ISSCA

International Society for the Study

and Conservation of Amphibians

(International Society of Batrachology)

SEAT

Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle, 25 rue Cuvier, 75005

Paris, France. — Tel.: (33).(0)1.40.79.34.87. — Fax: (33).(0)1.40.79.34.88. — E-mail: dubois@mnhn.fr.

BOARD

sident: W. Ronald HEYER (Washington, USA).

lain Dugois (Paris, France).

Depury Secrerarÿ: Annemarie OHLER (Paris, France).

: Jean-Louis DENIAUD (Paris, France).

surer: Rafael DE S (Paris, France).

Councillors: Brita GRiLLITSCH (Wien. Austria); C. Kenneth Dobp, Jr. (Gainesville, USA): Julio Mario

Hovos (Bogotä, Colombia); Thierry Lobé (Angers, France): Jon LOMAN (Lund, Sweden); Alain

PAGANO (Angers, France): Stephen J. RicHaRDs (Townsville, Autralia).

General Secretary

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(804) 289-8333) Source : MNHN, Paris

AINTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

November 2000 Volume 18, N° 1-2

Alytes, 2000, 18 (1-2): 1-14. 1

Un nouveau Leptodactvlodon

de la Dorsale camerounaise

(Amphibia, Anura)

J.-L. AMIET* & F. DOWSETT-LEMAIRE**

* 48 rue des Souchères, 26110 Nyons, France

** 12 rue des Lavandes, 34190 Ganges, France

A new species of Leptodactylodon is described from montane forest in

the Bakossi Mounts (Cameroun). This species is in the complex of small

Leptodactylodon with well-developed vomerine teeth. Its specific charac-

ters include very small size, narrow head, males without gular folds and

spiny skin, but with rather thick front legs. The voice is highly characteristic,

as the main calls are reminiscent of the noise produced by the cords of à

guitar being plucked.

Le pôle de diversification maximale des Astylosterninae Noble, 1927 se situe au Came-

roun, et plus précisément dans la partie occidentale du territoire, où se dressent les montagnes

et les plateaux de la Dorsale camerouna Les cinq genres de la sous-famille y sont

représentés et, parmi eux, les genres Astylosternus Werner, 1898 et Leptodactylodon

Andersson, 1903 comprennent chacun une dizaine d'espèces, pour la plupart orophiles

(AM 1977, 1980).

Pour ces deux genres, les variations climatiques quaternaires, en entraînant d'importants

changements dans les limites altitudinales des étages, et donc des extensions ou des morcel-

lements des aires de répartition, ont eu pour corollaire la formation d'espèces qui peuvent être

étroitement localisées (AMIET, 1975, 1987).

C'est probablement le cas pour le Leptodactylodon décrit ici. Cette espèce, trouvée

uniquement près des villages de Kodmin et d’Edib, dans les Monts Bakossi, était restée

ignorée malgré les prospections répétées effectuées par l’un de nous (JLA) sur le Mont

Manengouba, à une dizaine de kilomètres à vol d'oiseau de la première localité. Elle a été

découverte grâce à ses vocalisations, enregistrées en marge de recherches ornithologiques

(FDL): soumises pour identification, il apparut immédiatement qu'elles étaient produites par

BIBL. DU iothèque Centrale Muséum

MUSEUR

= JU

3 3001 0012

Source : MNHN, Paris

2 ALYTES 18 (1-2)

une espèce encore inconnue. Toutefois, leurs caractéristiques acoustiques étaient si inhabi-

tuelles qu'il ne fut même pas possible d'imaginer à quel genre leur auteur pouvait appar-

tenir !

Aucun spécimen n’avait été récolté, mais l'emplacement d’un groupe de mâles vocale-

ment actifs avait été bien repéré. Chris Wild, alors coordinateur scientifique des projets

Kupe-Bakossi-Takamanda (WWF Cameroun), put ainsi, à notre demande, y faire rechercher

cette espèce. Grâce à la collaboration de D. Menze et E. Abwe, 6 mâles et un juvénile purent

être capturés en décembre 1998. Ce matériel nous permet de décrire une nouvelle espèce que

nous sommes heureux de dédier à C. Wild, qui a contribué à résoudre l'énigme posée par ses

appels.

REMARQUES MÉTHODOLOGIQUES

(1) Dans les montagnes camerounaises, de nombreuses espèces d’anoures ont, comme les

Leptodactylodon, une activité vocale diurne (AMIET, 1989). Nous reproduisons ci-après le

passage d’une lettre de C. Wild (lettre du 21 janvier 1999 à JLA) décrivant le procédé qu'il a

utilisé pour séparer auditivement la nouvelle espèce de celles qui lui étaient associées: “(...) the

site was marked off with tape (about 5 m X 5 m), left undisturbed, and revisited (...) for

intensive microhabitat search involving the removal of all vegetation and debris. Forty two

(42) anurans were captured, comprising: Phrynobatrachus, Phrynodon, Wolterstorffina,

Arthroleptis and some small ‘Leptodactylodon-like’ specimens. AIl were brought back to my

house in Nyasoso and placed in a large container covered in muslim mesh with vegetation.

Within 24 hours the ‘guitar’ calls were heard. I then isolated the group by species to identify

which type was calling. Within the next 24 hours, the group calling with the exact same

guitar-like voices was conclusively determined.”

(2) Ainsi que l’un de nous l’a déjà souligné (AMIET, 1980), les rapports morphométriques

sont rarement discriminants chez les Leptodactylodon. Les quelques mensurations données ici

(tab. 1-2), ainsi que leurs abréviations, seront les mêmes que dans la “Révision” précitée :

L, longueur du corps, mesurée de l’extrémité du museau à l’entrejambe; T, largeur de la tête

derrière les yeux ; C, longueur de la cuisse, de l’entrejambe au genou ; J, longueur de la jambe,

de la saillie du genou à celle du talon ; P, longueur du pied, du talon à l'extrémité de l’orteil 4.

Leptodactylodon wildi n. sp.

Matériel étudié.

Holotype. - Mâle MNHN 2000.0607 de Kodmin (Cameroun, env. 1350 m), décembre

1998, C. Wild leg. Individu en livrée nuptiale (épines métacarpiennes bien développées).

Habitus : voir fig. 1. Mensurations: voir tab. 1.

Paratypes. — $ mâles en livrée nuptiale de même origine, capturés avec l’holotype et

déposés aux muséums de Paris (MNHN 2000.0608-0610), Bonn (ZFMK 72150) et Londres

(BMNH 1998.38).

Source : MNHN, Paris

AMIET & DOWSETT-LEMAIRE 3

Tab. 1.- Quelques mensurations (en dixièmes de mm) et proportions chez L. wildi. L'holotype

est le spécimen MNHN 2000.0607.

N° spécimen | L | T | cs |pP|TL|cL| y | PL

MNHN 2000.0607| 225 | 85 | 90 | 85 | 140 37,8) 40 | 37,8 | 62.2

MNHN 2000.0608| 230 | 85 | 80 | 90 | 150 | 37 348 | 39 | 65,2

MNHN 2000.0609! 215 | 80 | 85 | 80 : 135 | 37,2 | 39,5 | 37,2 | 62,8

MNHN 2000.0610| 235 | 85 | 85 | 95 . 140 | 36,2 | 36,2 | 40,4 | 59,6

BMNH 1998.382 | 225 | 80 | 85 | 95 | 140 | 35,5 | 37,8 | 42,2 | 62,2

ZFMK 72150 |235 80 | 85 | 95 135 | 34 | 36.2 | 40,4 | 57,4

Moyenne 230 82,5 | 87,5) 90 | 140 | 36,28) 37,42] 39,50 61,57

Ecarttype | 7.58 | 2.74 | 3,16 6.32 | 548 | 1,38 | 2.05 | 1.86 | 2.71

Tab. 2. - Données morphométriques relatives aux Leptodactylodon proches de L. wildi. Ep.

métac. : nombre d’épines métacarpiennes. Les données concernant les espèces autres que

L. wildi sont extraites de AMIET (1980).

Espèce L | T/L | JL | P/L |Ep. métac.

(nombre de spécimens) |

L. albiventris (9) 25,7 36,8 | 41,6 | 63,5 | 2

L. bueanus (6) 26 | 37,9 | 43,9 | 642 | 3

L. p. polyacanthus (7) | 25.2 | 388 | 438 | 66 | 5-6(7)

L. p. punctiventris (7)| 252 | 387 | 441 | 648 5-6

wildi (6) 23

362 | 39,5 | 615 | 4

Source : MNHN, Paris

4 ALYTES 18 (1-2)

Autre matériel. — (1) Un juvénile de même origine; (2) trois diapositives C. Wild de

spécimens vivants ou fraîchement sacrifiés ; (3) deux enregistrements sonores F. Dowsett-

Lemaire, effectués à Kodmin en mars-avril et décembre 1998.

Diagnose. — Leptodactylodon wildi se rapproche de L. albiventris (Boulenger, 190$) et L.

polyacanthus Amiet, 1971 par ses dents vomériennes en courtes crêtes transverses, sa face

ventrale non marbrée et sa taille inférieure à 30 mm. Il s’en distingue par sa taille plus faible

(21,5 à 23,5 mm chez les mâles), par sa tête plus étroite, et surtout par les caractères sexuels

secondaires du mâle : bras hypertrophiés (différence avec L. albiventris), épines métacarpien-

nes au nombre de 4, absence de plis gulaires (différence avec L. albiventris) et de spinosité

pectorale (différence avec L. polyacanthus). H se sépare aussi de ces espèces par divers traits de

coloration, et de toutes les autres espèces de Leptodactylodon par ses vocalisations.

Description.

Habitus, tégument, membres. — L. wildi (fig. 1 et 4) est une des plus petites espèces du

genre: les six mâles récoltés, adultes comme le montre le développement de leurs caractères

sexuels, mesurent de 21,5 à 23,5 mm, ce qui les situe dans le même ordre de grandeur que L.

bicolor Amiet, 1971, L. mertensi Perret, 1959, L. erythrogaster Amiet, 1971 et L. perreti Amiet,

1971. Comme ces derniers, il a une forme générale plutôt cylindrique, avec les flancs longue-

ment parallèles, mais il s’en sépare à première vue par sa tête relativement étroite (T/L de

l'ordre de 36 % : tab. 1) et par ses membres postérieurs encore plus courts et plus épais (fig. 2).

Cet habitus le distingue aussi d’un petit Leptodactylodon gabonais, L. blanci Ohler, 1999, dont

la tête est large et la forme plus trapue (OHLER, 1999: fig. 1).

Chez les individus étudiés, le tégument dorsal paraît lisse, ce qui pourrait être dû, au

moins en partie, au mode de fixation (ces spécimens étaient légèrement gonflés). Les photos de

€. Wild, prises sur le vivant ou sur des spécimens récemment sacrifiés, confirment que le relief

tégumentaire se limite à de vagues rides ou verrucosités à peine perceptibles, surtout sur les

côtés du corps.

Le museau est largement arrondi et dépourvu de canthus et de dépression loréale,

contrairement à ce qui s’observe chez L. albiventris et L. polyacanthus. Les narines ne sont

marquées par aucun relief ; elles se trouvent un peu plus près de l'extrémité du museau que des

angles palpébraux antérieurs. Les yeux, comme chez tous les Leprodactylodon, sont relative-

ment petits et peu saillants ; ils sont largement séparés, l’espace interoculaire représentant

environ 1,5 fois la plus grande largeur de la paupière (sur la tête vue en plan). Le tympan est

indiscernable, peut-être en raison du gonflement des spécimens, mais un faible repli supra-

tympanique est visible.

La main (fig. 3b) présente les caractères habituels chez les Leptodactylodon, avec des

doigts courts se rétrécissant régulièrement vers leur extrémité, qui est à peine élargie. Le doigt

Lest plus long que le doigt 2. Il y a deux tubercules sous-articulaires aux doigts 4 et 3, et un aux

doigts 2 et 1 : à chacun des doigts, les tubercules proximaux forment de larges coussinets peu

saillants. La paume présente deux plages d'aspect semblable, l’une externe, plus ou moins

dédoublée, l'autre interne, prolongeant la saillie où sont implantées les épines métacarpiennes

du mâle.

Source : MNHN, Paris

AMIET & DOWSETT-LEMAIRE 5

Fig. 1. — Leptodactylodon wildi n. sp. face dorsale de l'holotype, MNHN 2000.0607, de Kodmin

(Cameroun), 21.XI1.98, Wild leg. (dessiné après fixation).

Source : MNHN, Paris

6 ALYTES 18 (1-2)

77 L. albiventris L. bueanus

| À

L. p. polyacanthus L. wildi

_ Habitus des petits Leptodactylodon à dents vomériennes développées. L. albiventris, ILA 70.975,

le Wom: L. bueanus, ILA 78.155, de Bolifamba ; L. p. polyacanthus, JLA 70.941, de Bafut-

Ngemba : L. wildi, MNHN 2000.0608, de Kodmin. Remarquer la forme de la tête chez L. wildi.

Fig

Source : MNHN, Paris

AMIET & DOWSETT-LEMAIRE 7

Fig. 3. - Leptodactylodon wild: a, pied, face plantaire : b, main, face palmaire ; e, plafond de la cavité

buccale montrant la disposition des dents vomériennes.

Les membres postérieurs sont encore plus courts que chez les autres Leprodactylodon,

avec des cuisses volumineuses, presque ovoides. Les pieds (fig. 3a) sont larges et épais et

portent des orteils qui paraissent largement séparés à leur base car ils sont assez minces et

dépourvus de toute palmure interdigitale. Leurs extrémités sont faiblement dilatées ; les

tubercules sous-articulaires, peu saillants, sont au nombre de trois à l'orteil 4, de deux aux

orteils 3 et 5 et de un aux orteils 1 et 2. Il y a un tubercule métatarsien interne ovoïde,

aillant que les tubercules sous-articulaires.

relativement petit mais un peu plus s

Livrée. C. Wild a photographié un individu vivant (fig. 4) et deux individus fraîchement

sacrifiés, ce qui permet de connaître la coloration de cette espèce in vivo.

Source : MNHN, Paris

8 ALYTES 18 (1-2)

Fig. 4. — Leptodactylodon wildi en vie (photo C. Wild). Remarquer la maculation céphalique caractéris-

tique et l'épaisseur des membres postérieurs.

La face dorsale, moins foncée que chez L. polyacanthus, est brunâtre avec des marbrures

beige olivâtre et quelques taches d’un jaunâtre terne. Ces dernières dessinent un motif

céphalique caractéristique, bien visible même chez les individus fixés, comprenant : (1) sur le

museau, une tache à contour irrégulier, de forme variable ; (2) une large bande interoculaire

avec une dilatation médiane postérieure ; (3) deux bandes partant des yeux et convergeant vers

l'arrière en direction de la région scapulaire en dessinant un X ou un Y ouvert vers l'avant ;

toutes ces macules, mal circonscrites, ont un contour très diffus mais ressortent cependant

bien sur la teinte foncière. L'iris est piqueté d’argent sur un fond brun-noir. Sur le reste du

corps, quelques vagues macules jaunâtres se répartissent plus ou moins suivant deux lignes

latéro-dorsales (caractère chromatique pouvant se retrouver chez L. axillaris Amiet, 1971).

Le dessus des membres antérieurs, la région coccygienne, les talons, la face antérieure des

jambes et le dessus des pieds sont d’un jaune fauve terne, avec des traces de bandes transverses

incomplètes sur les cuisses et les jambes.

La région gulaire, brune, est piquetée de quelques gros points subcirculaires d’un blanc

bleuté, semblables à ceux de L. polyacanthus et L. albiventris. La coloration foncière de la

poitrine et de l'abdomen est d’un blanchâtre un peu translucide, fortement suffusé d’un voile

de chromatophores foncés. Le dessous des membres est d’un jaunâtre fauve, avec de petites

macules diffuses brunes sous les bras et les cuisses, et de larges macules nébuleuses foncées

sous les jambes et les pieds.

Source : MNHN, Paris

AMIET & DOWSETT-LEMAIRE 9

Caractères sexuels secondaires. — (1) Epines nuptiales. — La présence chez les mâles en

livrée nuptiale de deux groupes d’épines coniques, noires, sur la tranche interne de la main et

sur le doigt 1 représente un caractère générique des Leptodactylodon (AMIET, 1980). Chez L.

wildi, le groupe distal comprend 4 épines (5 à une main chez deux individus), comme chez

toutes les autres espèces. Le groupe proximal (“épines métacarpiennes”) en comprend 4. Le

nombre de ces épines métacarpiennes varie suivant les espèces et a de ce fait un intérêt

diagnostique. Dans le cas présent, L. wildi se situe, parmi les petits Leptodactylodon, entre L.

bueanus, à 3 épines, et L. polyacanthus, qui en possède 5 ou, presque aussi souvent, 6.

(2) Hypertrophie brachiale. - Chez de nombreuses espèces de Leptodactylodon, les mâles

se distinguent par l’hypertrophie de leurs bras et de leurs avant-bras. C’est le cas aussi chez L.

wildi, où ce caractère est bien marqué, avec formation d’un “faux coude”, dû à la saillie d’une

des crêtes aliformes de l’humérus (voir à ce sujet AMIET, 1980).

En revanche, L. wildi ne présente pas de protubérances axillaires (excroissances dures

situées à la racine des bras et produites par une apophyse basale de l’humérus). Ce caractère

négatif est cohérent avec ce qu’avaient déjà montré les autres espèces de Leptodactylodon : les

protubérances axillaires manquent chez celles dont les dents vomériennes sont bien dévelop-

pées.

(3) Spinosité cutanée. — Alors que les caractères précédents rapprochent L. wildi de L.

polyacanthus, aspect de la spinosité cutanée l'en sépare nettement. Contrairement à ce

dernier, il n’y a pas de productions fortement kératinisées mais seulement d’infimes spinules

incolores imperceptibles à l’œil nu. Ces productions tégumentaires se trouvent sur le pourtour

des mâchoires supérieure et inférieure, de part et d'autre de l'anus, sur les talons, sur la face

inféro-externe des tarses et sur la plante des pieds. La gorge et la région pectorale, fortement

spinescentes chez diverses espèces et en particulier chez L. polyacanthus, sont totalement lisses

chez L. wildi.

Cavité buccale. — Le développement et la disposition des dents vomériennes fournissent

chez les Leptodactylodon (contrairement aux autres Astylosterninae) de bons caractères

taxonomiques (AMIET, 1980). Ici, elles constituent deux bandelettes subrectilignes largement

séparées et plus courtes que chez les Leptodactylodon du groupe d'ovatus, leur extrémité

externe atteignant à peine le niveau des choanes (fig. 4c). Cette extension est similaire à celle

de L. albiventris et L. polyacanthus.

La langue, cordiforme, est étroitement et profondément échancrée à son extrémité ; elle

ne présente ni papille ni fossette sur sa face supérieure.

Vocalisations. — Les appels les plus souvent entendus sont des “toing” ou “toeng” espacés de

quelques secondes, qui ressemblent curieusement aux sons produits par le relâchement d’une

corde de guitare. Ils durent de 0,2 à 0,45 s et se présentent comme une série d'éléments serrés

dont la fréquence est légèrement relevée vers la fin (fig. 5). La première harmonique est plus

sonore que la fréquence de base (vers 2000 Hz) ; les harmoniques supérieures sont très faibles.

L'enregistrement réalisé le 30 mars 1998 ne comprend pas d'autre type de cri et FDL n'a rien

entendu d'autre sur la période du 30 mars au 3 avril. En décembre 1998, par contre, les appels

de type corde pincée étaient fréquemment suivis d'un ou deux “krier” plus courts (fig. 5).

Chez deux individus voisins dialoguant pendant plusieurs minutes, chacun émettait un

“toing” suivi d’un ou deux “krier” (avec un intervalle entre les notes d'environ une seconde);

Source : MNHN, Paris

10 ALYTES 18 (1-2)

Û 05 L 15 sec

Fig. 5. Leptodactylodon wildi: appel à timbre de corde pincée “toeng” suivi de l'appel plus court “krier”.

Sonagramme produit en bande large sur un spectrographe Key Electric Co. de type 7029A (enregis-

trement et sonagramme F. Dowsett-Lemaire).

l'intervalle de réponse d’un individu à son voisin variait de une à quatre secondes. Les “krier””

sont nettement plus courts, de l’ordre de 0,15 s et n’ont pas le timbre de corde pincée malgré

la similitude des fréquences ; les unités sont en revanche plus serrées que dans l’autre type

d'appel.

Le son de corde pincée est unique dans le genre Leptodactylodon, et même, de façon plus

générale, parmi les anoures camerounais (environ 160 espèces enregistrées par JLA). L'autre

cri, plus comprimé et plus “sec”, est d’un type plus banal chez les Leptodactylodon et rappelle

un peu, entre autres, celui de L. perreti (AMIET, 1980).

Distribution, écologie. — En 1998, l'espèce a été repérée en deux points situés tous deux à

1350 m d'altitude et à proximité du village de Kodmin (*Kumin” sur la feuille Buea-Douala

au 1/200.000; 9°42'E, 4°59°N), dans les monts Bakossi (fig. 6). Cet ensemble montagneux,

relativement peu élevé (il culmine à 1895 m) mais très accidenté, double à l’ouest l'axe

orographique majeur de la Dorsale camerounaise, constitué ici par l’alignement des monts

Koupé, Manengouba, Ekomane, etc.

Dans ce secteur, entre les monts Bakossi et le mont Manengouba tout proche, l'altitude

reste supérieure à 1200 m. Cela signifie que les deux massifs communiquent par un “pont”

d'étage submontagnard (sensu LETOUZEY, 198$), situation qui autorise le passage, dans un

sens ou dans l’autre, de nombreuses espèces orophiles, à l'exception de celles, inféodées à

l'étage montagnard, qui restent isolées sur les parties culminales du Manengouba.

Le premier site où la présence de L. wildi a été constatée est à moins d’un kilomètre du

village de Kodmin. Sa végétation correspond à une forêt basse croissant dans un petit ravin

humide, avec de nombreuses fougères arborescentes (Crathea manniana Hook) et de jeunes

Source : MNHN, Paris

AMIET & DOWSETT-LEMAIRE 11

CU L.p.punctiventris

% _L. albiventris

% L. bueanus

Yaoundé

Fig. 6.- Carte de la partie sud-oue:

du Cameroun, montrant la répartition des petits Leprodactylodon à

dents vomériennes développé

Source : MNHN, Paris

12 ALYTES 18 (1-2)

palmiers (Phoenix reclinata Jacq.). L'eau affleure par endroits, et un minuscule ruisseau coule

en fin de saison des pluies. Dans ce secteur, les espèces compagnes suivantes ont été enregis-

trées par FDL et identifiées auditivement par JLA : Phrynodon sp. 2 (sensu AMEr, 1975),

chantant exactement au même endroit, et, un peu plus bas, Phrynobatrachus cricogaster

Perret, 1957, P werneri (Nieden, 1910), Phrynodon sp. 1 (sensu AMIET, 1975), Leptodactylodon

mertensi Perret, 1959 (?) et Leptopelis brevirostris (Werner, 1898).

Le second site est à environ 1,5 km de Kodmin, en forêt plus fermée (Schefflera abyssinica

(Hochst. ex A. Rich.) Harms, Polyscias fulva (Hiern) Harms, Cyathea manniana), sur le

versant ombragé d’un petit ruisseau. Les espèces compagnes sont ici Phrynodon sp. 1 et sp. 2,

Leptodactylodon mertensi (?), Cardioglossa melanogaster Amiet, 1972, Leptopelis brevirostris,

L. calcaratus (Boulenger, 1906) et Afrixalus lacteus Perret, 1976 (même mode d'identification

que ci-dessus).

En mars 2000, de petites colonies ont été localisées par FDL en quatre endroits dans la

forêt primaire au sud-ouest du village d'Edib (4°57°N, 9°39°E), à des altitudes comprises entre

1000 et 1200 m. Le milieu consistait toujours en zones marécageuses ombragées, un peu à

l'écart des lits de ruisseaux, et à chaque endroit l’espèce co-existait avec Phrynodon sp. 2.

Les mâles de L. wildi chantent au sol, cachés dans des anfractuosités, sous les mousses ou

les racines. Au site où l’espèce a été enregistrée, il y avait au moins une vingtaine de chanteurs

dans un rayon de 10 m. Ils sont actifs à toute heure de la journée, mais davantage par temps

couvert, et peuvent également chanter la nuit, jusqu’à deux heures au moins après le coucher

du soleil.

Affinit Plusieurs espèces de Leptodactylodon sont caractérisées, comme L. wildi, par une

petite taille (inférieure à 30 mm), des dents vomériennes bien développées disposées en crêtes

transverses et une pigmentation terne et peu contrastée de la face ventrale. Il s’agit de L.

albiventris, L. bueanus (décrit initialement comme sous-espèce de L. albiventris mais qui paraît

en réalité spécifiquement distinct de ce dernier) et L. polyacanthus.

Les deux premiers ne sont pas orophiles mais sont localisés dans des régions accidentées

qui reçoivent d’abondantes précipitations, alors que L. polyacanthus est un orobionte qui a

une large répartition horizontale et verticale sur la Dorsale camerounaise. Ces trois espèces

pourraient représenter un ensemble monophylétique un peu comparable à celui que consti-

tuent, par exemple, Astylosternus batesi (Boulenger, 1900), A. diadematus Werner, 1898 et

A. montanus Amiet, 1977 (AMIET, 1977).

L. blanci, récemment décrit du Gabon sur un spécimen mâle (OHLER, 1999), peut être

rapproché de ces derniers par sa petite taille et ses crêtes vomériennes bien développées mais

s'en distingue par sa livrée ventrale vermiculée, patron qui n'était auparavant connu que chez

les “grands” Leptodactylodon; ses vocalisations sont inconnues, mais sa localité d’origine et la

liste des quelques espèces capturées dans le même site montrent qu'il s'agit d’une espèce

planitiaire.

Ilest tentant de considérer L. wildicomme un vicariant géographique de L. polyacanthus.

qui paraît manquer dans la partie sud du noyau principal de la Dorsale camerounaise (son

absence sur le Mont Manengouba est signalée par AMIET, 1980). L'absence de spinosité

corporelle kératinisée chez L. wildi constitue cependant un caractère différentiel important

par rapport à L. polyacanthus, d'autant plus que ce type de caractère sexuel secondaire a

Source : MNHN, Paris

AMIET & DOWSETT-LEMAIRE 13

généralement tendance à s’exacerber chez les anoures orophiles. Cette absence est encore plus

frappante quand on constate que L. wildi est, de tous les Leptodactylodon actuellement

connus, la seule espèce à membres antérieurs hypertrophiés qui soit dépourvue de protubé-

rances axillaires ou de spinosité gulaire et/ou pectorale. L'habitus de L. wildi, avec sa tête assez

étroite et ses membres postérieurs très épais, le distingue aussi des autres petits Leptodacty-

lodon à dents vomériennes développées.

Dans l’état actuel des connaissances, L. wildi peut être rapproché de ces derniers dans une

clé de détermination (voir ci-après) mais cela n'implique pas qu’il leur soit étroitement

apparenté. L'hypothèse qu’il représente à lui seul un petit phylum distinct ne doit pas être

écartée. Elle pourrait être corroborée par la qualité acoustique extraordinaire des vocalisa-

tions qui ont conduit à sa découverte.

CLÉ D'IDENTIFICATION DES PETITS LEPTODACTYLODON

À DENTS VOMÉRIENNES DÉVELOPPÉES

La clé d'identification suivante reprend une partie de celle de AMIET (1980), complétée

par OHLER (1999).

1. Mâles à bras non hypertrophiés, seulement un peu plus épais que ceux des femelles. Sac

vocal développé, sa présence marquée par des plis paramandibulaires. Dessous (sauf la gorge)

po maculé ou fortement vermiculé .

*. Mâles à bras hypertrophiés, sans plis paramandibulaires, Dessous plus ou moins maculé,

mais les taches mal circonscrites, diffuses, en faible contraste avec le fond ................ 4

2. Face ventrale foncée avec des vermiculations claires très apparentes. Taille du seul mâle

connu: 22 mm. Trois épines métacarpiennes L. blanci

2”. Face ventrale, sauf la gorge, non maculée, rose saumon in vivo. Taille moyenne des mâles :

26 mm. Deux ou trois épines métacarpiennes . 53

3. Doigts et orteils munis à leur extrémité de fins prolongements souples. Face dorsale d’un

brun noirâtre uniforme à l'exception d’une barre interoculaire plus claire. Métacarpe des

mâles avec 2 épines. L. albiventris

3. Doigts et orteils sans prolongements $ uples. Face dor: ale bicolore, la partie posté-

rieure du dos, la base des cuisses, etc., plus claires que le reste. Métacarpe des mâles avec 3

. L. bueanus

épines

4. Tête relativement étroite (T/L de l’ordre de 35 %). Taille plus faible : 21,5 à 23 mm chez les

mâles. Face dorsale d’un brun clair olivâtre. Métacarpe avec 4 ou 5 épines chez les mâles, qui

sont dépourvus de spinosité pectorale .… . L. wildi

4’, Tête plus large (T/L de l’ordre de 39 %). Taille plus grande: 24,5 à 26 mm chez les mâles.

Dessus d’un brun presque noir piqueté de points bleutés. Métacarpe avec 5 ou 6 épines chez les

âles, qui sont pourvus de fortes épines pectorales noires réparties en deux plages triangu-

…… L. polyacanthus: 5

Jaires.........,...,s.... ses.

Triangles d'épines pectorales larges mais ne se rejoignant pas sur le milieu de la poitrine.

Face ventrale plus ou moins assombrie mais dépourvue de points ronds éclairci

PES Re L. p. polyacanthus

Source : MNHN, Paris

14 ALYTES 18 (1-2)

5’. Triangles d'épines pectorales étroits, se rejoignant sur le milieu de la poitrine. Face ventrale

ponctuée de clair sur fond sombre, au moins dans la région gulaire ....... L. p. punctiventris

RÉSUMÉ

Un nouveau Leptodactylodon orophile du Cameroun, L. wildi, est décrit à partir de

spécimens trouvés dans les Monts Bakossi (Cameroun). Cette espèce fait partie des petits

Leptodactylodon à dents vomériennes bien développées, mais se distingue des taxons déjà

connus par divers caractères morphologiques (taille plus faible, tête étroite, mâles dépourvus

de spinosité cutanée et de plis gulaires mais à membres antérieurs hypertrophiés) et par ses

appels très différents de ceux des autres Leptodactylodon, comparables au son produit par le

relâchement d’une corde de guitare.

REMERCIEMENTS

Les recherches faunistiques de FDL dans les Monts Bakossi faisaient partie d'une enquête sollicitée

et financée par le Fonds Mondial de la Nature, WWF-Cameroun (représentant local, Dr. J. S. Gartlan),

sous la direction de Chris Wild. Nous remercions ce dernier et ses collaborateurs, D. Menze et E. Abwe,

pour leur efficace contribution. Notre reconnaissance va également à A. Ohler et A. Dubois pour leurs

suggestions concernant le manuscrit de ce travail.

RÉFÉRENCES BIBLIOGRAPHIQUES

Ar, J.-L. 1975. - Ecologie et distribution des Amphibiens Anoures de la région de Nkongsamba

(Cameroun). Ann. Fac. Sci. Yaoundé, 2: 33-107.

—_ 1977. - Les Astylosternus du Cameroun (Amphibia, Anura, Astylosterninae). Ann. Fac. Sci.

Yaoundé, 23-24: 99-221.

1980. — Révision du genre Leprodactylodon Andersson (Amphibia, Anura, Astylosterninae). Ann.

Fac. Sci. Yaoundé, 27: 69-224.

ce 1987. — Aires disjointes et taxons vicariants chez les Anoures du Cameroun : implications paléocli-

matiques. Alpes, 6 (3-4): 99-115.

1989. - Quelques aspects de la biologie des Amphibiens Anoures du Cameroun. Ann. biol., 28 (2):

73-136.

Lerouzey, R., 1985. - Notice de la carte phytogéographique du Cameroun au 1/500.000 (1985). Yaoundé,

Institut de la Recherche Agronomique, & Toulouse, Institut de la Carte Internationale de la

Végétation : 1-240.

Ouen, À., 1999. — Une nouvelle espèce du genre Leptodactylodon (Arthroleptidae, Astylosterninae) du

Gabon. Alyres, 17 (1-2): 73-80.

Corresponding editor: Alain Dunots.

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 15-50. 15

Systematics of Fejervarva limnocharis

(Gravenhorst, 1829)

(Amphibia, Anura, Ranidae)

and related species.

1. Nomenclatural status

and type-specimens of the

nominal species Rana limnocharis

Gravenhorst, 1829

Alain DUBois & Annemarie OHLER

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

A detailed analysis shows that the nominal species Rana limnocharis

was first made nomenclaturally available by Gravenorsr (1829), and then a

second time and independently by Wircmanx (1834). The consequences of

these facts regarding the name-bearing types of these two nominal taxa are

discussed and neotypes are designated for both of them. The status of the

following related nominal species are also discussed, and their type-

specimens are described: Rana cancrivora Gravenhorst, 1829; Rana

vittigera Wiegmann, 1834; Rana gracilis Wiegmann, 1834; Rana multi-

striata Hallowell, 1861; Rana wasi Annandale, 1917. Finally, on the basis

of several recent pieces of information, it is suggested that the group of

frogs usually known as “Rana limnocharis group” or “subgenus Fejerva-

rya” should be recognized as a distinct genus, Fejervarva Bolkay, 1915.

ABBREVIATIONS

MEASUREME)

SVL Snout-vent length.

Head

HW Head width.

HL Head length (from back of mandible to tip of snout).

MN Distance from back of mandible to nostril.

MFE Distance from back of mandible to front of eye.

MBE Distance from back of mandible to back of eye.

Source : MNHN, Paris

16 ALYTES 18 (1-2)

IFE Distance between front of eyes.

IBE Distance between back of eyes.

IN Internarial space.

EN Distance from front of eye to nostril.

EL Eye length.

SL Distance from front of eye to tip of snout.

NS Distance from nostril to tip of snout.

IUE Minimum distance between upper eyelids.

UEW Maximum width of upper eyelid.

Forearm

HAL Hand length (from base of outer palmar tubercle to tip of third finger).

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

Hindlimb

TL Tibia length.

TW Maximum tibia width.

FOL Foot length (from base of inner metatarsal tubercle to tip of fourth toe).

TFOL Length of tarsus and foot (from base of tarsus to tip of fourth toe).

FL Femur length (from vent to knee).

MTTF Distance from distal edge of metatarsal tubercle to maximum incurvation of web

between third and fourth toe.

TETE Distance from maximum incurvation of web between third and fourth toe to tip of

fourth toe.

MTFF Distance from distal edge of metatarsal tubercle to maximum incurvation of web

between fourth and fifth toe.

FETF Distance from maximum incurvation of web between fourth and fifth toe to tip of

fourth toe.

IMT Length of inner metatarsal tubercle.

ITL Inner toe length.

MUSEUMS AND PERSONS

AD Alain Dubois.

AMO Annemarie Ohler.

FMNH Field Museum of Natural History, Chicago, Illinois, USA.

MNHN Muséum National d'Histoire Naturelle, Paris, France.

NMW Naturhistorisches Museum, Wien, Austria.

RMNH Nationaal Natuurhistorisch Museum, Leiden, Netherlands.

ZMB Zoologisches Museum, Berlin, Germany.

ZSI Zoological Survey of India, Calcutta, India.

Source : MNHN, Paris

Duois & OHLER 17

INTRODUCTION

In most publications dealing with amphibians of south and south-eastern Asia, mention

is made of very common small frogs that occur in or around most paddy fields, small ponds

and open aquatic habitats of this region, and which are usually known under the name Rana

limnocharis. Following BOULENGER’S (19204) work, this group was long viewed as a single

species with four subspecies and this species was credited with a very wide distribution, from

Pakistan to China and Japan and to Indonesia. However, on the basis of the study of mating

calls and morphology, DuBois (1975b) showed that, in a very limited region (the small country

of Nepal), no less than four distinct species did occur and had been confused under the name

Rana limnocharis limnocharis. This author later showed that still other species were present in

southern India (DUBOIs, 1984b) and suggested that the whole group was composed of at least

15 species, probably many more (Dugois, 1987, 1992). He further proposed (Dugois, 1984b,

1987, 1992) to remove this group from the genus Rana as understood by BOULENGER (1918,

1920a-b) and to recognize it provisionally as the subgenus Fejervarya Bolkay, 1915 of the

genus Limnonectes Fitzinger, 1843. FI et al. (1991) and YE et al. (1993) elevated this group to

the rank of genus, but incorrectly under the generic name Euphlyctis Fitzinger, 1843 (a name

which in fact applies to another group of species from the Indian region, that are much more

aquatic than Fejervarya and that retain a lateral-line system in adults: see DuBois, 1992).

Finally, Dugois (2000), IsKANDAR (1998, 1999), Fer (1999) and MARMAYOU et al. (2000)

considered Fejervarya a distinct genus.

Within this frame, a question arises: to which species does the specific name Rana

limnocharis apply? Dusois (1984b) suggested that it applies to populations of Java (from

where the species was first described) and possibly of other regions, but that more work was

necessary to establish the range of the “true” Rana limnocharis. The electrophoretic and

morphometric data of Topa et al. (1998) and of VEITH et al. (2000) complicate this situation,

as they show that two different species of this group live in Java. Which one should bear the

name Rana limnocharis? Answering this question requires the clarification of the nomencla-

tural status and authorship of the name Rana limnocharis, to establish whether type-

specimens of this nominal species can be identified and studied, and, if the answer to the last

question is negative, to designate and describe a neotype for this taxon.

THE QUESTIONS

A great confusion exists in the literature regarding the nomenclatural status and author

of the name Rana limnocharis. This name has been credited so far with seven different

authorships: (1) “Kuhl” (without reference to a published text): (2) “Boie” (without reference

to a published text); (3) WIEGMANN (1834, often miscited as 1835": see ZHAO & ADLER, 1993:

411-412); (4) Boie in Wi£GMANN (1834); (5) Kuhl in GRAVENHORST (1829); (6) GRAVENHORST

(1829); (7) Boie in GRAVENHORST (1829).

Source : MNHN, Paris

18 ALYTES 18 (1-2)

() The name Rana limnocharis first appeared in GRAVENHORST (1829: 42), who credited

it to “Kuhl” and stated that this was a manuscript name appearing in an unpublished

manuscript by Boie, where a closely related species was also described under the name Rana

cancrivora.

@) Shortly after, several authors (WAGLER, 1830: 203; TscHuDi, 1838: 79; DUMÉRIL &

BIBRON, 1841: 376, 379) mentioned the name Rana limnocharis as a label name credited to

“Boie” that they had seen in the Leiden Museum. WIEGMANN (1834: 255-258; 1835: 277-278)

compared his new species Rana vittigera and Rana gracilis to “ Rana limnocharis H. Boje” or

“Rana limnocharis Boie”, a name for which he did not provide a reference. Then this name was

forgotten for over 20 years, and the name Rana gracilis Wiegmann, 1834 was used for the

species it denotes, until the name Rana limnocharis was resurrected by PETERS (1863: 77-78;

1871: 647), who cited WiEGMaNN’s (1834) text, credited the name to “Boie in Leyd[en]

Museum”, and stated that it should replace the name Rana gracilis. This was followed by

SToLiczKa (1872: 102; 1873: 112), who however introduced the incorrect subsequent spelling

Rana lymnocharis and credited it to “Boie”, still without a reference.

(3) After having used the name Rana gracilis (BOULENGER, 1882: 28), BOULENGER (1890:

450) resurrected the original spelling Rana limnocharis and credited authorship of this name

to WiEGMANN (1834). This interpretation was followed, among others, by STEINEGER (1907:

127, 1910: 95), BARBOUR (1912: 64), BOULENGER (1912: 236), SmirH (1916: 165), ANNANDALE

(1917: 132), KIRTISINGHE (1957: 38), NAKAMURA & UÉNO (1963: 49) and GORHAM (1974:

146).

(4) BOULENGER (19204: 28) presented a slightly different interpretation, since he credited

the species’s name to Boie in WIEGMANN (1834). This was accepted by many authors,

including VAN KAMPEN (1923: 167), Liu (1950: 315), INGER (1954: 267-268, 1966: 205), Liu &

Hu (1961: 139), Berry (1975: 73), ANONYMOUS (1977: 81), FRosT (1985: 500), MAEDA &

Matsui (1989: 108), Ft et al. (1991: 302), YANG (1991: 131), YE et al. (1993: 249), ZHAo &

ADLER (1993: 144) and DUTTA & MANAMENDRA-ARACHCHI (1996: 91).

(5) STEINEGER (1925: 27) was the first to point out that the name Rana limnocharis had

first been published by GRAVENHORST (1829), and that the latter had credited this name to

Kuhl. In the synonymy of this species, he therefore wrote its full original name as follows:

Rana limnocharis *Kuhf Gravenhorst”, This writing was also used by FANG & CHANG (1931:

111).

(6) However, many subsequent authors only mentioned GRAVENHORST (1829) as author

of the name, without mentioning Kuhl's “original authorship”: this was the , among

others, of GEE & BORING (1929: 30), PoPE (1931: 491), BORING et al. (193 CHaNG & HSÛ

(1932: 174), BORING (1934: 20, 1945: 82), PoPe & BORING (1940: 50), BOURRET (1942: 249),

TayLOR & ELBEL (1958: 1051), TAYLOR (1962: 380), OKaDA (1966: 112), DuBois (1984b: 143,

1992: 315), CHou & Lin (1997: 27), Durra (1997: 133), MANTHEY & GROSSMANN (1997: 97)

and FE1 (1999: 182).

(7) Finally, Dugots (1974: 382-383. 1981: 238) cited this name as * Rana limnocharis Boie

in GRAVENHORST, 1829”.

Despite this great diversity of interpretations, few discussions were clearly devoted to the

correct authorship of the name.

Source : MNHN, Paris

Dugois & OHLER 19

INGER (1954: 267-268) stated that the first “adequate description” of Rana limnocharis

was to be found in WIEGMANN's (1834) text, where the name was credited to Heinrich Boie.

INGER (1954) reported having seen a copy of the unpublished manuscript of Boie’s original

description of Rana limnocharis, and he concluded that the name Rana limnocharis should be

credited to Boie in WIEGMANN (1834).

Dugois (1974: 382-383) noted that the first published mention of the name Rana

limnocharis was in GRAVENHORST (1829), but that this latter author, although not very

explicitly, credited it to Boie: he therefore suggested to cite this name as “Rana limnocharis

Boie in GRAVENHORST, 1829”. Subsequently however (DuBois, 1984b), he realized that

GRAVENHORST (1829) was responsible both for first publication of the name and for satisfying

the criteria of its availability, and was therefore its sole author, in the technical nomenclatural

sense of this term. However, he did not provide a detailed explanation of these reasons to

reject INGER’s (1954) interpretation.

ZHao & ADLER (1993: 144) concurred with INGER (1954), and provided several reasons

for refusing to credit GRAVENHORST (1829) with the authorship of the valid name of this

species: “The name was introduced in Gravenhorst's synonymy of R. cancrivora, not as a

proper species name, but as a description of frogs living in small pools (hence his use of the

word ‘Range’ rather than Rana). Furthermore, we regard Gravenhorst's short description as

unidentifiable. Boie’s name, accompanied by a full description and a figure, is the first clear

association of the name À. limnocharis with this taxon.” In their synonymy of Rana limno-

these authors listed two distinct nominal species, each one with its own author and

first “ Ranae limnocharis Gravenhorst, 1829” and second “Rana limnocharis Boie in

MANN, 1834”. According to these authors, only the first of these two nominal species has

a clear type locality (Java), while for the second one they wrote: “Type locality: none given”.

Finally, they stated that the first name was a “nomen dubium” and they wrote the valid name

of this species as follows: “ Rana limnocharis Boie, 1834”.

charis

These discussions may appear exaggeratedly quibbling, if not gratuitous, but they are

not: according to the interpretation chosen, the nominal species Rana limnocharis may have

four different authors and two different dates, and more importantly, it may be based on four

different name-bearing types. If Kuhl is retained as author of the name, the type-specimens of

the nominal species will be the specimens collected by Kuhl & Van Hasselt, and distributed

later in several museums: if Boie is the author, only those specimens kept in Leiden when he

prepared his description and figure will be types; if Gravenhorst is the author, the name-

bearing type will be the specimens examined by this author in Breslau: finally, if Wiegmann is

the author, it will be the specimens examined by this author in Berlin. According to the

interpretation chosen, different specimens will have to be considered types, and in some cases

all types will be lost: this will have consequences regarding the possible choice of a lectotype

or neotype for the nominal species Rana limnocharis and the allocation of this name to one of

the two biological species occurring in Java. A detailed analysis of the history of the case and

of the various problems pointed out above regarding the availability of names is therefore in

order before any such designation of lectotype or neotype. To avoid criticism, this discussion

must be made strictly within the frame of the current Znternational Code of Zoological

Nomenclature (ANONYMOUS. 1999; cited below as “the Code”), which means that some

technical nomenclatural terms and rules will have to be mentioned below.

Source : MNHN, Paris

20 ALYTES 18 (1-2)

NOMENCLATURAL STATUS AND AUTHORSHIP OF THE NAME RANA LIMNOCHARIS

HISTORICAL SURVEY OF THE DISCOVERY, COLLECTING AND NAMING OF RANA LIMNOCHARIS

The first documented discovery and collection of Rana limnocharis was by H. Kuhl &

J.C. Van Hasselt, during their brief stay in Java (respectively 1820-1821 and 1S°1-1823) which

ended with the death of both of them (see e.g.: BRONGERSMA, 1942; ADLER, 159). These two

naturalists collected several specimens of a small species of frogs common around paddy

fields, for which they apparently coined the name Rana limnocharis. However, unlike for other

amphibian species (see e.g. DuBois, 1982), this name was not mentioned in the copies of the

letters sent by them to Europe that were published in three different zoological journals

(KuHL & VAN HASSELT, 1822a-b; VAN HASSELT, 1823; KUHL, 1824a-b). They sent specimens

of this species to the Rijksmuseum van Natuurlijke Historie (now the Nationaal Naturhisto-

risch Museum) in Leiden, where these were apparently labelled under two different names,

“Rana cancrivora” for the large ones and “ Rana limnocharis” for the small ones. Apparently,

in this collection the second of these names was credited to Kuhl, as is implied by the mention

of “Ranae limnocharis Kuhlii” in GRAVENHORST (1829: 42), while the name Rana cancrivora

seems to have been coined by Heinrich Boie, in the manuscript of his Erpétologie de Java: this

latter book, announced by SCHLEGEL (1826, 1827) and GRAVENHORST (1829), was never

published, although it had been sent to the printer in 1830 (see BRONGERSMA, 1942). The fact

that specimens labelled under the two above names were kept in the Leiden Museum was

reported by WAGLER (1830: 203) and TscHUDI (1838: 39, 79). Furthermore, according to

SCHLEGEL (1827: 282) and DUMÉRIL & BIBRON (1841: 379), some other specimens of this

group collected by Kuhl & Van Hasselt were also sent to other European Museums: this is

precisely documented at least in two cases, for two specimens in the Breslau (now Wroclaw)

Museum mentioned by GRAVENHORST (1829: 41-42) and for two specimens in the Berlin

Museum mentioned by WiIEGMANN (1834: 57-58).

The first publication of the name Rana limnocharis Was by GRAVENHORST (1829: 42).

This name was only briefly mentioned in the chapter dealing with a species described as new

under the name Rana cancrivora, where, after a Latin diagnosis of the latter species, one can

read: “Hujus speciei Javanensis de Haan duo individua mecum communicavit; unum, idque

majus, capite vix paulo obtusiore, corporis totius facie infera fusco- et fusco-ferrugineo-

maculata, sub nomine Ranae cancrivorae; minus, idque facie infera corporis albida immacu-

lata, sub nomine Ranae limnocharis Kuhlii. Conferantur quae ad Hylam leucomystacem, n.4,

monui.” (GRAVENHORST, 1829: 41-42). Above in the same volume, the following appears

under Hyla leucomystax: “Cum de Haan, conservator humanissimus musei Lugdunensis.

hanc Hylam leucomystacem Kuhlii javanensem, pluresque alias species, Javae indigenas, ad

me transmitteret, simul me certiorem faciebat, Boieum descriptiones et icones reptilium

novorum Javanensium, in peculiari Erpetologia, editurum esse. Inde harum specierum solas

diagnoses cireumscriptas proferam, ne auctori Erpetologiae temere antevertam. Utinam opus

exoptatissimum mox in lucem proderat! Conferas conspectum hujus Erpetologiae, quem

Schlegel in Bulletin des Sci. nat. 1826, n. 10, pag. 233-240 edidit.” (GRAVENHORST, 1829: 26).

Source : MNHN, Paris

Dupois & OHLER 21

These statements can be summarized as follows: (1) de Haan, Curator at the Leiden

Museum, had sent specimens, including several of the new frog species collected in Java by

Kuhl& Van Hasselt, to the Breslau Museum; (2) while doing so, he had informed Gravenhorst

in Breslau that, in a book already written but not yet published (and summarized by

SCHLEGEL, 1826), Boie had provided descriptions and figures of several new herpetological

species from Java; (3) in the expectation of the publication of this book, Gravenhorst decided

to publish only short diagnoses of the new Javanese species he had received from de Haan; (4)

among those were two frog specimens, a large one under the name Rana cancrivora and a

smaller one under the name Rana limnocharis; (5) Gravenhorst’s opinion was that these two

specimens, which also differed by the shape of the snout and the colour of the lower parts of

the body, belonged to a single species, for which he chose the name Rana cancrivora and

provided a Latin diagnosis.

As mentioned above, ZHAO & ADLER (1993) considered the status of the name Rana

limnocharis in GRAVENHORST (1829) as questionable, and decided not to apply this name to

the species. Let us consider their arguments.

AN ANALYSIS OF ZHAO & ADLER'S (1993) STATEMENTS

Several distinct reasons for not recognizing GRAVENHORST (1829) as the author of the

name Rana limnocharis can be sorted from ZHAO & ADLER'’s (1993) short statements (see

above). Let us distinguish them and clearly formulate them in precise technical nomenclatural

terms.

(1) Zao & ADLER (1993) first stated that the name Rana limnocharis “was introduced in

Gravenhorst's synonymy of R. cancrivora”. Although they did not discuss this point further,

this statement can be understood as meaning that the name Rana limnocharis was not made

nomenclaturally available in GRAVENHORST'S (1829) work for the mere reason that it had been

introduced there as a synonym.

(2) The next statement of ZHAO & ADLER (1993) is that the name Rana limnocharis was

proposed “not as a proper species name, but as a description of frogs living in small pool

Strictly taken, this statement does not mean much, since, of course, a new species name can

well be proposed for frogs living in small pools. What ZH4O & ADLER (1993) apparently meant

was that the two words Rana limnocharis were not proposed as the name of a new species-

group taxon, but merely as a statement aiming at giving some biological characterisation of a

frog species, which otherwise was remaining unnamed. In nomenclatural terms, this means

that in GRAVENHORST (1829) the combination Rana limnocharis was a kind of “formula”

without nomenclatural status, i.e. that the name Rana limnocharis Was nomenclaturally

unavailable in this text.

(3) In support of this interpretation, ZHAO & ADLER (1993) claimed that the use of the

word *Ranae° ead of Rana shows that GRAVENHORST (1829) referring to “frogs”

rather than to a frog species. In other words, and to put this in grammatical and nomenclatural

terms, they apparently believed that “Ranae limnocharis” was à nominative plural, and

therefore, for this mere reason, nomenclaturally unavailable under the Code for the name of a

new species-group taxon. Although they did not state this in full words, it seems that what

Source : MNHN, Paris

22 ALYTES 18 (1-2)

ZHao & ADLER (1993) had in mind is the fact that Article 11.9.1.1 of the Code requires that,

to be available, a new species-group name should be in the nominative singular.

(4) Statements (1) to (3) tend to show that ZHAO & ADLER (1993) regarded the name Rana

limnocharis as nomenclaturally unavailable. However, in their next sentence, as well as in the

synonymy of the species, they adopted another interpretation, since they insisted on the fact

that the short description of Rana limnocharis in GRAVENHORST (1829) was “unidentifiable”

and that this name was therefore a “nomen dubium”’. This interpretation is quite different

from, and actually contradictory to the previous one: according to the Code (ANONYMOUS,

1999: 111), a “nomen dubium” is a “name of unknown or doubtful application”, i.e. a name

nomenclaturally available but whose allocation to a biological taxon is impossible or doubtful.

For this reason, Recommendation 75.E of the 1985 edition of the Code (ANONYMOUS, 1985:

163) aptly suggested to designate neotypes for species-group nominal taxa “to clarify the

application of names when their continued existence as nomina dubia threatens the stability

of other names”, a formulation that has unfortunately disappeared in the last edition of the

Code.

To sum up, statements (1) to (3) of ZHAO & ADLER (1993) support the idea that the name

Rana limnocharis is nomenclaturally unavailable in GRAVENHORST's (1829) text, while their

statement (4), as well as their inclusion of this name in their synonymy of the species, supports

the opinion that this name is nomenclaturally available but of doubtful allocation to à

biological species, and therefore cannot be used. An important weakness of this interpreta-

tion is its failure to address the following problem: if two distinct nominal species are to be

recognized, the second name is a junior primary homonym of the first one and therefore an

unvalid name. In other words, if, as clearly implied by the end of their text, the name “Rana

limnocharis Gravenhorst, 1829” was both (1) an available name and (2) inappropriate for the

species, being a “nomen dubium””, then the species would have to bear another name: as the

only other synonym, beside “ Rana limnocharis Boie, 1834”, listed by ZHAO & ADLER (1993),

is Rana gracilis Wiegmann, 1834, which is also a primary homonym in the genus Rana (see e.

Dugois, 1984b: 154), no name would be lable for the species and a new name would have

to be coined for it!

IS THE NAME RANA LIMNOCHARIS IN GRAVENHORST (1829) AVAILABLE UNDER THE CODE?

Let us first consider the statements (1) to (3) of ZHao & AbrER (1993). According to these

statements, the name Rana limnocharis Would be nomenclaturally unavailable in GRAVEN-

HORST'S (1829) text for three distinct but complementary reasons: (1) this name was published

there as a synonym; (2) this name would not have been proposed to designate a frog taxon, but

merely to refer “informally” to a “kind” of frogs without “naming” them; (3) this name

would appear in GRAVENHORST'S text as a nominative plural, not nominative singular as

required by Article 11 of the Code.

(1) As correctly stated by ZHAO & ADLER (1993), the name Rana limnocharis was

first published by GRAVENHORST (1829) as a junior synonym of the name Rana cancrivora.

What are the nomenclatural consequences of this fact? The Code is quite clear about the

nomenclatural status of names first published as synonyms. Article 11.6 reads as follows:

“A name [...] first published [...] as a junior synonym [...] is not thereby made available” unless

it has been “treated before 1961 as an available name and either adopted as the name of a

Source : MNHN, Paris

Dugois & OHLER 23

taxon or treated as a senior homonym”; such a name “dates from its first publication as

a synonym”. The name Rana limnocharis mentioned in GRAVENHORST (1829) clearly

corresponds to this situation, since it has regularly been treated as an available name since

STEINEGER (1925); it therefore dates from its first publication as a junior synonym of Rana

cancrivora.

(2) Close examination of the whole book of GRAVENHORST (1829) also allows to

unambiguously reject interpretation (2). The name “ Ranae limnocharis” is composed of two

words, a generic name bearing a capital and a specific name starting with a lower-case letter,

as are all other scientific names of species in the book. This name is printed with wide spaces

between letters, which would correspond to italics in modern printing: in GRAVENHORST'S

book, such a way of printing is used only for scientific names of taxa and for some other words

that the author wanted to stress as particularly important in his text. Finally, this name is

followed by the mention “Kuhlii”, i.e. a genitive meaning “of Kuhl”, clearly indicating that in

GRAVENHORST'S mind Kuhl was the author of this name. The name “ Ranae limnocharis” was

thus clearly intended to designate a taxon. Nothing in this text gives the slightest support to

the interpretation that this name would be a “formula” informally designating a “kind” of

frogs.

(3) ZHao & ADLER (1993) are also incorrect when they consider the name “Ranae

limnocharis” to be a nominative plural meaning “frogs living in small pools”. As was

underlined by Bour & DuBois (1984), the Code’s requirement that species-group names, to be

nomenclaturally available, be published in the nominative singular, makes sense only when the

whole text is written in a language other than Latin: in a Latin text, the grammatical case of

words is determined by their place in the sentence, and only names occupying the place of

subjects can be written in the nominative; in all other situations, the grammatical case of

names will have to be different. We are here exactly in this situation: actually, considering the

structure of GRAVENHORST'S Latin sentence (quoted above), it is clear that the name “ Ranae

limnocharis”? was in the genitive singular and meant “of Rana limnocharis”. AN the text of

GRAVENHORST (1829) being written in Latin, in the sentence as it was written the use of the

genitive singular was compulsory. This case corresponds to the situation described in Article

11 of the Code: “A genus-group name proposed in Latin text but written otherwise than in the

nominative singular because of the requirements of Latin grammar is available, provided that

it meets the other requirements of availability, but it is to be corrected to the nominative

singular.” (Article 11.8.1); “An adjectival species-group name proposed in Latin text but

written otherwise than in the nominative singular because of the requirements of Latin

grammar is available provided that it meets the other requirements of availability, but it is to

be corrected to the nominative singular if necessary.” (Article 11.9.2).

Reasons (1) to (3) given by ZHAO & ADLER (1993) to refuse nomenclatural availability of

the name Rana limnocharis in GRAVENHORST (1829) are therefore to be rejected. Could there

be other reasons to refuse this availability? In other words, does this name meet the “other

requirements of availability” mentioned in the Code? The answer to this question is clearly

“yes”: all criteria mentioned in Articles 10, 11 and 12 of the Code are met with.

In conclusion of this section, the name Rana limnocharis as published in GRAVENHO

(1829) text is clearly available in zoological nomenclature. Let us now see to which taxon this

name applies.

Source : MNHN, Paris

24 ALYTES 18 (1-2)

IS THE NAME RANA LIMNOCHARIS IN GRAVENHORST (1829) À ‘NOMEN DUBIUM”?

To be sure, the “description or definition” (in the sense of Article 12 of the Code) given

to diagnose Rana limnocharis in GRAVENHORST (1829) is very short and vague, and liable to

cause problems for the aflocation of this name to a biological taxon, although it does not pose

a problem regarding the nomenclatural availability of the name. DuBois & OHLER (1995,

1997a-b) discussed the problems posed by old names based on incomplete or insufficient

descriptions. They remarked that the nomenclatural allocation of a name to a taxon does not

rely on its description, definition or diagnosis, but on its type-specimens, either actual or

potential, and through these specimens to the type-population from which these had been

collected. Actually, in frogs, many taxa named in the 18" and early 19° century were first

provided with very short and fully insufficient diagnoses or descriptions, and the types have

often been lost, but this has no bearing on the availability of names: in most of these cases, the

status of these names was later fixed through redescriptions by the same or other authors,

re-examination of holotypes or syntypes, or designation of neotypes. As short as it is, the

statement that Rana limnocharis is smaller than Rana cancrivora and has a different snout

shape and belly coloration is enough to make the former name nomenclaturally available,

even if it is not enough to ascertain the taxonomic allocation of this name to a biological

species. To solve the problem of the allocation of the name Rana limnocharis to a frog taxon,

it is necessary to identify the name-bearing type of this taxon. As mentioned above, the status

of this/these type-specimen(s) is directly linked to the authorship of this name. Before

addressing this question however, let us consider more generally the rules governing

authorship in zoological nomenclature according to the current Code.

AUTHORSHIP OF NAMES “BORROWED®” FROM MANUSCRIPTS OR COLLECTION LABELS

Article 50.1 of the Code provides the following definition of “author” in zoological

nomenclature: “The author of a name [...] is the person who first publishes it [..] in a way that

satisfies the criteria of availability However, if it is clear from the contents [of the

publication] that some person other than an author of the work is alone responsible both for

the name [...] and for satisfying the criteria for availability other than actual publication,

then that other person is the author of the name [...]."” Particularly important, and often

overlooked by taxonomists, are the terms “alone responsible”. These statements mean that

the author of a scientific name according to the Code is not any of the following: (1) the

person(s) who actually coined the name, or the person(s) who wrote the first unpublished

description or definition of the taxon, or provided any other information that could be an

indication making the name available under the Code, unless in the first valid publication of

the name it was made quite clear, in full words, that both the name and the published

description, definition or indication were directly copied, without any modification (i.e.,

“verbatim”), from this unpublished document: (2) any person who could have used this name

in conversations, meeting or unpublished documents, such as letters or labels attached to

specimens in zoological collections.

According to such stringent rules, the case is much rarer indeed than is often believed by

where a situation qualifies for authorship of a name being validly stated to be

many z00logi

Source : MNHN, Paris

Duois & OHLER 25

“XXX in ZZZ”. This applies to cases where a name was first published by an author who

stated clearly (1) that this name was provided to him/her privately by another colleague, but

(2) that he/she personally had never seen specimens of the taxon. A good example of this

situation is the European frog name Rana dalmatina, which was first validly published by

BONAPARTE (1838) in a text where he stated that he had not seen this species, but that the latter

had been mentioned to him, and briefly described, by Fitzinger in an unpublished letter: this

species must therefore be known as Rana dalmatina Fitzinger in BONAPARTE, 1838 (see

Dusois, 19844). But this situation is rather rare indeed. In most cases, the author who first

published a name had seen specimens of the taxon, and added (or may have added) some

observations or statements of his/her own concerning the latter: in such cases, even if this

author credited the new name to the person who had coined it, in the strict sense of the Code

the author is the person who published it. Such a practice of publishing manuscript names

credited to other authors was very common in the early 19° century. Several examples of this

situation can be found e.g. in TscHuDr's (1838) work. Tschudi “borrowed” a number of names

from DUMÉRIL & BIBRON'S (1841) then still unpublished book and from other manuscripts:

although he duly credited these names to their “proper” authors such as Bibron or

Schlegel, Tschudi had clearly examined personally specimens of these taxa, and used these

observations to write their brief descriptions or diagnoses, so that he is no doubt the author,

in the technical nomenclatural sense of the term, of these names.

A particular case must however be made for names first published as junior synonyms.

Article 50.7 of the Code reads as follows: “If a scientific name (taken, for example, from a label

or manuscript) was first published in the synonymy of an available name and became available

before 1961 through the provisions of Article 11.6, its author is the person who published it as

a synonym, even if some other originator is cited, and is not the person who subsequently

adopted it as a valid name [...]).” Therefore, in the case of a name first published as a junior

synonym and a posteriori validated through use by subsequent authors, in all cases the author

is the person who first published it, and in no case the writing “XXX in ZZZ” can be used.

WHO IS THE AUTHOR OF THE NAME RANA LIMNOCHARIS IN GRAVENHORST (1829)?

It was shown above that the name Rana limnocharis was nomenclaturally available from

the publication of GRAVENHORST (1829). The question now remains: who is the author, in the

technical sense of this term in the Code, of this name? Four possible authorships can be

considered: GRAVENHORST (1829), Kuhl in GRAVENHORST (1829), Kuhl & Van Hasselt in

GRAVENHORST (1829) and Boie in GRAVENHORST (1829).

While it is difficult at present to ascertain who really coined the name Rana limnocharis,

GRAVENHORST (1829) provided two different pieces of information in this respect: (1) this

name was attached to the label of one of the frog specimens sent by de Haan to the Breslau

Museum, and it was stated to be one of the names of taxa described as new by Boie in his

unpublished Erpérologie de Java: (2) however, by writing “Kuhlii” after this name, Graven-

horst clearly suggested that the name had been coined by Kuhl, not by Boie. The most likely

explanation of this apparent contradiction is that Boie himself, in his manuscript description

of the species, credited its name to Kuhl (or to Kuhl & Van Hasselt), possibly because this

name was already present in Kuhl's labels or manuscript notes sent with the specimens from

Source : MNHN, Paris

26 ALYTES 18 (1-2)

Java. Whatever the case may be, this is of purely historical but of no nomenclatural impor-

tance. The name Rana limnocharis having been published in GRAVENHORST (1829) as a junior

synonym and a posteriori validated by STEINEGER (1925) and other subsequent authors,

according to Article 50.7 the author of this name is clearly GRAVENHORST (1829), irrespective

of who had coined it before its first publication.

THE STATUS OF THE NAME RANA LIMNOCHARIS BETWEEN 1829 AND 1863

After the book of GRAVENHORST (1829), the first published occurrence of the name Rana

limnocharis was in WAGLER (1830: 230), who listed this species as valid among the species of

the genus Rana. WAGLER (1830) did not refer however to GRAVENHORST'S (1829) text, but to

an unpublished label by Boie in the Leiden Museum, and he provided no description,

diagnosis or indication characterising the species. It cannot therefore be argued that WAGLER

(1830) referred to the name Rana limnocharis Gravenhorst, 1829, and the name Rana

limnocharis in his text must be considered a nomen nudum, without status in nomencla-

ture.

The situation is different in WIEGMANN'S (1834) book, the second published text provid-

ing descriptive data on Rana limnocharis. WiEGMANN described two new species, Rana

vittigera and Rana gracilis, both of which he compared with “Rana limnocharis Boie”. He

stated that the Berlin collection possessed two specimens of the latter species: most probably,

although this is not stated in this text, these specimens were also part of those collected by

Kuhl & Van Hasselt in Java and had been obtained from the Leiden Museum. Later in the

same text, WIEGMANN (1834: 260) stated that he had received the new species “Hyla quadri-

lineata H. Boje in Mus. Lugd.” from Wagler (who was in München), thus clearly indicating the

presence in the Berlin collection of specimens from Boie’s material.

What is the status of the name Rana limnocharis in Wi£GMANN (1834)? Although he

credited the name to Boie, WIEGMANN (1834) did not cite the manuscript of the Erpétologie de

Java, nor GRAVENHORST'S (1829) book. The absence of any reference to the latter work in the

whole text precludes considering WiEGMANN (1834) as having used the name Rana limnocharis

Gravenhorst, 1829. Rather, this author used an unpublished label or manuscript name, which

he made nomenclaturally available, independently from GRAVENHORST (1829), by publishing

rather detailed descriptive data on this species. Therefore, WIEGMANN (1834) created a new,

distinct, nominal species. Who is the author, in nomenclatural terms, of this name? The

situation here is different from that discussed above for Rana limnocharis Gravenhorst, 1829,

since in WIEGMANN'S case the new name was not published as a junior synonym but as a valid

name, credited to Boie: in this case, Article 50.1 of the Code applies, and despite WIEGMANN'S

himself crediting this name to Boie, there is no doubt that the author is WIEGMANN (1834),

who described several precise morphological features of the species, clearly drawn from

personal examination of the two specimens in his hands.

WIEGMANN (1835: 277-278) summarized his 1834 work and mentioned again the name

Rana limnocharis. This name then appeared twice as a junior synonym in TscHUDI (1838: 79)

and in DUMÉRIL & BIBRON (1841: 376, 379), who referred to unpublished manuscripts or

labels, and was then apparently ignored by all authors until PETERS (1863) resurrected it and

Source : MNHN, Paris

Dugois & OHLER 27

cited WIEGMANN'S (1834) text. From 1863 this name was no longer used as a nomen nudum,

but as a name considered validly published in WIEGMANN (1834), or, after STEINEGER (1925),

in GRAVENHORST (1829).

STATUS OF THE ORIGINAL NAME-BEARING TYPES

As mentioned above, identification of the proper author, in the precise nomenclatural

sense of the term, of a scientific name, is crucial, as it implies identification of the proper

name-bearing type of the nominal taxon, which ultimately allows proper allocation of the

name to a biological taxon.

The analysis above has shown that two distinct nominal species bearing the name Rana

limnocharis, with different authors and dates, should be recognized: it results that both

nominal species have their own name-bearing type, which must be identified.

THE ORIGINAL NAME-BEARING TYPE OF RANA LIMNOCHARIS GRAVENHORST, 1829

The name Rana limnocharis Gravenhorst, 1829 was first published as a junior synonym

and therefore falls under the provisions of Article 72.4.3 of the Code: “The type series of a

nominal species-group taxon of which the name was first published as a junior synonym, but

was made available before 1961 under the provisions of Article 11.6, consists of the specimen

(or specimens) cited with that name in the published synonymy, or, if none was cited there,

denoted by that name when it was adopted as the name of a taxon”.

The situation in GRAVENHORST (1829) is particularly clear, as this author stated in full

words that he had received from de Haan a single specimen labelled Rana limnocharis. This

specimen was therefore the holotype by monotypy of Rana limnocharis Gravenhorst, 1829.

This specimen was kept in the Breslau (now Wroclaw) Museum, and apparently no informa-

tion about it was published posterior to GRAVENHORST's (1829) book. In reply to a request of

23 May 1997, on 6 June 1997 Prof. Andrej Wiktor (Muzeum Przyrodnicze, Uniwwersytet

Wroclawski) informed one of us (AD) by letter that the only remaining specimens of the

Gravenhorst collection are some insects, and that attempts to find specimens of other groups

before the Second World War were unsuccessful.

The holotype of Rana limnocharis Gravenhorst, 1829, collected in Java by Kuhl and Van

Hasselt between 1820 and 1823, must therefore be considered lost, and definitive stabilization

of the status of this name will require the designation of a neotype.

THE ORIGINAL NAME-BEARING TYPE OF RANA LIMNOCHARIS WIEGMANN, 1834

According to the anal presented above, the name Rana limnocharis Wiegmann, 1834

based on descriptive information published by WIEGMANN (1834) after examination of

two specimens of * Rana limnocharis Boie” in the Berlin Museum, presumably collected by

Kuhl and Van Hasselt in Java and received from the Leiden Museum. These two specimens

Source : MNHN, Paris

28 ALYTES 18 (1-2)

were therefore the original syntypes of this nominal species. These two specimens are

apparently lost: according to Rainer Günther (e-mail to AMO of 30 November 1998), the

Berlin Museum collection does not have a single specimen of Rana limnocharis collected by

Kuhl or Kuhl and Van Hasselt in Java. Final stabilization of the status of this name also

requires the designation of a neotype.

SPECIMENS AVAILABLE FOR NEOTYPE DESIGNATIONS

In order to definitely avoid possibilities of nomenclatural confusion, and to know which

of the two “sibling” species in Java should bear the name Rana limnocharis Gravenhorst, 1829

(see Verru et al., 2000), designation of a neotype for this nominal species is necessary. As for

the name Rana limnocharis Wiegmann, 1834, being a junior primary homonym it is an invalid

name and its existence does not threaten the stability of nomenclature, but, in order to know

in which synonymy it will have to stand, a neotype designation is also necessary. The most

logical action is to place it in the synonymy of Rana limnocharis Gravenhorst, 1829. Since

neotypes have to be designated for both nominal species, the most parsimonious solution is to

choose the same specimen as neotype of both: these two names will then be linked by an

objective synonymy and no further discussion of their status should arise in the future.

Which specimen would be best suited for this neotype designation? Both nominal species

were created on the basis of specimens collected in Java by Kuhl and/or Van Hasselt and sent

to the Breslau and Berlin Museums from the Leiden Museum. These specimens being lost, it

seems appropriate to look for other specimens collected in Java by these naturalists and kept

in the Leiden Museum or in other museums under the name Rana limnocharis, or possibly

also of Rana cancrivora (as both species were considered synonyms by some ancient authors,

including GRAVENHORST, 1829). Both SCHLEGEL (1827) and DUMÉRIL & BIBRON (1841) stated

that such specimens had been sent to several other European museums, but unfortunately

these authors did not specify which ones. Published and unpublished information was

therefore gathered about this question, with the following results: (1) no specimens under

these two specific names and collected in Java by Kuhl and/or Van Hasselt are to be found in

the old collections of the museums of Basel (M R, 1878, 1880, 1882, 1883, 1885, 1887,

1889, 1892, 1901), Frankfurt am Main (BOETTGER, 1892; MERTENS, 1967; AD & AMO,

personal observations), London (Nick Arnold, e-mail to AMO of 27 March 1998), München

(Frank Glaw, e-mail to AD of 31 March 1998), Paris (AD & AMO, personal observations)

and Wien (Heinz Grillitsch, e-mail to AD of 24 March 1998); (2) the Leiden Museum still has

a single specimen of this group, RMNH 4287 (Marinus S. Hoogmoed, e-mail to AD of 16

March 1998). This latter specimen, kept under the name Rana limnocharis, is stated to have

been collected by Kuhl in Java (no information is available on locality and date of collection)

ition, and this specimen, described below, is fully

appropriate for neotype designation, although unfortunately it has no precise locality. Given

the information provided in the letters sent by Kuhl and Van Hasselt from Java, it seems likely

that this specimen was collected in the vicinity of Buitenzorg, now Bogor: “En nog zijn wij

geen 20 uren ver van Buitenzorg gekomen” (“And until now we did not go further than 20

hours from Buitenzorg”) (KuHL & VAN HASsELT, 18224: 103).

It is a young female in rather good c

Source : MNHN, Paris

Dunois & OHLER 29

STATUS OF A FEW NAMES CLOSELY RELATED TO THE NAME RANA LIMNOCHARIS

As mentioned above, for a long time the name Rana limnocharis was applied indiscrim-

inately to small frogs from a wide area of south and south-eastern Asia. As a number of

different names had been proposed in the beginning of zoology for frogs of this complex,

these names were long considered either to be synonyms of Rana limnocharis or, at best, to

apply to subspecies of the latter species. During the second half of our century, in this group

and many others (see DuBois & OHLER, 1998), the strong “lumper philosophy” of INGER

(1954, 1966) had a drastic influence on the taxonomy adopted by most authors: thus, INGER

(954: 267-274; 1966: 205-206) treated the taxon Rana vittigera Wiegmann, 1834 from the

Philippines as a subspecies of Rana limnocharis: as for the name Rana was! Annandale, 1917,

although this name clearly applied to frogs of this complex and was based on a type-specimen

from Borneo, he ignored it altogether in his book on Bornean frogs (INGER, 1966). DuBois

(1975b, 1984b, 1987, 1992) showed that this complex was in fact composed of a number of

distinct species, and gave a list of names available for these frogs. He suggested that several

names until then considered as synonyms or as subspecific names did apply to some of these

species, and that other species remained to be named. We will provide elsewhere (DuBois &

OHLER, in preparation) an updated review of the taxonomy of this group. Here we will only

extend the discussion to the names which may still pose nomenclatural problems in relation

with the existence of two distinct species of this complex in Java.

Following Dugois’s (1984b) paper, few names remained as genuine synonyms of Rana

limnocharis. However, ZHAO & ADLER (1993: 144) still regarded the name Rana gracilis

Wiegmann, 1834 as a synonym of the latter. This synonymy deserves discussion. Besides, we

discuss here the status of four additional names: Rana cancrivora Gravenhorst, 1829; Rana

vittigera Wiegmann, 1834; Rana multistriata Hallowell, 1861; and Rana was! Annandale,

1917.

THE STATUS OF THE NAME RANA CANCRIVORA GRAVENHORST, 1829

In contrast with most of other names concerning frogs of this complex, the name Rana

cancrivora has long been considered to apply to a species distinct from Rana limnocharis, e.g.

by BOULENGER (1920a: 23), VAN KAMPEN (1923: 170), SurrH (1927: 205; 1930: 96), BOURRET

(1942: 245), TayLoR (1962: 377), ZHaO & ADLER (1993: 140), or even INGER (1954: 260; 1966:

175), who provided comparisons between Rana cancrivora and Rana “limnocharis” vittigera.

The fact that R. cancrivora Was almost universally considered distinct from À. limnocharis

rests apparently only on the comparative diagnosis provided by GRAVENHORST (1829) for

these two species, where this author stated that the former was “larger” than the latter: since

then, the name Rana cancrivora has been consistently applied to a large species of this

complex, occurring in Java and neighbouring regions. However this action has never been

based on the examination of a type-specimen, and apparently until now no author has tried

to trace such a specimen.

An incidental result of the above work is the verification that all type-specimens of

species described as new by GRAVENHORST (1829), including Rana cancrivora, must now be

Source : MNHN, Paris

30 ALYTES 18 (1-2)

considered lost. GRAVENHORST (1829) himself considered the names Rana limnocharis and

Rana cancrivora as synonyms, so that unequivocal allocation of the name Rana cancrivora to

a biological species also requires designation of a neotype. None of the collections mentioned

above is known to harbour any specimen under the name Rana cancrivora collected near

Buitenzorg in Java by Kuhl and/or Van Hasselt. Consequently, another specimen is described

below as neotype. For this, we chose a specimen in good condition, collected recently in a

precise locality near Bogor and that belongs to the species traditionally recognised under this

name. This specimen is part of those that were used as outgroup in the molecular study of this

group reported by VerrH et al. (2000). Designation of this specimen as neotype of Rana

cancrivora Gravenhorst, 1829 will preclude any confusion in the allocation of this name to a

biological species.

THE STATUS OF THE NAME RANA VITTIGERA WIEGMANN, 1834

WIEGMANN (1834: 255-257, pl. 21 fig. 1) described Rana vittigera, provided a good

drawing of a specimen, and compared this new species to Rana limnocharis. Subsequently, the

species Rana vittigera was considered as a synonym of Rana tigerina Daudin, 1802 by some

authors (e.g. STEINEGER, 1907: 139) and of Rana cancrivora Gravenhorst, 1829 by others (e.g..

BOULENGER, 19204: 23), until TAYLOR (1920: 236) resurrected this name for a species of the

Philippines. He was followed by SMITH (1927: 205-207) and INGER (1954: 267), who however

reduced this taxon to the rank of a subspecies of Rana limnocharis. None of these authors

examined the type-specimens of this taxon. INGER (1954: 267) stated that its type-locality was

“Laguna de Bay, Luzon”, which was incorrect because, as noted by TAYLOR (1920: 236), the

species had been described on the basis of specimens from two different origins.

As a matter of fact, according to WIEGMANN (1834: 257), the original description was

based on several specimens, some from Laguna de Bay (Luzon, Philippines), and some from

the market of Macao (now Aomen, Guangdong, China). PETERS (1863: 77) provided more

information in this respect: he stated that the Berlin Museum had two specimens (ZMB 3269)

from Laguna de Bay and two others (ZMB 3270) from China. Dugois (1984b: 151-152)

commented on this and restricted the type-locality of the species to Laguna de Bay. His

comment was misunderstood by DUELLMAN (1993: 229), who wrote: “Lectotypes: ZMB 3269,

designated by Dubois, 1984, Alytes, 3: 152.”. In fact, DuBois (1984b: 152) had not designated

a lectotype, but had stated that such a designation should be made, after examination of the

specimens: “In order to stabilize definitely the use of the name virrigera as proposed by INGER

(1954), it would be necessary to designate formally one of the two specimens ZMB 3269 as

lectotype of Rana vittigera Wiegmann, 1835, what I cannot do for the time being, as I have not

yet been able to examine these specimens.” (translated from the French text in Dugois, 1984b:

152). Because of this misunderstanding, DUELLMAN (1993) “almost” designated a lectotype

for this species, but of course he did not, because, to be valid, a lectotype designation must

point to an individual, and ZMB 3269 consists of two specimens.

On 21 December 1995, thanks to the hospitality of Rainer Günther, we had the

opportunity to examine the 4 known syntypes of this species in the Berlin Museum. When we

got the bottles containing these specimens for examination, these bottles were still sealed with

resin and had to be cut open with a scalpel, thus emitting a very pleasant smell of old

Source : MNHN, Paris

Dugois & OHLER 31

aromatized alcohol: it is very likely that these specimens had never been examined since the

19‘ century, perhaps since PETERS’s (1863) work.

These four specimens are in good condition. The two specimens from Laguna de Bay,

ZMB 3269, are two adult females (SVL 68.6 mm and 57.2 mm). The two specimens from

Macao, ZMB 3270, are also two adult females (SVL 58.5 mm and 55.9 mm). Comparisons of

these four specimens with fig. 1 of pl. 1 of WiEGMANN (1834) shows that the latter was drawn

from the largest of the two specimens from Laguna de Bay. This specimen is therefore here

designated as lectotype of Rana vittigera, which is consistent with the use of this name

introduced by TAYLOR (1920) and adopted by all subsequent authors. This lectotype is

described in detail and figured below.

THE STATUS OF THE NAME RANA GRACILIS WIEGMANN, 1834

WIEGMANN (1834: 257-258) described Rana gracilis on the basis of a single adult male

specimen, collected in China near the “Cap Syng-more” (now Kap Shui Mun, Lantau Island,

Hong Kong, China). He considered this species as very close to Rana limnocharis. PETERS

(1863: 78) stated that this species was “completely identical” (“stimmt ganz überein”’) with

Rana limnocharis and Rana vittigera. Since then, all authors have considered the name Rana

gracilis Wiegmann, 1834 as a subjective synonym of Rana limnocharis, and this synonymy was

still considered valid by ZHAO & ADLER (1993: 144), who however did not include Rana

vittigera in this synonymy.

During our stay in Berlin mentioned above, we examined the holotype of this species,

ZMB 3255. We provide below a redescription and a photograph of this specimen. We consider

that, by several important characters, this specimen is distinct from both species of this group

known from Java: frogs from China have significantly shorter heads, forelegs and hindlimbs,

and their inner metatarsal tubercle is shorter relative to the length of first toe. We will provide

more information on this question elsewhere, but, for the purpose of this paper, it is enough

to say that this Chinese species is distinct from both Javanese species of this group, and should

be removed from the synonymy of Rana limnocharis. This statement is also supported by the

results of the electrophoretic comparison of specimens froms Java and Hong Kong (Topa et

al., 1998).

However, the name Rana gracilis Wiegmann, 1834 cannot be resurrected for this Chinese

species, because this name is preoccupied in the genus Rana (see e.g. DuBois, 1984b: 154),

being a junior primary homonym of the name Rana gracilis Gravenhorst, 1829, a Sri Lankan

species of the subgenus Sy/virana Dubois, 1992 of the genus Rana Linnaeus, 1758 (see

Dugois, 1992: 326). According to the Code, a junior primary homonym is permanently

invalid, so that the name Rana gracilis cannot be resurrected for the Chinese species, even if

the two species bearing this name are no longer considered congeneric. As no junior synonym

of this name is currently known (see e.g. ZHAO & ADLER, 1993: 144), it would seem that we are

in à situation where, to designate this Chinese species, a new replacement name (nomen

novum) should be coined for the name Rana gracilis Wiegmann, 1834. However, we propose

below another, more “parsimonious”, solution to this problem.

Source : MNHN, Paris

32 ALYTES 18 (1-2)

THE STATUS OF THE NAME RANA MULTISTRIATA HALLOWELL, 1861

In a long and famous paper, HALLOWELL (1861) described several amphibian species

from Japan and Hong Kong. Several of these nominal species have never been allocated to

biological species since then, and their types seem to be lost (see e.g. ZHAO & ADLER, 1993:

280). However, these names are nomenclaturally available and their status should be clarified,

which can be done through the designation of neotypes from the same localities (see e.g.

Dusois & OHLER, 1997a-b). To be sure, HALLOWELL’s (1861) descriptions are too vague to

allow unambiguous allocation of these names. In order not to threaten the stability of

nomenclature, we think allocation of these names should be done following the three

following principles: (1) the biological species to which the name is allocated should be known

to be present in the area whence HALLOWELL’s specimens came; (2) it should not have

characters incompatible with HALLOWELL’s (1861) description; (3) this species should either

be still unnamed or be known under a name published before 1861, so that HALLOWELL'S

name becomes its junior subjective synonym: in the latter case, HALLOWELL’s name would

remain available for further taxonomic work, for example if a frog species from Hong Kong,

currently considered conspecific with other populations, was later shown to be a different

species.

In the light of these ideas, we propose the following interpretations of the three species

names proposed by HALLOWELL (1861) for frogs of Hong Kong, and which ZHAO & ADLER

(1993: 280) kept unallocated to biological species: Rana trivittata, Rana nebulosa and Rana

multistriata.

(1) Concerning the name Rana trivittata, in the light of the original description

(HALLOWELL, 1861: 504-505), we consider that it could well apply to the species now known

as Rana macrodactyla (Günther, 1859), a member of the subgenus Hylarana Tschudi, 1838 of

the genus Rana (see DuBois, 1992: 328), which occurs in Hong Kong (Lai & NG, 1972;

KARSEN et al., 1986). Definitive stabilization of the status of Rana trivittata as a junior

subjective synonym of Rana macrodactyla will require the designation as neotype of R.

trivittata of a specimen of the latter species collected in Hong Kong.

(2) As for the name Rana nebulosa, examination of the original description (HALLOWELL,

1861: 505) leads us to think that it could fit the species currently known as Rana livida (Blyth,

1856), a species currently placed either in the subgenus Odorrana Fei, Ye & Huang, 1991 or in

the subgenus Eburana Dubois, 1992 of the genus Rana (see FEt et al., 1991: 147; DuBois, 1992:

328; Fer, 1999: 188), which also occurs in Hong Kong (Laï & NG, 1972; Ka: etal., 1986).

In this case also, stabilization of this name in this synonymy will require the designation of a

neotype from Hong Kong.

(3) Finally, HaLLOwELL's (1861: 504-505) original description of the species Rana

multistriata could well apply to a species of the Rana limnocharis group, which is also present

in Hong Kong. As we have seen above, the holotype of Rana gracilis Wiegmann, 1834,

collected in Hong Kong, belongs to a species distinct from Rana limnocharis, and for which no

scientific name is currently available. We propose to take advantage of this situation to apply

the name Rana multistriata to this unnamed Chinese frog species, through designation as

Source : MNHN, Paris

Dusois & OHLER 33

neotype of the latter of the holotype of Rana gracilis, described and figured below: this

solution of the nomenclatural problems posed by both these names is an example of

“nomenclatural parsimony”, a concept that will be discussed at more length elsewhere

(Dugoïs, in preparation).

THE STATUS OF THE NAME RANA WASL ANNANDALE, 1917

ANNANDALE (1917: 131-132) erected the species Rana wasl for specimens from various

regions (Sarawak, Myanmar, Assam & Nicobar Islands). He stated that the holotype, ZSI

17282, was from Kuching (Sarawak, Malaysia, in the island of Borneo). BOULENGER (1920a:

28) placed this name in the synonymy of Rana limnocharis, where it has remained until now

(e.g.: VAN KAMPEN, 1923: 167; BOURRET, 1942: 250; GORHAM, 1974: 146), except for authors

who failed to mention it (e.g.: LIU, 1950: 315; TAYLOR, 1962: 380; INGER, 1966: 205). On 14

August 1973, DuBois (1984b: 155) was able to examine and measure the holotype of Rana

wasl in the Calcutta Museum: it is an adult female (SVL 56 mm; TL 31 mm; HW 19 mm; HL

18.5 mm; IUE 3 mm; UEW 4.5 mm; IN 5 mm), which is quite accurately shown in fig. 5 and

Sa of pl. 5 of ANNANDALE (1917), here reproduced as fig. 1. We are unable to provide here a full

redescription of this holotype, as the current loan policy of the Zoological Survey of India of

Calcutta is to refuse to send specimens abroad (Indraneil Das, e-mail to AD of 29 October

1998).

GENERIC CLASSIFICATION

A few words must be said here about the generic classification of the frogs related to Rana

limnocharis. Although long maintained in the genus Rana Linnaeus, 1758 (the type-species of

which is the European Rana temporaria Linnaeus, 1758: see DuBois, 1992: 333), these species

have often been referred to a particular “group”, “complex”, “section” or “subgenus” of this

genus. Thus, ANNANDALE (1917: 131) placed them in a “ Rana limnocharis group”, which he

considered distinct from a * Rana tigrina group”. In contrast, BOULENGER (1918: 115) united

both groups in a “groupe de R. tigrina et limnocharis” of his subgenus Rana s. Str.; he later

considered the same group as a “section” “ Ranae tigrinae” of this genus (BOULENGER, 19204:

9). DECKERT (1938) placed these species, as well as others, in the genus Dicroglossus Günther,

1860, which was recognized as a valid genus by LAURENT (1950), and later by DuBois (1974),

but as a subgenus of Rana. DuBois (1975a: 1112) pointed out that, for the latter group, the

name Euphlyctis Fitzinger, 1843 had priority. Dugois (1981 240) recognized several

species groups in the latter subgenus and designated Rana limnocharis as type-species of

Fejervarya Bolkay, 1915, in order to provide à genus-group name for this group. DuBois

(1984b) proposed to use this latter name as a subgeneric name within Rana. DUBois (1987: 61)

transferred this subgenus to the genus Limnonectes Fitzinger, 1843. Finally, Fet et al. (1991:

126) were the first to raise the Rana limnocharis group to the rank of a distinct genus, for

which, however, they used the incorrect name Euphlyctis (which applies in fact to Rana

cyanophlyctis Schneider, 1799 and related species, ie. a quite distinct group indeed: see

Dusois, 1992). Dugois (2000), ISKANDAR (1998, 1999), Fer (1999) and MARMAYOU et al.

(2000) followed this suggestion, except for its nomenclatural part, since Fejervarya is the valid

name for this group.

Source : MNHN, Paris

34 ALYTES 18 (1-2)

Fig. 1. -Rana was! Annandale, 1917, holotype, ZSI 17282: head in dorsal and lateral view (reproduced

from fig. 5 and Sa of pl. 5 of ANNANDALE, 1917).

Several reasons lead us to adopt FEt et al.’s (1991) proposal. This decision is supported

both by the important phenetic differences that exist between Fejervarya and Limnonectes,

such as the shape of the tips of di of adults (OnLER & DuBois, 1999), their types of male

secondary characters (BOULENGER, 19204), a higher morphometrical distance between the

adults of these genera than between them and those of other genera such as Phrynoglossus

Peters, 1867 (OnLEr & Dugois, 1999), or the differences in the mouthparts of their tadpoles

(Fel et al., 1991). More significantly even, the preliminary cladistic analyses, based on DNA

sequencing, provided independently by VENCES (1999), MarMaYoU et al. (2000) and BOssUYT

Source : MNHN, Paris

Dugois & OHLER 35

& MILINKOVITCH (2000), suggest that Fejervarya is not the sister-group of Limnonectes, but is

more closely related to other genera such as Hoplobatrachus Peters, 1863 and Sphaerotheca

Günther, 1859.

For all these reasons, we refer here all the species of the former “ Rana limnocharis group”

to a distinct genus Fejervarya Bolkay, 1915. We take this opportunity to point out the presence

in all species of this genus of a unique common derived character which seems to have escaped

the attention of all authors until now. This character was observed by us in all examined

species of this genus, but not in any other of a vast array of ranids from various groups

examined in this respect by us and also by Julio Mario Hoyos (personal communication). In

species of the genus Fejervarya, the ventro-lateral edge of the musculus pectoralis pars

abdominalis is slightly attached to the skin from armpit to groin, whereas usually in ranids it

is attached to muscles which are dorsal relative to it (musculus rectus abdominis and musculus

obliquus externus). This results in the presence, in adults of both sexes of all species of

Fejervarya, of a dark ventro-lateral line from armpit to groin, which is usually very clearly

conspicuous in live specimens, whose belly in this genus is usually bright white or yellowish

and unspotted. This dark line being characteristic of the species of the genus Fejervarya, we

propose to call it the “Fejervaryan line”. We consider this character as an autapomorphy of

the genus Fejervarya, that provides an apognosis for this genus (see DuBois, 1997).

This genus is still in need of an overall revision. For the time being, on the basis of the

information already published by Dugois (1984b, 1987, 1992) and provided in the present

paper, we recognize the following species as valid: Fejervarya andamanensis (Stoliezka, 1870);

Fejervarya cancrivora (Gravenhorst, 1829); Fejervarya greenii (Boulenger, 1904); Fejervarya

keralensis (Dubois, 1981) [synonym: Rana verrucosa Günther, 1876]; Fejervarya kirtisinghei

(Manamendra-Arachchi & Gabadage, 1996); Fejervarya limnocharis (Gravenhorst, 1829)

[synonyms: Rana limnocharis Wiegmann, 1834 and Rana was! Annandale, 1917]; Fejervarya

multistriata (Hallowell, 1861) [synonym: Rana gracilis Wiegmann, 1834]: Fejervarya nepalen-

sis (Dubois, 1975); Fejervarya nilagirica (Jerdon, 1853); Fejervarya pierrei (Dubois, 1975);

Fejervarya rufescens (Jerdon, 1853); Fejervarya syhadrensis (Annandale, 1919): Fejervarya

teraiensis (Dubois, 1984): Fejervarya vittigera (Wiegmann, 1834). Besides, the following

names, which are still unsufficiently characterized in published works, will also have to be

considered in any global revisionary work of this genus: Fejervarya altilabris (Blyth, 1855);

Fejervarya assimilis (Blyth, 1852); Fejervarya brama (Lesson, 1834); Fejervarya brevipalmata

(Peters, 1871); Fejervarva frithi (Theobald, 1868); Fejervarya moodiei (Taylor, 1920); Fejerva-

rya murthü (Pillai, 1979); Fejervarya mysorensis (Rao, 1922); Fejervarya nicobariensis (Stolic-

zka, 1870); Fejervarya parambikulamana (Rao, 1937); Fejervarya pulla (Stoliezka, 1870);

Fejervarya raja (Smith, 1930); Fejervarya sauriceps (Rao, 1937); Fejervarya schlueteri (Werner,

1893); Fejervarya verruculosa (Roux, 1911).

Source : MNHN, Paris

36 ALYTES 18 (1-2)

DESCRIPTIONS OF TYPE-SPECIMENS

NEOTYPE, BY PRESENT DESIGNATION, OF RANA LIMNOCHARIS GRAVENHORST, 1829 AND OF

RANA LIMNOCHARIS WIEGMANN, 1834 (FIG. 2-3)

RMNH 4287, young female, collected by H. Kuhl in 1821 near Buitenzorg [now Bogor]

(06°35S, 106°47°E), West Java, Java, Indonesia.

(A) Size and general aspect. — (1) Specimen of medium size (SVL 44.4 mm), body rather

slender.

(B) Head. — (2) Head of medium size, wider (HW 16.0 mm) than long (HL 14.6 mm;

MN 13.6mm; MFE9.8 mm; MBE 6.2 mm), convex. (3) Snout oval, protruding, its length (SL

7.78 mm) longer than horizontal diameter of eye (EL 5.19 mm). (4) Canthus rostralis

rounded, loreal region concave, acute. (5) Interorbital space flat, smaller ([UE 2.20 mm) than

upper eyelid (UEW 3.89 mm) and internarial distance (IN 3.05 mm); distance between front

of eyes (IFE 6.3 mm) more than one half of distance between back of eyes (IBE 10.9 mm).

(6) Nostrils oval, with small lateral flap, closer to tip of snout (NS 2.46 mm) than to eye (EN

4.02 mm). (7) Pupil rounded. (8) Tympanum (TY D 2.92 mm) distinct, oval, horizontal, about

half of eye diameter: tympanum-eye distance (TYE 1.55 mm) about half its diameter. (9)

Pineal ocellus present, between anterior border of eyes. (10) Vomerine ridge present, bearing

few small teeth, between choanae, with an angle of 45° to body axis, closer to choanae than

from each other, longer than distance between them. (11) Tongue large, cordate, emarginate.

(12) Supratympanic fold distinct, from eye to shoulder. (13) Parotoid glands absent. (14)

Cephalic ridges absent. (15) Co-ossified skin absent.

9.8 mm), not enlarged. (17) Fingers long, thin (TFL 5.77 mm). (18) Relative length of fingers,

shortest to longest: II < IV < I < III. (19) Tips of fingers pointed. (20) Fingers without dermal

fringe; webbing absent. (21) Subarticular tubercles prominent, rounded, single, all present.

(22) Prepollex oval, prominent; two oval, flat palmar tubercles; supernumerary tubercles

absent.

(D) Hindlimbs. — (23) Shank almost four times longer (TL 23.6 mm) than wide (TW

6.6 mm), longer than thigh (FL 20.6 mm), but shorter than distance from base of internal

metatarsal tubercle to tip of toe IV (FOL 24.6 mm). (24) Toes long, thin; toe IV long (FTL

14.4 mm), more than one third of distance from base of tarsus to tip of toe IV (TFOL

36.1 mm). (25) Relative length of toes, shortest to longest: 1 < II < V = III < IV. (26) Tips of

toes pointed. (27) Webbing moderate: 1 1 —2 II 1 —2 II 1-27, IV 27/,-1 1/7, V(WTF

4.80 mm; WFF 4.54 mm; WI 3.69 mm; WII 3.50 mm; MTTF 12.2 mm; MTFF 12.2 mm:

TETE 11.8 mm; FFTF 12.8 mm). (28) Dermal fringe along toe V present, from tip of toe to

base of metatarsus, well developed. (29) Subarticular tubercles prominent, rounded, simple,

all present. (30) Inner metatarsal tubercle rather short, prominent: its length (IMT 2.14 mm)

more than 2.5 times in length of toe I (ITL 5.57 mm). (31) Inner tarsal ridge present on distal

third of tarsus. (32) Outer metatarsal tubercle absent; supernumerary tubercles absent: tarsal

tubercle absent.

Source : MNHN, Paris

Dugois & OHLER 37

Fig. 2. -Rana limnocharis Gravenhorst, 1829, neotype, and Rana limnocharis Wiegmann, 1834, neotype,

RMNH 4287, young female (SVL 44.4 mm): dorsal view.

Fig. 3. - Rana limnocharis Gravenhorst, 1829, neotype, and Rana limmocharis Wiegmann, 1834, neotype,

RMNH 4287, young female (SVL 44.4 mm): right lateral view of head.

Source : MNHN, Paris

38 ALYTES 18 (1-2)

() Skin. — (33) Dorsal and lateral parts of head and body: snout and between the eyes

smooth; side of head with few glandular warts; back and upper part of flanks with glandular

folds; lower part of flanks with glandular warts. (34) Latero-dorsal folds absent. (35) Dorsal

parts of limbs: forelimbs smooth; thigh and shank with glandular warts: tarsus smooth. (36)

Ventral parts of head, body and limbs: throat, chest and belly smooth: thigh with glandular

warts. (37) No macroglands.

(F) Coloration in alcohol. — (38) Dorsal and lateral parts of head and body: fawn with a

large dirty-white mid-dorsal band and darker brown spots; upper flank coffee brown with

darker spots, lower part light fawn; loreal and temporal region fawn with a brown band on

canthus rostralis and tympanic fold and brown spots on upper lip; tympanunm light fawn with

its dorsal half dark brown. (39) Dorsal parts of limbs: forelimbs, thigh, shank and foot fawn

with darker bands: posterior part of thigh brown with white marbling. (40) Ventral parts of

head, body and limbs: throat, chest, belly and thigh light fawn; margin of throat light fawn

white with large brown spots; Fejervaryan line present.

(G) Female sexual characters. — (41) Oviduct translucent, folded. (42) Ovaries not

observed.

NEOTYPE, BY PRESENT DESIGNATION, OF RANA CANCRIVORA GRAVENHORST, 1829 (FIG. 4-5)

FMNH 256688 (field number MV.40), adult male, collected by Michael Veith on 5

February 1993 at Cianjur (06°49'S, 107°08°E), West Java, Java (Indonesia).

(A) Size and general aspect. — (1) Specimen of rather large size (SVL 68.2 mm), body

rather slender.

(B) Head. — (2) Head of medium size, narrower (HW 26.0 mm) than long (HL 29.7 mm:

MN 27.2 mm; MFE 21.3 mm; MBE 15.6 mm), slightly convex. (3) Snout oval, protruding, its

length (SL 10.9 mm) longer than horizontal diameter of eye (EL 7.7 mm). (4) Canthus

rostralis rounded, loreal region concave, obtuse. (5) Interorbital space flat, smaller (IUE

3.3 mm) than upper eyelid (UEW 5.5 mm) and internarial distance (IN 4.4 mm); distance

between front of eyes (IFE 9.9 mm) more than one half of distance between back of eyes (IBE

15.8 mm). (6) Nostrils oval, with small lateral flap, closer to tip of snout (NS 5.4 mm) than to

eye (EN 6.9 mm). (7) Pupil rounded. (8) Tympanum (TYD 4.8 mm) distinct, oval, horizontal,

about two thirds of eye diameter; tympanum-eye distance (TYE 2.7 mm) about half its

diameter. (9) Pineal ocellus present, between anterior quarter of eyes. (10) Vomerine ridge

present, bearing a few small teeth, between choanae, with an angle of 45° to body axis, closer

to choanae than from each other, longer than distance between them. (11) Tongue large,

cordate, emarginate. (12) Supratympanic fold distinct, from eye to shoulder. (13) Parotoid

glands absent. (14) Cephalic ridges absent. (15) Co-ossified skin absent.

(HAL 15.3 mm), not enlarged. (17) Fingers rather long, thin (TFL 7.9 mm). (18) Relative

length of fingers, shortest to longest: 11 < IV < 1 < III. (19) Tips of fingers pointed. (20)

Fingers Il and III with dermal fringe, webbing absent. (21) Subarticular tubercles prominent,

rounded, single, all present. (22) Prepollex oval, indistinct: palmar tubercles indistinct:

supernumerary tubercles absent.

Source : MNHN, Paris

Dugois & OHLER

Fig. 4.-Rana cancrivora Gravenhorst, 1829, neotype, FMNH 256688, adult male (SVL 68.2 mm): dorsal

view.

5. -Rana cancrivora Gravenhorst, 1829, neotype, FMNH 256688, adult male (SVL 68.2 mm): right

ateral view of head.

Source : MNHN, Paris

40 ALYTES 18 (1-2)

(D) Hindlimbs. — (23) Shank about three times longer (TL 35.7 mm) than wide (TW

12.9 mm), longer than thigh (FL 33.6 mm), but shorter than distance from base of internal

metatarsal tubercle to tip of toe IV (FOL 37.8 mm). (24) Toes long, thin; toe IV long (FTL

21.6 mm), more than one third of distance from base of tarsus to tip of toe IV (TFOL

52.9 mm). (25) Relative length of toes, shortest to longest: I < II < V < III < IV. (26) Tips of

toes pointed. (27) Webbing moderate: I 1-1 %111-2111-21V2-1 V(WTF7.0mm; WFF

7.3 mm; WI 6.7 mm; WII 5.3 mm). (28) Dermal fringe along toe V present, from tip of toe to

base of metatarsus, well developed. (29) Subarticular tubercles prominent, oval, simple, all

present. (30) Inner metatarsal tubercle oval, prominent; its length (IMT 3.7 mm) less than

2.5 times length of toe I (ITL 8.8 mm). (31) Inner tarsal ridge present on distal ?/, of tarsus.

(32) Outer metatarsal tubercle absent; supernumerary tubercles absent; tarsal tubercle absent.

(Ë) Skin. — (33) Dorsal and lateral parts of head and body: snout and between the eyes

shagreened; side of head with small glandular warts: back and upper part of flanks with

glandular folds; lower part of flanks with glandular warts. (34) Fine, narrow, interrupted

latero-dorsal folds on 7/; of back. (35) Dorsal parts of limbs: forelimbs, thigh, shank and

tarsus with glandular warts and folds. (36) Ventral parts of head, body and limbs: throat, chest

and belly smooth. (37) No macroglands.

(F) Coloration in alcohol. — (38) Dorsal and lateral parts of head and body: brown with

indistinct darker brown spots around the folds; canthus rostralis and tympanic fold of same

brown color; tympanum brown with inferior half clearer, lighter than head; three wide bands

from eye to upper lip; a wide light brown mid-dorsal band continuous from tip of snout to

vent. (39) Dorsal parts of limbs: forelimbs, thigh, shank and foot brown with darker bands;

posterior part of thigh dark brown with white marbling. (40) Ventral parts of head, body and

limbs: throat light brown with dark brown vocal sacs on both sides; belly and underside of

shank white with indistinet light brown spots; margin of throat white with large brown spots:

Fejervaryan line not visible (specimen dissected).

(G) Male sexual characters. — (41) Unique pad of numerous small grey brown nuptial

spines on prepollex and finger I. (42) Vocal sacs present.

LECTOTYPE, BY PRESENT DESIGNATION, OF RANA VITTIGERA WIEGMANN, 1834 (FIG. 6-8)

Largest of the two specimens under number ZMB 3269, adult female, collected by

FE. JF Meyen in Laguna de Bay (14°10°N, 121°20'E), Luzon, Philippines.

(A) Size and general aspect. — (1) Specimen rather large size (SVL 68.6 mm), body rather

slender.

(B) Head. — (2) Head of medium size, narrower (HW 21.6 mm) than long (HL 24.6 mm:

MN 21.1 mm; MFE 16.0 mm; MBE 9.8 mm), convex. (3) Snout oval, protruding, its length

(SL 10.1 mm) longer than horizontal diameter of eye (EL 7.2 mm). (4) Canthus rostralis

rounded, loreal region concave, obtuse. (5) Interorbital space flat, smaller (IUE 3.70 mm) than

upper eyelid (UEW 5.23 mm) and internarial distance (IN 3.63 mm); distance between front

of eyes (IFE 9.5 mm) two thirds of distance between back of eyes (IBE 13.8 mm). (6) Nostrils

oval, closer to tip of snout (NS 4.40 mm) than to eye (EN 5.93 mm). (7) Pupil indistinct.

(8) Tympanum (TYD 4.61 mm) distinct, oval, horizontal, about half of eye diameter:

Source : MNHN, Paris

Dugois & OHLER 41

74 /

Lana Here à #

Fig. 6. -Rana vittigera Wiegmann, 1834, lectotype, largest of the two specimens under number ZMB

3269: dorsal view (reproduced from fig. 1 of pl. 21 of WiFGMaNN, 1834)

tympanum-eye distance (TYE 2.44 mm) about half its diameter. (9) Pineal ocellus present,

between anterior border of eyes. (10) Vomerine ridge present, bearing few small teeth,

between choanae, with an angle of 45° to body axis, closer to choanae as from each other,

longer than distance between them. (11) Tongue not observed. (12) Supratympanic fold

distinct, from eye to shoulder. (13) Parotoid glands absent. (14) Cephalic ridges absent. (15)

Co-ossified skin absent.

12.3 mm), not enlarged. (17) Fingers rather long, thin (TFL 7.0 mm). (18) Relative length of

fingers, shortest to longest: IV < II < 1 < TEL. (19) Tips of fingers pointed. (20) Fingers without

dermal fringe; webbing absent. (21) Subarticular tubercles prominent, rounded, single, all

present. (22) Prepollex oval, indistinct; two oval, flat palmar tubercles: supernumerary

tubercles absent.

(D) Hindlimbs. — (23) Shank three times longer (TL 31.4 mm) than wide (TW 11.4 mm),

thigh (FL not measured, femur broken) about distance from base of internal metatarsal

tubercle to tip of toe IV (FOL 31.6 mm). (24) Toes rather short, thin; toe IV long (FTL

11.9 mm), less than one third of distance from base of tarsus to tip of toe IV (TFOL 46.5 mm).

(25) Relative length of toes, shortest to longest: 1 < IT < V = III < IV. (26) Tips of toes pointed.

(27) Webbing extensive: 10-—1110-1/2110-1/2IV1 2-0 V(WTEF 8.3 mm; WFF 6.6 mm:

Source : MNHN, Paris

4 ALYTES 18 (1-2)

7. -Rana vittigera Wiegmann, 1834, lectotype, largest of the two specimens under number ZMB

269, adult female (SVL 68.6 mm): dorsal view.

t of the two specimens under number ZMB

3269, adult female (SVL 68.6 mm): right lateral view of head

Fig. 8. -Rana vittigera Wiegmann, 1834, lectotype, larg

Source : MNHN, Paris

Dugois & OHLER 43

WI 7.2 mm; WII 4.8 mm; MTTF 16.6 mm; MTFF 17.4 mm; TFTF 12.9 mm; FFTF

13.8 mm). (28) Dermal fringe along toe V present, from tip of toe to base of metatarsus,

slightly developed. (29) Subarticular tubercles prominent, oval, simple, all present. (30) Inner

metatarsal tubercle short, very prominent: its length (IMT 2.23 mm) more than 3.5 times in

length of toe I (ITL 8.16 mm). (31) Inner tarsal ridge absent. (32) Outer metatarsal tubercle

absent; supernumerary tubercles absent; tarsal tubercle absent.

(£) Skin. — (33) Dorsal and lateral parts of head and body: snout and between the eyes

smooth; side of head with few glandular warts; back and upper part of flanks with short and

long glandular folds (the longest half of length of back); lower part of flanks with faded

glandular warts. (34) Latero-dorsal folds absent. (35) Dorsal parts of limbs: forelimbs

smooth; thigh with glandular warts and horny spinules: shank and tarsus smooth. (36)

Ventral parts of head, body and limbs: throat, chest and belly smooth; thigh with glandular

warts. (37) No macroglands.

(F) Coloration in alcohol. - (38) Dorsal and lateral parts of head and body: brown with

large, dense darker brown, rounded, confluent spots; light mid-dorsal line, slightly broadened

to the right in the mid of the back; shoulder pads continued by a clear band on the flanks:

three dark spots on upper lip; canthus rostralis and tympanic zone dark brown. (39) Dorsal

parts of limbs: forelimbs, thigh, shank and foot brown with outlines of darker bands:

posterior part of thigh dark brown with white marbling. (40) Ventral parts of head, body and

limbs: throat, chest, belly and thigh dirty white; margin of throat with some brown spots:

Fejervaryan line present.

(G) Female sexual characters. — (41) Oviduct large, folded. (42) Ovaries with small brown

and whitish eggs.

HOLOTYPE, BY MONOTYPY, OF RANA GRACILIS WIEGMANN, 1834 (NEC GRAVENHORST, 1829)

AND NEOTYPE, BY PRESENT DESIGNATION, OF RANA MULTISTRIATA HALLOWELL, 1861

(FIG. 9-10)

ZMB 3255, adult male, collected by F. J. F. Meyen near “Cap Syng-more”, now Kap Shui

Mun (22°2/°N, 114°03'E), Lantau Island, Hong Kong, China.

(A) Size and general aspect. — (1) Specimen of rather small size (SVL 33.0 mm), body

rather slender.

(B) Head. — (2) Head of medium size, longer (HL 12.6 mm) than wide (HW 10.0 mm:

MN 10.8 mm; MFE 8.2 mm; MBE 4.4 mm), convex. (3) Snout oval, slightly protruding, its

length (SL 5.44 mm) longer than horizontal diameter of eye (EL 4.61 mm). (4) Canthus

rostralis rounded, loreal region concave, obtuse. (5) Interorbital space flat, smaller (IUE

1.73 mm) than upper eyelid (UEW 2.74 mm) and internarial distance (IN 2.33 mm); distance

between front of eyes (IFE 5.5 mm) more than two thirds of distance between back of eyes

(BE 7.6 mm). (6) Nostrils oval, closer to tip of snout (NS 2.00 mm) than to eye (EN 2.80 mm).

(7) Pupil rounded. (8) Tympanum (TY D 2.04 mm) distinct, oval, horizontal, about half of eye

diameter; tympanum-eye distance (TYE 0.67 mm) about one third its diameter. (9) Pineal

ocellus absent. (10) Vomerine ridge present, bearing few small teeth, between choanae, with

an angle of 45° to body axis, less close to choanae than from each other, longer than distance

Source : MNHN, Paris

44 ALYTES 18 (1-2)

2 . À ‘

Fig. 9. - Rana gracilis Wieemann, 1834, holotype, and Rana multistriata Hallowell, 1861, neotype, ZMB

3255, adult male (SVL 33.0 mm): dorsal view.

Fig. 10. Rana gracilis Wiegmann, 1834, holotype, and Rana multistriata Hallowell, 1861, neotype, ZMB

3255. adult male (SVL 33.0 mm); right lateral view of head.

Source : MNHN, Paris

Dugois & OHLER 45

between them. (11) Tongue not observed. (12) Supratympanic fold indistinct, from eye to

shoulder. (13) Parotoid glands absent. (14) Cephalic ridges absent. (15) Co-ossified skin

absent.

7.5 mm), not enlarged. (17) Fingers long, thin (TFL 4.33 mm). (18) Relative length of fingers,

shortest to longest: IV < II <1< III. (19) Tips of fingers rounded. (20) Fingers without dermal

fringe; webbing absent. (21) Subarticular tubercles prominent, rounded, single, all present.

(22) Prepollex oval, prominent; one round, distinct internal palmar tubercle beside a very

small external palmar tubercle; supernumerary tubercles absent.

(D) Hindlimbs. — (23) Shank four times longer (TL 15.8 mm) than wide (TW 3.5 mm),

longer than thigh (FL 14.1 mm), but shorter than distance from base of internal metatarsal

tubercle to tip of toe IV (FOL 17.5 mm). (24) Toes long, thin; toe IV long (FTL 10.6 mm),

more than one third of distance from base of tarsus to tip of toe IV (TFOL 26.0 mm). (25)

Relative length of toes, shortest to longest: I < IT < V = III < IV. (26) Tips of toes rounded. (27)

Webbing moderate: 1 1-2111-21I11 %4-22/,1V27/,-1 V(WTE 3.35 mm; WFF 3.21 mm;

WI 3.10 mm; WII 2.37 mm; MTTF 8.65 mm; MTFF 8.86 mm; TFTF 7.74 mm; FFTF

8.51 mm). (28) Dermal fringe along toe V present, from tip of toe to base of metatarsus,

scarcely developed. (29) Subarticular tubercles prominent, oval, simple, all present. (30) Inner

metatarsal tubercle very short, very prominent: its length (IMT 1.16 mm) almost 4 times in

length of toe I (ITL 4.44 mm). (31) Inner tarsal ridge absent. (32) Outer metatarsal tubercle

absent; supernumerary tubercles absent; tarsal tubercle absent.

(E) Skin. — (33) Dorsal and lateral parts of head and body: snout and between the eyes

smooth; side of head with few glandular warts; back and upper part of flanks with rather

short glandular folds (the longest about size of eye-length); lower part of flanks almost

smooth. (34) Latero-dorsal folds absent. (35) Dorsal parts of limbs: forelimbs and thigh

smooth; shank and tarsus with horny spinules. (36) Ventral parts of head, body and limbs:

throat, chest and belly smooth: thigh with glandular warts. (37) No macroglands.

(F) Coloration in alcohol. — (38) Dorsal and lateral parts of head and body: colours

faded; brown with a large clearer mid-dorsal band and darker brown spots; shoulder spots

indistinct; four brown spots on each side of upper lip. (39) Dorsal parts of limbs: forelimbs,

thigh, shank and foot with dark bands: posterior part of thigh brown with white net forming

a light longitudinal line on the back side of each thigh. (40) Ventral parts of head, body and

limbs: chest, belly and thigh dirty white; greyish spots on side of throat continuous in the

middle; margin of throat beige white with large brown spots; Fejervaryan line present.

(G) Male sexual characters. - (41) Nuptial spines present, one single patch on prepollex

and finger I: numerous, very small, cream-coloured spines. (42) Vocal sacs present, greyish,

folded skin on the two sides of the throat: slit-like openings in posterior part of mouth floor.

(43) Fine horny spinules on the anterior border of the throat.

RÉSUMÉ

Une analyse détaillée du statut nomenclatural de l'espèce nominale Rana limnocharis

montre qu’elle a été rendue disponible pour la première fois par GRAVENHORST (1829), puis

Source : MNHN, Paris

46 ALYTES 18 (1-2)

une deuxième fois indépendamment par WiEGMANN (1834). Les conséquences de ces faits en

ce qui concerne les types porte-noms de ces deux espèces nominales sont discutées et des

néotypes sont désignés pour celles-ci. Le statut des espèces nominales suivantes, voisines de

Rana limnocharis, est aussi discuté et leurs spécimens-types sont décrits: Rana cancrivora

Gravenhorst, 1829; Rana vittigera Wiegmann, 1834; Rana gracilis Wiegmann, 1834; Rana

multistriata Hallowell, 1861; Rana wasl Annandale, 1917. Finalement, sur la base de plusieurs

informations récentes, il est suggéré que le groupe de grenouilles habituellement désigné

comme “groupe de Rana limnocharis” ou “sous-genre Fejervarya” devrait être reconnu

comme un genre distinct, Fejervarya Bolkay, 1915.

ACKNOWLEDGEMENTS

We would like to thank very specially the curators who allowed us to study their collections: Rainer

Günther (Zoologisches Museum Berlin), Marinus Hoogmoed (Nationaal Natuurhistorisch Museum,

Leiden) and Alan Resetar (Field Museum of Natural History, Chicago). Nick Arnold (Natural History

Museum, London), Indraneil Das (Institute of Biodiversity and Environmental Conservation, Kota

Samarahan), Heinz Grillitsch (Naturhistorisches Museum Wien), Günther Kôhler (Senkenbergstiftung

Frankfurt am Main) and Andrej Wiktor (Muzeum Przyrodnicze, Uniwwersytet Wroclawski) provided

information concerning collection data helpful to this work. Roger Bour (Muséum national d'Histoire

naturelle, Paris) gave us his support at various stages of the work. The staff of the Bibliothèque Centrale

in our Museum has our gratitude for helping to access the old literature and giving us permission to

publish plates copied from these antique books. This is publication N° 00-29 of PPF “Faune et flore du

sud-est asiatique” (publication N° 00-28 see DEUVE, 2000).

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Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 51-61. 51

Dentigerous bones and dentition

in the paedomorphic plethodontid

salamander Eurvycea neotenes:

Günter CLEMEN * & Hartmut GREVEN **

* Institut für Spezielle Zoologie und Vergleichende Embryologie der Universität Münster,

Hüferstr. 1, 48149 Münster, Germany

** Institut für Zoomorphologie und Zellbiologie der Universität Düsseldorf,

Universitätsstr. 1, 40225 Düsseldorf, Germany?

Tooth-bearing bones and dentition of males and females of the paedo-

morphic plethodontid salamander Eurycea neotenes are described using

alizarin transparencies and SEM micrographs. Dentition is strictly mono-

stichous at the upper jaw (premaxillae only), the lower jaw (dentaries,

splenials) and the palate (vomers, palatopterygoids). Teeth in both sexes are

monocuspid, conical, and pedicellate, but the weak zone between the

dentine crown and the pedicel is not very distinct and does not progress

beyond a late larval stage. In addition, we found some sex-linked characters.

Compared with females, males possess stronger premaxillae with separated

processus faciales, fissured inner margins of the dentated parts of the

palatoptervgoids that overlap the parasphenoïd, and larger splenials bear-

ing more teeth. In addition, males appear to have longer teeth, especially on

the anterior portion of premaxillae.

INTRODUCTION

In Urodela, tooth bearing bones, dentition and the form of teeth undergo a more or less

specific developmental sequence and remodeling during metamorphosis (WiLDER, 1925;

STADTMÜLLER, 1924; GREVEN, 1988; CLEMEN & GREVEN, 1977, 1994). Paedomorphic species

often possess a mosaic of larval and metamorphic traits which reflect their “degree” of

paedomorphosis; such traits can be found also in the tooth systems (e.g., GREVEN & CLEMEN,

1980; CLEMEN & GREVEN, 1988; REILLY 1994).

In plethodontids, paedomorphosis has pl: ant role in influencing evolution-

ary patterns and larval as well as paedomorphic features of the tooth systems have been

described (e.g., LARSEN, 1963: WAKE, 1966; MUTZ & CLEMEN, 1992).

1. Dedicated to Prof. Dr. H. Hartwig, Cologne, on the occasion of his 90th birthday

2. Address for correspondence.

Source : MNHN, Paris

52 ALYTES 18 (1-2)

In the present paper we examine the lower and upper jaw as well as the palate of the

plethodontid Eurycea neotenes. This species was considered to be probably paedogenetic

(progenetic) and not neotenic (BRUCE, 1976; SwEET, 1977; for terminology and the use of the

term paedomorphosis to specify any retention of juvenile features by adult descendants, see

Goup, 1977; WakE, 1980; RrILLY, 1994). Our observations broaden the hitherto available

studies on this topic in the genus Eurycea (see STEWART, 1958; LARSEN, 1963; WAKE, 1966;

SWEET, 1977; MUTZ & CLEMEN, 1992) and draw the attention to some sexual dimorphic

characters.

MATERIALS AND METHODS

A total of 5 adults of Eurycea neotenes (3 females, 2 males) of unknown origin were

examined. Specimens that died in captivity were preserved in 70 % ethanol. Total length of

males was approximately 68 mm, snout-vent length approximately 33 mm with only a

negligible range. The respective measurements in females were 60 mm and 34 mm. Males had

three lobes per testis and in females vitellogenesis was in progress or finished. According to

BRUCE (1976), males therefore were in the third reproductive year or fourth year of life and

females were maturing or mature.

Specimens were postfixed in Lillie buffered formalin for 3 days. Cleared specimens were

stained only for bone employing the method of Park & K1M (1984).

After drawing, the skeletal structures of the head of the specimens were transferred into

an enzyme solution of 30 ml saturated aqueous sodium borate, 70 ml distilled water and 1 g

pancreatin for several days to remove the soft tissue. Then the delicate dentigerous bones were

extracted, dehydrated in ethanol, critical-point dried and mounted on metal plates. They were

sputter-coated with gold and viewed in a SEM (Hitachi S-530).

The length of the head of two males and two females was measured from the midst of the

premaxillary arcade to the posterior end of the parasphenoïd and from the premaxillae to the

condylus. The width of the parasphenoid was determined at the level of the quadrate. Teeth

were measured directly on the SEM micrographs.

RESULTS

Length of the male’s head was 6 mm from the premaxillae to the posterior end of

the parasphenoïid and 6.5 mm to the condylus. The largest width of the parasphenoid was

2.2 mm. In the females the head measured 5 mm and 5.5 mm in length, and the parasphenoid

2.0 mm in width.

The upper jaw is composed only of the unpaired (fused) premaxilla. Maxillae are entirely

lacking. The tooth bearing partes dentales of the premaxillae form only a short arcade. Two

long processus faciales (p. praemaxillares according to CLEMEN & GREVEN, 1994) arise

Source : MNHN, Paris

CLEMEN & GREVEN

1mm

Fig. 1.- Ventral view of the anterior part of the skull of an adult female of Eurycea neotenes showing the

dentigerous and non dentigerous dermal bones of the upper jaw and the palate (black) and the

ossified quadrate (hatched). ch, choana: pg. palatopterygoid: pm, premaxillae with anteriorly

separated processus faciales (p. praemaxillares) (asterisk); ps, parasphenoid: v, vomer with anterior

process (white point). The posterior portion of the parasphenoid and other cartilaginous and

ossified elements of the skull are omitted

Fig. 2. - Ventral view of the anterior part of the skull of an adult male of Eurycea neotenes. For further

explanations and abbreviations, see fig. 1. Compared with the female (fig. 1). the pars palatina of

premaxillae (pm) is missing, the processus faciales (asterisks) are separated along their entire length

the vomers possess larger anterior processes (white point) and the fissured inner margins of the

dentated parts of the palatopterygoid (pg) overlap the broad parasphenoïd (ps)

Source : MNHN, Paris

54 ALYTES 18 (1-2)

1mm

Fig. 3. - Dorsal view of the dentigerous and non dentigerous dermal bones of the lower jaw (black) and

parts of the mostly cartilaginous hyobranchial apparatus (hatched) of an adult female of Eurycea

neotenes. d, dentary: par, prearticular: s, splenial: t, tongue

Fig. 4. - Dorsal view of the lower jaw (black) and parts of the hyobranchial apparatus (hatched) of an

adult male of Eurycea neotenes. For further explanations and abbreviations, see fig. 2. Compared

with the female (fig. 3), the splenials (s) are larger and the anterior portions of the dentaries (d) are

smaller.

Source : MNHN, Paris

CLEMEN & GREVEN 55

immediately below the tooth bearing portion (fig. 1-2). Two third of these processes are fused

posteriorly in females; in males they are separated for their entire length (fig. 5-6). The

toothless bony ledge along the lingual side (pars palatina) of the premaxillae is relatively large

in females, but absent in males (fig. 1-2).

The ventral components of the skull include fully ossified paired vomers anteriorly and

a median large parasphenoid posteriorly (fig. 1-2). The anteriorly directed processes of the

vomers are small in females, but broad in males (fig. 10). In the former they do not attach to

the premaxillae, in the latter they do and cover a larger portion of the processus faciales.

The posterolateral margin of each vomer forms approximately two thirds of the bony

margin of the inner choana. The inner toothless partes palatinae of the vomer overlap the

anterior part of the parasphenoiïd (fig. 7, 10). Vomers are separated widely from each other

posteriorly (fig. 1-2). The ossified paired palatopterygoids begin at the posterior margin of the

choanae and extend backwards to, but do not articulate with, the inner surface of the ossified

quadrate. In one specimen, however, an adhesion to the quadrate was observed on the left side.

The inner margin of the palatinal portion of the palatopterygoid is highly fissured in both

sexes (see fig. 7). This is obviously more pronounced in males, where this region overlaps the

broad parasphenoid. In two females the fissured region was not completely ossified on one

side, leaving separated bony patches (not pictured).

The lower jaw is composed of the dentaries that both form a broadly rounded arch. The

lingual side of the Meckel's cartilage is invested by the prearticular (fig. 3-4). In females,

dentaries are broader at their anterior end; the prearticulars extend more anteriorly than in

males. A small splenial (coronoid) occurs on the inner margin of each dentary overlying the

prearticular; it is very minute in females (fig. 8) and longer in males (fig. 9).

Premaxillae, dentaries, splenials, vomers and the palatinal portions of the palatopte-

rygoid bear strictly monostichous teeth (fig. 2-3, 7, 9-10). Teeth vary considerably in height.

Tooth height depends on the site of attachment and on sex. Teeth of males are longer than

those of females. These differences appear to be most obvious in the most anterior pre-

maxillary teeth. In addition, males have more tooth loci at the vomers, the palatinal portions

of the palatopterygoids and, in particular, at the splenials (tab. 1).The small number of

animals examined, however, does not allow further interpretation of the measurements and

counts of functional teeth (tab. 1).

Teeth are separated from each other and are slightly recurved towards the esophagus (fig.

6,11). They are similar in form, but decrease gradually in height and size posteriorly. Teeth are

ankylosed at the jaws in a distinct pleural condition (fig. 11) and at the labial face of the

vomers in a slight pleural condition (fig. 10). The majority of teeth of the labial face of the

palatinal portion of the palatopterygoid is attached horizontally (females) or more or less

pleurally (males). Only a few (2-3) posterior teeth in males are attached horizontally (fig. 7,

right side), whereas in females some anterior teeth are attached pleurally due to the different

decrease of height of the pars dentalis.

Each tooth is composed of a monocuspid crown and a pedicel (fig. 5-12). Both elements

are connected by collagenous fibres (fig. 13). À prominent dividing zone, however, as in most

transformed urodelan teeth consisting of à broad lingual and a small labial deepening is

Source : MNHN, Paris

56 ALYTES 18 (1-2)

missing (fig. 11-12). Some teeth show resorption pits mainly in the pedicel and signs of wear

apically. AIl teeth have at their lingual base one large opening to the pulp cavity and additional

small holes around the base (fig. 7-8, 11).

DISCUSSION

In Eurycea neotenes, the developmental state of dentigerous bones and teeth does not

progress beyond the larval condition. Similar traits in the tooth systems were found also in

other paedomorphic taxa (for review, see GREVEN, 1988; see also CLEMEN & GREVEN, 1977,

1988; GREVEN & CLEMEN, 1979).

Maxillae, which seem to appear relatively late during ontogeny in plethodontids (WiL-

DER, 1925; WaKkE, 1966), and processes forming the posterior vomerine tooth patches that are

typical for advanced metamorphosed plethodontids (STEWART, 1958; WakE, 1966; MUTZz &

CLEMEN, 1992; see fig. 12 in CLEMEN & GREVEN, 1994), are completely missing. Both

structures are, however, present in transformed Eurycea neotenes (see below).

We found no signs of disintegration of the palatopterygoids. This disintegration is

considered as one of the key factors indicating the onset of metamorphosis by some authors

(ReiLLY, 1986, 1987; ReiLLy & ALTIG, 1996). Obviously the disintegration of the splenials

runs parallel to that of the palatopterygoid, but dental laminae of the palatinal portion of the

palatopterygoid as well as of the splenials degenerate far earlier, indicating likewise the onset

of metamorphosis (GREVEN & CLEMEN, 1985; MUTZ & CLEMEN, 1992).

The larval condition holds also for the dentition. In our specimens, dentary, pre-

maxillary, vomerine, palatinal and splenial teeth are strictly monostichous. Generally, in early

urodelan larvae, at least the palatinal portion of the palatopterygoid and the splenials bear

more than one line of teeth, thus pos: g tooth patches (“Zahnfelder”). In transforming

Eurycea species, reduction of the dentition starts with the formation of a monostichous

Fig. $.- Fused monostichously dentated premaxillae of a female of Eurycea neotenes with partially fused

partes faciales.

Fig. 6. - Fused monostichously dentated premaxillae of à male of Eurycea neotenes. The partes faciales

are separated for their entire length.

Fig. 7. Vomer (asterisk) and palatinal portion of the palatopterygoid (point) of a male of Eurycea

neotenes. Note the only slight pleural attachment of teeth and the fissured palatinal portion

(arrowhead).

Fig. 8. - The minute splenial of the female of Eurycea neotenes bears only one tooth and overlies the

prearticular.

Fig. 9. - The splenial of the male of Eurycea neotenes is larger having up to eight tooth loci.

Fig. 10. — Vomer of the female of Eurycea neotenes With anterior processes (asterisk) and the labio-

marginal tooth-line.

Fig. 11. Distinet lingual dividing zone of vomerine teeth, male of Eurycea neotenes. Note the openings

to the pulp.

Fig. 12. - Dividing zone of a premaxillary tooth, labial side, male of Eurycea neotenes.

Fig. 13. - Collagenous fibers of the labial dividing Zone of a dentary tooth, female of Eurycea neotenes.

Source : MNHN, Paris

58 ALYTES 18 (1-2)

Table 1. — Height of teeth and number of tooth loci per side in two males and two females of

Eurycea neotenes as measured on SEM micrographs. Labial height is the distance from

the margin of the bone to the tip of the tooth crown, lingual height the distance from

the base of the pedicel to the tip of the crown.

Dentigerous Height of teeth in um Number of tooth loci (per side)

bones Male Female Male Female

Premaxillae

Ant É -25

Labial lerior 325-336 234-254

Posterior 212-245 205-217

1-14 14-16

à Anterior 407-415 325-336

Lingual

Posterior 299-340 387-230

Vomers

56- z

Labiar LAMterior 256-287 227-233

Posterior 228-238 183-190

12-14 10-12

Anterior 349-354 247-262

Lingual

Posterior 310-315 206-213

Palatopterygoids

Labia LAnterior 120-125

Posterior 87-94

9-11 6-8

Anterior 222-230 175-182

Lingual

Posterior 147-166 138-141

Dentaries

Labiar LAnterior 248-252 195-203

Posterior 234-241 196-205

_ 24-26 23-25

Anterior 309-328 308-315

Lingual

Posterior 245-257 219-224

Splenials 165-205 126-130 7-8 12

pattern and ends with the complete disintegration of these bones (WiLDER, 1925; MuTz &

Cu 1992). Therefore monostichy on the palatinal portions of the palatopterygoids and

the splenials is regarded as a late larval state. In contrast to other larval urodeles (see CLEMEN

& GREVEN, 1977, 1994; GREVEN & CLEMEN, 1985; MUTZ & CLEMEN, 1992; AMEND & GREVEN,

1996), vomerine teeth even of early plethodontid larvae (and paedomorphic plethodontids)

hitherto investigated are strictly monostichous (WAKE, 1966; MUTz & CLEMEN, 1992). Thus,

development of the vomerine dentition differs considerably from the general urodelan

scheme. Contrary to transformed urodeles, teeth of the vomer and the palatine are ankylosed

at the outer (labial) face of the bones, emphasizing again the late larval condition (for review,

see CLEMEN & GREVEN, 1994).

Source : MNHN, Paris

CLEMEN & GREVEN 59

Also form and size of the teeth are undoubtedly larval. Teeth are slightly recurved,

monocuspid, conical and show a dividing zone that never reaches the condition found in

transformed teeth. The typical developmental sequence documented in urodelan teeth is from

undivided monocuspid to divided monocuspids in the late-stage larvae to divided bicuspid

teeth during or immediately after (partial) metamorphosis (for review, see GREVEN, 1988).

Teeth in adult paedomorphic E. neotenes therefore can be classified as late larval stage.

Transformed adult plethodontids such as Eurycea bislineata (e.g., STEWART, 1958) or

Desmognathus fuscus (NOBLE, 1931 and further references herein) show a remarkable differ-

ence in the premaxillary teeth of males and females. Depending on the testosterone level,

males have more elongate and secondarily monocuspid teeth during the breeding period and

a subsequent marked loss of these teeth. In addition, males lack the anterior vomerine

dentition (NOBLE, 1931). Our few measurements and counts indicate that males of Æ. neotenes

have longer teeth, mainly on the premaxillae. Further studies using more specimens captured

in different seasons should confirm this observation and help clarify the regulation of this

possible sex dimorphism.

Some other obviously sex-linked differences are worth noting. Males also possess broad-

er premaxillae lacking a prominent pars palatina, fully separated processus faciales, larger

anterior processes of the vomer, that overlap the parasphenoid by the fissured margins of the

toothless palatinal portion of the palatopterygoid, and large splenials.

According to DUELLMAN & TRUEB (1985: 194), “ Euryceae neotenes and Typhlotriton

spelaeus are facultative neotenes intermediate ecologically between obligate neotenes and

those species of Eurycea that undergo normal metamorphosis in surface waters”. E. neotenes

transforms not only following treatment with thyroxine, but also naturally (for literature see

Sweer, 1977). Paedomorphosis, which includes neoteny, however, is evolutionarily fixed

(GouLp, 1977). Therefore terms as “obligate paedomorphosis” and “facultative paedomor-

phosis” should be avoided (REILLY, 1994), but are in common use. As outlined by REILLY

(1994), investigations regarding interspeficic heterochrony (“patterns of phyletic change that

have become fixed in independent phylogenetic lineages”) and intraspecific heterochrony

(variation of individuals in response to environmental change”) could help elucidate the

paedomorphic state of E. neotenes. However, discussion on this is beyond the topic of our

paper.

The skull of metamorphosed individuals (see figure 6 in SWEET, 1997) corresponds in

most respects to the skull of other transformed Eurycea as characterized by WAKE (1966) and

MuTz & CLEMEN (1992). As we studied maturing and mature E. neotenes, which reproduced

in captivity (HAKER, personal communication), we do not believe that the differences found

between sexes should be attributed to different stages of development towards metamorpho-

sis, but reflect real sex specific characters. We do not exclude, however, that an increasing

number of breeding cycles (and, thus, briefly elevated thyroxine levels) may enhance meta-

morphic effects in older specimens as seen on other species (unpublished).

Source : MNHN, Paris

60 ALYTES 18 (1-2)

ACKNOWLEDGEMENTS

We thank Mr. K. Haker, Hilden, for the animals, Mrs. Dipl. Biol. M. Fasel, Münster, for the

drawings, Mr. J. Lange, Münster, for the help with the SEM and Prof. Dr. D. Sever, Notre Dame, for

linguistic advice.

LITERATURE CITED

AMEND, R. & GREVEN, H., 1996. - Zur Ossifikation des Skeletts intra- und extrauteriner Larven des

Feuersalamanders Salamandra salamandra (L.) (Amphibia, Urodela). Abk. Berichte Naturkde,

Magdeburg, 19: 31-67.

BRUCE, R. C., 1976. - Population structure, life history and evolution of paedogenesis in the salamander

Eurycea neotenes. Copeia, 1976: 242-249.

CLEMEN, G. & GREVEN, H., 1977. - Morphologische Untersuchungen an der Mundhôhle von Urodelen.

IL. Die Munddachbezahnung von Ambystoma mexicanum Cope (Ambystomatidae, Amphibia).

Zool. Jb. Anat., 98: 95-136.

ee 1988. - Morphological studies on the mouth cavity of Urodela. IX. Teeth of the palate and the

splenials in Siren and Pseudobranchus (Sirenidae: Amphibia). zool. Syst. Evol.-forsch., 26:

135-143.

—— 1994. — The buccal cavity of larval and metamorphosed Salamandra salamandra: structural and

developmental aspects. /n: H. GREVEN & B. THIESMEIER (ed.), Biology of Salamandra and Merten-

siella, Mertensiella, 4:

DuELLMAN, W. E. & TRUEB, L., 1985. — Biology of amphibians. New York, McGraw-Hill, 1986": i-xix +

1-670.

GouLp, S. J., 1977. - Ontogeny and phylogeny. Cambridge, Mass., Belknap Press: 1-501.

GREVEN, H., 1988. - Teeth of extant Amphibia, Morphology and some implications. Fortschr. Zool., 35:

451-455.

GREVEN, H. & CLEMEN, G., 1980. - Morphological studies on the mouth cavity of urodeles. VI. The teeth

of the upper jaw and the palate in Andrias davidianus (Blanchard) and A. japonicus (Temminck)

(Cryptobranchidae, Amphibia). Amphibia- Reptilia, 1: 49-59.

-- 1985. - Morphological studies on the mouth cavity of urodeles, VIIL. The teeth of the upper jaw and

the palate in two Hynobius-species (Hynobiidae: Amphibia). Z. ool. Syst. Evol.-forsch., 23

136-147.

LARSEN, J. H., 1963. — The cranial osteology of neotenic and transformed salamanders and its bearing on

interfamilial relationships. PhD Dissertation, Seattle Univ. Washington. [Not seen].

Murz, T. & CLEMEN, G., 1992. — Development and dynamics of the tooth systems in Eurycea and

comparison of the definite dentition of the palate of Gyrinophilus (Urodela: Plethodontidae). Z.

cool. Syst. Evol.-forsch

K., 1931 he biology of the Amphibia. New York, Dover: i-xviii + 1-577.

k, E. H. & KIM, D.S., 1984. — A procedure for staining cartilage and bone of whole vertebrate larvae

while rendering all other tissues transparent. Srain Technol., 59: 269-272.

ReiLLY, S. M., 1986. — Ontogeny of cranial ossification in the eastern newt, Norhopthalmus viridescens

(Caudata: Salamandridae), and ist relationship to metamorphosis and neoteny. Z Morph., 188:

215-326.

" 1987. — Ontogeny of the hyobranchial apparatus in the salamanders Ambystoma talpoideum

(Ambystomatidae) and Notophthalmus viridescens (Salamandridae): the ecological morphology

of two neotenic strategies. J Morph.. 191: 201-214.

D 1994. - The ecological morphology of metamorphosis: heterochrony and the evolution of feeding

mechanisms in salamanders. /n: P. C. WAINWRIGHT & S. M. REILLY (ed.), Ecological morphology:

integrative organismal biology, Chicago, Univ. Chicago Press: 319-338

Source : MNHN, Paris

CLEMEN & GREVEN 61

REILLY, S$. M. & ALTIG, R., 1996. - Cranial ontogeny in Siren intermedia (Caudata: Sirenidac): paedo-

morphic, metamorphie, and novel patterns of heterochrony. Copeia, 1996: 29-41.

SrewarT, M. 1958. Seasonal variation in the teeth of the two lined salamander. Copeia, 1958: 190-196.

STADMÜLLER, F., 1924. - Studien am Urodelenschädel. I. Zur Entwicklungsgeschichte des Kopfskeletts

der Salamandra salamandra. Z. Anaï. Entw Gesch., 75: 149-225.

Sweet, S.S., 1977. - Natural metamorphosis in Eurycea neotenes, and the generic allocation of the Texas

Eurycea (Amphibia). Herpetologica, 33: 364-375.

Wake, D. B. 1966. - Comparative osteology and evolution of the lungless salamanders, family Pletho-

jidac. Mem. South. Calif. Acad, 4: 1-11.

. 1980. - Paedomorphosis. J. Herp., 14: 80-81.

Wiuoer, L. W., 1925. - The morphology of amphibian metamorphosis. Northampton, Massachusetts,

Smith College Publ.: 1-100.

Corresponding editor: Franco ANDREONE.

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 62-72.

La larva de Alsodes gargola Gallardo, 1970

(Leptodactulidae, Telmatobiinae)

Maria Susana PILLADO, Cecilia A. ALONSO & Carmen A. ÜUBEDA

Centro Regional Universitario Bariloche,

Universidad Nacional del Comahue,

Unidad Postal Universidad,

8400 San Carlos de Bariloche, Argentina

The tadpole of Alsodes gargola is described on the basis of specimens

collected at the type locality (Nahuel Huapi National Park and Reserve,

Province of Rio Negro, Argentina) and nearby areas. The tadpole of A.

gargola exhibits the diagnostic characters of the genus Alsodes (e.g. oral

disc not modified, emarginate and with rostral gap; intramarginal mental

papillae present; keratodont formula normal, 1:<1>/<1>:2; proctodeal

tube present, vent opening dextrally; exotrophic aquatic larvae). The mo:

phology of the tadpoles of this species evidences adaptations to a lotic-

benthic habit, a semi-sedentary life, and slow swimming (e.g. depressed

body; dorsolateral eyes; ventral oral disc: low fins). Features of the tadpole

of À. gargola are compared with those of other species of the genus.

Remarkable homogeneity exists among morphological characters of known

tadpoles of the genus Alsodes and those of A. gargola. The most striking

differences occur between À. gargola and A. nodosus.

INTRODUCCION

El género A/sodes contiene 12 especies distribuidas en Argentina y Chile (LAVILLA 1992,

1994; F FoRMaS et al., 1997, 1998). Hasta el presente han sido descriptas las

larvas de À. australis (FORMAS et al., 1997), 4. barrioi (VELOSO et al., 1981: LAVILLA, 1983;

Diaz PÉREZ, 1984; Diaz & VALENCIA, 1985), 4. montanus (BUSSE, 1980; Diaz PÉREZ, 1984), À

monticola (FORMAS, 1975; LAVILLA, 1983; Diaz PÉREZ, 1984; Diaz & VALENCIA, 1985), À.

nodosus (FORMAS, 1975; BUSSE, 1980; LavVILLA, 1983; Diaz PÉREZ, 1984; Diaz & VALENCIA,

1985), À. pehuenche (Cri & Ro1G, 1965; Crt, 1980; LAVILLA, 1983), À. rumultuosus (LAVILLA,

1983; Diaz PÉREZ, 1984; Diaz & VALENCIA, 1985) y À. verrucosus (Diaz & NÜREZ, 1988). Las

larvas de À. kaweshkari, A. vanzolinit Y A. vittatus se desconocen hasta el presente.

Alsodes gargola es una especie endémica del noroeste patagônico, citada hasta el

momento sélo para Argentina. Habita cuerpos de agua permanentes léticos y lénticos en

ambientes montañosos hasta los 2000 metros de altitud (ÜBrba et al., 1998). Fue descripta

por GALLARDO (1970) sobre ejemplares colectados en la Laguna Tonchek (41°12°S, 71°30°W:

Source : MNHN, Paris

PILLADO, ALONSO & ÜBEDA 63

1750 m) del Cerro Catedral, Parque y Reserva Nacional Nahuel Huapi (Argentina) (fig. 1).

GALLARDO (1970) también mencioné algunas caracteristicas de la larva. Posteriormente, CET

(1976) propuso la existencia de dos subespecies, À. gargola gargola para la localidad tipo y A.

gargola neuquensis para una nueva forma proveniente de la Meseta de Lonco Luan (Provincia

del Neuquén, Argentina). C1 (1980) realizé un comentario sobre la morfologia externa de la

larva de À. g. neuquensis y LAVILLA (1983) amplié la descripcién de la larva de esta subespecie

sobre ejemplares colectados en Primeros Pinos, localidad cercana a la Meseta de Lonco Luan.

En el presente trabajo se describe la larva de A. gargola (A. gargola gargola sensu Cet, 1976)

sobre ejemplares provenientes de la localidad tipo y de localidades proximas.

ABREVIATURAS

IZUA: Instituto de Zoologia, Universidad Austral de Chile.

MACN: Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos

Aires, Argentina.

MATERIALES Y MÉTODOS

Se realizaron muestreos (1996-1998) en cuerpos de agua permanentes, situados tanto en

bosques como en ambientes altoandinos del Parque y Reserva Nacional Nahuel Huapi y

zonas aledañas. Las localidades de muestreo fueron: Laguna Tonchek en el Cerro Catedral

(localidad tipo), Cerro Challhuaco, Valle de los Perdidos, Cerro Carbôn y Arroyo Nireco

(Provincia de Rio Negro, Argentina) (fig. 1).

Luego de confirmar visualmente la presencia de adultos de A/sodes gargola se verificé su

identidad especifica mediante el reconocimiento de sus caracteres diagnésticos y posterior-

mente se colectaron larvas con redes de mano. Los ejemplares se fijaron en formalina 10 %

neutra. Algunas larvas se mantuvieron vivas en acuario hasta completar la metamorfosis para

verificar su identidad especifica, describir la coloracién y observar su comportamiento.

Los estadios larvarios se determinaron estableciendo una equivalencia con la tabla

propuesta por GOsNER (1960). La descripciôn se realizé sobre el anälisis de los caracteres

morfolégicos externos de 28 ejemplares fijados y 10 vivos entre los estadios 27 y 39. Para la

morfometria utilizada en la descripciôn y la tabla 1 se midieron 18 larvas del lote fijado,

comprendidas entre los estadios 31 y 36, que en general presentan una constancia en los

caracteres morfolégicos. Los caracteres cualitativos y el anälisis morfométrico se determina-

ron conforme a LAVILLA (1983, 1988) y LAvILLA & SCROCCHI (1986). Para la nomenclatura de

las estructuras cérneas bucales se utilizé la presentada por vaN Duk (1966). La formula

dentaria se expresé segün Dugois (1995). Las mediciones de los ejemplares se realizaron con

calibre de lectura digital (resoluciéôn 0.01 mm) y con un ocular con lentilla micrométrica Carl

Zeiss (distancia entre lineas 100 ym) incorporado a un microscopio estereoscépico Bausch &

Lomb Stereo Zoom 7. Los dibujos fueron realizados con microscopio estereoscépico Leica

Wild M3C tipo MTR 31, provisto de cämara clara Wild 308700.

Source : MNHN, Paris

ar so 30 m0

Provincia

Ubicaciôn relativa

fo —_— Provincia

f n

/ De Rio Negro

(T-D) 81 SHLATV

(ARGENTINA

Co. Chaluaéo

2062

Escala Gräfica

Fig. 1. - Localidades de procedencia del material de A/sodes gargola en el Parque y Reserva Nacional Nahuel Huapi (Argentina) y zonas aledañas. (1)

Laguna Tonchek, Cerro Catedral: (2) Cerro Challhuaco; (3) Valle de los Perdidos: (4) Cerro Carbôn; (5) Arroyo Nireco. La linea de guiones indica

el limite provincial: la linea de guiones y puntos representa el limite internacional. Las alturas de los cerros se expresan en metros sobre el nivel mar.

Source : MNHN, Paris

PILLADO, ALONSO & UÜBEDA 65

Las caracteristicas de la larva de A/sodes gargola fueron contrastadas con las que

LAviLLa (1988) señalé como diagnésticas para el género Alsodes.

Las caracteristicas larvales de A/sodes gargola fueron comparadas con las caracteristicas

de las larvas conocidas de otras especies del género en base a especimenes y/o descripciones

publicadas. Se examinaron ejemplares de 4. monticola colectados en el Lago Espejo (Provin-

cia del Neuquén, Argentina) y de la coleccién IZUA (1098.A) y material de À. australis

colectado en el Arroyo Zanjén Hondo (Provincia del Chubut, Argentina) y de la colecciôn

IZUA (1629-1630).

Las larvas de A/sodes gargola MACN 36787-90 corresponden a los estadios 27, 32, 37 y

39 respectivamente.

DESCRIPCION DE LA LARVA DE ALSODES GARGOLA

(FIG. 2-3, TAB. 1)

Descripciôn basada en 38 ejemplares, estadios 27-39, incluidos los cuatro especimenes

MACN 36787-90. La larva alcanza gran tamaño (hasta 87 mm, estadio 39). El cuerpo (fig. 2)

es deprimido y de forma eliptica en vista dorsal; en vista lateral el contorno ventral es

ligeramente convexo:; el ancho mäximo se encuentra en el tercio medio. El extremo del hocico

es redondeado en vista lateral y levemente romo en vista dorsal. Las narinas son circulares,

con märgenes que presentan un suave reborde de igual nivel en todo su perimetro; se

encuentran en posiciôn dorsolateral y levemente mäs cerca de los ojos que del extremo del

hocico. Los ojos redondos, de mediano tamaño, estän en posicién dorsolateral; la distancia

interocular corresponde aproximadamente a 1/3 del ancho mäximo del cuerpo y es similar a

la distancia internasal. El tubo espiracular es ünico e izquierdo, estä en posiciôn laterodorsal,

separado del cuerpo en su extremo distal y orientado posterodorsalmente. El tubo proctodeal

es conspicuo; nace hacia la derecha y se abre en la misma direcciôn; su abertura puede ser

visible dorsalmente. La cola es de mediana longitud (aproximadamente 60 % del largo total),

con el eje recto, la musculatura desarrollada y el extremo redondeado; la altura de la cola es

algo menor que la altura del cuerpo; las aletas dorsal y ventral son subparalelas entre si y al eje

axial. La aleta dorsal se origina en la unién cola-cuerpo y el nacimiento de la aleta ventral estä

enmascarado por el tubo proctodeal.

El disco oral (fig. 3) estä en posicién subterminal ventral; su ancho corresponde aproxi-

madamente à la mitad del ancho mäximo del cuerpo; es intra-angular (emarginado) y

presenta claro rostral cuyo ancho es mayor que la mitad del ancho mäximo del disco oral;

posee una ünica hilera de papilas marginales que en vista dorsal pueden sobresalir del

contorno lateral de la cabeza, una hilera de papilas intramarginales mentonianas y papilas

laterales en la regiôn supra e infra-angular en nümero variable (1-4) o ausente. La férmula

dentaria es 1:<1>/<1>:2; los labios superiores siguen una linea curva y los inferiores una linea

recta; los queratodontes son numerosos; los rostrodontes estän fuertemente queratinizados y

sus bordes libres finamente aserrados; el infrarrostrodonte tiene forma de “V” y el suprarros-

trodonte de arco.

Source : MNHN, Paris

66 ALYTES 18 (1-2)

10 mm

Fig. 2. - Larva de A/sodes gargola (MACN 36789, estadio 37, Cerro Challhuaco, 2 marzo 1996): (a) vista

lateral: (b) vista dorsal; (c) vista ventral.

Source : MNHN, Paris

PILLADO, ALONSO & ÜBEDA 67

adio 37, Cerro Challhuaco, 2 marzo

Fig. 3.- Disco oral de la larva de A/sodes gargola (MACN 36789, es

1996).

La coloraciôn de los especimenes vivos es variable, desde negro a castaño claro con

manchas irregulares mäs oscuras y brillos dorados (fig. 2a). El color de la regiôn dorsal del

cuerpo presenta la pigmentaciôn mäs intensa, alcanzando el negro uniforme (fig. 2b), en tanto

que el vientre estä menos pigmentado y permite visualizar las asas intestinales (fig. 2c); las

narinas y el espiraculo tienen un anillo subterminal negro: el iris es castaño oscuro o negro con

un punteado dorado cobrizo: el disco oral muestra la mayor pigmentacién en la cara dorsal de

las papilas marginales y las estructuras cérneas son de color castaño oscuro a negro; el tubo

proctodeal exhibe una pigmentaciôn similar a la del vientre; la cola presenta una coloraciôn

variable con un patrôn de manchas mäs grandes y nitidas que las del cuerpo: la pigmentaciôn

de las aletas es menor que la de la musculatura caudal. Algunos ejemplares tienen una linea

vertebral clara que se extiende desde el hocico hasta el tercio medio de la cola; esta linea se

conserva en el juvenil y en el adulto. En ejemplares fijados la coloraciôn es castaño grisäceo a

ados; el borde de las narinas, del espiräculo y las

gris oscuro, sin manchas ni brillos doi

estructuras corneas varian de negro à castaño oscuro.

Source : MNHN, Paris

Tabla 1. — Promedios + desvios eständar de las medidas y relaciones morfométricas de la larva de Alsodes gargola. Los estadios corresponden a los

propuestos por GOSNER (1960). Las medidas estän expresadas en mm.

(T-1) 8 SALAIV

Estadio 31 32 33 34 35 36

Nümero de ejemplares o) 3 1 2 3 4

Largo total 47,92+4,93 | 54454248 | 43,85 | 5868415290] 57,8149,54 | 64,314 8,07

Largo cuerpo 19,87+2,29 | 2209+0,84 | 19,10 | 22,6543,70 | 21,53+1,39 | 24,394 1,64

Altura mâxima cuerpo 9554140 | 10,50+1,02 | 9,57 | 10,49+013 | 11,03+1,66 | 12,424+1,48

Altura cola 7,70 4 1,44 9324029 | 7,44 8,62 +0,91 8,46 + 0,60 9,98 + 1,74

[ess mäximo cuerpo 10,98+1,50 | 11,56+0,42 | 11,78 | 12,61+1,02 | 12,02+1,52 | 13,82+1,78

Diämetro ojo 1,50 + 0,35 1674023 | 1,50 1,784 0,18 2,03 40,15 2,12 +0,49

Distancia interocular 3,76 +0,42 4164020 | 3,24 4,27 4 0,62 4,2340,25 4,43 40,43

Distancia internasal 3,63 +0,66 41340,12 | 4,10 4374033 3,8740,35 4,19 +0,14

Distancia narina-hocico 2,62 40,43 2844026 | 2,70 3,05 +0,92 2,57+0,31 3,30 +0,99

Distancia ojo-hocico 5,17 40,68 564+0,72 | 4,90 6,05 +2,05 5,30 +0,26 6,35+0,95

Distancia espiräculo-hocico 11,16+1,14 | 12,8440,91 | 9,75 | 12,88+3,23 | 12,24+1,54 | 13,924 1,39

Ancho disco oral 5,05 +0,38 5764006 | 5,49 5,82 + 0,40 5,754 0,19 6,34 +0,38

Ancho claro rostral 3,08 +0,19 3224011 | 3,32 3,08 +0,32 3,394 0,07 3.144035

Largo cuerpo/largo total 0,41 +0,02 041+0,02 | 0,4 0,39 4 0,04 0,384 0,05 0,38 + 0,02

Ancho mäximo cuerpo/altura mäxima cuerpo 1,16 +0,18 1,1140,08 | 123 1,20 + 0,08 1,09 + 0,05 1,11 40,05

Distancia espiräculo-hocico/largo cuerpo 0,56 + 0,02 0,5840,04 | 0,51 0,56 + 0,05 0,57 + 0,04 0,57 + 0,02

Largo cuerpo/altura méxima cuerpo 2,11 40,30 21140,19 | 2,00 2,16 +0,33 1974021 1,97+0,13

Distancia interocular/distancia internasal 1,05 +0,12 1014005 | 0,79 0,97 + 0,07 1,10 + 0,09 1,06 + 0,13

[pisanca interocular/ancho mäximo cuerpo 0,34 40,03 0364001 | 0,28 0,34 + 0,02 0,35 +0,02 0,32 +0,03

Ancho claro rostral/ancho disco oral 0,61 +0,07 0,56+0,02 | 0,60 0,53 + 0,09 0,59 + 0,02 0,49 + 0,03

Distancia narina-hocico/distancia narina-ojo 1,17 # 0,17 1,3540,06 | 1,23 1,17+0,04 1,22 +0,09 1,39 +0,43

Distancia hocico-espiräculo/distancia espiräculo-base tubo proctodeal 1294011 1404024 | 1,04 1,3140,27 1324021 1334011

Source : MNHN, Paris

PILLADO, ALONSO & ÜBEDA 69

BREVE COMENTARIO SOBRE EL COMPORTAMIENTO DE LA LARVA EN ACUARIO

En acuario, las larvas frecuentan el fondo y las paredes del recipiente. Utilizan las rocas

para descansar o encontrar refugio al percibir disturbios y también para alimentarse del

epiliton. En el medio artificial las larvas pueden ser alimentadas con pienso escamado para

peces de origen vegetal o con carne molida cruda: el alimento es râpidamente detectado.

Ocasionalmente se observé a las larvas consumiendo los tejidos blandos de individuos

muertos. Las larvas cuando se alimentan varian sus posiciones y movimientos segün el

sustrato; si se alimentan contra el fondo del recipiente disponen el cuerpo paralelo a éste y

realizan movimientos lentos; al alimentarse sobre las paredes, los movimientos son mäs

enérgicos y el cuerpo forma un ängulo agudo con respecto al sustrato. Pueden captar el

alimento sobre la superficie del agua colocändose boca arriba y dejando el cuerpo suspendido.

DISCUSIÔN Y CONCLUSIONES

La larva de A/sodes gargola presenta las caracteristicas morfolégicas y meristicas (tab. 1)

que LAVILLA (1988) reconocié como diagnésticas para el género A/sodes: ancho del disco oral

menor que los 2/3 del ancho mäximo del cuerpo; disco oral emarginado, excepto 4. monticola

y con claro rostral; papilas intramarginales mentonianas; rostrodontes ms anchos que altos;

férmula dentaria 1:<1>/<1>:2; nivel de la abertura de las narinas no elevado: relaciôn entre

la distancia narina-hocico y narina-ojo > |; relaciôn entre la distancia hocico-espiräculo y

espiräculo-base del tubo proctodeal > 1: espiräculo izquierdo lateral; tubo proctodeal con

abertura dextral; aletas de tipo normal; desarrollo larvario acuätico y larva con alimentaciôn

activa. La larva de À. gargola también presenta todas las caracteristicas que Diaz (1992)

consideré como tipicas de las larvas del género 4/sodes (disco oral > 2 del ancho mäximo del

cuerpo; äreas comisurales del disco sin papilas; papilas intramarginales mentonianas; rela-

ciôn entre la distancia interocular y la distancia internasal < 1,6) (tab. 1).

La comparacién entre la larva de A/sodes gargola ÿ las restantes larvas descriptas del

género muestra algunas diferencias y similitudes.

Todas las especies descriptas del género A/sodes tienen el disco oral con constricciones

angulares, incluyendo a 4. monticola a diferencia de lo mencionado por LAVILLA (1988). En

À. gargola el ancho del disco oral es similar a la mitad del ancho mäximo del cuerpo al igual

que en À. barrioi, A. monticola, A. nodosus y A. tumultuosus: en À. pehuenche es menor y en À

verrucosus es mayor. A/sodes gargola, como ocurre con 4. australis, A. barrioi, A. montanus, A.

monticola, A. tumultuosus, A. pehuenche y A. verrucosus, presenta una hilera de papilas

intramarginales mentonianas, a diferencia de 4. nodosus que posee dos hileras de papilas.

La longitud de la cola en relacién con la del cuerpo se presenta con bastante variaciôn

dentro del género, desde 1,2 veces la longitud del cuerpo (A/sodes barrioi) a 1,6 veces (4.

australis, A. gargola, A. montanus y À. verrucosus) y hasta 1,7 a 1,8 veces (4. monticola, À

nodosus y A. tumultuosus). Todas las especies descriptas incluyendo à 4. gargola tienen el

extremo de la cola redondeado, excepto 4. nodosus que lo tiene suavemente puntiagudo.

Source : MNHN, Paris

70 ALYTES 18 (1-2)

Segün las descripciones y datos propios, las especies exhiben una coloracién similar,

excepto A/sodes barrioi (VELOSO et al., 1981) y À. monticola que tienen en el dorso dos

manchas irregulares amarillentas a ambos lados de la linea media.

Entre las especies comparadas, la larva de A/sodes gargola presenta las mayores diferen-

cias morfolôgicas externas con la larva de 4. nodosus, en cuanto al nümero de hileras de

papilas intramarginales mentonianas y a la longitud y la forma del extremo de la cola. En

cuanto a la longitud de la cola con respecto al cuerpo, la mayor diferencia la presenta con 4.

barrioi.

La larva descripta en este trabajo (A/sodes gargola gargola sensu Cet, 1976) no presenta

diferencias en cuanto a la morfologia externa y a la coloracién con la larva de la subespecie 4.

gargola neuquensis.

De la gran similitud que presentan las caracteristicas comparadas se pone de manifiesto

la remarcada homogeneidad y la constancia generalizada en la mayoria de los caracteres

morfolégicos externos de las larvas del género A/sodes, hecho también señalado por VELOSO

et al. (1981) y Diaz & VALENCIA (1985).

Por ültimo, merece destacarse que la descripcién de la larva de A/sodes verrucosus fue

realizada sobre ejemplares provenientes de la Isla Wellington, Chile (Diaz & NÜREZ, 1988),

donde, segün el estado del conocimiento actual, también existen otras dos especies del género,

A. monticola y A. kaweshkari (GRANDISON, 1961; FORMAS et al., 1998), no existiendo certeza

sobre la asignaciôn especifica de esas larvas.

La larva de Alsodes gargola por su morfologia externa corresponde al tipo “central

generalizado” segün la clasificacién propuesta por ORTON (1953) y pertenece al gremio

lôtico-benténico (Secciôn I, B7) segün ALTIG & JOHNSTON (1989). Algunas de las caracteris-

ticas de la larva como el cuerpo deprimido, los ojos dorsolaterales, el disco oral en posiciôn

ventral y las aletas bajas, evidencian adaptaciones al häbito semisedentario y de nataciôn

lenta. Todas las larvas descriptas del género A/sodes presentan häbitos similares a À. gargola

y, al igual que ésta, habitan cuerpos de agua permanentes lôticos con lecho de grava y arena,

aguas cristalinas y escasa vegetaciôn acuätica. A/sodes pehuenche también habita cuerpos de

agua sulfur Alsodes gargola presenta un largo periodo larvario y metamérfico (ÜBEDa et

1998), caracteristica corroborada en los ejemplares mantenidos en acuario, que superaron

los 24 meses de vida como larvas. La etapa larvaria de larga duraciôn es un fenémeno comün

en otros telmatobiinos.

RESUMEN

Se describe la larva de A/sodes gargola sobre la base de especimenes colectados en la

localidad tipo (Parque y Reserva Nacional Nahuel Huapi, provincia de Rio Negro, Argen-

tina) y en zonas aledañas. La larva de 4. gargola presenta los caracteres larvales diagnésticos

dados para el género (e.g. disco oral no modificado, emarginado y con claro rostral: papilas

intramarginales mentonianas presentes; férmula de queratodontes normal, 1:<1>/<1

tubo proctodeal presente, con abertura dextral: larva acuätica exotrôfica). La morfologia

exhibe adaptaciones al häbito lético-benténico y a una vida semisedentaria y de nataciôn

lenta (e.g. cuerpo deprimido, ojos dorsolaterales: disco oral ventral: aletas bajas). Se compa-

Source : MNHN, Paris

PILLADO, ALONSO & ÜBEDA 71

ran caracteristicas de la larva de À. gargola con las de otras especies del género, poniéndose de

manifiesto una remarcada homogeneidad de los caracteres morfolégicos. Las principales

diferencias se presentan con la larva de À. nodosus.

AGRADECIMIENTOS

A la Administraciôn de Parques Nacionales que autorizé realizar los estudios de campo en el Parque

y Reserva Nacional Nahuel Huapi. A D. D. Echeverria y a E. O. Lavilla por el asesoramiento y

sugerencias brindados para la realizaciôn de este trabajo, a J. R. Formas por permitir y facilitar la revision

del material del Instituto de Zoologia de la Universidad Austral de Chile, À K. R. Lips, C. A. Sheil y a un

revisor anénimo por las sugerencias realizadas que enriquecieron el presente trabajo. Este estudio fe

financiado con un subsidio para la investigaciôn otorgado por la Universidad Nacional del Comahue

{Proyecto 04/B083) y con el subsidio PICT 01-03698.

LITERATURA CITADA

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modes, morphologies, and habitats. Herp. Mon. 3: 81-109.

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Beschreibung seiner Larve (Amphibia: Leptodactylidae). Amphibia- Reptilia, À (2): 113-125

Cr, J. M., 1976. - Remarks on some neotropical amphibians of the genus A/sodes from Southern

Argentina (Anura Leptodactylidae). Ari Soc. ital. Sci. nat. Museo civ. Stor. nat. Milano, 117 (3-4):

159-164.

Æ 1980. - Amphibians of Argentina. Monit. zool. ital., (n.s.), Monogr. 2: [i-xii] + 1-609.

Ce, J. M. & RoG, V. G., 1965 . - The systematic status and biology of 7elmatobius montanus La

(Amphibia, Leptodactylidae). Copeia, 1965 (4): 421-425.

Diaz, N. F., 1992. - Phenetic and phylogenetic relationships of the Chilean A/sodes and Telmatobius

(Amphibia, Leptodactylidae) and proposal of a new genus. Stud. neotrop. Fauna & Environm., 27

(} 1-8.

F. & NÜKEZ, H., 1988. - Nuevo hallazgo de A/sodes verrucosus (Philippi, 1902) en Chile y

cripciôn de su larva (Anura: Leptodactylidae). Bol. Mus. nac. Hist. nat. Chile, 41: 87-94.

Diaz, N. F. & VALENCIA, JL. 1985. - Larval morphology and phenetic relationships of the Chilean

Alsodes, Telmatobius, Caudiverbera and Insuetophrynus (Anura: Leptodactylidae). Copeia, 1985

175-181.

Diaz PÉREZ, N. F., 1984. - Biosistemätica y relaciones filogenéticas de las especies chilenas de Leptodac-

tylidae (Amphibia: Salientia). enfoque multidisciplinario. Tesis Doctoral, Fac. Ciencias Bäsicas y

Farmacéuticas, Univ. Chile: [i-xxxiii] + 1-174.

Dunors, À., 1995. - Keratodont formulae in anuran tadpoles: proposals for a standardization. J. :ool.

Syst. evol. Res.. 33 (1): i-xv.

FORMAS, J. R., 1975. - Las larvas de las especies chilenas pertenecientes al género Eupsophus, grupo

nodosus (Anura: Leptodactylidae). Bol. Soc. Biol. Concepciôn, 49: 231-237.

= 1995. — Anfibios. Zn: J. A. SimonErTI. M. T. K. ARROYO, A. E. SPOTORNO & E. LOZADA (ed

Diversidud biolôgica de Chile, Santiago. Chile, Comité Nacional de Diversidad Biolôgica, Comi-

siôn Nacional de Investigaciôn Cientifica y Tecnolé 314-325.

FORMAS, J. R., CUEVAS, C. & NUREZ, J., 1998. — À new species of A/sodes (Amphibia: Anura: Leptodac-

lidae) from southern Chile. Proc. biol. Soc. Wash.. I (3): 521-530.

FoRMAS, J. R. UBepa, C. Cuevas, C. & NUREZ. J., 1997. — Alsodes australis, a new species of

leptodactylid frog from the temperate Norhofagus forest of southern Chile and Argentina. Stud.

neotrop. Fauna & Environm., 32: 200-211

ste

Source : MNHN, Paris

72 ALYTES 18 (1-2)

GaLLarDo, J. M., 1970. — A propésito de los Telmatobiinae (Anura, Leptodactylidae) patagénicos.

Neotropica, 16 (50): 73-85.

Gosxer, K. L., 1960. — À simplified table for staging anuran embryos and larvae with notes on

identification. Herpetologica, 16: 183-190.

Graxpiso, À. G. C., 1961. - Chilean species of the genus Eupsophus (Anura: Leptodactylidae). Bull.

brit. Mus. (nat. Hist.), Zoo. 8 (8): 111-149, pl. 1-7, 1 map.

LaviLLA, E.O,, 1983. - Sistemätica de larvas de Telmatobinae ( Anura: Leptodactylidae). Tesis Doctoral,

Fac. Ciencias Naturales, Univ. Nac. Tucumän, Tucumän: [i-v] + 1-354.

Pen 1988. — Lower Telmatobiinae (Anura: Leptodactylidae): generic diagnoses based on larval charac-

ters. Occ. Pap. Mus. nat. Hist. Univ. Kansas, 124: 1-19.

ae 1992. - Tipos portadores de nombre y localidades tipo de anfibios de Argentina. Acta zool. lilloana,

42 (1): 61-100.

NS 1994. - Comentarios sobre el género Telmalsodes Diaz, 1989 (Anura: Leptodactylidae). Bol. Asoc

Herp. arg, 10 (1): 15-18.

LaviLa, E. O. & ScroccHi, G. J., 1986, - Morfometria larval de los géneros de Telmatobinae (Anura:

Leptodactylidae) de Argentina y Chile. Physis, (B), 44 (106): 39-43.

Ortow, G. L., 1953. — The systematics of vertebrate larvae. Syst. Zool., 2 (2): 63-75.

UÜnrpa, €. ALONSO, C. & PILLADO, M. $., 1998. - Alsodes gargola, un anfibio endémico patagénico con

adaptaciones a la vida en altura. Ju: Fauna andinopatagénica: aportes a su conocimiento, Patagonia

Silvestre, Serie Técnica, 2: 1-9.

VaN Dur, D. E., 1966. — Systematic and field keys to the families, genera and described species of

Southern African Anuran tadpoles, with preliminary inclusion of certain undescribed and inade-

quately described tadpoles. Ann. Natal Mus., 18 (2): 231-286.

VELoso, A., Diaz, N., ITURRA, P. & PENNA, M., 1981. - Descripciôn de una nueva especie de Telmatobino

del género Alsodes (Amphibia, Leptodactylidae) de la cordillera de Nahuelbuta (sur de Chile).

Medio Ambiente, 5 (1-2): 72-7.

Corresponding editor: Karen R. LiPs

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 73-80. 73

The tadpoles of Hyla oliveirai

and Hyla decipiens

with notes on the Hyla microcephala

group (Anura, Hvylidae)

Adriana PUGLIESE!, Ana Claudia Reis ALVES

& Sergio Potsch DE CARVALHO E SILVA

Departamento de Zoologia, Universidade Federal do Rio de Janeiro,

Caixa Postal 68044, 21944-970 Rio de Janeiro, RJ, Brazil

The previously unknown tadpole of Hyla oliveirai is described and

illustrated based on a population from Alagoas State, northeastern Brazil.

The tadpole of Hyla decipiens is briefly redescribed and illustrated from

Rio de Janeiro State, southeastern Brazil, and compared with that of H.

oliveirai. The latter has a single, wide brown irregular stripe on the tail,

while H. decipiens has three or four narrower, more regular stripes. The

oral disc of both species does not bear denticles and has a single row of

papillae extending ventrally and laterally. Hyla oliveirai has just one ridge

between the beak and the papillae on the lower lip, while H. decipiens has

two ridges. The tadpoles of H. oliveirai and H. decipiens are similar to that

of H. berthalutzae in the general shape, colour pattern and oral disc. These

characteristics, added to adult characteristics, may suggest a close rela-

tionship between these species. Comparisons between these tadpoles and

other species from the Hyla microcephala group show remarkable differ-

ences, suggesting that H. oliveirai, H. decipiens and H. berthalutzae do

not belong in the H. microcephala group.

INTRODUCTION

The small species of Æyla have been arranged into groups by COCHRAN (1955), COCHRAN

& GoiN (1970), DUELLMAN (1970) and Lurz (1973). All of these species that have been studied

karyologically have 30 chromosomes (DUELLMAN & TRUEB, 1983). The Hyla microcephala

group is one of these groups and includes small, yellowish-tan species. Highly variable

phenetic arrangements have been proposed for this group (e.g., DUELLMAN & FOUQUETTE,

1968; COCHRAN & GoiN, 1970; Basros & POMBAL, 1996). Much of the confusion in these

different arrangements has been the result of a lack of knowledge on the osteology, tadpoles,

mating calls, and biology of the included species, making it difficult to determine more

sharply the relationships among these small hylid frogs.

1. Present address: Departamento de Vertebrados, Museu Nacional, Quinta da Boa Vista, 20940-040

Rio de Janeiro, RJ, Brazil.

Source : MNHN, Paris

74 ALYTES 18 (1-2)

As recognized by POMBAL & BASTOS (1998), the Hyla microcephala group contains 15

species, seven of them being known from the Atlantic Forest and one from the “cerrado”

(sensu AB’sABER, 1977) in Brazil (. berthalutzae Bokermann, 1962; H. branneri Cochran,

1948; II. decipiens À. Lutz, 1925; H. haddadi Bastos & Pombal, 1996; H. meridiana Lutz, 1973;

H. oliveirai Bokermann, 1963; H. werneri, Cochran, 1952; and H. cruzi Pombal & Bastos,

1998, the latter from the “cerrado”). None of the Brazilian species, except A. werneri, were

traditionally recognised in the Hyla microcephala group (DUELLMAN & FOUQUETTE, 1968:

DUELLMAN, 1970).

Hyla oliveirai and Hyla decipiens are very similar species with known distribution from

northeastern to southeastern Brazil (FROST, 1985; Basros & POMBAL, 1996). The tadpole of

H. decipiens was described, without illustrations, from Rio de Janeiro State by LUTz (1973).

Herein, we describe the tadpole of A. oliveirai and compare it to that of A. decipiens. Notes

about other closely related species are added.

MATERIAL AND METHODS

All specimens examined in this study are deposited in the collection of Departamento de

Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro (ZUFRIJ). Adults

and tadpoles of H. oliveirai (ZUFRJ 7326-7328, 7446) were collected from a population in

Alagoas State (Municipio de Quebrangulo; 9°19’S, 36°28°W) in November 1997. Adults and

tadpoles of H. decipiens (ZUFRJ 4441) were collected from a population in Rio de Janeiro

State (Municipio de Itaguai; 22°44'S, 43°42°W) in March 1991. Coordinates were taken from

local maps.

Adults were anaesthetised in 0.25 chloretone, preserved in 10 % formalin, and stored

in 70 % alcohol. Tadpoles were anaesthetised in 0.1 % chloretone and stored in 5 % formalin.

Two males and a female of A. oliveirai were kept alive and put in a plastic bag where eggs were

obtained. The tadpoles that emerged from these eggs were reared until stages 33-37 (GOSNER,

1960) for study. Tadpoles collected in the field were identified by means of comparisons with

those obtained from eggs.

The tadpoles of Æ. oliveirai obtained from eggs and others collected in temporary ponds

of an open area were raised in captivity in a plastic box (measurements: 262 X 77 X 147 mm)

with about 1.5 1 of water and a dense layer of leaves on the bottom. Eleven tadpoles were

maintained in the box. Fish food was regularly provided. The water temperature and pH were

not controlled. The studied tadpoles of A. decipiens were not raised. However, they were

compared with other specimens previously reared by SPCS from the eggs to metamorphosis.

Tadpoles reared from the eggs until stages 36-37 were used in the descriptions and

measurements. No changes were observed in the oral morphology or general shape of reared

tadpoles. Eventually other stages were used for comparisons and to provide a better account

of colour pattern development. Measurements taken according to the methods pro-

posed by ALTIG (1970) and DUELLMAN (1970), except interorbital distance which was taken

between the inner margins of eyes. An ocular micrometer in a stereomicroscope was used for

measurements, except for the total length, which was taken with a caliper. Drawings were

made using a stereomicroscope with a camera lucida.

Source : MNHN, Paris

PUGLIESSE, ALVES & DE CARVALHO E SILVA 75

RESULTS

HYLA OLIVEIRAI BOKERMANN, 1963 (FIG. la-d)

Description. - Mean total length at stage 37, 20.7 + 0.3 mm (7 = 7; tab. 1). Body oval in lateral

view, snout rounded in dorsal view. Body width 65 % (60-70 %) of body length. Eyes lateral,

visible ventrally; interorbital distance about twice of eye diameter. Nostrils about six times in

eye diameter, rounded, directed ventrally on snout tip. Internostril distance similar to inter-

orbital distance. Spiracle sinistral, short, slightly projecting; distance from spiracle to snout

approximately 67 % (60-74 %) of body length. Anal tube short, dextral, attached to ventral

fin. Tail length 64 % (62-67 %) of total length, tail musculature tapering gradually to the tip.

Dorsal fin starting at final portion of body, slightly arched in lateral view; ventral fin

rectilinear in lateral view. Lateral line system invisible. Oral disc anteroventral, its width

approximately 23 % (21-27 %) of body width, with single row of rounded papillae extending

ventrally and laterally, without denticles; lower lip with one dermal ridge between beak and

papillae. Beak strong, finely serrated: upper jaw slightly arched and low jaw approximately

U-shaped.

Colour in 5 % formalin. - Body brown in dorsal view, in lateral view with narrow cream stripe

extending from snout to lower margin of eye. Body in ventral view brown with marble-like

aspect in anterior region and slightly transparent in posterior region. Narrow cream stripes

Hyla oliveirai (ZUFRI 7446). Tadpole in (a) lateral, (b) dorsal and (c) ventral views: (d) oral disc.

Fig. 1

Source : MNHN, Paris

76 ALYTES 18 (1-2)

Table 1.- Measurements (x + s) in millimeters of tadpoles in stage 36-37.

Measurement Hyla oliveirai (n = 7) Hyla decipiens (n = 10)

Total length 20.7 + 0.3 20.8 + 0.4

Body length 7.5+02 7.7#0.1

Body width 49+#0.1 4.6+0.1

Body height 41+0.1 4.1+#0.1

Tail height 40+0.1 4.5+0.1

Dorsal fin 1.2+0.1 1,5+0.1

Ventral fin 1.240. 14+#0.1

Spiracle-snout distance 64+0.2 4.5+0.1

Interorbital distance 2.6+0.0 2.5+0.1

Internostril distance 25+0.0 2.4 0.0 |

Eye diameter 1.24 0.0 1.3 +0.0

Nostril diameter 0.2+0.0 0.2+0.0

Mouth width 1.1+0.0 1.0 + 0.0 |

usually visible on both sides of oral disc, extending to lower margin of eyes in ventral view,

forming an inconspicuous inverted “V”. Two rounded, slight cream to yellowish-brown

depressions under mouth between eyes. Tip of spiracle cream. Tail in lateral view cream to

yellowish with base of dorsal fin and tail musculature brown and an irregular, transverse

brown stripe in posterior half on which yellowish spots (usually one on each fin) may be

present.

Colour in life. - Tadpoles in life similar to preserved ones, but with more brilliant colours. Iris

coppery. Newly metamorphosed frog with a nacreous triangle on snout and two broad

nacreous bands disposed laterally from posterior margins of eyes to end of body; area

between bands with subrectangular, grayish-brown mark enclosing grayish-brown middle

line.

HYLA DECIPIENS A. LUTZ, 1925 (FIG. 2a-d)

Description. —- Mean total length at stage 37, 20.8 + 0.4 (n = 10; tab. 1). Body width

corresponding to 61 % (55-63 %) of body length. Tail length approximately 63 % (60-65 %) of

total length, tail musculature tapering abruptly to the tip; fins with rectilinear outline. Oral

disc with approximately 21 % (17-23 %) of body width, with single row of slightly rounded

and short papillae extending ventrally and laterally; papillae considerably attached to the lip:

lower lip with two dermal ridges between beak and papillae. Otherwise as described above for

Hyla oliveirai.

Colour in 5 % formalin. — Body brown in dorsal view, in lateral view with narrow cream stripe

extending from snout to lower margin of eye. Body in ventral view brown with marble-like

aspect. Distinct, narrow cream stripes visible on both sides of oral disc, extending to lower

Source : MNHN, Paris

PUGLIESSE, ALVES & DE CARVALHO E SILVA 77

ET]

Fig. 2. - Hyla decipiens (ZUFRJ 4441). Tadpole in (a) lateral, (b) dorsal and (c) ventral views: (d) oral

disc.

margin of eyes in ventral view, forming a conspicuous inverted “V”. Tip of spiracle cream.

Tail in lateral view cream to yellowish with base of dorsal fin and tail musculature brown and

three or four irregular, transverse brown stripes.

DISCUSSION

The tadpoles of Hyla oliveirai and Hyla decipiens have a similar colour pattern and are

found in temporary ponds in open areas. Both species present a narrow cream stripe extending

from the snout to the lower margin of the eye in lateral view and tail with transversal stripes,

alternating brown and cream. They can be distinguished from each other by the number of

transverse brown stripes in the tail: one wide stripe with a very irregular outline in A. oliveirai

and three or four narrower and more regular stripes in Æ. decipiens. The tail narrows

gradually in A. oliveirai, while it narrows abruptly in H. decipiens. The oral disc of both

species is similar: reduced, without denticles, and with a single row of papillae extending

ventrally and laterally. Hyla oliveirai has just one dermal ridge between the beak and the

papillae in the lower lip, while F. decipiens has two dermal ridges. The papillae in the oral disc

of H. decipiens are less rounded, shorter and more attached to the lip than in Æ. oliveirai.

The tadpoles of H. oliveirai and H. decipiens are similar to that of H. berthalurzae in the

general shape and colour pattern. As in the former two species, the oral disc of the tadpole of

Source : MNHN, Paris

78 ALYTES 18 (1-2)

H. berthalutzae lacks denticles and has a single row of papillae extending ventrally and

laterally, and two ridges are present between the beak and the papillae in the lower lip

(BOKERMANN, 1963). Hyla berthalutzae and H. decipiens, as well as H. branneri, have very

similar reproductive habits, although the latter two occur in open areas (LUTZ, 1947; BOKER-

MANN, 1963; BasTOs & POMBAL, 1996). They deposit the spawn on small dry sticks or leaves

hanging above the ponds where the tadpoles will develop. Based on the characteristics of

tadpoles, we agree with Basros & PomBAL (1996) about the close relationship between H.

oliveirai, H. decipiens and H. berthalutzae. According to those authors, it would also be

possible to relate H. haddadi to these three species based on the adult morphology.

Comparisons between tadpoles of species traditionally included in the Hyla microce-

phala group (see DUELLMAN & FOUQUETTE, 1968; DUELLMAN, 1970) with those of F. decipiens

and A. oliveirai show remarkable differences. The typical tadpole of that group (based on /.

bipunctata Spix, 1824, H. meridiana, H. microcephala Cope, 1886, H. nana Boulenger, 1889,

H. phlebodes Stejneger, 1906, H. rhodopepla Günther, 1859 and H. sanborni Schmidt, 1944

tadpoles: BOKERMANN, 1963; KENNY, 1969: DUELLMAN, 1970, 1972; LAviLLA, 1990; CRUZ &

Dias, 1991) has an elongate and depressed body in lateral view (ovoid in A. decipiens and H.

oliveirai), eyes almost in the middle of the body length (in the anterior third in A. decipiens and

H. oliveirai), dorsal fin starting at the posterior third of the body (starting at the end of the

body in A. decipiens and H. oliveirai), tail remarkably xiphicercal (just pointed in H. decipiens

and A. oliveirai), and oral disc without papillae and ridges between beak and lower lip (with

a single row of papillae and one or two ridges in H. decipiens and H. oliveirai).

DUELLMAN & TRUEB (1983) assumed that the small hylid frogs with 30 chromosomes

form a monophyletic complex, emphasizing that the major evolutionary trends in this

complex involve the larval structure, especially the mouthparts. They provided a hypothesis of

the cladistic relationships among seven species groups of Hyla (H. labialis group, H. colum-

biana group, H. minuta group, H. marmorata group, H. parviceps group, H. leucophyllata

group and A. microcephala group). The H. microcephala group, previously defined by

DUELLMAN (1970), appeared in their study as a monophyletic group with two synapomorphies

(both involving larval characteristics): body depressed and labial papillae absent. The remark-

able differences (specially those involving oral structure) between the tadpoles of Æ. decipiens,

H. oliveirai and H. berthalutzae and those of the other species traditionally recognized in the

H. microcephala group suggest that the former ones do not belong to this group.

RESUMÉ

Le têtard de Hyla oliveirai est décrit et illustré à partir de spécimens provenant d’une

population de l’état d'Alagoas, au nord du Brésil. Un têtard de Hyla decipiens provenant du

Rio de Janeiro, au sud du Brésil, est brièvement redécrit, illustré, et sa morphologie est

comparée avec celle de A. oliveirai. Le têtard de H. oliveirai a une unique bande marron large

etirrégulière sur la queue, tandis que Æ. decipiens a trois où quatre bandes plus étroites et plus

régulières. Le disque oral de ces deux espèces n'a pas de denticules et a une seule rangée

ventrale et latérale de papilles. Le têtard de A. oliveirai a un seul repli entre le bec et les papilles

Source : MNHN, Paris

PUGLIESSE, ALVES & DE CARVALHO E SILVA 79

sur la lèvre inférieure, tandis que H. decipiens en a deux. Les têtards de H. oliveirai et de H.

decipiens sont semblables à ceux de . berthalutzae en ce qui concerne leur forme d’ensemble,

leurs couleurs et leur disque oral. Ces caractéristiques, ajoutées à celles des adultes, suggèrent

une proche parenté entre ces deux espèces. En revanche, la comparaison de ces têtards avec

ceux d’autres espèces du groupe de Hyla microcephala montre l'existence de nettes différences,

suggérant que Æ. oliveirai, H. decipiens et H. berthalutzae n’appartiennent pas au même

groupe que FH. microcephala.

ACKNOWLEDGEMENTS

We are grateful to Carlos J. E. Lamas, Gabriel Mejdalani (Museu Nacional, UFRJ), Marcia R.

Gomes (IB, ÜFRJ) and two anonymous reviewers for their useful comments on the manuscript. Ana C.

T. Bonecker (1B, UFRI) lent the ocular micrometer, Financial support for fieldwork was provided by

Anita Studer from the Association Nordesta.

LITERATURE CITED

An'Sa, A. N., 1977. - Os dominios morfoclimäticos na América do Sul. Geomorfologia, 52: 1-21.

ALTIG, R., 1970. À key to the tadpoles of the continental United States and Canada. Herperologica, 26

(2): 180-207.

Basros, R. P. & POMBaL, J. P, Jr, 1996. - A new species of Hyla (Anura: Hylidae) from castern

Brazil. Amphibia-Reptilia, 17: 325-331.

BOKERMANN, W. C. A., 1963. - Girinos de anfibios brasileiros. I (Amphibia, Salientia). An. Acad. Bras

Ci, 35 (3): 465-474.

Cocnran, D. M., 1955. - Frogs of southeastern Brazil. U. S. nat. Mus. Bull., 206: 1-423.

Cocnran, D. M. & Goin, C. J., 1970. - Frogs of Colombia. U. S. nat. Mus Bull., 288: 1-655.

Cruz, C. A. G. & Dias, A. G., 1991. - Girinos do grupo “microcephala” do Estado do Rio de Janeiro

(Amphibia, Anura, Hylidae). Rev. bras. Zool., 7 (4): 679-683.

DurLLMax, WE. 1970. - The hylid frogs of Middle America. Mon. Mus. nat. Hist. Univ. Kansas, 23 (1):

1-753.

— 1972, — The systematic status and life history of Hyla rhodopepla Günther. Herpetologica, 28:

369-375.

DurLLMAN, W. E. & FOUQUETTE, M. 3. Jr., 1968. — Middle American frogs of the Hyla microcephala

group. Univ. Kansas Publ. Mus. nat. Hist., 17 (12): 517-557.

DuELLMAN, WE. & TuEB, L., 1983. - Frogs of the Hyla columbiana group: taxonomy and phylogenetic

relationships. Jr: A. G. J. RoDIN & K. Miyaïa (ed.), Advances in herpetology and evolutionary

biology, Cambridge, Mus. comp. Zool. Havard Uni

Frost, D. R. (ed.), 1985. - Amphibian species of the world.

Ain] + iv + 12732.

Goswer, K. L., 1960. — A simplified table for staging anuran embryos and larvae with notes on

identification. Herpetologica, 16: 183-190.

KENNY, LS. 1969. — The Amphibia of Trinidad. Stud. Fauna Curacao Caribb. Isl., 108: 1-78

LAVILLA, E. O., 1990. - The tadpole of Hyla nana (Anura: Hylidae). J Herp., 24 (2): 207-209.

Lurz, B., 1947. Trends towards non-aquatic and direct development in frogs. Copeia, 1947 (4): 242-2

Bre of Hyla. Austin & London, Univ. Texas Press: 1-262.

PoMBaL, JP. Jr. & BASTOS, R. P. 1998. - Nova espécie de Ayla Laurenti, 1768 do centro-oeste brasileiro

e à posiçäo taxonômica de Hyla microcephala wérneri Cochran, 1952 e Hyla microcephala

meridiana B. Lutz, 1952 (Anura, Hylidae). Bol. Mus. nac., (n.s), Zool., Rio de Janciro, 390: 1-14.

awrence, Allen Press & Assoc. Syst. Coll.:

Source : MNHN, Paris

80 ALYTES 18 (1-2)

APPENDIX 1

ADDITIONAL SPECIMENS EXAMINED

Hyla berthalutzae Bokermann, 1962. - Brazil, Rio de Janeiro State, Municipality of

Magé (22°39'S, 43°02°W). Adults: ZUFRJ 2283-2284, 2887, 3137, 5266, 5268-5269, 6613-

6619, 6621, 6624-6625, 6628. Tadpoles: ZUFRIJ 6629, 7754.

Hyla decipiens A. Lutz, 1925. - Brazil, Rio de Janeiro State, Municipality of Itaguai

(22°%4#S, 43°42°W). Adults: ZUFRJ 4463, 4465, 4471, 4475, 4500, 4502, 4506, 4516, 4591-

4592, 4594. Tadpoles: ZUFRIJ 4543.

Hyla oliveirai Bokermann, 1963. - Brazil, Bahia State, Municipality of Maracäs

(13°26'S, 40°25°W). Adult: MZUSP 74148.

Corresponding editor: Esteban O. LAVILLA.

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 81-89. 81

Euglenoids living in the intestines

of microhylid tadpoles of Argentina

Dinorah D. ECHEVERRIA * & Visitaciôn CONFORTI **

+ Laboratorio de Vertebrados

** Laboratorio de Protistologia

Departamento de Ciencias Biolégicas, Facultad de Ciencias Exactas y Naturales,

1428 Buenos Aires, Argentina

E-mail: echeverria@bg.fcen.uba.ar, conforti@bg.fcen.uba.ar

Large numbers of undamaged euglenoids (Euglena ehrenbergii var.

baculifera, E. spiroides, Lepocinclis fusiformis, L. salina, Phacus curvi-

cauda, Trachelomonas bacillifera var. minima, T. pusilla and T. volvo-

cina) were found in the intestinal contents of tadpoles of Dermatonotus

muelleri and Elachistocleis bicolor. Nine eggs of Elachistocleis bicolor

were reared in the laboratory in plastic pools until the larvae reached stage

24. Seven of these tadpoles were placed in a glass container with water and

a sample of phytoplankton rich in flagellated euglenoids (Euglena ehren-

bergii var. baculifera and E. intermedia var. klebsii). Two control tadpoles

were kept in the absence of euglenoids. Intestinal contents of the tadpoles

were observed at stages 25, 27 and 36 (sensu Gosxrr, 1960). E. intermedia

var. klebsii was found in the intestinal tract of tadpoles of E. bicolor. AIl

euglenoids were intact and had typical features of free-living cells in nature,

with large accumulations of paramylon granules. We conclude that these

euglenoids are not part of the diet of these tadpoles and that they are not

digested.

INTRODUCTION

Protozoans, algae and nematodes have been reported or sugg s food items in diets

of anuran tadpoles (HEYER, 1973: INGER, 1986). Most diet studies have been based on the

taxonomic composition of items found in the intestinal contents of fixed specimens. We have

found large numbers of euglenoids in the intestines of tadpoles of Dermatonotus muelleri and

Elachistocleis bicolor collected in lentic environments from Argentina (unpublished data).

These observations led us to study the euglenoïid fauna in the larvae of Elachistocleis bicolor

obtained from eggs developed in our laboratory and fed with freshwater euglenoids.

There are few reports on diet of microhylid tadpoles (Li & Lin, 1935; SAVAGE, 1952;

HEYER, 1973; INGER, 1986: INGER et al., 1986; WaNG et al., 1989). Almost all microhylid

tadpoles lack keratinized mouthparts and are “filter feeding tadpoles, type 2” of ALTIG &

JOHNSTON (1989). Li & Lin (1935) confirmed living euglenoids in the intestines of Kaloula

Source : MNHN, Paris

82 ALYTES 18 (1-2)

borealis and commented on the relationship between the protozoa and the tadpoles. Living

euglenoids can survive passage through the intestines (HEGNER, 1926). Li & Lin (1935)

noticed some digestion of euglenoids. Euglenamorpha and Hegneria are euglenoids that live in

the hindgut of tadpoles (BRUMPT & LAVIER, 1924; WENRICH, 1924). The purpose of this paper

is to determine whether tadpoles feed on euglenoids present in their intestines.

MATERIALS AND METHODS

Two samples of tadpoles were examined and staged according to GOsnER (1960). The

tadpoles have developed mouthparts and functional intestines in all stages used in this study.

Sample A. — Four tadpoles of Dermatonotus muelleri (stages 32, 33, 34 and 36) and two

tadpoles of Elachistocleis bicolor (stages 36 and 38) were collected from different freshwater

pools in Santiago del Estero and Misiones provinces. They were fixed in 10 % formalin.

Sample B. — Nine eggs of Elachistocleis bicolor from Corrientes province were reared in

the laboratory in a plastic pool with water from the environment where they were collected.

Water was changed several times during the next seven days until the larvae reached stage 24.

Seven of these tadpoles were placed in a glass container with 250 ml water, and 10 ml of

phytoplankton rich in euglenoids (Euglena ehrenbergii var. baculifera and Euglena intermedia

var. Klebsii) were added. The remaining two control tadpoles were reared separately in another

glass container and were fed with commercial fish food and yeast (Saccharomyces cerevisiae).

The euglenoids’ sample and the tadpoles were maintained at 18°C with a photoperiod of

12:12 LD.

The intestinal contents of all tadpoles were observed at stages 25, 27 and 36, and from 24

hours to 15 days from the beginning of the experiment. Temporary preparations from the

foregut, midgut and hindgut (including the cloaca and vent tube) were made and observed by

light microscopy. In the temporary preparations, the wall of a small part of the gut was slit

and placed on a glass slide, 1-2 drops of water were added, and a cover slip was placed over the

material. In sample À, intestinal contents and buccopharyngeal cavities were also analyzed by

scanning electron microscopy (SEM). Tadpoles preserved in 10% formalin were entire

critical-point dried, and the intestines were removed and broken over a piece of double-sided

tape placed on a microscope stub. The intestinal contents were coated with gold-palladium. A

Philips 515 scanning electron microscope, a vacuum evaporator (ION Sputtering Balzers

SCD 040) and a critical point dryer (Balzers CPD 030) were used.

In sample B, in vivo observations of the intestinal contents were made. The algae and

protists living in the pond water were identified by light microscopy. Two control tadpoles at

stage 31, living in the pond, were examined to verify that euglenoids were present in their

intestines.

The following bibliography was used for the taxonomic identification of euglenoids:

Goubics (1953), HUBER-PESTALOZZI (1955), STARMACH (1983) and TELL & CONFORTI (1986).

The eggs of E. bicolor were collected with the permission of and under the rules of the

Administraciôn de Parques Nacionales.

Source : MNHN, Paris

ECHEVERRIA & CONFORTI 83

Table 1. - Euglenoids found in the gut contents of Dermatonotus muelleri and Elachistocleis bicolor

tadpoles. Parts of intestine: 1, foregut; 2, midgut; 3, hindgut and cloaca.

Taxon PEER Observations

1 2 3:

Euglena ehrenbergii var. baculifera + undamaged

Euglena intermedia var. klebsii + + # alive

Euglena oxyuris + + + undamaged

Euglena spiroides + undamaged

Lepocinclis fusiformis + undamaged

Lepocinclis salina + + undamaged

Phacus sp. + + undamaged

Phacus curvicauda + undamaged

Trachelomonas bacillifera var. minima + undamaged

Trachelomonas pusilla + undamaged

Trachelomonas volvocina + + undamaged

RESULTS

Intact cells of Trachelomonas volvocina, T. bacillifera var. minima and T. pusilla were

found in the anterior zone of the intestines of Dermatonotus muelleri. Phacus sp., Lepocinclis

fusiformis, L. salina and Trachelomonas volvocina were found undamaged in the hind gut of

fixed tadpoles (sample A) of D. muelleri and Elachistocleis bicolor. Large numbers of

undamaged Lepocinclis salina, L. f nd Trachelomonas volvocina were found in the

cloaca. An intact cell was observed inside the lorica of Trachelomonas volvocina. Phacus

curvicauda, Euglena oxyuris, E. ehrenbergi var. baculifera, Lepocinclis salina and L. fusiformis

were found in the posterior part of the intestines of D. muelleri. AI euglenoids showed an

accumulation of paramylon granules. E ehrenbergiï var. baculifera also contained many

carotenoid granules irregularly distributed throughout the cell. SEM observations of the

intestinal contents and buccopharyngeal cavities of the tadpoles confirmed the euglenoid

taxa identified with light microscopy. Phacus sp., Trachelomonas sp. and Euglena spiroides

were observed in the roof of the buccopharyngeal cavity of Dermatonotus muelleri (fig. La-b).

Intact euglenoids were found along with other protists in the anterior and mid-zone of the

tadpole intestines. AIl these cells were undamaged (fig. Ic-d) and they had typical features of

free-living cells in nature (fig. ). Euglenoids were the only cells that were identified from

the hindgut (fig. 2d; tab. 1). Living Euglena intermedia var. klebsii and other algae (species of

Source : MNHN, Paris

84 ALYTES 18 (1-2)

vity of Dermatonotus muelleri. Scale line,

10 m. (1550 x). (b) Euglena spiroides in the buccopharyngeal cavity of D. muelleri. Scale line, 10 ym.

(1200 x). (c) SEM view of several euglenoids in the foregut of D. muelleri. Scale line, 10 ym. (50 x)

(d) Trachelomonas pusilla and several dinoflagellates (Peridinium sp.) from the midgut of D. muelleri

Scale line, 10 4m. (3100 x).

Source : MNHN, Paris

ECHEVERRIA & CONFORTI 85

Fig. 2. — (a) Phacus sp. (arrow) on Euglena oxyuris and a contracted Euglena sp. from the foregut of

D. muelleri. Scale line, 10 um. (1000 X). (b) Euglena spiroides from the foregut of D. muelleri. Scale

line, 10 um. (1000 x). (c) Lepocinclis salina from the midgut of Elachistocleis bicolor. Scale line, 10 um.

(2400 x). (d) Euglena oxyuris in the hindgut of D. muelleri. Scale line, 10 m. (1000 x).

Source : MNHN, Paris

86 ALYTES 18 (1-2)

Chlorophyceae: Scenedesmus sp. and Ankistrodesmus sp.; Dinophyceae: Peridinium sp.; and

Bacillarophyceae: Nitzschia sp.) were observed in the foregut and midgut of Elachistocleis

bicolor (sample B). Only E. intermedia var. klebsii was found in the hindgut and cloaca. No

algae were found in the cloaca other than the living euglenoids. These results were obtained 48

hours, 72 hours and 12 days after the freshwater sample was added to the containers with the

tadpoles. The euglenoids were very active, moving inside the intestine, especially in the mid

and hindgut, and they occurred in groups over the intestine walls. The intestinal contents of

one tadpole of sample B was analyzed 24 hours after placement in the glass container with the

phytoplankton. No evidence of E. ehrenbergii var. baculifera and E. intermedia var. klebsii

were found.

Intestinal contents of four control tadpoles were observed. Two tadpoles were fixed

immediately after collection (pond control) from the pond where the eggs of Elachistocleis

bicolor were obtained. Euglena spiroides, Phacus sp. and several species of Trachelomonas sp.

were observed in their intestinal contents. Two embryos were separated from the nine eggs and

kept in the absence of euglenoids (intestinal control). Ten days after the experiment began, no

euglenoids were observed in the intestinal contents of the developing tadpoles.

DISCUSSION

Although we found euglenoids in the intestines of the tadpoles of Dermatonotus muelleri

and Elachistocleis bicolor, these protists were not digested by the tadpoles of £. bicolor They

were alive and intact inside the length of the intestines. This conclusion specifically applies to

D. muelleri because all individual euglenoids examined with SEM were undamaged. These

tadpoles could be non-selective of the kind of organisms they ingest or digest, which agrees

With CAROTHERS & JAKSIC (1984). A few authors considered euglenoids to be part of the diet

of microhylid tadpoles, but the ability of a given tadpole to use euglenoids as food may be

determined by whether the tadpole has a gut laminarase that can degrade paramylon granules

(BULL & CHESTERS, 1966, fide WaLNE & Kivic, 1990). Euglenoids in £. bicolor had an excess

of paramylon granules, which suggests that the protists stored paramylon, and the intestines

did not limit nutrient availability. This agrees with CONFORTI (1998) and her results of the

study on euglenoids’ development in an environment with organic enrichment. Tadpoles are

microphagous feeders and the size of Euglena is within the range of ingested food particles,

but the pellicle and the lorica could be an impediment to digestion. Tadpoles have a non-acid

intestinal pH and a long intestine with weak peristalsis (THRALL, 1972, fide ALTIG &

JOHNSTON, 1989). Several enzymes were detected in the intestines of microhylid larvae (ALTIG

& MCDEARMAN, 1975), but euglenoids seem not to be affected by the internal gut conditions.

Euglenoids in the intestines had features typical of cells in samples from the field, including

pigmented plastids. These features are probably only possible in lighted conditions. The

ventral body wall of these tadpoles is translucent, at least in stage: to 31, and few coils of

the intestines can be seen through the body wall. We presume that sufficient light to promote

growth in the protozoa can pass through the tadpole tissues. The large size of paramylon

granules indicates that the level of light and nutrients in the intestines provides favorable

conditions for the euglenoids. Large accumulations of carotenoid granules in euglenoids were

Source : MNHN, Paris

ECHEVERRIA & CONFORTI 87

reported by BoROWITZKA (1988) as a signal of nitrogen deprivation. Euglena ehrenbergii var.

baculifera in the intestinal contents of D. muelleri had a great number of carotenoid granules

in the cytoplasm (this was not in the case in E. bicolor).

The number of euglenoids present in the last part of the intestines of D. muelleri and E.

bicolor tadpoles suggests that euglenoids could enter the intestines via tadpole’s vent tube.

The vent tube (and the cloaca) have no muscles nearby. A fecal strand usually extends outside

the body and could attract protists inside the cloaca. Euglena ehrenbergii may be able to locate

the fecal strands by chemoreception of a particular substance, for example, the nitrogenous

wastes of E. bicolor metabolism. Once inside the intestines, euglenoids could move along the

short and transparent intestines coils. In sample B of E. bicolor, Euglena erhrenbergii var.

baculifera was absent and only Euglena intermedia var. klebsii was present. The absence of E.

erhenbergii var. baculifera could be related to its size and/or its ecomorphological type. E.

erhenbergii var. baculifera (188-198 X 19.8-20 ym) is longer than E. intermedia var. klebsii

(78-90 x 7-15 um). Smaller euglenoids may be more effectively captured by the branchial

structures than larger ones, or £. erhenbergii var. baculifera may not be harvested by these

suspension-feeding tadpoles because of its benthic habitat. The results of our in vivo study

provide new information on the diet of two filter feeding tadpoles of Argentina.

CONCLUSIONS

(1) Euglenoids, along with other algae (several species of Chlorophyceae, Dinophyceae

and Bacillariophyceae) were found alive and undamaged in the intestinal contents examined

by optical microscopy.

(2) At least the observed euglenoids are not digested because they were always found

intact and alive inside the gut (anterior, posterior and cloacal portions). Euglenoid flagellates

were the only protists found living in the cloaca.

(3) The storage of paramylon granules in euglenoids suggests that the intestinal condi-

tions were favorable for these organisms, and that they encountered no nutrient limitation.

(4) The presence of live, undamaged euglenoids indicates that they are not part of the diet

of these tadpoles.

RÉSUMÉ

Dans le contenu intestinal de têtards de Dermatonotus muelleriet de Elachistocleis bicolor

provenant de divers environnements aquatiques, nous avons trouvé une grande quantité

d’euglénoïdes flagellés (Euglena ehrenbergii var. baculifera, E. spiroides, Lepocinclis fusiformis,

L. salina, Phacus curvicauda, Trachelomonas bacillifera var. minima, T. pusilla et T. volvocina).

Au laboratoire, 9 oeufs de Elachistocleis bicolor, récoltés dans des environnements naturels, se

sont développés jusqu'au stade 24 (selon GosnER, 1980). Les têtards ont été alimentés avec du

phytoplancton très riche en euglénoïdes flagellés qui contenait Euglena intermedia var. klebsiüi

Source : MNHN, Paris

88 ALYTES 18 (1-2)

et Euglena ehrenbergii var. baculifera. Après 48 h, 72 h et 12 jours, nous avons enregistré la

présence de E. intermedia var. klebsii vivante à l’intérieur de l'intestin. Etant donné que les

plastides étaient intacts et que les corps de paramylon étaient similaires à ceux qu'on trouve

dans la nature, les algues ne semblent pas affectées par le milieu intérieur de l'intestin. Ces

résultats nous permettent de conclure que les euglénoïdes étudiés ne font pas partie du régime

alimentaire des larves de ces Microhylidés et supportent sans problème apparent les condi-

tions internes de la cavité intestinale.

RESUMEN

En el contenido intestinal de renacuajos de Dermatonotus muelleri y de Elachistocleis

bicolor hallamos gran acumulacién de euglenoideos flagelados (Euglena ehrembergii var.

baculifera, E. oxyuris, E. spiroides, Lepocinclis fusiformis, L. salina, Phacus curvicauda,

Trachelomonas bacullifera var. minima, T. pusilla and T. volvocina), provenientes de distintos

cuerpos de agua. En el laboratorio, se dejaron desarrollar 9 huevos de Elachistocleis bicolor,

recogidos en ambientes naturales, hasta el estadio 24 (segün GOsNer, 1980). Los renacuajos

fueron alimentados con fitoplancton muy rico en euglenoideos flagelados que contenia

principalmente Euglena intermedia var. klebsii y E. ehrembergii var, baculifera. A las 48 horas,

72 horas y 12 dias se registré la presencia de E. intermedia var. klebsii, vivas en el interior del

intestino. Ellas no mostraron signos de ser afectadas por el medio interno del intestino ya que

presentaron los plästidos intactos y cuerpos de paramilon similares a los hallados en la

naturaleza. Nuestros resultados permiten concluir que los euglenoideos estudiados no for-

marian parte de la dieta de los renacuajos de microhylidos mencionados, y que soportan sin

perjuicio aparente las condiciones internas de la cavidad intestinal.

ACKNOWLEDGEMENTS

We thank the Director of Parques Nacionales and the staff of the Area Protegida Mburucuyä from

Corrientes for allowing us to collect the eges of E. bicolor for this study; Dr. Ronald G. Altig (Mississippi

State University) for advice and comments: and Mr. Dante Gimenez (SEM Service of CITEFA) for

technical assistance, This work was supported in part by funds provided by the BID 802/OC-AR

(PICTO0$27 to D. D. Echeverria).

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TeLz, G. & Conrorri, V., 1986. — Euglenophyta pigmentadas de la Argentina. Berlin, Biblioteca Phycolo-

gica, 75: 1-301.

THRALL, J.H., 1972. - Food, feeding and digestive physiology of the larval bullfrog, Rana catesbeïana Shaw.

PhD Dissertation, Illinois State University, Bloomington.

WALxE, P. L. & Kivic P. A., 1990. - Phylum Euglenida. /n: JONES & BARTLETT (ed.), Handbook of

Protoctista, Boston, 15: 270-287.

WanG, C. S., Wu, S. H. & Yu, W. T., 1989. - Notes on Microhyla inornata Boulenger (Anura,

Microhylidae) in Taiwan. J. Herp., 23 (4): 342-349.

Wexri, D. H., 1924. - Studies on Euglenamorpha hegneri n. g., n. sp., à euglenoïd flagellate found in

tadpol ol. Bull., 47: 149-174.

assers, Stuttgart, 16

Corresponding editor: Janalee P. CALDWELL.

Source : MNHN, Paris

90 ALYTES 18 (1-2)

Dumerilia

Publication de l'Association des Amis du Laboratoire des Reptiles et Amphibiens

du Muséum national d'Histoire naturelle de Paris

‘The name Dumerilia is a tribut 10 the memory of André-Marie-Constant DuMÉRIL (1774-1860), one of the founders of Herpetology,

senior author of the masterly treatise entitled Erpétologie générale (1834-1854)

‘The aim of Dumerilia is to publish the results of original researches dealing with all aspects of the study of Amphibia and Reptlia,

from biology to zoogeography, including systematics, evolution, conservation, or even pure nomenclatural problems.

We must here stress upon the uniqueness of the editoral policy of this journal. The comments of the referees are distributed in two

categories: objective criticisms, pointing to factual mistakes in the manuscript (incomplete or eroneous bibliographic information,

methodological errors, obscure writing, et); and subjective crticisms, the referee expressing an opinion different from or complementary to the

author's one regarding the interpretation of certain facts.

Objective eriticisms are transmited 10 the author who is asked 10 correct the manuscript in order 10 solve the problem. For lack of

correction, the paper cannot be acceptable for publication in Dumerilia.

As concems subjective criticisms, we introduced in Dumerilia a novelty: in case of acceplance of the paper, these remarks are

published, signed by their authors, in an appendix after the paper. The publication, with the consent of their authors who wil thus assume their

patemity, of these comments, offers a new possibility for specialists consulted for reviews 10 express thcir opinion, and is also a way 10 thank

them for their collaboration.

Morcover, the publication of these additional remarks, which may in some cases be in complete disagreement with the opinion of the

author of the paper, allows 10 pinpoint the most tricky or contentious questions and to stress the character still provisional of some data, analyses

or interpretations.

The following issues are presently published

Volume 1. - Monography: Patrick Davib. Liste des reptiles actuels du monde. 1. Chelonii. (French and English language).

Volume 2. - 6 papers, by 7 authors: R. BOUR; ER. BRYG00; À, DuBoïs; G. PASTEUR; J.-M. PROBST; S. RIBES; R. G, WE.

Volume 3. Monography: Patrick DAVID & Ivan INEICH. Les serpents venimeux du monde: systématique ct répartition. (French and English

language).

Volume 4, published in 3 separate fascicles. - Fascicle (1) published, 1 paper by A. DUBOIS & A. OHLER. - Fascicle (2) published, 4 papers by 13

authors: R. BOUR; U. CARAMASCHI; C. CHIMSUNCHART, P. DANGSEE; P. DaviD; W. DERECK; A. DUBOIS; C. KUMPS; O.-A. LAOHAWAT; P.

MADERSON, O, PAUWELS; C, PUANGIT; H. ZAHER. - Fascicle (3) in preparation:

Prices, including surface postage:

French Francs US. Dollars

Volume 1 260 a

Volume 2 280 56

Volume 3 360 nm

Volume 4 (complete set) 400 s0

AALRAM Secretaryÿ's office: Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle, 25, rue Cuvier, 75005

Paris, France, Tél.: (33) 1 40 79 34 87 - Fax: (33) 1 40 79 34 88 -

Payment in U. $. Dollars: by cheque payable to *AALRAM", sent 10 Dr. John B. IVERSON, Department of Biology, Earham College,

Richmond IN 473744095, U.S.A.

mail: reptamph@mnhn.fr

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 91-94. 91

A landmark publication

on the amphibians of northern Eurasia

C. Kenneth Dopp, Jr.

US. Geological Survey, Florida Caribbean Science Center,

7920 Northwest 71* Street, Gainesville, Florida 32653, USA

Sergius L. KUZMIN. — The amphibians of the former Soviet Union. Sofia, Bulgaria, Pensoft Publishers,

1999: 1-538. 598.00 (US). Hardbound. ISBN-954-642-045-X.

In the early 1990%, 1 was contacted by Kraig Adler of Cornell University to see if I would be

interested in working with a Russian colleague on a book on the status of amphibians in the territory of

the recently dissolved Soviet Union. I did not know Sergius KUZMIN at that time, and I only reluctantly

agreed in order to learn more about the amphibians of that vast territory which until recently had been

terra incognita to most western batrachologists. Our collaboration proceeded harmoniously, however,

and we eventually co-edited a series of publications on the amphibians of Russia and the countries

making up the former Soviet Union (KUZMIN et al., 1995; KUZMIN & Dopp, 1996, 1997).

During the course of our collaboration, Sergius asked me to help edit another venture, a book on the

biology of the amphibians of the former Soviet Union. Previous field guides on Soviet herpetology were

badly out-of-date. In addition, because of the language barriers and the difficulty in obtaining references,

much of the primary literature was inaccessible to scientists outside of Russia and the 14 other

present-day countries comprising the former Soviet Union. In order to fill this vacuum, he had written a

German-language book entitled Die Amphibien RuBlands und angrenzender Gebiete (1995), and he hoped

to publish an English version to further reach scientists in the West. The original English language edition

was never published, although the work was completed in the mid-1990%. Perhaps that was just as well

The present book, greatly expanded from the original version, contains a wealth of information on nearly

every aspect of the biology and life history of the 41 species now known from the Baltic Sea to the Russian

Far East, from Siberia to the high mountains and deserts of Kazakhstan. This is truly a remarkable work.

The book is organized into chapters as follows: Chapter 1, History; Chapter 2, Faunistics and

gcography: Chapter 3. Ecology: Chapter 4, Anthropogenic pressure and conservation: and Chapter 5, Keys

for species identification. Chapter 6is actually a series of detailed accounts for all41 species of amphibians

found throughout the northern Eurasian continent; this section, comprising 300 pages, makes up the

greatest part of the book. The book has 65 pages of references covering hundreds of articles. Titles in

Russian or other languages of the former Soviet Union (e.g., Georgian, Azerbaïjanian) are translated into

English. Maps of habitats, species richness and amphibian distributions (based on an atlas grid) are

followed by color plates showing habitats and species, including multiple life-history stages, a glossary of

terms, species and subject indexes, and a list of species and subspecies with their original authors.

Although the color plates are sequestered at the end of the book, black and white photos are liberally

placed throughout the main body of the text, especially in the species accounts.

Inthe Preface, KUZMIN describes the purpose of the book (1.e., a state-of-the art rendering of Soviet

batrachology in such a manner as to introduce non-Soviet scientists to the literature and scope of research

on amphibians) in order to set the stage for the information that follows. He provides line drawings and

Source : MNHN, Paris

92 ALYTES 18 (1-2)

descriptions of the morphology of both adult and larval amphibians. These serve as a guide to

morphological nomenclature and to the methods by which amphibian measurements are presented in the

text (e.g., salamanders were measured from the snout to the anterior portion of the vent). Chapter 1

outlines the history of amphibian research in Russia, beginning with folk legends and paleo-art and

extending through the great explorations of Pallas and Kessler, to the foundations of modern amphibian

studies, that is, the works of Strauch, Nikolsky and Terentjev. The chapter concludes with a summary of

modern amphibian research throughout Russia and the former Soviet Union; this section is invaluable to

western researchers as a guide to the extent of amphibian research currently underway and who is

conducting it. The history of herpetological research is usually passed over superficially in regional or

national texts as various researchers’ publications and contributions are discussed. I often wonder about

who these people were, what motivated them to chase amphibians at times of great difficulties in travel

and social upheavals. After reading Chapter 1, I wish I had the language skills to read some of the

biographies available on Russian herpetologists (e.g.. MAZURMOvICH, 1983), especially those working

during and in the decades immediately after World War Il.

Chapter 2 accords an extensive review of biogeograph al assemblages and habitat types

throughout northern Eurasia. I found this section somewhat tedious, with lists and lists of cold northern

habitats and the species that live within them. KUZMIN documents long-term changes in distribution

patterns with explanations of how species can disperse or populations can be extirpated in rather short

periods of time. He generally rejects rapid dispersion as accounting for new distribution records in favor

of an explanation involving the discovery of relictual populations. Such an explanation seems reasonable,

although later in the species accounts he notes some rather dramatic range expansions associated with

human activity, such as the expansion of Rana ridibunda in southern European Russia and Kazakhstan.

Chapter 3 summarizes general amphibian ecology, from activity cycles and reproduction to feeding

(Kuzmin’s particular specialty) and population structuring. As a semi-tropical resident, 1 found this

section interesting because of its discussion of life histories in, of necessity, extremely cold climates. The

diverse ways in which amphibians complete the vital functions of growth and reproduction in a short

period of time in a cold climate often mirror the ways they adapt to an unstable (in terms of hydroperiod

and seasonal breeding patterns) subtropical environment. Similar developmental and life history plas-

ticity sometimes occur in response to northern cold and subtropical drought, despite our differences in

temperature. For example, amphibians living in unstable or extreme environments in Russia and Florida

both respond to stress by a decrease in the length of the larval period, and salamandrids in both regions

sometimes remain neotenie, when hydroperiods permit, in order to take advantage of favorable feeding

and thermal conditions.

1 was intrigued by KUZMIN'’s assertion that “the duration of active life probably is genetically

determined to a greater extent than the total life span which includes inactive periods of hibernation”. If

this is true, would not northern populations of a species live longer than more southern populations?

Unfortunately, the lack of references to empirical data impedes examination of this hypothesis (see

below). 1 was also intrigued to learn that exceptionally large larval Rana ridibunda become sexually

mature after overwintering and prior to metamorphosis. Perhaps gonads could enlarge prior to meta

morphosis, but to say that such tadpoles are truly “’sexually mature” indicates imprecise terminology (R.

ALTIG, personal communication). In any case, endocrinologists and evolutionary biologists should be

interested in à frog that develops a mature hormonal system, in part, prior to metamorphosis and when

breeding is impossible.

In Chapter 4, KUZMIN presents information on the general status of amphibians in the former Soviet

Union. As in most of the industrialized world, habitat destruction and altération have led to localized

declines, and habitat loss has à potential for greatest impact in regions containing species of limited

distribution. In addition, collection for pets and teaching, the ubiquitous presence of pollution and toxic

chemicals, highway mortality, and the general litany of the ills of modern society have substantially

impacted many species, especially now that legal protection has eroded as à result of a lack of

enforcement due to a crumbling economic situation. Declines do not seem Lo be mysterious, as they have

been represented in North and Central America and Australia, nor, surprisingly, does disease appear to

have figured prominently in declines. Likewise, malformations do not seem widespread, although they

oceur occasionally, such as in the vicinity of Chernobyl. Because of the remoteness of much of the former

Soviet Union and the limited number of amphibian enthusiasts monitoring populations, is the lack of

Source : MNHN, Paris

Dobp 93

such reports good news or does it merely reflect sampling bias? KUZMIN provides an extensive review of

the conservation of all taxa, including their status in the former republics and their presence in nature

reserves. Most species are found in at least a few reserves, except, unfortunately, for the rare salamander

Ranodon sibiricus. The likely negative effects of extensive social upheaval on the protection and manage-

ment of the reserves, however, does not invoke cause for optimism.

Chapter 5 is a short chapter devoted to identification. Keys are given not only for mature adults, but

also for egg masses and larvae. The keys are accompanied by line drawings of high quality (by N. V.

Panteleev), although the lack of color makes distinguishing one gray tadpole from another difficult.

Color plates would have greatly enhanced the utility of the keys. Unfortunately, the larvae of some of

these species are virtually impossible to distinguish, and there is a great deal of regional color and

morphological variation. The numbering system used in the supposedly dichotomous keys is confusing

and difficult to follow: I was at a loss in the latter parts of it. Further, there are no drawings of tadpole

mouth parts, although these are sometimes used as key characters.

The species accounts are what makes the book invaluable. Nearly everything that one might want to

know is covered in depth, at least as far as research in the former Soviet Union is concerned. Each account

contains sections on synonymy, names in several languages, taxonomie notes, description, karyology,

distribution, subspecific differentiation and variability, ecology (habitats and abundance, thermobiology

and activity cycles, reproduction, development, feeding, natural enemies, parasites and diseases),

influence of anthropogenic factors, status and conservation, and references (by name and date). The

topics are covered comprehensively, and KUZMIN presents alternative points-of-view, such as when

describing taxonomic interpretation of the Rana macrocnemis complex or the enigma of Hynobiu

turkestanicus. There is a wealth of information here, information which will give Eurasian batrachologist:

a great deal to ponder in the coming decades (a subterranean Proteus in the Crimean Peninsula,

perhaps?). In the species accounts, Kuzmin has certainly fulfilled his promise to present the scope of

Soviet research on amphibians available to the English-speaking world. The sheer volume of the literature

summarized also leads to the book’s major flaw, however.

Given the extensive review of the Soviet literature presented by KUZMIN, it is unfortunate that it is

impossible to determine the source of information in the text. This is because all literature citations are

included at the back of the chapters or species accounts, rather than in the text. Thus, the readers do not

know, nor can they determine, whether they are evaluating Kuzmin's personal opinion, the results of his

research. or statements based on the empirical research or opinions of others. Fortunately, the tables and

some of the figures contain original literature citations. Still, the book would have been much more useful

had all the citations been included directly in the text so that the original papers could be connected with

the data presented.

There are a few additional problems. For example, à map of the former Soviet Union is presented on

page 466: it should have been placed earlier in the book. Although country boundaries are lightiy

outlined, there are no maps of provinces, districts, or the autonomous republics often mentioned in the

text. One not familiar with the geography and political units of Russia, especially, cannot determine

where some of the locations referred to in the text are found. The species distribution maps are not labeled

at all in terms of place. The text could have been edited better, as there are a number of words that either

aren'’t correct or don't quite make sense: e.g.. type “territories"”" for type localities (p. 2), “semi-flowing”

waters (meaning streams with only seasonal flow?), “Ecologycal” (p. 43), “fisheries” for hatcheries (p.

382), “forest rides” (?). The lack of articles (the, a, an) in the Russian language sometimes carries over into

the text, and there are a few words (e.g.. “synanthropization”) that I have never heard of! Still, for such a

large work on a complex subject, The amphibians of the former Soviet Union is à remarkably easy and

enjoyable book to read, especially when compared 10 most English language works emanating from the

former Soviet Union. When the entire book is considered, these criticisms are minor.

The amphibians of the former Soviet Union is a landmark publication in the history of the herpetol-

ogy of the former Soviet Union, an invaluable reference source, and à much needed compilation of the

biology of amphibians across northern Eurasia. It belongs in the library of everyone interested in the

biology of Eurasian amphibians, and has much to offer batrachologists throughout the world. I highly

recommend it despite the price, which is steep by North American standards, Sergei KUZMIN has provided

Source : MNHN, Paris

94 ALYTES 18 (1-2)

the foundation upon which modern studies of the amphibians of this vast territory will be based for a

long time to come. Fortunately for biologists fluent in Russian, a Russian language version was issued by

the Russian Academy of Sciences in 1999.

LITERATURE CITED

KUzMIN, S. L., 1995. — Die Amphibien Ruflands und angrenzender Gebiete. Magdeburg, Germany, Westarp

Wissenschaften: 1-274.

KuzMi, S. L. & Don, C. K., Jr. 1996. - Advances in amphibian research in the former Soviet Union. Volume 1.

Sofia, Bulgaria, Pensoft Publishers: i-vi + 1-220.

_— 1997. — Advances in amphibian research in the former Soviet Union. Volume 2. Sofia, Bulgaria, Pensoft

Publishers: i-vi + 1-176.

Kuzmix, S. L., Dop, C. K., Jr. & PiuLIK, M. M., 1995. — Amphibian populations in the Commonwealth of

Independent States: current status and declines. Moscow, Russia, Pensoft Pulishers: 1-159.

MAZURMOVICH, B. N., 1983. — Alexander Mikhailovich Nikolskii. Moscow, Russia, Nauka: 1-75.

Corresponding editor: Annemarie OHLER.

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 95-96. 95

A synthesis

of our knowledge about tadpole biology

Stéphane GROSIEAN

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

Roy W. McDiarib & Ronald ALTIG (ed). — Tadpoles: the biology of anuran larvae. Chicago & London,

The University of Chicago Press, 1999: i-xiv + 1-444, 115 fig., 27 tables. ISBN 0-226-55762-6.

For a long time, the majority of studies in batrachology were focused on adult morphology or

anatomy. Since the last mid-century the increase of our overall knowledge led the researchers to focus

their investigations on new research fields or disciplines such as the study of advertisement calls of adult

frogs, the study of tadpoles or of ecology and interactions of different species in a population or

ecosystem. The numerous studies about tadpoles published during the last decades revealed an extraor-

dinary diversity of forms, modes of development and adaptations to various habitats. AI the data

published until now, scattered in various periodicals, needed to be compiled in a single book providing a

synthesis of available information concerning the biology of anuran larvae. Such a reference textbook

should be useful both to the novice and to the confirmed professional. This aim was largely reached by the

present book.

This collective book includes a general introduction dealing with the significance of tadpoles in the

research world; a glossary which provides accurate redefinition for terms that were more or less currently

employed, but whose meaning depended on the author or remained sometimes obscure; an abundant

bibliography: an author index, a subject index and à taxonomic index which allow to find easily any theme

treated in the book.

Between the introduction and the glossary, the multiple aspects of tadpole biology are divided in

eleven chapters. All chapters (including the introduction) have a summary and some provide an exhaus-

tive list of species with the reference to original publications relative to any precise topic. The first one is

devoted to the materials and techniques, and review all the herpetologist needs for studying tadpoles,

from the collect of these remarkable vertebrates to the scientific material necessary to study them through

fixation methods, stage and ecomorphological determination. The four following chapters are relative to

general organization of tadpoles. The first of them is devoted 10 the external morphology of tadpole

with an emphasis on the oral apparatus morphology and a discussion about functional and evolutionary

aspects of tadpole morphology. The last three chapter deal with tadpoles’ internal organisation and

gather together rare and scattered data about cranial and axial musculoskeleton, viscera and endocrines,

and nervous and sensory systems.

aper is devoted to all cases of endotrophy in anurans (six among the 21 developmental

guilds defined by ALTIG & JOHNSTON, 1989), certainly the most peculiar and captivating evolutionary

mechanism in Anura.

The last part of this volume deals with the relations of anuran larvae with their environment

throughout four chapters. The aspects of physiology of tadpoles treated are those relevant to their

ecology, such as respiration, thermal relations, and ion and water balance, Two closely related chapters

Source : MNHN, Paris

96 ALYTES 18 (1-2)

gather information about intra- and interspecific relations, social behavior, the repartition of species in

the multiple microhabitats available in a biotope, resource use and predation.

The problem of the maintenance and evolution of complex life cycles in anurans first developed by

WassERSUG (1974, 1975) is then discussed in the last but one chapter.

The twelfth and last chapter intends to show the diversity of anuran larvae and gives the most

obvious characteristics (geographic range, ecomorphological guild and gross morphological features of

oral disk, vent, spiracle, colour and pattern, snout-vent length, and finally authors of the information) for

each family and each genus. An illustration is given for certain genera. This overall panorama is

completed by a key which helps to find the identification of an unknow tadpole at the family or subfamily

level.

The compilation work done by the different authors is considerable, the presentation is very

meticulous and illustrations are of excellent quality. Inevitably some lacunae appear when attempts are

made for providing exhaustive lists as for instance in tables 12.2 and 12.3 (chapter 12: 297-298), where a

list of misidentified and undetermined tadpoles is given. However this kind of weakness is amply

excusable in face of the amount of data to gathered. The systematics adopted by the authors in the

chapter dealing with diversity mixes recent classificatory schemes (as for example recognizing the family

Megophryidae) with older classifications (as the family Rhacophoridae and its three subfamilies Buer-

geriinae, Rhacophorinae and Mantellinae). I regret also the scarcity of data about the first stages of

ontogeny. The only real criticism that I have do to this book is the lack, in the literature surveyed and cited,

of representatives of Asiatic species as well as the omission of several important Asiatic authors and of

some of their very useful contributions to the knowledge of the Asiatic herpetology.

In conclusion, this book is very impressive by the amount of data gathered and by the work of the

authors who succeeded in treating most aspects of larval anuran biology with clarity and pedagogy. This

book establishs itself as a major work for all batrachologists and must have its place in your library beside

the henceforth essential DUELLMAN & TRUEB (1985).

LITERATURE CITED

ALTIG, R. & JOHNSTON, G. F., 1989. — Guilds of anuran larvae: relationships among developmental

modes, morphologies, and habitats. Herp. Mon., 3: 81-109

ELLMAN, W. E. & TRUEB, L., 1985. Biology of amphibians. New York, McGraw-Hill, “1986:

1-670.

WASsERSUG, R., 1974. — Evolution of anuran life cycles. Science, 185: 377-378.

1975. - The adaptative significance of the tadpole stage with comments on the maintenance of

complex life cycles in anurans. Am. Zool., 15: 405-417.

Corresponding editor: Annemarie OHLER.

BIBL. Di

MUSEUM:

Source : MNHN, Paris

ATES

International Journal of Batrachology

published by ISSCA

EDITORIAL BOARD

Chief Editor: Alain Dusos (Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle, 25

rue Cuvier, 75005 Paris, France).

Deputy Editor: Thierry Loi (Laboratoire d'Ecologie animale, Université d'Angers, 2 boulevard Lavoisier,

49045 Angers Cedex, France):

Editorial Board: Jean-Louis ALBARET (Paris, France); Franco ANDRFONE (Torino, Italy): Lauren E. BROWN

(Normal, USA); Janalee P. CALDWELL (Norman, USA): Ulisses CARAMASCHI (Rio de Janeiro, Brazil):

Günter GOLLMANN (Wien, Austria); Heinz GiLLITsCH (Wien, Austria); Tim HALLIDAY (Milton Keynes.

United Kingdom); W. Ronald HEvER (Washington, USA); Esteban O. LAvILLA (Tucumän. Argentina):

Karen R. Lies (Canton, USA); Masafumi MaTsut (Kyoto, Japan): Alain PAGANO (Angers, France) : John

C. PoyNTON (London, England) : Miguel VENCES (Paris, France).

Technical Editorial Team (Paris, France): Alain Dusois (texts): Roger BouR (tables): Annemarie OHLER (figures).

Book Review Editor: Annemarie OHLER (Paris, France)

SHORT GUIDE FOR AUTHORS

(for more detailed Instructions 10 Authors, see Alvtes, 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 béhaviours, etc.

The title should be followed by the name(s) and addresses) 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,

acknowiedgements, 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 reduction. The 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 follos

BoURRET, R.. 1942. - Les batraciens de l'Indochine. Hanoï, Institut Océanographique de l'Indochine: i-x + 1-547,

pl. 1-4.

Graf, J-D. & PoLLs PELAZ, 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.

INGER, RE, Voris, H. K. & Voris, H. H. 1974, - Genetic variation and population ecology of some Southeast

Asian frogs of the genera Bufo and Rana. Biochem. Gener., 12: 121-145.

Manuscripts should be submitted in triplicate cither 10 Dunois (address above) if dealing with

amphibian morphology, systematies, biogeography, evolution, genetics, anomalies. pathology or developmental

biology, or to Thierry Lobé (address above) if dealing with amphibian population genetics, ecology, ethology,

life history, conservation biology or declining populations. Acceptance for publication will be decided by the

editors following review by at least two referees.

IF possible, after acceptance, a copy of the final manuscript on a floppy disk (3 should be sent 10

the Chief Editor. We welcome the following formats of text processing: (1) pret Fable AS Word 1060 DOS

or Windows), WordPerfect (4.1 to 5.1, DOS or Windows) or WordStar (3.3 10 7.0): (2) less preferably. formated

DOS (ASCII) or DOS-formated MS Word for the Losh (on à 3 24 high density 1.44 Mo floppy disk 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 oflered 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 Natur France)

Directeur de la Publication: Alain Dusois.

Numéro de Commission Paritaire: 64851

Source : MNHN, Paris

Alytes, 2000, 18 (1-2): 1-96.

Contents

J.-L. AMIET & F. DOWSETT-LEMAIRE

Un nouveau Leptodactylodon

de la Dorsale camerounaise (Amphibia, Anura) ....................... 1-14

Alain Dugois & Annemarie OHLER

Systematics of Fejervarya limnocharis (Gravenhorst, 1829)

(Amphibia, Anura, Ranidae) and related species.

1. Nomenclatural status and type-specimens of the

nominal species Rana limnocharis Gravenhorst, 1829 . 15-50

Günter CLEMEN & Hartmut GREVEN

Dentigerous bones and dentition in the paedomorphic

plethodontid salamander Eurycea neotenes ............................ 51-61

Maria Susana PILLADO, Cecilia A. ALONSO & Carmen A. ÜBEDA

La larva de A/sodes gargola Gallardo, 1970

(Leptodactylidae, Telmatobiinae) 62-72

Adriana PUGLIESE, Ana Claudia Reis ALVES & Sergio Potsch DE CARVALHO E SILVA

The tadpoles of Hyla oliveirai and Hyla decipiens

with notes on the Hyla microcephala group (Anura, Hylidae) ........... 73-80

Dinorah D. ECHEVERRÉA & Visitaciôn CONFORTI

Euglenoids living in the intestines of microhylid tapoles of Argentina ... 81-89

Announcement

Dumerilia 90

Book reviews

C. Kenneth Dobp, Jr.

A landmark pulication on the amphibians of northern Eurasia ......... 91-94

Stéphane GROSIEAN

A synthesis of our knowledge about tadpole biology ................... 95-96

Alytes is printed on acid-free paper.

Alytes is indexed in Biosis, Cambridge Scientific Abstracts, Current Awareness in Biological

Sciences, Pascal, Referativny Zhurnal and The Zoological Record.

Imprimerie F. Paillart, Abbeville, France.

Dépôt légal: 4° trimestre 2000.

Source : MNHN, Paris