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AIVVTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

November 1999 Volume 17, N° 1-2

Source : MNHN, Paris:

sp ee

ISSCA

International Society for the Study

and Conservation of Amphibians

(International Society of Batrachology)

SEAT

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Source : MNHN, Paris

AIMTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

November 1999 Volume 17, N° 1-2

Alytes, 1999, 17 (1-2): 1-2. 1

Editorial

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

This issue of A/ytes contains three papers that deal with the taxonomy and nomenclature

of ranoid frogs, a fascinating research subject that still promises many novelties in the coming

decades. Two of these papers include discussions of nomenclatural matters, that were written

when the third edition of the /nternational Code of Zoological Nomenclature (ANONYMOUS,

1985) was in force. Actually, this edition is still is in force at the time of this publication, but for

a few months only: in September 1999, the fourth edition of the Code (ANONYMOUS, 1999) was

published, whose provisions are to replace those of the third edition as of 1 January 2000.

Some of the nomenclatural discussions of these two papers will then become irrelevant. Thus,

according to the new Article 16, after that date, any new species name will be nomenclaturally

available only if “explicitly indicated as intentionally new” and accompanied in the original

publication by the explicit fixation of a holotype or syntypes, and by reference to the

collection of deposition of this or these specimen(s). These new rules are highly welcome, as

they will limit seriously the risk of publication of “phantom names” as defined below in this

issue by VENCES et al. (1999). Let us note however that they do not apply to names published

before 1 January 2000.

The new edition contains other important changes regarding some articles of the Code,

that will no doubt be discussed by zoologists worldwide in the coming years. One of them

deserves particular attention: the new Article 23.9 introduces the concepts of “reversal of

precedence”, of “prevailing usage” and of “nomen protectum”. This article states that,

whenever two names are considered synonyms or homonyms, “prevailing usage” must be

maintained when “the senior synonym or homonym has not been used as a valid name after

1899”, and “the junior synonym or homonym has been used for a particular taxon, as its

presumed valid name, in at least 25 works, published by at least 10 authors in the immediately

preceding 50 years and encompassing a span of not less than 10 years”. Had the word

available been used instead of valid in this article, the latter would have raised no major

problem and would have been welcomed virtually by all zoologists. But the use of the term

valid opens the door for possible abuses and for future problems and discussions. It is no

mystery for any experienced taxonomist that many names treated once as “invalid” because

they were then considered junior subjective synonyms were later “resurrected” when this

subjective synonymy was demonstrated to be wrong. The new article “moderates” the

application of the Principle of Priority in this case, to replace it (without naming it) by a

so-called “principle of usage”. However, there is no doubt that, unlike priority, usage can be

“deliberately rigged or manipulated” (Dumois, 1995b, 1997). A tendency already exists for

some zoologists, when describing a new taxon, to coin a new name for it even if names are

Bibliothèque Centrale Muséum

QUIL

00088

MNHN, Paris

2 ALYTES 17 (1-2)

already available and may even be widely known but sometimes “hidden in synonymies” (for

recent examples in amphibians, see e.g.: DUBOIs, 1995a, 1998, 1999a-b ; DuBois & OHLER,

1995, 1998, 1999). The new Article 23.9 may be received by some authors as an encourage-

ment for hasty and careless work, or even for deliberate omission of names published prior to

1900, in order to create “their” names. Ten years is a very short period in taxonomy indeed,

and such poor nomenclatural actions may be quickly “validated” through this new article.

This would not only, as some previous recent actions and statements, be an insult to the

zoologists of the past (some of whom were at least as careful and competent as recent ones),

and “to the thousands of authors who have followed the principle of priority (...) and thanks

to whom stability has been reached for the very large majority of names” (Dugois, 1995c).

More importantiy, perhaps, this would be liable to strengthen the current progressive growth

of a lax attitude of neglect or ignorance of the basic nomenclatural rules in zoology (see e.g.

Dusois & OHLER, 1997, 1999), that might rather quickly lead to a chaotic situation in this

field, as discussed below in this issue (DUBOIS, 1999b).

In view of these potential problems, the greatest attention will be paid, in the coming

years, during the review process, to the nomenclatural aspect of papers describing new

amphibian taxa submitted for publication to 4/ytes: such papers will be checked to provide all

necessary information showing that a careful analysis of the situation has been carried out

and that no earlier name is available for any such taxon. Hopefully, all other zoological

journals worldwide will follow the same editorial policy.

LITERATURE CITED

ANONYMOUS [International Commission on Zoological Nomenclature], 1985. - Code international de nomencla-

ture zoologique. Third edition. London, International Trust for zoological Nomenclature: i-xiv + 1-328.

_— 1999. — International code of zoological nomenclature. Fourth edition. London, International Trust for

zoological Nomenclature: i-xxix + 1-306.

Dumoïs, A. 1995a.- The valid scientific names of the Italian treefrog, with comments on the status of some early

scientific names of Amphibia Anura, and some articles of the Code concerning secondary homonyms.

Dumerilia, 2: 55-71.

= 1995b. — Discussion draft of the fourth edition of the International Code of Zoological Nomenclature:

comments. (3). Bull. 2001. Nom. 52 (4): 299.

ca 1995c. - Comments on the proposed conservation of Hemidactyliini Hallowell, 1856 (Amphibia, Caudata).

Bull. zool. Nom. 52 (4): 337-338.

--— 1997. - Proposals concerning the conditions needed for a name being eligible for conservation. In: DuBoIs

& OnLer (1997): 317-320.

-— 1998. — List of European species of amphibians and reptiles: will we soon be reaching “stability”?

Amphibia-Reptilia, 19 (1): 1-28.

1999a. - South Asian Amphibia: a new frontier for taxonomists. Invited editorial / Book review. J South

Asian nat. Hist., 4 (1): 1-11.

= 1999h. — Miscellanea nomenclatorica batrachologica. 19. Notes on the nomenclature of Ranidae and

related groups. Alyres, 17 (1-2): 81-100.

Dusors, A. & OuLer, À. 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,

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

LE 1997. - Early scientific names of Amphibia Anura. L. Introduction. Bull. Mus. natn. Hist. nat., (4), 18 (3-4):

297-320.

= 1998. — À new species of Leprobrachium (Vibrissaphora) from northern Vietnam, with a review of the

taxonomy of the genus Leprobrachium (Pelobatidae, Megophryinae). Dunerilia, 4 (1): 1-32.

1999. — Asian and Oriental toads of the Bufo melanostictus, Bufo scaber and Bufo stejnegeri groups

(Amphibia, Anura): a list of available and valid names and redescription of some name-bearing 1ypes. J

South Asian nat. Hist., 4 (2): 133-180.

VENCES, M. GLAW, F. & BÜHME, W., 1999. - À review of the genus Mantella (Anura, Ranidae, Mantellinac)

taxonomy, distribution and conservation of Malagasy poison frogs. Alyres, 17

ISSCA 1999

Source : MNHN, Paris

Alytes, 1999, 17 (1-2): 3-72. 3

A review of the genus Mantella

(Anura, Ranidae, Mantellinae):

taxonomv, distribution and conservation

of Malagasy poison frogs

Miguel VENCES *, Frank GLAW ** & Wolfgang BÔHME *

* Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany

** Zoologische Staatssammlung, Münchhausenstr. 21, 81247 München, Germany

In this paper, 17 species of the genus Mantella are recognized and the

genus is partitioned into six species groups which can be distinguished by

combination of bioacoustic, morphological, osteological and coloration

characters. The following species and species groups are recognized:

Mantella betsileo group (Mantella betsileo, Mantella viridis, Mantella

expectata, Mantella sp. 1, and one new species described herein); Man-

tella laevigata group (Mantella laevigata); Mantella cowani group (Man-

tella cowani, Mantella baroni, Mantella aff. baroni, Mantella harald-

meieri, Mantella nigricans); Mantella bernhardi group (Mantella

bernhardi);, Mantella madagascariensis group (Mantella madagasca-

riensis, Mantella pulchra); Mantella aurantiaca group (Mantella auran-

tiaca, Mantella crocea, Mantella milotympanum). This partition is of

rather high resolution, and some of the groups may also be regarded as

superspecies or species complexes.

A detailed type re-examination showed that M. madagascariensis and

M. baroni represent two different species which are very similar in dorsal

coloration, but M. madagascariensis can be distinguished by some charac-

ters of ventral coloration (horseshoe marking on the throat, reddish color

ventrally on femur) and morphology (large inner metatarsal tubercle) from

M. baroni.

Specimens from Marojezy preserved in the Paris museum are cata-

logued as M. cowani nigricans and must therefore be considered as

syntypes of this taxon. The syntype series is heterogeneous, also containing

specimens of M. laevigata. The name nigricans is stabilized by designation

of a lectotype corresponding to a M. cowani group form from Marojezy. The

name Mantella cowani nigricans Guibé, 1978 is revalidated and raised to

species rank as Mantella nigricans.

A big problem in Mantella systematics is that, in recent years, hob-

byists increasingly tend to publish “phantom” scientific names without type

n several cases lead to involuntary but nomenclaturally

available new nominal taxa. Two phantom names which must be considered

as nomenclaturally available are Mantella aurantiaca milotympanum

Staniszewski, 1996 and Mantella aurantiaca rubra Staniszewski, 1996.

We consider the name rubra as synonym of M. aurantiaca, but prelimina-

rily attribute specific status to M. milotympanum.

Lectotypes (in addition to M. nigricans) are designated for M. cowani,

M. aurantiaca, M. betsileo, Mantella attemsi (synonym of M. betsileo),

M. aurantiaca rubra (synonym of M. aurantiaca) and M. milotympanum

Source : MNHN, Paris

4 ALYTES 17 (1-2)

(from published figure). Clarifications on types and type series are provided

for several species.

We provide a key to the species of the genus Mantella, and describe and

discuss their color variability. In several species, a large intraspecific color

variability was recorded (M. aff. baroni, M. nigricans, M. crocea). À

detailed review of all published Mantella localities and the corresponding

voucher specimens results in updated distribution maps. Sympatric and

syntopic occurrence was reliably only found in species from different

species groups, the species within each group being allopatrically distri-

buted. Future studies on contact and hybrid zones may demonstrate that

some of the species recognized herein should possibly better be regarded as

subspecies; however, for practical reasons, we here regard all taxa as

species.

In an attempt to provide an estimate of the conservation status of each

Mantella species, we combined data on distribution (maximum locality

distance, number of known localities), habitat (primary forest restriction),

trade intensity and attractiveness to the pet trade. We group the species in

various classes, according to their potential vulnerability, and outline priori-

ties of research needed to get a more reliable data basis for such estimates.

INTRODUCTION

The ranoid subfamily Mantellinae currently contains two genera, both endemic to

Madagascar (GLAW & VENCES, 1994): the type genus Mantella, and the large and heterog-

eneous Mantidactylus with currently 63 species. Mantella are small, largely diurnal and often

colorful frogs, which were named Malagasy (or Madagascan) poison frogs due to the presence

of alkaloid toxins in their skin (e.g. DaLy et al., 1996).

Accounts on the genus were published by GUIBÉ (1964, 1978) and BUssE (1981).

BLOMMERS-SCHLÔSSER & BLANC (1991) largely relied on BUSsE's revision which they comple-

mented by detailed distribution maps. The description of four new species by PINTAK &

BÔHME (1988, 1990), Bi & BÔHME (1992) and VENCES et al. (1994) demonstrated, however,

that those accounts were far from being complete. While GuiBé (1978) listed only four species

and one subspecies, GLAW & VENCES (1994) already accepted 13 different species.

One of the major problems in Mantella systematics has been weak morphological

differentiation. Since early workers generally studied only preserved material, they had to rely

largely on color pattern for species diagnoses. GurBé (1964, 1978) and especially BUSSE (1981)

considered single species (named M. cowani or M. madagascariensis, respectively) as highly

variable in coloration, but they never proved this variability in specimens from a single locality

(Day et al., 1996). Without definite knowledge of intra- and interpopulational color

variability, the attribution of type specimens of early name: M. cowani,

M. baroni,

M. madagascariens

1. pulchra) \argely depended on the subjective impression of the corr

author, causing large confusion in the usage of these names in scientific and non-

literature.

In the following we report the main results on taxonomy, distribution and color variabi-

lity of Mantella which were gathered in the framework of a comprehensive study of the genus.

Contributions to the morphometry, osteology, tadpole morphology, reproduction, karyology,

as well as bioacoustic and allozyme differentiation within Mantella are being published

elsewhere. The aim of the present paper is mainly to clarify the taxonomy and nomenclature

Source : MNHN, Paris

VENCES, GLAW & BÔHME $:

of Mantella species as well as their distribution, in order to give a more stable basis for future

investigations of these frogs. We divide the genus into phenetic species groups, and use our

new scheme of Mantella systematics to discuss biogeographical subjects and to summarize

conservation needs.

MATERIAL AND METHODS

SPECIMENS EXAMINED

The present review is mainly based on preserved material of the following collections:

The Natural History Museum, London (BMNHD); Field Museum of Natural History, Chi-

cago (FMNH); Museum of Comparative Zoology, Cambridge (MCZ); Muséum National

d'Histoire Naturelle, Paris (MNHN);: Museo Regionale di Scienze Naturali, Torino

(MRSN/MZUT); Naturhistorisches Museum Basel (NMB): Naturhistorisches Museum

Wien (NMW); Transvaal Museum, Pretoria (TM); Zoëlogisch Museum Amsterdam (ZMA);

Museum für Naturkunde der Humboldt-Universität zu Berlin (ZMB); Zoologisches For-

schungsinstitut und Museum Alexander Koenig, Bonn (ZFMK). Specimens were examined

in detail and their color patterns and morphology recorded. Locality and collector are

generally litterally given according to the corresponding catalogue. Abbreviations used are:

CS, cleared and stained specimens; TE, tissue extracted for electrophoresis, specimens only

partly preserved (generally liver extracted and two limbs amputated); NIL, specimens not

individually labeled. The term “ex” is used in the sense of “formerly” to characterize old

collection numbers.

LOCALITIES AND DISTRIBUTION MAPS

The examined material is the basis of the locality maps and the statements on color

ies are numbered, the numbers corresponding to those in the respective

distribution maps. A star behind the locality number marks the localities which were con-

firmed by FG (and partly by MV) in the field. The type locality, in the nomenclatural account

on each taxon, is given in quotation marks litterally as in the original description: additional

discussions, when necessary, are provided in the Comments sections.

D

RIPTION OF COLOR PATTERNS

Variation of color patterns is described in a standardized way and generally refers to live

coloration of adult specimens. Terms which we use to refer to certain color elements are

defined as follows: (1) dorsolateral color border: a sharp longitudinal border between the

color of the flanks (darker) and the dorsum (lighter); (2) frenal stripe: a light longitudinal

stripe along the upper lip: (3) rostral stripe: a light (yellowish, greenish or brownish) stripe

running from anterior head tip and nostril above the eye to a point behind the eye:

Source : MNHN, Paris

6 ALYTES 17 (1-2)

(4) diamond marking: a central (dark) marking on the back of more or less distinct double-

rhomboid shape; (5) flank blotches: light markings of varying extension which are located

posterodorsally around the forelimb insertion and anterodorsally around the hindlimb

insertion; they mostly can be seen as an extension of the dorsal humerus/femur color on the

flanks; (6) flashmark: a sharply delimited, bright orange or red marking on the posterodorsal

femur, knee hollow and ventral tibia which in some species can cover the ventral tibia nearly

entirely; (7) horseshoe marking: a light (generally whitish blue) continuous marking on the

throat, running more or less broadly along the lower lip and thus horseshoe-shaped. The

terms femur, tibia, and tarsus, as used in the sections on coloration, do not refer to the skeletal

elements but to the external coloration of the corresponding hindlimb sections.

MORPHOMETRY AND MORPHOLOGY

Measurements taken were: SVL: snout-vent length; HW: maximum head width; HL:

head length, measured from snout tip to forelimb insertion (not to maxilla articulation); Eye:

horizontal eye diameter; Tym: horizontal tympanum diameter; Eye-Ns: distance between eye

and nostril; Ns-St: distance between nostril and snout tip; ForL: forelimb length; HaL: hand

length; HiL: hindlimb length; FoTL: foot length including tarsus; FoL: foot length; ToL1:

length of first toe; FW3: width of third finger just before terminal finger disk; DW3: width of

terminal disk of third finger; IMTL, IMTH, IMTW: length, height and width of inner

metatarsal tubercle. All measurements were made by the senior author with a precision

calliper to the nearest 0.1 mm, except FW3, DW3, IMTL, IMTH, IMTW which were

measured using a binocular with measuring device to the nearest 0.01 mm or, when no

binocular was available, with a calliper to the nearest 0.1 mm. Original measurements in the

present paper are only given for type specimens, but the size ranges and morphometric ratios

in the species accounts refer to a total of about 400 measured specimens.

In the text, besides SVL, we use the abbreviations IMT for inner metatarsal tubercle, and

TTA for tibiotarsal articulation. The size (SVL) is given as range of adult specimens, followed

where possible by the range recorded in the males and females which could be reliably sexed.

Since in many cases specimens could not be sexed with a sufficient reliability, known adult size

range may be wider than that recorded in males and females separately.

DESCRIPTION OF CALLS

Detailed call descriptions will be published elsewhere; here we tentatively distinguish

four different general call types: (1) double click calls are series of notes which each are

composed of two emphasized and very short “metallic” clicks; (2) single click calls are series

of notes which each are composed of one emphasized and very short “metallic” click; (3) trill

calls are (irregularly repeated) notes composed of up to 10 short clicks: (4) chirp calls consist

of (irregularly or regularly repeated) notes with a less “metallic” appearance than in click calls

as used above (a note is often composed of 2-3 emphasized pulses).

Source : MNHN, Paris

VENCES, GLAW & BÔHME 7

SYNONYMIES

For each Mantella species, we present a synonymy and chresonymy (for the definition of

the term chresonymy, see SmirH & SmirH, 1973), following the scheme used by DaviD &

VOoGEL (1996). The overwhelming number of publications in which at least one species of

Mantella is mentioned makes it impossible to provide a complete chresonymy. Instead, we

present a selection of references (partial chresonymy) which either (1) discuss intrageneric

taxonomy and systematics, (2) provide original data for at least one species, (3) include

pictures of at least one species, or (4) were published before GUIBÉ’S (1964) revision of the

genus (the latter, however, must be seen with reservation since it is often difficult to understand

to which species the author actually referred). Page numbers are only given if necessary to

locate a deviating name usage or a figure. Only publications which contain either original data

or figures are listed in the chresonymies of the species. Exceptions are the works of GUIBÉ

(1964, 1978), BussE (1981), BLOMMERS-SCHLÔSSER & BLANC (1991) and GLAW & VENCES

(1992a, 1994), which are here considered as monographie accounts on the genus. AII names

used in these works are listed in the corresponding synonymies. Generally, taxa which were

defined in a publication in a way that, according to present definition, they were in fact

composed of several species, are listed as “partim-chresonyms” (“part.”?) in the chresonymies

of each of these species (in the case of monographs) or of the species which were shown or

explicitly meant (in the case of other papers). Nomenclatural validity of names is discussed

according to the /nternational Code of Zoological Nomenclature (ANONYMOUS, 1985; cited

below as “the Code”).

RESULTS

THE GENUS MANTELLA

Definition of the genus

Following the data of GUIBÉ (1978), BLOMMERS-SCHLÔSSER & BLANC (1991), BLOMMERS-

SCHLÔSSER (1993), DaLy et al. (1996), GLAW et al. (1998b), PINTAK et al. (1998), VENCES &

KNIEL (1998) and VENCES et al. (1998, 19994), the genus Mantella can be defined by the

combination of the following characters:

(1) Eight presacral vertebrae; (2) vertebral centrae procoelous: (3) sacral diapophyses not

enlarged; (4) atlantal cotyles widely separated; (5) three free distal tarsals; (6) six free distal

carpals; (7) terminal phalanges slightly Y-shaped; (8) hyoid with anterolateral and posterolat-

eral processes: (9) anterior processes of hyalia forming complete arch in some specimens of

most or all species; (10) palatines present; (11) maxillary and premaxillary teeth absent: (12)

13) dentigerous process of vomer (and thus vomerine teeth) absent; (14)

squamosal with reduced zygomatic process: (15) frontoparietals anteriorly convex-shaped

and separated along their whole length: (16) process of pars fascialis of maxilla reduced; (17)

shoulder girdle firmisternal; (18) ossified sternum and omosternum present; (19) sternum

Source : MNHN, Paris

8 ALYTES 17 (1-2)

shorter than omosternum; (20) omosternum forked at its base; (21) complete ventral circum-

marginal groove on terminal finger and toe expansions; (22) SVL of adults 18-31 mm; (23)

tibiotarsal articulation reaching between forelimb insertion and nostril; (24) tympanum

visible externally, mean tympanum/eye ratio 1/2 to 2/3; (25) lateral metatarsalia connected;

(26) no webbing between fingers nor toes; (27) inner and outer metatarsal tubercle present;

(28) no dorsal “scutes” on finger and toe tips: (29) karyotype 2 n = 26, with 5 pairs of large and

8 pairs of small chromosomes which are meta- or submetacentric; (30) tongue very slightly

notched; (31) microphagous and myrmecophagous feeding; (32) skin alkaloids present; (33)

long prey-capture jumps absent; (34) colorful pattern at least ventrally (black/blue, yellow or

orange), often also dorsally; (35) activity largely diurnal; (36) calls consisting of short clicks,

chirps or trills; (37) no strong mating amplexus; (38) eggs generally laid outside of the water;

(39) eggs unpigmented; (40) tadpoles with horny beak and keratodont formula 1:2+2/3 to

1:5+5/3 (formula according to DuBois, 1995); (41) no tadpole transport; (42) no external gills

in early larval stages; (43) egg clutches consisting of less than 200 eggs: (44) no externally

prominent femoral glands as in many Mantidactylus, but granular thigh patches present (see

also DaLY et al., 1996), most distinct in males (exact structure of these patches will be subject

to a forthcoming publication).

Character states 9, 11, 14, 30, 31, 32, 33 and 34 are, as far as known, not found in

Mantidactylus, the second genus of the Mantellinae. They all can be considered as derived in

Mantella (based on outgroup comparison with other ranid frogs, e.g. the Malagasy rhaco-

phorines of the genus Boophis). However, states of characters 11, 13, 14, 16, 30, 32, 33, 34 and

35 are all part of a character complex related to microphagy (character 31), which reduces

their value as independent characters for the assessment of phylogenetic relationships (see

VENCES et al., 1998). In fact, most of them are also found in the Dendrobatidae which, too, are

microphagous but clearly differ from Mantella in other characters (different states in charac-

ters 5, 6, 7, 18, 20, 28, 29, 30, 40, 41 and 42; for references, see VEN et al., 1998).

Apomorphic states supporting the status of Mantella as a monophyletic (holophyletic) group

within the Mantellinae are thus the microphagy character complex (see above) and the hyoid

structure (character 9).

Etymology of the generic name

The genus Mantella was erected by BOULENGER (1882) to accomodate the species bersileo,

madagascariensis and ebenau; in an addendum he described the new species cowanii. The type

species is Mantella betsileo, as designated by LieM (1970). No etymology was given in the

original description of the genus. The generic name is most probably a diminutive of mantis

(Classical Greek mantis, prophet) which was used with the meaning “treefrog” in the sense of

a weather prophet by HesycHios. This meaning of mantis is included in several Greek-

German dictionaries (e.g. PAPE, 1888) but was not found in Greek-French or Greek-English

dictionaries (see GLAW & VENCES, 1994: 400). The term mantis was often used for generic

anuran names; BOULENGER himself erected in 1895 the genus Mantidactylus for several

Madagascan frogs which today are included together with Mantella in the Mantellinae.

A second etymology for Mantella, however, cannot be totally excluded. One of the early

subjects of BOULENGER'S studies were dinosaur fossils found in Belgium, which belonged to

the genus /guanodon. The first Jguanodon fossils had been found by an English doctor,

Source : MNHN, Paris

VENCES, GLAW & BÔHME 9

G. MANTELL, and his wife, and were subsequently described as /guanodon mantelli (see

BULTYNCK, 1987). Still less probable is a derivation from the Italian word mantella (cloak)

which is sometimes used to describe animal (mammal) color patterns.

DEFINITION OF SPECIES GROUPS

Although several authors have stressed similarities between selected Mantella species

and erected species groups within the genus (GLAW & VENCES, 1994; ZIMMERMANN, 1996a;

STANISZEWSKI, 1996), no comprehensive attempt has so far been published to partition the

whole genus into such groups, and to explicitly list the characters distinguishing them. We

here divide the genus into six phenetic species groups, a subdivision of rather high resolution;

in fact, some groups could also be characterized as superspecies or species complexes. The

differential characters between species groups are summarized in tab. 1.

Mantella betsileo group (contains: Mantella betsileo, M. sp. 1, M. viridis, M. expectata,

and one new species described herein). - This group is characterized by the combination of

several characters which, however, are each also present in at least one other species group:

double click call (also in M. laevigata), horseshoe marking (also in several other groups),

frenal stripe (also in M. crocea and some M. madagascariensis), hindlimbs ventrally black

without orange and red (also in M. Jaevigata and M. nigricans).

Mantella laevigata group (contains: Mantella laevigata).— The classification of Mantella

laevigata in a separate species group is clearly justified by its unique habits (partly arboreal,

tree hole breeding, single eggs) and its distinctly enlarged finger tips. It is the only species with

a double click call which lacks a horseshoe marking.

Mantella cowani group (contains: Mantella baroni, M. aff. baroni, M. cowani, M. nigri-

cans, M. haraldmeieri). — A group characterized by light (mostly yellow or red) flank blotches

of variable extension (also found in the M. madagascariensis group and in M. bernhardi) and

single click calls (exclusive to this group).

Mantella bernhardi group (contains: Mantella bernhardi). — Classification of M. bern-

hardi in a separate species group is mainly based on its relevant allozyme differentiation

(VENCES et al., 1999b) and its trill calls.

Mantella madagascariensis group (contains: Mantella madagascariensis, M. pulchra). -

The species included in this group are mainly characterized by a very large IMT (see diagnosis

of M. pulchra in Guisé, 1964, 1978). Light flank blotches of varying extension, horseshoe

markings and flashmarks are present. Calls, as far as known, are chirp calls.

Mantella aurantiaca group (contains: Mantella aurantiaca, M. crocea, M. milotympa-

num). — Species of this group are characterized by a rather stout body shape, distinct

flashmarks and a chirp call. In contrast to species of the M. madagascariensis group, there are

no flank blotches and the IMT is smaller. Two species (M. aurantiaca, M. milotympanum) are

characterized by a largely uniform yellow to red dorsal and ventral coloration. M. crocea is

included in this group since specimens with color pattern intermediate between M. crocea and

M. milotympanum are known (GLAW & VEN 1998), and juvenile coloration of M. crocea

and M. aurantiaca is Very similar (personal observation). The close relationships between the

Source : MNHN, Paris

Table 1. - Differential characters between Mantella species groups. Not all characters have been ascertained in all species of the groups. See Definition

of species groups section for more information. Stemum shape is given according to VENCES et al. (1999a). IMT, inner metatarsal ubercle.

Mantella Mantella Mantella Mantella

Mantella Mantella

Character betsileo laevigata cowani bernhardi madagascariensis aurantiaca

group group group group group group

es mostly double click, | double click, single click, till, chip, chip,

series series series mostly no series series mostly no series!

Sternum forked forked unforked unforked forked forked

Horseshoe marking present absent absent present present present/absent

Frenal stripe present absent absent absent absent (present) present/absent

Flank blotches absent absent large/small small large absent

Orange/red ventral absent absent present (absent)* present present present

color on hindlimbs

Habitat terrestrial partly arboreal terrestrial terrestrial terrestrial terrestrial

Eggs laid as clumps single eggs clumps clumps clumps clumps

Egg feeding of absent present absent absent absent absent

tadpoles

IMT small small small small large small

! Only ascertained in M. aurantiaca, M. crocea calis are also chirp calls (personal observation), but notes may be arranged more often in series than in M. aurantiaca.

? A frenal stripe may be present in certain specimens of the “variable” morph of M. madagascariensis.

? Orange/red ventral color on hindlimbs is present in all species of the M. cowani group except M. nigricans.

OI

(T-D LI SALATV

Source : MNHN, Paris

VENCES, GLAW & BÜHME 11

species of the M. aurantiaca group were supported by chromosome morphology (PINTAK et

al., 1998) and by studies on allozyme variation (VENCES et al., 1999b). ZIMMERMANN (1996)

also mentioned a M. aurantiaca group which included M. aurantiaca and M. crocea.

SPECIES ACCOUNTS

In the following, we list Mantella species separately for each species group; within the

groups, species are arranged alphabetically. Photographs of living specimens of all species are

shown in fig. 1-3, dorsal and ventral views of holotypes and lectotypes (all photographed

1992-1999) in fig. 4-5, and variation of ventral pattern in fig. 6-8. Distribution maps are shown

in fig. 9.

Mantella betsileo group

Mantella betsileo (Grandidier, 1872)

Dendrobates betsileo Grandidier, 1872. - Name-bearing type: lectotype, by present designation, MNHN

1895.278, sex unknown due to bad state of preservation, SVL 19.0 mm. - Type locality: “Pays des Betsileos”

according to original description and MNHN catalogue. — Other types: paralectotype, following present

lectotype designation, MNHN 1895.279. — Etymology: named after the type locality, the region Betsileo.

Mantella betsileo: BOULENGER, 1882, 1888; VAILLANT, 1885; WERNER, 1901; MOCQUARD, 1909;

METHUEN & HewrT, 1913; MizLor & Guimé, 1951; Guiné, 1964, 1978; LiEM, 1970; BACHMANN &

BLOMMERS-SCHLOSSER, 1975; BLOMMERS-SCHLÔSSER, 1978, 19794; MEIER, 1980 (part.: p. 353, third

figure from above), 1986 (Abb. 3): Busse, 1981 (part.: see M. haraldmeierÿ}, PINTAK, 1990;

BLOMMERS-ScHLÔSSER & BLANC, 1991 (part.), 1993 (plate 19.101); ANDREONE, 1992 (pl. III fig.

3-4); GLaw & VENCES, 19924 (part.; see localities), 1992 (fig. p. 29), 1994 (part; see localitics);

ZAMMERMANN & ZIMMERMANN, 1992 (fig. 5.17); GARRArFO et al., 1993; HERRMANN, 1993 (fig.

, 1993; BarTLerT, 1995 (fig. p. 26); HENKEL & SCHMIDT, 1995 (fig. p. 50); Caris

, 1995 (fig. p. 42); VENCES et al., 1996, 1998; DaLy et al., 1996: STANISZEWSKI, 1997a (fig,),

1997b (fig.); LARSEN, 1997; PiNTAK et al., 1998: VENCES & KNIEL, 1998.

Dendrobates ebenaui Boettger, 1880. - Namu-bearing type: lctotype, by designation of MERTENS (1967: 44),

SMF 7323 (ex 1141, la), adult female. - Type localit

tion. — Other types: possibly one paralectotype, FMNH 18236 or 183237 (see Marx, 1958, and comment

below). - Etymology: named after C. EBëNAU who provided the type material

Dendrobates Ebenaui: MocquaRD, 1909 (syn. betsileo): MERTENS, 1922 (syn. hetsileo); MERTENS, 1967

(syn. betsileo).

Dendrobates ebenaui: Guisé, 1964, 1978 (syn. betsileo), BUSSE, 1981 (syn. betsileo): BLOMMERS-

ScuLôsser & BLANC, 1991 (syn. betsileo); GLAW & VENCES, 1994 (syn. hetsileo: p. 411).

Mantella ebenaui: BOULENGER, 1882: WERNER, 1901; METHUEN & HEWITT, 1913 (syn. betsileo).

Mantella attemsi Werner, 1901. - Name-bearing type: lectotype, by present designation, NMW 20837, female,

SVL 25.6 mm. — Tÿpe locality: uncertain, but (in original description) was speculated 10 be probably

“Madagascar oder Nossi-Bé”. — Other types: paralectotype, following present lectotype designation, ZMB

16588. — Etymology: named after C. ATTEMS who provided the type specimens from Zanzibar.

Mantella Attemsi: MOCQUARD, 1909

Mantella attemsi: GUwÉ, 1964, 1978 (syn. betsile

& BLANC, 1991 (syn. bersileo); GLAW & VE

(syn. besileo).

USSE, 1981 (syn. betsileo); BLOMMERS-SCHLÔSSER

, 1994 (syn. hetsileo: p. 412) : HäuPL et al., 1994

— Day et al. (1996) were concerned about the fact that the type locality of M.

betsileo (see below) is outside the known range of the species. They questioned whether the

name is currently correctly applied. A re-examination of the types (see below) leads us to

Source : MNHN, Paris

12 ALYTES 17 (1-2)

conclude that they (1) cannot be conspecific with any species having red or orange ventral

color on the hindlimbs, (2) are morphologically different from M. laevigata and M. nigricans,

and (3) are smaller than M. viridis, M. sp. 1 and M. expectata. It seems therefore likely that the

name is currently (e.g. GLAW & VENCES, 1994) correctly applied.

Comments. — (1) The taxon betsileo was originally based on the syntypes MNHN 1895.278-

279. The lectotype MNHN 1895.278 (SVL 19.0 mm; sex unknown) is larger and in slightly

better state of preservation. The paralectotype MNHN 1895.279 is probably a subadult

specimen (SVL 15.7 mm). In both types, coloration has become a contrastless, nearly uniform

brown. The dorsolateral coloration border mentioned in the original description cannot be

unequivocally recognized. Since the hindlimbs were folded in both specimens, the pattern is

less faded on the posteriorly directed (ventral) part of the tibia which was not exposed to light.

Here, a distinct light crossband can be recognized, as is typical for species of the M. betsileo

group (and for M. laevigata and M. nigricans). Based on this character it can be excluded that

the types are conspecific with Mantella species having red or orange color ventrally on the

hindlimbs. — (2) The type locality of M. betsileo is a large region in central Madagascar. Up to

now, no Mantella betsileo specimens are known to have been collected in the eastern forests

south of Nosy Boraha. As discussed by DALY et al. (1996), the travel routes of GRANDIDIER

are rather well documented. Maybe, the types were not collected in the eastern Betsileo forests

but in western Betsileo, where the occurrence of M. betsileo seems more probable due to the

existence of several localities in western Madagascar. It also cannot be excluded that the type

locality is wrong. — (3) According to the original description (BOETTGER, 1880: 281), Dendro-

bates ebenauï was based on two syntypes, a male and a female. However, in his 1892 catalogue,

BOETTGER (1892: 21) mentioned “numerous specimens” (“Zahir. Ste”) of this species from

Nossibé, kept in the Frankfurt Museum under number SMF 1141,1a: presumably these

specimens included the two original syntypes and several other non-type specimens. TENS

(1922: 166) stated that the “Typus” of this species was bearing the number SMF 1141,1a, but

since this number was used by BOETTGER (1892) as a collective number for a series, this

mention cannot be considered as a lectotype designation under article 47 (b) of the Code.

Designation of an individual specimen, SMF 7323, as lectotype of this species, was made by

MERTENS (1967: 44). Thus, only one paralectotype exists. It might be one of the two FMNH

“paratypes” listed by Marx (1958), which were presumably part of the series mentioned by

BOETTGER (1892). Further clarification of the status of these two specimens is necessary. —

(4) The description of Mantella attemsi was based on two specimens (WERNER, 1901),

corresponding to the specimens NMW 20837 and ZMB 16588. Both are today in a rather bad

state of preservation. Color patterns are largely faded, only the dorsolateral color border

still recognizable. In the lectotype NMW 20837, a few ventral color patterns (light vermicu-

lated markings on the posterior venter) are still faintly recognizable. The paralectotype ZMB

16588 is most probably a male.

Material examined. - Difficulty of identification of specimens as M. betsileo is enhanced by the existence

of a very similar, undescribed species (M. sp. 1, see below). Since this spe neraily larger than

M. betsileo, size was one of the major diagnostic characters for preserved specimens with faded

coloration. However, we cannot totally exclude that some specimens may be wrongly identified and in fact

be subadults of M. sp. L. In parentheses, we give SVL for most specimens.

ly be assigned to M. betsileo: BMNH 84.11.3.4 (Nosy Be: purch.

28 (Nosy Be; “Senckenberg Museum”; NIL; SVL 23.7 mm,

The following specimens can

from “Linnaea”); BMNH 86.2

Source : MNHN, Paris

VENCES, GLAW & BÔHME 13

20.7 mm, 22.2 mm, 23.3 mm); BMNH 1909.10.19.21 (Nosy Be; P. KRerFT); BMNH 1926.10.27.4-7

(Antongil forest, Maroantsetra; purch. ROSENBERG; NIL: largest female 25.3 mm); BMNH

1952.1.1.55-56 (Rantabe, Antongil bay, Maroantsetra; coll. C. S. WeBB; female 55: 25.7 mm); ZFMK

17604-9 (Maroantsetra; leg. H. MEIER 1.1976); ZFMK 27680 (Maroantsetra; leg. H. MEIER X.1979);

ZFMK 28867-8 (Nossi-Be; through Linnaea 1886; originally Museum Gôttingen); ZFMK 46004 (Nosy

Boraha [Ile Ste. Marie]; leg. F. W. HENKEL I1.1987; CS); ZFMK 46781-3 (Nosy Be: Loucoubé; leg. R.

Serpp IV.1987); ZFMK 47007 (Nosy Bé: Loucoubé; leg. F. W. HENKEL & J. SAMEIT 11.1987); ZFMK

47218 (Sahafary; leg. F. GLawW XI.1987); ZFMK 47289 (Nosy Boraha [Ile Ste. Marie]; leg. H. MEIER

XL.1987); ZFMK 48257-8 (Nosy Bé: Loucoubé:; leg. W. ScHMipT 1987); ZFMK 52744 (Nosy Boraha [Ile

Ste. Marie]; leg. F. GLaw & M. VENCES IIL.1991); ZFMK 52745 (Nosy Bé: leg. F. GLAW & M. VENCES

1111991); ZFMK 51818 (Nosy Boraha [Ile Ste. Marie]; leg. F. W. HENKEL et al. 1988-89); ZFMK

53708-10 (Nosy Bé; leg. F. GLAW & J. MÜüLLER I1.1992; juveniles); ZFMK 59978-9 (Kirindy; leg. F.

GLAW 1.1995); ZFMK 62685-7 (locality unknown; CS); ZFMK 62689-96 (locality unknown; TE);

ZFMK 62688 (Nosy Be; leg. K. SCHMIDT; TE); MRSN A0064.1-4 (Maroantsetra; leg. F. ANDREONE

23.1V.1990); MRSN A0068.1-2 (Kirindy; leg. R. NINCHERI 22.XIT.1992).

The following specimens are assigned to M. betsileo based on size, general appearance and morpho-

metric characters such as relative hindlimb length: BMNH 94.2.27.21 (Madagascar, coll. LAST, purch.

GERARD; pattern totally faded); BMNH 1930.7.1.54-57 (valley 3/4 miles W of Ampoza, 15 miles E of

Ankazoabo, SW Madagascar: pres, Ware; rather small specimens; NIL). MNHN 1884.603-4 (Nossi Bé;

SVL 25 mm [603], 21 mm [604)); MNHN 1885.34-7 (locality unknown: SVL 21 mm [34], 18 mm [35], 20

mm (36), 18 mm [37]); MNHN 1885.48 (Nossi Bé); MNHN 1895.278-9 (lectotype and paralectotype:

Pays des Betsileos); MNHN 1896.435-6 (Madagascar, “acquis de l’Institut Linnaea”; color totally faded:

SVL 24 mm [435, female?], 23 mm [436]); MNHN 1900.15 (Pays Mahafaly, au Sud: with remark: mâle

”; SVL 20 mm); MNHN 1929.225 (source de Namoroko [Ambongo]: juv?, SVL 16 mm): MNHN

MNHN 1991.1795 [originally 129a] (forêt d'Manjaba; TTA reaches eye center [1795], SVL 25

mm [129], 24 mm [1795], few ventral markings, absent on breast [1795]; MNHN 1953.131, MNHN

1991.1796 [originally 131a] (Bas Manongarivo; SVL 17 mm [131], 16 mm [1796]; MNHN 1953.130,

MNHN 1953.13 (locality unknown; SVL 23 mm [130, 133]}; MNHN 1953.134 (Nossi-Bé — Lokobe,

Manjoky: juv., SVL 13 mm); MNHN 1962.895 (Namoroka, grotte de Bemahara; R. PAULIAN IX.52:

TTA reaches eye center, SVL 22 mm); MNHN 1962.896-7 (Anove, forêt littorale; A. DOMERGUE L.1961;

TTA reaches eye center (896, 897], SVL 21 mm [896], 22 mm [897]); MNHN 1976.200-2 (Nosy Komba:

SVL 21.1 mm [200], 19.7 mm [201], 19.3 mm [203]): TM 9858-67 (Eastern Region, Madagascar: coll.

HERSCHELL-CHAUVIN).

The following MNHN specimens with unknown locality are here assigned to M. betsileo only based

on their size which is given in parentheses: MNHN 1976.181-2 (SVL 22.3 mm [181], 22.0 mm [182]);

MNHN 1976.187 (SVL 22.2 mm); MNHN 1976.206 (SVL 19.5 mm); MNHN 1976.222-3 (SVL 20.9 mm

1222}, 17.6 mm [223]); MNHN 1976.225 (SVL 22.1 mm); MNHN 1976.227 (SVL 22.1 mm); MNHN

1976.230 (SVL 22.8 mm).

The status of two specimens is not suficiently clarified. They have enlarged disks on fingers and toes

and thus resemble A. laeviguru: MNHN 1953.192, MNHN 1991.1707 [originally 132a] (Tsaratanana;

Distribution. — Except the type locality “Pays des Betsileos”, all known localities are located in

lowlands (altitude between 0 and ca. 500 m), generally near the coast. Also, all east coast

localities are in an area north of Betsileo (see DaLY et al., 1996: 19). Confirmation of the type

locality would therefore be important.

The species is common along the east coast in the Maroantsetra region and on Nosy

Boraha, and in the Sambirano region; it also occurs along the west coast. Localities are as

follows: [1*] Nosy Boraha (<10 m altitude); [2*] Voloina (GLAW & VENCES, observations in

1991); [3] Maroantsetra; [4] Rantabe; [5*] Sahafary; [6] Anove; [7] Antanambaobe, and

Ambavala near Sandrakatsy in the Mananara reserve (DALY et al., 1996; 100-200 m altitude);

[8] Farakaraina near Maroantsetra (DALY et al., 1996; 30 m altitude); [9*] Nosy Be; [10*]

Source : MNHN, Paris

14 ALYTES 17 (1-2)

Nosy Komba; [11*] Benavony (F. GLAW & J. MÜLLER, observations in 1992); [12] Ankify

(village near ferry docks N Ambanja, personal communication of W. B. LOvE); [13] Manon-

garivo; [14] Tsaratanana (SCHIMMENTI, personal communication); [15*] Kirindy (Amborom-

potsy, see KUCHLING, 1993); [16] Namoroka; [17] Tsingy de Bemaraha (SCHIMMENTI, personal

communication); [18] Mahafaly (?); [19] Ampoza (not traced and therefore not included in the

distribution map).

The Mahafaly locality (MNHN 1900.15) in the very arid South-Western Region needs

confirmation but seems corroborated by the specimens from Ampoza in SW-Madagascar,

and is therefore accepted here in a preliminary way. Specimens from the Anosy mountains in

southern Madagascar identified as M. betsileo by BUSSE (1981) in fact belong to M. harald-

meieri (see below). MiLLOT & GUIBÉ (1951) mentioned the species from the “forêt de Bevia”

near Fort Dauphin, but we could not find voucher specimens for this locality in the Paris

Museum. Also HENKEL & SCHMIDT (1995) gave no vouchers for their locality Tolagnaro. We

consider these localities in need of confirmation, and do not accept them here.

We did not find voucher specimens for seven additional localities which were listed by

BLOMMERS-SCHLÔSSER & BLANC (1991). Of these, Nosy Mangabe probably was based on

BussE (1981: 29) who stated that specimens of M. laevigata collected by H. MEIER (on Nosy

Mangabe) lived parapatrically or sympatrically with M. betsileo. We did not find M. betsileo

on Nosy Mangabe, and the locality thus needs confirmation. The population from Montagne

des Français (near Antsiranana) is here referred to M. viridis (see below), and those from

Morondava, Androatsabo and Tongahybe to M. sp. 1 (see below). Baly probably refers to

Tongahybe, since in the Paris Museum catalogue the additional remark “Baly Ouest” is given

for the corresponding specimens (see section on M. sp. 1). We did not find vouchers

from Andranoboka in the Paris Museum and therefore consider this locality in need of

confirmation.

Diagnosis. — (1) Morphology: À small, relatively slender Mantella. SVL, males 18-21 mm,

females 19-26 mm. TTA mostly reaching eye center. Terminal disks of fingers and toes

expanded. Tympanum/eye ratio generally 1/2 to 3/5. IMT medium sized (ratio width/length

about 2/3). - (2) Dorsal color and pattern: Dorsal head surface and dorsum yellowish to

orange or light brown, mostly with a diamond marking and a sharp dorsolateral color border

to the largely black flanks. White frenal stripe present. Limbs brown to grey, with at least one

dark brown crossband on femur, tibia and tarsus. Iris with light pigment in its upper part. (3)

Ventral color and pattern: Black with blue markings of different size and extension often

showing vermiculated patterns and fusipg with each other. Breast region generally with light

markings, smaller than those on posterior venter. Distinct horseshoe marking present, of

larger extension in males than in females, and sometimes including a central stripe.

Mantella expectata Busse & Bôhme, 1992

Mantella expectata Busse & Bôhme, 1992. - Name-bearing type: holotype by original designation (BUSSE &

BÔHME, 1992: 58), ZFMK 53540, male, SVL 23.4 mm. — Type localil 20 km southeast of Toliara

(=Tuléar), W-Madagascar” according to original description. - Other types: paratypes ZFMK 53541-2,

ZFMK 59095-8 and four (lost) additional paratypes (see comment below). - Etymology: derived from Latin

expectare (to await); rediscovery of this species was awaited during several years after MEtER (1986) first

published a picture of it.

Source : MNHN, Paris

VENCES, GLAW & BÔHME 15

Mantella expectata: GLAW & VENCES, 1992, 1994; HERRMANN, 1993 (fig.); LE BERRE, 1993 (fig. p. 20):

HexkeL & Scmir, 1995 (fig. p. 52); BARTLETT, 1995 (fig. p. 26); VENCES et al., 1996; DaLY et al

1996; Sraniszewski, 1996 (plate p. 18), 1997a (fig. p. 16), 1997 (fig.), 1998a (fig): VENCES &

KNIEL, 1998.

Other chresonyms:

Pictured in Meter (1986: fig. 8) as “Eine noch unbekannte Mantella-Form”.

Comment. — In the original description (BussE & BÔHME, 1992), beside the catalogued

specimens ZFMK 53541-2, “eight living specimens which will be incorporated in the ZFMK

collection later” were also designated as paratypes. Of these captive stock, only four speci-

mens were eventually preserved and catalogued as ZFMK 59095-8; the remaining four

paratypes must be considered as lost.

Material examined. -ZFMK 53540 (SE Tulear: leg. G. GorrLeee II.1992; holotype); ZFMK 53541-2 (SE

Tulear: leg, G. GorrLere II.1992; paratypes): ZFMK 59095-8 (SE Tulear, through pet trade, paratypes);

ZFMK 62713-5, ZFMK 62789 (locality unknown; TE); ZFMK 62716-7 (locality unknown; CS).

Distribution. —- Known from: [1] the type locality, 20 km SE of Toliara; [2] the area around

Morondava, based on a picture made by a German development aid worker and published by

Mate (1986); [3] the Isalo massif (altitude ca. 800 m), based on a personal communication of

A. PEYRIERAS and on DALY et al. (1996). The locality Mandena in south-eastern Madagascar,

given by GLAW & VENCES (1994), was based on an erroneous information of G. HALLMANN

and was corrected by VENCES et al. (1996).

Diagnosis. —-(1) Morphology: À medium-sized, stout Mantella. SVL 20-26 mm. TTA someti-

mes reaching only the tympanum, but generally reaching the eye center. Terminal disks of

fingers and toes expanded. Mean tympanum/eye ratio nearly 3/5. IMT medium sized (ratio

width/length slightly more than 2/3). — (2) Dorsal color and pattern: Head and dorsum dirty

yellow to lemon yellow with a sharp dorsolateral color border to the black flanks. Limbs grey

to bright metallic blue. A thin bluish white frenal stripe present. Iris with light pigment in its

upper part. —(3) Ventral color and pattern: Black with irregularly shaped blue markings which

can fuse to form a blue-black marbling. Blue markings present on the breast. Throat largely

blue, as a very extended horseshoe marking.

Mantella manery n. sp.

Mantella sp: GLAW & VENCES, 1994; VENCES et al., 1996.

Mantella “maro) fconditional name] Ki, 1996, 19974, 1997b (eg. p. 16-18).

Mantella “marojezt” [conditional name]: STANISZEWSKI, 1997b (p. 61).

Mantella *’marojezy" [conditional name]: LARSEN, 1997.

Name-bearing type. — A single specimen of this species was preserved and deposited in the

herpetological collection of the Zoological Institute of the Antananarivo University, Mada-

gascar (leg. F. GLAW, N. RABIBISOA & O. RAMILISON, 27.11.1994) and is here designated as

holotype. The following description is based on color slides of this specimen:

Type locality. - Réserve Naturelle Intégrale Marojezy, near Camp 1, ca. 300 m altitude.

Other types. - None.

Source : MNHN, Paris

ALYTES 17 (1-2)

Fig. 1.- Photographs of Mantella species. (a) M. betsileo from Nosy Be (specimen not pres

v

(b) M. sp. 1 from Ankarana (specimen not preserved), 1995: (c) M. viridis, specimen without locality

data (not preserved), 1995: (d) A. expectata, specimen without locality data (not preserved), 1995:

(@) M. manery from Marojezy (holotype: deposited in the herpetological collection of the Antana-

narivo University), 1994: (1) M. laevigata from Marojezy (specimen not preserved), 1995; (g-h) M.

nigricans from Marojezy with and without greenish dorsal color patterns, 1995.

Source : MNHN, Paris

VENCES, GLAW & BÔHME 17

Identity. — Color patterns of this species differ from the remaining species of the M. betsileo

group. Its occurrence in rainforest also differs from most other species of the group (except

M. betsileo). It was considered a distinct species by GLAW & VENCES (1994). Unfortunately, no

specimen of this form was available for detailed examination, as the only preserved specimen

is stored in the herpetological collection of the University of Antananarivo. Until present, we

thought that the description of this species should wait until new material was collected, and

new data on its variation, calls, ecology and osteology became available. However, several

hobbyist authors (e.g., STANISZEWSKI, 1996; LARSEN, 1997) have made reference to this form

as “Mantella marojezyi”, “ Mantella marojezi” or “ Mantella marojezy”, providing diagnoses

which were entirely based on our previously published data. All authors who previously used

these names wrote them in quotation marks; these usages thus must be seen as conditional

names which are not nomenclaturally available according to article 15 of the Code. However,

it can be expected that sooner or later the name will be used without quotation marks in

any of the increasingly published hobbyist accounts on Mantella (see Discussion below),

accompanied by a diagnosis, and will thus become valid. We therefore prefer to name the form

by a formal preliminary description, designating the specimen stored in the Antananarivo

collection as holotype. Our preliminary account should be complemented as soon as the

holotype (currently not available to us) is examined in detail, and new field observations are

made.

Etymology. — Derived from the Malagasy verb manery (to force, forced), here used as an

invariable substantive standing in apposition to the generic name. We were forced to describe

and name this form in a preliminary way to avoid it being named without proper diagnosis in

a hobbyist publication.

Distribution. — Only known from the type locality: [1*] Marojezy massif, near Camp 1.

Diagnosis. - The new species is a member of the M. betsileo group based on the presence of a

horseshoe marking, frenal stripe, dorsolateral color border, and lack of orange/red color

ventrally on the hindlimbs. It differs from all species of that group by the rounded light ventral

spots (generally at least partly vermiculated in the other species of the group), the brownish

posterior dorsum (of same color as anterior dorsum in the other species) and the dark brown

dorsal color of fore- and hindlimbs (lighter in the other species). It further differs from M.

betsileo and M. sp. 1 by the greenish rather than brown dorsum; from M. viridis by the entirely

dark brown flanks; and from M. expectata by the lack of bluish dorsal color on the dorsal

surface of the limbs and the lesser extent of the light ventral spots and markings. — (1)

Morphology of the holotype: Unknown, estimated SVL 25 mm. - (2) Dorsal color and pattern

of the holotype: Head and anterior part of dorsum yellowish green. Posterior part of dorsum

and flanks dark brown. Sharp dorsolateral color border present anteriorly. The yellowish

green dorsal color posteriorly ending straight (not semicircularly) and not covering the

posterior part of the dorsum. A thin, light, partly interrupted frenal stripe present. Limbs

dark brown with a very fine, irregular black dotting. Two dark crossbands on the hindlimb.

Iris with light pigment in its upper part. See also color pictures in GLAW & VENCES (1994) and

VENCES et al. (1996). — (3) Ventral color and pattern of the holotype: Black with a relatively large

number of small, regularly rounded blue markings which become smaller anteriorly. Horse-

shoe marking present.

Source : MNHN, Paris

18 ALYTES 17 (1-2)

Mantella sp. 1

Mantella n. sp. 3: CLARK, 1994.

Chresonyms:

Mantella betsileo: BLOMMERS-SCHLÔSSER & BLANC, 1991 (part.): GLAW & VENCES, 1992a (part.; see

localities), 1994 (part.; see localities).

Mantella cf. betsileo: VENCES et al, 1996.

Identity. — VENCES et al. (1996) first mentioned the presence of this form in Ankarana, based

on the observations of J. KÔHLER. R. NUSSBAUM (personal communication) found it in the

spiny desert of southern Madagascar and considered it as a species distinct from M. betsileo.

Comment. — No scientific name is currently disponible for this form. Formal description of

this species will be the subject of a forthcoming paper.

Material examined. — ZFMK 61238-41 and ZFMK 62197-9 (Ankarana; leg. J. STEINBRECHER 1995;

61241: CS). Several MNHN specimens can also be referred to this species: MNHN 1973.484-96

(Androatsalo): MNHN 1973497 (Tongahybe [Baly-Ouest}}; MNHN 1973.498 (Androatsalo);

MNHN 1976.214-8 (Morondava; SVL 26.4 mm [214], 23.3 mm [215], 26.1 mm [216], 21.7 mm [217], 22

mm (218). The specimens MNHN 1973.217-8 may also belong to M. betsileo which is known from

Kirindy near Morondava (KUCHLING, 1993; GLawW & VENCES, 1994). Due to the large size of the

specimens MNHN 1976.214 and MNHN 1976.216, we here refer the whole series to M. sp. L. We also

refer to the species in a preliminary way a BMNH series: BMNH 74.10.29.1-4 (Mohambo; purch.

M. BoucarD; NIL; four females, SVL 26.5 mm, 25.9 mm, 25.4 mm, 21.0 mm, no light spots in thorax

region).

Several other MNHN specimens with unknown localities may be referred to M. sp. 1 based on their

large size: MNHN 1976.183-6 (SVL 23.2 mm [183], 24.6 mm [184], 26.6 mm [185], 26.9 mm [186]):

MNHIN 1976.18 (SVL 23.6 mm); MNHN 1976.191 (SVL 24.2 mm); MNHN 1976.193 (SVL 27.3 mm);

MNHN 1976.194 (SVL 25.1 mm); MNHN 1976.197 (SVL 24.5 mm): MNHN 1976.198 (SVL 26.4 mm);

MNHN 1976.199 (SVL 27.1 mm).

Distribution. — [1] Ankarana; [2] Tongahybe; [3] Morondava; [4] Androatsalo (Androatsabo

according to BLOMMERS-SCHLÔSSER & BLANC, 1991); [5] Mohambo (locality not traced

and not included in map). According to NUssBAUM (personal communication; see also

CLARK 1994), large populations of this species occur in the spiny desert of south-western

Madagascar.

Diagnosis. — (1) Morphology: A large, rather stout Mantella. SVL 22-30 mm, males 25 mm,

females 29-30 mm. TTA reaching the posterior eye margin in small specimens (males),

between forelimb insertion and tympanum in large females. Terminal disks of fingers and toes

slightly expanded. Mean tympanum/eye ratio nearly 3/5. IMT medium sized (ratio

width/length about 2/3). — (2) Dorsal color and pattern: Dorsal head surface and dorsum

yellowish to light brown, mostly without diamond marking. Sharp dorsolateral color border

anteriorly present. Flanks black, with fiery red color extending posteriorly. White frenal stripe

present. Limbs brown to red-brown, with at least one dark brown crossband on femur, tibia

and tarsus. Iris with light pigment in its upper part. (3) Ventral color and pattern: Black with

blue markings, of different size and extension but often showing vermiculated patterns and

fusing with each other. Breast region generally without or with only very small light markings.

Distinct horseshoe marking present, sometimes including a central stripe.

Source : MNHN, Paris

VENCES, GLAW & BÜHME 19

Mantella viridis Pintak & Bôhme, 1988

Mantella viridis Pintak & Bôhme, 1988. — Name-bearing type: holotype by original designa-

tion (PINTAK & BÔHME, 1988: 120), ZFMK 47900, female, SVL 30.3 mm. — Type locality: “südlich

Antseranana (= Diego Suarez), Nord-Madagaskar”, according to original description. — Other ypes:

11 (lost) paratypes (see comment below). - Etymology: derived from Latin viridis (green).

Mantella viridis: PiTaK, 1990; Ouvert, 1990 (fig.); BLOMMERS-SCHLÔSSER & BLANC, 1991 (p. 274);

ZimMERMANN, 1992; GaRRAFFO et al., 1993; ANDREONE, 1992 (plate III fig. 5-6); GLAW & VENCES,

1992a, 1994; HERRMANN, 1993 (fig.); Le BERRE, 1993 (fig. p. 20); ZIMMERMANN & ZIMMERMANN,

1994; Barrierr, 1995 (fig. p. 17): HeNKeL & ScHminr, 1995 (fig. p. 57); CarissiMI-PRIORI, 1995

Gig. p. 43); Vences et al., 1996, 1998; DaLv et al., 1996, 1997a; STANISZEWSKI, 19976 (fig. p. 6);

LaRsEN, 1997; PrNTAK et al., 1998; VENCES & KNIEL, 1998.

Mantella spec.: Van ToMME, 1988 (fig. 2).

Other chresonyms:

Mantella betsileo: Buse, 1981 (part); BLOMMERS-ScHLôssER & BLANC, 1991 (part);

GLaw & Vences, 19924 (part.; see localities), 1994 (part; see localities).

Mantella expectata: SraNiszEWskt, 1997a (fig. p. 12).

Comment. — In the original description (PINTAK & BÔHME, 1988), 11 living, uncatalogued

specimens (four males and seven females, with same locality data as holotype) were desig-

nated as paratypes. No specimens of this captive stock were eventually preserved and cata-

logued: all paratypes must therefore be considered as lost.

Material examined. - ZFMK 47900 (according to catalogue: Mtge. d'Ambre, S of Diego [Antseranana];

leg. D. Berz 1987; holotype); ZFMK 48038-53 (Antseranana [Diego Suarez}; leg. H. MEIER 111.198;

48048: CS); ZFMK 62708-9 (locality unknown: CS); ZFMK 62710-2 (locality unknown: TE); MRSN

A416 (locality unknown: through the pet trade)

Three specimens (MNHN 1976.211-3; Montagne des Français) are also referred to M. viridis based

on their locality, size and relative hindlimb length. They clearly cannot be attributed to M. bersileo as

in the MNHN catalogue, but due to the faded colors we cannot completely exclude their belonging to M.

sp. 1. MNHN 1992.4820 (locality unknown) is here also referred to M. viridis based on size and relative

hindlimb length, although color patterns are not recognizable any more.

Distribution. — Only known from the northern tip of Madagascar. The published type locality

is south of Antsiranana. The only reliable localities known are: [1] 13 km south of Antsira-

nana (DaLv et al., 1996): [2*] Montagne des Français (GLAW & VEN 1994; ca. 100-300 m

altitude), south of Antsiranana. ZFMK specimens with the locality “Antseranana” were

most probably collected in the Montagne des Français. ANDREONE (1992) showed pictures of

Mantella viridis from “area of Montagne d’Ambre National Park” (plate II fig. 5-6), but

previously stated (p. 423) that he had not observed the species in nature and that locality

information was based on PINTAK & BÔHME (1988). Most probably the species is not present

in the Montagne d’Ambre National Park since recent surveys failed to find it (RAXWORTHY &

NUSSBAUM, 1994: GLAW & VENCES, 1994).

Diagnosis. —(1) Morphology: À large, rather stout Mantella. SVL, males 22-25 mm, females

27-30 mm. TTA reaching the eye center in many specimens (mostly males), but only the

forelimb insertion in large females. Terminal disks of fingers and toes expanded. Mean

tympanum/eye ratio about 2/3. IMT medium sized (ratio width/length slightly less than 2/3).

— (2) Dorsal color and pattern: Head, dorsum and largest (posterior) part of the flanks light

green to yellowish. Anterior part of the flanks black, this color reaching in many specimens to

a point around the forelimb insertion. A sharp dorsolateral color border present in this area.

Source : MNHN, Paris

20 ALYTES 17 (1-2)

Distinct, white to light green frenal stripe present. Color of limbs generally similar to dorsum,

without a dark crossband, but hindlimbs with a metallic blue shade in some specimens. Iris

with light pigment in its upper part. (3) — Ventral color and pattern: Black with bluish white

markings decreasing in size from posteriorly to anteriorly, generally absent on the breast.

Markings of irregular shape, often vermiculate and fusing with each other. Distinct horseshoe

marking present on the throat.

Mantella laevigata group

Mantella laevigata Methuen & Hewitt, 1913

Mantella laevigata Methuen & Hewitt, 1913. - Name-bearing type: holotype by original designation (METHUEN

& HewiTT, 1913: 58), TM 10074 (ex 1214), sex unknown, SVL 22.5 mm. - Tÿpe locality: “Folohy”,

according to original description and TM catalogue. — Other types: six paratypes according to original

description (ex 1212, 1215-9), TM 10085-8, TM 10090 and MCZ 10815 (see comment below). - Etymology:

derived from Latin faevigare (to make smooth)

Mantella laevigata: Gui, 1964 (syn. cowant), 1978 (syn. cowani); BUSSE, 1981; Mr1ER, 1986: PINTAK,

1990; BLOUMERS-SCHLÔSSER & BLANC, 1991; ANDREONE, 1992 (plate IV fig. 1-2); GLAW & VENCES,

1992a-b, 1994; GararrO et al., 1993; HERRMANN, 1993 (fig); Le BeRRE, 1993 (fig. p. 20); HENKEL

& Scmibr, 1995 (fig. p. 54); BarTLerT, 1995 (fig. p. 24); DALY et al., 1996; STANISZEWSkI, 1997a

(fig.), 19976 (fig.), 1998a (fig); LARSEN, 1997; GLAW et al., 1998a; PINTAK et al., 1998; VENCES ct

al, 1998; VENCES & KNiEL, 1998

Other chresonyms:

Mantella cowani: GUWBÉ, 1964 (part.), 1978 (part.).

Mantella madagascariensis: BLOMMERS-SCHLÔSSER & BLANC, 1991 (part; included in syntype series

of subspecies M. m. nigricans, locality Marojezy; same applies to BUSSE, 1981, and GLAW &

VENCES, 1992a, 1994)

Pictured in Meter (1980: fig. p. 353 below) as “Bisher nicht cindeutig einzuordnende Mantella-

Art”.

Comments. —(1) The holotypeis in good state of preservation. The pattern is largely faded but

still recognizable on head and anterior dorsum. A few of the light ventral spots are still

recognizable; they are small and rounded. The paratypes TM 10085-6 are in a rather bad state

of preservation; the pattern contrast is largely faded, and the ventral pattern is not recog-

nizable, TM 10088 and 10090 are also in bad state of preservation with faded pattern, but they

can clearly be assigned to M. lacvigata by their broad finger disks. TM 10087 is in good state

of preservation; pattern contrast is weak, but both dorsal and ventral (small rounded spots)

pattern is still recognizable. — (2) Seemingly, the type locality “Folohy forest” does not exist

any more; its location was traced by BLOMMERS-SCHLÔSSER & BLANC (1991) immediately

north of Toamasina (Tamatave), and we follow this placement in our distribution maps. — (3)

Barbour & Loveridge (1929) mentioned the existence of one “syntype” in the TM and onein

the MCZ (MCZ 10815). However, the original description (a) clearly stated that it was based

on “seven examples”, listed as “1212, 1214-1219” and (b) separately mentioned specimen

1214 once more, as “type”. Although not explicitely stated, this infers the existence of one

holotype (ex 1214, today TM 10074); we consider all additional specimens listed in the

original description as paratypes. MCZ 10815 is almost certainly the specimen formerly

numbered TM 10089, which was exchanged with MCZ, according to the TM catalogue, on

Source : MNHN, Paris

VENCES, GLAW & BÔHME 21

8.VIL.1925. - (4) Whereas the locality of the holotype and of the paratypes TM 10087-8 and

MCZ 10815 is Folohy, that of the paratypes TM 10085-6 and 10090 is only “E Madagascar”,

according to the TM catalogue.

Material examined. - TM 10074 (Folohy, Eastern Madagascar; coll. by HERSCHELL-CHAUVIN, 1911;

holotype); TM 10085-6 and 10090 (Eastern Madagascar resp. East region, Malagasy Republic; coll.

HERSCHELL-CHAUVIN, 1912; paratypes); TM 10087-8 (Folohy, E-Madagascar; coll. HERSCHELL-

CHAUVIN, 1912; paratypes); BMNH 1952.11.53-54 (Mangabe island, Antongil bay: coll. C. S. WeBr;

NIL); MNHN 1973.534-40 (Marojezy, 300 m; paralectotypes of M. nigricans); MNHN 1973.542-7

(Marojezy, 300 m; paralectotypes of M. nigricans); MNHN 1973.549 (locality unknown; paralectotype

of M. nigricans); MNHN 1973.557-8 (Marojezy, 600 m; paralectotypes of M. nigricans); two juvenile

specimens of the MNHN collection most probably also belong to M. laevigata: MNHN 1973.517

(Marojezy 300 m; SVL 12.3 mm; paralectotype of M. nigricans), MNHN 1973.548 (Marojezy 300 m;

SVL 12.1 mm; paralectotype of M. nigricans), ZFMK 19298 (Maroantsetra: leg, H. MEIER 1976); ZFMK

48660 (Nosy Mangabe: leg. R. ZoBeL VI.1988); ZFMK 52747-51 (Nosy Mangabé; leg. F. GLAW & M.

VENCES IIL.1991; 52749 CS); ZFMK 59911 (Marojezy Camp L: leg. F. GLAW & O. RAMILISON II. 1995);

ZFMK 59912 (Marojezy Camp 2; leg. F. GLAW & O. RAMILISON I1.1995; juvenile), ZFMK 59913-4

(Marojezy Camp 3; leg. F. GLAW & O. RAMILISON II.1995); ZFMK 62786-8 (locality unknown: TE):

MRSN A0065.1-3 (Nosy Mangabe: leg. F. ANDREONE 24.1V.1990); MRSN A1826 (Tsararano Chain,

Camp I; leg. F. ANDREONE 4.XI1.1996), MRSN A1827, MRSN A1828.1-2 (Tsararano Chain, Camp 2;

leg. F. ANDREONE 13-14.XI1.1996).

Distribution. [1] Type locality Folohy. Recent localities from the East and North-East are:

[2*] the small island Nosy Mangabe (100-300 m altitude); [3] the Tsararano chain (700 m

altitude); [4*] the Marojezy massif (300-700 m altitude). Two additional localities from the

northern part of the Eastern Region are found in DALY et al. (1996): [5] Ambodimanga and

Varary, both in the Mananara reserve (ca. 100 m altitude). The locality Maroantsetra (based

on ZFMK 19298; see BUSSE, 1981) does almost certainly not refer to the town Maroantsetra

itself but to a nearby locality (most probably Nosy Mangabe) and is therefore not accepted

here.

Diagnosis. (1) Morphology: À medium-sized to large Mantella With a generally very slender

appearance. Terminal disks of fingers and toes largely expanded. SVL 22-29 mm. TTA

reaching generally the eye center and slightly beyond the eye in some specimens.

Tympanum/eye ratio between 1/2 and 3/5. IMT medium-sized (ratio width/length about 3/5).

— (2) Dorsal color and pattern: Head and anterior part of dorsum covered by a sharply

delimited yellow mark, posteriorly either ending semicireularly or prolonged as a pointed

triangle to the cloacal region, with a sharp dorsolateral color border to the black flanks and

sides of head. Variation in shape of dorsal yellow mark not corresponding to sexual dimor-

phism. Limbs deep black (exceptionally copper brownish). Hands and finger tips often with

blue spots. No frenal stripe, but single yellowish spots sometimes present under the eyes. No

flashmarks. Iris completely black without light pigment. —(3) Ventral color and pattern: Venter

and limbs black with small, rounded, bluish or bluish-grey spots. Throat generally completely

black without pattern (few light spots sometimes present). No red, orange or yellow pattern

on hindlegs.

Source : MNHN, Paris

22 ALYTES 17 (1-2)

Fig. 2. - Photographs of Mantella species. (a) M. haraldmeieri from Nahampoana, 1991; (b) M. cowani,

specimen without locality data (not preserved), 1994: (c) M. baroni, specimen without locality data

(not preserved, but belonging to the same series as ZFMK 62718-25), 1997: (d) M. af. baroni from

Andringitra (ZMA 6754), photograph taken from BLOMMERS-SCHLÔSSER & BLANC (1993); (e) M.

madagascariensis, specimen without locality data (not preserved, but belonging to the same series as

ZFMK 62732-41), 1997; (1) M. pulchra, specimen without locality data (not reserved), 1994; (g) M.

bernhardi from near Tolongoina (holotype, ZFMK 57164), 1994; (h) M. crocea, specimen without

locality data (ZFMK 52746), 1991

Source : MNHN, Paris

VENCES, GLAW & BÔHME 23

Mantella cowani group

Mantella baroni Boulenger, 1888

Mantella_Baroni Boulenger, 1888. — Name-bearing type: holotype by monotypy, BMNH 1947.2.7.19

{ex 84.12.2250), male (according to the original description), SVL 27.2 mm. - Tÿpe locality: not specified in

the original description: “Madagascar” without further specifications according to the BMNH catalogue.

— Etymology: named after the collector of the type, Reverend P. BARON.

Mantella Baroni: MOCQUARD, 1909.

Mantella baroni: WERNER, 1901; METHUEN & HewiTT, 1913; PARKER, 1925; GUIBÉ, 1964, 1978 (syn.

cowani); BUSSE, 1981 (syn. madagascariensis): BLOMMERS-SCHLÜSSER & BLANC, 1991 (syn. mada-

gascariensis), GLAW & VENCES, 1994 (syn. madagascariensis: p. 412); DALY et al., 1996, 1997b;

PinTak et al., 1998; VENCES et al., 1998; VENCES & KNIEL, 1908: STANISZEWSKI, 1998a (ig.).

Phrynomantis maculatus Thominot, 1889. — Name-bearing type: lectotype (designated by GLAwW & VENCES,

1994), MNHN 1991.2845 (ex 6807a), sex unknown, SVL 27.0 mm. - Type locality: “l'île de La Réunion”

according to original description (probably erroneous; see comment below). — Other types: paralectotypes

MNHN 1991.2846 (ex 6807b), MNHN 1991.2847 (ex 6807c) and MNHN 6807. — Etymology: derived from

Latin maculatus (spotted).

Phrynomantis maculatus: Guisé, 1964, 1978 (syn. cowani); Busse, 1981 (syn. madagascariensis);

BLOMMERS-SCHLÔSSER & BLANC, 1991 (syn. madagascariensis); GLAW & VENCES, 1994 (syn.

madagascariensis; p. 413).

Other chresonyms:

Mantella cowani: GUÉ, 1964, 1978 (part.); Marz, 1975 (fig.); Meter, 1975 (fig. 1-2); Meier, 1980

(üg. p. 352); OserLe, 1981 (pl. 29); Le Berke, 1993 (part.; outer fig. p. 21).

Mantella madagascariensis: BEUTELSCHIESS & BEUTELSCHIESS, 1987; PiNTAk, 1990 (part.); OLIVETTI,

1990 (fig.); BLOMMERS-SCHLÔSSER & BLANC, 1991 (part.); ANDREONE, 1992; GLaW & VENCES,

1992b (fig. p. 28); ZIMMERMANN & ZIMMERMANN, 1992 (fig. 5.21); GARRAFFO et al., 1993; CLARK,

1994 (fig. p. 10 below); VENCES et al., 1994 (fig. p. 390); GLAW & VENCES, 1994 (part.}; BARTLETT,

1995 (fig. p. 18 below right}; HeNkeL & ScHmiDr, 1995 (fig. p. 55); CARISSIMI-PRIORI, 1995

(üg, p. 42); Sraniszewskt, 1996 (plate p. 16-17); 1997a (fig. p. 12), 1997b (fig.); ZIMMERMANN,

1996b (fig. 14).

Mantella madagascariensis sensu stricto: GLAwW & VENCES, 19924 (part.; see localities).

Mantella madagascariensis madagascariensis: BUSSE, 1981 (part.); Meter. 1986 (fig. 6); VAN TOMME,

1988 (fig. 5-6); ANDREONE & GAVETTI, 1993 (p. 105).

— BUSSE (1981) defined M. madagascariensis as a very variable species containing

several junior synonyms, including M. baroni. One main problem with this definition was the

bad state of preservation of the M. madagascariensis types (see below) which made reliable

attribution of this name to any specific morph impossible. Recent studies have shown that

many of the forms previously summarized under the name M. madagascariensis do in fact

belong to separate well-defined, valid species (GLAW & VENCES, 1994). Two morphs (here

named A and B) remained without an unequivocal definition: morph A, figured on plate 61 in

GLAW & VENCES (1994), was considered as M. madagascariensis, Whereas the *

morph B, figured on plates 58-59 in GLaw & VENCES (1994), was considered as M. “loppei” in

a preliminary way. GLAW & VENCES (1994) mentioned that morph A corresponds to the type

of M. baroni which they listed as synonym of M. madagascariensis. DALY et al. (1996),

referring to this definition, argued that M. baroni should be revalidated as a valid name (for

morph A), whereas the name madagascariensis Should be seen as “nomen dubium”” due to the

bad state of the type specimens.

We here follow these conclusions as far as the definition of morph A as Mantella baroni

is concerned. M. madagascariensis, however, is not “unidentifiable” (DALY et al., 1996); a new,

detailed examination of the lectotype of that taxon showed that it corresponds to morph B

(see corresponding section).

Source : MNHN, Paris

24 ALYTES 17 (1-2)

Comments. — (1) The holotype of M. baroni is in rather bad state of preservation, but the

relevant color patterns can still be recognized. - (2) According to BLOMMERS-SCHLÔSSER &

BLANC (1991), the number BMNH 1947.2.7.19 defines “syntypes” of M. baroni; however,

BOULENGER (1888) in his original description mentioned explicitely “a single male specimen”.

We found no indications on the existence of types other than a single holotype in the BMNH

collection and catalogue. The specimens ZFMK 28770-28772 cannot be seen as M. baroni

“paratypes” as was suspected by BUSSE (1981); their collecting data agree with those of the

series BMNH 95.7.4.34-6 and 96.12.2.28-31, but not with those of the holotype. -(3) The type

locality of Phrynomantis maculatus, according to the original description, is “Île de La

Réunion”. BUSSE (1981) first gave the locality “Nosy Cumba-Nosy Be” without providing

additional information nor his source of information. This locality was subsequently also

given by BLOMMERS-SCHLÔSSER & BLANC (1991) but was questioned by GLAW & VENCES

(1994). According to A. OHLER (in litteris, 1997), a second MNHN catalogue informs that the

specimens were supplied by the “Com. scientifique de Bourbon” (Bourbon is an old name for

the island of La Réunion). This explains the wrong locality information “Réunion”, which

was later corrected to “Nossi-Be et Nossi-Cumba” in one MNHN catalogue, and to “Mada-

gascar” in a second catalogue. We consider also the Nosy Be — Nosy Komba locality

information as wrong (see below).

Material examined. - BMNH 1947.2.7.19 (holotype; Madagascar: leg. R. BARON); BMNH 95.7.4.34-6,

BMNH 96.12.2.28-31 (Ambohimitombo forest; coll. FoRsYrH MAJOR [1 specimen exch. Vienna 1912],

NIL); BMNH 1925.7.13.1-6 (Madagascar; coll. ForsyTH Major; NIL); BMNH 1925.7.2.57 (Antsiha-

naka; purch. ROSENBERG); BMNH 1930.2.2.1 (Analamazoatra forest, environs of Périnet; purch. ROSEN-

BERG): BMNH 1953.1.5.42-5 (Madagascar, pres. G. W. ALLAN); MNHN 6807, MNHN 1991.2846-7 (ex

MNHN 6807; “Nosy Komba”; paralectotypes of Phrynomantis maculatus); MNHN 1991.2845 (ex

MNHN 6807A; lectotype of Phrynomantis maculatus; “Nosy Komba”); MNHN 1883.584 (locality

unknown; ded. HumBLor); MNHN 1902.335 (Ikongo; M. Bensch}; MNHN 1907.161-2, MNHN

1991.1813 (locality unknown; obtained from the “section de Madagascar à l'Exposition coloniale de

Marseille”; ex MNHN 1907.162, 162A); MNHN 1931.14 (locality unknown); MNHN 1991.1807-9

(locality unknown; ex MNHN 1931.14.A-C); MNHN 1931.15 (Moramanga); MNHN 1991.1810-2

(Moramanga; ex MNHN 1931.15.A-C); MNHN 1931.16-7 (SE Fianarantsoa; DECARY 1926-1930):

MNHN 1933.247 (Ruisseau d'lorantjatsy: Distr, Fianarantsoa, alt, 1000 m); MNHN 1936.40-2 (Forêt de

Tsianovoha); MNHN 1936.43-6 (probably “forêt de Tsianovoha”; HriM); MNHN 1953.135 (Anosibe

[Moramanga]} MNHN 1972.775-6 (Moramanga); MNHN 1976.233-4 (locality unknown); MNHN

1988.7599 (locality unknown; don. O. BenRA III.1988); MNHN 1993.1441-2, MNHN 1993.1444,

MNHN 1993.1446-7 (locality unknown); ZFMK 14208 (Niagarakely; leg. H. Mer 1972); ZFMK

28870-2 (Ambohimitombo forest; leg. ForsyrH-Masor 1903; originally Museum Gôttingen): ZFMK

460358 (locality unknown: through pet trade; 46035 CS); ZFMK 47008-9 (Moramanga; leg. R. Serpr

IV.1987); ZFMK 48054-60 (120 km S Moramanga: Marolamba; leg. H. MER III.1988; 48055 CS.

ZFMK 50161-3 (Moramanga; leg. H. MEiER I1.1989); ZFMK 50551 (Moramanga: leg. F. W. HENKEL, W.

Scumibr & V. MüLLER 1989); ZFMK 56165-9 (through pet trade, ded. F GLAW XI.1993); ZFMK 62242

(Mantady, le LAW 11.1996); ZFMK 62718-21 (locality unknown, CS), ZFMK 62722-$ (locality

unknown; TE): ZFMK 62287-8 (juveniles) and 64139-40 (all Vohiparara: leg. F. GLAW, D. RAKOTOMA-

LALA & F. RANAIVOIAONA I11.1996; TE); MRSN A0061.1-4 (Andasibe, Amalonabe: leg, F. ANDREONI

2.XI1.1991); MRSN A0066.1-5 (Vatoharanana-Ranomafana, lo lfanadiana; leg. F. ANDRFONE

8.11.1993; NIL); MRSN A0067.1-5 (Vohiparara: leg. F. ANDRFONE 9.11.1993); TM 9890, 9896, 9900

(Analamazoatra: leg. METHUEN).

The following somewhat deviating specimens are also attributed to M. haroni in a preliminary way

(see discussion below): TM 9888-9, 9892, 9895, 9898-9 (Folohy: coll. MErHuEN): TM 9894 (Folohy: coli.

HERSCHELL-CHAUVIN); BMNH 1986.2 (Camp 4, Zahamena, 17°40'S, 48°50°W; leg. C. J. RAXWORTHY

8.IX.1985).

Source : MNHN, Paris

VENCES, GLAW & BÔHME 25

Distribution. — The species of the M. cowani and M. madagascariensis groups (as defined

in the present study) were insufficiently distinguished in previous works. The corresponding

distribution maps (mainly in BLOMMERS-SCHLÔSSER & BLANC, 1991, as M. madagascariensis

and M. cowani) did not contain references to literature records or voucher specimens.

GLaw & VENCES (1994) assigned some localities to the species haraldmeieri, cowani,

pulchra and “loppeï”, but most localities remained without reliable attribution to any

species.

The distribution map of BLOMMERS-SCHLÔSSER & BLANC (1991) was mainly based on

MNHN voucher specimens. All of these were examined by us. This allows us for the first time

to outline the distribution of the different species with a certain reliability. M. baroni occurs in

the central Eastern Region, mainly at mid-altitude localities: [1] Antsihanaka; [2*] Ankeni-

heny (ca. 1000 m altitude); [3*] An’Ala (ANDREONE, 1993; DaLy et al., 1996; personal

observation at ca. 840 m altitude); [4*] Analamazoatra; [5] Anosibe (Anosibeanala); [6]

Niagarakely; [7] Marolamba (120 km S Moramanga; probably identical with Marolambo,

which is situated about 100 km S Moramanga; see BLOMMERS-SCHLÔSSER & BLANC, 1993); [8]

Ambohimitombo; [9] Ikongo: [10] Ruisseau d’Iorantjatsy; [11] Forêt de Tsianovoha; [12*]

Ranomafana National Park (ANDREONE, 1992; GARRAFFO et al., 1993; personal observation

near Vohiparara, ca. 1000 m altitude); [13*] Mantady. Additional localities were published by

DaLy et al. (1996): [14] Sahavondrona (near Ranomafana; ca. 1000 m altitude); [15] 30-35 km

south of Moramanga.

Two additional localities, [17] Folohy and [18] Zahamena (TM and BMNH vouchers, see

above) are attributed to M. baroni only in a preliminary way. These specimens, which

unfortunately have largely faded color patterns, show a deviating coloration which resembles

M. nigricans in many respects. In the Folohy sample, the ventral side including the femur is

dark with small (not large as usually in M. baroni) rounded light spots (no horseshoe

marking). The tibia and the foot are light (except TM 9888 which has a dark tibia). The flank

blotches are large and rounded, as typical for M. baroni. The rostral stripe appears indistinct

without sharp borders, and the head surface may have been lighter than the back in life. The

single known Zahamena specimen, according to the attached field label, had the following life

coloration “Back and legs vivid bright green, flanks black, lower back and legs brown, belly

black with pale blue spots, iris black.” In preservative, the pattern is largely faded. Femur

and tibia are dark, but the foot is light ventrally and dorsally. The existing information

on these specimen does not allow for further statements; in the distribution map, we list the

two localities as intermediate between M. baroni and M. nigricans. The color and pattern

information given below for M. baroni applies to all populations except for Folohy and

Zahamena.

The remaining localities listed by BLOMMERS-SCHLÔSSER & BLANC (1991) for M. mada-

gascariensis can be assigned as follows: Marojezy refers to M. nigricans; Antsihanaka is the

type locality of M. pulchra: Marolambo is the type locality of M. loppei (junior synonym of

M. madagascariensis according to the present study), and seems also to be a locality of M.

baroni (see above: if Marolambo and Marolamba are identical); Ambalavato is the type

locality of M. madagascariensis; Iremo, Ambatodradama and Betafo refer to M. cowani,

Chaînes Anosyennes, Ambana, Bekazaha and Soavala refer to M. haraldmeieri; Xvohibe and

Marovitsika refer to M. aff. baroni which is here considered separately (see below)

Source : MNHN, Paris

26 ALYTES 17 (1-2)

We propose to delete the localities Nosy Be and Nosy Komba (which are based on a

dubious locality information referring to the types of Phrynomantis maculatus:; see above). As

discussed by GLAW & VENCES (1994), these localities are in the Sambirano region where recent

extensive surveys have only yielded records of species of the M. betsileo group. We also

propose to ignore the locality Ambohidratrimo, located 20 km NW of the Malagasy capital

Antananarivo (VIETTE, 1991) near the Ivato airport. No vouchers for this locality were found

in the MNHN. The presence of habitat structures suited for species of the M. baroni group or

M. madagascariensis group is not probable at this locality for the last 100 years.

Diagnosis. -(1) Morphology: A large, slender Mantella. SVL 22-30 mm. TTA mostly reaching

the eye center but at least the tympanum. Terminal disks of fingers and toes expanded.

Tympanum/eye ratio generally 3/5. IMT small (ratio width/length about 4/5). - (2) Dorsal

color and pattern: Head, dorsum and flanks deep black, without dorsolateral color border.

Frenal stripe absent. Yellowish rostral stripe present, generally not in contact with flank

blotch. Forelimb (except the mostly black fingers) and femur yellow to greenish. This color

continuing onto the flanks, forming relatively large, rounded flank blotches. These sometimes

dorsally expanding onto the back, not being delimited by the dorsolateral border. Size of

blotches variable, but in none of the examined specimens blotches of opposite flanks

contacting each other on the back. Tibia, tarsus and foot orange with irregular black

crossbands and markings. No flashmarks. Iris completely black without light pigment. — (3)

Ventral color and pattern: Venter, throat and limbs black with few relatively large, rounded

light markings which are generally not blueish but yellow to greenish. No horseshoe marking,

throat with only a single rounded marking, sometimes completely black. Tibia, tarsus and

foot orange as dorsally, but mostly without black patterns. The orange color sometimes

reaching the distal part of the femur but not further proximally. Exceptionally, single

specimens with a nearly complete horseshoe marking (observed in one specimen of the series

MRSN A0066).

Mantella aff. baroni (from Andringitra)

Chresonyms:

Mantella cowani: Gus

Mantella madagascariens

VENCES, 1994 (part.).

Mantella madagascariensis sensu stricto: GLAW & VENCES, 19924 (par

Mantella madagascariensis madagascariensis: BUSSE, 1981 (part. fig. 5).

1964 (part. fig. 4-6), 1978 (part.).

: BLOMMERS-SCHLÔSsER & BLANC, 1991 (part.), 1993 (pl. 19 fig. 104); GLaw &

e localities and fig. 180).

Identity. — Within and between the known populations of Mantella baroni, the dorsal and

ventral coloration of adults as described above is rather uniform (see also ANDREONE, 1992;

Glaw & Vences, 1994). On the contrary, specimens from Andringitra (south of all other

known localities of M. baroni) differed by an enormously variable dorsal pattern (see below).

We here consider the Andringitra population as a separate form Mantella aff. baroni which

clearly is very closely related to M. baroni. Final clarification of its status is not possible at

present.

Comment. — No scientific name is currently disponible for this form.

Source : MNHN, Paris

VENCES, GLAW & BÔHME 27

Material examined. — The following specimens can clearly be assigned to this form due to their largely

extended dorsal green-yellow pattern: MNHN 1953.136 (Col d’Ivohibe, Andringitra}; MNHN

1991.1800-4 (Col d'Ivohibe, Andringitra; ex MNHN 1953.136.A-E); MNHN 1972.67, MNHN

1972.769-72, MNHN 1972.74, MNHN 1972.777-8 (Col d’Ivohibe, forêt Marovitsika).

Several other specimens differ from typical M. baron only by a gradually larger extension of the

yellow pattern. These are: MNHN 1991.1805-6 (Col d'Ivohibe, Andringitra; ex MNHN 1953.136.G-H);

MNHN 1972.763-6 (Col Ivohibe, forêt Marovitsika); MNHN 1972.768, MNHN 1972.773 (Col Ivohibe,

forêt Marovitsika). MNHN 1991.1805 is most similar to typical M. baroni by dorsal pattern. MNHN

1972.763-5 are very large and stout specimens, probably females.

Distribution. — Only known from the Col d’Ivohibe [1] in the Andringitra massif.

Diagnosis. —(1) Morphology: A large, slender Mantella. SVL 27-31 mm, females 30 mm. TTA

reaching eye center in some specimens, but only to forelimb insertion or slightly beyond in

large females. Terminal disks of fingers and toes expanded. Tympanum/eye ratio generally 1/2

to 3/5. IMT small. — (2) Dorsal color and pattern: In some specimens similar to typical M.

baroni, but with a larger extension of the yellow flank blotches which reach widely onto the

dorsum (coloration observed in all specimens identified as females). Other specimens, by

general body proportions possibly mainly males, showing a broad dorsal contact of the flank

blotches, or a further increase of these, resulting in a nearly uniformly yellow pattern dorsally

(see GUIBÉ, 1964: fig. 4-6; BUSsE, 1981: fig. 5), with the yellow color also extending onto the

tibia, which is otherwise orange with black. No flashmarks. Iris ssemingly with some light

pigment in its upper part according to the color plate in BLOMMERS-SCHLÔSSER & BLANC

(1993; here reproduced in black-and-white on fig. 2) which shows a specimen relatively similar

to typical M. baroni, with a (very indistinct) dorsolateral color border. Rostral stripe present

and generally in contact with the flank blotches. In specimens with large extension of yellow

color, the rostral stripe is the sharp border between yellow dorsal and black lateral color of the

head. — (3) Ventral color and pattern: Similar to M. baroni, but with a higher number and

smaller size of light markings (intermediate between M. baroni and M. haraldmeieri). Infor-

mation on the color of the light markings in life not available.

Mantella cowani Boulenger, 1882

Mantella cowani Boulenger, 1882. — Name-bearing type: lectotype, by present designation, BMNH 194

{ex BMNH 82.3.16.38), female according 10 BOULENGER (1882), SVL 28.2 mm. — Type localit

Betsileo”, according 10 the original description and the BMNH gue. — Other types: parak

following present lectotype desienation, BMNH 1947 2.7.5 (ex BMNFI 82.3.16.30), female according to

BOULENGER (1882). - Erymology: named after the collector of the type series, Reverend W, Deans COWAN.

Mantella cowanii: DaLY et al., 1996: STANISZEWSKI, 1998a (fig).

Mantella Cowani MOCQUARD, 1909.

Mantella cowani: Werner, 1901; METHUEN & HEWITT, 1913; PARKER, 1925: Guibé, 1964, 1978 (part.):

Busse, 1981 (syn. madagascariensis); BLOMMERS-SCHLOSSER & BLANC, 1991 (part.); BOHME et al..

1993; VENCES et al., 1994; GLAW & VENCES, 1994; BARTLETT, 1995 (fig. p. 24): CARISSIMI-PRIORI,

1995 (fig. p. 43}: SraniszEwskt, 1997a (fig.), 1997b (fig.): LARSEN, 1997; VENCES & KNIEL, 1998.

Mantella cowant s. Str.: ANDREONE, 1992 (p. 438).

Other chresonyms:

Mantella madagascariensis: BLOMMERS-SCHLÔSSER & BLANC, 1991 (part)

Maniella madagascariensis madagascariensis: BUSSE, 1981 (part.).

Mantella madagascariensis nigricans: BUSSE, 1981 (part; MNHN 9594 on p. 33).

Mantella madagascariensis (color morph Mantella “cowant” ): GLAW & VENCES, 19924.

Source : MNHN, Paris

28 ALYTES 17 (1-2)

Identity. — See BÔHME et al. (1993) and VENCES et al. (1994) for the confusing taxonomic

history of the taxon. The species is well distinguished by its typical pattern.

Comments. — (1) Lectotype and paralectotype are in excellent state of preservation. The

typical pattern is still recognizable, although the red color has largely faded. — (2) Although

the specific name was written cowanii in the original description, we here continue using the

spelling cowani which was used by most subsequent authors, since the Code allows both

spellings to be used.

Material examined. - BMNH 1947.2.7.4-$ (lectotype and paralectotype; E-Betsileo: leg. W. D. CowAN):

MNHN 1906.171, MNHN 1991.2844 (Betafo); MNHN 1973.528-9 (Ambatomenaloha, massif Itremo);

MNHN 9594 (Ambatodradama, 2000 m; ARNOULT 11.XI1.1962); ZFMK 59822 (locality unknown; ded.

F. GLaw IV.1995); ZFMK 62726-7, ZFMK 62729-31 (locality unknown; TE); ZMB 10404, ZMB

50106-7 (East-Betsileo; leg. HILDEBRANDT).

Also the following specimens with unknown locality are considered as M. cowani based on unpub-

lished electrophoretic and morphometric data: ZFMK 62728 (TE; color in life yellow/black instead of

red/black); ZFMK 62719, ZFMK 62721 (TE; color in life yellow/black instead of red/black, and

extension of yellow color intermediate between M. cowani and M. baroni).

Distribution. — The type locality “East Betsileo” comprises a large area and does not represent

a concrete locality. According to a personal communication of A. PEYRIERAS, the species

occurs: [1] in forested regions of the highlands SE of Ambatolampy and [2] near Antoetra.

MNHN vouchers corroborate the localities [3] Betafo, [4] Itremo and [5] Ambatodradama.

See the discussion in the section on distribution of M. baroni.

Diagnosis. — (1) Morphology: A large, slender Mantella. SVL 22-29 mm. TTA mostly not

reaching the eye but between forelimb insertion and tympanum (only reaching forelimb

insertion in a few specimens). Terminal disks of fingers and toes nearly not expanded.

Tympanum/eye ratio generally 1/2-3/5. IMT medium-sized (ratio width/length about 2/3). -

(2) Dorsal color and pattern: Head, dorsum and flanks deep black. Rostral and frenal stripes

absent. Proximal part of femur and humerus generally red (exceptionally orange or yellow).

This color extending on the flanks as small flank blotches, and also present as a broad band on

tarsus and foot (sometimes disrupted by black markings). A light spot below the eye

sometimes present. AIl remaining dorsal surface uniformly black. No flashmarks. Iris com-

pletely black without light pigment. —(3) Ventral color and pattern: Black with relatively large,

cireular whitish-blue markings. Single markings on throat, but no horseshoe marking. Limbs

also black with whitish-blue markings, except broad red bands on tibia, tarsus and foot which

correspond to those on the dorsal surface.

Mantella haraldmeïeri Busse, 1981

Mantella madagascariensis haraldmeieri Busse, 1981. — Name-bearing type: holotype by original designation

(BUSSE, 1981: 34), ZFMK 25351, male, SVL 22.0 mm. - Tipe locality ar”,

according 10 original description and ZFMK catalogue. - Other 1}

25352. - Etymology: named afte man amateur herpetologist Harald Mr1Er who collected the types.

Mantella madagascariensis haraldmeieri: Bone & BiSCHOrF, 1984; MEIER, 1986,

Mantella haraldmeïeri: PiNTAK, 1990: ANDREONE, 1992 (plate IV, fig. 5-6): GLAW & VENCES, 19924,

1992 (fig. p. 28): ANDREONE, 1903 (fig. 2); BÔHME et al., 1993; HERRMANN, 1993 (fig): VENCI

al, 1994 (fig. p. 392): GLaw & VENCES, 1994: HekeL & ScHMiDT, 1995 (fig. p. 53): STANISZ

1997b (fig.): LARSEN, 1997; Pak et al.. 1998: VENCES et al., 1998: VENCES & KNIEL, 1998.

Source : MNHN, Paris

VENCES, GLAW & BÜHME 29

Other chresonyms:

Mantella Cowani: MOCQUARD, 1902.

Mantella cowani: BACHMANN & BLOMMERS-SCHLÔSSER, 1975; BLOMMERS-SCHLÔSSER, 1978; 19794;

BLOMMERS-ScHLÔSSER & BLANC, 1991 (part.), 1993 (pl. 19 fig. 103).

Mantella betsileo: Meier, 1980 (part: p. 353, second fig. above); BUSsE, 1981 (part.: tab. 1, specimens

from Anosyennes).

Identity. — M. haraldmeieri has been generally considered as a separate species in recent years

(see BÔHME et al., 1993), mainly based on color patterns as (1) a light dorsum sharply

bordering the dark flanks, (2) dorsally uniform hindlimb coloration and (3) small, beige flank

blotches. MNHN specimens from the Anosy mountains (Chaînes Anosyennes) in southern

Madagascar, near the haraldmeieri type locality Tolagnaro, were erroneously identified as

Mantella betsileo by BUSSE (1981). A detailed re-examination of this large series showed that

all specimens are clearly to be assigned to M. haraldmeieri based on color patterns (1)-(3) as

defined above, and further on (4) presence of an unforked sternum (VENCES et al., 19994), (5)

lack of a horseshoe marking, (6) presence of small, rounded light spots on the venter, and (7)

ventrally uniformly light colored tibia, tarsus and foot.

In most MNHN specimens, the dorsolateral coloration border is very indistinct or

absent; we presume that the dorsal darkening was caused by the formalin fixation to which the

specimens most probably have been exposed (see section on M. nigricans). In fact, in at least

one specimen (MNHN 1973.511), the coloration border is still clearly recognizable.

In several MNHN specimens, the flank blotches are larger than described until present

for M. haraldmeieri (see fig. 10), indicating the close relationships of M. haraldmeieri with the

remaining species of the M. cowani group.

Comments. —(1) M. haraldmeieri, according to our personal observations, does not occur in

the coastal town Fort Dauphin (Tolagnaro), the type locality, itself, but in nearby rainforest

remains near Nahampoana. — (2) Probably due to a typing error, BUSsE (1981) did not

mention the specimen ZFMK 25353 which has similar collection data as the holotype and

paratypes, and was listed in the appendix of BussE’s (1981) work. Although this specimen was

originally catalogued as paratype, it cannot therefore be considered as such (and was not listed

in the account of BôHME & BISCHOFF, 1984).

Material examined. - MNHN 1901.232 (Fort Dauphin; envoi de M. ALLUAUD: pigments totally faded,

identification by size, locality and medium-sized IMT); MNHN 1973.49 (Soavala-Ambana, Chaînes

Anosyennes); MNHN 1973.500 (Beampigaratra, Nord Bekazaha, alt. 950 m); MNHN 1973.501 (Camp

IV?) MNHN 1973.502-11 (Camp IV, Ambana); MNHN 1973.512-16 (Camps IV et III bis; MNHN

1973.518-20 (Camps IV et III bis); MNHN 1973.521-7 (Ambana); ZFMK 21805-7 (Fort Dauphin; leg.

H. Mare 1.1978; paratypes): ZFMK 25351 (Fort Dauphin: leg. H. ME1ER 1978; holotype): ZFMK 25

(Fort Dauphin; leg. H. MEiEr 1978; paratype): ZFMK 25353 (Fort Dauphin; leg. H. MER 1978);

ZFMK 47831-3 (Fort Dauphin: leg. H. Meter 11988): ZFMK 48181-2 (Fort Dauphin: leg. F. W

HexkeL & R. Sep 1988); ZFMK 52741-3 (Nahampoana near Fort Dauphi FE. GLAwW & M. VENCES

1991; 52741 CS); MRSN A0062.1-2 (Nahampoana: leg. F. ANDREONE 14.1V.1990); MRSN A0063

(Nahampoana: leg. R. NINCHERI 14.1V.1990).

Distribution. - ZFMK specimens with a reliably known locality were collected in [1*] near

Nahampoana. A. PEYRIERAS (personal communication) found the species in [2] Mahatalaha.

MNHN vouchers demonstrate that the species is the only Mantella so far known in the Anosy

mountain chain. Locali are: [3] Chaînes Anosyennes: [4] Ambana; [5] Bekazaha; [6]

Soavala. See the discussion in the section on distribution of M. baroni.

Source : MNHN, Paris

30 ALYTES 17 (1-2)

Diagnosis. (1) Morphology: À medium sized to large, relatively slender Mantella. SVL 21-27

mm. TTA mostly reaching the eye center but in some specimens only the tympanum. Terminal

disks of fingers and toes expanded. Tympanum/eye ratio generally 3/5. IMT medium-sized

(ratio width/length about 2/3). (2) Dorsal color and pattern: Dorsun light brown with three

regular dark brown patterns: (a) an either triangular or inversely Y-shaped marking in the

shoulder region; (b) a larger, heart-shaped marking at the center of the dorsum; and (c) two

spots in the anal region. Flanks dark brown, with a sharp dorsolateral color border. Hind-

limbs yellowish-brown with indistinct darker crossbands. Forelimbs cream to beige. Color of

limbs extending as mostly rather small flank blotches on the flanks. No flashmarks; postero-

dorsal part of femur and knee hollow orange, but without contrast to the surrounding

surface. Upper part of iris light. — (3) Venrral color and pattern: Forelimb, femur, venter and

throat black with many small rounded whitish blue spots. On the throat, these whitish blue

spots sometimes are arranged semicircularly along the lip, but they are not fused (not forming

a closed horseshoe marking). Foot, tarsus and tibia orange-red. This color sometimes

extending onto the distal part of the femur.

Mantella nigricans Guibé, 1978

M{antella] cowani nigricans Guibé, 1978. — Name-bearing type: lectotype, by present designation, MNHN

1973.55$, female, SVL 26.3 mm. - Tÿpe locality: “massif du Marojezy”, according to original description.

— Other types: paralectotypes, following present lectotype désignation, MNHN 1973.517, MNHN

1973.530-54, and MNHN 1973.556-9. - Erymology: derived from Latin nigricare (to darken towards black);

referring to the uniformly dark color of the type series which, however, was most probably caused by

fixation in formalin.

Mantella madagascariensis nigricans: BUSSE, 1981 (part. not MNHN 9594); BLOMMERS-SCHLÜSSER &

BLANC, 1991; GLAW & VENCES, 1994 (nomen dubium: p. 412).

15: VENCES & KNIEL, 1998; STANISZEWSKI, 1998

Mantella “negristara” [conditional name]: LARSEN, 1997.

Mantella new species: SraniszEwskt, 19974 (fig. p. 11 and 16).

Mantella sp.: VENCES et al., 1998.

Other chresonyms:

Mantella madagascariensis: GLAW & VENCES, 1994 (part.).

Mantella madagascariensis sensu stricto: GLAW & VENCES, 19924 (part.; locality Marojezy).

Identity. — The name was erected by GUIBÉ (1978: 84) as the subspecies Mantella cowani

nigricans. No types were designated. The original description was véry short and superficial:

“Parfois, au contraire, les taches claires de la racine des membres se réduisent considérable-

ment et finissent par disparaître, le corps et les pattes sont alors uniformément noirs. De tels

individus mélaniques se rencontrent en particulier dans le massif du Marojezy, ils correspon-

dent à une sous-espèce: M. cowani nigricans n. Subsp.”.

BUSsE (1981) and BLOMMERS-SCHLÔSSER & BLANC (1991: 274) accepted the subspecies in

a preliminary way. BUssE (1981), however, doubted the locality Marojezy and assigned

MNHN vouchers from Betafo and Ambatodradama to nigricans (these specimens, however,

belong to M. cowani; see above).

During examination of Mantella voucher specimens in the MNHN we noted that all

specimens from Marojezy are identified as Mantella cowani nigricans in the catalogue. The

whole series w talogued in 1973, while the batrachological MNHN collection was curated

Source : MNHN, Paris

VENCES, GLAW & BÔHME 31

by Jean Guisé. In all these specimens, the light color pattern is largely faded, very probably

due to a previous formalin fixation, giving the impression of melanistic specimens. There is

little doubt that GUIBÉ's description was based on these specimens, which must therefore be

considered as syntypes.

Unfortunately, the syntype series is not homogeneous; it contains some specimens of M.

laevigata as well as a rather large sample of specimens of a M. cowani group species which

differs from all other members of the group (see below). In order to reach stability of the

name, we here designate one of these specimens as lectotype. This avoids the necessity of

creating a new name for the Marojezy populations belonging to the M. cowani group.

Mantella nigricans belongs to the M. cowani group based on: (1) single click calls (GLAW,

personal observation); (2) unforked sternum (VENCES et al., 19994); (3) lack of horseshoe

marking; (4) rounded and isolated ventral spots; (5) lack of a frenal stripe; (6) presence of

flank blotches. It differs from all other members of the group by lacking red ventral color on

the hindlimbs. Furthermore, it differs from M. cowani and M. baroni by smaller ventral spots

and a different dorsal extension of light (green) color, and from M. haraldmeieri by a different

dorsal coloration. M. nigricans is most similar by dorsal coloration to some specimens of M.

aff. baroni.

Comment. — Of the paralectotypes, only the specimens listed in the Material examined section

are conspecific with the lectotype; see section of M. laevigata for the remaining specimens.

Description of lectotype. - MNHN 1973.555, female specimen with nearly mature oocytes.

Specimen in good state of preservation with a longitudinal central cut along the venter.

Stomach and intestine removed for content analysis and stored separately in small tubes. For

measurements, see tab. 2. Body slender; head not broader than body; snout slightly pointed in

dorsal, rounded in lateral view: nostrils directed laterally, not protuberant, nearer to tip of

snout than to eye; canthus rostralis weak, straight; loreal region even; tympanum rather

indistinct, medium-sized, rounded, its diameter about half of eye diameter; supratympanic

fold weakly developed; tongue longish to ovoid, slightly bifid posteriorly; maxillary and

vomerine teeth absent; choanae small, rounded. Arms slender; subarticular tubercles single;

outer metacarpal tubercle rounded, inner metacarpal tubercle rounded, both rather distinct

and of similar size; fingers without webbing; finger length 1<2<4<3, finger 4 distinctly longer

than 2; finger 2 only slightly longer than l; faintly developed but distinct terminal finger disks.

Legs moderately robust; tibiotarsal articulation reaching posterior eye margin; feet with

small, slightly elliptical inner and rounded outer metatarsal tubercles; subarticular tubercles

single, rounded; toe disks faintly developed but distinct. Foot without webbing. Lateral

metatarsalia connected; toe length 1<2<3<5<4, toe 3 distinctly longer than 5. Skin on the

upper surface smooth: ventral surface smooth, except for granular thigh patches (“femoral

glands”) extending from the anus ca. 6 mm distally (max. width 3.6 mm). Color in life

unknown; in preservative almost uniformly dark brown, with very little pattern contrast

(probably due to formalin fixation). Contours of moderately large light flank blotches faintly

recognizable. Venter and ventral side of fore- and hindlimbs, including humerus, fibula,

femur, tibia, tarsus and foot, uniformly dark with small rounded light spots. Six spots

positioned on the throat along the lip, but not fused to form a horseshoe-marking. No spots

in the breast area. No flashmarks.

Source : MNHN, Paris

32 ALYTES 17 (1-2)

Material examined. - MNHN 1973.555 (Marojezy, 600 m; lectotype): MNHN 1973.541 (Marojezy, 300

m; paralectotype); MNHN 1973.530-3, MNHN 1973.550-4, MNHN 1973.56, MNHN 1973.559

(Marojezy, 600 m; paralectotypes): ZFMK 59887-8, ZFMK 59902 (Marojezy Camp 3: leg. F. GLAwW & O.

RAMILISON 11.199); MRSN A1822 (Tsararano Chain, Camp 2; leg, F. ANDRFONE 10.XI1.1996); MRSN

A1823.1-4, MRSN A1824.1-2, MRSN A1825 (Tsararano Chain, Camp l; leg. F. ANDREONE

XI-XI1.1996); MRSN A1829.1-8 (Analabe/Anjanaharibe, Camp 2: leg, F. ANDREONE II.1996); MRSN

A1830 (Analabe/Anjanaharibe, near Camp 2 at about 1200 m; leg. F. ANDREONE II.1996).

Distribution. — Known from: [1*] the Marojezy massif (North-Eastern region; 300-700 m

altitude); [2] Hiaraka (Jaraka) (Masoala peninsula; A. PEYRIERAS, personal communication);

[3] Tsararano (700 m altitude); [4] Anjanaharibe (1200 m altitude). See the discussion in the

section on distribution of M. baroni.

Diagnosis. — (1) Morphology: À medium sized to large, relatively stout Mantella. SVL 27-

28 mm. TTA reaching the forelimb insertion or the tympanum. Terminal disks of fingers and

toes rather largely expanded. Tympanum/eye ratio generally sightly below 3/5. IMT medium-

sized (ratio width/length 2/3 to 4/5). — (2) Dorsal color and pattern: Relatively variable. Some

specimens similar to M. pulchra (see below). Flanks black, with a sharp dorsolateral color

border. Limbs brown, except humerus and proximal femur; these light green to yellowish

green, this color extending as relatively large flank blotches onto the flanks. In other

specimens the green color making up the major part of the dorsal surface, including dorsum

and flanks (in one specimen the anterior two thirds of the dorsal surface were green). In these

cases, however, a strong dorsolateral color border remains on the head. No sharply delimited

rostral stripes and no flashmarks. Iris with light pigment in its upper part. — (3) Ventral color

and pattern: Black with small, rounded blue spots. On the throat these spots sometimes

arranged semicircularly along the lip, but only exceptionally fusing to form a closed horseshoe

marking.

Mantella bernhardi group

Mantella bernhardi Vences, Glaw, Peyrieras, Bôhme & Busse, 1994

Mantella bernhardi Vences, Glaw, Peyrieras, Bôhme & Busse, 1994. - Name-bearing type: holotype by original

désignation (VENCES et al., 1994: 391), ZFMK 57164, male, SVL 19.0 mm. - Type locality: *Regenwald nahe

Tolongoina, Provinz Fianarantsoa”, according to the original description. - Other types: none. -

logy: named after the German zoologist Bernhard M1ER

Maniella bernhardi: GLAW & VENCES, 1994; CarisstMi-PriORt, 1995 (fig. p. 43); STANISZEWSKI, 1996

(plate p. 26), 1997a (fig.). 1997 (fig. p. 21, 40-41, 60 above and middle: probably not fig. p. 37 and

60 below), 1998a (fig.); VENCES et al, 1998.

Material examined. - ZFMK 57164(S-Mad.: E-Betsileo [forest near Tolongoina fide PEYRIERAS]; ded. M.

VENCES, 11.194, leg. A. PEYRIERAS; holotype): ZFMK 59820-1 (near Tolongoina: leg. PEYRIERAS, ded. F.

GLAW IV.1995); ZFMK 62697-8 (locality unknown; CS); ZFMK 62699-707 (locality unknown; TE).

MRSN A1964 (Ambohimana next to Tolongoina: leg. F. ANDREONE 20.VIL.1995).

Distribution. — Until now, the species is only known from the type locality: [1] forest near

Tolongoina. This locality is corroborated by the observation of F. ANDREONE (personal

communication) who, however, found only one single specimen in the dry season.

Source : MNHN, Paris

Table 2. - Morphometric measurements of Mantella type specimens, and of a reference specimen of M. milowympanum (ZFMK 65626). Stat., Status;

HT, holotype; PT, paratype; LT, lectotype; PLT, paralectotype; TOT, topotype; M, male; F, female; TT, point that is reached by the tibiotarsal

articulation when limbs are adpressed along the body: 1, forelimb insertion; 2, nearly to tympanum; 3, tympanum; 4, between tympanum and eye;

5, posterior eye margin; 6, center of eye. See Materials and methods section for abbreviations of measurements. Most specimens could not be

reliably sexed, generally due to bad state of preservation. Measurements of inner metatarsal tubercle and disk width on third finger were only

taken from few equally well fixed specimens. Other lacking measurements are due to damage or bad preservation of the respective specimens.

Eye-

Collection number | Sex | Stat. | SVL | HW HL Eye | Tym Ns Ns-St | ForL | HaL | HiL | FoTL | FoL | ToLi | DW3 | FW3 | IMTL |IMTB}IMTH| TT

Wantella bersileo

MNHN 1895278 irfwoleilsz|arlialiolialueleo[#ilus]s7[ifos#|o#lfos» G

MNHN 1895.279 PLT | 157 10.4 25.3 6

Mantella attemsi (syn. Mantella betsileo) £

NMW 20837 F I LT |25.6 | 72 9.2 2.6 15 2.1 13 163 7.1 34.7 | 16.5 | 11.4 17 07 04 0.8 04 03 5 &

zmBiesss | M frirair| 64 [94 | 24 [7 | 20 [11 [ue] 64 [56148] 97 6 El

Mantella viridis ©

ZFMK 47900 F HT | 303 8.9 114 2.8 2.0 23 20 17.7 | 8.0 | 39.5 | 18.8 | 12.8 17 0.95 | 0.68 | 1.05 | 0.75 l CL

Mantella expectara £

ZFMK 53540 HT | 234 | 77 9.5 29 LS 2.0 15 15.8 | 6.7 | 34.8 | 17.1 | 11.4 11 0.78 | 0.43 | 0.85 | 0.50 | 0.30 5 Le

ZEMK 53541 Pr [2068 | 56/2414 [19/14 [14] 63 [as |ico[ios | 15 [075 [045 [075 [053 [035] 6 œ

ZFMK 53542 PT | 220 | 68 8.6 24 1.4 18 14 144 | 63 34.8 | 16.0 | 109 15 6 S

ZFMK 59095 Pr [use io 26 fisc fie liss| 74 [aceliro| na] 14 [ow|oclos|os]|02] 6 Ë

[zu 50096 Prfas(ei(os]2sfur|2o us (is2] 70 [ssifiés[ui (ii [os [os [os/[06|030] 5

IK 59097 PT | 234 LE 94 27 L6 17 L6 16.6 | 64 33.8 | 16.4 | 10.7 12 6

ZFMK 59098 PT |23.6 | 72 89 28 1.6 17 12 154 74 34.5 | 16.6 | 10.8 14 6

Mantella lacvigata

TM 10074 HT[25]6190[24[12[17l16/149/68 [321155 96/[18[11/04[07/03/03] 6

TM 10085 PT ERIKA 92 25: 12 1.6 14 17.3 82 38.2 | 17.6 | 10.4 E 11 0.4 1.0 0.4 0.4 $

TM 10086 fois fs os fe fs 53 Pi fes fa [ses Pics Pros Pis [2 Pos Fos [04 Pos [5 |

TM 10087 Pr [ailes flioof2efisfur is [iéo] 75 [368 [i6s [100 [| 20 | 08 | 03 | 10 [04 [os | 6

TM 10088 Pr [2462 |s6|24/iifir|iifiols7|[a6|iss] ss 6

TM 10090 PT 17.4 53. 78 20 09 15 10 11.5 | 46 | 264 | 108 | 69 5

Mantella baron

BMNH 19472719 M [HT [22/73 [io9 | 28 [16 | 22 [ir ii 71 [361172

E

Source : MNHN, Paris:

Collection number | Sex | Stat. | SVL | HW | HL | Eye | Tym à Ns-St | ForL | HaL | HiL | FoTL| FoL | ToLi | DW3 | FW3 [MTL |IMTB|mMTH| TT

Phrynomantis maculatus (syn. Mantella baroni)

MNHN 1991.2845 LT [270] 75 [109[ 29 [15 [ 20 [ 16 [177] 77 [404 [186/[124] 17 [06 [046 [08 CE 3

MNHN 1991.2846 PLT | 262 | 7.6 | 10.1 32 1.5 2.1 1.6 | 16.9 | 7.0 | 39.4 | 17.2 | 11.0 | 1.9 | 0.57 | 0.43 | 0.86 0.44 5

MNEN 1991.2847 Prfmo|sifoo [31132116 |icol 75 [54/83 [u7|is 5

MNEN 6307 _| F [Pir|284] 80 [102] 29 | 19 | 22 | 18 [172[ 73 [391 [175[n2|14 [oafosælfii o51| 3

Mantella cowani

BMNH 1947.2.7.4 LT | 282 | 75 10.1 28 13 19 16 | 16.5 | 7.3 | 35.8 | 17.4 | 11.5 | 1.6 06 0.5 10 04 04 12

BMNH 1947275 PLT [275] 75 [103] 27 [14 | 20 [ 16 [i67[ 71 [374[182[123[ 18 [07 [o6 [11 [04 [03 [12 |

Mantella haraldmeïeri Il

ZFMK25351 | M [ur [20/68 [86/24 [14/18] 14 [144[ 63 [58/[160[100[ 15 [os [04/[08/]03%/[0%] 6

ZEMK21805 | F | PT |268 | 75 [108 | 26 | 19 | 20 | 15 [162] 68 | 372|167 [109 | 17 3

ZFMK 21806 | M | PT [231] 70 25 [13/18] 14/158|60/|354[159 [106] 14 [os [05 [058 [04/02] 6

ZFMK 21807 M PT _|214 | 7.2 84 24 14 2.1 13 14.0 | 6.2 | 33.4 | 16.4 | 10.1 15 6

ZFMK 25352 PT [27/70/87] 25]14/|19|13 [145 w3[160[ 103 15 6

ZFMR 25353 ToT| 20] 70 [82 [24 [14 [19 | 15 [157] 65 [355 [163 | 103] 14 G

Mantella nigricans

MNEN 1973555 | F [LT [263[ 75 [104[ 28 [16 [19 [17 [171] 73 [#1[i88[123[ 19 [10 [05 [09 [05 [05] 5

1 Mantella bernhardi

ZFMK 57164 M HT 19.0 | 59 8.0 2.2 LA 17 12 [12.7] 5.1 | 29.1 | 13.5 | 8.6 14 | 0.50 | 0.33 | 0.65 | 0.53 | 0.23 5

Mantella madagascariensis

MNHN 1895.276 LT |218] 68 84 23 13 EE 17

IE INHN 1895.277 PLT | 17.9 22 12 13.8 | 89

Mantella loppei(syn. Mantella madagascariensis)

MNEN 1935-16 PT [286] 84 [iwo7[ 31 [18 [19 [18 [163] 68 [6/[187[126[ 15 [07%6[06[130[095[08] 2

Mantella pulchra

BMNH 1947.2.7.20 HT [247] 75 [103] 24 [ 14 [19 [ 18 [i58] 66 [wo [i62[109[ 18 [07 [05 [14 [06 [06 | à

BMNH 1947.2.7.27 101] 247] 69 | 96 | 25 | 14 | 16 | 16 [153 | 66 | 548 [166 [108 | 19 | 04 [03 [13 Joss |o6 ji

IBMNH 1947.2.7.28 TOT | 260 | 7.4 | 10.7 | 3.0 15 18 16 | 15.2 | 6.0 | 33.8 | 16.5 | 11.0 | 1.7 05 04 15 07 0.3 12

BMNH 1947.27 29] TOT| 243] 71 | 92 | 23 | 17 | 20 | 16 [145] 59 | 527|156| 08 | 15 | 07 | 05 | 13 | 05 | 03 | 3

BMNH 1947.2.7.30| Tor] 250] 74 | 99 [26 [15 | 18 | 13 [157] 61 [335 [156/ 103] 14 |045 [035 | 16 | o7 | 05 [12

BMNH 19472731 Tor] 219] 72 | 89 | 25 | 14 | 18 | 16 [143] 59 | 519 [155 [100] 15 | 05 | 04 [12 [07 | 06 | 3

ZMB 50105 OT|231|68/91/19 [13/21] 13136] 52 | 320 | 162

ZMB 30576 Tor] 245] 79 | 100] 27 | 17 | 18 | 13 [151] 67 |351[1i70/117] 14 [06 [045 [12804504 | 1

MNHN 19912843 TOT | 23.9 | 7.7 | 104 | 28 16 24 17 15.3 | 6.5 | 37.2 | 16.7 | 114 1.5 | 0.55 | 0.45 | 1.50 0.50 3

MNEN 1928.106 TOT | 248 | 73 [101[ 32 | 14 | 19 | 18 [148 | 63 | 332 [160 [104 | 14 3

LA

(T-1) LI SALATV

Source : MNHN, Paris

Es

collection number | Sex | sue [svL | nw | Hi Eye | Tym A Ns-St| ForL | HaL | Hil |FoTL| FoL | ToLi | DW3

Mantella aurantaca

MNHN 1899412 | M? | LT [212 | 60 19 Ti 55 [324 [154 [102 5

MNHN 1899413 | F |PLT]208 | 57 19 [12 55 [310] 142] 94

Mantella aurantiaca rubra (syn. Mantella aurantiaca)

ZFMK 68868 FIiT[246[78[06]726 [16/20/16 [ia] 56 [as [152[ 959 [15 [06 [os [os [06 [05] 3

Mantella crocea

ZFMK 45007 _] F [ur [225] 69 | 94 [ 27 [13 [16] 14 [135] 56 [307[ 144] 97 | 15 [058 [040 [06053 [0407] 3

K 45008 Pr [193 | 58 | 80 | 21 | 13 | 12 | 00 [i22| 51 [276129 | 79 | 14 5

ZFMRK 50173 PT [220] 60 | 82 | 20 | 14 11] 54 [311/142|] 96 | 14 3

ZFMK 50174 Prfnwofsaf77fisfuifis|izfuo] as |23ofi2o) 77 | 13 5 <

ZFMK 50175 PT [202] 62 | 87] 22 | 13 | 18 | 10 [us] 49 | 294 [134 | 88 | 15 5 2

ZFMK 50176 PT 211) 62 | 83 | 22 | 14 | 16 | 12 [129 | 54 [297142] 90 | 14 3 Q

ZFMK 50177 PT [w9|57]78|20 [11] 16 | 10 [133] 56 | 24 [142] 89 | 12 üi 5 #

ZFMK 50178 PT [28/62 | 82 [19 [1219 19] 52 [314 [144] 90 | 17 3 Q

ZFMK 50179 PT |207/61]79]21/[ 11/1612 [127] 52 |297| 138 | 90 | 13 4 >

ZFMK 50180 PT [204] 59 | 80 | 22 | 14 [ 14 [11 [i52/ 55 [275 [138] 89 | 13 4 £

ZFMK 0181 PT [195] 54 | 74 | 18 | 13 [16 118] 51 [290128] 83 | 10 G &

ZFMR 50182 PE [AS |52/220 [17 | 12 111] 48 | 263 [123] 90 | 12 5 w

ZFMK 30183 Prfiw3|s6|75|22/13|15/1o fus] 46]20/129] 87 [12 4 É

É ZFMK 50184 Pr [iss|s3 | 72/18 | 12/16 | 09 [116] 54 [258 [150 85 | 15 5 ë

ZFMRK 50185 PT | 203 130 290 [130] 88

ZFMR 50186 PT [196] 55 | 85 | 20 [ 12 | 13 | 12 [136 | 49 [254135] 90 | 15

ZFMRK 30552 PT [209 | 60 | 84 | 22 | 13 2/55 [307[148) 03 [15 3

ZFMR 50553 PT [208 | 60 | 79 | 22 | 13 | 15 | 12 [137] 64 | 307 | 142] 88 | 14 ü

ZFMK 50721 PT|20|67|s8i]2s [sis ]i2 |134] 58 [228145] 93 | 16 3

ZFMK 30722 PT [24/62 | 78 | 20 | 11 af 56 [4/51] 98 | 12 | 048 | 033 [068 | 038 [030 | 3

[ZFMK 50723 PT [14] 5168 | 18 | 11 | 12 | 11 | 9.6 | 40 | 240] 111] 70 | 09 5

ZFMRK 50724 PT [29 ) 64 | 95 | 24 | 14 | 17 | 12 [145] 55 | 309 | 147] 97 | 17 [045 [030 | 055 [045 [025] 3

ZFMK 50725 PT [24/61/85 |23/]12{14/[11f103]52[2%6[1m58/92{|1 5

Mantella milon mpanum

ZFMK 65626 M] -1]25]64[96[27[17[16]13]144]63]34[147[101[] 16 [05 [03] 08 [05105] 3

w

a

Source : MNHN, Paris:

36 ALYTES 17 (1-2)

Diagnosis. — (1) Morphology: The smallest known Mantella. SVL 19-22 mm, males 19 mm,

females 19-22 mm. TTA reaching the posterior eye margin or the eye center. Terminal disks of

fingers and toes slightly expanded. Tympanum/eye ratio generally 1/2 to 3/5. IMT small (ratio

width/length more than 4/5). — (2) Dorsal color and pattern: Dorsum and head dark grey or

brown. A fine light middorsal line sometimes present. Flanks black. Poorly contrasted

dorsolateral color border. No frenal stripe. Humerus yellowish beige, femur bright yellow, this

color extending slightly onto the flanks as small flank blotches. Fibula and hands, as well as

tibia and feet, brown with generally only one distinct dark crossband, respectively. No

flashmarks, but posterodorsal part of femur and knee hollow orange as ventral surface of

hindlimb. Iris with light pigment in its upper part. — (3) Ventral color and pattern: Venter,

throat and forelimbs black with few large whitish blue markings which can be irregularly

vermiculated, but always with very distinct, largely rounded borders. Throat with a distinct

horseshoe marking, often covering most of the throat surface in males, being smaller and

sometimes not continuous in females. Tibia and femur orange. Foot and tarsus are also

orange, but this color is mostly covered by irregular dark pigment.

Mantella madagascariensis group

Mantella madagascariensis (Grandidier, 1872)

Dendrobates madagascariensis Grandidier, 1872. - Name-bearing type: lectotype, by designation of GLAW &

Vexcrs (1994: 403), MNHN 1895.276, sex unknown due 10 bad preservation, SVL 21.8 mm. - Type locality:

“Forêt d’Ambalavatou, entre Mananzarine et Fianarantsoua” according Lo the original description, given

as “forêt d'Ambalavato, entre Mananjary et Fianarantsoa” by BLOMMERS-SCHLÔSSER & BLANC (1991). —

Other types: paralectotype, following lectotype designation of GLAW & VENCES (1994), MNHN 1895.277.

— Etymology: named after its geographic origin, Madagascar.

Mantella madagascariensis: Were, 1901; MOcqQUARD, 1909; BOULENGER, 1882; BUSSE, 1981 (part.):

Pinrak, 1990 (part.); BLOMMERS-SCHLOSSER & BLANC, 1991 (part.): GLAW & VENCES, 19924

(part.; see localities), 1994 (part.; see localities); HERRMANN, 1993 (fig.); STANISZEWSKI, 1997a (fig.

p. 12): LaRsEN, 1997 (fig.); VENCES & KNIEL, 1998.

Mantella m. madagascariensis: ME1ER, 1986 (fig. 5).

Mantella loppei Roux, 1935. - Name-bearing type: holotype as inferred from original description (Roux, 1935:

441; see comment below), NMB 4849, female (number and sex according to FORCART, 1946). = Tÿpe

locality: “Moroulambo, province de Vatomandry”, according to the original description. - Other types: two

paratypes according to original description, one corresponding to MNHN 1935.416, and the second

specimen probably stored in the La Rochelle Museum (see comment below). - Etymology: named after E.

Loppf, former director of the La Rochelle Museum

Mantella loppei. FORCART (1946); BUSSE, 1981 (syn. madagascariensis):. BLOMMERS-SCHLÔSSER &

BLANC, 1991 (syn. madagascariensis).

Mantella “loppeï”": GLAW & VENCES, 1994; STANISZEWSKI, 1997 (fig. p. 57).

“Mantella nasuta Sp.” (nomen nudum, referring to the “variable” color morph}: CLARK, 1994 (fig. p. 10 above

and p. 11 above).

Mantell sp [refèrring Lo the *variable* color morphl: CLARK, 1994 (fig p. 1 below); VENCE et al, 1994 (lg

LAW & VENCES, 1994 (plates 58-60)

cal pate tos * [conditional name, referring to the “variable” color morph]: BarTLerr, 1995 (fig. p. 18).

Other chresonyms:

Mantella pulchra: Guié, 1964, 1978 (part.).

Mantella cowani: WoLperT & MÜLLER, 1980

Mantella crocea: BARTLETT, 1995 (fig. p. 16 below) [referring to the “variable” color morph]

Identity. - Dorsal color patterns of this species are sometimes very similar to M. baroni, and

single specimens can only be identified by combination of several color characters. The

Source : MNHN, Paris

VENCES, GLAW & BÜHME 37

syntopic occurrence of M. baroni and M. madagascariensis as recorded by us in Vohiparara,

however, demonstrates that both must be regarded as separate species. As far as can be

concluded from large series of specimens exported from Madagascar in the pet trade, the

pattern is constant at some localities but may be extremely variable elsewhere.

M. madagascariensis was considered as “nomen dubium”” by several authors based on

the very bad state of preservation of the types and the short and little detailed original

description (GuIBÉ, 1964; DaLy et al., 1996). A detailed examination of the lectotype,

however, revealed one character which is still recognizable and can be used for a diagnosis. The

specimen’s dorsal and ventral color has nearly completely faded to uniform brownish. The

hindlimbs are separated from the body. The posteroventral part of the femur and the distal

part of the tibia, in the knee hollow area, still show some contrasting pattern with an

extension corresponding exactly to the flashmarks present in all specimens of the form here

attributed to M. madagascariensis (see fig. 11). Ventrally, the lectotype shows light color

extending onto the distal part of the femur, corresponding to the pattern generally present in

the form here attributed to M. madagascariensis but not in the otherwise rather similar M.

pulchra (fig. 12).

Comments. — (1) The paralectotype of M. madagascariensis is most probably a subadult, but

it may also be a M. bernhardi and thus not conspecific with the lectotype. — (2) Status of two

names coined in recent publications to refer to “variable color morphs” must be discussed

here. “Mantella mysteriosa” was used in quotation marks by BARTLETT (1995). The author

states explicitely (p. 20) that this name originated from a pet dealer’s list. Diagnosis, type

designation and type locality were not given. The name must thus be seen as documentation

of the usage of a conditional name in the pet trade, and is not nomenclaturally available.

“Mantella nasuta sp.” was used by CLARK (1994) in the captions of two figures. No unequi-

vocal diagnosis of the specimens figured is possible since neither dorsal pattern of hindlegs

nor ventral coloration were documented or described. Further diagnosis, type designation

and type locality were not given. No direct reference to the name is to be found in CLARK'S

(1994) text and key. Two common names, Mimic Mantella and Panther Mantella, are used in

the captions of the figures on p. 10-11 to refer to specimens named Mantella nasuta sp. Both

common names were also included in CLARK’S (1994) species list as “Mantella sp. A” and

“Mantella n. sp. 5”. The latter two names, on the other hand, are also found in his key. Thus,

two forms considered as different species are indirectly keyed as M. nasuta sp., and there is no

direct diagnosis related to this name, which we consider as a nomen nudum. — (3) Mantella

loppei, according to the original description (Roux, 1935), was based on “3 Amphibiens

appartenant au genre Mantella et qui représentent une espèce nouvelle … M. le Docteur Et.

Loppé a bien voulu nous autoriser à conserver pour le Musée de Bâle le spécimen-type de

l'espèce, tandis que deux autres exemplaires se trouvent au Musée de La Rochelle.” Although

not explicitely mentioned, this infers the existence of a holotype in the collection of the Basel

Museum (NMB 4849 according to FORCART, 1946), and two additional specimens which we

consider as paratypes (originally both in the La Rochelle Museum; one later exchanged with

the Paris Museum, catalogued as MNHN 1935.416).

Material examined. —- NMB 4849 (Prov. Vatomandry, Mouroulambo, coll. E. PICHON 1930); MNHN

1895.276-7 (Ambalavato: lectotype and paralectotype);: MNHN 1931.12 (Moramanga), MNHN 1931.13

(Moramanga?); MNHN 1935.416 (Vatomandry: J. Roux: “don du DR. Loprr, Conservateur du Musee

de la Rochelle”; paratype of M. loppeir; MNHN 1992.4821-2, MNHN 1993.1440, MNHN 1993.1445

Source : MNHN, Paris

38 ALYTES 17 (1-2)

(origine inconnue); ZFMK 14184-207 (Niagarakely; leg. H. Mrier 1972; 14186, 14188, 14196: CS);

ZFMK 14209-13 (Niagarakely: leg. H. Meter 1972); ZFMK 14325-30 (Niagarakely: leg. H. MEIER 1974);

ZFMK 14471-5 (Niagarakely; leg. H. Meter 1973); ZFMK 22107-12 (Niagarakely: leg, H. Mer 1973);

ZFMK 56153-4 (pet trade; ded. F. GLaw XI.1993); ZFMK 60132 (locality unknown, ded. F. GLAW

IV.1995; ZFMK 62740-1 (locality unknown; CS); ZFMK 62732-6, ZFMK 62738-9 (locality unknown;

TE); ZFMK 62737 (locality unknown; TE; pattern very similar to M. baroni); ZFMK 64138 (Vohiparara;

leg. F. GLAW, D. RAKOTOMALALA & F. RANAIVOYAONA III.1996; TE).

Distribution. - Type locality is [1] Ambalavato near Ranomafana. Type locality of the junior

synonym M. loppei is [2] Marolambo (Vatomandry). ZFMK vouchers were collected at [3]

Niagarakely. At [4*] Vohiparara (ca. 1000 m altitude; near Ranomafana), we found one

specimen syntopic with M. baroni. According to A. PEYRIERAS (personal communication),

populations of the “variable morph”, here included in M. madagascariensis, occur near [5]

Beparasy. See the discussion in the section on the distribution of M. baroni.

Diagnosis. — (1) Morphology: À medium-sized Mantella. Compared with M. baroni, general

body shape rather stout. SVL 20-27 mm, recorded lengths of males 21-22 mm, of females

24-25 mm. TTA rarely reaching the eye center, sometimes the posterior eye margin, mostly the

tympanum, and sometimes only the forelimb insertion. Terminal disks of fingers and toes

slightly expanded. Tympanum/eye ratio generally 1/2 to 3/5. IMT large (ratio width/length

less than 3/5). — (2) Dorsal color and pattern: Upper head surface, dorsum and flanks black,

generally without recognizable dorsolateral color border. Yellowish rostral stripe present.

Femur and humerus yellow to green, this color extending as large flank blotches onto the

flanks and sometimes onto the dorsum. Distinct orange flashmarks present. Tibia, tarsus and

foot orange with or without blackish crossbands and marblings. Iris mostly containing light

pigment in its upper part. Rostral stripe often in contact with flank blotch. In specimens of the

“variable morph”, yellow color in varying extension can sometimes be present on the dorsum.

AI intermediate states, from a few yellow spots to a reticulated yellow marbling or a dense

yellow speckling, are known. A greenish frenal stripe, often interrupted, can be present as

well. Specimens without reliable locality information are known which are nearly uniformly

yellow dorsally and ventrally, with only a few blackish spots and marblings. In these speci-

mens, the more distinct yellow surface in the flank blotch area is reminiscent of the typical

coloration, but it is not clear whether they really are conspecific with M. madagascariensis. —

(3) Ventral color and pattern: Venter, throat and forelimbs black with light markings (mostly

whitish-blue, sometimes yellow to green), these being generally rather large, rounded, and

situated posteriorly on the venter. Distinct horseshoe marking present, more extended in

males. Femur, tibia (except flashmark area), tarsus and foot often uniformly orange, in other

specimens with areas of black and yellow (the latter corresponding to yellow color on the

dorsal surface). Areas of femoral “glands” often darkly pigmented. In some specimens, femur

nearly totally black with blue spots. In “variable” specimens, typical ventral pattern some-

times replaced by a dense yellow marbling.

Mantella pulchra Parker, 1925

Mantella pulchra Parker. — Name-bearing type: holotype by monotypy, BMNH 1947.2.7.20 (ex

1925.7.2.58), female according to original description, SVL 24.7 mm. — Type locality: “Antsihanaka”,

according to original description. - Other types: none (see comment below). - Etymology: derived from

Latin pulcher (beautiful).

Source : MNHN, Paris

VENCES, GLAW & BÔHME 39

Mantella pulchra: Guwé, 1964, 1978 (part.}; GLAW & VENCES, 1994; HexKEL & ScHmipr, 1995 (fig.

p.56): BartLerr, 1995 (fig. p. 24 above left); van TuuL, 1995; CaissimI-PRIORI, 1995 (fig. p. 43):

SraniszEwskI, 1996 (pl. p. 23), 1997b (fig.), 1998a (fig.): DALY et al., 1996; LARSEN, 1997; VENCES

& KNIEL, 1998.

Other chresonyms:

Mantella madagascariensis: DALY et al., 1984; BLOMMERS-SCHLÔSSER & BLANC, 1991 (part.).

Mantella madagascariensis madagascariensis: BUSS, 1981 (part.).

Maniella sp. cf. madagascariensis: GARRAFFO et al., 1993.

Mantella madagascariensis (colour morph Mantella “pulchra”): GLAw & VENCES, 19924.

Mantella cowani pulchra: ANDREONE, 1992; GAvErrI & ANDREONE, 1993 (p. 105).

Mantella cowani: ZIRGENHAGEN, 1981: LE BERRE, 1993 (inner fig. on p. 21).

Mantella sp: SrANISzEwSkI, 1998a (fig.).

Comments. —(1) According to the BMNH catalogue, there were 22 “paratypes” (old numbers

BMNH 1925.7.2.59-80), one of which (ex BMNH 1925.7.2.80) was cleared and stained and

seemingly not given a new number when the types were re-numbered in 1947. Nine additional

specimens were exchanged according to this catalogue. Seven of these were located by us:

MNHN 1928.106, MNHN 1991.2843 (ex MNHN 1928.106.A), ZMA 5809-10 (according to

VAN TuuL, 1995), ZMB 50105, ZMB 30576, MZUT An.108 (all from Antsihanaka). Accord-

ing to vAN TuuL (1995), “paratype(s)” were also deposited in the MCZ collection. — (2) The

original description of M. pulchra was based on a single specimen (“Type specimen: a female

from Antsihanaka”; PARKER, 1925: 394), and contains no mention of other specimens.

Although the specimens listed above have similar collecting dates as the holotype, they can

therefore not be considered as paratypes. As already stated by GAVETTI & ANDREONE (1993),

they must be regarded as topotypes only.

Material examined. - BMNH 1947.2.7.20-32 (holotype and paratypes; all from Antsihanaka: coll. or

purch. ROsENBERG); MNHN 1928.106, MNHN 1991.2843 (“acquis par échange avec le British Mus.

[Nat. History] en 1927”; paratypes): MNHN 1993.1443 (locality unknown); ZFMK 52122-3 (locality

unknown; ded. D. KaRBE 1991); ZFMK 56155 (locality unknown: ded. F. GLaw XI.1995; CS); ZFMK

62258 (An'Ala bei Andasibe; leg. F. GLAW 3.I1.1996); ZFMK 62742-4 (locality unknown: CS); ZFMK

62645-59 (locality unknown; TE); ZMB 50105, ZMB 30576 (Antsihanaka; exchanged with BMNH in

111.1927; paratypes); MRSN A0059.1-4 (An’Ala [syntopic with M. baronif: leg. F. ANDREONE 4.1.1992

{sacrificed 14.X1.1992]); MRSN A444.1-3 (locality unknown); TM 9893, TM 9897, and possibly the

juvenile TM 9901 (Folohy; coll. METHUEN).

Distribution. — Type locality is [1] Antsihanaka. ANDREONE (1992) and DaLy et al. (1996)

collected the species near [2*] An’Ala (near Andasibe; ca. 850-1000 m altitude), and A.

PEYRIERAS (personal communication) in [3] Andekaleka (Rogez). Further localities within the

[4] Mananara reserve (ca. 100-200 m altitude) were published by DALY et al. (1996). Speci-

mens in the TM corroborate the occurrence in [5] Folohy. Exact location of the type locality

Antsihanaka is unknown; most probably, it was used in the past for a forested region near

Lake Alaotra (see VIETTE, 1991). BLOMMERS-SCHLÔSSER & BLANC (1991: map 4) locate

Antsihanaka, probably erroneously, east of Andasibe.

Diagnosis. — (1) Morphology: À medium-sized Mantella. General body shape rather stout.

SVL 21-25 mm, recorded length of males 22-23 mm. TTA often reaching the posterior eye

margin, sometimes the tympanum or the forelimb insertion. Terminal disks of fingers and

toes slightly expanded. Tympanum/eye ratio generally less than 3/5. IMT very large and

protruding (ratio width/length less than 1/2). - (2) Dorsal color and pattern: Dorsum and

flanks dark brown to black. On the upper head surface, the dark color of the dorsum

gradually fading into light brown. Dorsolateral color border present: indistinct in the inguinal

Source : MNHN, Paris

40 ALYTES 17 (1-2)

region, but very distinct in the head and shoulder region. Hand, fibula, foot, tarsus and tibia

light brown, with few dark brown crossbands. Humerus and femur yellow to green, in some

specimens (locality unknown) blue. This color extending as relatively large flank blotches

onto the flanks. Flank blotches delimited by the dorsolateral coloration border and not

extending onto the dorsum. Bright red flash marks present. Iris with light pigment in its upper

part. —(3) Ventral color and pattern: Venter, throat, forelimbs and femur dark brown to black

with small, generally regularly rounded whitish-blue spots and a distinct horseshoe marking,

which in males can cover nearly the complete throat. Tibia with a distinct orange marking,

sometimes continued on the knee, distal part of femur and foot. In preservative, this

coloration changes, becoming partly bright red and partly white, with a sharp border between

both colorations (see also DALY et al., 1996). A similar but less distinct change is also observed

in specimens of M. madagascariensis.

Mantella aurantiaca group

Mantella aurantiaca Mocquard, 1900

Mantella aurantiaca Mocquard, 19004. — Name-bearing type: lectotype, by present designation, MNHN

1899412, probably a male, SVL 21.2 mm. - Type locality: “une forêt entre Beforona et Moramanga”,

according to the original description. - Other types: paralectotype, following present lectotype designation,

MNHN 1899.413. - Etymology: derived from Latin aurantiacus (golden).

Mantella aurantiaca: MOcQUARD, 1900b, 1909; WERNER, 1901; METHUEN & HEwiTT, 1913; GuiBé,

1964, 1978; AuDY, 1973; MUDRACK, 1965, 1974; ARNOULT, 1966; MAT, 1975 (fig.); BACHMANN

& BLOMMERS-SCHLÔSSER, 1975; BLOMMERS-SCHLÔSSER, 1978, 19794; OosTvEEN, 1978a-b; MEIER,

1980 (fig. p. 353 above), 1986; Buse, 1981; DaLy et al., 1984, 1996, 1997a; UNFRIED, 1987; VAN

TOMME, 1988 (fig. 1); AMMER, 1989; SIEGENTHALER, 1989; PinraKk, 1990; Ouiverri, 1990 (fig.):

BLOMMERS-SCHLÔSSER & BLANC, 1991; PRESTON-MAFHAM, 1991 (fig. p. 79); ANDRE 1992 (pl.

I fig. 1-2); GLawW & VENCES, 19924, 1994 (part.), 1998; ZIMMERMANN, 1992, 1996a-b; ZIMMER-

MANN & ZIMMERMANN, 1992 (fig. 5.15-16), 1994; Le BERRE, 1993 (fi GarRAFFO et al., 1993;

HERRMANN, 1993 (fig.); CLARK, 1994 (fig. p. 7); Hay et al., 1995; BARTLETT, 1995 (fig. p. 18 below

left); HENKEL & SCHMIDT, 1995 (fig. p. 49); CarissimI-PRIORI, 1995 (part. fig. p. 41 above and

below right); SraNISzEwsKki, 1996 (pl. p. 23 and 26), 1997a-b (fig.), 1998a (fig.), 1998b; LARSEN,

1997; PiNTAK et al., 1994

Mantella aurantiaca rubra Staniszewski, 1996. - Name-bearing type: lectotype, by present designation, ZFMK

68868, female, SVL 24.6 mm. — Type locality: origin of lectotype unknown; taxon is said to occur in “forests

of Anosibe An'Ala” according 10 original description. — Other types: an unspecified number of (probably

lost) paralectotypes. — Etymology: derived from Latin ruber (red).

Mantella aurantiaca rubra: STANISZEWSKI, 1997b (fig.)

Identity. - Mantella aurantiaca is one of the early names in the genus, and its status as a

distinct species has never been questioned.

Comments. — (1) The lectotype specimen of M. aurantiaca is probably a male, with longitud-

inal, lateral cuts on both sides on the body, and is in slightly better state of preservation than

the paralectotype. The paralectotype is a female in rather poor state of preservation, with a

longitudinal cut through the ventral skin. — (2) STANISZEWSkI (1996) coined the name

Mantella aurantiaca rubra for specimens with a red (instead of yellowish-orange) color. His

diagnosis, although very short, gives in words one character (color) and should thus be

recognized as valid according to the Code: “The type orange form is located in Pandanus

Source : MNHN, Paris

VENCES, GLAW & BÔHME 41

Fig. 3. - Photographs of Mantella species. (a-b) M. milotympanum, specimen without locality data

(ZFMK 65626), dorsolateral and ventral view (c-d) M. aurantiaca, specimen without locality data,

reddish morph (ZFMK 65627), 1997, dorsolateral and ventral view.

forests around Andasibe [...] and the deep blood orange form [known as M. a. rubra] in the

forests of Anosibe An’Ala.” No figure was published together with this description, but

several color photographs were published later (STANISZEWSkI 1997b: 52-53) by the same

author. The assumed type locality Anosibe An’Ala given by STANISZEWSKI (1996) was

probably based on GLAW & VENCES (1994), but STANISZEWSKT'S captive specimens (including

the lectotype described below) almost certainly were obtained through the pet trade without

locality: consequently, the taxon rubra has currently no type-locality. — (3) Regarding the

validity of rubra, it must be stressed that, according to several authors (e.g. ZIMMERMANN &

ZIMMERMANN, 1994; DaLy et al., 1996), reddish aurantiaca morphs occur at several localities,

parapatrically with more orange populations. No evidence supports the status of rubra as

valid subspecies or species, and no genetic differences were found by allozyme electrophore:

between reddish and orange-colored aurantiaca specimens (M. VENCES, personal observa-

tion); we consider rubra as synonym of M. aurantiaca.

Description of the lectotype of Mantella aurantiaca rubra Sraniszewski, 1996. - ZFMK 68868,

adult female with developing oocytes, supplied by M. STANISZEWSKI in 1998 and said to

belong to the series on which the original description was based. Specimen in good state of

Source : MNHN, Paris

42 ALYTES 17 (1-2)

preservation with a longitudinal cut through right flank. For measurements see tab. 2. Body

rather stout; head not broader than body; snout rounded in dorsal and lateral view; nostrils

directed laterally, not protuberant, nearer to tip of snout than to eye; canthus rostralis weak,

straight; loreal region plain; tympanum rather indistinct, medium-sized, rounded, its diameter

about half of eye diameter; supratympanic fold weakly developed; tongue ovoid, only very

slightly bifid posteriorly; maxillary and vomerine absent; choanae small, rounded. Arms

moderately slender; subarticular tubercles single; outer metacarpal tubercle rounded,

inner metacarpal tubercle elliptical, both very weakly developed: fingers without webbing;

finger length 1<2<4<3, finger 4 only slightly longer than 2; finger 2 only slightly longer than 1:

terminal finger disks nearly not developed. Legs moderately robust; tibiotarsal articulation

reaching tympanum; feet with small, rounded inner and outer metatarsal tubercles; subarticu-

lar tubercles single, rounded; toe disks nearly not developed. Foot without webbing. Lateral

metatarsalia connected; toe length 1<2<3<5<4, toe 3 distinctly longer than 5. Skin on the

dorsal and ventral surface smooth. Color in life unknown; in preservative uniformly orange,

ventrally translucent orange. Flashmarks visible as yellowish areas. Iris black, pupil whitish

(due to fixation).

Material examined. - BMNH 1953.1.5,40-41 (Madagascar, pres. G. W. ALLAN); BMNH 1956.1.1.13 ([6

specimens NIL] Périnet District, L. Mason); MNHN 1899.412-3 (lectotype and paralectotype; forest

between Beforona and Moramanga); MNHN 1953.137 (Périnet, forêt}; MNHN 1984.117-23 (coll.

RaZARIHELISOA); MNHN 1988.7515-25 (pet trade); MNHN 1993.1435-9 (locality unknown); MNHN

1994.1105-10; ZFMK 22113-22 (Périnet: leg. H. Meier 1973; 22113, 22115, 22119: CS); ZFMK 56170-83

locality unknown; ded. F GLAW XI.1993); ZFMK 62776, ZFMK 62779, ZFMK 62780, ZFMK 627

ZFMK 62785 (locality unknown; TE): ZFMK 62774, ZFMK 62777 (locality unknown: TE; live

coloration orange); ZFMK 62775, ZFMK 62778, ZFMK 62781-2, ZFMK 62784 (locality unknown: TE;

live coloration reddish); ZFMK 68868 (locality unknown: lectotype of M. a. rubra; ded. M. STANIS-

ZEWSkI, 1998).

Additional specimens were not examined in detail; they are here listed according to the catalogue

entries: MNHN 1976.235-6, MNHN 1976.240-2 (locality unknown); MNHN 1976.237-9 (forêt de

Périnet [239 tadpoles according to catalogue); MNHN 1976.243-9 (forêt de Périnet); ZFMK 8861-70

(Périnet: leg. H. Meter 1973); ZFMK 9127-35 (Périnet: leg. H. Mrter 111.1973); ZFMK 13648-9 (Périnet;

leg. H. Meter 1972); ZFMK 14700 (Périnet; leg. H. Mrier XI.1974); ZFMK 14331-6 (Périnet; leg, H.

Me 11.1974); ZFMK 50170-2 (Andasibe [Périnet}; leg. H. MrtER I1.1989); ZFMK 51792 (Andasibe

[Périnet leg. F. W. HENKEL et al. 1988/89); MRSN A0736.1-4 (locality unknown: through the pet trade):

MRSN A0060.1-4 (Périnet [2]; leg. F. ANDREONE 24.1V.1990); TM 10047-50, 10053-5, 10057-59 (Amba-

toharanana; coll. P. A. Methuen [TM 10051, 10052, 10056 exchanged with MCZ]).

Distribution. — Occurrence in Andasibe is often quoted, but most probably the species does

not occur in the immediate vicinity of this village, records referring to single introduced

specimens. ZIMMERMANN & HETZ (1992) and ZIMMERMANN & ZIMMERMANN (1994) mapped

M. aurantiaca localities in the area of the Torotorofotsy swamps NW of Andasibe. They

found several (more or less isolated) populations, mainly in the northern part of the swamp,

one of these consisting mainly of red colored specimens.

Localities are [1*] the Torotorofotsy swamps (including also Antaniditra, see BLOMMERS-

SCHLÔSSER, 1979) and two other localities which are based on a personal communication of

A. PEYRIERAS: uniformly yellow or orange Mantella specimens are known from near [2]

Beparasy, whereas near [3] Anosibe An’Ala reddish specimens occur. METHUEN & HEWITT

(1913) reported the species from [4] Ambatodradama (Ambatoharanana according to TM

catalogue), which, according to their map, is located near Analamazoatra.

Source : MNHN, Paris

VENCES, GLAW & BÔHME 43

Detailed data on the distribution of the species were also included in the unpublished

report of BEHRA et al. (1995). These authors, beside delimiting the exact distribution area in

the Torotorofotsy area, listed several other localities of uniformly colored Mantella in the

central part of the Eastern Region. Considering the existence of another uniformly orange

species, M. milotympanum (see below), specific belonging of these populations is uncertain.

Uniformly orange specimens were also observed on the Rantsara plateau between Ihosy

and Ivohibe (A. PEYRIERAS, personal communication). This record, however, possibly

corresponds to M. aff. baroni which occurs on Pic Ivohibe. The locality “Fiherenana valley”

(see GLaw & VENCES, 1994) is here referred to M. milotympanum (see below). The map shown

by UNFRIED (1987), giving the whole of eastern Madagascar as the distribution area of M.

aurantiaca, must clearly be considered as pure fantasy.

Diagnosis. — (1) Morphology: À generally rather small and stout Mantella. SVL generally

19-24 mm, but some females can reach up to 31 mm. TTA reaching the forelimb insertion in

large females, the eye center in small specimens, but generally the tympanum or posterior eye

margin. Terminal disks of fingers and toes slightly expanded. Tympanum/eye ratio between

1/2 and 3/5. IMT medium sized (ratio width/length slightly less than 3/4). — (2) Dorsal color

and pattern: Uniformly yellow-orange, in some populations red-orange, often with a translu-

cent shade. Bright red flashmarks present. Iris nearly uniformly black, only a little light

pigment in its upper part. — (3) Ventral color and pattern: Uniform, similar to dorsal surface

but generally somewhat lighter, except red flashmark (extended nearly on the whole tibia).

Some inner organs visible through the slightly transparent ventral skin.

Mantella crocea Pintak & Bôhme, 1990

Mantella crocea Pintak & Bôhme, 1990. - Name-bearing type: holotype by original designation (PINTAK &

BÔHME, 1990: 59), ZFMK 45007, female, SVL 22.5 mm. — Type locality: “Andasibé (= Périnet), mittleres

Ostmadagaskar”, according to original description. — Other sypes: paratypes, ZFMK 45008, ZFMK

50173-86, ZFMK 50552-3, ZFMK 50721-5, and 10 (lost) additional paratypes (see comment below). -

Etymology: derived from Latin croceus (saffron yellow).

Mantella crocea: PINTAK, 1990; BLOMMERS-SCHLOSSER & BLANC, 1991 (p. 274): ZIMMERMANN, 1992;

ANDREONE, 1992 (pl. IV fig.3-4): GLAW & VENCES, 19924; ZIMMERMANN & ZIMMERMANN, 1992

(fig. 5.23); OTTENSMANN, 1993; GarrArrO et al., 1993; HERRMANN, 1993 (fig.); ZIMMERMANN &

ZIMMERMANN, 1994; GLaw & VENCES, 1994; BARTLETT, 1995 (fig. p. 16 above): HENKEL &

Scamipr, 1995 (fig. p. 51); ZIMMERMANN, 1996a-b; STaNISzEwSkI, 1996 (pl. p. 18 and 23); DALY et

al., 1996: SranszEWSkI, 1997a-b (fig.), 1998a (fig.); LARSEN, 1997; PiNTAK et al., 1998; GLAW &

VENCES, 1998.

Other chresonyms:

Mantella viridis: SraNiszewsk1, 19974 (fig. on p. 13 and 17), 19976 (fig. pp. 33, 49, 50), 1998a (fig).

Comments. — (1) Since the holotype was supplied by the pet trade, the exact location of the

type locality is uncertain. It seems rather probable, however, that it is roughly in the central

eastern rainforest region north of Andasibe (formerly Périnet). — (2) In the original descrip-

tion (PINTAK & BÔHME, 1990), beside the catalogued specimens, 10 living uncatalogued

specimens were designated as paratypes. No specimens of this captive stock were eventually

preserved and catalogued; all these additional paratypes must therefore be considered

as lost.

Source : MNHN, Paris

44 ALYTES 17 (1-2)

Material examined. - ZFMK 45007 (Périnet area [*}: through pet trade, 1986; holotype); ZFMK 45008

(Périnet area [?}; through pet trade, 1986; paratype); ZFMK 50173-86 (Moramanga: leg. H. MEIER

IL.1980; paratypes); ZFMK 50552-3 (Moramanga: leg. F. W. HENKEL, W. ScHmipr & V. MÜLLER

V.1989; paratypes): ZFMK 50721-3 (Moramanga: leg. H. Meter 1989; paratypes); ZFMK 50724-5

(Andasibé [Périnet}; paratypes); ZFMK 51480-2 (Andasibé [Périnet]: leg. O. PRONK 11.1990); ZFMK

51738-42 (Périnet; leg, H. ZIMMERMANN 1989); ZFMK 52746 (Andasibé? [Périnet]: ded. F. GLAwW & M.

VENCES 1991; CS); ZFMK 62760-1, ZFMK 62763, ZFMK 62766, ZFMK 62769 (locality unknown; TE):

ZFMK 62765, 62767 (locality unknown; TE: live coloration yellow; 62767 CS); ZFMK 62762, ZFMK

62764, ZFMK 62768 (locality unknown; TE: live coloration green); MNHN 1993.1448 (locality un-

known); MRSN A0058 (Andasibe [?}: leg. F. ANDREONE 4.1. 1992).

Distribution. — The type locality (Andasibe) could not be confirmed by recent surveys

(see above). Also the Moramanga locality (ZFMK vouchers) seems rather dubious. The

only reliable information of which we are aware is included in BEHRA et al. (1995), who

confirmed the occurrence of the species in the Bakozetra area north of Andasibe (located

immediately to the north of the known distribution area of M. aurantiaca in the Torotoro-

fotsy swamps).

Diagnosis. (1) Morphology: À small Mantella. Small specimens of slender appearance, large

specimens rather stout. SVL 17-24 mm, females 23-24 mm. TTA mostly reaching the tympa-

num or posterior eye margin, rarely theeye center. Terminal disks of fingers and toesexpanded.

Mean tympanum/eye ratio nearly 3/5. IMT medium sized (ratio width/length slightly more

than 2/3). — (2) Dorsal color and pattern: Head, dorsum and posterior part of flanks yellow,

orange or light green, sometimes (mainly in the yellowish specimens) with fine black spots.

Sometimes an indistinct dark middorsal line and traces of a diamond marking. Head laterally,

and anterior flanks generally black (black pattern can be largely reduced in some specimens),

with a sharp dorsolateral color border. Light frenal stripe present, often interrupted in the yel-

lowish specimens. Bright red flashmarks present. Iris with some light pigment in its upper part.

— (3) Ventral color and pattern: Black with a variable number and extension of grey to bluish-

white or yellowish markings, sometimes fusing to form an irregular network. Horseshoe mark-

ing present and mostly distinct, but poorly developed in some specimens. Hindlimbs sometimes

uniformly orange or reddish ventrally, except the red ventral flashmark extension on the tibia.

In other specimens, orange color only present on tibia (except flashmark area); foot, tarsus and

femur being black with grey-whitish markings. Pictures of dorsal and ventral coloration of the

different morphs were given in GLAW & VENCES (1998).

Mantella milotympanum Staniszewski, 1996

Mantella aurantiaca milotympanum Staniszewski, 1996. - Name-bearing type: lectotype, by present designation,

specimen figured on p. 18 of STANISZEWSKI (1996): this specimen was not preserved and must therefore be

considered as lost (STANISZEWSKI, personal communication). - Tipe locality: the taxons said Lo oceur in the

“Fiherenana Valley in central east Madagascar” according to the original description, but the locality of

kctotype is unknown. — Other types: an unspecified number of (probably lost) paralectotypes. -

Etymology: probably derived from classical Greek melas (genitive melanos). black (milo being a derived

spelling which possibly was originally created by pet dealers) and classical Greek rumpanon (latinized as

L'mpanum), drum (meaning eardrum), referring 1 the black tympanum color.

Mantella aurantiaca milorympanum: STANISZEWSKI, 19974 (fig).

Mantella aurantiaca “milotympanum": STANISZEWSKI 1997 (fg.)

Mantella “milorympanum”: LARSEN, 1997.

Source : MNHN, Paris

VENCES, GLAW & BÔHME 45

Other chresonyms:

Mantella aurantiaca: Le BERRE, 1993 (fig. p. 20); GLAW & VENCES, 1994 (part; “black tympanum®);

CarissiMt-PrioRt, 1995 (part.: fig. p. 41 below left).

Mantella cf. aurantiaca: GLAW & VENCES, 1994 (pl. 52).

Mantella sp. 3: VENCES & KNIEL, 1998.

“Black-eared mantella”: STANISZEWSKI, 1998a.

Mantella sp., Variante 3: GLAW & VENCES, 1998

Maniella sp.. Variante 4: GLAW & VENCES, 1998.

Identity. — The name milotympanum was, to our knowledge, first used in a publication by

STANISZEWSKI (1996) to name a form of M. aurantiaca previously referred to as “black

tympanum” variant (GLAW & VENCES, 1994). SrANISZEWSKI (in litteris, 1997) had no

intention to create a new scientific name, and his paper does not include a formal description

nor a type designation. However, it describes distinctive features of the form in a way that

must be regarded as a diagnosis:

“Tam in no doubt that a mantella currently defined as another subspecies of the golden

mantella should be raised to specific status. The black-eared golden mantella (Mantella

aurantiaca milotympanum) is so different in appearance and behaviour that it must merit this.

[...] The dorsal colour is a slightly drab orange (males brighter than females) while the venter

is a greenish yellow (orange yellow in M. aurantiaca). This species is overall much slimmer

than the golden mantella, the eyes are oblong rather than round and the skin is much more

granular. Significant raised veins are apparent on the hind limbs, as its name suggests the

eardrum (tympanum) is black as is the nostril region and there is a black line apparent from

the eye to the nostril [...]”. (STANISZEWSKI, 1996: 24).

According to our observations, the presumed slim habitus is not present in all specimens

(especially absent in large females), and the eyes are not of oblong shape (rounded as in other

Mantella). The presumed “semi-nocturnal behaviour” and “very nervous disposition” were

not confirmed by us in our captive group of this species. The same regards the observation of

eges “possessing a yellowish-brown nucleus and measuring only 1 mm in diameter”.

Nevertheless, a diagnosis of this form exists (see above), and the name was not used in a

conditional way. It must therefore be regarded as nomenclaturally available. Since this form

differs from typical M. aurantiaca and M. crocea, We here consider it as a full species in a

preliminary way (see section Specific status below).

Comment. — The locality information “Fiherenana valley” in the original description

almost certainly was based on a personal communication of A. PEYRIERAS as published in

GLAW & VENCES (1994). It is not sure that STANISZEWSKl's specimens were collected at this

locality.

Lectotype designation. - We here follow the procedure applied by Dusois & OHLer (1997a-b) to

stabilize old names for which no type material is preserved in scientific collections but figures

were published. The original description (STANISZEWSKI, 1996: 18) includes a color picture

which shows all characters currently known as characterizing the form (black pigment on tym-

panum and around nostril, rather granular skin, dorsal color not of translucent appearance).

We designate this figured specimen as lectotype. This specimen (as all specimens kept by M.

STANISZEWSKI until the description of milorympanum) was not preserved, and is therefore not

available for comparative purposes (STANISZEWSKI, in litteris 1997). À neotype designation is

Source : MNHN, Paris

46 ALYTES 17 (1-2)

postponed until specimens with reliable collecting data become available. In the following, we

describe one reference specimen from the ZFMK collection for comparative purposes.

Description of reference specimen. — Adult male specimen, ZFMK 65626, SVL 22.5 mm.

Specimen in excellent state of preservation, with longitudinal cuts along both flanks.

For measurements see tab. 2. Body slender; head not broader than body; snout slightly

pointed in dorsal, truncated in lateral view; nostrils directed laterally, not protuberant,

nearer to tip of snout than to eye; canthus rostralis weak, slightly concave; loreal region

even; tympanum rather indistinct, medium sized, rounded, its diameter about half

of eye diameter; supratympanic fold moderately developed; tongue longish, only very

slightly bifid posteriorly; maxillary and vomerine absent; choanae small, rounded. Arms

slender; subarticular tubercles single; outer metacarpal tubercle rounded, inner metacarpal

tubercle elliptical, both very weakly developed; fingers without webbing; finger length

1<2<4<3, finger 4 only very slightly longer than 2; finger 2 only slightly longer than 1;

terminal finger disks nearly not developed. Legs moderately robust; tibiotarsal articula-

tion reaches tympanum; feet with small, rounded inner and outer metatarsal tubercles;

subarticular tubercles single, rounded; toe disks faintly developed. Foot without webbing.

Lateral metatarsalia connected; toe length 1<2<3<5<4, toe 3 distinctly longer than 5. Skin

on the upper surface smooth, slightly granular on the flanks; ventral surface smooth,

except for granular thigh patches (“femoral glands”) extending from the anus ca. 5 mm

distally (max. width 3.1 mm). Color in life dorsally, and on flanks and upper surface of fore-

and hindlimbs deep orange, except for small black areas around the nostril and covering

the tympanum. bright red flashmarks. Ventral side orange except for the dirty blackish

“femoral gland” region. After one year in preservative, the orange color has changed to olive

greenish. The flashmark areas are yellowish. The ventral side is dirty olive except for

the hindlimbs which are yellowish. The “femoral gland” region is dark brown with small

whitish spots.

Maierial examined. = ZFMK. 62770 (locality unknown; CS); ZFMK 62771 (locality unknown: TE);

ZFMK 62772, 65626, 68868 (locality unknown; 62772: TE: live coloration red-orange): ZFMK 62773

(locality unknown; TE; live coloration yellow-orange}; MNHN 19924823 (locality unknown;

identification based on remains of dark pigments on tympanum and around nostril).

Distribution. — According to A. PEYRIERAS (personal communication in GLAW & VENCES,

1994) this species occurs in the Fiherenana valley, located about 50 km N Andasibe (not the

Fiherenana valley in the South-Western region, near Toliara).

Diagnosis. — (1) Morphology: Generally, a rather small and stout Mantella, although single

females can become relatively large. SVL generally 19-23 mm (females exceptionally up to

30 mm; personal observation, specimen not preserved). TTA reaching the tympanum or

posterior eye margin. Terminal disks of fingers and toes slightly expanded. Mean

tympanum/eye ratio slightly larger than 1/2. IMT rather large (ratio width/length less than

3/5).- (2) Dorsal color and pattern: Uniformly yellow-orange or red-orange, without translu-

cent shade, and with a black spot covering the tympanum and a little black pigment around

the nostril. Bright red flashmarks present. Iris nearly uniformly black, only a little light

pigment in its upper part. — (3) Ventral color and pattern: Uniform, similar to dorsal surface

but generally somewhat lighter. Area of “femoral glands” often speckled with blackish. Tibia

bright red.

Source : MNHN, Paris

VENCES, GLAW & BÜHME 47

Fig. 4. - Ventral and dorsal views of name-bearing types of Mantella species. (a) M. betsileo (lectotype,

MNHN 1895.278); (b) M. bersileo (lectotype of M. attemsi, NMW 20837): (c) M. expectata

(holotype, ZFMK 53540); (4) M. viridis (holotype, ZFMK 47900): (e) M. laevigata (holotype, TM

10074); (D M. madagascariensis (holotype of M. loppei, NMB 4849); (g) M. madagascariensis

Gectotype, MNHN 1895.276). Not to scale

Source : MNHN, Paris

48

ALYTES 17 (1-2)

Fig. 5. — Ventral and dorsal views of name-bearing types of Mantella species. (a) M. nigricans (lectotype,

MNHN 1973.55); (b) M. haraldmeieri (holotype, ZFMK 25351); (c) M. baroni (holotype, BMNH

947.2.7.19); (d) M. cowani (lectotype, BMNH 1947.2.7.4); (e) M. pulchra (holotype, BMNH

1947.2.7.20); (1) M. crocea (holotype, ZFMK 45007); (8) M. bernhardi (holotype, ZFMK 57164); (h)

M. aurantiaca (lectotype, MNHN 1899.412). Not to scale.

Source : MNHN, Paris

VENCES, GLAW & BÜHME 49

[Mantella betsileo | betsileo

LEE UT

ZFMK 17604 ZFMK 17605 GLaw & VENCES ZFMK 62688 ZFMK 62686

(1994: fig. 319)

LELXLEI

ÉELLE

MNHN 1973.496 MNHN 1973.494 MNHN 1973.489 MNHN 1973.487 MNHN 1973.491

Mantella expectata Mantella manery

ZFMK 53541 ZFMK 53540 ZFMK 59096 Holotype

Mantella viridis

ZFMK 47900 ZFMK 48046 ZFMK 48047 GLAW & VENCES

(1994: fig. 318)

Fig. 6. - Variation of ventral pattern in species of the Mantella betsileo group.

Source : MNHN, Paris

50 ALYTES 17 (1-2)

[Mantella laevigata | laevigata

ELLES

ZFMK 52751 ZFMK 59913 GLAW & VENCES GLAW & VENCES ZFMK 52750

(1994: fig. 322) (1994: fig. 323)

Mantella nigricans

MNHN 1973.555 MNHN 1973.551 ZFMK 59902 ZFMK 59887 ZFMK 59888

Mantella haraldmeieri

# 029 K f ÿ

Eye L EME fe LME

\ \) \ ;

$ eu es =

1 T Ti 4 TT

MNHN 1973.511 ZFMK 21807 ZFMK 21805 MNHN 1973.505 MNHN 1973.510

Mantella aff. baroni

f g

|, SM &

} l }

MNHN 1991.1602 MNHN 1972.771 MNHN 1972.768 MNHN 1972778 MNHN 1991.1804

Fig. 7. - Variation of ventral pattern in Mantella laevigara and some species of the M. cowani group.

Diagonally hatched areas represent light coloration which is different from the normal bluish or

greyish (exceptionally greenish-yellow) spots and markings on the black venter. A further differen-

tiation of the light color was not undertaken, partly because in many preserved specimens the color

is largely faded. The diagonally hatched areas thus comprise orange, vellowish, and light brown areas

as well as the flashmark areas of some species which in life are vivid red.

Source : MNHN, Paris

VENCES, GLAW & BÔHME SÏ

MNHN 1933.247 MNHN 1991.1805 MNHN 1936.41 MNHN 1972775 ZFMK 28870

Mantella cowani Mantella crocea

MNHN 1973.528 MNHN 9594 MNHN 1973.529 ZFMK 52746 ZFMKG62764 ZFMK 62761

Mantella puichra Mantella bernhardi

LLL

7

ZFMK62258 ZFMK52122 GLAW & VENCES GLAW & VENCES ZFMK 62702 ZFMK 62705

(1994: fig. 330) (1994: fig. 324)

Mantella madagascarien

Denon, st LT

\ \

$ \ s

s 7

ZFMK 56154 ZFMK 14327 ZFMK 14326 ZFMK 56153 GLAW & VENCES

(1994: fig. 331)

Fig. 8. — Variation of ventral pattern in some species of the Manrella cowani group, and in species of the

M. madagascariensis group, M. bernhardi group and M. aurantiaca group. Se also legend of fig. 7

Source : MNHN, Paris

52 ALYTES 17 (1-2)

CO CE:

es #2 +

1e 1 7

1 16 e

1 1 Ta

2 2 2

2 22° 22

24°! 24 24

12

1e

16

18

| me

| |

: #

æt [TT j 2 æ Le

/ J |

— ÿ ! | @ m mada-

a Lu JE gascarienss | | 24e

} CE | | @ 4 puictra

. { M , { M M bomnard | |

46 4 So ca CRT

Fig. 9. - Distribution maps of Mantella species as distinguished in the present paper. Positioning of

localities in the maps is only approximate and mainly based on BLOMMERS-SCHLOSSER & BLANC

(991).

Source : MNHN, Paris

VENCES, GLAW & BÜHME 53

KEY TO THE CURRENTLY KNOWN SPECIES OF Mantella

The following key should allow identification of all currently known Mantella species by

their live coloration. Examination of both dorsal and ventral patterns is necessary for a

reliable identification. Where useful, we also give morphological, ecological or bioacoustic

characters as additional identification aids. A reliable identification of preserved specimens is

not always possible, especially in formalin fixed individuals with faded pattern contrast, and

in hybrid or rare intermediately colored specimens.

1. Ventral surface of hindlimbs partly or completely orange, yellow and/or red... 2

Ventral surface of hindlimbs black with blue, whitish-blue or greyish markings, without

orange or red elements. 12

2. Dorsal coloration uniformly green, yellow, orange or reddish, with only rudimentary,

dispersed'black:elements 1522.46 4e 0 arantian nn bep néilenareste tres Sels 24e 3

Dorsally with distinct black or dark brown elements, often covering the largest part of

dorsum and/or flanks

3. Ventral surface generally black with light markings, at least with some distinct black

patterns 4

Ventral surface uniformly yellow or orange. 5

4. Flank blotch area more densely covered by green/yellow than remaining flanks:; flashmarks

presents ur unent heu ut en UOR A. M. madagascariensis, variable morph

Flank blotch area not more densely covered by green/yellow than remaining flanks:

horseshoe marking and flashmarks present. ............................... M. crocea

Flank blotch area more or less densely covered by green/yellow than remaining flanks;

horseshoe marking and flashmarks absent ............... M. aff. baroni (Andringitra)

5. Black pigment absent; skin often with a translucent shade.............. M. aurantiaca

Black pigment present on tympanum and around nostril; skin without translucent

shade M. milotympanum

6. Frena

Frenal stripe absent

stripe present .

7. Flank blotches present, often integrated in an irregular network pattern of green/yellow

and black M. madagascariensis, variable morph

Flank blotches absent: flanks anteriorly black, posteriorly of same color as

COUR Ses PRE Peben Fame te nue Led R eee ENS DS MT M. crocea

8. Horseshoe marking present; chirp or trill calls.................................... 9

Horseshoe marking absent: single click calls

9. Small species (adult SVL 18-22 mm); IMT small; flank blotches very small, flanks thus

nearly uniformly black; dorsum grey, with sharp but little distinct dorsolateral color

border 5e tentes Poe ee are ones 2 tn ed pres ener al M. bernhardi

Source : MNHN, Paris

54 ALYTES 17 (1-2)

Larger species (adult SVL 20-27 mm); IMT large; flanks black with large yellow, greenish

or blue fank:blotches 00e. sn iteue ee eue une cs es perereponeners de ie 10

10. Dorsum, and especially dorsal head surface, brown, with a distinct dorsolateral

color border to the black flanks; femur ventrally generally without red/orange

color M. pulchra

Dorsum and head surface black (sometimes with green/yellow); femur ventrally generally

with red/orange patterns M. madagascariensis

11. Dorsolateral color border present; flank blotches small, beige; hindlimbs dorsally

Don re nées nete tm ne ne ED ORe M. haraldmeieri

Dorsolateral color border absent; flank blotches medium-sized, generally red; hindlimbs

dorsally black with red

Dorsolateral color border absent; flank blotches large and yellow or greenish; tibia, tarsus

and foot dorsally orange with black.................... M. baroni and M. aff. baroni

13

M. betsileo group, 14

13. Throat generally uniformly black, without or with very few light markings; flank blotches

absent; fingers and toes with largely expanded terminal disks; double click calls; partly

arboreal habits M. laevigata

Throat black with light markings; flank blotches present; fingers and toes with moder-

M. cowani

12. Frenal stripe and horseshoe marking absent

Frenal stripe present; horseshoe marking generally present .

ately expanded terminal disks; single click calls; terrestrial habits... M. nigricans

14. Flanks anteriorly black, posteriorly greenish, no dark crossband on tibia... M. viridis

Flanks anteriorly black, posteriorly brownish-red .......................... M. sp. 1

Flanks generally uniformly black or dark brown... 15

15. Dorsum brownish; dark crossband on tibia present. M. betsileo

Dorsum yellowish; limbs blue to grey, dark crossband on tibia absent... M. expectata

Dorsum yellowish; limbs brown ........................................ M. manery

DISCUSSION

RELIABILITY OF PUBLISHED DATA AND TREATMENT OF **PHANTOM NAMES”

During our survey of literature for the present paper, we became aware of many errors,

especially regarding locality data. Furthermore, we noted that during the last years, hobbyists

increasingly published unreliable or fantasy data on distribution, behaviour, variation and

reproduction of Mantella species. With this statement, we do not want to downgrade

publications of amateur herpetologists to Mantella knowledge in general. Several important

contributions were published e.g. by ZIMMERMANN (1992, 1996a-b), MEIER (1975, 1980, 1986)

and STANISZEWSKI (1998b), among others. However, distributional data such as those of

UNFRIED (1987), data on reproduction such as those of LE BERRE (1993; M. laevigata tadpoles

Source : MNHN, Paris

VENCES, GLAW & BÔHME 55

Table 3. - Phantom names of Mantella forms, their identity and current status. Additionally, the

following phantom names (with clear mention of their conditional status) were listed by

STANISZEWSKI (1998a): Mantella spezei, Mantella crocea calxis, Mantella verronique,

Mantella tulaï, Mantella mangabe.

Name History of name Taxonomic status Nomenclatural status

Mantella “mysteriosa": not used any more | M madagascariensis, conditional name

BARTLETT, 1995 “variable morph” (not available)

Mantella nasuta Sp.: not used any more |. M: madagascariensis, nomen nudum

CLARK, 1994 “variable morph” (not available)

name used in several

Mantella aurantiaca rubra | er habbyist synonym of Mantella A

Si ki, 1996 hear aurantiaca

Pros publications

Mantella aurantiaca | name used in several "

milotympanum other hobbyist " FR available name

Staniszewski, 1996 publications

name used in several

“Mantella marojezyi”: Mantella manery, conditional name

other hobbyist

à d he t available)

STANISZEWSKI, 1996 Tüblications described herein (noi )

Mantella “marojezy": Rs Mantella manery, sn

bed h ot available

LARSEN, 1997 publications described herein LL (n )

Mantella “negristata": | name not yet used again | Mantella nigricans conditional name

(not available)

LARSEN, 1997

developing within two weeks), habitat data such as those of STANISZEWSKI in his 1997b

booklet (e.g. Mantella crocea and M. cowani occurring in lowland forests), and lists of

assumed new species as given in CLARK (1994), lack of any reliable data basis and must largely

be seen as inventions of the authors or their informants. Especially the work of Andrew

CLARK (1994) must be read with extreme caution in this respect. So, the information of a

single specimen collected at high altitude on the Marojezy mountains which belongs to a new

species and possibly new genus, quoted by CLARK (1994: 12) as personalcommunication of R.

NussBAUM, is false; in fact, no such species was collected, and no such information provided

to A. CLARK (NUSSBAUM, in litteris 1997). The major problem is that new scientific names are

constantly coined in these papers. New Mantella names used without proper description and

type designation for the taxon are here referred to as “phantom” names.

As discussed in the corresponding sections and summarized in tab. 3, most phantom

names used until now are nomenclaturally not lable since they must regarded as nomina

nuda due to the lack of a diagnosis, or as conditional names due to the use of quotation

marks. Unfortunately, this does not apply to two of the names coined by STANISZEWSKI

(1996), M. aurantiaca rubra and M. milotympanum, which are stabilized by lectotype designa-

tions in the present paper.

Source : MNHN, Paris

56 ALYTES 17 (1-2)

As a conclusion, editors of hobbyist journals should not permit their authors usage of

new scientific names to name undescribed or undetermined morphs unless the names are

accompanied by a formal description and type specimens are deposited in a publicly available

scientific collection. Instead of phantom names, authors should be advised to refer to

unknown morphs with numbers, letters or localities in quotation marks (e.g. Mantella sp. A.,

Mantella sp. 1, Mantella sp. “Marojezy”).

According to the official information available in December 1998 on the ICZN webpage

(www.iczn.org), the fourth edition of the Code will include the following requirements for new

specific names proposed after 1999 to become available (slightly shortened in the following):

(1) the new name must be explicitely indicated as being new (preferably by a term such as “sp.

nov.”); (2) the description will have to include the explicit fixation for it of a name-bearing

type (a holotype or a syntype series); (3) when the name-bearing type of a species-group taxon

proposed after 1999 consists of a preserved specimen or specimens, the proposal will be

required to include a statement naming the collection(s) in which the name-bearing type is to

be found.

Based on our experiences with Mantella phantom names, we strongly support these new

requirements (as compared to the third Code edition currently in force) to valid species

descriptions, which will at least avoid “accidental” taxa descriptions in hobbyist journals and

pet dealer lists in the near future.

SPECIFIC STATUS

It must be stressed that the taxonomic status of several of the species as defined in the

present paper is not yet totally clarified. This concerns M. manery, for which basic data on

morphology and variation are lacking, the species of the M. aurantiaca group which appear to

be very similar genetically (VENCES et al., 1999b), and M. pulchra which may be a subspecies

of M. madagascariensis. I also concerns M. aff. baroni, R. NUSSBAUM (personal communi-

cation) collected specimens referable to this form at a locality south of Andringitra,

confirming that it occupies a range between those of M. baroni and M. haraldmeieri. The

specific status of these and the remaining taxa of the M. cowani group (all apparently

distributed allopatrically) must still be confirmed. Specimens with intermediate color patterns

are known which possibly are hybrids of M. baroniand M. cowani (personal observation), and

others which may represent intermediates between M. baroni and M. nigricans (specimens

from Zahamena and Folohy; see section on M. baroni).

Generally, more detailed data of the species’ distribution, variability and genetic differ-

entiation in contact (hybrid?) zones are necessary. Some available data, however, already

indicate a substantial amount of differentiation between the taxa mentioned above, so that

attribution of specific status to them seems currently the most consistent hypoth Our

proposal to consider all these forms as distinct species is based (1) on several biological

indications, and (2) on practical reasons.

(1) Arguments for the specific distinctness between M. haraldmeieri, M. cowani and M.

baroni are (a) the chromosomal differences between M. baroni and M. haraldmeieri (PINTAK et

al., 1998), (b) the morphological differentiation of M. cowani (personal observation), and (c)

Source : MNHN, Paris

VENCES, GLAW & BÔHME SF

a relevant genetic differentiation between M. baroniand M. cowani (VENCES et al., 1999b). The

specific status of the closely related, probably allopatric forms M. madagascariensis and M.

pulchra is currently only corroborated by color differences and by a certain genetic differen-

tiation detected by allozyme electrophoresis (VENCES et al., 1999b), but it cannot be excluded

that M. pulchra is in fact a northern subspecies of M. madagascariensis. The very low genetic

differentiation between all three species of the M. aurantiaca group (VENCES et al., 1999b;

determined by allozyme electrophoresis) as well as the rather large color variability of M.

crocea Would support their status as color morphs of one single species. However, (a) the

status of crocea as separate species was corroborated by chromosomal differences to auran-

tiaca (PINTAK et al., 1998), (b) relevant chromosomal differences were also found between M.

aurantiaca and M. milotympanum (G. ODIERNA, personal communication), and (c) hybridi

zations in captivity between M. aurantiaca and M. milotympanum resulted in less vital

offspring than simultaneously reared young of M. aurantiaca (personal observation).

(2) Mantella species are attractive animals which are often kept in captivity and traded in

rather large numbers (BEHRA, 1993; GoRZULA, 1996). To get an overview of the extent of

trade and possibly necessary protection efforts and trade restrictions, it is often useful to have

scientific names which can easily and reliably be assigned to forms with a certain, character-

istic coloration. For example, M. aurantiaca as presently defined has been in the center of

conservation efforts and discussions on trade restrictions (e.g. ZIMMERMANN, 1996a), and the

inclusion of M. crocea and M. milotympanum as junior synonyms (respectively their posterior

resurrection, since detailed future studies will possibly corroborate their specific distinctness)

would cause confusion in conservation organizations and administrations, as for example in

CITES authorities. These practical considerations are an additional support for our decision

to assign species status to all currently distinguishable Mantella forms.

COLOR VARIABILITY

Our results allow for a first time to draw definitive statements on intrapopulational color

variability in Mantella species. Earlier analyses (e.g. GUIBÉ, 1964: fig. 2-6) are confusing in this

respect since they mixed several populations, belonging to different species, to demonstrate a

presumed large variability in single taxa. In the following, we first summarize the current

knowledge about intrapopulational color variability, and subsequently the known variability

among different populations of the same species. Finally, we discuss deviating color morphs

without reliably known localities.

Color and pattern variability within populations

(1) According to our data, dorsal and ventral coloration is rather uniform within

populations of M. baroni, M. betsileo, and also in the one population of M. aurantiaca which

we observed in the area of the Torotorofotsy swamps. — (2) A slight variability is known in M.

laevigata (Nosy Mangabe population), mainly regarding the posterior extension of the

yellow-greenish dorsal color (GLAW & VE 1992b). In M. haraldmeieri, the extension of

flank blotches can vary between individuals (fig. 10). - (3) An important variability is

observed in the dorsal pattern (extension of yellowish/green color) of M. nigricans (Marojezy

Source : MNHN, Paris

58 ALYTES 17 (1-2)

MNHN 1973.507

MNHN 1973.505 MNHN 1973.511

Fig. 10. - Size variation of flank blotches in Mantella haraldmeïeri from the Chaînes Anosyennes. The

dorsolateral color border is not sufficiently recognizable in the figured specimens and is therefore not

included in the drawings.

population; see fig. 1g-h). In M. sp. 1 from Ankarana, the extension of the fiery red flank color

is very variable (VENCES et al., 1996). Even more extreme variability is found in the dorsal

pattern of M. aff. baroni as it is corroborated by MNHN vouchers which reliably were

collected at the same locality. — (4) Too little is known for reliable statements on intrapopu-

lational variation of the remaining species.

Color and pattern variability among populations

(1) According to the existing data, differences are rather low between populations of M.

baroni (see also DALY et al., 1996 and ANDREONE, 1993), except for the deviating specimens

from the localities Folohy and Zahamena at the probable northern distribution edge. Simi-

larly, no differences are known between M. laevigata populations. — (2) Slight differences are

known in M. betsileo; the Kirindy population differs from the east coast and Sambirano

populations by reddish brown crossbands on the hindlegs, and a lighter leg color (VENCES et

al., 1996). — (3) Too few data are available on most other species; a high variability among

populations may be found in the M. aurantiaca group when more extensive fieldwork is

ried out on these specie me is true for M. madagascarie: ee below), in which the

observed high variability may also be due to intrapopulational variation.

Color and pattern variability in specimens without reliable locality information

(1) In some cases, deviating colorations have been observed in single specimens. One M.

laevigata specimen from the pet trade had brown instead of black legs (GLAW et al., 1998). —

Source : MNHN, Paris

VENCES, GLAW & BÔHME

my) «À

M. madagascariensis lectotype

MNHN 1895.276

4

_s

M. madagascariensis

ZFMK 62741 (left)

ZFMK 62740 (right)

F

+

>

M. baroni

MNHN 1991.1805

M. haraldmeieri

MNHN 1973.508

M. madagascariensis

MNHN 1935.416 (paratype loppei)

© Ÿ

1Y

M. aff. baroni

MNHN 1991.1804

w Lo

M. cowani

MNHN 1973.528

59

Fig. 11.- Pattern on posterodorsal femur and knee hollow in the lectotype of Mantella madagascariensis

and in se

Madagas

al Mantella species which occur in the Eastern, Central and South-Eastern Regions of

The pattern of the lectotype clearly corresponds best to that of the paratype of M.

loppei (to be considered as junior synonym of M. madagascariensis) and to other specimens here

considered as M. madagascariensis. The dotted line on the femur of the ZFMK specimens marks the

(sharp) color border between yellow (above) and orange (below) which is only visible in lif

after preservation. Regarding dorsolateral color border of M. haraldmeieri, see caption of f

or shortly

Source : MNHN, Paris

60 ALYTES 17 (1-2)

EL _ 2 D

M. madagascariensis lectotype M. madagascariensis

MNHN 1895.276 MNHN 1935.416

M. baroni M. aff. baroni

MNHN 1991.1805 MNHN 1991.1804

,

y y /

{

\

M. cowani M. haraldmeieri

MNHN 1973.528 MNHN 1973.508

Se

À :

CE CE

CES

M. bernhardi

ZFMK 57164 (holotype)

Fig. 12. Ventral pattern on femur and tibia in the lectotype of Mantella madagascariensis and in several

Mantella species which occur in the Eastern, Central and South-Eastern Regions of Madagascar

The pattern of the lectotype clearly corresponds best to that of the paratype of M. loppei

(MNHN 1935.416), but not to M. bernhardi which has a ventrally uniformly light femur

Source : MNHN, Paris

VENCES, GLAW & BÔHME 61

(2) Specimens with intermediate coloration (possibly in some cases due to hybridization) are

known between M. baroni and M. cowani (personal observation), and between M. nigricans

and M. baroni (specimens from Folohy and Zahamena). Also, M. crocea specimens are known

which have a nearly uniform (greenish or yellowish) dorsal color, with only remains of a dark

ventral pattern, and thus appear very similar to M. milotpmpanum (GLAW & VENCES, 1998).

— (3) DaLy et al. (1996) were right in stating that information based on specimens from the pet

trade should be seen with caution, but large series of specimens seen in the cages of the same

dealer at the same time (personal observation) allow, in our opinion, the conclusion of

important variability (dorsally and ventrally) in M. madagascariensis. Whether this variability

is between different uniform populations, or within single variable populations, cannot be

decided at the current state.

Causes of variation

As in dendrobatids (Myers & DaLy, 1983), the evolutionary mechanisms causing the

observed intrapopulational variation (contrasting with the uniformity in other populations) in

some species are not yet understood. Considering the presence of skin alkaloids in Mantella

(Day etal., 1996), their coloration can be seen as largely aposematic. Itis thus possibly subject

to strong predatory selective pressures, and phenomena of Müllerian mimicry, which seem to

be exceptional among anurans (DUELLMAN & TRUEB, 1985), may also be involved.

SYNToPY

According to the data presented herein, the following reliable cases of syntopic occur-

rence of different Mantella species are known (the possible syntopic occurrence of

M. expectata, M. betsileo and M. sp. 1 near Morondava needs confirmation): (1) M. baronil M.

pulchra (An'Ala; ANDREONE, 1993, DaLY et al., 1996; personal observation); (2) M. baronil M.

madagascariensis (Vohiparara, personal observation; Niagarakely, based on ZFMK

vouchers); (3) M. nigricans/M. laevigata (Marojezy, Camp 3; personal observation);

(4) M. laevigatal M. manery (Marojezy, Camp 1; personal observation); (5) M. laevigatal M.

betsileo (Mananara; DALY et al., 1996); (6) M. betsileolM. pulchra (Mananara, DALY et al.,

1996).

It is remarkable that these few cases all refer to species of different species groups

occurring syntopically. On the other hand, in several groups the species appear to be

allopatrically distributed. This is most distinct in the M. cowani group (see fig. 9). Also the two

taxa of the M. madagascariensis group seem to be distributed in an allopatric north-south

pattern, whereas the species of the M. aurantiaca group are probably distributed para-

patrically in swamp forest and rain forest areas in the east. Only in the M. besileo group are

the areas of different forms (M. betsileo, M. sp. 1, M. expectata) known to overlap. It is not

known whether in these overlap areas the distribution patterns are at least locally of close

syntopy or always of parapatry.

Source : MNHN, Paris

62 ALYTES 17 (1-2)

Table 4. - Regional endemism in Mantella species.

Region Number of species | Number of endemic species Endemism

South-West 2-3 0 0%

West 2-3 0 0%

South-East 1 1 100 %

East 10 8 80%

Center 1 1 100 %

North-East 4 2 50%

Northern Center L 0 0%

North 2 1 50%

Sambirano (N.-W.) 1 0 0%

BIOGEOGRAPHY

The almost complete re-examination of the historical voucher specimens and review of

recent field data in the present paper enabled us to present updated distribution maps. The

resulting distribution patterns of many species, especially those of the M. cowani group, are

very different from those presented by BussE (1981) and BLOMMERS-SCHLÔSSER & BLANC

(991).

AIT Mantella species are exclusively distributed on Madagascar and its adjacent islets

(Nosy Be, Nosy Komba, Nosy Boraha, Nosy Mangabe). Records of Mantella species on La

Réunion island (THOMINOT, 1889; GUIBÉ, 1964) or the Seychelles (STANISZEWSKI, 1997b) are

not corroborated by reliable voucher specimens, and must be considered as wrong.

Most Mantella species inhabit areas of tropical rainforest but at least three species (M.

expectata, M. betsileo, M. sp. 1) are known from arid regions in western Madagascar.

Although there are no reliable altitude data for most localities, it can be stated that they are

mostly in-between sea level and ca. 1000 m altitude. Only M. cowani is known to occur at much

higher altitudes (Ambatodradama: 2000 m).

According to ANGEL (1942), as modified by BRYGO0 (1971), GLAW & VENCES (1994) and

RAXWORTHY & NUSSBAUM (1995), Madagascar was herpetogeographically divided into the

Eastern Domain and the Western Domain, each consisting of various regions. The Western

Domain contains the Western and South-Western Regions, the Eastern Domain contains the

South-Eastern, Eastern, Southern Central, Central, North-Eastern, Northern Central,

Northern and Sambirano (North-Western) Regions. Here we follow the delimitation of

regions in the map 3 of GLAW & VENCES (1994).

Source : MNHN, Paris

VENCES, GLAW & BÔHME 63

In contrast to other terrestrial vertebrate groups as the dwarf chameleons of the genus

Brookesia (see RAXWORTHY & NUSSBAUM, 1995), the northern biogeographic regions (North-

West, North, Northern Center, North-East) do not appear to be a diversity center for

Mantella (as compared to the Eastern Region; see tab. 4). AII six species groups defined herein

have representatives in the Eastern Region, whereas only three (M. laevigata group, M.

betsileo group, M. cowani group) have representatives in one of the northern regions. The

Eastern Region harbours at least 10 Mantella species, whereas only between one and four

species are known from each of the northern regions (six species altogether). None of the

species groups is endemic to the northern regions, whereas three species groups are endemic to

the East. Three species (M. viridis, M. manery, M. nigricans) are endemic to the northern

regions, whereas eight species are endemic to the East.

However, these counts may draw a biased picture since many species of the East show in

fact a very low genetic differentiation (VENCES et al., 1999b), and some species complexes may

better be seen as single units for biogeographic comparisons. Counting the M. aurantiaca

group and the M. madagascariensis group as single units, and seeing M. aff. baroni as closely

related to M. baroni, reduces the importance of the Eastern Region as center of diversity and,

especially, endemism of Mantella. It also is interesting that the northern regions are mainly

inhabited by species which are considered as relatively basal within the genus (PINTAK et al.,

1998; VENCES et al., 1999a-b): M. laevigata and the M. betsileo group. Also M. nigricans, due

to the lack of reddish ventral hindleg color, can be seen as the most basal representative of the

M. cowani group.

CONSERVATION

Among the anurans of Madagascar, and beside the tomato frogs (Dyscophus antongili

and D. guineti), Mantella is certainly the group most attractive to the pet trade. According to

BEHRA (1993), a total of 10597 Mantella specimens were legally exported from Madagascar in

the first half of 1990. Mantella species have been subject of discussions on trade restrictions

and CITES inclusion. During several years, Mantella aurantiaca was the only species included

in the CITES regulation (appendix 2) due to its assumed restricted distribution and vulnera-

bility. In 1997, inclusion of several other species (M. haraldmeieri, M. bernhardi, M. cowani

and M. viridis) was discussed. Also, Mantella have been used as key species for the justification

of expansion or implementation of natural reserves (e.g. ZIMMERMANN, 1996).

The basis of all these discussions were the published distributional data and species

definitions, as well as some unpublished reports. For statements on vulnerability by excessive

collecting or habitat destruction, and identification of conservation priorities, a comparative

assessment of the status of all Mantella species is necessary. In the following we analyze five

different factors which may influence the status of Mantella species.

(1) Geographical distribution of the species. — We estimated the extent of the distribution

area and the density by which it is populated by a certain species by the total number of

localities known and the largest distance in kilometers between two locality records attributed

to the species. Species can be classified as follows: (a) common species with a large distribution

Source : MNHN, Paris

64 ALYTES 17 (1-2)

area (> 10 localities, and > 400 km distance between the most distant localities): M. betsileo,

M. baroni; (b) more localized species with a large distribution area (£ 5 localities, > 400 km

distance): M. sp. 1; (c) relatively common species with a moderate distribution area

5 localities, 100-400 km distance): M. laevigata, M. madagascariensis, M. cowani; (d) more

localized species with a moderate distribution area (< 5 localities, 100-400 km distance): M.

expectata, M. pulchra; (e) species with a small distribution area (> 3 localities, 50-100 km

distance): M. nigricans, M. haraldmeieri, M. aurantiaca; (f) localized species which are only

known from one or two localities (distance < 50 km): M. manery, M. viridis, M. bernhardi, M.

crocea, M. milotympanum.

(2) Number of nature reserves and protected areas in which a species is known to occur. - At

present, this is known to apply to the following species and localities: M. betsileo, Tsarata-

nana, Mananara, Masoala, Lokobe, Manongarivo, Tsingy de Bemaraha; M. sp. 1, Ankarana;

M. expectata, Isalo; M. manery, Marojezy; M. laevigata, Mananara, Nosy Mangabe,

Anjanaharibe-Sud, Marojezy; M. baroni, Analamazoatra, Mantady, Ranomafana, probably

Zahamena; M. aff. baroni, Ivohibe; M. nigricans, Anjanaharibe-Sud, Marojezy, probably

Masoala; M. haraldmeïeri, possibly Andohahela; M. madagascariensis, Ranomafana; M.

pulchra, Mananara; M. aurantiaca, not yet known from any protected area (would occur

within the limits of Analamazoatra if this reserve was expanded as suggested by ZIMMER-

MANN, 1996b).

(3) Restriction of the species to primary (forest) habitat. — Field data are lacking or

insuficient for most Mantella voucher specimens examined in the present study. However,

some authors give reliable habitat data of Mantella species, which are here combined with our

personal observations. Species which are until now only found in primary rainforest are

Mantella laevigata (localities Nosy Mangabe, Marojezy: personal observation; Anjanaharibe,

Tsararano: personal communication of F ANDREONE), M. baroni (several localities:

ANDREONE, 1993; DaLy et al., 1996; personal observation), M. haraldmeieri (pristine and

degraded primary forest near Nahampoana:; personal observation), M. nigricans (Marojezy,

Tsararano, Anjanaharibe; personal communication of F. ANDREONE and personal observa-

tion), M. manery (personal observation), M. madagascariensis (Ranomafana; personal obser-

vation), M. pulchra (several localities; ANDREONE, 1993, DALY et al., 1996, personal observa-

tion), M. aurantiaca (swamp forest near Andasibe; personal observation, ZIMMERMANN et al.,

1990), M. crocea (swamp forest; DaLy et al., 1996), and M. bernhardi (a single specimen found

in degraded primary forest rests near rice fields; personal communication of F. ANDREONE).

Species known from more arid forest are M. viridis (personal observation at Montagne des

Français; see also DALY et al., 1996), M. expectata (Isalo; DaLY et al., 1996) and M. sp. 1

(Ankarana; personal communication of J. KôHLER). Only M. betsileo is known to occur

regularly outside primary habitats (personal observation on Nosy Be, Nosy Komba, Nosy

Boraha and near Maroantsetra). For the remaining species, no reliable field observations are

available to us; however, it is to be expected that M. milotympanum is restricted, as M.

aurantiaca, to swamp forests.

(4) Extent of trade of the species. — Although trade statistics do exist, a comparison of

numbers of traded specimens between species is not possible due to taxonomic confusion in

the past. In many cases, it is not possible to state which species actually was traded under a

certain name. Therefore we prefer to summarize our subjective impressions made between

Source : MNHN, Paris

VENCES, GLAW & BÔHME 65

Table 5. - Conservation status and trade of Mantella species. For each species we give: the number

of known localities; the maximum distance between the most distant known localities

(+ 20 km) measured on a 1:2,000,000 map (Carte routière, Foiben Taosarintanin’I Madagasi-

Kara [Institut National de Géodésie et Cartographie, Madagascar]) as very rough estimate of

the distribution area; the number of nature reserves in which the species is known to occur; its

known restriction to primary forest habitat (+ restricted to primary forest; - not restricted to

primary forest); the frequency in which we have seen it in trade (only our subjective

impressions between 1990-1997: - not exported in relevant numbers, + exported, ++ often

exported); and the potential attractiveness for hobbyists and the pet trade (+ not very

attractive, ++ attractive, +++ very attractive). Status is coded as follows: OK, not threatened;

CT, commercially threatened (potential danger of overcollecting exists at least locally); R,

rare; K, insufficiently known: I, indeterminate; V, vulnerable. Research needs are coded as

follows: 1, distribution; 2, taxonomic status and validity; 3, variation; 4, habitat.

Number of | Maximum | Number | Restriction | Research

Mantella species | Jocalities [locality distance| | to primary |Traded| Atractiveness | Status |" Leeds

reserves | forest

M. betsileo 17(18) |840(1220)km| 6 Es + + Ok É

M: sp. 1 5 1260 km 1 © 3 + K 2,3

M. viridis 2 <20 km © @ us un R 1

M expectata 3 340 km 1 oO + +++ R | 14

M. manery 1 0 km ü + = 4+ K |1,2,3,4

M laevigaia 5 360 km 4 + + + CT e

M. nigricans 4 80 km 3 + n + CT 2

M haraldmeïeri 6 50 km 1? + - + R 2

M. baroni 16 (18) 420 km 36) = + FF CT =

M aff. baroni 1 0km 1 7 = ++ K_[1,2,3,4

M. cowani 5 160 km o a ++ + R [1,2,3,4

| M Dernhardi 1 0km o # + + Var

M pulchra 5 320 km 1 + + + CFA |e2:3

M madagascariensis 5 260 km 1 + + + CT | 3,4

[ M. crocea 1? Okm 0 #? + + 1 [12,3,4

[M aurantiaca 4 60 km 0 + +4 + V 1,3

[ M. milotympanum 1 0 km 0 ? + +++ 1 |1,2,3,4

1990 and 1997. In these years, we monitored several times the exhibitions of specialized pet

dealers in Germany as well as in Madagascar, and thus got some indications on extent of trade

of certain species which are summarized in tab. 5. Our impressions are relatively well in

accordance with the data of GORZULA (1996), who reported the incidence of Mantella species

among a sample of 69 European hobbyists: M. aurantiaca, 15.9 %; M. madagascariensis

(probably partly referring to M. baroni) and M. crocea, 14.5 % each; M. cowani (possibly also

largely referring to M. baroni or M. madagascariensis), 11.6 %; M. pulchra, 4.4 %:, M. viridis,

2.9 %; M. betsileo and M. haraldmeïeri, 1.5 %. Also the list of BEHRA (1993) of Mantella

exported in 1990 from Madagascar does not contradict our observations: M. aurantiaca,

Source : MNHN, Paris

66 ALYTES 17 (1-2)

30.5 %; M. viridis, 14%; M. betsileo, 3 %; M. cowani (probably largely referring to M. baroni),

29 %; undetermined species, 23 %.

(5) Potential subjective attractiveness 10 hobbyists, estimated by amount of colorful

pattern and interest of breeding biology (in M. laevigata).

To summarize these data, we tried to assign status categories to Mantella species. We

followed categories used in the European CITES regulations (ANONYMOUS, 1996), except the

category CT (“commercially threatened”) which we used in a modified way as specified below,

and the abbreviation OK which we used for non-threatened species.

(1) OK (not threatened). — Not threatened at present is M. betsileo, which has a low

attractiveness, a very large distribution area, and also occurs outside primary forest.

(2) K (insufliciently known). — M. manery and M. aff. baroni are expected to belong to

one of the categories below (probably R), but basic information is lacking. M. sp. 1 does not

seem to be threatened at the moment due to its low attractiveness and apparently large

distribution area; this species, however, may be more locally restricted than M. betsileo, and

more dependent on a threatened habitat type (dry forest). Also in this case, more data are

needed.

(3) CT (commercially threatened). — This category is here used for species which may be

locally and potentially affected by overcollecting due to their high attractiveness, but which

are not yet threatened in their whole distribution area. In this category, we include M.

laevigata, M. nigricans, M. baroni, M. pulchra and M. madagascariensis.

(4) R (rare). - Species with restricted distribution areas which are not yet vulnerable or

endangered, but are at risk. In this category, we include M. viridis, M. expectata and M.

cowani.

(5) V (vulnerable). — Species likely to become soon endangered by extinction if causal

factors continue operating. At present, we only include M. aurantiaca and M. bernhardi in this

category.

(6) I (indeterminate). — Species known to be endangered, vulnerable or rare, but for

which there is not enough information to say which of the categories is appropriate. We

include M. crocea and M. milotympanum in this category.

We do not yet assign any known Mantella species to the “endangered” category (species

facing a very high risk of extinction in the wild in the near future), but some species might

move to this category within the next ten years.

Considering the lack of basic knowledge on distribution, variation, and taxonomic

status of many Mantella species, and the vulnerability of several of them (see tab. 5), we

propose the following research priorities:

(1) Clarification of taxonomy and distribution of the species of the M. aurantiaca group,

by detailed mapping of color morph occurrence and genetic studies along hybrid zones.

Single voucher specimens from each recorded locality should be deposited in publicly

available scientific collections.

(2) Habitat descriptions and mapping of M. expectata, M. cowani and M. bernhardi.

Source : MNHN, Paris

VENCES, GLAW & BÔHME 67

(3) Clarification of the taxonomic status of M. aff. baroni and of M. baroni from the

Zahamena area.

(4) Studies on variability in the M. madagascariensis group (status of M. pulchra, identity

of the “variable morph(s)” of M. madagascariensis).

(5) Formal description and naming of M. sp. 1.

(6) Comparative studies on the microhabitat and ecology of all Mantella species.

ACKNOWLEDGEMENTS

We wish to thank Franco ANDREONE (Torino; MZUT), Rainer GÜNTHER (Berlin; ZMB), Barry T.

CLARKE, Colin McCaRTHY and Edwin N. ARNOLD (London; BMNH), Berthus VAN TUuL (Amsterdam;

ZMA), Franz TIEDEMANN and Heinz GRizLrrscH (Wien; NMW), Wulf HaACKE (Pretoria; TM), and

Eugen KRAMER (Basel; NMB) who made possible the examination of specimens held in their care. Nirhy

RABiBisoA, Domoina RAKOTOMALALA, Olivier RAMILISON and Fara RANAIVOJAONA assisted during the

field work. John W. DaLy (Bethesda, Maryland) provided much useful information. Kathrin SCHMIDT

(ZFMK, Bonn) admirably organized keeping and breeding of live stocks of several Mantella species.

Franco ANDREONE (Torino), Jôrn KôHLER (ZFMK, Bonn), Giovanni SCHIMMENTI and Riccardo JESU

(Genova), Ronald NUSSBAUM (Ann Arbor), Bill Love (Alva, Florida) and Gaetano ODIERNA (Napoli)

provided unpublished data. Special thanks are due to Annemarie OHLER and Alain DUBOIS (Paris;

MNHN) for fruitful discussions and important advice. W. Ronald HEYER and one anonymous referee

critically revised an earlier draft of the manuscript.

Fieldwork of FG was made possible by a cooperation accord between the University of Antanana-

rivo (Madagascar) and the ZFMK (Bonn), and financially supported by the Deutscher Akademischer

Austauschdienst (DAAD).

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Source : MNHN, Paris

Alytes, 1999, 17 (1-2): 73-80. 73

Une nouvelle espèce

du genre Leptodactvlodon

(Arthroleptidae, Astyulosterninae) du Gabon

Annemarie OHLER

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 7500$ Paris, France

E-mail: ohler@mnhn.fr

A new species of astylosternine frog is described from Lopé Fauna

Reserve, Gabon. The new species is close to Leptodactylodon albiventris

and is characterized by its relatively long vomerine ridges, its emarginate

tongue, its brownish dorsal color with small white spots, and, in adult male,

its metacarpal with three spines and its first finger with four spines; throat

is black-violet, vent dark brown with white marbling and the underside of

thigh is vellowish. This is the first record of the genus Leptodactylodon in

Gabon; this genus is known therefore from Nigeria, Cameroon, Equatorial

Guinea and Gabon.

INTRODUCTION

Le genre Leptodactylodon Andersson, 1903 (Arthroleptidae, Astylosterninae) inclut

actuellement onze espèces et quatre sous-espèces: L. albiventris albiventris (Boulenger, 1905);

L. albiventris bueanus Amiet, 1980; L. axillaris Amiet, 1971; L. bicolor Amiet, 1971; L.

boulengeri Nieden, 1910; L. erythrogaster Amiet, 1970; L. mertensi Perret, 1959; L. ovatus

Andersson, 1903; L. perreti Amiet, 1971; L. polycanthus polycanthus Amiet, 1970; L. polycan-

thus orientalis Amiet, 1971; L. ventrimarmoratus (Boulenger, 1904). Il n’est connu que du

Cameroun, où AMIET (1980) a reconnu 1 1 espèces et 4 sous-espèces, de l’est du Nigeria, où se

rencontre une seule de ces onze espèces, et de Guinée Equatoriale, où DE LA Riva (1994) a

signalé L. albiventris. AMIET (1980) a fait une révision du genre et a discuté les relations

évolutives et biogéographiques au sein de celui-ci. Il a également proposé une interprétation

de la position de Leptodactylodon parmi les Astylosterninae.

Leptodactylodon est donc maintenant un genre bien étudié pour lequel nous disposons

d'une révision systématique de qualité. Quand nous avons observé, parmi d’autres récoltes

faites par Charles P. BLANC dans la Réserve de faune de la Lopé au Gabon, un spécimen mâle

adulte du genre Leptodactylodon, la tâche de détermination nous a semblé aisée. Nous nous

sommes ainsi très rapidement aperçue que cette petite grenouille ne peut être attribuée à

aucune des formes reconnues par AMIET (1980). Nous sommes ainsi dans la situation

embarrassante de devoir décrire une nouvelle espèce sur un spécimen unique, travail qui n'est

Source : MNHN, Paris

74 ALYTES 17 (1-2)

possible que par rapport au riche matériel étudié en détail par AMIET, et également parce que

ce spécimen est un mâle adulte et montre plusieurs caractères importants en systématique du

genre Leptodactylodon.

Leptodactylodon Andersson, 1903

Leptodactylodon Andersson, 1903: 141. - Espèce-type: Leptodactylodon ovatus Andersson, 1903, par

monotypie.

Bulua Boulenger, 1904: 262. - Espèce-type: Bulua ventrimarmorata Boulenger, 1904, par monotypie.

Synonymie: ANDERSSON, 1905: 22.

Leptodactylodon blanci sp. nov.

(ig. 1-3)

Holotype. - MNHN 1996.8876, mâle adulte, récolté par Charles P. BLANC en mars 1995.

Localité-type. —- Campement SOFORGA (0°36’30°S, 11°31°59°E), Réserve de faune de la

Lopé (voir BLANC, 1998), Gabon.

Diagnose. — Petite espèce de Leptodactylodon, mâle adulte de 22,1 mm; dents vomériennes en

deux séries assez longues atteignant le bord extérieur des choanes; langue échancrée; méta-

carpien avec trois épines, doigt I avec 4 épines; coloration dorsale brun uni avec petits points

blancs et une zone plus claire près du cloaque: gorge noir violacé avec taches blanches

indistinctes, ventre brun foncé avec marbrures blanches nettes; dessous des cuisses jaune clair

uni.

Description de l'holotype. - MNHN 1996.8876, mâle adulte de petite taille (longueur museau-

anus 22,1 mm), corps ramassé (fig. 1). Tête plus large (10,0 mm) que longue (9,2 mm); museau

arrondi, dépassant à peine le bord antérieur de la bouche, sa longueur (3,76 mm) plus longue

que le diamètre de l’œil (2,72 mm); canthus rostralis arrondi, concave et région canthale

faiblement inclinée. Espace interorbitaire plat, plus large (2,92 mm) que la largeur de la

paupière (2,01 mm), mais bien moins large que la distance internasale (3,95 mm); narines plus

proches de l'œil (1,16 mm) que de l'extrémité du museau (2,14 mm). Pupille arrondie. Tympan

présent, plutôt indistinct, son diamètre (1,00 mm) un tiers du diamètre de l'œil, aucun vestige

d’ocelle pinéal. Crêtes vomériennes (fig. 2) présentes avec une dizaine de petites dents chacune,

perpendiculaires à l’axe du corps, postérieures aux choanes, et touchant celles-ci, les rangées

de dents bien plus longues que la distance les séparant. Langue de taille moyenne, en forme de

cœur avec une petite échancrure. Repli supratympanique peu distinct, de l'œil à l'épaule.

Bras court; avant-bras (5,64 mm) plus long que la main (5,38 mm) (fig. 3b); doigts plutôt

longs et plutôt larges; doigt I égal au IT, le IT plus court que le IV, le doigt II le plus long

(3,05 mm); extrémités de tous les doigts pointues, non élargies, avec ventouses portant des

sillons ventro-latéraux; sillons proximaux absents; bords dermiques le long des doigts I à II

des 2 cotés; palmure absente; tubercules sous-articulaires distincts, arrondis, simples, tous

présents; tubercules métacarpiens ovales, bien développés; deux tubercules palmaires ovales,

bien développés: tubercules surnuméraires à la base des doigts absents.

Source : MNHN, Paris

OHLER 75

Fig. 1. — Leptodactylodon blanci, holotype MNHN 1996.8876, mâle adulte. À gauche, vue dorsale ; à

droite, vue ventrale.

e

Na

Jambe trois fois plus longue (10,2 mm) que large (3,5 mm), plus courte que la cuisse

(10,7 mm), et que la distance entre la base du tubercule métatarsien interne et l'extrémité de

l'orteil IV (11,5 mm) (fig. 3a); longueur relative des orteils: 1 < II < V < III < IV. Extrémités de

tous les orteils pointues, faiblement élargies, portant des ventouses avec sillons ventro-

latéraux, sillons proximaux absents. Palmure absente. Bord dermique sur l’orteil V absent.

Tubercules sous-articulaires nets, ovales; tubercule métatarsien interne court, proéminent,

longueur (2,20 mm) contenue 2,5 fois dans la longueur de l’orteil I (5,44 mm); tubercule

métatarsien externe absent; tubercules surnuméraires absents; tubercules tarsiens absents; pli

tarsien absent.

Source : MNHN, Paris

76 ALYTES 17 (1-2)

a

Fig. 3. — Leptodactylodon blanci, holotype MNHN 1996.8876, mâle adulte. (a) pied, vue ventrale; (b)

main, vue ventrale.

Dessus et coté de la tête ainsi que dos lisses; flancs avec pustules effacées; membres

antérieurs lisses; membres postérieurs avec pustules peu nettes; face inférieure lisse; présence

de spinules cornées au bord antérieur de la gorge.

Dos, dessus et côté de la tête, haut du flanc et avant-bras brun foncé avec petits points

blancs, barre blanche reliant le tiers antérieur des yeux et quart postérieur du dos brun plus

Source : MNHN, Paris

OHLER 77

clair; dessus de la cuisse et partie proximale de la jambe brun foncé avec petits points blancs,

partie distale de la jambe et tarse beiges avec points bruns, séparée de la partie brun foncé par

une large bande brun noirâtre; partie postérieure de la cuisse blanchâtre couverte de mouche-

tures brunes denses. Gorge noir violacé avec quelques taches blanches indistinctes; bord de la

gorge brun avec taches blanches; poitrine et ventre brun avec vermiculations blanches nettes.

Dessous des cuisses jaunâtre uniforme.

Caractères sexuels secondaires mâles. — Trois épines nuptiales sur le métacarpien et quatre sur

le doigt I, de couleur noire; petites spinules transparentes sur le bord antérieur de la gorge; sac

vocal externe unique; ouvertures arrondies de chaque coté en arrière dans la bouche.

Ecologie. — BLANC (1998) décrit le milieu d’origine de l’holotype comme suit: “forêt den:

sous-bois; ruisselet; camp abandonné; pistes”. Sur le même site, 8 autres espèces d’Amphi-

biens ont été capturées: A/exteroon s] rthroleptis sylvaticus (Laurent, 1954); Bufo latifrons

Boulenger, 1900; Conraua crassipes (Buchholz & Peters, 1875); Dimorphognathus africanus

(Hallowell, 1857); Phrynobatrachus cornutus (Boulenger, 1906); Phrynobatrachus sp.; Ptycha-

dena perreti Guibé & Lamotte, 1958.

Etymologie. - L'espèce est dédiée à Charles P. BLANC qui a récolté le spécimen et nous l’a

confié pour description.

DISCUSSION

AMIET (1980) présente une clef de détermination du genre Leptodactylodon que nous

avons reprise ici (Annexe 1), en insérant la nouvelle espèce, sur la suggestion d’un lecteur qui

pensait qu’il serait souhaitable de la mettre à disposition d’un plus large public, la clef

originale étant parue dans un journal absent dans de nombreuses bibliothèques.

Le genre Leptodactylodon peut être divisé en quatre groupes d'espèces: (1) un groupe

(groupe de L. mertensi) qui comporte des espèces dont les mâles portent des protubérances

axillaires et qui se caractérisent par l’atrophie des dents vomériennes (L. mertensi, L. erythro-

gaster, L. perreti, L. axillaris); (2) un groupe (groupe de L. bicolor), comportant une seule

espèce, caractérisée par l’atrophie des dents vomériennes et l’absence des protubérances

axillaires; (3) un groupe (groupe de L. ornatus), comportant une seule espèce, montrant une

langue non-échancrée et dont les mâles présentent une seule épine sur le métacarpe: et (4) un

groupe (groupe L. ovatus) montrant plusieurs épines sur le métacarpe et une langue échancrée

(L. ovatus, L. boulengeri, L. ventrimarmoratus, L. albiventris, L. polycanthus).

La nouvelle espèce se place dans ce dernier groupe. Dans sa clef, AMIET (1980) y a

distingué deux sous-groupes par la taille des espèces et par la formation de leurs crêtes

vomériennes. On ne peut pas assigner L. blan ilement à l’un des deux sous-groupes. La

nouvelle espèce est certes de petite taille, mais par l'extension de ses crêtes vomériennes elle

semble intermédiaire. Sa livrée dorsale et sa taille ressemblent à celles de L. bicolor, mais elle

se distingue de cette espèce par la présence de dents vomériennes et par l'absence de tout

dessin sur les cuisses; les taches blanches sur le ventre sont plus denses et de taille inférieure

Source : MNHN, Paris

78 ALYTES 17 (1-2)

chez L. blanci. Cette livrée dorsale “bicolore” se rencontre également dans la sous-espèce L.

albiventris bueanus. Morphologiquement, cette forme semble plus proche encore de la nou-

velle espèce, notamment par la présence de dents vomériennes et son nombre d'épines

métacarpiennes de 3. Les deux formes se distinguent néanmoins par la coloration ventrale:

chez les deux sous-espèces d’alboventris, seule la gorge est foncée avec des taches plus claires.

Ce spécimen témoigne de la présence de Leptodactylodon au Gabon et amène à augmen-

ter l'aire de distribution du genre vers le sud. Ainsi cette aire inclut actuellement le Nigéria, le

Cameroun, la Guinée Equatoriale et le Gabon.

RÉSUMÉ

Une nouvelle grenouille de la sous-famille Astylosterninae est décrite de la Réserve de

faune de la Lopé au Gabon. Cette nouvelle espèce se rapproche de Leptodactylodon albiventris

et se caractérise par des crêtes vomériennes assez longues, sa langue échancrée, la présence de

trois épines sur le métacarpe, la présence de quatre épines sur le premier doigt et le dos de

couleur brune avec des petits points blancs; la gorge est violet noirâtre, le ventre brun foncé

avec des marbrures blanches et le dessous des cuisses est jaunâtre. Ce spécimen représente la

première mention du genre Leptodactylodon au Gabon; le genre est donc connu du Nigéria,

du Cameroun, de la Guinée Equatoriale et du Gabon.

REMERCIEMENTS

Charles P. BLANC a collecté le spécimen et me l’a confié pour étude. Je remercie Alain

Dugoïis pour ses remarques, comme toujours très qualifiées, sur le manuscrit.

RÉFÉRENCES BIBLIOGRAPHIQUES

Amie, J.-L. 1970. - Espèces nouvelles ou mal connues de Leprodacrylodon (Amphibiens Anoures) de la

Dorsale camerounaise. Ann. Fac. Sci. Cameroun, 5: 57-81.

—— 1971 - Leptodactylodon nouveaux du Cameroun. Ann. Fac. Sci. Cameroun, 7-8: 141-172.

ision du genre Leptodactylodon Andersson (Amphibia, Anura, Astylosterninae). Ar

Fac. Sci. Yaoundé, 27: 69-224.

ANDERSSON, L. G., 1903. - Neue Batrachier aus Kamerun von den Herren Dr. Y. Sjôstedt und Dr. S.

Junger gesammelt. Verh. Zool. Bor. Ges. Wien, 53: 141-145.

— 1905. — Batrachians from Cameroon collected be Dr. Y. Sjôstedt in the years 1890-1892. 4rk. Zool.,

2) 20:

BLANC, C., 1998. — Rapport scientifique de mission. Mission ECOFAC GABONIAGRECO-GEIE. Reptiles

et Amplhibiens. Réserve de faune de la Lopé (09 mars 1995 - 09 avril 1995). Montpellier: 1-6

BouLenGer, G. A., 1904. — Descriptions of two new genera of frogs of the family Ranidae from

Cameroon. Ann. Mag nat. Hist. (7), 13: 261-262.

Source : MNHN, Paris

OHLER 79

1905. Descriptions of new West-African frogs of the genera Perropedetes and Bulua. Ann. Mag. nat.

Hist. (D), 15: 281-283.

DE La Riva, L., 1994. — Anfbios anuros del Parque Nacional de Monte Alen, Rio Muni, Guinea

Ecuatorial. Rer. esp. Herp., 8: 123-139.

NtEDEN, E, 1910. - Neue Reptilien und Amphibiens aus Kamerun. Arch. Naturgesch., 76: 234-246.

PerReT, J.-L., 1959. - Etudes herpétologiques africaines. Bull. Soc. neuchât. Sci. nat., 82: 247-253.

ANNEXE I

CLEF DE DÉTERMINATION DES ESPÈCES ET SOUS-ESPÈCES DE LEPTODACTYLODON

1. Dents vomériennes bien développées, disposées en bandelettes de forme variable

- Dents vomériennes atrophiées, formant 2 petites saillis arrondies, ou absentes .

2. Dents vomériennes disposées en accent circonfiexe; langue non échancrée à son extrémité; une bande

dorsale sub-rectangulaire foncée; côtés de la tête, dessous des pieds et marques para-anales foncées; 1

seule épine métacarpienne chez le mâle (groupe de L. ornatus) .…

- Dents vomériennes en bandelettes rectilignes ou arquées; langue échancrée; pattern dorsal et latéral peu

marqué ou indistinct; plusieurs épines métacarpiennes chez le mâle (groupe de L. ovatus) 4

3. Face ventrale, membres postérieurs exceptés, vermiculée de blanc et noir. L. ornatus ornatus

Face ventrale, y compris les membres postérieurs, noire avec de grosses taches blanches plus ou moins

arrondies L. ornatus permaculatus

4. Taille grande (35 à 44 mm env.); dents vomériennes en longues bandelettes dépassant les choanes vers

re ‘extérieur et étroitement rapprochées; face ventrale, au moins la gorge, ponctuée ou marbrée de blanc et

; 3, 4 ou 5 épines métacarpiennes chez le mâle 5

- Taille médiocre (au max. 30 mm chez les femelles, en moyenne 25-26 mm chez les mâles): dents

vomériennes en bandelettes courtes, écartées et ne dépassant pas ou à peine les choanes vers l'extérieur;

livrée ventrale souvent uniforme ou brun avec vermiculations blanches; 2-3, 4 ou 5-6 épines métacarpien-

nes chez le mâle 8

5. Région inguinale marbrée ou largement maculée de sombre sur fond clair, ce pattern pouvant s'étendre

à la face antérieure des cuisses (L. ovatus) 6

— Région inguinale non marbrée 7

6. Face ventrale entièrement mouchetée de clair sur fond sombre ......... .. L ovatus ovatus

— Gorge seule mouchetée de clair sur fond ombre, le reste de la face ventrale 1 rose in vivo, avec de rares

DORCUATIONS LATE NÉE R TR PTE APR SNA tire cr ere L. ovatus orientalis

7. Face ventrale entièrement couverte de vermiculations noires et blanches; fascie dorsale peu apparente,

réduite à un petit triangle céphalique . L. ventrimarmoratus

- Face ventrale entièrement claire, rose ou jaunâtre in vivo, scie dorsale assez nette,

en triangle étiré jusqu’au milieu du dos . L. boulengeri

8. Gorge des mâles présentant de gros plis paramandibulaires ou sac vocal très devéloppé; pas d’hyper-

trophie brachiale ni de spinosité pectorale; 2, 3 ou 4 épines métacarpiennes chez le mâle . ARETO

— Pas de plis gulaires chez les mâles; spinosité pectorale très développée; hypertrophie brachiale et 5-6

épines métacarpiennes chez le mâle (L. polyacanthus) .… ANA R il

9. Taille moyenne 26 mm chez les mâles; dessous, sauf la gorge, d'un rose saumon uniforme; gorge des

mâles présentant de gros plis paramandibulaires: pas d'hypertrophie brachiale ni de spinosité pectorale:

2 ou 3 épines métacarpiennes chez le mâle (L. albiventris).… mr … 10

= Taille du seul mâle connue 22 mm: dessous brun avec vermiculations blanches; sac vocal très développé

chez les mâles: pas d'hypertrophie brachiale ni de spinosité pectorale; 3 épines métacarpiennes

ie Re a LEA … L blanc

Source : MNHN, Paris

80 ALYTES 17 (1-2)

10. Orteils munis de prolongements acuminés souples; dessus entièrement sombre sauf 2 fascies para-

anales oranges; 2 épines métacarpiennes chez le mâle L albiventris albiventris

— Orteils sans prolongements; partie postérieure du dos, base des cuisses, moitié distale des jambes et base

des tarses éclaircis, beige in vivo; 3 épines métacarpiennes chez le mâle. … L. albiventris bueanus

11. Face ventrale plus ou moins assombrie, mais dépourvue de points ronds éclaircis (bleutés in vivo);

triangles d’épines pectorales larges, ne se joignant pas sur le milieu de la poitrine .

L. polyacanthus polyacanthus

[Face ventrale ponctuée de clair sur un fond assombri; triangles d'épines pectorales étroits, se joignant

sur le milieu de la poitrine L. polyacanthus punctiventris

12. Face dorsale bi- ou tricolore; face ventrale noire avec de grosses taches blanches arrondies ou

bacilliformes; pas de protubérances axillaires; pas d’hypertrophie brachiale; 2-3 épines métacarpiennes

chez le mâle (groupe de L. bicolor) L. bicolor

- Face dorsale en générale unicolore (ou mouchetéc: L. axillaris); face ventrale non tachée de blanc sur

fond noir; des protuberances axillaires et une forte hypertrophie brachiale chez le mâle (groupe de L.

mertensi) 13

13. Dessus brun foncé (mâles) ou rouge ou rougeâtre (femelle); dessous vivement coloré avec pigment

rouge plus ou moins étendu; 3-4 épines métacarpiennes chez le mâle 14

— Pas de dichromatisme sexuel; dessous blanchâtre avec taches nébuleuses sombres plus ou moins

étendues, aucun pigment rouge; 2 ou 5-6 épines métacarpiennes chez le mâle 15

14. Dessous à pigmentation très contrastée, en général blanc avec grosses marbrures noires et bords rouge

vif .. L. mertensi

- Dessous finement moucheté de brun noir sur fond rougeâtre, rosé ou blanchâtre (mâles) ou rouge cerise

ou rouge et blanc (femelles) . L. erythrogaster

15. Taille petite (moyenne des mâles: 23 mm; femelles: 28 mm); dessus brun ou roussâtres sans taches

jaunes; tégument peu granuleux; protubérances axillaires subconiques et 2 épines métacarpiennes chez le

mâle . .. L perreti

= Taille grande atteignant 38 mm chez les mâles qui sont plus grands que les femelles (27-33 mm); dessus

brun violacé foncé, avec souvent des mouchetures jaunes; tégument très granuleux dorsalement; protu-

bérances axillaires aplaties et tronquées et 5-6 épines métacarpiennes chez le mâle ..… L. axillaris

Corresponding editor: John C. POYNTON.

© ISSCA 1999

Source : MNHN, Paris

Alytes, 1999, 17 (1-2): 81-100.

81

Miscellanea nomenclatorica batrachologica.

With more than 750 species (GLAW et al.,

Ranidae” is one of the largest amphibian higher taxa. Its taxonomy

“family

19. Notes on the nomenclature

of Ranidae and related groups

Alain DUBOIS

Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The need of “working taxonomies”, as tools providing a framework for

alpha-taxonomic revisionary works and hypotheses for phylogenetic analy-

ses, is pointed out, especially in groups with wide distribution and high

number of species. Even during the transitional period, non-ambiguous

communication between zoologists requires that use of names for taxa

strictly follows the international rules of zoological nomenclature. Several

cases of recent nomenclatural problems are pointed out in the “family

Ranidae”. Rediscovery of the generic name Chilixalus Werner, 1899

provides an opportunity for discussing several careless treatments of the

generic and infrageneric taxonomy of frogs of the “genus Rana” by recent

authors. The name Indiraninae Blommers-Schlôsser, 1993 is shown to be

an invalid junior synonym of Ranixalinae Dubois, 1987, and, on this

occasion, the fact that family-group names are regulated by the rule of

priority is reminded once again. In order to help knowing the valid name

among several names published simultaneously, in the case they are

lered subjective synonyms, two tables of first-reviser actions in the

Ranidae and related groups are presented. Finally, discussion of the

nomenclatural status of the name “Rana duboisi” recently published by

Emerson & War (1998) allows to point to the problems posed by the

publication of data taken from unpublished manuscripts by colleagues,

either submitted to review by an editor, or privately communicated by the

author or another person. This case is also the basis for the discussion of

two more general questions, which are likely to appear again on several

occasions in the years to come: is a species name rendered nomenclaturally

available by publication, either (1) of a Genbank catalogue number, or (2) of

a cladogram including this species? The answer to both questions is clearly

no”, at least under the current Code. Allocation of names to taxa is not

based on definitions, diagnoses or descriptions, but on the taxonomic

allocation of name-bearing type-specimens: the recent proposal of at-

taching the names to “phylogenetic definitions of taxon names” is therefore

based on a major misunderstanding and entertains an unnecessary confu-

sion between taxonomy and nomenclature, as the current nomenclatural

system is liable to accomodate any kind of taxonomy, including “phyloge-

netic”’ ones.

1998) distributed almost worldwide, the

still very

problematic, if not really “in a state of chaos”? (DUELLMAN & TRUEB, 1985: 544). Resolution

of all the problems it raises will be a long task, as it will require a high number of works of

Source : MNHN, Paris

82 ALYTES 17 (1-2)

various kinds (morpho-anatomical, molecular, cytogenetic, bioacoustic, ethological,

ecological), both at local scale and at world scale, and dealing both with alpha-taxonomy

and with higher classification. Only when this is done can we hope to have a reasonably good

knowledge of the species of the family and understanding of their phylogenetic rela-

tionships. Given the high number of species concerned, and the extremely large distri-

bution of the group, it is impossible for any researcher to work on the whole of the family

at once. Before applying any technique or carrying out any survey, choices must be made

among the hundreds of species of the family. This choice can be made along three major lines:

(1) on a geographical basis, i.e. studying the ranids of a given region of the world; despite the

fact that this approach is clearly unstatisfactory (see e.g. DuBois, 1981, 1987a, 1992), for

material reasons this has been the case of the vast majority of works dedicated to the

taxonomy of this family until now; (2) according to the availability of specimens in one or

several museum or other collection(s), which may be a little better when rich, largely

representative collections are used, but remains unsatisfactory in most cases; (3) on a

provisional taxonomic basis, which is clearly the best starting point for any revisionary

taxonomic work (see e.g. MAYR, 1981). Some zoologists (e.g., INGER, 1996) seem to think that

a taxonomy is only a result, and that taxonomies should only be established, or changed, when

biologists have “final” data on the (cladistic or other) relationships between the species. This

is a very reductory and misleading view of the rôle of taxonomy in biology, for two reasons at

least:

(1) Most data on which taxonomies are based are conventional (i.e., based on sub-

jective choices, e.g. as to which kind of information should be provided by the classifi-

cation) and/or hypothetical (e.g., cladograms as hypotheses of cladistic relationships). As a

consequence, no taxonomy is or will ever be the “final” one, for any group of living

beings. Asking to postpone the establishment or change of taxonomies until we have “final”

data is just a way to say that no taxonomy can ever be established, or that the existing

taxonomies (often inherited from “tradition”, both in its best and worse senses), should never

be changed, for reasons of “stability of nomenclature”. However, no nomenclature can ever

be completely stable, if taxonomy is to remain a living, i.e. evolutive, science (see DUBOIS,

1998).

(2) Such requests ignore one of the functions of taxonomies, 1.e. their “heuristic value”:

a taxonomy is not only a result, it is also a starting point. It is a hypothesis, that can be tested

by further research and modified. This is particularly true in speciose and complex groups

with large distributions, that cannot be comprehensively studied at once, such as the Ranidae:

in these groups, at least if one really wishes to improve the existing taxonomies, provisional

groupings as “phenetic taxa” (such as e.g. the “phenetic groups” recognized in toads of the

genus Bufo by some authors: DUELLMAN & SCHULTE, 1992; DuBois & OHLER, 1999) are

necessary to have a “working taxonomy”. Such groups of a reasonable size, chosen neither on

a geographical nor on “availability” grounds, will allow real, although partial, revisionary

works. This is demonstrated by the fact that most of those who recently really tried to improve

the taxonomy of ranids above the species level (and not only to “comment” on the work of

others), had to start from subsets of the whole family, which, although they might not have

stated it, were those proposed as “working taxa” by previous authors (see e.g. EMERSON &

BERRIGAN, 1993; EMERSON, 1996; EMERSON & WaRD, 1998).

Source : MNHN, Paris

Dusois 83

The request for stability of taxonomies and nomenclatures, that some authors (e.g.

INGER, 1996) wish to apply to the Ranidae!, is relevant only for some zoological groups, either

of very small size (e.g., higher primates) or already very much studied (e.g., birds), for which

an enormous wealth of information is already available, and in which competing taxonomic

schemes only or mostly depend on subjective choices as to the major criteria to be taken into

account in the building of classifications (classification or “cladification”: MAYR, 1997;

Dusois, 1997), on different weightings of the characters, etc. But in poorly known groups like

the Ranidae, we strongly need provisional, working taxonomies, to really help progress of

research and to guide future alpha-taxonomic works and phylogenetic studies. Such tempo-

rary taxonomies can be progressively modified and replaced by better ones, as information

becomes available, but just to obtain this information may be very time-consuming. Pending

its obtention, zoologists cannot be left in a “non-taxonomic land”. They need “working

taxonomies” and “working nomenclatures”. In such groups, a fascination, or a quasi-

religious respect, for “stability” of taxonomy and nomenclature, can work as a break against

increase and improvement of our knowledge. Of course, in such enormous groups as the

Ranidae, where no researcher in the world can have access to all, or even to a high proportion

of, the described species, and where many species are known only by a very low number of

specimens, sometimes only in the adult stage (the tadpoles being unknown), such a provision-

al taxonomy can only be based on a heterogeneous combination of various sources of

information: some based on field work, some on detailed anatomical studies, some on

examination of specimens limited to external characters, and some on data published by

previous authors. Because of this unavoidable diversity of sources of information, the data set

is bound to be incomplete, as some character states (particularly those requiring dissection

and anatomical study, or those of tadpoles) will be known for some taxa only: such data

cannot therefore be used to build up a matrix and carry out a phylogenetic analysis, but can

allow partial, provisional definitions of phenetic groups, diagnosed by characters shared only

by their included species and that can in a first step be hypothesized to be synapomorphies of

the latter. Of course, such a work is likely to include some mistakes, but then a useful

contribution of subsequent workers will be to correct these and improve the provisional

1. Actually, the motivation for writing this paper (INGER, 1996) are difficult to understand. Although this author

ha blished numerous papers on the Oriental, Asiatic and African frogs for more than half a century, he has

never shown real interest in the phylogeny and supraspecific taxonomy of these groups, as he never provided a

significant contribution to this field but merely perpetuated BOULENGER'S ideas and schemes in this respect.

Some of the information provided in his recent paper could have been proposed as a constructive contribution

to the taxonomy of ranids, and will no doubt be used as such in the future. This is indeed the kind of information

I was expecting when I wrote my “proposals” (Durois, 1992), which are clearly a basis for discussion and

improvements, not a “final system”: although my paper was the result of research over a 20-year period, itis clear

that I could not have examined all ranid groups worldwide. However, instead of proposing these comments as

positive elements for correcting and improving my proposals, INGER'S (1996) paper is only negative and

ag ; and does not offer alternate proposals but “waiting for more data”, to paraphrase KOTTELAT'S (1997:

2,4) nice words (see also DuBois & OHLER, 1999: 135). TI some doubts on the real intentions of its author

(and of the editors of the journal where it was published). The aim of this publication was clearly not to open a

debate (which could well have been published in the “points of view” of this journal): this is why I did not think

itworthwhile to write a reply. Replies will come slowly but surely as the much needed works on the ranids are

progressing (see e.g. MARMAYOU et al.. in press). In the meantime, following INGER's “traditional” taxonomy is

not likely to help authors who are really interested in the relationships within the Ranidae, and who need

subgroups to deal with the hundreds of species still allocated by some to the genus Rana: presenting these species

in a publication by alphabetical order of specific names is highly misleading, as is the use of arbitrary groups

based on grossly incomplete data, as well exemplified by the recent “sections” recognized by Tin et al. (1995) in

this genus, which ignore many important pieces of information published after BOULENGER'S Works.

Source : MNHN, Paris

84 ALYTES 17 (1-2)

taxonomy, rather than reject it altogether without using its good parts. This is the way science

usually progresses, particularly in biology (see e.g. MAYR, 1982, 1997).

Taxa, even provisional, must be named (Dugois, 1988), and the fact that they are

provisional does not mean that their nomenclature should not be rigorous. Strictly following

the rules of the /nternational Code of Zoological Nomenclature (ANONYMOUS, 1985; quoted

below as “the Code”) is a guarantee of non-ambiguous, stable, automatic and universal

allocation of names to taxa, which is much more important than the (highly praised by some)

“stability of taxa and names”. Among these rules, the rule of priority is an important one. À

recent tendency has developed among some zoologists (e.g., SAVAGE, 1990a-b, 1991; Bocx,

1994) and in decisions of the International Commission on Zoological Nomenclature (quoted

below as “the Commission”) to severely limit its application in zoology. In so doing, the

Commission has not properly played its rôle of “Keeper of the Law” (DuBois & OHLER, 1997:

299), and has encouraged neglect of all the rules by zoologists. No doubt such attitudes

contribute to the current weakening of the binding legislative status of the Code for the

establishment of the valid names of taxa in the eyes of many taxonomists. Such a movement

may have important negative consequences in the long run regarding the existence of a unique

international nomenclatural system, and therefore the unity and universality of zoology as a

science (DUBOIS, in preparation).

In the recent years, and largely as a result of this tendency, a number of nomenclatural

problems in amphibians have been pointed out (see e.g. DuBois, 1987a-b, 1995, 1998; Dugois

& OHLER, 1995, 1998). The purpose of this paper is to present a few new such problems in the

Ranidae and their solutions, within the frame of the current taxonomy of this family. This

does not preclude the possibility that the nomenclature of the taxa discussed below may have

to be changed in the future, as the taxonomy of these groups evolves: nomenclature being at

the service of taxonomy (and not the reverse), names will necessarily have to change as taxa

are modified, suppressed or created. The particular cases presented below will also provide the

opportunity to discuss several more general problems of zoological nomenclature and of

scientific publications.

CuiLiXALUS WERNER, 1899

ScaMipr (1857: 11) published a preliminary diagnosis of /xalus warszewitschii. Shortly

after, the same author (SCHMIDT, 1858: 241-242, 258, pl. 1) provided a more detailed

description of this species, for which he gave a precise type-locality, now situated in Panama

(see Hizuis & DE SÀ, 1988: 15). For the name of the species, he then used two spellings: on

page 258, the correct original spelling /xalus warszewitschit, and on page 241, on two

occasions, the spelling /xalus warschewitschii, which must therefore be viewed as an incorrect

subsequent spelling, as noted by HiLis & DE SA (1988: 16). Most subsequent authors (e.g.,

DUNN, 1931:416; TAYLOR, 1952: 896; GORHAM, 1974: 153; FRosT, 1985: 520) used this latter

spelling, until HiLis & DE SA (1988: 1) resurrected the correct original spelling.

Scamipr (1858: 242) stated that FITZINGER had privately suggested to him that this

species should be placed in a new genus, but he refrained from doing so until field work could

Source : MNHN, Paris

Duois 85

bring more data about it. However, another, subsequent author did not have as many scruples

and created a nominal genus for this species without having additional information: WERNER

(1899: 117) proposed the new generic name Chilixalus for the nominal species “/xalus

warszewiczii Schmidt”. He had apparently not seen the publications of SCHMIDT (1857, 1858)

on this species, as he stated that he had found this name on a bottle in the Krakow Museum

containing a frog specimen from “Neu Granada”. Fortunately, he mentioned the collection

number (1006) of this specimen, which is the same as that reported by HiLis & DE SA (1988:

15) for ScamibT's (1857) holotype, so that there is no doubt about the fact that WERNER (1899)

actually dealt with the same species as SCHMIDT (1857, 1858). WERNER'S (1899) spelling of the

specific name, repeated twice in his paper, was clearly intentional, and should be regarded

either as an unjustified emendation of SCHMIDT's original name, or, because of WERNER’S

ignorance of SCHMIDT’s publications, as the name of a new nominal species: in both cases, the

name Chilixalus warszewiczii has an independent status in nomenclature and is a junior

objective synonym of Jxalus warszewitschii Schmidt, 1857.

Although published by a well-known zoologist in a major journal, the name Chilixalus

has been almost completely forgotten by subsequent authors. It was mentioned by BOULEN-

GER (1900b: 28; 1910: 152) and NEAVE (1939: 691), but ignored altogether in all major works

dealing with the classification of Ranidae and Rhacophoridae, or with the ranids of Central

America: e.g. GÜNTHER (1900), BOULENGER (1920), AHL (1931), NOBLE (1931), TAYLOR

(1952), GorHAM (1974), Dugois (1981, 1992), DUELLMAN & TRUEB (1985), FROST (1985) and

Huzuis & DE SA (1988). Although close by its spelling to the generic names Chirixalus

Boulenger, 1893 (Ranidae, Rhacophorinae) and Callixalus Laurent, 1950 (Hyperoliidae,

Hyperoliinae), the name Chilixalus differs from these names by one or two letters, and is

therefore not their homonym. The type-species of this nominal genus is now considered a

member of the genus Rana Linnaeus, 1758, as Rana warszewitschii (Schmidt, 1857) (HiLis &

DE SÀ, 1988).

Fortunately, rediscovery of the name Chilixalus does not have disturbing effects on

nomenclatural stability, whatever the classification scheme chosen. Three classification

schemes are currently applied by different authors to the group of ranids including the

nominal species /xalus warszewitschii Schmidt, 1857: (1) for authors who do not recognize

subgenera in Rana, it is a member of the Rana palmipes group; (2) for some authors, it is a

member of the subgenus Lithobates Fitzinger, 1843 of Rana; (3) according to the provisional

classification of ranids proposed by DuBois (1992), it is a member of the subgenus 7ryphe-

ropsis Cope, 1868 of Rana.

Unfortunately, a certain amount of taxonomic and nomenclatural vagueness applies to

several recent works dealing with the “Lithobates section” (Dugois, 1992: 323, 329) of the

genus Rana. Although the title and abstract of their paper only referred to the “ Rana palmipes

group”, Hizis & DE SA (1988: 16-17) suddenly mentioned a “subgenus Lithobates”, without

stating its author, date and content, and without reference to a publication where this would

appear. In support of this use, they gave two references, one to a paper (HiLLis & Davis, 1986)

where the “subgenus Lithobates”" was not at all mentioned, and one to an unpublished thesis

(His, 1985), where a subgenus “ Lithobates Fitzinger” (without date) was briefly mentioned

(p. 266-267), without any reference allowing to identify this name. As a matter of fact, as of

1988, the status of the name Lithobates had been discussed in only one publication (DuBois,

Source : MNHN, Paris

86 ALYTES 17 (1-2)

1981: 249-250), not quoted by these authors, where this name was considered a synonym of

Rana. Therefore, Hizzis & DE SÀ (1988) resurrected the generic name Lithobates for a new

subgenus for which they did not provide a diagnosis, and without discussing, even briefly, the

status of the other subgenera recognized until then in Rana (DuBois, 1981, 1987a).

The taxonomy of the American species, species-groups and subgenera currently referred

to the genus Rana will not be definitively clarified until their relationships with non-American

(i.e., European and East Asian) species of this “genus” are studied in detail, and any current

taxonomic scheme can only be considered as a provisional, working taxonomy. I provide

below synonymies of the subgenera provisionally recognized by DuBois (1992: 329-331) in his

“Lithobates section” of the genus Rana: these are phenetically diagnosable groups for which

a hypothesis of cladistic relationships has been proposed by HizLis & DE SÀ (1988: 18). Under

this scheme, the generic name Chilixalus appears as a junior subjective synonym of Tryphe-

ropsis. Under the other classification schemes mentioned above, it is either a junior subjective

synonym of Lithobates or a junior subjective synonym of Rana.

Lithobates Fitzinger, 1843

Lithobates Fitzinger, 1843: 31. - Type-species by original designation: Rana palmipes Spix, 1824: 29.

Ranula Peters, 1859: 402 (nec Schumacher, 1817: 77). — Type-species by monotypy: Ranula gollmerit

Peters, 1859: 402.

Pohlia Steindachner, 1867: 15. — Type-species by monotypy: Rana palmipes Spix, 1824: 29.

Sierrana Dubois, 1992

Sierrana Dubois, 1992: 33

385.

— Type-species by original designation: Rana sierramadrensis Taylor, 1939:

Trypheropsis Cope, 1868

Tiypheropsis Cope, 1868: 117. — Type-species by original designation: Ranula chrysoprasina Cope, 1866:

129.

Levirana Cope, 1894: 197. - Type-species by monotypy: Levirana vibicaria Cope, 1894: 197.

Chilixalus Werner, 1899: 117. - Type-species by monotypy: Chilixalus warszewiezit Werner, 1899.

Laevirana Günther, 1900: 206. - Unjustified emendation of Levirana Cope, 1894.

Zweifelia Dubois, 1992

Ziweifelia Dubois, 199

416.

= Type-species by original designation: Rana tarahumarae Boulenger, 1917b:

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Dusois 87

OTHER INCONSISTENCIES IN RECENT TREATMENTS OF RANID TAXONOMY

AT GENUS AND SUBGENUS LEVEL

The case of the “subgenus Lithobates” just discussed is not unique in the recent literature

dealing with ranid taxonomy. To tell the truth, it is difficult in many cases to be sure of what

infrageneric taxonomy is indeed followed by many current authors within the genus Rana,

which clearly points to a general uneasy feeling in this respect. Thus, in a paper dealing with

some Asian frogs, EMERSON & BERRIGAN (1993) mentioned a “subgenus Limnonectes (Fitzin-

ger)” in their title, but did not clearly state in the text which species they included in this

subgenus, nor in other “subgenera” of their very comprehensive “genus Rana”. They

introduced (p. 23) the new combination “Rana {Occidozyga) cyanophlyctis” without any

comment, which seems to imply that they recognized a new subgenus Occidozyga in the genus

Rana. However, they did not propose a diagnosis or definition of the latter, nor did they define

its content. Did they mean that all species shown in the consensus tree of their figure 8 should

be included in this subgenus, or should the latter be understood as comprising only some

species of this tree, namely /ima (type-species of Occidozyga: see DUBOIs, 1981), limnocharis,

cancrivora and cyanophlyctis? This information is not to be found in their paper. It is

surprising to see publication of such non-professional treatments of taxonomic and nomen-

clatural matters in a well-known herpetological journal, but this is only one example of a

recent tendency for zoological publications, even of high level, to ignore the basic taxonomic

and nomenclatural rules.

In a later paper, EMERSON (1996: 279) first expressed high concern for the proper use of

scientific names in biological publications: “until a proper systematic treatment is completed,

it seems premature and potentially confusing to use the name Limnonectes in the literature. In

this paper, members of that group will be referred to as the fanged frogs and their relatives.”

However, a few pages below in the same article, she seemed to have forgotten these good

resolutions, as she presented quite confusing information. In page 281, she wrote: “Egg size

was measured in adult females of 19 species of fanged frogs and 16 species of outgroup ranids

belonging to the genus Hylarana”. In the legend of her figure 2 (p. 282), “outgroup species of

the genus Æylarana” appear again, but the text of the same page mentions “outgroup ranids

belonging to the subgenus Hylarana”. Genus or subgenus? Actually, until now, while most

zoologists working in Africa consider Hylarana Tschudi, 1838 as a genus, no author working

on Asian frogs has treated Hylarana as a full genus, except FEret al. (1991) and YEet al. (1993),

in two works not cited in the References of EMERSON’S (1996) paper, so that treating Oriental

Hylarana as a full genus would seem to have required at least a short comment. Furthermore,

Dusois (1987a: 42) pointed out that Asian species of “ Hylarana” did in fact represent several

clearly distinct groups, and later (DUBoIs, 1992) distributed these species in several sections

and subgenera of the genus Rana. It would therefore be important to know which are the “16

species of outgroup ranids belonging to the genus Hylarana” studied. Unfortunately, EME

soN’s (1996) paper does not contain a list of the species, not to say of the specimens, examined.

Contrary to her initial statement, EMERSON’s (1996) taxonomic treatment of the Ranidae is

very difficult to understand and highly confusing. On one hand, she recognized a genus (or

subgenus?) Hylarana distinct of Rana for a heterogeneous group of frogs that by all skeletal

and morphological characters have long been known to be rather closely related to the group

including the type-species of Rana (Rana temporaria Linnaeus, 1758). But on the other hand,

Source : MNHN, Paris

88 ALYTES 17 (1-2)

allegedly to avoid “potential confusions” linked to the use of the name Limnonectes, she

maintained in Rana, without comment and without mention of subgenera, several groups of

ranids long known to be only distantly related to the latter group, including some that have

been for more than 150 years (since TsCHUDI, 1838) placed in one or two genera (Occidozyga

Kuhl & Van Hasselt, 1822 and sometimes Phrynoglossus Peters, 1867) distinct from Rana. AI

these taxonomic and nomenclatural novelties were presented in this paper without explana-

tion or discussion, and published in a famous journal whose title claims interest in “systematic

biology”.

INDIRANINAE BLOMMERS-SCHLÔSSER, 1993 AND

NYCTIBATRACHINAE BLOMMERS-SCHLÔSSER, 1993

T'have on several occasions already (e.g., DuBois, 1984: 6, 1987b: 117-126, 1987c: 48-52)

pointed out that, contrary to what some zoologists seem to believe, family-group names in

zoology are regulated by the Code and must follow the rule of priority, just like species-group

and genus-group names. This means in particular that the valid name of a family-group taxon

is the first published one based on a generic name included in the taxon, whatever the current

status of this generic name (valid name or invalid junior synonym); this is exactly parallel to

the situation in the genus-group, where a genus name can be valid even if its type-species is a

junior synonym. This rule is by far the best one for the stability of names, for reasons that were

explained in detail already by MYERS & LEVITON (1962), and that may be well illustrated by a

simple example (see Dugois, 1984). GÜNTHER (1858) established a family Polypedatidae,

based on the generic name Polypedates Tschudi, 1838. This family was recognized under this

name by many authors for many years, including in the title of a volume of the famous series

Das Tierreich (Au, 1931), but its name was changed into Rhacophoridae by HOFFMAN (1932)

because the genus Polypedates was then considered a synonym of Rhacophorus Kuhl & Van

Hasselt, 1822. Since then however, a tendency has developed to revalidate Polypedates as a

valid genus: it would clearly have been better, for the sake of nomenclatural stability, to keep

using the name Polypedatidae for the family, whatever the status of the generic names. A quite

similar case concerns the use of Microhylidae Günther, 1858 instead of Gastrophrynidae

Fitzinger, 1843 (see Dugois, 1984). In both these cases, by virtue of Article 40 of the Code, the

senior names (Polypedatidae and Gastrophrynidae) cannot be resurrected now, as they were

replaced because of a synonymy of the type-genus before 1961, but it is important to note that

the same would not be true if this replacement had taken place after 1960.

A number of recent authors do not understand this rule, or deliberately refuse to follow

it, and they tend to change the name of a family-group taxon as soon a: type-genus is

considered an invalid junior synonym, even when this was first done much after 1960; they

may even clearly argue in favour of this non-respect of the Code (see e.g. SAVAGE, 1986).

Strikingly, in several recent cases, their attitude was supported by the Commission itself, in

accepting to “suppress” some senior synonyms in the family-group in order for the corre-

sponding taxa to bear names based on valid generic names. If followed by all, this movement

would lead to suppress the rule of priority for family-group names, thus going back to a

situation where local groups of zoologists, the most “powerful” ones probably, could try and

Source : MNHN, Paris

Dugois 89

impose “their” nomenclature to the whole international scientific community — a most

retrograde step indeed.

Interested readers can find a list of such problems in living amphibian family-group

nomenclature in DuBois (1984, 1987a: 11-12, 1987b: 121-122, 1987c: 48-52). Particularly

striking are the cases of two invalid names that are still used by some authors (and accepted

by the editors of some journals, even well-known ones), despite these repeated mentions of

their invalidity, and although in these cases Article 40 does not apply and does not permit to

keep them, i.e. the invalid Xenopodinae Fitzinger, 1843 instead of the valid Dactylethrinae

Hogg, 1838, and the invalid Bombininae Fejérväry, 1921 instead of the valid Bombinatorinae

Gray, 1825.

There wish to point out a new case of the same kind. Dugois (19874: 66) established a

tribe Ranixalini for the ranid genera Nannophrys Günther, 1869, Nyctibatrachus Boulenger,

1882 and Ranixalus Dubois, 1986. Some months later, however, the same author (DUBOIs,

19874) discovered that the generic name /ndirana Laurent, 1986 was a senior subjective

synonym of Ranixalus and should replace it; however, he remarked that, according to the

Code, the name Ranixalini did not have to be changed and remained the valid one for the tribe.

Dusois (1992: 334) raised this tribe to the rank of subfamily, under the name Ranixalinae.

BLOMMERS-SCHLÔSSER (1993) presented a new hypothesis of cladistic relationships within the

Ranidae and proposed two new subfamilies, the Indiraninae and the Nyctibatrachinae. The

first of these names is clearly an invalid one, being a strict junior subjective synonym of

Ranixalinae: if subsequent authors wished to adopt BLOMMERS-SCHLÔSSER’S (1993) taxono-

mic scheme, they should replace the name Indiraninae by Ranixalinae in this classification. As

for the name Nyctibatrachinae, it is also likely to be a synonym of Ranixalinae, but here for

taxonomic, not nomenclatural, reasons. BLOMMERS-SCHLÔSSER (1993) did not take into

account in her analysis the characters pointed out by Dugois (1987a, 1992) and that suggest

that Indirana, Nannophrys and Nyctibatrachus most likely constitute a holophyletic group,

such as the presence of femoral glands in males of ndirana and Nyctibatrachus, the highly

derived terrestrial tadpoles showing several probable synapomorphies in /ndirana and Nan-

nophrys, and the characters shared by Nannophrys and Nyctibatrachus according to CLARKE

(1983: 395). Despite these data, BLOMMERS-SCHLÔSSER (1993) referred these three South

Indian and Sri Lankan genera to three distinct subfamilies (she included Nannophrys in her

African subfamily Cacosterninae). Therefore, the whole phylogenetic, taxonomic and

nomenclatural scheme proposed by this author seems highly questionable and will have to be

re-evaluated.

RELATIVE PRIORITY BETWEEN SIMULTANEOUS NAMES

As shown by these examples, it is clear that the taxonomy of the Ranidae is still not

stabilized and will show important changes in the future. As argued above, the existence of

provisional taxonomic schemes, if well understood, can provide a strong help for the progress

of our works on this taxonomy. Taxonomic changes will in their turn result in nomenclatural

changes, but, if the proper care is taken, these latter changes will be automatic and should not

pose any problem to taxonomists. The previously published catalogues of genus-group and

Source : MNHN, Paris

90 ALYTES 17 (1-2)

Table 1. — Past first-reviser actions concerning simultaneous (i.e., published at the same date) genus-

group and family-group names in the families Hyperoliidae, Phrynobatrachidae and Ranidae (as

defined by DUBOIS, 1992). The sign > means “afforded priority over”

Original

publication

Relative priority afforded to simultaneous names

by first-reviser action

First-reviser

TSCHUDI, 1838 _| DUMÉRIL & BIBRON, 1841: 515 Polypedates > Boophis + Buergeria

TSCHUDI, 1838 STEINEGER, 1907: 143 Polypedates > Buergeria + Theloderma

FITZINGER, 1843 FITZINGER, 1843: 31 Pelophylax > Euphlyctis + Limnophilus + Phrynoderma

FITZINGER, 1843 DUBOIS, 1976: 1112 Euphlyctis > Limnonectes + Phrynoderma

[BouzEvorR, 1882 DUBOIS, 1987: 68 Nyctibatrachus > Nannobatrachus

NOBLE, 1931 DuBoIS, 1982: 135-136 Petropedetinae > Cacosterninae

LAURENT, 1944 | LAURENT & COMBAZ, 1950: 277 Afrixalus > Acanthixalus + Heterixalus

family-group names available in the Ranidae (Dugois, 1981, 1984, 19874, 1992) should allow

any author to find if names are already available for any newly defined taxon, which one has

priority, or if a new name has to be coined. The only cases when nomenclatural changes will

not be automatic are those where different names, initially published in the same work or in

different works but at the same date (“simultaneous names”), are considered subjective

synonyms. In such cases, according to the Code, relative priority among these names is fixed

by a first-reviser action. Once published, a first-reviser action is definitive and cannot be

modified by subsequent authors. It is therefore important to be able to trace all first-reviser

actions ever taken in the nomenclature of a given zoological group, but it is a difficult work for

anyone who is not very well acquainted with all the literature dealing with the taxonomy of the

group.

In order to help future workers on the taxonomy of Ranidae and related groups

(recognized as the families Arthroleptidae, Astylosternidae, Hemisotidae, Hyperoliidae,

Mantellidae and Phrynobatrachidae by Dugois, 1992: 309), I provide in tab. 1-2 a list of

publications where several simultaneous genus-group and family-group names currently

referred to these groups were published, with information on first-reviser actions and on the

resulting relative priority among these names. Only names created in the same publication

were surveyed. Problems of priority may also occur between names published in different

works of the same year, but then a careful study of the case, with research of information on

exact dates of publication, must be carried out, which was beyond the scope of the present

work. Table 1 gives information on first-reviser actions that have already been taken by

previous authors in the past. When no such first-reviser action has already been published, I

hereby take such an action (tab. 2), so that in the future any allocation of genus-group or

family-group name in these families should be automatic (except in the rare possible cases of

names published in different works of the same year, as mentioned above) and not liable to be

complicated by subsequent “inadvertent” first-reviser actions in obscure publications (e.g..

due to subjective synonymisation of two names, one being cited in the synonymy of the other)

that may escape the attention of some colleagues. The choice of the order of priority among

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Dugois 91

Table 2. - New first-reviser actions concerning simultaneous (i.e., published at the same date) genus-

group and family-group names in the families Arthroleptidae, Astylosternidae, Hemisotidae,

Hyperoliidae, Mantellidac, Phrynobatrachidae and Ranidae (as defined by DUBOIS, 1992). The

sign > means “afforded priority over”. Names followed by an asterisk were also concerned by

past first-reviser actions (see tab. 1); in such cases, the new first-reviser actions below are

compatible with these earlier actions.

Relative priority afforded to simultancous names by present first-reviser action

KUHL & VAN HASSELT, 1822

Rhacophorus > Occidozyga

TSCHUDI, 1838

Hylarana > Polypedates* > Boophis* > Pyxicephalus > Buergeria* > Strongylopus >

Theloderma® > Cornufer > Oxyglossus > Eucnemis > Orchestes > Oxydozyga

DUMÉRIL & BIBRON, 1841

Tomopterna > Limnodytes > Ixalus

FITZINGER, 1843

Pelophylax* > Euphiyctis* > Limnonectes* > Lithobates > Hydrophylax >

Tachyenemis > Trachyhyas > Eremiophilus > Limnophilus* > Phrynoderma*

GISTEL, 1848

Philautus > Zoodioctes > Buccinator > Dendricus > Phyllodytes > Epipole

Sur, 1849 Arthroleptis > Stenorhynchus

GüNTHER, 1859 Platymantis > Leptopelis > Hemisus > Sphaerotheca

PETERS, 1863 Hoplobatrachus > Hemimantis > Leptoparius

COPE, 1865 Amolops > Staurois

PETERS, 1867 Phrynoglossus > Leptomantis

GÜUNTHER, 1869 Nannophrys > Megalixalus

HOFFMANN, 1878

Fergusonia > Aemolops > Cruminifera

BOULENGER, 1882

Mantella > Nyctibatrachus* > Nannobatrachus*

SCHULZE, 1890

Crotaphitis > Baliopygus

BOULENGER, 1893

Chirixalus > Phrynoderma

PALACKY, 1898

Rhacoforus > Nannofrys

BOULENGER, 19004

Trichobatrachus > Cardioglossa > Scotobleps > Gampsosteonyx > Dilobates

BOULENGER, 1917a

Prychadena > Aubria

METHUEN, 1920

Gephyromantis > Trachymantis

AHL, 1924

Pseudocassina > Tornierella

Hylarthroleptis > Pararthroleptis:

Arthroleptella > Microbatrachella > Microbatrachus

Palmatorappia > Dendrobatorana

Petropedetinae* > Cacosteminae* > Comuferinae > Phrynopsinae

DECKERT, 1938

Micrarthroleptis > Pseudarthroleptis

LAURENT, 1940

Coracodichus > Abroscaphus > Arthroleptulus

LAURENT, 1941

Taphriomantis > Elaphromantis > Heteropelis

LAURENT, 1944

Afrixalus* > Heterixalus* > Acanthixalus*

LAURENT & COMBAZ, 1950

Crprothylax > Phlyctimantis

LAURENT, 1972

Kassinini > Leptopelini

Dusois, 1987a

Ingerana > Kirtixalus > Taylorana > Amietia > Gorhixalus > Liurana > Bourretia

Dusois, 1987a

Ranixalini > Pychadenini > Tomopternini

PERRET, 1988

Alexteroon > Arlequinus > Chlorolius

CHANNING, 1989

Buergerinae > Tachycneminae

Flet al., 1991

Odorrana > Glandirana > Rugosa > Pseudorana > Tenuirana > Unculuana >

Quadrana > Tigrina

DuBois, 1992

Amo > Sylvirana > Nidirana > Afrana > Amnirana > Ombrana > Sierrana >

Nasirana > Pulchrana > Amerana > Pantherana > Humerana > Papurana >

Sanguirana > Blommersia > Brygoomantis > Guibemantis > Spinomantis >

Quasipaa > Annandia > Eripaa > Gynandropaa > Feirana > Chalcorana >

Aquarana > Zweifelia >Aurorana > Eburana > Tylerana

DuBoïs, 1992

Limnonectini > Paini > Conrauini

BLOMMERS-SCHLOSSER, 1993

Nyctibatrachinae > Indiraninae

GLAW & VENCES, 1994

Phylacomantis > Chonomantis > Pandanusicola > Ochthomantis

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92 ALYTES 17 (1-2)

several simultaneous names was based on the following rationale: in most cases, names

currently in use were afforded priority over names currently considered invalid, and well-

known names over poorly known names; priority was usually given to names designating

groups of larger size (with more species) than others, and to names designating genera over

names proposed for subgenera; junior homonyms and objective synonyms (e.g., unjustified

emendations) of potentially valid names were given lowest priority; all other things being

equal, I have preferred euphonious or elegant names to disgracious ones.

Of course, most of these first-reviser actions will have no bearing on future nomencla-

tures in these families, because synonymies between simultaneous names will be rather rare,

even for the authors who currently advocate a very strong “lumper” approach to higher

taxonomy: thus, to take just one example, it is highly unlikely that the names Buergeriinae

Channing, 1989 and Tachycneminae Channing, 1989 will ever compete for synonymy!

However, in other cases the problem will certainly arise, and it will be simpler and easier to

refer to a single couple of tables to know the relative situation of two given names, rather than

having to embark on long and difficult researches, so that these tables embrace all cases of

“simultaneous” family-group and genus-group names in these families.

“RANA DUBOIST” IN EMERSON & WARD (1998)

Peer review by colleagues before acceptance of a paper for publication in a scientific

periodical or journal is now a common practice worldwide. Manuscripts thus submitted for

advice to scientists, who usually work in the same research field as the author of the paper, are

sent to them under the (usually tacit) agreement that the reader will not make a private use of

the information contained in the paper and will not publish this information, or information

derived from it, prior to the publication of the submitted paper. With some shocking

exceptions, this rule is usually followed by reviewers. But a particular problem may arise when

the submitted paper is rejected, at least in its original form, and is never published, or only

published after a considerable delay. The risk exists that the referee, either by inadvertency or

by lack of request for information, might consider that, after a certain time has elapsed, the

paper was actually published, and might feel free to use the information it contained, or to

refer to this information as if it had been published. In some cases, this merely has the

consequence of publishing only the final result of a work, without all the accompanying data

that allow to ascertain that these results were obtained in a serious scientific manner: in some

other cases, this may have nomenclatural consequences.

An example of the first kind is KURAMOTO’s (1990) mention of some of the results of a

work by IsKANDAR et al. (unpublished) that he quoted as being in “A/ytes (in press)”, but that

was actually never published or sent to press. The manuscript in question was indeed

submitted to the journal A/yres on 4 April 1989, accessioned under number 89.156 and sent

for review to two referees, including Mitsuru KURAMOTO, on 12 April 1989. On 31 August

1989, after receipt of the reports of the two readers, copies of these reports were sent to the

first author of the manuscript with an accompanying letter stating that, on the whole, the

manuscript was very interesting and useful, but asking for a few minor modifications before

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Dugois 93

the paper could be published. Despite subsequent requests for the final manuscript, sent to the

first author on 9 March 1990 and 18 October 1991, no second version of this paper was ever

sent to Alytes. To the best of my knowledge, this paper was never published elsewhere. All the

information contained in the original manuscript has therefore remained unpublished. Men-

tion by KURAMOTO (1990) of some of this information is equivalent to mention of unpub-

lished data obtained from a colleague through “personal communication”, with the differ-

ence that in this case the communication was not direct between two colleagues, but went

through the “mediation” of a journal editor. I suggest that colleagues who might wish to use

these data should quote them as “ISKANDAR et al. in KURAMOTO (1990)”.

In some cases, publication of previously unpublished taxonomic information may have

nomenclatural consequences. This does not occur when no name is associated with the

taxonomic information. Thus, EMERSON (1996: 279) wrote: “a new species has been discovered

in Sulawesi in which the females retain fertilized eggs and the tadpoles develop in the body of

the female (ISKANDAR, 1996).” This information seems quite interesting indeed, and readers

may wish to know more about it. In the References of EMERSON's (1996) paper, the title of a

paper by “ISKANDAR, 1996” appears, followed by the mention “A/ytes (in press)”. However, as

of today (16 April 1999), no manuscript under this title (or an approaching one) has ever been

submitted to Alytes for publication, so that this reference (ISKANDAR, unpublished 4) could

well be qualified as a “phantom reference” (see below). This may be quite frustrating for the

reader but at least, from a nomenclatural point of view, there is no disturbing consequence, as

the name of the “new species” was not mentioned.

The situation is different in the case of EMERSON & WaRD's (1998) article on frogs of the

“Rana grunniens species group”. This paper starts (p. 538) with a table 1 presenting a list of

species referred to this group. This table has a striking particularity: scientific names of species

appear there “nude”, i.e. without their authors and dates. As was well explained by NG (1994),

citation of author and date is not only a tradition in zoology: it is important as it allows

unambiguous identification of the nominal species at stake. Absence of such a basic infor-

mation in a table published in a journal having “Linnean” in its title is an interesting

illustration of a recent trend for zoological publications to neglect or fully ignore the basic

rules, recommendations and needs of nomenclature. In this case, reference to Dugois's

(1987a) work can allow the reader to avoid confusion, but without going to this reference it is

impossible to know e.g. if the nominal species referred to in this table as “ Rana microtympa-

num is Rana microtympanum Van Kampen, 1907 (a member of Limnonectes) or Rana

microtympanum Boulenger, 1919 (a member of Hildebrandtia). This table contains 14 of the

15 names listed by DuBois (19874: 63) as members of his Limnonectes (Limnonectes)

grunniens group, but the name Rana macrodon has disappeared from this list without

explanation. Transfer of all these species from the genus Limnonectes to the genus Rana would

have required a change of the grammatical gender of some of the specific names, which was

not donein alle so that the list contains two incorrect spellings (Rana modestus for Rana

modesta, Rana visayanus for Rana visayana). Later in the paper, p. 540, at the beginning of the

Material and methods, two additional names suddenly appear for two species of this species-

group: Rana macrodon and * Rana duboisi”, This latter name is given without any explanation

or reference to its source. It appears again on three occasions in the paper (p. 545, 546. 553),

without further information. No publication proposing this name for a new species was ever

published from 1758 to 1998, year of publication of EMERSON & WaRD's (1998) work, so that

Source : MNHN, Paris

94 ALYTES 17 (1-2)

this name in this paper must be considered a new species name — unless it was borrowed from

some unpublished manuscript, not cited in the References of their paper.

Actually, I am aware of two unpublished manuscripts where this name, or a related one,

was proposed as the name of a new species: in the first one (DAS, unpublished), the name

“Rana duboisi” was proposed for a new species of Rana (Sylvirana) (sensu Duois, 1992)

from above Kallar (Kerala, India); in the second one (IskANDAR, unpublished b), a new

species of Limnonectes (Limnonectes) (sensu DuBois, 1992) from Kamarora (Lore Lindu

National Park, Central Sulawesi, Indonesia) was described as “Limnonectes duboisi”. To the

best of my knowledge, none of these two papers has been published so far, and it is not even

certain that they were ever submitted for publication. At the head of the manuscript of

ISKANDAR'S paper (a copy of which was presented to me by Georges PASTEUR), it was stated

that this paper was intended for submission to the journal Alytes, but as of today this has not

yet been done. However, it is very likely that the name “Rana duboisi” was borrowed (and

modified, through change of generic allocation) from this second manuscript, as the origin

given for the specimens of this species studied by EMERSON & WARD (1998: 553) is the same

(except for the misspelling “Linu” for Lindu) as that of the type-locality of “Limnonectes

duboisi” in ISKANDAR (unpublished b), and as EMERSON & WARD (1998: 551) thank Djoko

ISKANDAR for providing them with “tissue samples of Southeast Asian ranids”. It would thus

appear that EMERSON & WaRD (1998), by publishing the name “Rana duboisi before

ISKANDAR, became the authors (in the technical sense of this term according to the Code) of

this nominal species. However, this is not true, because this name is a nomen nudum in their

paper: no character is provided to distinguish this species from related ones, nor is there any

reference to a “bibliographic reference to such a published statement” (Article 13.a of the

Code). This case is interesting, however, as it allows discussion of two questions that are likely

to be raised again later in zoological nomenclature: is a species name rendered nomencla-

turally available by publication, either (1) of a Genbank (or other similar data base) catalogue

number reference for a sequence of this species, or (2) of a cladogram showing the hypothe-

sized relationships of this new species to related taxa?

(1) EMERSON & WaRD’s (1998) paper does not contain any table or figure giving the

sequences obtained for portions of the 12S and 16$ ribosomal RNA genes of the specimens

studied in their work. If it was the case, the sequences associated with the new name “Rana

duboisi” would clearly qualify as diagnostic characters making this latter name nomencla-

turally available, just like mating call characteristics (see e.g. SCHNEIDER & SINSCH, 1992;

Duois & OHLER, 1995: 179) or any other non-morphological character of an animal species.

EMERSON & WaRD (1998: 541) stated that the sequences obtained in their work were entered

in the Genbank data base, and provided their catalogue numbers. As such a procedure is likely

to become more and more common in evolutionary biology, it is important to know whether

such Genbank cataloguing qualifies as a publication as defined by the Code. This is clearly not

the case in the edition of the Code currently in force. Article 8 of this edition allows for a work

containing a new name or a nomenclatural act to be regarded as published even if “produced

after 1985 by a method that does not employ ink on paper in conventional printing”, but only

if it contains “a statement by the author that any new name or nomenclatural act within it is

intended for permanent, public, scientific record”. This does not apply to names entered in the

Genbank, so that the latter must be considered as “unpublished” in the eyes of the Code.

Caution will however have to be given to the precise wording of Article 8 in the final,

Source : MNHN, Paris

Dugois 95

published version of the next edition of the Code, to check if this provision has not been

changed. Sequences entered in the Genbank cannot therefore be used as diagnostic characters

for new taxa, but it is important to note that, as soon as an author (either the person who

established the sequence, or another colleague) publishes this sequence in a printed work, this

sequence can become an excellent diagnostic character making a new name nomenclaturally

available.

(2) Recent proposals have been made (see e.g. DE QUEIROZ & GAUTHIER, 1994) to modify

drastically the philosophical basis of zoological nomenclature by attaching the names to

“phylogenetic definitions of taxon names”, that would be more in agreement with a phyloge-

netic system of taxonomy than other kinds of “definitions”. These proposals are in my

opinion based on a major misunderstanding and entertain a confusion between taxonomy

and nomenclature. In the system of zoological nomenclature currently in force, allocation of

zoological names to taxa is not at all based on définitions, diagnoses or descriptions, but on the

taxonomic allocation of name-bearing type-specimens or onomatophores, which constitute an

objective, material and stable connection between the real world of animal populations and

the world of language, whereas definitions are liable to change (for more details, see Dugois &

OHLER, 1997). In this system, definitions or diagnoses only contribute to the nomenclatural

availability of names, but not to their allocation to taxa. On the other hand, definitions or

diagnoses are crucial for the qualification of taxa, and in this domain one may well wish to use

“phylogenetic definitions”, but this is a matter of taxonomy, not of nomenclature. Nomen-

clature is a system allowing a non-ambiguous, stable, automatic and universal allocation of

names to taxa, under a given taxonomy, and the current rules of nomenclature are fully

compatible with any taxonomic system, including the “phylogenetic taxonomy” (or “clado-

nomy” sensu DUBoIs, 1997) advocated by DE QUEIROZ & GAUTHIER (1990, 1992).

Under a nomenclatural system like that suggested by DE QUEIROZ & GAUTHIER (1994),

the association of a name with a given clade in a cladogram, as is the case of the name “ Rana

duboisi” in figure 3 of EMERSON & WaRD’s (1998) paper, could possibly be considered enough

to provide a “phylogenetic definition” of this name and to make it nomenclaturally available,

but this is not true under the Code currently in force for all zoologists. Cladograms are

hypotheses of relationships but, although built on the basis of a character analysis, they alone

do not provide the characters of the included taxa. Under the Code, the presence of a

diagnosis or definition, i.e. a statement regarding characters (not relationships), is necessary

for a name to be nomenclaturally available: a taxon name published only with information on

the supposed cladistic relationships of this taxon is therefore not available under the present

Code.

In conclusion, the name “ Rana duboisi” published by EMERSON & WARD (1998) associ-

ated with a reference to the Genbank and with a position in a cladogram, but without any

diagnostic character, is, according to the current Code, a nomen nudum.

To avoid the frequent repetition of similar nomenclatural puzzles in the future, itis highly

desirable that authors try their best not to publish new names borrowed from unpublished

manuscripts or from personal communications from colleagues (see e.g. DuBois, 1998: 20).

Any name inadvertently published in such conditions may qualify, like in the present case, as

a nomen nudum, i.e. a name devoid of nomenclatural status, which has no real nomenclatural

consequences. But it may also happen to be a validly published name, if it was associated in its

Source : MNHN, Paris

96 ALYTES 17 (1-2)

first publication with descriptive or diagnostic data, for which e.g. paper-printed gene se-

quences would fully qualify. Under the Code currently in force, such inadvertent publication

of new names associated with gene sequences would give birth to nomenclaturally available,

although unvoluntarily so, names, aptly qualified by VENCES et al. (1999) as “phantom

names”. This will remain so as long as the current Code is in force. Seemingly, as reported by

VENCES et al. (1999), the next edition of the Code will include the following, highly desirable,

new rules, that would greatly reduce the inadvertent creation of such names: (1) the need for

an explicit statement that the new name applies to a newly defined species-group taxon: (2) the

need for a clear designation of a name-bearing type, deposited in an identified collection.

Such rules appear very reasonable and “obvious” for all experienced taxonomists, and, once

in force, they would certainly be beneficial for the future of zoological nomenclature. But it is

greatly to be hoped that these rules will only be prospective (ï.e., applying to works published

after the new edition of the Code) and will have no retroactive effect: otherwise, this might

have dramatic consequences regarding nomenclatural universality and stability, as many

names now considered valid by all zoologists were first published (either very long ago or

more, sometimes much more, recently) without respect for these rules.

ACKNOWLEDGEMENTS

For information, advice and comments on previous drafts of this paper, I am most grateful to Franco

ANDREONE (Torino), Roger BOUR (Paris), Alessandro MINELLI (Padova), Annemarie OHLER (Paris),

Georges PASTEUR (Montpellier) and Miguel VENCES (Bonn).

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Corresponding editor: Franco ANDREONE.

©ISSCA 1999

Source : MNHN, Paris

AIVTES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD

Chief Editor: Alain Dusois (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France).

Deputy Editor: Janalee P. CaLoWELL (Oklahoma Museum of Natural History, University of Oklahoma,

Norman, Oklahoma 73019, USA).

Editorial Board! Jean-Louis ALBARET ais France); Franco ANDREONE (Torino, Italy): Günter GOLLMANN

(Wien, Austria); Tim HALLIDAY (Milton Keynes, United Kingdom); W. Ronald HEvER (Washington,

USA): Esteban O. LAviLLA (Tucumän, Argentina): Karen R. Lies (Canton, USA); Thierry LODÉ

(Angers, France); Masafumi Matsut (Kyoto, Japan); John C. PoynroN (London, England); Erik R.

WiLD (Dubuque, USA).

Technical Editorial Team (Paris, France): Alain Dupois (texts); Roger BouR (tables); Annemarie OHLER

(figures). ; ;

Book Review Editor: Annemarie OHLER (Paris, France).

SHORT GUIDE FOR AUTHORS

(for more detailed Instructions to Authors, see Alytes, 1997, 14: 175-200)

Alytes publishes original papers in English, French or Spanish, in any discipline dealing with amphibians.

Beside articles and notes reporting results of original research, consideration is given for publication to synthetic

review articles, book reviews, comments and replies, and to papers based upon original high quality illustrations

(such as colour or black and white photographs), showing beautiful or rare species, interesting behaviours, etc.

The title should be followed by the names) and address(es) of the author(s). The text should be

typewritten or printed double-spaced on one side of the paper. The manuscript should be organized as follows:

English abstract, introduction, material and methods, results, discussion, conclusion, French or Spanish

abstract, acknowledgements, literature cited, appendix.

Figures and tables should be mentioned in the text as follows: fig. 4 or tab. 4. Figures should not exceed

16 x 24 cm. The size of the lettering should ensure its legibility after cure legends of figures and tables

should be assembled on a separate sheet. Each figure should be numbered using a pencil.

References in the text are to be written in capital letters (BOURRET, 1942; GRAF & PoLLs PELAZ, 1989;

INGER et al., 1974). References in the Literature Cited section should be presented as follows:

Bouraer, R. 1942. Les barraciens de l'dochine. Hanoï Institut Océanographique del'Indochine:x + 1-547,

pl.

Grar, D. & PoLLsPeLAz, M. 1989. — Evolutionary genetics of the Rana esculenta complex. In:R. M. DAWLEY

& 1. P. BoGaRT (ed.), Evolution and ecology of unisexual vertebrates, Albany, The New York State

Museum: 289-302.

IxGeR, RE, Vorss, H. K. & Voris, H. H., 1974. - Genetic variation and population ecology of some Southeast

Asian frogs of the genera Bufo and Rana. Biochem. Genet., 12: 121-145.

Manuscripts should be submitted in triplicate either to Alain Dupois (address above) if dealing with

amphibian morphology, systematics, biogeography, evolution, genetics or developmental biology, or to Janalee

P. CaLDWELL (address above) if dealing with amphibian population genetics, ecology, ethology or life history.

Âcceptance for publication will be decided by the editors flic lowing review by at least two referees.

If possible, after acceptance, a copy of the final manuscript on a floppy disk (3 4 or 5 ) should be sent

to the Chief Editor. We welcome the following formats of text processing: (1) preferably, MS Word (1.1 to 6.0,

DOS or Windows), WordPerfect (4.1 to 5.1, DOS or Windows) or WordStar (3.3 to 7.0); (2) less preferably.

formated DOS (ASCID or DOS -formated MS Word for the Macintosh (on a 3 % high density 1.44 Mo floppy

isk only).

Page charges are requested only from authors having institutional support for this purpose. The

publication of colour photographs is charged. For each published paper, 25 free reprints are offered by ISSCA

to the author(s). Additional reprints may be purchased.

Publish with the support of AALRAM

(Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum National d'Histoire Naturelle, Paris, France).

Directeur de la Publication: Alain Dusois.

Numéro de Commission Paritaire: 64851.

©ISSCA 1999

Source : MNHN, Paris:

Alytes, 1999, 17 (1-2): 1-100.

Contents

Alain DuBois

Editorial

Miguel VENCES, Frank GLAW & Wolfgang BÔHME

A review of the genus Mantella (Anura, Ranidae, Mantellinae):

taxonomy, distribution and conservation of Malagasy poison frogs .…

Annemarie OHLER

Une nouvelle espèce du genre Leptodactylodon

(Arthroleptidae, Astylosterninae) du Gabon .