 3: 21-56, figs 1-8.

GUINOT-DUMORTIER D., 1960. — Révision des genres Euxanthus Dana et Hypocolpus Rathbun (Crust. Decap.

Brach.). Remarques sur les cavités sous-hépatiques et les coaptations des Hypocolpus. Mém. Mus. nain.

Hist, nat., nlle série, A, Zool. 20 (2) : 153-218, figs 1-5, pis 1-12.

Haan W. de, 1833-1850. — Crustacea. In P. F. DE SlEBOLD, Fauna Japonica sive Descriptio animalium. quae

in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava Imperium tenent.

Source : MNHN, Paris

— 564 —

suspecio, an il is 1823-1830 collegil, notis. observationibus el adumbrationibus illusiravit. Lugduni

Batavorum fasc. 1-8: 1-XXI + VII-XVII + IX-XVI + 1-243, pi. 1-55, A-Q, circ., pi. 2.

HENDERSON J. R., 1893. — A Contribution to Indian Carcinology. Trans. Linn. Soc. Lond., 2' sér. 5: 325-458,

pis 36-40.

LAURIE R. D., 1906. — Report on the Brachyura collected by Prof. Herdman, at Ceylon, in 1902. In W. A.

HERDMAN, Report to llie Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar.

Part V. Suppl. Rep. n°40: 349-432, fig. 1-12, pis 1-2.

MlERS E. J., 1884. — Crustacea. In Report on the zoological collections made in the Indo-Pacific Ocean during

the voyage of H.M.S. Alert 1881-1882. Part I. The collections from Melanesia. Part II. The collections

from the Western Indian Ocean. London: 178-322, 513-575, pis 18-32, 46-51.

RATHBUN M. J., 1894. — Description of two new species of crabs from the western Indian Ocean, presented to

the National Museum by Dr. W. L. Abbott. Proc. U.S. natn. Mus. 17: 21-24.

— 1909. — New crabs from the Gulf of Siam. Proc. hiol. Soc. Wash. 22: 107-114.

SERÈNE R., 1984. — Crustacés décapodes brachyoures de l’océan Indien occidental et de la mer Rouge. Xan-

thoidea : Xanthidae et Trapeziidae. Avec un addendum par A. Crosnier : Carpiliidae et Menippidae. Faune

tropicale 24 : 1-400, fig. A-C + 1-243, pis I-XLVIII.

Strahl C., 1861. — Carcinologische Beitràge. A. Ueber die Dekapoden Gattung Euxanthus Dana. B. Ueber

Cancer panope Herbst. Arch. Naturgesch., sér. 27 1 : 101-106.

TAKEDA M., 1980. — A new Xanthid Crab from the Ryukyu Islands. Bull, biogeogr. Soc. Japan 35 (3): 39-44,

figs 1-6.

VANNINI M., 1982. — Notes on somalian species of the genus Hypocolpus (Decapoda, Brachyura, Xanthidae)

with the description of a new species. Crustaceana 42 (1): 101-105, figs 1-2, pi. 1.

YamaGUCHI T. & Baba K., 1993. — Crustacean Specimens Collected in Japan by Ph. F. Von Siebold and

H. Biirger and Held by the Nationaal Natuurhistorisch Museum in Leiden and other Museums. In

T. YAMAGUCHI (ed.). Ph. von Siebold and Natural History of Japan. Crustacea. Carcinological Society of

Japan: 145-570, figs 1-200 D + II A-F + 3 fig. n.n. + III A-D.

Source : MNHN, Paris

Bulletin

l'Histoire

naturelle, Paris, 4' série, 18, 1996

Section A, n“ 3-4 : 565-570

Redescription des types de Naushonia perrieri (Nobili, 1904)

(Crustacea, Decapoda, Laomediidae)

par Nguyen NGOC-HO

Résumé. — Les types de Naushonia perrieri (Nobili, 1904), récoltés à Djibouti, en mer Rouge sont décrits

et figurés. Ils sont déposés au Muséum national d'Histoire naturelle, à Paris et comprennent un mâle désigné

comme lectotype et une femelle paralectotype. Tous les deux sont en très mauvais état et n’avaient jamais été

réexaminés depuis leur description originale par Nobili, qui comporte plusieurs erreurs. Une clé de comparaison

avec les autres espèces du genre Naushonia est présentée.

Mots-clés. — Crustacea, Decapoda, Thalassinidea, Laomediidae, taxonomie, Naushonia , mer Rouge.

Redescription of the types of Naushonia perrieri (Nobili, 1904)

(Crustacea, Decapoda, Laomediidae)

Abstract. — The type material of Naushonia perrieri (Nobili, 1904) from Djibouti, Red Sea is described

and figured. The newly designated lectotype male and paralectotype female are deposited in the Muséum national

d'Histoire naturelle, Paris. Both specimens are in very poor condition and had never been reexamined since their

original description by Nobili, which contains a number of mistakes. A key to the known species of Naushonia

is provided.

Key-words. — Crustacea, Decapoda, Thalassinidea, Laomediidae. taxonomy, Naushonia, Red Sea.

N. Ngoc-Ho, Muséum national d’Histoire naturelle. Laboratoire de Zoologie IArthropodes), 61 rue de Buffon, F-75231 Paris

cedex 05.

INTRODUCTION

Le genre Naushonia Kingsley, 1897 comprend actuellement six espèces : N. crangonoides

Kingsley, 1897, N. portoricensis (Rathbun, 1901), N. perrieri (Nobili, 1904), N. macginitiei (Glas-

sell, 1938), N. panamensis Martin & Abele, 1982 et N. lactoalhida Berggren, 1992. Quatre d’entre

elles sont américaines et proviennent soit de la côte atlantique (N. crangonoides ), soit de la côte

pacifique (N. macginitiei), de la mer Caraïbe (N. portoricensis) ou du Panama ( N. panamensis).

Seules N. perrieri et N. lactoalhida ont été récoltées dans l’océan Indien, la première à Djibouti,

en mer Rouge, la deuxième en Mozambique.

N. perrieri a d’abord été décrite par Nobili (1904, 1906) sous le nom de Coralliocrangon

Perrieri et prend le nom actuel à la suite de la mise en synonymie de Coralliocrangon avec

Naushonia par Chace (1939). Le matériel-type de N. perrieri est déposé au Muséum national

d’Histoire naturelle, Paris et comprend deux spécimens de petite taille, un mâle désigné ici comme

lectotype et une femelle paralectotype. Tous les deux sont en très mauvais état et n’ont jamais

Source : MNHN, Paris

— 566 —

été réétudiés depuis la description originale de Nobili, description qui, comme ses illustrations,

comporte plusieurs erreurs.

Les types de N. perrieri sont redécrits et figurés afin de permettre une comparaison avec

les autres espèces du genre Naushonia. Les dimensions indiquées se rapportent à la longueur

de la carapace (l.c.) mesurée de l’extrémité du rostre au bord postérieur de la carapace.

Genre NAUSHONIA Kingsley, 1897

Naushonia perrieri (Nobili, 1904)

(Fig. 1)

Coralliocrangon Perrieri Nobili, 1904: 234; 1906: 82; DE Man 1920: 257.

Naushonia perrieri ; Chace 1939 : 527; GURNEY & LEBOUR 1939 : 609; Goy & Proven-

ZANO 1978 : 258; 1979 : 353, figs 6-8; Martin & Abele 1982 : 482; BERGGREN 1992 : 522.

Matériel examiné. — Djibouti, H. Coutière coll. 1897: 1 <3, Le. 6,5 mm (lectotype, MNHN-Th202a),

fragmenté en trois morceaux, mandibule et maxillule droites et premiers péréiopodes présents: 1 $, Le. 4,5 mm

(paraiectotype, MNHN-Th202b), fragmenté en deux, maxillule et maxille droites, premier et deuxième maxilli-

pèdes droits, pléopodes 1-5 présents.

Les spécimens ont été colorés par une solution diluée de noir de Chlorazol pendant une demi-heure environ

avant l'observation.

Description

Carapace (Fig. la), avec linea thalassinica distincte mais sillon cervical très faible, parcourue

longitudinalement, de la base du rostre environ jusqu'au bord postérieur de la carapace, par une

fine crête médiane accompagnée de chaque côté de deux crêtes latérales obliques, plus courtes,

ces dernières sont limitées à la partie antérieure de la carapace, en avant du sillon cervical.

Rostre large (Fig. If), cachant presque entièrement les yeux, arrondi en avant et bordé d’une

rangée de denticules, avec un faible sillon médian ; dents postorbitaires simples, fortes. Abdomen

(Fig. lb) à pleurons arrondis postérieurement, ceux du deuxième segment recouvrant en partie

ceux du premier et du troisième.

Pédoncules antennulaires (Fig. If, g) séparés à leur base par une longue épine du proépi-

stome, visible en vue ventrale; premier article avec une épine ventro-distale, deuxième article

avec deux épines dorso-distales ; flagelles courts.

Pédoncules antennaires (Fig. If, g) : troisième article avec une épine mésio-subdistale; qua¬

trième article avec une épine externe subdistale et une mésio-distale, face dorsale recouverte

d’une grande écaille antennaire avec quatre petites dents externes et bordée de soies du côté

interne.

Mandibule (Fig. lh) : bord sécant muni de petites dents obtuses, palpe à trois articles.

Premier maxillipède (Fig. li) : exopodite composé d’une partie basale élargie et d’une partie

distale étroite, se terminant par un fouet muni de soies ; endopodite à extrémité élargie en spatule

triangulaire; épipodite probablement cassé laissant une trace.

Deuxième maxillipède avec épipodite en lame triangulaire.

Source : MNHN, Paris

— 567 —

Source : MNHN, Paris

— 568 —

Premier péréiopode mâle (Fig. le, d) : mérus avec des denticules au bord distal et une ex¬

pansion arrondie denticulée ventro-externe, recouvrant une partie du carpe. Carpe triangulaire

muni de denticules au bord disto-exteme. Propode deux fois plus long que le mérus et environ

deux fois plus long que large, avec une grande dent vers le milieu du bord ventral, une grande

dent subdistale précédée par trois dents plus petites et des denticules; bord ventral finement

serrulé entre les deux grandes dents; une spinule externe distale au niveau de l’articulation avec

le dactyle. Dactyle d’une longueur environ égale aux deux tiers de celle du propode, arqué,

recourbé, inerme, à extrémité effilée.

Péréiopodes 2-5 manquant chez le lectotype, tous manquant chez le paralectotype, certaines

branchies persistent mais sont difficiles à observer, toutefois les épipodites sont présents sur les

P1-4.

Pléopode 1 absent chez le lectotype, présent, très petit, à deux articles chez le paralectotype.

Pléopodes 2-5 perdus chez le lectotype, présents chez le paralectotype, leurs deux rames lan¬

céolées.

Telson (Fig. le) légèrement plus long que large, inerme, à bord postérieur arrondi.

Uropodes (Fig. le) : basipodite bilobé; endopodite et exopodite à peu près de même lon¬

gueur, à bord postérieur arrondi ; endopodite avec une carène longitudinale, exopodite avec deux

carènes, toutes très faibles; endopodite et exopodite pourvus au tiers postérieur d’une suture

transversale munie de spinules et se terminant par une épine externe plus forte.

Remarques

L’examen des types de cette espèce permet d’accepter pleinement son placement par Chace

dans le genre Naushonia. Elle permet aussi de mieux définir l’espèce et de corriger certains des

caractères morphologiques mal observés dans la description originale de NOBILI.

Naushonia perrieri peut être définie comme une espèce à bord rostral antérieur arrondi,

denticulé; épines postorbitales simples; line a thalassinica distincte; sillon cervical faible; une

crête longitudinale médiane et deux paires de crêtes obliques sur la carapace ; écaille antennaire

avec quatre petites dents externes; palpe mandibulaire à trois articles; péréiopode 1 à mérus

inerme avec une expansion distale, denticulée, ventro-externe, propode inerme au bord dorsal,

bord ventral avec, sur la moitié distale, deux grandes dents, des dents plus petites et des den¬

ticules, dactyle inerme; telson inerme, à bord postérieur arrondi; uropodes avec exopodite et

endopodite à peu près de même longueur, à sutures complètes, munies de spinules. La formule

branchiale complète de l’espèce ainsi que la morphologie des P1-4 ne peuvent pas être précisées

actuellement.

À l’exception des épines postoculaires, simples, et des denticules du bord ventral entre les

deux grandes dents du propode des PI, N. perrieri ne présente pas de caractères communs avec

N. lactoalbida qui en est géographiquement l’espèce la plus proche. Elle se distingue aisément

des espèces américaines comme le montre la clé ci-dessous.

Clé de détermination des espèces du genre Naushonia

(adaptée de Martin & Abele 1982 et Berggren 1992) :

1. Épine postorbitaire bifide ou trifide . 2

— Épine postorbitaire simple . 4

Source : MNHN, Paris

— 569 —

2. Écaille antennaire avec dix dents externes ou plus; exopodite des uropodes avec cinq épines

externes, telson inerme. N. crangonoides

— Écaille antennaire avec moins de dix dents externes ; exopodite des uropodes avec deux épines

externes, telson muni de dents aux bords latéraux . 3

3. Telson avec une dent aux bords latéraux; dent terminale de l’écaille antennaire fortement

recourbée médialement. N. portoricensis

— Telson avec trois dents aux bords latéraux; dent terminale de l’écaille antennaire droite, non

recourbée médialement. N. macginitei

4. Rostre pointu vers l’avant. 5

— Rostre à bord antérieur arrondi ; carapace avec une crête longitudinale médiane et deux paires

de crêtes obliques; mérus de PI avec une expansion distale ventro-externe; telson inerme

. N. perrieri

5. Propode de PI avec deux fortes dents proximales ventrales, dactyle avec dents proximales

dorsales; telson avec une dent aux bords latéraux. N. panamensis

— Propode de PI sans dents proximales ventrales, dactyle inerme au bord dorsal; telson avec

quatre dents aux bords latéraux . N. lactoalbida

Remerciements

L'auteur remercie Michèle de Saint-Laurent qui a relu le manuscrit.

RÉFÉRENCES

BERGGREN M., 1992. — Naushonia lactoalbida, new species (Decapoda: Thalassinidea: Laomediidae), a mud

shrimp from Inhaca Island, Moçambique. J. Crust. Biol. 12 (3) : 514-522, figs 1-6.

CHACE F. A. Jr., 1939. — On the systematic status of the crustacean genera Naushonia. Homoriscus and Coral-

liocrangon. Ann. Mag. Nat. Hist. ser. II (3): 524-530, figs 1-14.

GLASSELL S. A., 1938. — New and obscure decapod Crustacea from the West American coasts. Trans. San Diego

Soc. Nat. Hist. 8 (33): 411-454.

GOY J. W. & PROVENZANO A. J., 1978. — Larval development of the rare burrowing mud-shrimp Naushonia

crangonoides Kingsley (Decapoda: Thalassinidea: Laomediidae). Biol. Bull. 154 : 241-261.

— 1979. — Juvenile morphology of the rare burrowing mud-shrimp Naushonia crangonoides Kingsley, with

a review of the genus Naushonia (Decapoda: Thalassinidea: Laomediidae). Proc. Biol. Soci. Wash. 92 (2):

339-359, figs 1-8.

GURNEY R. & Lebour m. V., 1939. — The larvae of the Decapod Genus Naushonia. Ann. Mag. Nat. Hist. ser.

11 (3): 609-614, figs 1-14.

KINGSLEY J. S., 1897. — On a new genus and two new species of macrurous Crustacea. Bull. Essex Inst. 27:

95-99.

MAN J. G. DE, 1920. — The Decapoda of the Siboga Expedition. Part IV. Families Pasiphaeidae, Stylodactylidae,

Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psalidopodidae, Gnathophyllidae, Proces-

sidae, Glyphocrangonidae and Crangonidae. Siboga Exped. Monogr. 39 (a3): 1-318, pis 1-25.

MARTIN J. W. & ABELE L. G., 1982. — Naushonia panamensis, new species (Decapoda: Thalassinidea: Laome¬

diidae) from the Pacific coast of Panama with notes on the genus. Proc. Biol. Soc. Wash. 95 (3): 478-483,

figs 1-3.

Source : MNHN, Paris

— 570 —

NOBILI G., 1904. — Diagnoses préliminaires de vingt-huit espèces nouvelles de Stomatopodes et Décapodes

macroures de la mer Rouge. Bull. Mus. nail d’Hist. nat.. Paris 10 (5) : 228-238.

— 1906. — Faune carcinologique de la mer Rouge : Décapodes et Stomatopodes. Ann. Sci. Nai. Zool. 9 (4) :

1-347, pis. 1-11.

RATHBUN M. J., 1901. — The Brachyura and Macrura of Porto Rico. Bull. U.S. Fish. Comm, for 1900. 2: 1-127.

Source : MNHN, Paris

Bulletin du Muséum

national d'Histoire naturelle, Paris, 4 e série, 18, 1996

Section A, n" 3-4:571-587

Découverte du genre Archipolydesmus en Algérie :

description de trois espèces nouvelles

(Diplopoda, Polydesmida, Polydesmidae)

par Ourida ABROUS-KHERBOUCHE et Jean-Paul MAURIÈS

Résumé. — Le genre Archipolydesmus, jusqu'à présent connu par cinq espèces de France, d'Espagne et

du Maroc, est récolté pour la première fois en Algérie. Trois espèces nouvelles sont décrites. Elles ont été dé¬

couvertes dans deux parcs nationaux du Nord de l’Algérie : une dans celui de Chréa (sud d’Alger) et les deux

autres dans celui du Djurdjura (est d’Alger). Une liste récapitulative des vingt-cinq genres et cinquante et une

espèces de diplopodes cités d’Algérie est donnée, avec leurs références bibliographiques.

Mots-clés. — Archipolydesmus, Diplopoda, Algérie.

Discovery of genus Archipolydesmus in Algeria :

description of three new species

(Diplopoda, Polydesmida, Polydesmidae)

Abstract. — The genus Archipolydesmus , until now known by five species from France, Spain and Morocco,

is discovered for the first time in Algeria, in two national parks. Three new species are described : one of them

in the National Park of Chrea (south of Algiers), and two in the National Park of Djurdjura (east of Algiers).

A list of the Millipeds (twenty-five genera - fifty-one species) cited until today from Algeria is given, with their

bibliographic references.

Key-words. — Archipolydesmus, Diplopoda, Algeria.

O. Abrous-Kherbouche, Laboratoire d'écologie animale, I.S.N., U.S.T.H.B., B.P 32. El Alia. Bah-Ezzouar, Alger (Algérie).

J.-P. Mauriès. Muséum national d'Histoire naturelle. Laboratoire de Zoologie (Arthropodes), 61 rue de Buffon, F-75231 Paris

cedex 05.

INTRODUCTION

Durant l’étude effectuée par l’un de nous (O. A.-K.) sur la pédofaune du parc national de

Chréa, de décembre 1989 à décembre 1990 (Abrous-Kherbouche 1996), et sur celle du parc

national du Djurdjura, de novembre 1992 à novembre 1994, douze espèces de Diplopodes ont

été récoltées (tableau ci-après).

Source : MNHN, Paris

— 572 —

Liste des espèces et nombre d’individus récoltés dans les parcs nationaux de Chréa et du

Djurdjura.

Espèces

P. N.

Chréa

P. N.

Djurdjura

Cylindroiulus (Phalloiulus) distinctus (Lucas, 1846)

244

Cylindroiulus (Phalloiulus) aff. algerinus (Lucas, 1846)

Ommatoiulus gauthieri (Brôlemann, 1931)

Ommatoiulus aumalensis (Brôlemann, 1925)

347

Brachyiulus sp.

2 j.

Glomeris conspersa C.L. Koch, 1847

Glomeris flavomaculata Lucas, 1846

Polydesmus (Brachydesmus) superus Latzel, 1884

Polydesmus (Brachydesmus) proximus Latzel, 1889

Archipolydesmus chreensis n.sp.

Archipolydesmus kabylianus n.sp.

Archipolydesmus fodili n.sp.

210

Dans cette liste, les trois dernières sont des espèces inédites qui appartiennent à un genre

qui n’était encore connu que de l’extrême Sud de la France, de l’Espagne et du Maroc :

Archipolydesmus Attems, 1898 (= Hispaniodesmus Verhoeff, 1910 = Miradoria Attems, 1952).

Ce genre qui, dans la famille des Polydesmidae (cf. HOFFMAN 1980) est proche du grand

genre Polydesmus, s’en distingue par un caractère morphologique très visible extérieurement :

la présence sur le dos des métatergites, non de simples mamelons plus larges que longs, peu

proéminents et peu nombreux, mais de nombreux petits tubercules serrés, plus longs que larges,

et surmontés d’une soie (Verhoeff 1910). Du point de vue des caractères gonopodiaux, Archi¬

polydesmus se distingue de Polydesmus par l’absence de pulvillum et la forme, particulière au

genre, très allongée et sinueuse de la vésicule séminale.

La première espèce décrite l’a été du Maroc, en même temps que le genre : Archipolydesmus

maroccanus Attems, 1898. Puis, ont été décrites successivement :

— panteli (Brôlemann, 1900) du Centre-Est de l’Espagne (prov. Cuenca), type de Hispa¬

niodesmus Verhoeff, 1910;

— hedeli (Brôlemann, 1902) d’Espagne centrale (prov. Segovia);

— ribauti (Brôlemann, 1926) de l'extrême Sud de la France (Pyrénées-Orientales) et d’Es¬

pagne du Nord (prov. Gérona);

— terreus (Attems, 1952) du Pays basque espagnol, prov. Guipuzcoa (= Miradoria terrea)',

— ose liai Ceuca, 1968, du Nord de l’Espagne (prov. Huesca).

Source : MNHN, Paris

— 573 —

Archipolydesmus s’ajoute aux vingt-quatre genres de Diplopodes déjà connus de la faune

d’Algérie; quant au nombre des espèces recensées, il s’élève à cinquante et un, compte tenu

des quelques taxa douteux et des trois nouveaux. Ce nombre, qui est le plus fort des trois pays

du Maghreb, (vingt-quatre pour la Tunisie, quarante-cinq pour le Maroc) est relativement modeste,

comparé à celui d’autres pays circumméditerranéens, comme l’Espagne (150-170), la France (250-

300), ou l’Italie (450-500). Si cette faiblesse s’explique en partie par la pondération minorante

due à la situation géographique, notamment la proximité de la zone saharienne, elle indique

néanmoins que la faune myriapodologique de l’Algérie offre encore un potentiel de découvertes

assez important.

Trente-huit de ces espèces étaient déjà citées par Brolemann 1921, et quarante-deux par

Schubart 1953. Ensuite, pour ce qui concerne les diplopodes, les citations concernant les dix

autres espèces, nouvelles pour la Science ou non, citées d’Algérie, sont dispersées dans une

dizaine de travaux ponctuels, soit sur le plan taxinomique, soit sur le plan géographique. C’est

pourquoi il nous a paru nécessaire, dans un souci de mise à jour, de publier ici, en fin d’article,

la liste des diplopodes d’Algérie (dans la nomenclature actuelle), avec les références biblio¬

graphiques les concernant.

MATÉRIEL ÉTUDIÉ

Le matériel récolté récemment par le premier auteur dans les parcs nationaux mentionnés

plus haut, est le support principal du présent travail. Une partie (matériel-type notamment) a été

intégrée dans la «Collection Myriapodes du Muséum national d’Histoire naturelle», et le reliquat,

propriété du premier auteur, est mentionné sous l’abréviation «Coll. A.-K.». La présente étude

inclut en outre quelques matériaux issus des collectes « Biospeologica» qui avaient été récoltés

autrefois dans le Djurdjura par Peyerimhoff (n°436A, 715 et 916) et qui n’avaient jusqu’à ce

jour jamais été étudiés; ces derniers spécimens se rapportent tous à l’une des trois espèces dé¬

crites, la deuxième, A. kabylianus.

DESCRIPTION DES ESPÈCES NOUVELLES

Les trois espèces algériennes décrites ci-après ne se distinguent pas de leurs congénères

par leurs caractères externes; le nombre des aires polygonales métatergales est le même que

chez quatre des six espèces (seuls ribauti et panteli en ont davantage).

En revanche, les gonopodes des mâles, qui possèdent le court solénomérite prolongeant

l’ampoule spermatique (vésicule séminale) très allongée et sinueuse, caractéristique du genre,

se singularisent par une plus grande simplicité de l’ensemble du gonopode et par un relativement

grand développement, et en largeur, et en longueur, de la partie tibiotarsale.

L’importance en nombre du matériel examiné a permis d’apprécier l’inhabituelle grande

variabilité de la structure de ces gonopodes, qui se traduit de deux manières :

Source : MNHN, Paris

— 574 —

1) chez les trois espèces, par de notables différences intraspécifiques dans le développement

des découpures en lobes ou dents du tibiotarse et des différents processus dentiformes qui ornent

le fémur ;

2) chez la première espèce décrite, chreensis, où il existe deux formes sympatriques qui,

du point de vue des caractères externes, ne se distinguent que par la taille (on observe quelques

grands individus, et de nombreux petits), les gonopodes de la petite forme sont plus simples

que ceux de la grande.

Concernant les caractères des femelles, il faut rappeler que ceux d’une seule espèce sont

connus, ceux de la plus septentrionale, A. ribauti. La comparaison avec les espèces algériennes

est très décevante, car les variations morphologiques observées sont si faibles qu’elles n’appa¬

raissent pas comme caractéristiques ou propres à définir telle ou telle espèce : les vulves sont

identiques. Le profil des crêtes pleurostemales transverses montre une assez grande variabilité

intraspécifique, et les P.2 chez qui, rappelons-le, coxites et sternite sont soudés en une plaque

coxosternale subtrapézoïdale formant clapet au-dessus des vulves, ne montrent que de faibles

variations dont il est difficile d’apprécier la nature et la valeur taxinomique.

En outre, se pose pour nous, un autre problème, qui concerne plus précisément les deux

espèces du Djurdjura, qui sont sympatriques. Curieusement en effet, toutes les femelles récoltées

sur les différents sites de ce massif sont toutes de petite taille, contrairement aux mâles qui, en

plus des caractères gonopodiaux, se distinguent aisément par leur taille : kabylianus est plus

grand que fodili. S’il est relativement aisé d’attribuer certaines de ces femelles à A. fodoli, on

ne peut en attribuer aucune autre à kabylianus, à moins d’admettre que chez cette espèce, les

femelles sont nettement plus petites que les mâles, ce qui nous paraît peu vraisemblable et en

tout cas serait une «rareté» chez les myriapodes. C’est pourquoi nous ne donnons pas ici les

caractères femelles de kabylianus, le problème qu’ils posent nécessitant des investigations sup¬

plémentaires.

Archipolydesmus chreensis n.sp.

Localité-type. — Algérie, Wilaya de Blida, atlas de Blida, parc national de Chréa : Pic Abd-el-Kader,

versant nord, ait. 1600 m, pièges en forêt de Cedrus atlantica, 19.11.1990, 1 d holotype (petite forme) (MNHN

Paris, Collection Myriapodes, JC 291).

Topotypes. — 18.XI1.1989, 30 dd (28 petits, 2 grands) et 3 2 2 ; 27.1.1990, 10 dd (9 petits, 1 grand),

6 2 2 (MNHN Paris, Collection Myriapodes, JC 291).

Caractères morphologiques externes

Coloration claire, souvent dépigmentée, souvent ochracée, plus claire antérieurement et ven-

tralement; pattes jaunâtres.

Source : MNHN, Paris

— 575 —

1 B

FIGS 1-4. — Archipolydesmus chreensis n.sp. S holotype, <3 lopotypes, 9 paralype : 1, moitiés droites des huitièmes (IA) et

neuvièmes (IB) anneaux d'un â paratype. 2, gonopode gauche du <J holotype. en vue caudale. 3. le même en vue médiale.

4. gonopode gauche d'un â topotype (du 18.XII.1989), en vue médiale.

Source : MNHN, Paris

— 576 —

Corps bombé, à côtés parallèles, à carènes faiblement saillantes : parmi les mâles récoltés

sur la localité type, il a été constaté la coexistence de petits (de beaucoup les plus nombreux,

dont le mâle holotype et trente-sept des quarante mâles examinés) et de plus grands individus

qui ne se distinguent ni par l’ornementation métatergale, ni par d’autres caractères externes.

Mensurations du mâle holotype, de deux mâles (un petit et un grand) et d’une femelle

topotype (en mm) :

S holotype

Petit S

Grand 6

Longueur du corps

8,50

7,90

10,50

Largeur de la tête

0,81

0,84

1,20

1,15

Largeur du collum

0,60

0,66

0,90

T.7 : largeur

1,10

1,15

1,30

1,25

T.7 : hauteur du métazonite

0,80

0,85

1,20

T.7 : diamètre du prozonite

0,70

1,10

0,90

Tête courte, convexe, couverte d’une pilosité courte et dense. Bourrelet sou s-an tennaire

arrondi en arrière.

Antennes assez longues, à sept antennomères en cylindres modérément claviformes, mesurant

0,10 + 0,16 + 0,26 + 0,20 + 0,21 + 0,28 + 0,08 = 1,19 mm chez le mâle holotype.

Stipes mandibulaires bombés.

Collum (T.l) plus étroit que la tête, en ovale-réniforme (très convexe en avant, faiblement

échancré en arrière). Sa surface est légèrement bombée et découpée en petites aires polygonales

faiblement convexes portant chacune en son centre une petite soie claviforme; ces polygones

sont répartis en quatre rangs : un rang antérieur de 7 + 7 polygones disposés en arc de cercle

épousant le bord antérieur; le deuxième rang est moins arqué et compte 4 + 4 polygones; les

troisièmes et quatrièmes rangées sont sinueuses et comptent respectivement 4 + 4 et 3 + 3 po¬

lygones.

Si le collum (surtout) et le T.3 sont plus étroits que la tête (joues comprises), le T.2 est

légèrement plus large. A partir du T.4, qui a la même largeur que la tête, la largeur des anneaux

croît assez rapidement pour atteindre sa largeur maximale au T.7. Les carènes sont un peu re¬

poussées vers l’avant sur les anneaux antérieurs, notamment les T.3 et T.4.

A partir du T.2, tous les anneaux comptent trois rangs de polygones sétifères (Figs IA, B).

Ces polygones, en général au nombre de 6 + 6 (quelquefois 5 + 6 ou même 5 + 5 sur les anneaux

antérieurs ou postérieurs) sont plats, de forme allongée sur les premiers et deuxièmes rangs, de

forme arrondie sur les troisièmes où ils sont peu saillants, sauf les deux ou trois plus externes,

qui forment des cônes saillants en arrière, à l’instar de l’angle postérieur des carènes.

Sur les anneaux non porifères, le bord externe des carènes est faiblement pluri-échancré

en une (ou deux ou trois) faible(s) denticulation(s) antérieure(s) suivie(s) de trois lobules sétifères

dont le plus postérieur forme l’angle caudal externe de la carène, qui est séparé du cône le plus

Source : MNHN, Paris

— 577 —

Source : MNHN, Paris

— 578 —

Source : MNHN, Paris

— 579 —

externe du rang postérieur de polygones sétifères par une échancrure peu profonde sur les anneaux

antérieurs, en angle droit sur les moyens, en angle plus aigu sur les postérieurs.

Les carènes des anneaux porifëres (Fig. IB) diffèrent par l’existence d’un lobule sétifère

supplémentaire, les deux lobes postérieurs constituant une grosse verrue globuleuse saillante dans

laquelle s’ouvre latéralement l’ozopore.

L’angle postérieur des carènes est aigu sur les T. 16, T. 17 et T. 18 et, sur les deux derniers

(T. 17 et T. 18) ne dépasse que légèrement le bord postérieur; sur le T. 19, il ne dépasse pas et

la verrue porifère est très réduite et non saillante.

Les soies tergales, toujours très petites sont généralement en massue ou en bâtonnet; elles

sont parfois, sur les polygones antérieurs, plutôt spiniformes.

Le telson est conique, à pointe émoussée, orné de verrues polygonales et de quatre rangs

transverses de dix, cinq, deux et deux soies dont les postérieures sont les plus longues. Valves

anales peu bombées, à rebords saillants. Valve sous-anale en triangle arrondi à sommet muni

d’une paire de soies longues.

Pattes assez longues, mesurant 1,15 mm de long chez le mâle holotype, dont 0,44 mm pour

le tarse, qui est grêle et se termine par un uncus simple et court.

Caractères sexuels du mâle

Gonopodes (Figs 2-4, 5-10) plus proches, par leur conformation générale [brièveté du so-

lénomérite (s) et simplicité du tibiotarse (tt)] des formes franco-espagnoles (panteli, ribauti et

osellai) que de l’espèce marocaine, type du genre. Ils se signalent par la plus grande longueur

relative du tibiotarse qui est élargi en raquette et se termine en une dent rétroarquée, mais surtout

par la partie fémorale. Celle-ci se distingue bien du tibiotarse (tt) par sa plus forte chitinisation

et est limitée distalement, à l’intérieur par deux dents érigées de longueur inégale (f et g), à

l’extérieur par un ressaut angulaire (h) d’où se détache le court solénomérite (s) et, ventralement

et plus proximalement, une dent aiguë (e) inclinée vers la base. À signaler que les gonopodes

des grands mâles présentent quelques différences (voir Figs 4-7), notamment la présence d’un

denticule (v) sur la face concave du tibiotarse et la réduction de la dent fémorale (g).

Hanches des P.7 plus écartées que celles de la P.6, à stemite excavé.

Caractères sexuels de la femelle (Figs 11-16)

Coxosternite des P.2 illustré figure 13 : le stemite (st) et les hanches (k), bien que distincts,

sont soudés en une seule plaque subtrapézoïdale qui vient s'ajuster au-dessus du vestibule vulvaire

et le ferme comme un clapet.

Au-dessus du vestibule vulvaire, l’arc pleuro-sternal s’élargit en un bandeau orné de chaque

côté d’un large lobe arrondi proéminent et, sagittalement, d'une courte pointe (Fig. 11).

Vulves (Figs 14-16) très semblables à celles de ribauti, par leur forme générale en hémi¬

sphère très fortement aplati parasagittalement, à profil latéral arrondi, à valves latérales externe

(ve) et interne (vi) fortement développées et très symétriques, à valves antérieure (opercule :

op) et postérieure (gorgerin : g) petites et situées très basalement. Si ce dernier (g) ne présente

aucune particularité qui le distingue des valves latérales, l’opercule (op) forme un clapet sub¬

trapézoïdal dont une aire subcarrée plus chitinisée est garnie de quatre très fortes soies (comme

chez ribauti). Dans le large espace non chitinisé ménagé entre les deux valves latérales, le cimier

(ci), marqué d’une quinzaine de canelures transverses, de forme ovale allongée (en vue distale-

Source : MNHN, Paris

— 580 —

orale, Fig. 14), est partagé en deux zones subégales par une crête longitudinale fortement pro¬

éminente dans et le long de laquelle court la gouttière apodématique.

P.3 (Fig. 12) se signalant par la présence, en avant et entre les coxites, d’une paire d’ex¬

croissances sternales triangulaires (ts).

Archipolydesmus kabylianus n.sp.

Localité-type. — Algérie, Wilaya de Tizi-Ouzou, massif et parc national du Djurdjura, à Tala-Guilef, versant

nord, ait. 1420 m, pièges en forêt de Cedrus atlantica, 17.V.1993, 1 6 holotype, 1 6 paratype (MNHN Paris,

Collection Myriapodes JC 292), 25 d 6 (Coll. A.-K.).

Topotypes. — 20.V1.1993. 1 6 (Coll. A.-K.).

Autres localités. — Algérie, Wilaya de Tizi-Ouzou : massif et parc national du Djurdjura, à Tala-Guilef,

versant nord, ait. 1420 m, pièges en pâturage. 17.V.1993, 1 6 ; 16.V. 1994, 3 66 (Coll. A.-K.); 16.V. 1994,

2 6 6, (MNHN Paris, Collection Myriapodes JC 292);

— massif et parc national du Djurdjura, à Tala-Guilef, versant sud, ait. 1420 m, pièges en pâturage,

17.V.1993, 4 (îd (Coll. A.-K.);

— Draa El Mizan, Tesserefts Tissoukdel, 31.X.1912, Biospeologica 715, 1 6 (MNHN Paris, Collection

Myriapodes JC 292) ;

— massif et parc national du Djurdjura, crête du Djebel Haiser, novembre 1912, Biospeologica hors série,

2 6 6 (MNHN Paris, Collection Myriapodes JC 292) ;

— Aïn El Hammam. Tala n'Tahalouant, 13.X.1916, Biospeologica 916, 1 6 (MNHN Paris, Collection

Myriapodes JC 292).

Les localités ci-après (dont certaines se retrouvent ci-dessus) ont fourni des femelles dont on n'est pas

certain qu’elles appartiennent bien à A. kabylianus. D’après leur taille elles seraient plutôt à classer parmi les

fodili (voir plus haut, dans l’introduction du § description des espèces) :

— massif et parc national du Djurdjura, à Tala-Guilef, versant nord, ait. 1420 m, pièges en forêt de Cedrus

atlantica , 17.V.1993, 15 9 9 topotypes (Coll. A.-K.);

— massif et parc national du Djurdjura, à Tala-Guilef, versant nord, ait. 1420 m, pièges en pâturage,

17.V.1993, 1 9 (MNHN Paris, Collection Myriapodes JC 294); 16.V.1994, 5 99 (Coll. A.-K.); 16.V.1994,

2 9 9 (MNHN Paris, Collection Myriapodes JC 294) ;

— Aïn El Hammam. Djebel Azerou Tidjer, Ifri (grotte) Maareb, 10.VII. 1910, I 9, 4 66 juv., 3 j.,

Biospeologica 436A (MNHN Paris, Collection Myriapodes JC 294).

Caractères morphologiques externes

D’aspect très semblable à la précédente, mais plus pigmentée, et possédant des gonopodes

assez voisins quoique distincts, cette espèce se signale par sa taille plus grande (c’est la plus

grande des trois espèces algériennes), voisine de 15 mm, et par quelques faibles différences de

la morphologie externe.

Mensurations (en mm) :

6 holotype

<î

Longueur du corps :

14,70

13,2 à 17,9

Longueur antenne :

2,20

1,60 à 2,07

Largeur du collum :

0,90

Largeur du T.4 :

1,40

Largeur max. du T.7 :

1,75

1,52 à 1,65

Source : MNHN, Paris

— 581 —

Source : MNHN, Paris

— 582 —

Les proportions observées sur les antennes sont semblables à celles de chreensis.

Au collum, le nombre d’aires polygonales est un peu plus grand : 8 + 8 au premier rang,

et respectivement 6 + 6, 5 + 5 et 4 + 4 aux trois autres rangs.

Comme dans l’espèce précédente, le collum (surtout) et le T.3 sont plus étroits que la tête

(joues comprises) et le T.2 est légèrement plus large. A partir du T.4, qui a la même largeur

que la tête, la largeur des anneaux croît assez rapidement pour atteindre un maximum au T.7.

Sur les anneaux, porifères ou non, les aires polygonales ont la même disposition que chez

chreensis, et leur nombre est tout aussi variable (5/6 + 5/6). La seule différence tient aux échan¬

crures des parties latéro-caudales, qui sont moins marquées, et au fait que les aires polygonales

médiodorsales du rang postérieur sont plus ou moins effacées.

Caractères sexuels du mâle

Gonopodes (Figs 17-23) assez variables se distinguant de ceux de chreensis par un port

plus robuste, l’atténuation de la dent tibiotarsale distale, la réduction de l’une (g) des deux dents

fémorales internes, et la réduction ou l’absence de dent fémorale ventrale (e).

Caractères sexuels de la femelle

Une étude et des prélèvements supplémentaires sont nécessaires pour fixer les caractères

de la femelle de kabylianus (voir plus haut, l’introduction du § description des espèces).

Archipolydesmus fodili n.sp.

Localité-type. — Algérie, Wilaya de Tizi-Ouzou, massif et parc national du Djurdjura, à Tala-Guilef, versant

nord, ait. 1420 m, pièges en forêt de Cedrus atlantica, 20.X1.1993. 1 tî holotype, 6 6 6, 2 $2 paratypes

(MNHN Paris, Collection Myriapodes JC 293), 48 66,2 2 2 paratypes (Coll. A.-K.).

Topotypes. — 17.V.1993, 6 6 6 , 5 2 2 (MNHN Paris, Collection Myriapodes JC 293), 43 6 6 , 22 2 2

(Coll. A.-K.); 20.VI.1993, 4 22 (Coll. A.-K.); 18.X.1993, 2 66 (Coll. A.-K.).

Autres localités. — Algérie, Wilaya de Tizi-Ouzou : massif et parc national du Djurdjura, à Tala-Guilef,

versant sud, ait. 1420 m, pièges en pâturage, 17.V.1993, 3 6 6 , 2 2 2 (Coll. A.-K.) ;

— massif et parc national du Djurdjura, à Tala-Guilef, versant sud, ait. 1420 m, pièges en forêt de Cedrus

atlantica, 17.V.1993, 4 66; 18.VII.1993, 1 6; 18.IX.1993, 1 2; 18.X.1993, 1 2 (Coll. A.-K.);

— massif et parc national du Djurdjura, à Tala-Guilef, versant nord, ait. 1420 m, pièges en pâturage,

17.V.1993, 1 6,2 22; 20.VI. 1993, 1 2 ; 20.XI.1993, 10<î<3,3 22; 16.V.1994, 5 6 6, 11 22; 15.VI.1994,

1 2 (Coll. A.-K.).

Étymologie. — Espèce dédiée au conjoint du premier auteur, en remerciement pour son aide très efficace

sur le terrain.

Caractères morphologiques externes

D’aspect et de coloration très semblable à la précédente, mais possédant des gonopodes

bien distincts, cette espèce ne se distingue extérieurement que par sa taille qui est nettement

plus faible, variant autour du centimètre ; elle est cependant un peu plus grande que chreensis.

Source : MNHN, Paris

— 583 —

Figs 24-29. — Archipolydesmus fodili n.sp. 6 holotype, 6 $ paratypes. 9 paralype : 24, gc

vue latérale externe. 25, la paire de gonopodes du S holotype : le droit est en place, e

déplacé par accident, montre sa face médiale. 26, gonopode droit d'un â topotype, vue

et partie distale des vulves d'une 9 paratype, en vue orale. 28, excroissances sternales d

de la même, en vue orale. 29, P.2 isolées de la même, vue orale.

onopode gauche du 6 holotype, en

-n vue orale, tandis que le gauche,

caudale. 27, bandeau pleurostemal

u quatrième anneau et base des P.3

Source : MNHN, Paris

— 584 —

Quelques mensurations (en mm) :

holotype

Longueur du corps :

9,50

8,70 à 10,30

10,40 à 12,50

Longueur antennes :

1,65

1,60 à 1,84

1,66 à 2,05

Largeur max. du T.7 :

1,20

0,90 à 1,30

1,26 à 1,40

Aux antennes, les proportions sont semblables à celles relevées sur les deux autres espèces.

Au collum, le nombre d’aires polygonales est identique à celui de kabylianus : 8 + 8 au

premier rang et, respectivement, 6 + 6, 5 + 5 et 4 + 4 aux trois autres rangs.

Caractères sexuels du mâle

Gonopodes (Figs 24-26) construits sur le même modèle que les deux autres espèces algé¬

riennes, mais se distinguant nettement par son tibiotarse, qui est ici divisé en deux branches,

(tt) et (u), d’épaisseur et de longueur inégales, par une large et profonde échancrure au fond

arrondi, située en position distale-exteme. La partie fémorale se signale par l'allongement plus

important de la dent proximale-ventrale (e) et de l’une des deux dents internes (0, l’autre (g)

étant réduite à un vague moignon.

Caractères sexuels de la femelle

Le coxostemite des P.2 (Fig. 29) ne se signale que par sa forme un peu plus arrondie que

chez chreensis.

Le bandeau pleuro-stemal du T.3 ne possède que les deux proéminences lobulaires latérales

(Fig. 27).

Les vulves ne se distinguent pas de celles de chreensis. C’est pourquoi elles ne sont pas

figurées ici. Elles ne se distinguent que par la relative faiblesse du marquage en canelures du

cimier; mais comme ce dernier est non chitinisé, il s’agit certainement de différences liées à

un état physiologique ou à un artefact de fixation.

Aux P.3, les deux triangles sternaux (ts) sont plus volumineux, moins aigus que chez chreen¬

sis et soudés sagittalement jusqu’à mi-hauteur (Fig. 28).

LISTE DES ESPÈCES DE DIPLOPODES CITÉES D’ALGÉRIE

Les citations bibliographiques concernant la présence en Algérie d’un taxon décrit d’un

autre pays sont entre {crochets} ; quelques-uns des taxons anciens, dont l’identité reste douteuse,

sont entre «guillemets».

Polyzonida

Hirudisoma remyi Schubart, 1964

Dolistenus savii Fanzago, 1874 (Pocock, 1892}

Source : MNHN, Paris

— 585 —

Craspedosomida

Meinerteuma edoughensis Mauriès, 1982

«Craspedosoma polydesmoides Leach, 1814» {Lucas, 1849)

Iulida

Thalassisobates littoralis (Silvestri, 1903) {Brolemann 1925}

Archichoneiulus drahoni (Giard, 1899)

Archichoneiulus brevicornis Brolemann 1921, 1923

Archichoneiulus crebresulcata Brolemann 1921, 1923

Microchoneiulus gracilis Brolemann 1921, 1923

Microchoneiulus baboricola Brolemann 1921, 1923

«Blaniulus corticalis Lucas, 1846 & 1849», Brolemann, 1897 (Microchoneiulus probable)

Ommatoiulus albosignatus Brolemann, 1931

Ommatoiulus fuscounilineatus (Lucas, 1846, 1849) Pocock, 1892, Brolemann, 1897

Ommatoiulus aumalensis Brolemann, 1925

Ommatoiulus lapidarium (Lucas, 1846, 1849), Brolemann, 1897

Ommatoiulus (diplurum) appendiculatum Brolemann, 1925

Ommatoiulus gauthieri Brolemann, 1931

Afropachyiulus oraniensis (Verhoeff, 1901)

Cylindroiulus truncorum (Silvestri, 1896) {Schubart, 1964}

= Cylindroiulus africanus Brolemann, 1897

Cylindroiulus algerinus (Brolemann, 1897)

Cylindroiulus distinctus (Lucas, 1846, 1849) Pocock, 1892, Brolemann, 1897

Cylindroiulus distinctus gauthieri Brolemann, 1931

Brachyiulus lusitanus (Verhoeff, 1898) (Schubart, 1964}

Polydesmida

Oxidus gracilis (C. L. Koch, 1847) {Schubart, 1964}

Oranmorpha guerini (Gervais, 1836)

Oranmorpha guerini atlanticum Brolemann, 1894

«Strongylosoma pallipes Olivier, 1792 {Lucas, 1849}»

Stosatea tropifera (Attems, 1903)

Melaphe blainvillei (Eydoux & Gervais, 1838), Lucas, 1849

= ? Melaphe mauritanica Lucas 1844, 1849

= ? Melaphe mauritanica geniculata Brolemann, 1910

Haplocookia mauritanica Brolemann, 1915

Polydesmus (Brachydesmus) proximus Latzel, 1889

= Brachydesmus insculptus Pocock, 1892

Polydesmus (Brachydesmus) superus Latzel, 1884

Polydesmus dismilus Berlese, 1891 {Brolemann, 1910}

Macellolophus rubromarginatus (Lucas, 1846, 1849)

= ? Macellolophus excavatus Verhoeff, 1931 {Ceuca, 1988}

Source : MNHN, Paris

— 586 —

Glomerida

Eupeyerimhoffia algerina Brôlemann, 1913

Glomeris conspersa C. L. Koch, 1847 = Gl. marmorata Brandt, 1840. 1841

Glomeris fuscomarmorata Lucas, 1846, 1849, Pocock, 1892

Glomeris flavomaculata Lucas, 1846, 1849, Pocock, 1892

Glomeris pustulata Latreille, 1804, (Brandt, 1840, 1841)

Glomeris sublimbata Lucas, 1846, 1849, Brôlemann, 1913

«Glomeris pustulata anisosticta (Brandt, 1840, 1841)»

Polyxenida

Phryssonotus platycephalus (Lucas, 1846, 1849)

Macroxenus rubromarginatus (Lucas, 1846, 1849), Brôlemann, 1917

Lophoproctus lucidus Chalande, 1888 {Seurat, 1930)

Polyxenus lagurus (L.) {Marquet & Condé, 1950)

Lophoproctinus inferus maurus Marquet & Condé, 1950

RÉFÉRENCES

ABROUS-KHERBOUCHE O., 1996. — Étude systématique et écologique des myriapodes dans le parc national de

Chrea (atlas Blidéen). Algérie. In J. J. Geoffroy, J.-P. MaurièS, M. Nguyen Duy Jacquemin (eds).

Acta Myriapodologica (9° Int. Congress), Mém. Mus. natl Hist. nat. : 163-174.

ATTEMS C., 1898. — System der Polydesmiden. I. Teil. Denks. Math.-Naturw. Kaiser!. Akad. Wiss., Wien 67 :

221-482.

— 1903. — Beitrâge zur Myriopodenkunde. Zoo!. Jahrb. Jena 18 (1) : 63-154.

— 1952. — Myriopoden der Forschungreise Dr. H. Franz in Spanien 1951 nebst übersicht über die Gesamte

iberische Myriopodenfauna. Eos. Madrid 28 (4) : 323-366.

Brandt J. F.. 1840. — Rapport sur les Oniscides et les Myriapodes de la régence d’Alger. Rev. Soc. cuvierienne,

— 1841. — Ueber die in Regenschaft Algier beobachteten Myriapoden. In Wagner's Reisen in der Regenschaft

Algier III. Leipzig.

BRÔLEMANN H. W., 1894. — Contributions à la faune myriapodologique méditerranéenne. Mém. Soc. Zool.

France : 437-462.

— 1897. — lulides d'Algérie. Ann. Sci. Nat., Zool. & Paléont. : 253-276.

— 1900. — Myriapodes recueillis en Espagne par le P. J. Pantel. Bull. Soc. Entom. France 6 : 131-133.

— 1902. — Description d’un nouveau Polydesme d’Espagne. Bull. Soc. Entom. France 1 : 13-14.

— 1910. — Symphyles, Pselaphognathes, Polydesmoïdes et Lysiopétalides (Première série). Arch. Zool. Exp.

gén., Biospeologica XVII 14 (7) : 99-453.

— 1913. — Glomerides, Myriapodes (Première série). Arch. Zool. Exp. gén., Biospeologica XXXI 25 (6) :

387-445.

— 1915. — Description d’un genre nouveau et d’une espèce nouvelle de myriapode d’Algérie. Bull. Soc. Hist.

Nat. Afr. Nord, Alger, juin 1915 7 (6) : 88-94.

— 1917. — Macroxenus. nouveau genre de myriapodes pselaphognathes. Bull. Soc. Hist. nat. Afr. Nord 8:

114-118.

BROLEMANN H. W.. 1921. — Liste des myriapodes signalés dans le nord de l’Afrique. Bull. Soc. Sci. nat. Maroc

1 (3-6): 99-110.

Source : MNHN, Paris

— 587 —

— 1923. — Blaniulidae, Myriapodes (Première série). Arch. Zool. Exp. gén., Biospeologica XLVIII 61 : 99-453.

— 1925. — Races nouvelles de Schizophyllum algériens. Bull. Soc. Hist. Nat. Afr. Nord. Alger, nov. 1925 16 :

245-253.

— 1926. — Myriapodes des Pyrénées-Orientales. Bull. Soc. Hist. Nat. Toulouse. 1926 (1927) 54 (2) : 233-267.

— 1931. — Myriapodes recueillis par M. le Dr H. Gauthier en Algérie. Bull. Soc. Hist. Nat. Afr. Nord 12 :

121-134.

Ceuca T.. 1968. — Archipolydesmus osellai (Diplopoda, Polydesmidae) une nouvelle espèce montagnarde de

l'Espagne. Mem. Mus. Civ. Stor. Nat. Verona 68 : 137-140.

— 1988. — Sur quelques diplopodes de la Péninsule ibérique et du nord de l’Afrique. Stud. Univ. Bahes-Bolyai.

Biologia 33 (2) : 40-48.

CONDE B., 1954. — Diplopodes Pénicillates d’Afrique septentrionale. Bull. Mus. Hist. Nat., Paris, 2 e sér. 26

(4) : 496-500.

EYDOUX F. & GERVAIS P.. 1838. — Myriapodes du voyage autour du monde de la Favorite, Zool., 2 e partie,

Paris: 177-180.

GlARD A., 1899. — Sur un myriapode cavernicole du Djurdjura, Blaniulus Drahoni n.sp. Ass. Fr. Avanc. Sci.,

27 e session, Nantes : 170.

HOFFMAN R. L., 1980. — Classification of Diplopoda. Muséum d’Histoire Naturelle, Genève, 1979 : 1-238.

Lucas H., 1846. — Notes sur quelques nouvelles espèces d’insectes (myriapodes) du nord de l’Afrique. Rev.

Zool. Soc. de Cuvier, Paris 9 : 283-289.

— 1849. — Histoire naturelle des animaux articulés. Tome I. Crustacés, Arachnides, Myriapodes, In Exploration

scientifique de l'Algérie pendant les années 1840-41-42, Paris.

MARQUET M. L. & CONDÉ B., 1950. — Contribution à la connaissance des diplopodes pénicillates d'Afrique et

de la région Madécasse. Mém. Inst. Sci. Madagascar, A 4 (1) : 113-134.

MAURIÈS J. P., 1982. — Une famille nouvelle et deux genres nouveaux de Cleidogonoidea, avec notes sur la

classification de la superfamille (Diplopoda, Craspedosomida). Steenstrupia, Copenhaguen 8 (6) : 165-176.

POCOCK R. I., 1892. — On the Myriapoda and Arachnida collected by Dr Anderson in Algeria and Tunisia. Proc.

Zool. Soc. London 1892 : 24-28.

SCHUBART O., 1953. — Diplopoden aus Marokko gesammelt vom Institut Scientifique Chérifien. Bull. Soc. Sci.

nat. Maroc 32 : 199-225.

— 1964. — Ueber einige Diplopoden aus Algier. Bull. Soc. Sci. Nat. & Phys. Maroc 1963 (1-2) : 79-94.

SEURAT L. G., 1930. — Exploration zoologique de l’Algérie de 1830 à 1930. Coll. Centenaire de l’Algérie, Paris

(Masson) : 1-708.

VERHOEFF K. W„ 1901. — Beitrage zur Kenntniss palaarktischer Myriopoden. XVII. Aufsatz : Diplopoden aus

dem Mittelmeergebiet. Arch, f Naturgesch. 1901 1 (1) : 79-102.

— 1910. — Ueber Diplopoden. 42. Aufsatz : Neue Polydesmiden aus Mitteleuropa und ihre Verwandten. Zool.

Anz. Leipzig 36 : 132-145.

Source : MNHN, Paris

Bulletin du Muséum national d Histoire naturelle. Paris, 4 e série, 18, 1996

Section A, n“ 34 : 589-594

Psalistops gasci n.sp., première Barychelidae de Guyane française

(Araneae, Mygalomorphae)

par Patrick MARÉCHAL

Résumé. — Psalistops gasci. de la famille des Barychelidae. est une nouvelle espèce décrite de Guyane

française. C’est la première fois qu'un mygalomorphe de cette famille est trouvé dans cette partie de la région

néotropicale. Cette espèce vit au niveau du sol des forêts humides.

Mots-clés. — Araneae, Mygalomorphae, Barychelidae, Guyane française, nouvelle espèce.

Psalistops gasci n.sp., first Barychelidae from French

Guiana (Araneae, Mygalomorphae).

Abstract. — One species of Barychelidae, Psalistops gasci. is described from French Guiana. This is the

first record of this mygalomorph family from this neotropical country. The species is a ground dweller occurring

in wet tropical forests.

Key-words. — Araneae, Mygalomorphae, Barychelidae. French Guiana, new species.

P. Maréchal, Muséum national d'Histoire naturelle. Laboratoire de Zoologie (Arthropodes). 61 rue de Buffon. 75231 Paris

cedex 05.

INTRODUCTION

La Guyane française est une région relativement peu explorée du point de vue des études

arachnologiques. Si l’on excepte les travaux de LOURENÇO (1983, 1991) sur les scorpions, les

autres ordres d’arachnides sont peu ou pas connus. Ainsi, le dernier travail d’importance sur les

araignées, publié par Caporiacco (1954) ne recense-t-il que 357 espèces pour ce département

dont quatorze mygalomorphes répartis en trois familles : les Idiopidae, les Dipluridae et les

Theraphosidae.

La description d’une nouvelle Barychelidae porte donc à quatre le nombre de familles de

mygales présentes en Guyane française.

Les Barychelidae américaines

Les Barychelidae du continent américain comprennent neuf genres (Raven 1994). Leur aire

de répartition englobe l’Amérique du Sud, les Caraïbes et une partie de l’Amérique centrale, le

Costa Rica constituant la limite nord de leur distribution (Valerio 1986). La présence de cette

Source : MNHN, Paris

— 590 —

famille n’a donc rien d’exceptionnel en Guyane française, où vivent certainement plusieurs autres

espèces.

Le genre Psalistops

Créé par Simon (1889), il a été initialement caractérisé par la position relative des yeux

les uns par rapport aux autres, critère trop vague pour des mygalomorphes. Redéfini par Raven

( 1985), les araignées du genre Psalistops se caractérisent par la présence d’une seule rangée de

dents sur les griffes tarsales paires, une aire oculaire deux fois plus large que longue, des tarses IV

sans scopula et un labium portant plus de dix denticules apicaux. Le genre Psalistops comprend

actuellement douze espèces présentes dans les Caraïbes, la moitié sud de l’Amérique centrale

et le nord de l’Amérique du Sud.

Psalistops gasci n.sp.

Matériel examiné. — Holotype 9, rivière Arataye, Saul Pararé (4°05'N - 52°40'W). Capturée le 18 février

1990 (P. Maréchal). Paratype 9, Montagne de Kaw (4°40’N - 52°15’W). Capturée en juin 1993 (C. Marty).

L’holotype et le paratype sont déposés au Muséum national d’Histoire naturelle de Paris.

Étymologie. — Cette espèce est dédiée au Prof. J.-P. Gasc grâce auquel j'ai pu prospecter le territoire

guyanais à deux reprises dans le cadre de l'action spécifique «Guyane» du Muséum national d’Histoire naturelle.

Diagnose

Psalistops gasci diffère de toutes les autres espèces du genre par la présence de longs poils

orangés disposés sur toute la surface des tibias et des métatarses III et IV, donnant à ces pattes

l’aspect d’écouvillons.

L’espèce diffère de P. melanopygius Simon, 1889 par des scopulas entières sur les tarses I

et II, et par l’article apical digitiforme des filières latérales postérieures.

Description

Holotype 9

Longueur totale sans les chélicères, 25 mm.

Couleur. Céphalothorax brun s’éclaircissant vers l’avant de la partie céphalique. Chélicères,

pattes, sternum et pièce labiale, marron clair, presque orange. Abdomen noir sur la face dorsale

avec de long poils blanchâtres, beige sur les côtés et sur la face ventrale.

Carapace. Longueur 9,5 mm; largeur 8,25 mm. Recouverte d’une pubescence blanchâtre

et clairsemée, plus dense et composée de poils plus longs sur les bords et vers la fovéa.

Fovéa. Profonde et nettement procurvée.

Yeux. (Fig. lb) Tubercule distinct. Aire oculaire occupant un quart de la partie céphalique

en largeur. Yeux disposés en un rectangle compact, largeur 1,6 mm; longueur 1,15 mm. Yeux

antérieurs en ligne procurvée, les médians contigus: yeux postérieurs en ligne récurvée.

Chélicères. Glabres à leur base, mais recouvertes d’une pubescence blanchâtre dense sur

le dessus, plus clairsemée sur les côtés. Elles possèdent, en position dorso-interne, des soies

Source : MNHN, Paris

— 591 —

Fig. 1. — Psalistops gasci n.sp., holotype 9. a, sternum, lames maxillaires et labium, vue ventrale, b, aire oculaire, vue dorsale,

c, filières, vue ventrale. Echelle : 1 mm.

Source : MNHN, Paris

— 592 —

noires à la base s'éclaircissant progressivement, en disposition plus dense sur la partie antérieure

de l'article basal. Marge interne munie de dix à douze dents coniques. Sillon ventral pourvu à

la base d'une plage comprenant une quinzaine de denticules. Pas de rastellum.

Labium. (Fig. la) De forme trapézoïdale. Longueur 1,6 mm; largeur à la base 2,3 mm.

Muni de trente-cinq à quarante denticules apicaux. Suture labio-sternale composée de deux

dépressions communicantes.

Lames maxillaires. (Fig. la) Longueur antérieure 3,15 mm; longueur postérieure 3,95 mm;

largeur 2,2 mm. Pourvues de cinquante à soixante denticules dans leur angle interne.

Sternum. (Fig. la) Longueur 4,7 mm; largeur 4,4 mm. Sigillés ronds, les postérieurs les

plus gros, les deux paires antérieures peu distinctes.

Pédipalpe. Muni d'une griffe tarsale mutique, en partie cachée par des fascicules unguéaux

denses et irisés. Scopula peu dense et localisée sur les deux tiers apicaux des tarses.

Pattes. Griffes paires munies d’une rangée médiane de cinq à six dents. Fascicules unguéaux

moins denses sur les tarses III et IV. Scopula entière sur les tarses et la moitié apicale des

métatarses I et II, largement coupée par une bande médiane de longues soies noires sur les

tarses III, absente sur les tarses IV. Tibias et métatarses III et IV portant, sur toute leur surface,

de longs poils orangés donnant à chacune de ces pattes l’aspect d'un écouvillon (Fig. 2).

Mensurations (mm)

III

Fémur

6,5

7,5

Patelle

4,5

3,25

Tibia

5,5

3,75

5,5

3,75

Métatarse

3,25

5,5

7,5

Tarse

2,75

2,5

Total

22,5

21,25

21,5

27,5

Épines. Pédipalpe. Tibia, face ventrale : deux apicales et une prolatérale.

Patte I. Métatarse, face ventrale ; une apicale et une médiane.

Patte II. Métatarse, face ventrale : trois apicales et deux médianes. Tibia, face ventrale :

une apicale et une médiane; face dorsale : une prolatérale au tiers antérieur du segment.

Patte III. Métatarse, face ventrale ; trois apicales, deux médianes et trois prolatérales; face

dorsale : trois prolatérales et deux rétrolatérales. Tibia, face ventrale : deux apicales, une médiane

et une prolatérale; face dorsale ; deux prolatérales et deux rétrolatérales.

Patte IV. Métatarse, face ventrale : trois apicales, deux médianes et deux prolatérales; face

dorsale : trois prolatérales et trois rétrolatérales. Tibia, face ventrale : deux apicales, une médiane

et une prolatérale; face dorsale : deux prolatérales et deux rétrolatérales.

Filières. (Fig. le) Longueur des postérieures médianes 0,98 mm. Longueur des postérieures

latérales 5 mm; article basal 2 mm; article médian 1,4 mm; article apical 1,6 mm. L’article

apical est digitiforme.

Source : MNHN, Paris

— 593 —

Paratype 9

Plus grande, cette femelle est certainement plus âgée et possède au moins un stade de

développement supplémentaire.

Longueur totale sans les chélicères 29 mm. Carapace, longueur 11 mm; largeur 9,5 mm.

Longueur de l’abdomen 18 mm.

Deux caractères diffèrent légèrement par rapport à l’holotype. La marge interne des chéli¬

cères porte douze à treize dents et les denticules apicaux du labium sont un peu plus nombreux.

Ces valeurs évoluant au cours des stades de développement chez Acanthogonatus francki (Ne-

mesiidae) (CALDERON corn, pers.), les écarts constatés sont vraisemblablement dûs à la différence

d’âge des deux individus.

L’observation du nombre des épines montre également de très légères différences; l’épine

prolatérale sur la face ventrale du tibia manque sur les deux pédipalpes. La face ventrale du

tibia III porte, dans sa partie basale, une épine surnuméraire fine et effilée. Le tibia IV possède

trois épines apicales sur sa face ventrale. Le nombre et la répartition des autres épines sont

strictement identiques à ceux observés sur l’holotype.

Les deux individus sont en tout point semblables pour l’ensemble des autres caractères.

Le mâle est inconnu.

Habitat

L’holotype a été trouvé sous une branche tombée au sol à quelques mètres de la rivière

Arataye, au niveau du saut Pararé. Aucune construction, tissage ou terrier, n’a été observée.

Juvénile, l’individu a été élevé au laboratoire où il a subi trois mues avant d’atteindre l’âge

adulte en novembre 1990. Cette femelle est morte un mois plus tard.

FIG. 2. — Psalislops gasci n.sp., vue dorsale de la patte droite; métatarse (Mt) et tibia (t) III. Échelle; 1 mm. (Cliché Dr

J. KOVOOR).

Source : MNHN, Paris

— 594 —

Le paratype a été collecté dans la montagne de Kaw et maintenu quelque temps en captivité.

La biologie de cette espèce reste inconnue. Cependant, les observations d’élevage et de

collecte laissent supposer la confection d’une loge soyeuse au niveau du sol en utilisant des

abris ou anfractuosités naturels, car ces deux individus ont rempli leur enceinte d’élevage d’un

abondant tissage. Psalistops gasci ne semble pas creuser de terrier, hypothèse appuyée par l’ab¬

sence de rastellum, structure généralement présente chez les espèces fouisseuses de cette famille.

Remerciements

Mes plus vifs remerciements au Dr R. J. Raven pour sa patience, sa disponibilité et ses nombreux

conseils relatifs à la systématique des araignées mygalomorphes.

RÉFÉRENCES

CAPORIACCO L. DI, 1954. — Araignées de la Guyane française du Muséum national d’Histoire naturelle de Paris.

Commentai, pontif Acad, scient. 16 (3) : 45-193.

LOURENÇO W. R., 1983. — La faune des scorpions de Guyane française. Bull. Mus. nail Hist. nat.. Paris, sér. 4,

A 5 (3) : 771-808.

— 1991. — La «province» biogéographique guyanaise; étude de la biodiversité et des centres d’endémismes

en vue de la conservation des patrimoines génétiques. C. R. Soc. Biogéogr. 67 (2) : 113-131.

Raven R. J., 1985. — The spider infraorder Mygalomorphae (Araneae): Cladistics and systematics. Bull. Am.

Mus. nat. Hist. 182 : 1-180.

— 1994. — Mygalomorph spiders of the Barychelidae in Australia and the Western Pacific. Mem. Qld. Mus.

35 (2): 291-706.

SIMON E., 1889. — Voyage de M. E. Simon au Vénézuela (Décembre 1887-Avril 1888). 4 e mémoire. Ann. Soc.

ent. Fr. 9 (6) : 169-220.

Valerio C. E., 1986. — Mygalomorph spiders in the Barychelidae (Araneae) from Costa Rica. J. Arachnol. 14:

93-99.

Source : MNHN, Paris

itional t l'Histoire naturelle. Paris. 4 e série. 18. 1996

Section A. n 05 3-4 :595-609

Liste des types d’oiseaux des collections du Muséum national

d’Histoire naturelle de Paris. 4. Hérons (Ardeidae)

par Claire VOISIN & Jean-François VOISIN

Résumé. — Les types de dix-huit taxons de Hérons ( Ardeidae) sont passés en revue de façon critique.

Des lectotypes et des paralectotypes sont désignés pour A idea novae-hollandiae Vieillot, 1817 et pour Ardea

australasie Vieillot, 1823 (les deux descriptions sont fondées sur les mêmes spécimens), ainsi que pour Ardea

Inline ata Cuvier in Lesson 1831. Quatre spécimens supplémentaires, portant la mention «type» mais qui n'en

sont pas au sens du Code, ont aussi été examinés.

Mots-clés. — Oiseaux, types, Ardéidés.

List of the types of birds in the collections of the Muséum national d’Histoire naturelle of Paris.

4. Herons (Ardeidae).

Abstract. — The types of eighteen heron ( Ardeidae ) taxa are reviewed critically. Lectotypes and paralec¬

totypes are designated for Ardea novae-hollandiae Vieillot. 1817, Ardea australasie Vieillot, 1823 (both descrip¬

tions are based on the same specimens), and for Ardea bilineata Cuvier in Lesson, 1831. Four additional

specimens, indicated as « types ». but which are not according to the meaning of type in the Code of Zoological

Nomenclature, were also examined.

Key-words. — Birds, types. Ardeidae.

C. Voisin & J.-F. Voisin. Muséum national d' Histoire naturelle. Laboratoire de Zoologie. Mammifères et Oiseaux, 55 rue de

Buffon. F-75231 Paris Cedex 05.

CATALOGUE

Ce travail fait suite à trois articles précédents concernant les types d’oiseaux du Muséum

national d’Histoire naturelle (J.-F. Voisin 1992, 1995; C. Voisin 1993) et suit les mêmes conven¬

tions.

Pour chaque spécimen nous avons noté successivement :

- le nom sous lequel il a été décrit;

- la référence de la description;

- la catégorie de type à laquelle il appartient;

- le numéro d’inscription au Catalogue Général ;

- le nom qu’il porte dans la nomenclature actuelle. Nous avons en général suivi Hancock

& Elliott (1978).

Les mentions qui se trouvent sous le socle ont toutes été recopiées en commençant, dans

la mesure où il est possible de les identifier, par les plus anciennes. Elles s'échelonnent pour

Source : MNHN, Paris

— 596 —

certaines de la fin du xvm e siècle à nos jours. Les mentions portées sur les étiquettes des socles

ont également été recopiées. Les retours à la ligne sont indiqués par le symbole /. Nous en

avons respecté l’orthographe.

Remarques

Les numéros de catalogue compris entre 1 à 15566 et non précédés de la mention N. C.

(Nouveau Catalogue) correspondent à l’ancien catalogue établi au XIX e siècle. Les numéros pré¬

cédés de la mention N. C. correspondent au catalogue des oiseaux montés de la Grande Galerie,

établi durant la première moitié du XX e siècle par le Professeur J. Berlioz. Les numéros précédés

de la mention C. G. et d’un millésime correspondent au Catalogue Général actuellement en

vigueur. Tous ces documents sont conservés au laboratoire des Mammifères et Oiseaux.

Les socles de quelques spécimens portent la mention «Type de Buffon» ou bien «type de

la planche de Buffon. » Bien qu'ils aient servi à Buffon pour ses descriptions et pour l’illustration

de son Histoire Naturelle des Oiseaux dont la première édition date de 1771-1786, ils ne sauraient

être considérés comme des types au sens du Code de Nomenclature Zoologique (1985) car

Buffon n’employait pas la nomenclature binominale de Linné.

TYPES DE TAXONS DONT LA DÉNOMINATION EST ENCORE EN VIGUEUR

Ardea heliosylus Lesson, 1828

Lesson, in Duperrey (ed.) Voyage de la Coquille 1822-1825 ; LESSON & Garnot Zool.

1: 722, Atlas, Zool. Oiseaux planche 44.

Type par monotypie.

C. G. 1995 - 236.

Zonerodius heliosylus (Lesson, 1828).

Inscriptions sous le socle :

Botaurus heliosylus / (Less.) / Lesson et Gamot / N.Guinée / Exped. Duperrey / Type

de l’espèce / [écriture ancienne] Butor tigré n. Guinée / Zonerodius heliosylus / (Less.)/

(Type) / La Coquille - N. Guinée / N. C. 158 [écriture plus récente]

Inscriptions sur l’étiquette :

Butor tigré de N. Guinée / Zonerodius heliosylus / (Less.) / (Type) / La Coquille

N. Guinée.

Remarques : la description de Lesson, précise et détaillée, comprend une page entière dans

le livre et une planche en couleur du spécimen dans l’atlas. L’oiseau a été tiré par M. Roland

«sur les rivages du hâvre de Doréry, Nouvelle Guinée». Lesson a fait partie de l’expédition

Duperrey sur la corvette la Coquille. L’auteur de la description est donc également le collecteur.

Buphus bacchus Bonaparte, 1855

Bonaparte, Consp. Gen. Avium : 127.

Type par monotypie.

Source : MNHN, Paris

— 597 —

C. G. 239 - 239.

Ardeola bac chus (Bonaparte, 1855).

Inscriptions sous le socle :

Ardea bacchus. (Bp.) / (Type) / Calcutta / par M Barbe / juillet 1840 / [écriture ancienne]

/ 13948 B [écrit au crayon] N. C. n° 100 / Crabier de Chine / Ardeola r. bacchus / (Bp.)

/ type / M. Barbe / Calcutta / [écriture plus récente].

Inscriptions sur l’étiquette :

Crabier de Chine / Ardeola r. bacchus / (Bp.) / Type / M. Barbe / Calcutta.

Remarque : dans sa diagnose Bonaparte précise que le spécimen décrit a bien été rapporté

par M. Barbe.

Ardea leuconotus Wagler, 1827

Wagler, Systema Avium , Ardea : 189 sp. 33.

Type par monotypie.

C. G. 1995 - 245.

Gorsachius leuconotus (Wagler, 1827).

Inscriptions sous le socle :

Ardea leuconotos, Wag. type de / Wagler. / Ardea leuconotos. (Tem) / Bihoreau à dos

blanc / [écriture ancienne] Sénégal par Watrin. 1817 / N. C. 124 Nycticorax leuconotus /

(Wag.) Type / 14013 Bihoreau à dos blanc [mention au crayon] Bihoreau à dos blanc /

Nycticorax leuconotus (Wagl.) / M. Watrin. Sénégal [écriture récente].

Inscriptions sur l’étiquette :

Bihoreau à dos blanc / Nycticorax leuconotus / (Wagl.) Type / M.Watrin Sénégal.

Remarques : Wagler a séjourné à Paris en 1825 lors de la rédaction de son livre Systema

Avium (Stresemann 1951). Dans sa description, l'auteur précise que le type se trouve au Muséum

de Paris et provient de Sénégambie. La référence à Temminck sous le socle sans la mention

«type» et sans renvoi à une planche coloriée ou à une description montre que ce spécimen n’est

pas un type de Temminck.

Nycticorax goisagi Temminck, 1835

Temminck, Planches coloriées , Livre 98. Description et planches en couleurs, planche 582.

Type par monotypie.

C. G. 1995 - 241.

Gorsachius goisagi (Temminck, 1835).

Inscriptions sous le socle :

Botaurus melanolophus / (Raffl.) / Gorsakius typus / Puch. type. / échangé à Temminck /

1836 [écriture ancienne] / 14006 / décrit Consp. Av. II p. 138 / V.pl. Col. 582 / [écriture

ancienne] / N. C. n° 148 / Petit butor de Malaisie / Gorsachius goisagi / (Temm) / Japon :

[écriture plus récente].

Inscriptions sur l’étiquette :

Petit Butor du Japon / Gorsachius goisagi / (Temm.) / Japon.

Source : MNHN, Paris

— 598 —

Remarques : c'est ce spécimen qui a servi de modèle à la planche coloriée de Temminck

lors de la figuration de l’espèce. Il fut désigné comme «type de la planche» dès cette époque,

d’où les mentions sous le socle : «V. PI. Col. 582» et «échangé à Temminck».

Par la suite Bonaparte (1857) s’est mépris sur l’identité de Gorsachius melanolophus (Raf¬

fles) qu’il considère comme l’adulte de Gorsachius goisagi (Temm.), alors qu’il en voit la forme

juvénile dans Gorsachius goisagi lui même (voir également la partie : espèces considérées à tort

comme types). Ainsi lorsque Bonaparte crée le genre Gorsachius, il ne cite que l’espèce goisagi.

La confusion qu’il fait avec Gorsachius melanolophus n’empêche pas de considérer Gorsachius

goisagi comme espèce-type du genre Gorsachius car, bien qu’il ait cru avoir affaire à un individu

juvénile, Bonaparte a correctement identifié ce taxon. Pucheran ne semble jamais avoir décrit

le genre Gorsachius, qu’il faut donc attribuer à BONAPARTE.

Ardea cinerea monicae Jouanin & Roux, 1963.

Jouanin & Roux, Oiseaux et Rev.fr. Orn. 33 : 103-106.

Premier spécimen, holotype par désignation originale.

C. G. 1961 - 4.

Ardea cinerea monicae Jouanin & Roux, 1963.

Etiquette rouge :

Ardea cinerea monicae / Jouanin et Roux 1963 / C. G. 1961 n° 4 / Ardea cinerea monicae

Jouanin et Roux 1963 / O.R.F.O. 33 : 104.

Étiquette blanche :

Ardea cinerea monicae / Jouanin et Roux / Loc. : île Arel, banc d’Arguin, 20 juin 1960 /

adulte nicheur / Type / Coll. : Mr. J. Dragesco / Loc. : Mauritanie / C. G. 1961 n° 4.

Remarque : ce spécimen désigné comme «type» dans la description est en fait l’holotype

par désignation originale (Code international de Nomenclature Zoologique, 3 e éd., 1985, article

73 A 1).

Second spécimen, paratype par désignation originale implicite

C. G. : 1961-770.

Ardea cinerea monicae Jouanin & roux, 1963.

Étiquette rouge :

Ardea cinerea monicae / Jouanin et Roux / C. G. 1961 n°770 / paratype.

Étiquette blanche :

Col. Mr. Dragesco / Loc. Mauritanie / C. G. 1961 n° 770 / Ardea cinerea monicae /

Jouanin et Roux / Loc. île Kiaone-ouest / Banc d’Arguin / juin 1960 / adulte nicheur.

Remarque : ce second spécimen, mentionné mais non désigné dans la publication originale,

a lui aussi servi à établir la description de l’espèce et constitue donc un paratype (Code inter¬

national de Nomenclature Zoologique, 3 e éd., 1985, article 72 B V).

Ardea purpurea bournei de Naurois, 1966.

de Naurois, L’Oiseau et Rev. fr. Orn. 36 : 89.

Source : MNHN, Paris

— 599 —

Premier spécimen, holotype par désignation originale.

C. G. 1966-891.

Ardea purpurea bournei de Naurois, 1966

Étiquette rouge :

Ardea purpurea bournei / de Naurois 1966 / C. G. 1966 n° 891 / Ardea purpurea / boumei

de Naurois 1966 / ORFO 36: 89.

Étiquette blanche :

Coll. Abbé R. de Naurois / Loc. Ile du Cap Vert / C.G. 1966-891/ Ardea purpurea boumei

Naurois / Loc. Ilha de Santiago / Type / 1964.

Second spécimen, paratype par désignation originale.

C. G. 1966-892.

Ardea purpurea bournei de Naurois, 1966.

Étiquette rouge :

Ardea purpurea / boumei Naurois / C. G. 1966 n° 892 / paratype / Ardea purpurea bour¬

nei / O.R.F.O. 1966 / 36 : 89.

Étiquette blanche :

Coll. Abbé R. de Naurois. / Loc. Archipel du Cap Vert / C. G. 1966 n° 892 / Ardea

purpurea bournei / Naurois / Loc. Ile de Santiago / 31.X. 1965. Femelle.

Troisième spécimen, paratype par désignation originale.

C. G. 1966-895.

Ardea purpurea bournei de Naurois, 1966.

Étiquette rouge :

Ardea purpurea boumei Naurois / C. G. 1966 n° 895 / Paratype / Ardea purpurea boumei

Naurois / O.R.F.O. 1966 36 : 89.

Étiquette blanche :

Coll. Abbé R. de Naurois / Loc. Iles du Cap Vert / C. G. 1966 n° 895 / Ardea purpurea

bournei / Naurois / Loc. Ilha de Santiago / 1964 Sexe?

Quatrième spécimen, paratype par désignation originale

C. G. 1966-897.

Ardea purpurea bournei de Naurois, 1966.

Étiquette rouge :

Ardea purpurea boumei Naurois / C. G. 1966 n° 897 / Paratype / Ardea purpurea boumei

Naurois / O.R.F.O. / 1966 36 : 89.

Étiquette blanche :

Col. Abbé R. de Naurois / Loc. Iles du Cap Vert : C. G. 1966 n° 89 / Ardea purpurea

bournei : Naurois / Loc. île Santiago / 1964 sexe?

Remarque : la sous-espèce Ardea purpurea bournei de Naurois est incontestablement valable.

Immédiatement identifiable, elle diffère largement autant de la sous-espèce nominale que ne le

fait la sous-espèce asiatique Ardea purpurea malinensis Meyen.

Source : MNHN, Paris

— 600 —

Ardeola podiceps Bonaparte, 1855.

Bonaparte, Consp. Gen. Avium, tome 2 : 134.

Type par monotypie.

C. G. 1995 - 250.

Ixobrychus minutus podiceps (Bonaparte, 1855).

Inscriptions sous le socle :

13982 Ardeola podiceps. Bp.type / de Madagascar / 1894 / par M. Bernier / [écriture

pâle] N. C. N°214 Blongios nain. / Ixobrychus m. podiceps / Type (Bp.) / M. Bemier.

Madagascar [écriture plus récente, très nette].

Remarque : dans sa diagnose BONAPARTE précise que l’exemplaire décrit provient de Mada¬

gascar et qu’il a été rapporté par Monsieur Bemier.

Cancroma coromanda Boddaert, 1783.

Boddaert, Table Planches Enlum. : 5.

Type par monotypie.

C. G. 1995 - 233.

Bubulcus ibis coromanda (Boddaert, 1783).

Inscriptions sous le socle :

heron a [sic] cou jaune rapporté par Sonnerat en 1770 / Duf. Enl 910 / aigrette dorée

[écriture très ancienne] monté en 1807 [autre écriture très ancienne] / ardea comata B,

Gm. / ardea russata, Tem. / de la côte de / Coromandel / par Sonnerat. enl. 910 / selon

toute probabilité / le type de la planche / de Buffon / N° 48 / 6-5 [toutes ces mentions,

bien qu’anciennes sont encore bien lisibles] Bubulcus coromandus (Bodd) [cette dernière

mention est récente et écrite au crayon].

Inscriptions sur l’étiquette :

Bubulcus coromandus / (Bodd.) / Type / Sonnerat Coromandel

Remarques : Boddaert a décrit Cancroma coromanda d’après BUFFON 1780 Hist. Nat.

Oiseaux 7 : 393 et Le Crabier de la côte de Coromandel de Daubenton 1765-1781, planche

enl. n°910. D’après l’inscription sous le socle ce spécimen est celui représenté par Daubenton.

Considéré comme «type de Buffon» il est aussi celui de l’espèce, d’après le Code international

de Nomenclature de Zoologie, 3 e éd., 1985, article 72 c 5 : «dans le cas d’un taxon nominal

du groupe-espèce fondé sur l’illustration ou une description, ou une référence bibliographique

à une illustration ou à une description, le spécimen illustré ou décrit et non l’illustration ou la

description» constitue un Type porte-nom acceptable.

Ardea agami Gmelin, 1789.

Gmelin, Systema Naturae, Aves : 629.

Type par monotypie.

C. G. n° 1995 - 246.

Agamia agami (Gmelin, 1789).

Source : MNHN, Paris

— 601 —

Inscriptions sous le socle :

Remis les plateaux à réparé [sic] en 1849 / Le plateau portait pour renseignement /

Cayenne, par Martin / l’Etiquette, l’amerique / [mot illisible]. Je crois que c’est le type

[mot illisible] / la pl. enl. 859 de Buffon. / 1349-D / [écriture ancienne]. Ardea agami /

(Gm) / Type de / Buffon / N. C. n° 213 / [écriture récente].

Inscriptions sur l’étiquette :

Ardea agami / (Gm). Type de Buffon / Cayenne?

Remarques : GMELIN a décrit Ardea agami d’après le texte de Buffon : Hist. Nat. Oiseaux

7 : 382 et la planche enlum. pl. 859 «Le Héron Agami de Cayenne» dessinée par Daubenton

1765-1781. D’après l’inscription sous le socle ce spécimen est bien celui représenté par DAUBEN-

ton. Considéré comme «type de Buffon» il est aussi le type de l’espèce d'après le Code

international de Nomenclature Zoologique, 3 e éd. (1985) article 72 c 5 (voir ci-dessus).

TYPES DE TAXONS TOMBÉS EN SYNONYMIE

Ardea atricollis Wagler, 1827.

Wagler, Systema Avium, Ardea : 174 sp. 4.

Type par monotypie.

C. G. 1995 - 238.

Ardea melanocephala Vigors & Children, 1826.

Inscriptions sous le socle :

Ardea atricollis Wagl. / du Sénégal / Ech. à m. Fl. Prévost l[illisible]27 Type de / Wagler/

13864 / 388 Manuel / Degland [écriture ancienne; les trois dernières mentions sont ins¬

crites au crayon] Ardea melanocephala / (Vig. et Chil.) / Sénégal / A. atricollis Wagler

Type [écriture plus récente] Ardea melanocephala V.& Ch. / (= Ardea atricollis type de

Wagler) / Fl. Prévost. Sénégal. N.C.18 [écriture plus récente]

Inscriptions sur l’étiquette :

Ardea melanocephala V. & Ch. / (= Ardea atricollis Type de Wagler) / Fl. Prévost Sénégal.

Remarques : cet exemplaire représente bien le type de Wagler car cet auteur précise, à

la suite de sa description, que le spécimen décrit est originaire de Sénégambie et qu’il provient

de la collection de Fl. Prévost. La description d 'Ardea melanocephala par VIGORS & Children

a la priorité sur celle d’A. atricollis Wagler, 1827, car elle est parue en 1826, un an avant celle

de Wagler. Ardea atricollis Wagler en est donc un synonyme plus récent.

Ardea novae-hollandiae Vieillot, 1817.

Vieillot, Nouveau Dictionaire d'Histoire Naturelle, Tome XIV : 436.

Premier spécimen, lectotype, présente désignation.

C. G. 1995 - 251.

Nycticorax caledonicus hilli Mathews, 1912.

Source : MNHN, Paris

— 602 —

Inscriptions sous le socle :

Bihoreau Canel / Différent de novae-hollandiae [illisible] c’est l’ardea caledonica [illi¬

sible] / Ardea [illisible] [écriture épaisse ancienne, en partie effacée].

Deux étiquettes sont collées sous le socle. Etiquette n° 1 : asie aust. Ctte [pour corvette]

le nat.ste [pour naturaliste] / expédit. du Cap. Baudin / an 11 / N.h. par Lesueur/ Etiquette

n° 2 : Bihoreau de la nelle hollande / Ardea novae hollandiae / [les deux derniers mots

sont rayés avec au-dessus une mention illisible] Vieill. nouv. diet. / d’hist. nat. 1817. il

a échappé une erreur / dans le texte. On a dit gris pour la couleur / du dos et des côtés

du col. Type de Vieillot [encre de Chine très pâlie], Nycticorax caledonicus Australasiana

[illisible] [écriture un peu effacée mais pas très ancienne] / 14015 / N. C. n° 127 /

Bihoreau cannelle / Nycticorax caledonicus (Gm.) / A. australasiana (type de Vieillot.)

/ M. Péron. Australie [écriture plus récente].

Inscriptions sur l’étiquette :

Bihoreau cannelle / Nycticorax caledonicus / (G m) / A. australasiana (Type de Vieillot)

/ M. Péron. Australie.

Second spécimen, paralectotype, présente désignation.

C. G. 1995 - 253.

Nycticorax caledonicus hilli Mathews, 1912.

Inscriptions sous le socle :

héron de la n elle holland [sic] / ardea novae-hollandiae Vieill. [le nom latin précédent

est barré et remplacé par australasia] / [illisible] glissé une erreur dans [illisible] [écriture

ancienne, texte en grande partie effacé] Nycticorax caledonicus (Gould) / A. australiasana

(V.) Type / N. C. 126 / 14014 / Bihoreau cannelle / Nycticorax calédonicus / (A. aus¬

tralasiana (Type Vieil.) / M. Péron - Australie.

Inscriptions sur l’étiquette :

Bihoreau cannelle / Nycticorax caledonicus / (Gm.) / - A. australasiana (Type de Vieil¬

lot) / M. Péro / Australie.

Troisième spécimen, paralectotype, présente désignation.

C. G. 1995 - 252.

Nycticorax caledonicus hilli Mathews, 1912.

Inscriptions sous le socle :

Nycticorax caledonicus, Gm. / A. australiasana (V.) Type / Péron et Lesueur / exp. du

C.ne Baudin / d’Australie / 14016 / [écriture ancienne] Bihoreau cannelle / Nycticorax

caledonicus (Gm) (A. Australasiana - type de Vieil.) / Mr. Péron - Australie / N. C. -

n° 128.

Inscription sur l’étiquette :

Bihoreau cannelle / Nycticorax caledonicus / (Gm) / A. australasiana. (Type de Vieillot) /

M. Péron - Australie

Quatrième spécimen, paralectotype, présente désignation.

C. G. 1995 - 254.

Nycticorax caledonicus hilli Mathews, 1912.

Source : MNHN, Paris

— 603 —

Inscriptions sous le socle :

Nycticorax caledonicus (Gm.) / A. australasiana (V.) Type / jeune de l’année / exp. Bau¬

din / Péron et Lesueur / [écriture ancienne] 14017 [écrit au crayon] N. C. n° 129 / Bi¬

horeau cannelle / Nycticorax caledonicus / A. australasiana. type de Vieil. / Mr. Péron

- Australie [écriture plus récente]

Inscription sur l’étiquette :

Bihoreau cannelle / Nycticorax caledonicus / (Gm.) / - A. australasiana. (Type de Vieil¬

lot) / M. Péron / - Australie

Remarque : les spécimens ci-dessous sont les mêmes que les précédents car Vieillot les

a redécrits sous un autre nom.

Ardea australasie Vieillot, 1823.

Vieillot, Tableaux encyclopédiques méthodologiques. Ornithologie volume 3: 1130.

Premier spécimen, lectotype, présente désignation.

C. G. 1995 - 251.

Nycticorax caledonicus hilli Mathews, 1912.

Second spécimen, paralectotype, présente désignation.

C. G. 1995 - 253.

Nycticorax caledonicus hilli Mathews, 1912.

Troisième spécimen, paralectotype, présente désignation.

C. G. 1995 - 252.

Nycticorax caledonicus hilli Mathews, 1912.

Quatrième spécimen, paralectotype, présente désignation.

C. G. 1995 - 254.

Nycticorax caledonicus hilli Mathews, 1912

Remarques concernant Ardea novae-hollandiae Vieillot et Ardea australasie Vieillot :

Vieillot a décrit successivement les quatre spécimens ci-dessus en 1817 et 1823 sous les noms

de novae-hollandiae et d’ australasie respectivement. Il s’agit de deux adultes et de deux jeunes :

les numéros 126 et 127 concernent les adultes, les numéros 128 et 129 les jeunes.

Ainsi que l’a déjà fait remarquer PuCHERAN (1851), la description que donne Vieillot

pour Ardea australasie (1823) ne diffère de celle qu’il a déjà donnée pour Ardea novae-hollandiae

que par l’addition d’une diagnose latine qui n’apporte rien de plus. Il résulte de cette double

description que le nom subspécifique de novae-hollandiae a priorité sur celui A'australasie. Tou¬

tefois, un Ardea novae-hollandiae ayant déjà été employé par Latham (1790), A. novae-hollan¬

diae Vieillot en est un homonyme objectif et la stricte application des règles du Code aboutit

au contraire à conserver australasie. Dans les deux cas Vieillot s’est trompé en décrivant comme

grises les parties qui sont normalement brun-roux chez Nycticorax caledonicus.

En 1912 Mathews considère que l’Australie est habitée par deux sous-espèces de

N. caledonicus, à savoir N. caledonicus australasiae. Vieillot (il change la dénomination aus¬

tralasie de Vieillot en australasiae, ce qui ne serait plus possible de nos jours) et N. caledonicus

Source : MNHN, Paris

— 604 —

hilli Mathews (1912). La description de cette dernière sous-espèce se résume à : «Diffère de N.

c. australasiae par sa coloration plus pâle». La description de VIEILLOT étant erronée, celle de

Mathews l’est aussi. Lors de la rédaction du troisième volume de son «Birds of Australia»

(1913-1914), Mathews, après avoir pris connaissance de la description de Vieillot, exprime

l’avis que ce dernier ne pouvait pas avoir eu en main d’authentiques N. caledonicus et a supprimé

la sous-espèce australasiae pour ne conserver pour l’Australie (y compris la Tasmanie) que la

sous-espèce hilli. La stricte application du Code conduirait à désigner la sous-espèce australienne

sous le nom de australasie puisque malgré l’erreur de Vieillot nous avons bien des spécimens

qui sont d'authentiques N. caledonicus rapportés par l’expédition Baudin en 1800-1804. Cepen¬

dant, ce choix serait sans doute malheureux pour la stabilité de la nomenclature car ce nom n’a

plus été employé depuis la parution de Birds of Australia et est donc devenu un nomen oblitum.

Ardea bilineata Cuvier in Lesson, 1831.

Lesson, Traité d'Ornithologie : 574.

Premier spécimen, lectotype, présente désignation.

C. G. 1995 - 242.

Ixobrychus flavicollis flavicollis (Latham, 1790).

Inscriptions sous le socle :

Ardea bilineata cv / Par M Labillardière / de Java [écriture ancienne] / 13958 / [très

pâle] N. C. N° 217 / [très net] Ardea flavicollis / (Cuv. Type) / [écriture plus récente].

Inscriptions sur l’étiquette :

Ardea flavicolli / A. bilineata (Cuv.) T. / Labillardière Java.

Second spécimen, paralectotype, présente désignation.

C. G. : 1995 - 243.

Ixobrychus flavicollis flavicollis (Latham, 1790).

Inscriptions sous le socle :

heron de Java / Par M. Leschenault / 1807 [écriture ancienne]/ Ardea bilineata (cuv.)

[autre écriture ancienne] type [écriture plus récente] / Ardea flavicollis (Lath) / 13959 /

[même écriture que celle de type] / N.C.- N°218

Inscriptions sur l’étiquette :

Ardea flavicollis / A. bilineata (Cuv.) T. / Labillardière Java.

Remarques : la description originale d 'Ardea bilineata se trouve dans le traité de LESSON

(Cuvier ne l’ayant jamais publiée lui-même). Cependant Lesson attribue bien cette espèce à

CUVIER puisqu’il la nomme Ardea bilineata, Cuv. Dans ce cas la dénomination d 'Ardea bilineata

Cuv. est gardée (C.I.N.Z. 3 e édition, 1985 : 90, article 50.a.). La diagnose très succinte de LESSON

se termine par «De Java. (Labillardière).» Ce sont donc bien les spécimens ci-dessus qu’il a

examinés. Le nom spécifique de bilineata Cuv. n’est cependant qu'un synonyme plus récent de

celui de flavicollis Latham (1790).

Source : MNHN, Paris

— 605 —

Ardea australis Cuvier in Lesson, 1831.

Lesson, Traité d' Ornithologie : 572.

Type par monotypie.

C. G. 1995 - 244.

Ixohrychus flavicollis flavicollis (Latham, 1790).

Inscriptions sous le socle :

Ex(tion) Baudin / Ardea flavicollis (Lath). / A. australis (Cuv.) type / [cette inscription

presque effacée] / Ext Baudin [écrit au crayon] / 13960 [presque effacé] N. C. N°219

[très net].

Pas d’étiquette.

Remarques : comme pour le taxon précédent la description originale se trouve dans le traité

de Lesson (1831) et CUVIER ne l’a jamais publiée lui-même. Cependant Lesson attribue ce

taxon à CUVIER puisqu’il le nomme Ardea australis Cuv. On garde donc la dénomination d'Ardea

australis Cuv. (C.I.N.Z. 3 e édition, 1985 : 90, article 50.a.). Cette diagnose très succinte ne com¬

prend que quelques lignes et Lesson n’a pas remarqué que Ardea bilineata et Ardea australis

représentaient en fait la même espèce. Dans sa description LESSON précise que le spécimen qu’il

a examiné est celui du «voyage de Péron» (Péron était un des membres de l’expédition Baudin).

Le nom spécifique d ’australis Cuv. n’est qu’un synonyme plus récent de celui de flavicollis

Latham (1790).

Ardea nivea. Cuvier in Lesson, 1831.

Lesson, Traité d'Ornithologie : 575.

Type par monotypie.

C. G. 1995 - 235.

Egretta intermedia intermedia (Wagler, 1829).

Inscriptions sous le socle :

reçu en 1818 chenault pondicherie [texte ancien repassé à l’encre de chine par la suite]

N° 398 / Ardea alba [presque effacé] / Ardea nivea, / Cuv. (type) / Jeune avec [effacé] /

mais [effacé] / A. egrettoides / les tarses plus longs qu’[illisible], / le bec blanc tandisqu’il

est noir / dans le garzetta; et taille moindre / que ardea timoriensis cv qu’il égale pour /

la blancheur. Val. [écriture ancienne] / N. C. N° 53 [autre écriture] / Ardea intermedia /

A. Nivea (Cuv.) Type / 13886 [écriture récente] 13886 [noté deux fois au crayon]

Inscription sur l’étiquette :

Moyenne Aigrette / Egretta intermedia / Type (Wagl.) / M. Leschenault Pondichéry.

Remarques : comme dans les cas précédents la description originale d ’Ardea nivea se trouve

dans le traité de Lesson (1831) qui attribue cette espèce à Cuvier sans doute à cause de la

mention sous le socle. Dans sa diagnose LESSON note que le spécimen décrit se trouve à la

Galerie de Paris, qu’il provient de Pondichéry et qu’il a été raporté par Leschenault, ce qui

correspond bien au spécimen C. G. 1995 - 235. La diagnose de LESSON, très succinte, est pos¬

térieure à celle de Wagler publiée en 1827 et Ardea nivea Cuv. en est donc un synonyme plus

récent.

Source : MNHN, Paris

— 606 —

Ardea pealii Bonaparte, 1826.

Bonaparte, Annals of the Lyceum of Nat. Hist, of New-York II. : 154.

Type par monotypie.

C. G. 1995 - 234.

Egretta rufescens rufescens (Gmelin, 1789).

Inscriptions sous le socle :

Individu / figuré American / Ornithology / 4 [illisible] 26 fig 1 [texte presque effacé] /

Ardea peali, des Etats unis. / Floride / Bonap. / (type de l’espèce) de la collection /

donnée par le Prince / Charles Bonaparte / en 1854. / (Cat n° 1199). / Ardea rufa /

(Bodd.) / N. C. n° 212 / Dicromanassa rufescens (= Ardea Pealii / type de Bonap.) /

coll. Ch. Bonap. Floride.

Inscriptions sur l’étiquette :

Aigrette roussatre / Dichromanassa rufescens / (Gm) (Ardea Pealii Type de Bp.) / Coll.

Ch. Bonaparte Floride.

Remarques : ce spécimen est le type par monotypie d 'Ardea pealii Bn. Il a fait l’objet

d’une représentation graphique dans Bonaparte 1833 ( American Ornithology vol. IV : 96,

pl. 26 fig.l.) Le nom spécifique de pealii Bn. n’est qu’un synonyme plus récent de rufescens

Gmelin (1789), décrit bien antérieurement.

Ardea ardesiacea Lesson, 1831.

Lesson 1831, Traité d’Ornithologie : 575

T^pe par monotypie.

C. G. 1995 - 247.

Egretta caerulea (Linné, 1758).

Inscriptions sous le socle :

Ardea Coerulea, L/ de Cayenne par M. Poiteau avril 1822 / (n° 135) / Ardea / ardesiacea, /

Lesson / (type) / 13911 [écriture ancienne presque effacée] N. C. 211 / Petit heron cendré

d’Amérique / Florida caerulea (L) / (= Ardea ardesiacea, type de Lesson / M. Poiteau

- Cayenne) [écriture récente]

Inscriptions sur l’étiquette :

Petit Héron cendré d’Amérique / Florida coerulea (L.) / (= Ardea ardesiacea type de

Lesson) / M. Poiteau Cayenne.

Remarques : d’après les mentions portées sous le socle, ce spécimen est bien celui décrit

par LESSON qui, de plus, précise dans son ouvrage qu’il provient de Cayenne. La description

à'Ardea caerulea Linné (1758) étant bien antérieure à celle de LESSON, Ardea ardesiaca Lesson

n’en est qu’un synonyme plus récent. C’est de plus un homonyme objectif plus récent de Ardea

ardesiaca Wagler, 1827.

Ardea callocephala Wagler, 1827.

Wagler, Systema Avium, Ardea : 189 sp. 34.

Type par monotypie.

C. G. 1995 - 255.

Nycticorax violaceus (Linné, 1758).

Source : MNHN, Paris

— 607 —

Inscriptions sous le socle :

le bihoreau de Cayenne, buff. Enl 899 / [écriture très ancienne : les 4 mentions qui suivent

sont de la même écriture mais ont été rayées par la suite] ardea cayennensis Gm. / ardea

violacea. Will vol 8 pl 65 fig 1 / le bihoreau à [mot illisible] / ardea sexsetacea Vieill.

Diet, [l’écriture suivante est ancienne et la même que celle qui a rayé les mentions pré¬

cédentes] / Ardea violacea L. / A. violacea et cayennensis, Gm. / A. violacea et A. sex¬

setacea Vieill. / A. callocephala Wagl. (T), [l’écriture suivante est plus récente] Martin

/ De Cayenne [rayé] / de Cayenne / par / Martin / Nycticorax violaceus (L.) / a. callo¬

cephala (Wagl.) [autre écriture récente] / Bihoreau à tête jaune / Nyctanassa violacea

(L.) / A. violacea (type de Wagl.) / M. Martin. Cayenne [chiffre presque effacé, très

ancien et repassé au crayon] 14034.

Inscriptions sur l’étiquette :

Bihoreau à tête jaune / Nyctanassa violacea (L.) / A. violacea (type de Wagl.) / M.

Martin. Cayenne.

Remarques: dans son travail, Wagler (1827) mentionne que le spécimen qu’il décrit se

trouve au Muséum de Paris. Après sa diagnose il donne une liste de synonymes à savoir : Ardea

violacea Wil., Bihoreau de Cayenne Buff., Ardea cayennensis Gm. et Ardea sexsetacea Vieillot.

Dans ces conditions on ne sait pour quelles raisons WAGLER a donné à ce taxon le nouveau

nom d 'Ardea callocephala, qui n’est qu’un synonyme plus récent d 'Ardea violacea Linné.

SPÉCIMENS CONSIDÉRÉS À TORT COMME DES TYPES

Ardea mokoho Vieillot, 1817.

Vieillot, Nouveau Dictionaire d’Histoire Naturelle Tome XIV Ardea : 440.

C. G. 1995 - 237.

Botaurus lentiginosus (Rackett, 1813).

Inscriptions sous le socle :

New-york / par M. milberts 1818 / Ardea minor orn. am. / 8 pl.65 fig 3 / Ardea stellaris

Gm Edw 136 / Ardea lentiginosa mont, et Shaw [écriture ancienne sur une étiquette

collée sous le socle] / placé à tort parmi les oiseaux d’Europe. / [2 mots illisibles] 13995 /

Butor à bandes noires [écriture ancienne] 397 cat. Degland [écrit au crayon] Ardea /

mokoho / Vieill. / Wagl. [écriture ancienne] Botaurus lentiginosus / (mont.) / A. mokoko

[sic] / type / N. C. n° 207 [écriture plus récente] Butor de la baie d’Hudson / Botaurus

lentiginosus (mount.) / A. mokoko [sic] Type de Vieillot / M. Milbert Etat Unis.

Inscriptions sur l’étiquette :

Butor de la Baie d’Hudson / Botaurus lentiginosus / (Mont.) (A. mokoko [sic] Type de

Vieill.) / M. Milbert Etats-Unis.

Source : MNHN, Paris

— 608 —

Remarques : le Nouveau Dictionnaire d'Histoire Naturelle ayant paru en 1817 et ce spécimen

ayant été rapporté du Nouveau Monde par M. Milbert en 1818, il ne s’agit probablement pas

de l’exemplaire décrit par Vieillot. Plus tard, il a été examiné par Wagler qui s’est contenté

de reprendre le nom que lui avait donné VIEILLOT. Il ne s’agit donc pas d'un type au regard

du Code. Dans sa description originale, Vieillot, qui savait qu’il s’agissait d’un oiseau amé¬

ricain, l’a nommé mokoho, d’après une appellation indigène, nom repris par WAGLER (1827).

L'orthographe «mokoko», utilisée par nombre d’auteurs du xix e siècle, est erronée.

C’est à Rackett (1813) qu’il faut attribuer le nom de lentiginosus et non à MONTAGU

(1813). En effet, la seconde édition du Catalogue de Pultney, dans laquelle Rackett décrit cet

oiseau qu’il croyait européen, est parue un peu avant le travail de MONTAGU. Le nom spécifique

de lentiginosus Rackett, 1813 a donc la priorité. La mention «placé à tort parmi les oiseaux

d’Europe» se rapporte sans doute à la place que ce spécimen occupait dans la Galerie.

Ardea melanoptera Cuvier in Pucheran, 1851.

Pucheran, Rev. Mag. de Zool. : 375.

Premier spécimen, nomen nudum

C. G. 1995 - 248

Ixobrychus sinensis (Gmelin, 1789).

Inscriptions sous le socle :

Ardea melanoptera cv / 13966 [ancien, presque effacé] / Ardea sinensis / Gm / Ard.

lepida / Horsf. Wagl. / des Mariannes Exp. uon / freycinet 1820 [écriture ancienne] / Ardetta

sinensis (Gm) / A. melanop. / (Cuv.) / type [écriture plus récente] / Blongios de Chine

/ Ixobrychus sinensis / Type A. melanoptera / (Gm) / L’Uranie. I. Mariannes / N. C.

N° 215.

Inscriptions sur l’étiquette :

Blongios de Chine / Ixobrychus sinensis / (Gm) / Type / L’Uranie I. Mariannes / [ajouté

à la main sur l’étiquette imprimée après la mention : type] A. melanoptera (Cuv.).

Second spécimen, nomen nudum.

C. G. 1995 - 249.

Ixobrychus sinensis (Gmelin, 1789).

Inscriptions sous le socle :

Ardetta sinensis (Gm) / A. melanoptera (Cuv. type) / Exp. freycinet 1820 / Iles Mariannes /

13970 [écriture pâle] / N. C. N°216 / Blongios de Chine / Ixobrychus sinensis (Gm) /

Type A. melanoptera (Cuv.) / L’Uranie. I. Mariannes [écriture plus récente].

Inscriptions sur l’étiquette :

Blongios de Chine / Ixobrychus sinensis / Type (Gm.) / A. melanoptera (Cuv.) [La mention

A. melanoptera (Cuv.) est ajoutée à la main sur l’étiquette imprimée] / L’Uranie I.

Mariannes.

Source : MNHN, Paris

— 609 —

Remarque : Cuvier n’a pas publié de description de ce taxon. PUCHERAN se contente de

le citer et l’attribue à Cuvier (sans doute à cause des mentions sous le socle), également sans

en donner de description. Il s’agit donc d’un nomen nudum et ces deux spécimens ne sont pas

des types au sens du Code.

Gorsachius melanolophus Raffles, 1822.

C. G. 1995 - 240

Gorsachius melanocaphalus Raffles, 1822.

Inscriptions sous le socle :

Héron de l’Inde / par Brassard / fev. 1829 / 14007 / Nycticorax Goisagi Tem [presque

effacé volontairement] Individu décrit / Consp. / Avium vol ii p. 138 [écriture ancienne)

Type / Botaurus melanolophus / (Raff.) / Gorsakius typus Pucheran / N. C. 143 / [écriture

plus récente] Petit Butor de Malaisie / Gorsachius melanolophus / (Raff.) / M. Brassard

- Inde, [écriture récente]

Inscription sur l’étiquette :

Petit Butor de Malaisie / Gorsachius melanolophus / (Raffl.) / M. Brassard Inde.

Remarques : ce spécimen est un mâle en plumage nuptial. Nous avons déjà indiqué plus

haut (voir goisagi) que Bonaparte (1855) s’était mépris à son sujet en le décrivant comme

étant l’adulte de Gorsachius goisagi (Tem.). La mention «type» apposée sous le socle signifie

seulement que ce spécimen était considéré, avec celui de goisagi, comme type du genre

Gorsachius Bonaparte.

Remerciements

Nous remercions ici le Prof. W. J. Bock (New York) ainsi que le Prof. J. Vuillemier (New York)

pour leurs critiques constructives du manuscrit de cet article.

RÉFÉRENCES

(autres que celles des descriptions)

BONAPARTE C. L., 1855. — Conspectus Generum Avium. Éd. Brill, Leyde.

Hancock J. & Elliott H., 1978. — The herons of the world. London Editions. Londres.

Mathews G. M., 1912. — A reference list to the birds of Australia. Novit. Zool. 18: 171-455.

— 1910-1927. — The Birds of Australia, 12 vol. Witherby, Londres.

MAYR E. & COTTRELLE G. W. (eds)., 1979. — Peter’s Check-list of Birds of the World, vol. I, 2nd éd. Cambridge,

Muséum of Comparative Zoology.

STRESEMANN E., 1951. — Die Entwicklung der Ornithologie. F. W. Peters, Berlin W 15.

VOISIN C., 1993. — Liste des spécimens types d'ibis (Threskiornithinés) de la collection du Muséum national

d'Histoire naturelle de Paris. Oiseau et R.f.O. 63 : 45-53.

VOISIN J.-F.. 1992. — Liste provisoire des types d’oiseaux des collections du Muséum national d’Histoire naturelle

de Paris. 1. Pélécaniformes. Oiseau et R.f.O. 62 : 162-172.

— 1995. — Liste des types d’oiseaux des collections du Muséum d’Histoire naturelle de Paris. 3. Podici-

pédiformes. Bull. Mus. nail Hist, not., série 4. A 17 (3-4) : 345-351.

Source : MNHN, Paris

LOUIS-JEAN

avenue d'Embnin, 05003 GAP cedex

Tél. : 04.92.53.17.00

Dépôt légal : 936 — Décembre 1996

Imprimé en France

Date de distribution le 20 décembre 1996

erratum :

Le fascicule n° 1-2 1996 a été distribué le 17 juillet 1996

Source : MNHN, Paris

Mémoires du Muséum national d’Histoire naturelle

Tome 140 : Dubois, A., 1988. — The genus in zoology: a contribution to the theory of evolutionary

systematics. 124 p. (ISBN : 2-85653-151-2) 150 FF.

Tome 146 : Erard, Ch., 1990. — Écologie et comportement des gobe-mouches (Aves : Muscicapinae,

Platysteirinae, Monarchinae) du Nord-Est du Gabon. Volume 2 : Organisation sociale et écologie des

Muscicapinae. 234 p. (ISBN : 2-85653-178-4) 320 FF.

Tome 148 : Matile, L., 1990. — Recherches sur la systématique et l’évolution des Keroplatidae

(Diptera, Mycetophiloidea). 704 p. Relié/Hardback. (ISBN : 2-85653-173-3) 650 FF.

Tome 157 : Chazeau, J. & S. Tillier (eds), 1993. — Zoologia Neocaledonica, volume 3. 218 p.

(ISBN : 2-85653-205-5) 280 FF.

Tome 159 : Robbe, P., 1994. — Les Inuit d’Ammassalik, Chasseurs de l’Arctique. 398 p. (ISBN :

2-85653-207-1)360 FF.

Tome 163 : Crosnier, A. (ed.), 1995. — Résultats des campagnes MUSORSTOM, volume 13.

518 p. (ISBN : 2-85653-224-1) 550 FF.

Tome 166 : Jamieson, B. G. M., J. Ausio & J.-L. Justine (eds), 1995. — Advances in Spermatozoa!

Phylogeny and Taxonomy, 550 p. Relié/Hardback. (ISBN : 2-85653-225-X) 440,80 FF.

Tome 167 : Bouchet, Ph. (ed.), 1995. — Résultats des campagnes MUSORSTOM, volume 14, 654 p.

Relié/Hardback. (ISBN : 2-85653-217-9) 600 FF.

Tome 169 : Geoffroy, J.-J., Mauries, J.-P. & Nguyen-Duy-Jacquemin, M. (eds), 1996. — Acta

Myriapodologica. 682 p. Relié/Hardback. (ISBN : 2-85653-502-X) 538,69 FF.

A partir de 1993, les Mémoires du Muséum national d'Histoire naturelle ne paraissent plus en quatre

séries séparées (A, B, C, D), mais en une seule série. Le nom de la revue devient Mémoires du Muséum

national d'Histoire naturelle (ISSN 1243-4442). La numérotation commence au tome 155.

Vente en France

Sales Office (France excluded)

MUSÉUM NATIONAL D’HISTOIRE NATURELLE

UNIVERSAL BOOK SERVICES

ÉDITIONS SCIENTIFIQUES DU MUSÉUM. PARIS

Dr. VP. Backhuys

Diffusion Delphine Henry

57, rue Cuvier, 75005 Paris, France

P.O. Box 321 2300 AH Leiden

Tél. : [33] 01.40.79.37.00

Fax : [33[ 01.40.79.38.40

Internet http://www.mnhn.fr

The Netherlands

Tel. : [31] (71) 5 17 02 08

Fax : [31] (71)5 17 18 56

Internet backhuys@euronet.nl

: TVA 2.10%

Source : MNHN, Paris