SOCIÉTÉ BATRACHOLOGIQUE DE FRANCE

(Société pour l’Étude et la Protection des Amphibiens)

SIÈGE SOCIAL

Laboratoire des Reptiles et Amphibiens, Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France.

CONSEIL D'ADMINISTRATION POUR 1987

Président : Jean-Jacques MORÈRE.

Vice-Président : Jean-Louis AMIET.

Secrétaire général (renseignements et demandes d'adhésion) : Alain DUBOIS.

Trésorière : Dominique PAYEN,

Membres : Alain CoLLENoT, Michel DELAUGERRE, Edouard LEMÉE, Luck MARTIN-BOUYER, Manuel PoLLS

PELAZ et Jean-Paul RiscH.

ADHÉSION

La S.B.F. est ouverte à toute personne française ou étrangère intéressée par l'étude et la protection des Am-

phibiens ; écrire au Secrétaire général. La cotisation inclut le service du Bulletin d'information Circalyes.

TARIFS 1987

Lieu de résidence

France Etranger

Membres de la S.B.F. :

Cotisation seule .. 100 F 100 F

Cotisation + abonnement à Alyes 170 F 180 F

Cotisation membre associé (conjoint, et 40F 40F

Abonnement à Ayes pour les non-membre:

Individus 100 F 130 F

200 F 260 F

Supplément pour expédition d'Alytes par avion (membres et non-membres) … 5 50F

Achats au numéro et rachats d'anciennes séries d’Alyres : écrire au Sécrétaire général pour information.

MODALITÉS DE RÈGLEMENT

FRANCE. - Par chèque postal ou bancaire à l’ordre de “Société Batrachologique de France”, adressé à notre

Trésorière, où par virement postal sur notre C.C.P. : “Société Batrachologique de France”, C.C.P.

7976 90 K, Paris.

EUROPE. - Exclusivement par virement postal ou mandat postal, libellé en Francs Français et adressé à

notre Compte Chèques Postal : “Société Batrachologique de France”, C.C.P. 7976 90 K, Paris.

OUTSIDE EUROPE. - Please write to our General Secretary for information.

Source : MNHN, Paris

AIRTTES

INTERNATIONAL JOURNAL OF BATRACHOLOGY

Trimestriel Volume 6

Mars-Juin 1987 Fascicules 18&2

Alytes, 1987, 6 (1-2) : 1-9. 1

Miscellanea taxinomica batrachologica (II)

Alain Dugois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The Oriental frog genus Microhyla is shown to be heterogeneous. This genus is

here divided into two subgenera, one of which includes two distinct species groups, and

à new genus is erected for M. inornata and related forms.

Ordre ANURA Rafinesque, 1815 (Hogg, 1839)

Sous-ordre RANOIDEI Wilbrand, 1814 (Sokol, 1977)

Superfamille RANOIDEA Rafinesque-Schmaltz, 1814 (Gray, 1825)

(Fitzinger, 1826) (Bolkay, 1919)

Famille MICROHYLIDAE (Fitzinger, 1843) Günther, 1858

(Noble, 1931) (Parker, 1934)

Sous-famille MICROHYLINAE (Fitzinger, 1843) Günther, 1858

(Noble, 1931)

LAURENT (1986) a suggéré de séparer en deux sous-familles, l’une orientale (Micro-

hylinae) et l’autre américaine (Gastrophryninae), la sous-famille des Microhylinae de PARKER

(1934). En attendant une analyse phylogénétique plus approfondie de ces groupes, nous le

suivons partiellement ici, en accordant à ces taxons le rang de tribus, et les noms respectifs

de Microhylini et de Gastrophrynini.

Bibliothèque ii Hus

3001 001115 LI INHN, Paris

2 ALYTES 6 (1-2)

Tribu MICROHYLINI (Fitzinger, 1843) Günther, 1858

(Noble, 1931) (emend. nov.)

Genre MicROHYLA Tschudi, 1838

Espèce-type. - Microhyla achatina Tschudi, 1838, par monotypie.

Discussion. — Dans sa révision des Microhylinae, PARKER (1934) incluait dans le genre Mi-

crohyla 15 espèces orientales et 11 espèces américaines. CARVALHO (1954 : 12-13) a suggéré

que ce groupe était un rassemblement artificiel d’espèces issues de différentes lignées évo-

lutives de Microhylinae dans lesquelles les clavicules et les procoracoïdes ont été perdus, et

cet auteur a retiré les espèces américaines du genre Microhyla. Même restreint aux espèces

orientales, ce genre est encore hétérogène, en termes morphologiques, comme l’ont déjà sou-

ligné plusieurs auteurs (PARKER, 1928 ; CARVALHO, 1954 : 13 ; LAURENT, 1986 : 753-754).

Ayant eu l’occasion, de 1970 à 1984 au Népal et en Inde, et en 1986 en Thaïlande, d’obser-

ver en vie diverses espèces rapportées à ce genre (M. annectens, M. berdmorei, M. butleri, M.

heymonsi, M. ornata, M. pulchra et M. inornata), nous avons été frappé par les différences

d’habitus, de comportement et de chant entre M. inornata d’une part et les autres espèces

de l’autre. Ces différences viennent s’ajouter à celles déjà mises en évidence par les auteurs

cités ci-dessus concernant la morphologie, et nous amènent à placer M. inornata dans un genre

distinct de Microhyla. De plus, nous proposons de diviser ce dernier genre en deux sous-

genres, dont un comportant lui-même deux groupes d’espèces distincts. On trouvera ci-des-

sous de brèves diagnoses préliminaires de ces nouveaux taxons, qui feront ultérieurement

lobjet de travaux plus approfondis.

En ce qui concerne l’espèce malgache décrite par GUIBÉ (1974 : 1188) sous le nom de

Microhyla palmata, nous partageons l'avis de LAURENT (1986 : 744) selon lequel elle appar-

tient à un autre genre et même sans doute à une autre sous-famille. Lors d’une visite récente

au Muséum de Paris, Rose M.A. BLOMMERS-SCHLÔSSER nous a informé de son intention de

nommer ce genre, qui ne sera donc pas discuté ici.

Diagnose. - CARVALHO (1954), suivi par NELSON (1972) et par la totalité des auteurs depuis,

a retiré de Microhyla toutes les espèces américaines maintenues par PARKER (1934) dans ce

genre, pour les placer dans plusieurs genres distincts, pour la plupart dans Gastrophryne.

NELSON (1972) a donné une liste des différences entre Microhyla et Gastrophryne, sur laquelle

il est inutile de revenir. Quant aux différences entre Microhyla et le nouveau genre décrit ci-

dessous, elles sont mentionnées dans la diagnose de ce dernier.

Deux sous-genres peuvent être reconnus au sein du genre Microhyla dans sa nouvelle

acception : on trouvera leurs synonymies, diagnoses et contenus ci-dessous.

DuMÉRIL & BIBRON (1841 : 613) ont créé le nom de remplacement Micrhyla, qu’ils

ont francisé en “Micrhyle”, pour ce genre. Nous adoptons ici ce dernier nom, allégé en “Mi-

cryle”, pour désigner ces animaux en français.

Sous-genre MICROHYLA Tschudi, 1838

Microhyla Tschudi, 1838 : 28. — Espèce-type par monotypie : Microhyla achatina Tschudi, 1838 : 71.

Micrhyla Duméril & Bibron, 1841 : 613. - Nomen novum pro Microhyla Tschudi, 1838.

Dendromanes Gistel, 1848 : xi. - Nomen novum pro Microhyla Tschudi, 1838.

Source : MNHN, Paris

Dusois 3

Diagnose. - Ce sous-genre se distingue du suivant par la présence à l’extrémité des doigts et

orteils de disques différenciés, habituellement divisés par un sillon longitudinal médian sur

leur face supérieure ; les phalanges terminales (au moins des orteils) sont en forme de T ; la

palmure est généralement assez importante.

On peut encore reconnaître, à la suite de PARKER (1928, 1934), deux ensembles dif-

férents au sein de ce sous-genre, auxquels on peut attribuer la rang de groupes d’espèces.

Dans le groupe berdmorei, le plus primitif, les palatins sont présents, les disques digitaux

généralement bien développés et la palmure moyenne à importante. Dans le groupe acha-

tina, probablement issu du précédent, les palatins sont absents, et le cartilage de la partie

postérieure de la capsule nasale est partiellement ossifié ; les disques digitaux sont petits et

la palmure réduite ; de plus, les deux têtards connus de ce groupe, ceux de M. achatina et

de M. heymonsi (SMITH, 1916 ; PARKER, 1928, 1934 ; POPE, 1931), se distinguent de tous les

autres têtards connus de Microhyla par leur bouche en entonnoir. On trouvera ci-dessous une

liste des espèces que nous reconnaissons pour l'instant au sein de ces deux groupes d’es-

pèces.

Espèces incluses. — (1) Groupe de Microhyla (Microhyla) berdmorei (Blyth, 1855) : Microhyla

(Müicrohyla) annamensis Smith, 1923 ; Microhyla (Microhyla) annectens Boulenger, 1900 ; Mi-

crohyla (Microhyla) berdmorei (Blyth, 1855) ; Microhyla (Microhyla) borneensis Parker, 1926 ;

Microhyla (Microhyla) butleri Boulenger, 1900 ; Microhyla (Microhyla) fowleri Taylor, 1934 ;

Microhyla (Microhyla) mixtura Liu & Hu, 1966 ; Microhyla (Microhyla) palmipes Boulenger,

1897 ; Microhyla (Microhyla) perparva Inger & Frogner, 1979 ; Microhyla (Microhyla) petri-

gena Inger & Frogner, 1979 ; Microhyla (Microhyla) superciliaris Parker, 1928.

(2) Groupe de Microhyla (Microhyla) achatina Tschudi, 1838 : Microhyla (Microhyla)

achatina Tschudi, 1838 ; Microhyla (Microhyla) chakrapanü Pillai, 1977 ; Microhyla (Micro-

hyla) fusca Andersson, 1942 ; Microhyla (Microhyla) heymonsi Vogt, 1911 ; Microhyla (Mi-

crohyla) zeylanica Parker & Hill, 1948.

Sous-genre DiPLOPELMA Günther, 1859

Siphneus Fitzinger, 1843 : 33 (nec Brants, 1827 : 19). - Espèce-type par désignation originale : Engys-

toma ornatum Duméril & Bibron, 1841 : 745.

Diplopelma Günther, 1859 : 50. - Nomen novum pro Siphneus Fitzinger, 1843.

“Scaptophryne” Fitzinger, 1860 : 416, nomen nudum. — Espèce-type par monotypie : “Scaptophryne la-

byrinthica” Fitzinger, 1860 : 416, nomen nudum.

Copea Steindachner, 1864 : 286. - Espèce-type par monotypie : Copea fulva Steindachner, 1864 : 286.

Ranina David, 1871 : 76 (nec Lamarck, 1801 : 156). — Espèce-type par monotypie : Ranina symetrica'

David, 1871 : 7

Diagnose. - Ce sous-genre se distingue du précédent par l'absence de disques à l’extrémité

des doigts et orteils ; les phalanges terminales sont simples ; la palmure est généralement

réduite.

1. Davin (1871 : 76, 77 ; 1872 : 85) a toujours écrit ce nom “symerrica”, et non pas “symmerica”, comme l'ont écrit

la plupart des auteurs ultérieurs, à la suite de BOULENGER (1882 : 166).

Source : MNHN, Paris

4 ALYTES 6 (1-2)

C’est probablement du groupe achatina du sous-genre Microhyla que le présent sous-

genre est issu : comme chez celui-ci, le palatin est absent, et le cartilage de la partie posté-

rieure de la capsule nasale est partiellement ossifié (PARKER, 1928) ; toutefois les têtards de

Diplopelma sont dépourvus de la bouche spécialisée en entonnoir que possèdent les têtards

connus du groupe achatina (PARKER, 1934).

Espèces incluses. — Microhyla (Diplopelma) okinaventris Stejneger, 1901 ; Microhyla (Diplo-

pelma) ornata (Duméril & Bibron, 1841) ; Microhyla (Diplopelma) picta Schenkel, 1901 ; Mi-

crohyla (Diplopelma) pulchra (Hallowell, 1860) ; Microhyla (Diplopelma) rubra (Jerdon, 1853).

Genre MICRYLETTA nov.

Espèce-type. - Microhyla inornata Boulenger, 1890 : 37.

Espèces incluses. - Micryletta inornata (Boulenger, 1890), avec les sous-espèces Micryletta inor-

nata inornata (Boulenger, 1890) et Micryletta inornata lineata (Taylor, 1962) ; Müicryletta

steinegeri? (Boulenger, 1909).

Etymologie du nom générique. - Ce nom rappelle celui du genre voisin Microhyla ; la désinence

-etta est destinée à évoquer le comportement sautillant de ces animaux.

Diagnose. - Les Micrylettes (Micryletta) se distinguent des Micryles (Microhyla) par un grand

nombre de caractères, notamment les suivants.

() Chez Micryletta, le palatin est absent, et l’ethmoïde est très développé, sa zone an-

térieure bordant l’arrière et les côtés du choane interne (PARKER, 1928). Chez Microhyla, le

palatin est présent (groupe berdmorei) ou absent (groupe achatina, sous-genre Diplopelma) ;

Fig. 1. — Têtes de profil. (A) Microhyla heymonsi, mâle adulte, MNHN 1987.2197, entre Nam Tok et

Sai Yok, Kanchanaburi, Thaïlande. (B) Micryletta inornata, mâle adulte, MNHN 1987.2442, Khao

Chong, Trang, Thaïlande.

2. BOULENGER (1909 : 494) a écrit le nom de cette espèce “steinegen”, ce qui signifie qu’il avait latinisé le nom de

STEJNEGER en STEINEGERUS, comme il en avait le droit le plus absolu. Le nom “stejnegeri” employé par la plupart

des auteurs ultérieurs est une émendation injustifiée du nom original, dont l'auteur est NIFDEN (1926 : 28, 35).

Source : MNHN, Paris

Dugois 5

Fig. 2. — Têtes en vue dorsale. Mêmes spécimens que dans la fig. 1.

lethmoïde est toujours séparé du choane interne par le palatin (quand il existe) ou par une

zone de cartilage partiellement ossifié, et il ne s’étend pas latéralement par rapport au choane

interne (PARKER, 1928).

(2) La forme de la tête est différente dans les deux genres. Le museau est plus long

que l’oeil et nettement proéminent chez les Micryles, tandis qu’il est plus court que l’oeil et

peu proéminent chez les Micrylettes (fig. 1). En vue dorsale, la tête des Micryles s’élargit

régulièrement depuis l'extrémité du museau jusqu’en arrière des yeux, alors que chez Mi-

cryletta la tête cesse de s’élargir au niveau des yeux (fig. 2). La région loréale est plus ou

moins oblique chez les Micryles, et quasi-verticale chez Micryletta.

(3) Le tympan est nettement visible chez Micryletta, alors qu’il est caché sous la peau

chez Microhyla (fig. 1).

(4) La main a une forme différente dans les deux genres. Chez Micryletta, le premier

doigt n’est que légèrement plus court que le second, qui n’est lui-même que modérément

plus court que le troisième (fig. 3). Chez les Micryles, le troisième doigt est très long, et le

premier est bien plus court que le second (fig. 3) ; la tendance au raccourcissement du pre-

mier doigt va même très loin chez deux espèces de Bornéo du groupe berdmorei, M. perparva

et M. petrigena, où ce doigt est quasiment absent (INGER & FROGNER, 1979).

(5) La paume de la main des Micrylettes est couverte de nombreux tubercules sem-

blables aux tubercules sous-articulaires des doigts et aux tubercules palmaires habituels des

Anoures (fig. 3) : des tubercules similaires ont été décrits chez divers Ranidae orientaux par

INGER (1954, 1966), qui estime qu’ils doivent faciliter le comportement grimpeur de ces es-

pèces. Des tubercules surnuméraires semblables sont également parfois visibles, quoique

moins nets et développés, sur le métatarsien de certains orteils chez certains spécimens de

Micryletta. Chez les Micryles en revanche, de telles structures sont généralement absentes

sur les mains et les pieds, parfois seulement esquissées sur les mains (fig. 3).

(6) Les extrémités des doigts et orteils des Micrylettes sont dépourvues de disques, et

les phalanges terminales simples (PARKER, 1928), comme dans le sous-genre Diplopelma, alors

que dans le sous-genre Microhyla s. str. des ventouses et des phalanges terminales en T sont

présentes.

Source : MNHN, Paris

6 ALYTES 6 (1-2)

Fig. 3. — Mains en vue ventrale. (A) Microhyla butleri, mâle adulte, MNHN 19872284, Khao Chong,

Trang, Thaïlande. (B) Microhvla heymonsi, mâle adulte, MNHN 1987.2197, entre Nam Tok et Sai

Yok, Kanchanaburi, Thaïlande. (C) Microhvla berdmorei, femelle juvénile, MNHN 1987.2246, Khao

Yai, Nakhon Nayok, Thaïlande. (D) Micrylerta inornata, mâle adulte, MNHN 1987.2442, Khao

Chong, Trang, Thailande.

(7) La palmure est totalement absente chez les Micrylettes, alors qu’elle est toujours

présente, quoique parfois réduite à un rudiment de membrane, chez les Micryles.

(8) Les pattes postérieures sont plus longues chez les Micryles que chez les Micry-

lettes.

(9) Le pattern dorsal de toutes les espèces de Microhyla suit un même “modèle”, avec

un dessin médio-dorsal symétrique plus ou moins développé et net, mais toujours recon-

naissable, et parfois une ligne médio-dorsale fine. Ce dessin n’est pas reconnaissable, même

“déformé”, chez les Micrylettes : en revanche chez ces dernières il existe sur le dos des lignes

ou bandes foncées longitudinales plus où moins complètes, qui n’ont pas d’équivalent chez

les Micryles.

Source : MNHN, Paris

Dugois q

(10) En vie, le ventre des Micryles est entièrement pigmenté, blanc ou jaune nacré,

habituellement sans taches (sauf sur les bords), tandis que celui des Micrylettes est totale-

ment ou partiellement (petites taches sur fond blanc nacré) translucide rosâtre, les viscères

apparaissant à travers ces zones de peau non pigmentée.

(11) Les animaux des deux groupes ont non seulement des habitus différents, qui

frappent dès le premier abord sur le terrain, mais également des comportements fort dissem-

blables. Le déplacement habituel des Micryles se fait par petits bonds, alors que les Micry-

lettes marchent plutôt, à la manière de certains Bufo. Lorsque ces animaux sont inquiétés,

les Micryles fuient généralement à terre à l’aide d’un très grand bond (parfois de plus d’un

mètre, chez les grandes espèces comme M. berdmorei où M. pulchra), à l’arrivée duquel l’a-

nimal se tient totalement immobile sur le sol, où il est souvent impossible de le retrouver et

de le distinguer de la terre et de la végétation, tant il est bien camouflé par sa coloration ;

dans la même situation en revanche, les Micrylettes s’enfuient généralement par une suc-

cession très rapide de petits sauts non séparés par des haltes, et se dirigent souvent vers une

mare où elles plongent et disparaissent sous l’eau.

(12) Le poste de chant des Micryles est varié : souvent simplement à même le sol, gé-

néralement dans une zone herbeuse, parfois assez loin de l’eau ; plus rarement perché dans

la végétation riveraine ou aquatique, ou même parfois flottant dans l’eau (LIU, 1950 : 251).

Les Micrylettes chantent plus rarement à terre, mais plus souvent perchées, parfois très haut,

dans la végétation du bord de l’eau ou, préférentiellement semble-t-il, dans la végétation

aquatique dont les tiges ou branches émergent de l’eau et surplombent celle-ci. Les chants

des Micryles sont variés, mais consistent tous en une répétition plus ou moins rapide de notes

généralement graves : les bandes de fréquences dominantes des chants de six espèces de Mi-

crohyla décrits par HEYER (1971), NELSON (1973) et DuBois (1977) se situent entre 1200 et

4500 Hz. En revanche, le chant de Micrylerta inornata a un “aspect” très différent : il se pré-

sente à l'oreille humaine comme une sorte de sifflement aigu, que HEYER (1971 : 67) compare

à un chant de grillon. HEYER (1971) a présenté une analyse de ce chant, dont la bande de

fréquence dominante se situe entre 4400 et 6500 Hz.

Discussion. - Comme cela a été souvent souligné (p. ex. LAURENT, 1987 : 2), l’atrophie ou

la perte d’une structure ne constitue pas un bon critère de proche parenté phylogénétique

entre espèces différentes, puisqu'elle peut s'être produite indépendamment dans des lignées

distinctes. Le genre Microhyla tel que le concevait PARKER (1934) reposait principalement

sur l'absence des clavicules, des procoracoïdes et de l’omosternum. Comme l’ont souligné

CARVALHO (1954) et LAURENT (1986), la perte de ces os a pu avoir lieu à plusieurs reprises

et dans différentes lignées de Microhylinae. Les caractères (morphologie, comportement,

chant) de Micryletta sont fondamentalement différents de ceux de Microhyla et nous pensons

que les deux genres sont probablement d’origine phylogénétique distincte au sein de la sous-

famille, contrairement à ce qu’estimait PARKER (1928), pour qui M. inornata représentait l’a-

boutissement d’une lignée dont les espèces que nous plaçons dans le groupe de M. berdmorei

seraient les représentants actuels les plus primitifs. Une analvse cladistique de la sous-famille

sera nécessaire afin de préciser les affinités phylogénétiques des deux genres.

La nature distincte des deux genres est particulièrement frappante pour qui a l’occa-

sion de les observer en vie dans la nature, et c’est peut être ce qui explique que GRESSITT

(1938) ait ressenti le besoin de placer le spécimen unique de Micrylette qu’il avait récolté à

Source : MNHN, Paris

8 ALYTES 6 (1-2)

Formose dans un autre genre que Microhyla. Il rapporta ce spécimen au genre Rana, et le

décrivit comme espèce nouvelle sous le nom de Rana gracilipes. MATSUI & BUSACK (1985)

ont récemment placé ce nom dans la synonymie de M. inornata. Pour notre part, nous sui-

vons ici provisoirement BOULENGER (1909) pour reconnaître la forme de Taiwan comme une

espèce distincte Micrylerta steinegeri, nom dont Rana gracilipes est synonyme. Nous adoptons

également pour l'instant, en attendant une étude plus approfondie de la variation géogra-

phique chez Micryletta inornata, les deux sous-espèces inornata et lineata reconnues par Tay-

LOR (1962) au sein de celle-ci.

REMERCIEMENTS

Nous avons plaisir à remercier M. Jarujin NABHITABHATA pour l’aide qu’il nous a apportée en

Thaïlande, notamment en nous donnant la possibilité d’étudier les Amphibiens dans plusieurs parcs

nationaux. Le travail sur le terrain dans ce pays a bénéficié de la collaboration de Mile Annemarie OH-

LER et de MM Jarujin NABHITABHATA et Jean-Paul RiscH. Les dessins qui illustrent ce travail ont été

réalisés par Mlle Annemarie OHLER.

RÉFÉRENCES BIBLIOGRAPHIQUES

BOULENGER, G. A., 1882. - Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British

Museum. London, Taylor & Francis : i-xvi + 1-503, pl. I-XXX.

ES 1890. - List of the Reptiles, Batrachiai nd freshwater Fishes collected by Professor Moesch and

Mr. Iversen in the district of Deli, Sumatra. Proc. zool. Soc. Lond., 1890 : 31-40.

ee 1909. — Descriptions of four new frogs and a new snake discovered by Mr. H. Sauter in Formosa.

Ann. Mag. nat. Hist., (8), 4 : 492-495.

BRANTS, À., 1827. — Het geslacht der muizen door Linnaeus opgesteld, volgens de tegenswvoordige toestand

der wetenschap in familien, geslachten en soorten. Berlyn, Akademische Bockdrukkery : i-xii + 1-

190 + ii, 1 pl.

CARVALHO, À. L. DE, 1954. — À preliminary synopsis of the genera of American microhylid frogs. Occ.

Pap. Mus. Zool. Univ. Michigan, 555 : 1-19, pl. I.

Davip, A., 1871. - Rapport adressé à MM les Professeurs-Administrateurs du Muséum d'Histoire na-

turelle. Bull. Nouv. Arch. Mus., 7 : 75-100.

= 1872. — Journal d’un voyage dans le centre de la Chine et dans le Thibet oriental. Bull. Nouv.

Arch. Mus., 8 : 3-128.

Dugois, A., 1977. - Chants et écologie chez les Amphibiens du Népal. /n : Himalaya. Ecologie-Eth-

nologie, Paris, CNRS, Colloques internationaux du CNRS, 268 : 109-118.

DuMéÉRIL, À. M. C. & BIBRON, G., 1841. — Erpétologie générale ou histoire naturelle complète des Reptiles.

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

FITZINGER, F. L., 1843. - Systema Reptilium. Fasc. 1. Amblyglossae. Vindobonae, Braumüller & Sei-

del : 1-106 + i-ix.

me. 1860. - Die Ausbeute der ôsterreichischen Naturforscher an Säugethieren und Reptilien während

der Weltumsegelung Sr. Majestät Fregatte Novara. Sber. Akad. Wiss. Wien, 42 : 383-416.

GisTEL, J., 1848. - Naturgeschichte des Thierreichs für hôhere Schulen. Stuttgart, Hoffmann : i-xi + 1-

216 + i-iv, pl. I-XXXII.

GRESSITT, J. L., 1938. - Some amphibians from Formosa and the Ryu Kyu Islands, with description

of a new species. Proc. biol. Soc. Wash., 51 : 159-164.

GUIBÉ, J., 1974. - Batraciens nouveaux de Madagascar. Bull. Mus. Nat. Hist. nat., (3), 171 (Zool. 116) :

1169-1192.

GUNTHER, 1859. - Catalogue of the Batrachia Salientia in the collection of the British Museum. Lon-

don, lor & Francis : i-xvi + 1-160, pl. I-XIT.

Source : MNHN, Paris

Dusois 9

Hever, W. R., 1971. - Mating calls of some frogs from Thailand. Fieldiana : Zool., 58 : 61-82.

INGER, R. F., 1954. - Systematics and z0ogeography of Philippine Amphibia. Fieldiana : Zool., 33

183-531.

—— 1966. — The systematics and zoogeography of the Amphibia of Borneo. Fieldiana : Zool., 52 : 1-

402.

INGER, R.F. & FROGNER, K. J., 1979. - New species of narrow-mouth frogs (genus Microhyla) from

Borneo. Sarawak Mus. J., 27 : 311-322.

LAMARCK, J.B., 1801. — Système des animaux sans vertèbres, ou tableau général des classes, des ordres et des

genres de ces animaux. Paris, Deterville : + 1-432.

LAURENT, R. F., 1986. — Sous-classe des Lissamphibiens (Lissamphibia). Systématique. /n : P.-P.

GRassé & M. DELsOL (éds.), Traité de zoologie, Tome XIV, Batraciens, Fasc. 1-B, Paris, Mas-

son : 594-797.

os 1987. — The systematic position of the genus Afrixalus Laurent (Hyperoliidae). Alytes, 5 : 1-6.

Liv, C.-C., 1950. - Amphibians of Western China. Fieldiana : Zool. Mem., 2 : 1-400, pl. 1-10.

MarsuI, M. & BUSACK, S.D., 1985. — A reassessment of the taxonomic status of Rana gracilipes Gres-

sitt, 1938 from Taiwan. Herpetologica, 41 : 159-160.

NELSON, C. E., 1972. Systematic studies of the North American microhylid genus Gastrophryne. J.

Herper., 6 : 111-137.

ne 1973. - Mating calls of the Microhylinae : descriptions and phylogenetic and ecological conside-

rations. Herpetologica, 29 : 163-176.

NIEDEN, F., 1926. - Amphibia. Anura II. Engystomatidae. Das Tierreich, 49 : i-xvi + 1-110.

PARKER, H. W., 1928. — The brevicipitid frogs of the genus Microhyla. Ann. Mag. nat. Hist., (10), 2 :

473-499.

=. 1934. — À monograph of the frogs of the family Microkylidae. London, British Museum : i-viii + 1-

208.

Porr, C. H., 1931. - Notes on amphibians from Fukien, Hainan and other parts of China. Bull. Am.

Mus. nat. Hist., 61 : 397-611, pl. I-X.

SmrrH, M., 1916. — Descriptions of five tadpoles from Siam. 7. mat. Hist. Soc. Siam, 2 : 37-43, 2 pl.

STEINDACHNER, F., 1864. - Batrachologische Mittheilungen. Verhandl. zool.-bor. Ges. Wien, 14 : 239-

288, pl. IX-XVII.

TayLor, E. H., 1962. - The amphibian fauna of Thailand. Kansas Univ. Sci. Bull., 63 : 265-599.

TscHUDI, J. J., 1838. — Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Ab-

theilung der Reptilien. Neuchâtel, Petitpierre : 1-102, pl. I-VI.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2) : 10.

Miscellanea nomenclatorica batrachologica (XVI)

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

A replacement name is proposed for Xenobius Zhang & Hu. 1985 (Hynobiidae).

which is preoccupied by Xenobius Borgmeier, 1931 (Coleoptera).

ZHANG & Hu (1985) ont récemment nommé Xenobius un nouveau genre d’Urodèles

Hynobiidae de Chine. Malheureusement, ce nom ne peut être conservé pour ce genre, car

le même nom avait été donné par BORGMEIER (1931 : 358) à un genre de Coléoptères du

Brésil. Nous proposons le nom de remplacement suivant, qui évoque l’origine chinoise de

celui-ci, pour le genre d’Urodèles :

SINOBIUS nom. nov.

Nomen novum pro Xenobius Zhang & Hu, 1985 (nec Borgmeier, 1931).

Espèce-type, par désignation originale sous Xenobius Zhang & Hu, 1985 : Sinobius me-

lanonychus (Zhang & Hu, 1985).

Diagnose : voir ZHANG & Hu (1985).

RÉFÉRENCES BIBLIOGRAPHIQUES

BORGMEIER, T., 1931. — Sobre alguns coleopteros ecitophilos do Brasil (Staphylinidae). Rev. Entom.,

Säo Paulo, 1 : 355-367.

ZmaxG, F. & HU, Q., 1985. — The aquatic evolution of Hynobiidac of China, with description of a

new genus and a new species from Western Anhui. Acta herpet. sin., 4 : 36-40.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2): 11-17. il

Le régime alimentaire des Amphibiens :

méthodes d’étude

Pierre JoLY

Laboratoire de Biologie animale et Ecologie,

UA CNRS 367 Ecologie des Eaux douces,

Université Claude Bernard Lyon 1,

69622 Villeurbanne, Cédex,

France

For the ecologist, it is important to preserve the studied populations. The stom-

ach-flushing method provides the stomach content without killing the animal and thus

allows the researcher to know the amphibian diet without destroying the natural pop-

ulations. After a review of the literature, the paper gives à detailed description of an

equipment which has been extensively tested in the field for stomach flushing of uro-

deles and anurans. Such an equipment can be adapted to particular situations, like those

involving a long walk. Advices are given for prey fixation and determination.

INTRODUCTION

Le régime alimentaire d’une espèce dans un milieu donné est un paramètre souvent

indispensable pour la compréhension de la dynamique de la population concernée. La mé-

thode la plus ancienne d’obtention des données consiste à sacrifier les animaux et à recueillir

le contenu du tube digestif par dissection. L’avantage de cette méthode est de permettre une

lecture directe du taux de remplissage de l'estomac. Mais elle présente par contre un incon-

vénient majeur d’ordre méthodologique : la progression récente des connaissances amène le

chercheur à travailler de plus en plus à l'échelle des groupes fonctionnels de reproduction

(dèmes) ou au niveau des individus. Chez les amphibiens, dans de nombreux cas, ces groupes

ne comptent que quelques dizaines ou centaines d’individus. out prélèvement important

peut alors entraîner rapidement des modifications profondes de la structure du groupe, voire

la disparition de celui-ci.

Depuis une cinquantaine d’années ont été développées des méthodes non destructrices

d’analyse du régime alimentaire dans plusieurs groupes zoologiques. On peut les classer se-

lon trois types :

(1) Méthodes d’observation directe de l’animal en train de chercher sa nourriture ou

pendant le transport d’une proie ; ce sont essentiellement des travaux sur les mammifères

de grande taille (lion : SCHALLER, 1972) ou sur des oiseaux chassant en milieu découvert

(chevalier gambette : Goss-CusTARD, 1977) ou photographiés au moment du retour au nid

(mésange charbonnière : RoyAMA, 1970).

(2) Méthodes d’analyse des réjections orales ou anales : examen des pelotes de réjec-

tion des rapaces (LiBois, 1984) ou de sacs péritrophiques de larves de libellules (CHOVET,

1976).

Source : MNHN, Paris

12 ALYTES 6 (1-2)

(3) Méthodes de lavage gastrique : elles ont surtout été développées chez les poissons

(ROBERTSON, 1945 ; SEABURG, 1957 ; GAUDIN, MARTIN & CAILLÈRE, 1981).

Les deux premières méthodes ne permettent pas de définir correctement le régime ali-

mentaire chez les amphibiens. L'observation directe est difficile car la plupart des espèces

sont nocturnes, ou chassent sous terre ou dans l’eau. On n’observe jamais de transport de

proie. L’examen des excréments révèle la présence de certaines parties sclérifiées des proies

(pièces buccales, griffes, capsules céphaliques, élytres) mais ne rend pas compte de l’inges-

tion de plathelminthes, annélides ou mollusques.

Le lavage de l’estomac ne présente par contre aucune difficulté majeure et a déjà été

pratiqué par divers auteurs tant chez les Urodèles (FRASER, 1976 ; CHACORNAC & JoLY, 1985 ;

GRIFFITHS, 1986), que chez les Anoures (LEGLER & SULLIVAN, 1979 ; OPATNRY, 1980 ; GIT-

TINS, 1987). La cavité buccale et l’estomac ne forment pas entre eux de coude, ce qui permet

le passage aisé d’un tube, même rigide. A la différence de nombreux poissons Cyprinidés,

les amphibiens n’ont pas de dents pharyngiennes qui peuvent s’opposer au passage d’un tube

(exemple du goujon : CASTRO, 1986). Enfin le cardia n’oppose pas de résistance à l'injection

d’eau.

Le but de cet article est de présenter une revue des méthodes et du matériel utilisé,

accompagnée de critiques et de conseils.

LE VIDANGEUR

Il se compose de trois parties :

(1) un réservoir de mise en pression de l’eau;

(2) un injecteur;

(3) un dispositif récepteur du contenu de l'estomac.

Le réservoir de mise en pression peut être une simple seringue (OPATNRY, 1980 ;

GRIFFITHS, 1986), dont le faible encombrement facilite le transport sur le terrain. Un réser-

voir plus volumineux permet des manipulations plus sûres dans de meilleures conditions, en

particulier pour les animaux de grande taille, ou bien lorsqu’on souhaite limiter la durée des

manipulations (la digestion progresse chez les animaux que l’on a capturés et qui attendent

d’être traités). Lorsque la situation du lieu de travail n’implique pas un long trajet à pied,

j'utilise un pulvérisateur pour horticulture d’une capacité de 5 litres. Une pompe manuelle

et une valve permettent d’ajuster la pression de façon à obtenir à la sortie de l’injecteur un

débit qui ne doit pas dépasser 4 ml/s, sous peine de provoquer une déchirure de la paroi

stomacale. La poignée de la lance fixée sur une planchette constitue une pédale qui permet

de contrôler avec le pied d’injection d’eau et d’avoir les mains libres pour la manipulation

des animaux. Pour un travail dans les Alpes qui nécessitait un long portage, nous avons uti-

lisé pour gagner en encombrement et en légereté un réservoir de polypropylène souple de

200 ml. Deux valves anti-retour sont alors nécessaires pour éviter le reflux dans l’appareil

lors de la décompression (voir fig. 1).

Une aiguille de seringue prolongée d’un cathéter ou un trocard dont l’extrémité est en

bourrelet (pour ne pas léser les organes) sont suffisants. Je préfère la deuxième solution car

Source : MNHN, Paris

JoLy 13

Fig. 1. Schéma d’un dispositif léger de vidange gastrique : (1) réservoir de polypropylène souple de

200 ml ; (2) valves anti-retour ; (3) embout injecteur de laiton à deux conduits : (4) pilulier récep-

teur ; (5) cheminée d'évacuation avec filtre ; (6) réserve d’eau. D’après CHACORNAC & JOLY (1985).

la rigidité du trocard permet un meilleur contrôle du mouvement et d’éviter de pénétrer le

poumon. Ce type d’injecteur est irremplaçable pour les animaux de petite taille (Triturus hel-

veticus, T. italicus, T. vulgaris). La réception du contenu de l’estomac se fait dans une boîte

de Petri ou sur un petit tamis (maille 0,5 mm) qui retient les proies. Le contenu est ensuite

transvasé dans un pilulier, éventuellement à l’aide d’un pinceau à gouache.

Avec des animaux de plus grande taille, utilisé un injecteur qui assure en même

temps la réception du contenu stomacal dans un pilulier, ce qui diminue les manipulations.

Ce système dérive de celui décrit par SEABURG (1957) et utilisé chez les poissons Cottidés

(GAUDIN, MARTIN & CAILLÈRE, 1981) et Salmonidés (GEORGES & GAUDIN, 1984). L'eau est

injectée par un fin tube de laiton (1 mm), soudé sur un tube plus large, de diamètre adapté

à la taille de l’animal. Ce tube, par lequel va transiter le contenu de l’estomac, est relié au

pilulier par l'intermédiaire d’une couvercle troué à cet effet. Sur le même couvercle, une

cheminée équipée d’un filtre assure l’évacuation de l’air ou de l’eau excédentaire (fig. 2).

Le tableau I donne pour quelques espèces le diamètre de l’embout à utiliser.

UTILISATION

Il est possible de pratiquer la vidange sur l’animal éveillé. Il est néanmoins préférable

d’opérer sur l’animal anesthésié. Pour ce faire, les animaux sont immergés dans une solution

soit de phénoxyéthanol, dosée à 1 ml pour 2 1 d’eau, soit de MS 222, dosée à 1 g par litre

(dans ce dernier cas, la solution doit être maintenue à l’abri de la lumière, qui dégrade le

produit). L’anesthésie demande plusieurs minutes. Dès leur perte d’équilibre, les animaux

doivent être retirés et placés dans un récipient d’eau pure, afin d’éviter toute intoxication par

une concentration interne trop élevée.

Source : MNHN, Paris

14 ALYTES 6 (1-2)

Fig. 2. Schéma du dispositif de vidange gastrique : (1) pulvérisateur horticole ; (2) vanne-pédale ; (3)

trocard ; (4) boîte de Petri ; (5) embout-injecteur ; (6) pilulier récepteur.

L'animal est maintenu dans une main dans une position verticale, tête en bas. L’in-

jecteur est introduit dans la cavité buccale en passant sur la langue (la rabattre activement

si nécessaire). Dans le cas des embouts pour grands animaux, le passage du pharynx oppose

une résistance qui doit demeurer légère, sinon il faut utiliser un embout de diamètre infé-

rieur. Il n’est pas nécessaire d’enfoncer l’embout plus profondément. Par contre dans le cas

du trocard, il est préférable d’atteindre l'estomac. Au moment de l'injection, l'animal ne doit

pas se gonfler d’eau ; un tel cas signifie que l'injection se fait dans le poumon. Il faut alors

stopper et laisser l’animal récupérer (ce qui se passe bien le plus souvent). Dans le cas des

gros embouts, une résistance à l’écoulement signifie qu’une proie trop grosse obstrue le tube

d'évacuation. Il faut alors retirer l’embout et saisir la proie, qui peut se trouver soit dans le

pharynx, soit à l’extrémité de l’embout, avec une pince douce pour entomologie.

Cesser l'injection lorsque l’eau de vidange s’écoule librement sans transporter de proies.

EFFICACITÉ ET IMPACT

J'ai pu vérifier l’efficacité de la méthode par dissection de l'estomac après vidange chez

quelques animaux de laboratoire, Triturus alpestris, T. cristatus et Rana esculenta. Elle est to-

tale si toutes les précautions précédemment décrites ont été prises (extraction de grosses proies

avec une pince). Seul est resté collé sur la paroi de l'estomac d’un triton crêté un morceau

de feuille morte, sans doute ingéré avec une proie, et dont la forme particulièrement aplatie

a pu ne pas donner prise au courant d’eau.

La mortalité est nulle si l’on veille :

— à respecter la limite de débit conseillée : j’ai pu observer une déchirure de l’estomac

Source : MNHN, Paris

Jory 15

Tableau I. - Correspondance entre espèce ou taille et diamètre des injecteurs.

Espèce Diamètre (mm)

T. helveticus, T. vulgaris trocard

T. alpestris

T. cristatus, T. marmoratus

Rana sp. 30-40 mm

Rana sp. 40-50 mm

Rana sp. 50-60 mm

Rana sp. 60-70 mm

œsauas

chez des tritons avec une pression d’injection trop élévée ; pour un matériel donné, le débit

sera une fonction définissable de la pression dans le pulvérisateur, cette dernière contrôlable

par un manomètre;

— à appliquer une méthode d’anesthésie rigoureuse. L’anesthésique est toujours un

produit toxique qui peut entraîner la mort. Les animaux ne doivent séjourner dans la so-

lution d’anesthésique que le temps nécessaire à leur perte de motricité, et doivent être en-

suite placés dans de l’eau pure, éventuellement aérée. L’excrétion de l’anesthésique par la

peau est ainsi rendue possible.

Afin d’estimer le stress infligé aux animaux, j'ai, sur le terrain, offert des larves de

chironomes à une dizaine de tritons alpestres, après capture et vidange gastrique. Tous ont

mangé dans les trois heures qui ont suivi l’opération. Il semble ainsi que les tritons sup-

portent assez bien cette méhode. Je ne dispose cependant pas de données équivalentes pour

des amphibiens anoures.

FIXATION DES CONTENUS

Dans le cas d’une étude des biomasses, le meilleur fixateur est le formaldéhyde (for-

mol). Deux ou trois ml ajoutés à l’eau qui baigne le contenu assurent correctement la conser-

vation. L'avantage de ce fixateur est de ne pas altérer sensiblement la composition des proies

et donc de permettre la mesure de leur poids, frais ou sec. Son inconvénient est de provo-

quer des irritations de la muqueuse nasale lorsqu'on doit l’inhaler pendant une longue pé-

riode ; aussi est-il recommandé de rincer les contenus sur un petit tamis avant d’effectuer

les déterminations. L'alcool procure un meilleur confort de travail mais présente l’inconvé-

nient d’être un solvant des graisses et de modifier de façon sensible le poids des animaux

L'alcool est donc un bon fixateur dans le cas d’une étude essentiellement qualitative. L’é-

tiquette doit être rédigée au crayon à papier et glissée dans le pilulier : c’est le plus sûr moyen

de limiter les risques de confusion.

DÉTERMINATION DES PROIES

La précision de la détermination dépend du but que l’on se fixe, mais aussi du temps

dont on dispose, car le coût en temps augmente de façon exponentielle avec la précision.

Source : MNHN, Paris

16 ALYTES 6 (1-2)

Mais cette dernière n’apporte pas toujours une information pertinente. Une grande précision

implique aussi des tableaux de chiffres trop grands pour être traités aisément par l’infor-

matique. La détermination jusqu’au niveau de l’espèce est intéressante dans le cas par exemple

d’espèces proies qui présentent des caractéristiques éco-éthologiques précises (répartition dans

le milieu ou trajectoire de déplacement) pouvant fournir au prédateur des éléments de sé-

lection. Dans le cas plus général de la description du spectre alimentaire, les catégories de

proies doivent regrouper des espèces dont les caractéristiques stimulantes (taille, mouve-

ment, contraste, situation dans le milieu, odeur) sont proches.

RÉSUMÉ

Ne pas détruire les populations qu’il étudie est un impératif scientifique ou éthique

pour le chercheur en écologie. En procurant le contenu de l’estomac sans porter atteinte à

la vie de l’animal, la méthode de vidange gastrique permet chez les amphibiens l'étude du

régime alimentaire tout en préservant les populations. Après avoir comparé les méthodes

proposées par la littérature, l’article donne une description détaillée de matériels testés dans

des études de longue durée du régime d’urodèles et d’anoures. Un tel équipement peut être

adapté à des situations particulières comme celle exigeant un long transport à pied. Des

conseils sont donnés pour la fixation et la détermination des proies.

QUELQUES ADRESSES

Pulvérisateur : bons résultats avec un Berthoud Florally 7.

Confection de tamis : tissus nylon de divers vides de maille : Tissmetal, 138 bd de la Croix-Rousse, 69004

Lyon.

Détermination des proies : pour la France :

Ouvrages généraux : /ntroduction à l'étude des macroinvertébrés des eaux douces ; ouvrage collectif édité

par M. BOURNAUD, H. TACHET & P. RicHoUXx. Disponible auprès de : Association française de

Limnologie, 14 avenue de St Mandé, 75012 Paris. La mésofaune du sol de la région lyonnaise par

J. MATHIEU, P. RicHoux & H. TACHET, C.R.D.P. Lyon, 47 rue Philippe de Lassalle, 69004

Lyon.

Fascicules spécialisés édités par l'Association française de Limnologie : Coléoptères, Annélides,

Crustacés, Insectes divers : s'adresser à Prof. R. GINET / A.F.L., Université Claude Bernard Lyon

1, Laboratoire de Biologie animale et Ecologie, 69622 Villeurbanne Cédex.

Pour les autres pays, demander conseil auprès de l'Association de Limnologie du pays concerné.

RÉFÉRENCES BIBLIOGRAPHIQUES

CasTro, M., 1986. — Activité prédatrice du goujon Gobio gobio. Aspects eco-éthologiques. Thèse de troi-

sième cycle, Lyon : 1-197.

CHacorNac, J. M. & JoLv, P., 1985. - Activité prédatrice du triton alpestre (Triturus alpestris) dans un

lac alpin (2125 m, Alpes françaises). Acta Œcologica, Œcol. Gener., 6 : 93-103.

CHover, M., 1976. - L'alimentation de la larve de Cordulegaster boltoni Donov. 1807 (Odonates : Ani-

sopières) dans son milieu naturel. Thèse de troisième cycle, Lyon : 1-72.

Source : MNHN, Paris

Joy 17

FRASER, D. F., 1976. — Coexistence of salamanders in the genus Plethodon : a variation of the Santa

Rosalia theme. Ecology, 57 : 238-251.

GauniN, P., MARTIN, E. & CAILLÈRE, L. 1981. - Le tubage gastrique chez les poissons : mise au point

d’un équipement et test de la méthode chez le chabot : Cottus gobio L. Bull. Fr. Pisc., 282 : 8-

GEORGES, J. P. & GAUDIN, P., 1984. — Le tubage gastrique chez les poissons : expérimentation chez la

truitelle (Salmo trutta L.). Arch. Hydrobiol., 101 : 453-460.

GITTiNs, S. P., 1987. — The diet of the common toad (Bufo bufo) around a pond in Mid-Wales. Am-

phibia-Reptilia, 8 : 13-17.

Goss-CusTARD, J. D., 1977. - Optimal foraging and the size selection of worms by redshank Tringa

totanus. Anim. Behav., 2' 0-29.

GRIFFITHS, R.A., 1986. — Feeding niche overlap and food selection in smooth and palmate newts, Tri-

turus vulgaris and T. helveticus, at a pond in Mid-Wales, ÿ. Anim. Ecol., 55 : 201-214.

LEGLER, J. M. & SULLIVAN, L. J., 1979. - The application of stomach-flushing to lizards and anurans.

Herpetologica, 35 : 107-110.

Lisois, R. M., 1984. - Régime alimentaire de l’effraie et écologie des micromammifères. Cahiers Ethol.

Appliquée, Enquêtes et dossiers: 1-202.

OPATNRY, E., 1980. - Food sampling in live amphibians. West. cs. spolec. zool., 44 : 268-271.

ROBERTSON, O. H., 1945. — À method for securing stomach contents of live fish. Ecology, 26 : 95-96.

Royama, T., 1970. — Factors governing the hunting behaviour and selection of food by the great tit

(Parus major L.). 7. Anim. Ecol., 39 : 619-668.

SCHALLER, G. B., 1972. - The Serengeti lion. Chicago, Univ. Chicago Press : 1-480.

SEABURG, K. G., 1957. - A stomach sampler for live fish. Prog. Fish Cult., 19 : 137-139.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2): 18-22.

On the Triturus vulgaris schreiberi problem :

electrophoretic data

Miloë L. KALEZIC*, Georg DZuxiC**, Jelka CRNOBRNJA** &

Nikola TVRTKOVIC***

“Institute of Zoology, Faculty of Science, Studentski trg 16, 11000 Beograd, Yugoslavia

**Institute for Biological Research “Sinisa Stankovié”, 29 Novembra 142, 11060 Beograd, Yugoslavia

***Croatian Museum of Natural History, Demetrova 1, 41000 Zagreb, Yugoslavia

An electrophoretic study for genetic variation of a Triturus vulgaris population

from Ravni Kotari, an area whose populations were named T. v. schreiberi, revealed that

in contrast to previously analysed non-nominotypical subspecies, no gene loci showed

especial allele frequency patterns. This population is more similar to the nominotypical

subspecies, which is congruent with their morphological similarities (SCHMIDTLER &

ScHminrLer, 1983) and apparently denies a separate taxonomic position for the smooth

neur from Rauni Kotari.

INTRODUCTION

The smooth newt, Triturus vulgaris, underwent considerable evolutionary diversifica-

tion in the Balkan region. A substantial morphological and genetical differentiation, mainly

of populations restricted to small and marginal areas of the species range, occurred during

the series of Pleistocene glaciations (BOLKAY, 1928 ; STEWARD, 1969 ; KALEZIC, 1984). Not

surprisingly, subspecific status has been attached to most of these groups of populations (e.g.

FREYTAG, 1954 ; BRAME, 1967 ; THORN, 1969). One of the subspecies, Triturus vulgaris

schreiberi (WOLTERSTORFF, 1914), was described on a population sample from Bokanjaëko

Blato, near the town of Zadar (fig. 1). According to the original description, the individuals

of this subspecies are characterized by their small size, unspotted belly and by a low and

straight dorsal crest. However, SCHMIDTLER & SCHMIDTLER (1983), who studied morpho-

logical differentiation of the smooth newt populations inhabiting the Adriatic coast zone, de-

nied the existence of T. v. schreiberi. They considered that the populations from Bokanjac

(a village nearby Bokanjaëko Blato) and the surrounding area (Ravni Kotari) belong to the

nominotypical subspecies.

It has been revealed that smooth newt subspecies from Yugoslavia are characterized

by different allele frequency patterns at some gene loci (KALEZIC, 1984 ; KALEZIC & Tucié,

1984). Therefore, in order to yield additional insight into the taxonomic status of the Ravni

Kotari smooth newt populations, we made an electrophoretic study for genetic variation of

a population from the area in question.

Source : MNHN, Paris

KALEZziC, DZUKkIC, CRNOBRNJA, TVRTKOVIC 19

É [5661000

Fe Æj 1000-1500

Nadinsko blato EH > 1500

Benkovac

Fig. 1. Map showing collection locality (Smilëiéi) and distribution of Triturus vulgaris subspecies in Yugoslavia. © -

T. v. vulgaris, À - T. v. meridionalis, @ - T. v dalmaticus, ® - T. ». graecus.

MATERIAL AND METHOD

The sample used for electrophoretic analysis was obtained from a population in the

vicinity of the village of Smil&iéi, 20 km northeast of Zadar (fig. 1). Adult newts, 37 in num-

ber, were collected from a few holes dug nearby which are used as water supply sites for

agricultural purposes. Interestingly, 23 individuals were paedomorphic (mainly females) and

this again conformed to the statement that the Submediterranean area is an important centre

of the smooth newt paedomorphosis (KALEZIC & DZUKIC, 1986).

The electrophoretic techniques used to study protein polymorphism are thoroughly

described elsewhere (HEDGECOCK, 1976). Twenty-one presumptive gene loci were scored (see

Table I in KALEZIC, 1983). Genetic interpretation of allozymic data are based on criteria ela-

borated for Triturus newts by KALEZIC & HEDGECOCK (1980) and KaLEzIC & Tucic (1984).

Estimates of genetic identity are expressed in term of NEr’s (1972) J value.

RESULTS

The following gene loci were monomorphic in the smooth newt population analysed :

Acph-2, Est-1, Fum, Gdh, G-6Pdh, Idh, Ldh, Pgi, 6-Pgdh, Pgm, Prot-1, Prot-2, Prot-3, and

Source : MNHN, Paris

20 ALYTES 6 (1-2)

Table I. - Allele frequencies at four loci in the Smiléiéi sample, and weighted means of allele

frequencies at the same loci of four Triturus vulgaris subspecies (from KALEZIC, 1984).

Locus Alleles Smiléiéi T.v. vulgaris T.v. meridionalis T.v. dalmaticus T.v. graecus

Acph-2 95 0.000 0.075 0.044 0.753 0.000

100 1.000 0.889 0.144 0.000 0.073

105 0.000 0.036 0.812 0.247 0.927

Est4 96 0.435 0.137 0.134 0.000 0.000

100 0.565 0.863 0.866 0.000 1.000

110 0.000 0.000 0.000 1.000 0.000

Mdh-2 93 0.056 0.186 0.044 1.000 0.870

100 0.944 0.637 0.889 0.000 0.130

107 0.000 0.177 0.067 0.000 0.000

Me 98 0.042 0.015 0.015 0.242 0.149

99 0.250 0.307 0.116 0.459 0.461

100 0.653 0.486 0.756 0.299 0.373

102 0.056 0.192 0.113 0.000 0.017

Sod (for abbreviations see Table I in KALEZIC, 1983). Polymorphic loci which were the most

variable included Est-4 with an observed heterozygosity of 0.484, Me (0.444) and a-Gpdh

(0.300). The population from Smiléiéi has a mean heterozygosity per individual of

0.062+0.002, with averaged number of alleles detected per locus being 1.47+0.04. Meta-

morphic individuals appeared to have slightly more genetic variation than paedomorphic

ones ; heterozygosity per individual was 0.068 and 0.059, respectively. Genetically these two

subsamples (“paedomorphic” and “metamorphic”) are very similar ; NEPs Z value is 0.992.

In order to compare the overall genetic identity between the population in question

and populations of smooth newt subspecies which have been electrophoretically analysed so

far, we calculated the 7 values. NEIs measure was computed using weighted means of allele

frequencies of T. v. vulgaris, T. v. meridionalis, T. v. dalmaticus and T. v. graecus (presented

in Table I in KALEZIC, 1984). The population from Smiléiéi is genetically more similar to

the nominotypical subspecies (1=0.987), than to T. v. meridionalis (1=0.977), T. v. graecus

(1=0.907) and T. v. dalmaticus (1=0.865). For the same purpose, Table I shows allele fre-

quencies of four gene loci for the Smil&iéi population as well as for the above-mentioned

subspecies. These loci appeared to differentiate smooth newt subspecies which inhabit the

Yugoslav part of the species range.

DISCUSSION

In contrast to previously analysed T. vulgaris subspecies, none of the twenty-one as-

sayed gene loci of the population from Smiléiéi showed special allele frequency patterns. The

Source : MNHN, Paris

KaLezié, DZukic, CRNOBRNJA, TVRTKOVIÉ 21

absence of a relatively specific genetic entity apparently cannot candidate the smooth newt

population from Ravni Kotari as belonging to a different taxon. Allele frequencies of T. v.

vulgaris populations are the most similar to those of the population we studied, which is

congruent with their morphological similarities as suggested by SCHMIDTLER & SCHMIDTLER

(1983).

À rather broad zone of gene flow between T. v. vulgaris and T. v. meridionalis, pre-

sumably secondary, was found (see fig. 3 in KALEZIC, 1984). Populations from the zone had

Acph-21 and Acph-21% alleles in appreciable frequencies, while allele Acph-21 was by far

the most common in T. v. vulgaris and allele Acph-21% was so in T. 2. meridionalis (Table

D. In the population from Smiléiéi the Acph-2'% allele is fixed and this indicates that smooth

newt populations from Ravni Kotari have no contact with nearby populations of T. v. me-

ridionalis from Lika (see fig. 1). It was found that these groups of populations are separated

by a zone which for unknown reason(s) contains only the Alpine newt (Triturus alpestris).

Such a zone continues and also separates the Ravni Kotari populations from the nomino-

typical subspecies. In spite of isolation, the genetic structure of smooth newt populations

from Ravni Kotari has not changed much from the main body of T. v. vulgaris populations.

The relatively high level of genetic variability of the population in question might indicate

a nonfragmented population structure of smooth newts from Ravni Kotari, and also that these

populations have not suffered population size bottlenecks as has been suggested for some

other populations of the Submediterranean zone (KALEZIC, 1984).

The population from Smilèiéi is substantially different from more southern subspe-

cies, especially T. v. dalmaticus which comprises adjacent smooth newt populations (fig. 1).

Electrophoretically, T. v. dalmaticus is well differentiated from the Smil&iéi population due

to different alleles at loci Acph-2 and Est4, as well as substantial differences in allele fre-

quencies at loci Me and Mdh-2.

RÉSUMÉ

Une étude électrophorétique de 23 locus présumés de 37 individus d’une population

de Triturus vulgaris de Ravni Kotari, région dont les populations ont reçu le nom de T. v.

schreiberi, a montré que dans ce cas, contrairement aux autres sous-espèces non-nominatives

étudiées jusqu’à présent, aucun locus ne manifeste de fréquence allélique particulière. Cette

population est plus semblable à la sous-espèce nominative, ce qui est congruent avec les don-

nées morphologiques (SCHMIDTLER & SCHMIDTLER, 1983), et ce qui indique que les Tritons

vulgaires de Ravni Kotari ne méritent apparemment pas une position taxinomique séparée.

(Résumé traduit par A. DUBOIS)

LITERATURE CITED

Bozkay, S. J., 1928. - Die Schädel der Salamandrien mit besonderer Rucksicht auf ihre systematische

Bedeutung. Zeitsch. Anat. Entw., Munchen, Berlin, 86 : 259-319.

BRAME, À. H., 1967. — À list of the world’s recent and fossil salamanders. Herpeton, 2 : 1-26

Source : MNHN, Paris

22 ALYTES 6 (1-2)

FREYTAG, G. E., 1954. — Der Teichmolch. Wittenberg, Lutherstadt, A. Ziemsen Verlag.

HEDGECOCK, D., 1976. - Genetic variation in two widespead species of salamanders, Taricha granulosa

and Taricha torosa. Biochem. Genet., 14 : 561-576.

KaLeziC, M. L., 1983. - Geographical aspects of genetic variability in the smooth newt (Triturus vul-

garis). Genetika, 15 : 93-103.

KaLezié, M. L., 1984. - Evolutionary divergences in the smooth newt, Triturus vulgaris (Urodela, Sa-

lamandridae) : electrophoretic evidence. Amphibia-Reptilia, 5 : 221-230.

KaLezic, M. L. & DZuxic, G., 1986. — The frequent occurrence of paedomorphosis in the smooth

newt (Triturus vulgaris) populations from the Submediterrancan area of Yugoslavia. Amphibia-

Repiilia, 7 : 86-89.

KaLEziC, M. L. & HEDGECOCK, D., 1980. - Genetic variation and differentiation of three common Eu-

ropean newts (Triturus) in Yugoslavia. Brit. J. Herpetol., 6 : 49-57.

KaLeziC, M. L. & Tucic, N., 1984. — Genic diversity and population genetic structure of Triturus vul-

garis (Urodela, Salamandridae). Evolution, 3 : 389-401.

Net, M., 1972. - Genetic distance between populations. Amer. Natur., 106 : 283-292.

SCHMIDTLER, J. J. & SCHMIDTLER, J. F., 1983. - Verbreitung, Okologie und innerartliche Gliederung

von Triturus vulgaris in den adriatischen Küstengebieten. Spixiana, 1 : 229-249.

STEWARD, J. W., 1969. - The Tailed Amphibians of Europe. Newton Abbot, David & Charles.

THORN, R., 1969. — Les Salamandres d'Europe, d'Asie et d'Afrique du Nord. Paris, Lechevalier.

WOoLTERSTORFF, W., 1914. - Zwei neue Tritonenformen der paläarktischen Region. Abh. Ber. Mus.

Naturk. Vorgesch., Magdeburg, 2 (4) : 371-381.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2): 23-26. 23

On the identity of the so-called “algae like cells”

in tadpole cultures of European green frogs

(Rana ridibunda)

Manuel POLLS PELAZ

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

Green algae of the genus Chlorella, corresponding morphologically to the so-

called “algae like cells” of previous authors, were found in laboratory water conditioned

with tadpoles of European Green Frogs. The colour of these cells seems to depend on

the light conditions under which the tadpoles are raised. These algae were found in va-

ried culture conditions, including those where the tadpoles show rapid and synchroni-

zed development ; this suggests that these algae do not play a significant inhibitory role

in the development of tadpoles.

INTRODUCTION

In the course of studying the micro-organisms associated with the alimentary tract of

tadpoles of European Green Frogs (Rana ridibunda of hybrid origin), I found the same cel-

lular types that had been described and photographed by RICHARDS (1958, 1962), AKIN (1966),

LicxT (1967), and HEUSSER & BLANKENHORN (1973), cultured from other tadpoles of va-

rious American and European anurans. Some of these authors attributed an inhibitory role

to these cells during the growth of larvae. This paper is devoted to their identification, with

a comment on this proposed inhibitory role.

MATERIAL AND METHODS

The larvae studied came from a clutch deposited the night of 5 June 1985 in the la-

boratory by a pair of diploid Rana kl. esculenta specimens, which had been found in am-

plexus the same day in a pond in the Fontainebleau forest near Paris. These tadpoles were

Rana ridibunda of hybrid origin (GRAF & PoLLs, in prep.). They were reared in the Paris

Museum from June to September 1985 at various densities (5-30 tadpoles/liter), volumes of

water (0,5-1,5 liter), and exposed to varied intensities and sources (natural and artificial) of

light. The culture water was tap water which had remained standing for 24-28 hours for de-

chlorination ; it was replaced in part (1/3 of each volume) every 1-2 days. The tadpoles were

fed “ad libitum” with a progressive diet of boiled lettuce salad, Tetramin, and cooked egg

Source : MNHN, Paris

24 ALYTES 6 (1-2)

Fig. 1. Chlorella green algae cells found in tadpoles’ cultures, corresponding to the s0-called “algae like cells” of

previous authors.

yolk. Microscopic observations, measurements and photographs were made at magnification

of 400. Initial observations were made just after sampling, with samples isolated in sterilized

20 cm flasks, under natural North-oriented sun-light. Later observations were made on the

same samples, from a week to several months after isolation. Taxonomic identification of the

algae cells was confirmed by Dr. BOURRELY.

RESULTS

The same cell types that had previously been described in the American and European

literature (see above) were found in all water samples — with, or without, any inhibition on

growth of the tadpoles. The colour of these cells was variable ; cells from the samples with

natural light were green, while those from the samples with weak artificial light were un-

coloured. Independent of colour, all these cells had the morphological characteristics of green

algae belonging to the genus Chlorella. The cells often formed aggregations (fig. 1). The cha-

racteristics were the same as described by BOURRELY (1966) for all of probably more than

30 species in the genus :

— Spherical or elliptic cells.

— Cells containing a plast and a pyrenoid.

— Diameter from few to maximal 20 micrometers.

After one week of incubation of the uncoloured forms under strong sun-light and in

isolation from external contamination, major blossom of green Chlorella cells was observed,

with an almost total disappearance of the non-coloured forms. The same bottles, put in total

darkness, retained green algae for several months.

Source : MNHN, Paris

POLLS PELAZ 25

DISCUSSION

RicHARDS (1962) tentatively identified the same types of algae, which she found un-

coloured, as belonging to the genus Prototheca. The morphology of the two genera is very

similar, and they only differ with respect to colour — green (Chlorella), versus unpigmented

(Prototheca) ; however, some mutations are known in the genus Chlorella which affect the

number of pigment molecules (MARGALEF, 1983). The possibility exists that there are two

algal genera in my samples, but this would not alter any conclusions about the possible in-

hibitory role of these cells in tadpoles growth. It should be noted that, in the same culture

water where I found Chlorella, several species of algae (e.g. Selenastrum, Euglena, Phacus,

Cosmarium, Scenedesmus, Phacotus), protozoa and bacteria (e.g. Zooglea, Moraxella, Aero-

monas, Kurthia, Micrococcus) were also found (KAISER & POLLS, in prep.).

AKIN (1966) suggested that the “algae like cells” could execute an inhibitory role on

tadpoles as carriers of a molecule present in the intestine of anuran larvae. Following the

opinions of WEST (1960) and STEINWASCHER (1978) — who discusses earlier accounts by ROSE

& ROSE (1961), STEPANOVA (1974) and RUNKOVA, STEPANOVA & KOVAL’CHUK — , I consider

that, except possibly at extremely high concentrations, Chlorella plays a negligible role in the

initiation of growth inhibition.

ACKNOWLEDGEMENTS

I thank Dr. P. BOURRELY and Dr. R. POURRIOT for their help in algae identifications,

and Dr. A. Dugois and Dr. R. WAssERSUG for their interesting comments on the manus-

cript. The original French manuscript was kindly translated into English by Dr. A. DUBoIs.

RÉSUMÉ

Au cours d’un élevage de têtards de la Grenouille verte européenne, des algues vertes

appartenant au genre Chlorella ont pu être détectées ; leur morphologie était identique à celle

décrite par d’autres auteurs dans d’autres cultures de tétards, pour d’autres espèces d’Anoures

dans d’autres localités ; à cause de leur caractéristique incolore, ces algues avaient été ap-

pellées des “algae like cells”. Dans nos élevages la présence des cellules vertes (Chlorella) ou

des non colorées paraît dépendre des conditions d’éclairage. Des Chlorella et des cellules non

colorées ont été retrouvées dans différents cristallisoirs d’élevage des têtards, y compris ceux

où on a observé un développement bon et rapide, synchronisé pour le groupe.

De ce point de vue, le rôle inhibiteur de la croissance des têtards, attribué par d’autres

auteurs à ces algues, ne semble pas devoir seulement dépendre de la présence qualitative des

“algae like cells” ou des Chlorella.

Source : MNHN, Paris

26 ALYTES 6 (1-2)

LITERATURE CITED

AkIN, G. C., 1966. — Self-inhibition of growth in Rana pipiens tadpoles. Physiol. Zool., 39 : 341-356.

BOURRELY, P., 1966. - Les Algues d’eau douce. I. Algues vertes. Paris, Boubée & Cie : 1-511.

HEUSSER, H. & BLANKENHORN, H., 1973. - Crowding-Experiments mit Kaulquappen aus homo- und

heterotypischen Kreuzungen der Phänotypen esculenta, lessonae, und ridibunda (Rana esculenta -

Komplex, Anura. Amphibia). Revue suisse Zool., 80 : 543-569.

Licar, L. E., 1967. - Growth inhibition in crowded tadpoles : intra specific and inter specific effects.

Ecology, 48 : 736-745.

MARGALEF, R., 1983. — La Limnologia. Barcelona, Omega : i-xiv + 1-1010.

RicHarDs, C. M., 1958. — The inhibition of growth in crowded Rana pipiens tadpoles. Physiol. Zool.,

1 : 138-151.

ca 1962. — The control of tadpole growth by alga-like cells. Physiol. Zool., 35 : 285-296.

Rose, S. M. & ROSE, F. C., 1961. — Growth controlling exudates of tadpoles. Symposia of the Sociery

for Experimental Biology, 15 : 207-218.

RUNKOVA, G. G., STEPANOVA, Z. L. & KOVAL'CHUK, L. A., 1974. - Organ specificity in the action of

metabolites of amphibian larvae on their endogenous metabolism under condition of increased

population density. The role of protein in the regulating influence of metabolites. Akademia Nauk

SSSR Doklady Biological Sciences Section, 217 : 328-330. Not seen ; cited fide STEINWASCHER

(1978).

STEINWASCHER, K., 1978. — Interference and exploitations competition among tadpoles of Rana utri-

cularia. Ecology, 59 : 1039-1046.

STEPANOVA, Z. L., 1974. - The chemical nature of the products of metabolism of amphibian larvae,

excreted into the water. Soviet Journal of Ecology, 5 : 148-149. Not seen ; cited fide STEINWAS-

CHER (1978).

WEsT, L. B., 1960. - The nature of growth inhibitory material from crowded Rana pipiens tadpoles.

Phsyiol. Zool., 33 : 232-239.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2) : 27-55. 27

Again on the nomenclature of frogs

Alain DUBoIS

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

CONTENTS

Abstract …

Introduction . A

Suggested rules for the nomenclature of class-group taxa.

Distinction between different types of generic names and spellings.

Introduction …

Different types of generic names and spellings

New genus versus new replacement name.

Unijustified emendation versus incorrect subsequent spelling

Unjustified emendation versus new replacement name …

WAGLER’s generic replacement names

Other problems raised by SAVAGE (1986) in generic names.

Valid family-group names …

Classification used .

Conclusion …

Résumé.

Literature

ted.

The author answers the various criticisms recently expressed by SAVAGE (1986)

on his own works, and shows that these are due to severe misunderstandings of several

important rules of nomenclature. À particular attention is given to the problems raised

by the distinction of different types of names and spellings (new name. new replacement

name, unjustified emendation, incorrect subsequent spelling), and several examples are

treated in detail. Some of the suggestions of SAVAGE (1986) concerning family-group

names are shown to amount to a proposal of a return to pre-Code rules for these names,

since this author does not accept the principle of priority for them. The question of the

rules suggested by Dugois (1984) for the nomenclature of class-group taxa is also dis-

cussed, and the modifications recently proposed in this respect by LESCURE, RENOUS &

Gasc (1986) are criticized.

Note. — This paper pas judged 100 long by the editors of Copeia to whom it had first been sent.

In order 1 save space, the various papers by DuROIS cited below will be referred to using code

numbers, which are made explicit in the bibliography. The abbreviation ASW refers to FRoST's (1985)

checklist.

Acknowledgements. — 1 wish to thank Annemarie OHLER for her help in the preparation of this

paper and Ronald I. CROMBIE, Richard WASsERSUG and particularly Carl GANS and Jean-Jacques Mo-

RÈRE for reading the original manuscript and for their comments about it.

Source : MNHN, Paris

28 ALYTES 6 (1-2)

INTRODUCTION

The journal Copeia recently published a review by SAVAGE (1986) of a paper of mine

on the suprageneric nomenclature of anuran amphibians (D-26). It so happens that this au-

thor, who is a member of the ICZN, disagrees with me on several important nomenclatural

points, but that I am not at all convinced by his arguments. I am answering these comments

in detail below. As will be shown, some of the points in question are not unrelated to some

of the comments I made elsewhere (D-41) concerning ASW.

SAVAGE’s (1986) comments bear on different aspects of my work, which will be dis-

cussed successively below : suggested rules for the nomenclature of class-group taxa ; dis-

tinction between different types of generic names or spellings ; valid family-group names ;

classification used. As will be shown, SAVAGE’s (1986) disagreement with my work comes

from two major sources : disagreements on the interpretation of the /nternational Code of z0-

ological Nomenclature (ANONYMOUS, 1985 a) ; and misunderstanding of some parts of my

(French) text.

An initial comment. SAVAGE (1986 : 259) writes : “Unfortunately Dubois would have

been better served had he waited for the appearance of the revision (3rd ed.) of the Inter-

national Code of Zoological Nomenclature (1985) to appear (sic) instead of basing many of

his conclusions on the 2nd ed. (published in 1964 and amended in 1974).” This is a rather

strange statement indeed. The “short work” (as SAVAGE, 1986 : 259, calls it) in question was

the result of a 4 years study. It was finished and sent to printer in 1983, and published in

1984. On the other hand, the 3rd. edition of the Code had been announced for several years

already, and there was no assurance that it would appear in 1985 rather than in 1986 or even

later. SAVAGE (1986) is not serious when he says that everything should have stopped in the

world, or at least that no nomenclatural work should have been published “just before” the

publication of this 3rd. edition. The question in fact must be reversed. This 3rd. edition

introduces significant changes in some articles of the Code, and indeed I immediately cor-

rected my own work just after the new Code had appeared, in a paper (D-29) that SAVAGE

(1986) ignored, although it was published in August 1985. But, as was pointed out in this

paper, the changes introduced in these articles of the Code are largely open to criticism, since

they generate an unavoidable new cause of confusion and instability (see also below). It would

have been the responsibility of the authors of this new edition to avoid introducing such new

causes of disruption of nomenclatural stability, and not of systematists to stop working until

the new edition appears. Now, this new edition, which was clearly prepared too quickly, is

undoubtedly open to strong criticism (see e.g. DurPuis, 1984 ; DuBois, D-29), and will cer-

tainly have to be modified in some of its parts in the future. To follow SAVAGE’s suggestion,

should all systematists stop to work on nomenclatural matters, in the expectation of the

forthcoming 4th. edition of the Code? Let us hope they will not, although it is clear that a

“final” Code will never exist.

SUGGESTED RULES FOR THE NOMENCLATURE OF CLASS-GROUP TAXA

SAVAGE (1986) provided a shortened translation of some of the rules I proposed (D-26)

for the nomenclature of class-group taxa (i.e. all taxa above the family-group). This trans-

Source : MNHN, Paris

Dugois 29

lation is correct, although abridged, for most rules. Two errors of translation must however

be pointed out. In rule No. 4, the last part of SAVAGE’s sentence (‘‘and were not created for

taxa superior in rank to those of the class-group”) is completely invented : since I proposed

to consider all names above the family-group as belonging to the class-group, there exist no

taxa superior in rank to those of the class-group ; even the names of highest ranks, such as

those of phyla and kingdoms, do belong, in my opinion, to the class-group, for the reasons

explained below. This last part of the sentence should be replaced by the following : “even

when they were not the first ones to have been created for the higher taxa in question.” In

rule No. 6, the first part of SAVAGE’s sentence (“In the absence of an international consensus

among several old names that have had considerable usage it is preferable to choose”, ital-

ics mine) is incorrectly translated and should be replaced by : “In the absence of any inter-

national consensus and of old names having been used for a long period, it may be preferable

to choose..….”. As will be acknowledged by those who will look carefully at these sentences,

the errors of translation sensibly modified my proposals, making them more open to criti-

cism. Besides these errors of translation, it should be added that my proposals were not lim-

ited to these six rules. I also proposed to generalize the use of the type concept to the names

of the class-group, and suggested that the types of class-group taxa should be genera, not

families or other taxa. These type genera would be designated either by (original or subse-

quent) monotypy, or by subsequent designation among the originally (or subsequently) in-

cluded genera of the taxon.

Recently, LESCURE, RENOUS & Gasc (1986) suggested to modify my proposals, in two

respects : (1) they proposed to recognize not only one class-group above the family-group,

but a class-group and an order-group; (2) they suggested that types of class-group taxa should

be families, and types of order-group taxa should be classes. I disagree with these proposals,

for the following reasons.

To separate higher taxa (above the family-group) in two different groups would be ex-

tremely artificial, because many names first created e.g. for an order were later applied to a

class, or the reverse. If two distinct sets of names were to be recognized, that means that the

principles of priority and of homonymy would apply independently in these two groups. Thus,

when e.g. a name created for a taxon considered a super-order by its author is transferred

to the category subclass by a subsequent author, it would then have to be treated as a new

name, with its own author and date. This would complicate very much and most unneces-

sarily the matters. Furthermore and above all, if class-group and order-group names were

separated, the principle of homonymy could not apply between them : this would allow the

possibility that the same name be beared e.g. by a class and by an order (either included in

the class, or in another class, possibly in a quite different group). This would be a strong

source of confusion, and should be avoided. The principle of homonymy is an important

principle of the Code, which should not be underestimated. Due to the very high number

of species-group names, it cannot apply in an absolute way at this level, where it only applies

within a given genus. But at higher levels, it applies within the whole animal kingdom : no

two animal genera or families can have the same name, no matter how far these taxa may be

in the phylogeny and classification. I think the same should apply to all animal taxa above

the family-group. This is all the more justified that the number of taxa at this level of clas-

sification is extremely low as compared 10 the numbers of taxa at lower levels and there would

be no point in distinguishing two independent sets among this low number of names. Fi-

Source : MNHN, Paris

30 ALYTES 6 (1-2)

nally, if a distinct group of names was to be recognized for orders and related taxa, the same

should be true for phyla, kingdoms and other higher taxa.

In reality, there exists at present no theoretical reason for stating that a higher taxon

is, say, a super-order or a subclass. These categories are only successive steps in a hierar-

chical scale, and if new dichotomies are added, new taxa recognized, or, on the reverse, if

taxa are merged, the place of many taxa in the scale may well move up or down. It is true

that the same also occurs for taxa of the family-group and of the genus-group, but never-

theless, at least at the genus level, some rules may be proposed to reduce the part of arbi-

trariness in the allocation of taxa to given categories (see DuBois, D-12, D-14, D-30). There

seems therefore to exist neither practical nor theoretical justification for recognizing two dis-

tinct groups of names above the family-group. These principles had guided me in my pro-

posal of a single class-group for all these names, and I had to explain them here in full detail

only because this proposal was challenged.

It was also after reflection, and not by chance, that I suggested that nominal class-group

taxa should have types, and that these types should be nominal genera, not families, or or-

ders, etc. As a matter of fact, a hierarchical succession of types would only unnecessarily

complicate the nomenclature of higher taxa and would create additional difficulties at any

change of diagnosis of the higher taxa and at any transfer of included lower taxa from one

group to another. In the family-group, all taxa names are based on generic names, whatever

their rank : a familial name is not based on a subfamilial name, a subfamilial name on a tribal

name, etc. Thus all taxa of the class-group should, for more simplicity and clarity, be based

on names of the same category. Then, why not choose the family, rather than the genus, for

this category? For two reasons: (1) since family-group names cannot be, like class-group

names, entirely new names, but are based on generic names (and formed in a very strict

manner on the basis of the stem of these names), to designate a type family is strictly equiv-

alent to designating a type genus (on the name of which the family name is based) and brings

no additional information ; (2) many old class-group names were created at the beginning of

systematics, when the category family still did not exist or was not in common use (it began

so only quite after the beginning of the XIX century). Therefore, these taxa were created

without included families, while they were in most cases credited with included genera at

their foundation, the category genus being in use since LINNAEUS. Now, the proposal of

LESCURE, RENOUS & Gasc (1986) would make more difficult, in many cases, the determi-

nation of the type taxon of a given class-group taxon. In all the cases where the taxon was

created with no included family, the “originally included families” would be the first ones

to have been referred to this taxon by subsequent authors, and this information may be dif-

ficult to trace. Another possibility would be to consider that the originally included families

are those based on the generic names cited as included in the new taxon at its creation, but :

() this would be strictly equivalent to recognizing type genera (rather than type families)

for these taxa ; (2) the originally included genus (or genera) may not be the type genus (gen-

era) of (an) existing family-group name(s), which would cause an additional problem. For

all these reasons, I think it justified to maintain my initial proposals and to refuse the changes

proposed by LESCURE, RENOUS & Gasc (1986).

To come back now to SAVAGE’s (1986) comments, he seems to accept my proposals,

but he adds rightfully that to have universal value these rules should be adopted by the ICZN

after discussion : this is true, but before they could be discussed these rules had to be pro-

Source : MNHN, Paris

Dugois 31

posed by someone. SAVAGE (1986) does not seem to reject my rule No. 3 (“A class-group

name that is a junior homonym of another class-group name must be rejected”). Then, why

does he state that the names Archaeobatrachia Reig, 1958 and Neobatrachia Reig, 1958 are

“perfectly good names”, when I had shown (D-19, D-26) that these names are junior hom-

onyms (despite the one-letter difference) of SARASIN & SARASIN’s (1890) Archacobatrachi and

Neobatrachi? Also, he states that my rule No. 6 creates difficulties, and he seems to suggest

that a strict priority rule should be followed. However this proposal is contradictory to my

rule No. 4. To take only the example of anurans, if a rule of priority was to be followed,

neither the names of NOBLE (1931) nor those of STARRETT (1973) would be the valid ones

for the suborders of this order. Numerous much older names do exist (see KUHN'’s 1967 list,

which furthermore is incomplete), and strict application of the rule of priority to this case

would require the resurrection of names which have never or almost never been used and

which are almost completely forgotten now. This is the reason why I had expressly stated

(D-26 : 9) that, at the class-group level, the principle of priority should not be strictly ap-

plied.

Finally, it is the full right of SAVAGE (1986 : 261) to “like subordinal names such as

Archaeobatrachia or Lemmanura better than Discoglossoidei”, but let us note that class-group

names such as Discoglossoidei have not only been proposed by SokoL (1977) in Anura, but

also by LESCURE, RENOUS & Gasc (1986) in Gymnophiona, and are currently universally ac-

cepted as valid by all authors in Urodela (see e.g. ASW), so that, at least for amphibians,

names of this kind have tended to gain a growing acceptance among systematists.

DISTINCTION BETWEEN DIFFERENT TYPES OF GENERIC NAMES OR SPELLINGS

INTRODUCTION

As unpleasant as it may be to some modern taxinomists, systematics is not a young

discipline, and it has undergone many changes since its beginnings. These changes have been

particularly drastic in that part of systematics which deals with the names of taxa, i.e. no-

menclature. The beginning of “modern” nomenclature is arbitrarily fixed in 1758, at the

publication of LINNAEUS’ 10th. edition of the Systema Naturae, but it is clear that at this

time almost no rules existed. The need for nomenclatural rules appeared and developed only

with the increase of the number of known taxa and of their zoological names, and led to the

creation of international Rules of zoological Nomenclature, which have been the matter of

various modifications and of several editions, the last of which appeared very recently

(ANONYMOUS, 1985 a ; see pp. xv-xvi of this book for a very brief history of the current Code).

It is clear that works published prior to the establishment of any set of rules could not follow

them. For example, no principle or priority was followed by many authors in the XVIII and

XIX centuries, and even quite late in the XX century some authors did not follow it in all

cases : e.g. in amphibians BOULENGER as late as in 1920 still rejected some names (e.g. Po-

lypedates afghana) for “inappropriateness”, although he clearly acknowledged that they were

senior synonyms of names considered by him as valid.

The same applies to all the other rules of the current Code (principle of homonymy,

rules for the designation of types, rules for the formation and treatment of names, rules for

Source : MNHN, Paris

32 ALYTES 6 (1-2)

the authorship of names, etc.). This is important to bear in mind when in 1986 we use the

1985 edition of the Code to analyse old, pre-Code texts. I have already stressed on several

occasions (Dugois, D-21, D-42 ; Bour & DuBois, 1984) the fact that the application of the

Code to such texts “must be made with care, understanding and intelligence” (Bour & Du-

BOIS, 1984 : 357). For example, the concept (and term) of lectotype was created only at the

beginning of our century (SCHUCHERT & BUCKMAN, 1905) and became widely used only in

the second half of this period ; it is therefore irrelevant to reject a lectotype designation which

was made in the older times by the simple use of the term “type” (see e.g. BISCHOFF, 1982

and Dugois, D-42).

Similarly, to decide if a new name was proposed as the name of a new taxon, or as a

new replacement name (nomen novum) for a taxon already recognized by previous authors

requires a detailed and careful analysis when old texts are involved. Here we come to the

main criticism of SAVAGE (1986) concerning my work. SAVAGE (1986 : 260) writes : “[Du-

bois] apparently believes that the citation of a previously proposed name in the synonymy

or footnotes associated with the proposal of another name by a subsequent author makes the

latter name a new replacement name.” This statement is totally unfounded, since I never

wrote or even believed such things. My judgments concerning the status of old names never

rely on “general rules” like the strange rule invented by SAVAGE, but on a careful analysis

of the texts themselves, as is shown in my previous works on similar problems (D-8, D-11,

D-17, D-18, D-21, D-24, D-26, D-29, D-40). In this respect, I disagree with SAVAGE (1986)

and also with HoLTHUIS (1983) on the status of the generic name Dendrobates and of several

other generic names created by WAGLER (1830 b). To save space in the Bull. zool. Nom.,

and because I thought all interested biologists could go by themselves back to the original

texts, study them honestly, and come to the same conclusion as myself, I wrote : “Instead

of discussing this in detail in this Bulletin, I think it simpler to refer the readers to WAGLER’s

(1830) original text itself. Other arguments could also be found by studying the other pub-

lications of WAGLER.” (D-21 : 198). However, SAVAGE’s (1986) paper shows that I had been

too optimistic. Since despite my previous papers on this question (D-18, D-21), some people

remain unconvinced by my arguments, it is necessary to come back to the question in more

detail, as heavy and space consuming as it may seem.

Before going into the details of WAGLER’s works, however, it will be useful to discuss

these matters at a more general level.

DIFFERENT TYPES OF GENERIC NAMES AND SPELLINGS

When a systematist uses a generic name in a scientific publication, he may use an ex-

isting name, without changing it or with some modifications, or a new name. For the sake

of clarity, the different possible cases may be presented in the form of a dichotomic key.

I. The writer uses a new generic name for a genus which he considers new (even if

including already named species): both the taxon and the name are new, and must be cred-

ited to the author of the publication. In modern times, the new name is generally presented

with the indication “gen. nov.” or “n. gen.”. In older texts, indications like “mihi” or “nobis”

were sometimes used, but in many other cases the fact that the name and taxon are created

as new in the paper must be inferred from other direct or indirect sources of evidence.

Source : MNHN, Paris

Duois 33

Il. The writer deals with a genus which he credits to a previous author (even if he

modifies in part the diagnosis or the contents of this genus, e.g. in retiring from it or adding

into it some species). This reference to an already existing taxon is generally explicit, but in

some cases must be inferred from the context : for example, in older papers many batrach-

ologists used the names Rana, Bufo or Hyla without ever mentioning their authors, but it

was however clear that they were not creating new names or taxa. In this case the taxon is

not new (although it may be somewhat emended). As for the name, two possibilities appear.

A. The writer uses exactly the same name (same spelling) as the author who had

first recognized the taxon and named it. This situation is clear: neither the taxon nor the

name are new.

B. The writer uses a spelling slightly or totally different from that of the original

name. Two possibilities again appear here.

1. The new spelling differs slightly (e.g. generally by one letter or a few letters)

from the original spelling, and this difference in spelling is not intentional from the part of

the writer : it may be due to a misspelling of his part (e.g. a mistake in copying the original

text) or of the part of the printer (misprint). Such a spelling is an “incorrect subsequent

spelling” and it has no status in nomenclature. In this case like in the preceding one, neither

the taxon nor the name are new.

2. The writer uses intentionally a different name or spelling because he thinks,

for some reason, that the original name created by the first author is incorrect or invalid and

must be modified or replaced. In this case it is clear that the taxon is not new but that the

name is. In modern times, this new name is called a new replacement name or nomen novum

(ANONYMOUS, 1985 a), and it is generally presented with the indication “nom. nov.”. In older

times, it was sometimes presented as a new name in an explicit sentence, but in many other

cases the fact that it was such a name must be inferred from other direct or indirect sources

of evidence. This latter category may again be subdivided into two categories, although as

will be discussed in detail below, no objective or reliable criteria currently exist to distin-

guish between them in all cases.

a. The new name is a new replacement name for the original name, i.e. a com-

pletely new name, not derived from the original one.

b. The new name is an emendation of the original spelling of the name, i.e. only

the spelling of the name is changed but the new spelling is clearly derived from the original

one. The current Code here distinguishes again two subcategories : justified emendations

(which have no separate status in nomenclature) and unjustified emendations (which have a

separate status in nomenclature).

Since in all the older texts (XVIII and most of XIX century texts) the above distinc-

tions were often not clearly expressed in all words, we need some criteria to decide what

kind of name was used by a given author. In this respect it will be useful to study separately

several types of distinctions which may prove difficult in some cases : new genus name ver-

sus replacement name ; unjustified emendation versus incorrect subsequent spelling ; un-

justified emendation versus new replacement name.

Source : MNHN, Paris

34 ALYTES 6 (1-2)

NEW GENUS NAME VERSUS NEW REPLACEMENT NAME

SAVAGE (1986 : 259) states that according to the Code a name may be considered as a

new replacement name only when it is “a new name expressly proposed as a replacement for

an available name” (italics his). This is only partially correct. As a matter of fact, the words

“proposed expressly” appear in the Code only in Art. 13(a)(üi), which deals with names pub-

lished after 1930, but in Art. 12(b)(3), which concerns names published before 1931, the Code

only states that “the proposal of a new replacement name (nomen novum) for an available

name” is enough to make this name available. In the Glossary of the Code (p. 258), a ‘new

replacement name”, is defined as “A name established expressly to replace an already estab-

lished name”, which is more restrictive that the wording of Art. 12. In fact, one could dis-

cuss at length about the sense which should be given to the word “expressly” in these sen-

tences, but this would make little sense. I have always believed that, to solve a problem, it

is more useful to take time to think about it than to repeat or underline words. Now, when

dealing with old taxinomic words, how will it be possible to distinguish between a new ge-

neric name and a new replacement name for an existing genus?

The writers of many older texts did not indicate the authors of the generic names they

were using. For some well-known names (like Rana, Bufo or Hyla), this causes no problem,

but the same is not true for less common names. Furthermore, a very special, but real, prob-

lem arises from the fact that LINNAEUS (1758, 1761, 1766, 1767) had a very wide concept of

the genus, so that his genera were later subdivided. In many cases, the subsequent authors

named the new genera with terms which LINNAEUS had proposed for species. For example,

among the species placed by LINNAEUS (1758) in his genus Rana were R. pipa, R. bufo and

R. hyla. LAURENTI (1768) used the same terms for his genera Pipa, Bufo and Hyla. In some

cases however, this may raise some problems. For example, LINNAEUS’s (1761) species Rana

bombina served as the basis for OKEN’s (1816) genus Bombina. MERREM's (1820) genus name

Bombinator was obviously based on the same root. Is this latter name a new generic name or

a new replacement name (or an emendation) for Bombina? This can be established only

through a careful examination of the original publication. Fortunately, MERREM (1820) gave

long and detailed synonymies in his book, and, not only did he not cite the generic name

Bombina as a synonym of Bombinator, but, even more, he did not cite OKEN at all ; he pre-

sumably did not know OKEN’s (1816) Lehrbuch der Naturgeschichte. For this reason, I con-

sidered Bombinator Merrem, 1820 as a new generic name, not as a replacement name (or an

unjustified emendation) for Bombina Oken, 1816 (D-26 : 13-14 ; D-29). This example shows

that a careful analysis of the original texts is often required to ascertain the status of a name.

Às a general rule, assurance that a new name is not the name of a new taxon but a new

name for an existing taxon requires some evidence, in the original text, that this was the

intention of the author. This evidence may be of various types, as will be shown by some

examples studied in more detail below. The existence of such evidence answers to the qual-

ification “proposed expressly” which appears in the Code.

Source : MNHN, Paris

Dugois 35

UNJUSTIFIED EMENDATION VERSUS INCORRECT SUBSEQUENT SPELLING

Here again, SAVAGE (1986) cites and italicizes the phrase “demonstrably intentional”

without discussing its meaning in detail. However, the Code itself gives in its Art. 33(bXi)

additional information in this respect : “A change in the original spelling of a name may only

be interpreted as “demonstrably intentional” when in the work itself, or in an author’s (or

publisher’s) corrigenda, the original and the changed spelling are cited and the latter is

adopted in place of the former, or when two or more names in the same work are treated in a

similar way” (italics mine). Although this statement broadens already very much the concept

of ‘“‘demonstrably intentional” as compared to SAVAGE (1986), who seems to believe that this

phrase may only mean “stated in full words”, and is enough to show that this author is wrong

in considering names like Bombitator or Calamites as incorrect subsequent spellings or new

generic names (see below), it is in my opinion still too restrictive and should be modified in

future editions of the Code. As a matter of fact, the phrase “demonstrably intentional” can-

not be restricted to the two possibilities mentioned in this sentence (either citation of both

the original and the changed spelling, or a “similar” treatment of several names). These words

mean : (1) that the new spelling was proposed in the clear intention to replace the original

spelling of the name ; (2) that this intention was made clear by the writer in some way, not

necessarily the only two ones mentioned above. One of these ways may be that the new

spelling contains by itself this information. For example, when in older days a name was found

to have been incorrectly formed with respect to its etymology (badly latinized, etc.), it often

happened that a subsequent author changed the spelling in order to make it correct. Since

in these remote times all biologists were cultured people, who knew Latin and Greek names

or could easily trace them, it was often felt unnecessary to ‘“expressly” write in all words that

the name had been emended, because this was considered obvious.

In such cases, the evidence that the intention of the author was to replace a (suppos-

edly) incorrect spelling by another one may be found in the new spelling itself, when it was

correctly formed from the Latin or Greek root whereas the original spelling was not, or could

be considered so. As we shall see in detail below, such a case clearly applies to the names

Megalophrys versus Megophrys, Bombitator versus Bombinator, Myiobatrachus versus Myoba-

trachus, etc. If this is believed by some to be contrary to the wording of the Code, then this

simply means that this wording is wrong, and should be corrected. If the ICZN decided that

Megalophrys, Bombitator and Myiobatrachus are not unjustified emendations (but incorrect

subsequent spellings having no status in nomenclature), because their authors did not write

in full words that they were so, then I frankly declare it would be a stupid decision, because

there can be no doubt in such cases about the intention of the authors of such spellings to

correct the original names according to the Latin or Greek grammar.

This does not mean at all, as SAVAGE (1986) seems to believe, that I am ready to accept

any subsequent spelling as an emendation. Some evidence that the intention to change the

spelling must exist in the original texts, and in this respect it is useful to give an indicative,

non limiting, list of such potential evidence. To be deemed an emendation, a name must meet

at least either of the following conditions :

(C1) the name is presented in words as such;

(C2) both the original and the modified spellings are given, and the second one is re-

tained by the writer as the valid one;

Source : MNHN, Paris

36 ALYTES 6 (1-2)

(C3) several names are treated in a similar way, e.g. corrected according to the etym-

ology, or according to some, possibly arbitrary, rule which is evident from the context;

(C4) the modified spelling is clearly etymologically justified and correctly formed, while

the original spelling was not, or could be considered not to be (see below for additional com-

ments);

(CS) the modified spelling is introduced by the very author of the original spelling,

either in a “corrigenda”, as stated by the Code (which however does not precise if the latter

must have been published in the same time as the original name, e.g. as an addenda to it,

or may have appeared later), or in later publications, especially when the new spelling is used

repeatedly in subsequent works and the original spelling definitively abandoned by the au-

thor;

(C6) the modified spelling is introduced by a subsequent author and used repeatedly

in the same or, better, in subsequent works, whereas the original spelling is definitively

abandoned by this author (especially when before introducing the new spelling he had made

use of the original spelling in previous publications).

À few examples of these criteria shall be given and discussed below in the comment

of some of SAVAGE’s (1986) statements ; examples of the other ones may be given here.

Thus, the type (C3) of evidence may be illustrated by PALACKŸ’s (1898) paper in which

this author used a lot of emended spellings, according to a strange rule of his own, which is

clearly evident in the text itself, although it is never stated in words : he systematically re-

placed the letters “ph”, when they appeared within a generic name, by the letter “P” ; on the

other hand, he did not modify the generic names which were beginning by “Ph”. Strange and

unjustified as this “rule” may be, PALACKŸ’s (1898) action was clearly intentional and all these

modified spellings are unjustified emendations, which have a separate status in nomencla-

ture. Since only a few of these names have already been mentioned in recent works (DUBoIs,

D-11, D-17, D-26 ; CLARKE, 1983), I am giving here in Table I a list of PALACKŸ’s (1898)

emendations of generic names, along with the original spellings of these names. Besides these,

PALACKŸ (1898) also proposed a few emendation of specific names (Rana cyanoflyctis, p. 378 ;

Hyla stefeni, p. 379 ; Bufo filipinicus, p. 380 ; Bufo dialofus, p. 381).

An example of the type (C6) of criteria may be provided by the name Ptychadaena Par-

ker, 1930. PARKER (1930 a) introduced the new spelling Piychadaena without any explana-

tion ; this spelling appeared twice in this paper, but the spelling Ptychadena Boulenger, 1917

was not mentioned. In his subsequent publications (e.g. 1930 b, 1932, 1936 a, 1936 b, 1937),

PARKER always used the spelling Piychadaena, but he never apparently mentioned the spell-

ing Ptychadena, which he could not ignore, since he was well acquainted with BOULENGER’s

works. Despite the absence of any “explicit” statement in words in this respect, it is clear

that PARKER (1930 à) intentionally modified the spelling Piychadena, and that Piychadaena

must be considered an unjustified emendation, with its own status in nomenclature.

Now that we have discussed the major types of evidence for the fact that a new spell-

ing was intentionally created by an author, it may be useful to see which are the principal

clues to the reverse situation, i.e. to “incorrect subsequent spellings”:

(C7) no explanation is given for the modified spelling, and the latter has no clear etym-

ological justification (on the contrary, it may often be incorrectly formed while the original

spelling was correctly formed);

Source : MNHN, Paris

Dugois 37

Table I. - Unjustified emendations of generic names of amphibians created by PALACKŸ

(1898).

Unjustified emendation Page in Original generic name Recent

PALACKŸ reference to

(1898) PALACKY”S (1898)

name

Amfignathodon 375 Amphignathodon Boulenger, 1882

Asterofiys 380 Asterophrys Tschudi, 1838

Calofrynus 374 Calophrynus Boulenger, 1882, an unjustified

emendation of Kalophrymus Tschudi, 1838

Chlorofilus 374 Chlorophilus Baird, 1854

Cofofiyne 379 Cophophryne Boulenger, 1887

Eupemfix 376 Eupemphix Steindachner, 1863

Genyofiyne 375 Genyophryne Boulenger, 1890

Glyfoglossus 379 Glyphoglossus Günther, 1868

Megalofys 379 Megalophys Wagler, 1830, an unjustified D-17

emendation of Megophrys Kuhl & Van Has-

selt, 1822 a

Nannofrys 375 Nannophrys Günther, 1869 D-11

Nectofryne 374 Nectophryne Buchholz & Peters in PETERS,

1875

Pseudofryne 374 Pseudophryne Fitzinger, 1843

Rhacoforus 374 Rhacophorus Kuhl & Van Hasselt, 1822 a D-11

Rhombofryne 378 Rhombophryne Bocttger, 1880

Scafopus 381 Scaphiopus Holbrook, 1836

Scaforhina 378 Scaphiophryne Boulenger, 1882

Stenofiyne 380 Sphenophryne Peters & Doria, 1878

(C8) the original spelling is not mentioned, or, if it is, this is in a different part of the

text, as a valid name, and no choice is made between both spellings;

(C9) the modified spelling appears only once in the text, and is never used again by

the same author in subsequent publications, while this author may revert to the use of the

original spelling in these papers;

(C10) the author of this modified spelling is known for having made numerous such

mistakes (because of his carelessness regarding these matters, of his bad handwriting, which

was difficultly read by the printer, of the carelessness of the printer, etc.).

A few examples here also may illustrate these general criteria.

Thus, besides the unjustified emendations mentioned above, PALACKŸ (1898) also in-

troduced modified spellings which are certainly incorrect subsequent spellings according to

type (C8) of evidence above, since he used in the same paper two, or even three, different

spellings, including the original one, and dit not choose between them : Caluela, p. 375,

Calluella and Calluela, p. 380 ; Megalixalus, p. 375, and Megalixcelus, p. 378 ; Batrachy-

lodes, p. 375, and Batrachhylodes, p. 381. In a few other cases, he used generic or specific

names which differ from the original names by no clear rule of transformation, which appear

only once in the text, and which are therefore, according to criteria (C7), (C8) and (C9), best

considered as incorrect subsequent spellings, without status in nomenclature (although their

modification was possibly intentional in some cases): Phrynomantes, p. 376 ; Grypinus and

Source : MNHN, Paris

38 ALYTES 6 (1-2)

Leiuporus, p. 382 ; Rana mascarenensis, p. 377 ; Ixalus kakhynensis, p. 379 ; Cofofryne sik-

kimenus, p. 379.

As I have already had the occasion to stress it (D-17), J.E. GRAY may be considered

as a top specialist in the field of incorrect subsequent spellings : he often used different spell-

ings for the same name in a given work, modified without any reason names previously pro-

posed by other authors or even by himself, etc. For this reason, of the type (C10) of evi-

dence, I suggested that the name Megalophys, which appears in GRAY (1842) and is considered

by SHERBORN (1928) as an unjustified emendation of Megalophrys Wagler, 1830, is in fact an

incorrect subsequent spelling, without status in nomenclature. In the same paper (D-17), I

showed that the spelling Megaphrys, credited by SHERBORN (1928) to GRAY (1825), was also

an incorrect subsequent speling. These examples show that, contrary to what SAVAGE (1986)

seems to believe, I have never considered all subsequent spellings as emendations, but based

my opinion on a careful analysis of the texts themselves.

UNJUSTIFIED EMENDATION VERSUS NEW REPLACEMENT NAME

This problem is not tackled by SAVAGE (1986), but, for the sake of completeness, it

will be briefly discussed here. À more complete discussion of this question may be found in

a previous paper (D-29) ignored by this author.

The need for a distinction between these two types of names is a new one, which was

created by the modifications introduced in Art. 32, 35 and 39 of the new edition of the Code.

Before, both types of names had the same status, or rather, an unjustified emendation was

only considered as a particular case within the general category of “new replacement name”

(see e.g. D-11, D-17, D-18, D-26). As a matter of fact, as is shown in terms of logic in the

dichotomic key presented above, these two types of names are very closely related, being

only subdivisions of the category II(B)2) above : in both cases, a subsequent author replaces

an existing name, which he deems to be incorrect or invalid for some reason, by a new re-

placement name. Of relatively minor importance in this case is the fact that this new re-

placement name may be “completely new”, or obtained by a modification of the spelling of

the original name. Furthermore, not only is this distinction of little relevance, but it is also,

in many concrete cases, difficult to make, as is shown in more detail elsewhere (D-29).

The definition given by the Code (Art. 33) of an emendation (“any demonstrably in-

tentional change in the original spelling of a name”) does not give us any criterion to allow

deciding in a clear, objective and indisputable manner, when one is confronted with a new

spelling which “resembles” that of an already existing name, if it is an “intentional change

in the original spelling” of the name, or a completely new name, based e.g. on a related but

different etymology. When one thinks to this problem in some detail, one realizes that sev-

eral criteria could be proposed for recognizing emendations (e.g.: given number of letters of

difference with the original spelling ; changes in the radical or in the termination of the name ;

recourse to a more exact etymology ; maintenance of the pronunciation of the name ; etc.),

but that none of them is objective and general enough to account for all cases of emenda-

tions. The following examples, discussed in detail by DuBois (D-29), illustrate this complex

problem : Megophrys montana Kuhl & Van Hasselt, 1822 and Megophrys monticola Smith,

1931 ; Kaloula Gray, 1831 and Callula Günther, 1864 ; Kassina Girard, 1853 and Cassina

Source : MNHN, Paris

Dugois 39

Cope, 1864 ; Occidozyga Kuhl & Van Hasselt, 1822 and Oxydozyga Tschudi, 1838 ; Bufo

Laurenti, 1768 and Batrachus Rafinesque-Schmaltz, 1814 ; Rana Linnaeus, 1758 and Ran-

aria Rafinesque-Schmaltz, 1814 ; Hyla Laurenti, 1768 and Hylaria Rafinesque-Schmaltz,

1814 ; Triton Laurenti, 1768 and Triturus Rafinesque, 1815 ; Caecilia Linnaeus, 1758 and

Cecilia [Rafinesque-Schmaltz, 1814].

But, if the Code gives no precise clue for distinguishing emendation from nomen no-

vum in these cases, why should this distinction be important? It only became important with

the following new parts of Art. 32, 35 and 39 of the 1985 Code, which did not exist in the

previous editions :

Art. 32(cX

ily-group name, it (.

Art. 3(dXii): “A family-group name based upon an unjustified emendation of a generic name

is an unjustified original spelling and must be corrected (...)”.

“An original spelling is an ‘incorrect original spelling” if (...) in the case of a fam-

s based (...) on an unjustified emendation of a generic name (...)”.

Art. 39(a): “Effect of unjustified emendations. - If an unjustified emendation of the name of the

type genus becomes itself the replacement name, the family-group name is then to be based upon it by

correcting the name to the spelling required by the stem of the name of the replacement type genus ;

the author and date of the family-group name remain unchanged.”

At first reading, these new rules may appear trifling. In fact, as I have shown (D-29,

D-41), in amphibians these modifications entail a change of spelling, author and date for one

family-group name (Cycloramphini ; sole case noticed by SAVAGE, 1986, although on the ba-

sis of a wrong analysis, since he considers Cyclorhamphus as an incorrect subsequent spell-

ing ; see below), and a change of author and date for 12 other family-group names (Hylidae,

Hylinae, Hyloidea, Megophryinae, Microhylidae, Microhylinae, Pipidae, Pipinae, Pipoidea,

Ranidae, Raninae, Ranoïidea ; none of these changes is remarked by SAVAGE, 1986). Similar

changes have been necessary in the higher nomenclature of chelonians (BoUR & DuBois, 1986)

and will be necessary in all other groups. As justly stressed by LAURENT (1986 b), it is sur-

prising that it was the ICZN, the function of which is to provide rules for allowing the great-

est universality and stability possible in nomenclature, which introduced in 1985 these new

rules which seriously threaten the stability of family-group names. Furthermore, these new

rules are also liable to cause additional confusions and raise new problems even more diffi-

cult to solve, and for which rather artificial solutions must be found, if stability is to be pre-

served : see in this respect the difficulties raised by the new Art. 39(a) of the Code in the

case of the family-group name Triturinae (D-29).

In conclusion, I repeat here my previously expressed opinion (D-29) that the new

changes brought in Art. 32, 35 and 39 of the Code are not good and should be suppressed.

I contend that unjustified emendations should still be considered as only particular cases of

replacement names, and not be treated differently, even when they are the basis of family-

group names. Furthermore, the particular treatment of these names suggested by the Code

in such cases is contradictory with the fact that unjustified emendations continue to be con-

sidered by Art. 33(bY(iii) of the new Code as names having their own status in nomenclature

and available with their own authors and dates.

On the other hand, if these modifications were to be maintained, then the ICZN must

devise and publish precise criteria and rules to allow taxinomists to objectively decide if a

given name is to be considered as an unjustified emendation or as a new replacement name.

I predict that this would not be an easy task.

Source : MNHN, Paris

40 ALYTES 6 (1-2)

WAGLER’S GENERIC REPLACEMENT NAMES

Now that we have seen the things rather generally, let us look more carefully at the

problems raised by some of the generic names used by WAGLER (1830 b). SAVAGE (1986),

following HoLTHUIS (1983), is mainly concerned with the problem of the name Dendrobates,

because in my first paper on this question (D-18) I had presented an interpretation at vari-

ance with that accepted in all the previous applications published in the Bull. zool. Nom. on

this case, one of which was even co-signed by SAVAGE (SILVERSTONE, 1971 ; MYERS & DALY,

1971 ; CUELLAR et al., 1972 ; PETERS et al., 1972 ; LESCURE, 1982). For some reason, maybe

because both are members of the ICZN, HoLTHUIS (1983) and SAVAGE (1986) want to pre-

serve this first interpretation, but unfortunately they are wrong. Because of the relative im-

portance of this case as an examplar one (since it has now been discussed by several authors

and that no agreement seems to be currently reached among the supporters of opposed opin-

ions), this question is dealt with here in some detail.

In WAGLER’s times, the phrases “new replacement name” or “‘nomen novum”, with

the precise meaning they now have in the Code, did not exist. If the word “expressly” is

taken literally (i.e. if these phrases must appear as such in the text), it is clear that no name

published at the time of WAGLER, or even later, will ever qualify as a “‘nomen novum”. But

if it is taken, as suggested above, in the sense that the author must clearly show, somehow

or other, that his intention is to propose a new name for an already existing taxon, and not

for a new taxon, it will be clear that many names of this period qualify as such.

As I had already pointed out (D-18), the names proposed by WAGLER (1830 b) to re-

place already existing names are all presented in a similar way, with a footnote giving (1) the

etymology of the new name, (2) the replaced name, with its author and sometimes its date

and reference, and (3) sometimes, additional comments. What is important in my argumen-

tation (D-18) is that I stressed this similarity of presentation and considered that all pairs of

names which appear in this way in WAGLER's text (e.g., in Amphibia, Asterodactylus for Pipa,

Dendrobates for Hylaplesia, Enydrobius for Hylodes, Systoma for Engystoma, Bombitator for

Bombinator) are to be treated similarly as a couplet composed of a replacement name and of

a replaced name. Strangely however, neither HOLTHUIS (1983) nor SAVAGE (1986) discuss

this aspect of my argument. Both of them refuse to consider Dendrobates as a replacement

name for Hylaplesia, and SAVAGE (1986) further refuses to consider Asterodactylus and Bom-

bitator as replacement names for respectively Pipa and Bombinator. None of them however

discusses the cases of Systoma or of Enydrobius (for which MyERs, 1962, and LyNCH, 1971,

had already adopted the same interpretation as me).

SAVAGE’s (1986) insistance on the fact that Asterodactylus is not a replacement name for

Pipa is all the more incomprehensible to me that I had already stressed (D-21) that this name

had first been proposed by WAGLER, not in his 1830 book, but in a 1827 paper, where he

wrote : “(Asterodactylus m. Pipa Auctor)”. To refuse to interpret such a presentation as a

statement that the first of both these names is “proposed expressly as a new replacement

name” for the second one makes really no sense, since this mode of presentation for new

replacement names was very common in these times. To take examples dealing with am-

phibians, a similar presentation appears in the works of RAFINESQUE-[SCHMALTZ] (1814, 1815)

and of GISTEL (1848) (see below). If SAVAGE refuses to consider Asterodactylus as a replace-

ment name of Pipa, he should logically also refuse to consider e.g. Triturus Rafinesque, 1815

Source : MNHN, Paris

DuBols 41

as a replacement name for Triton Laurenti, 1768, or Philautus Gistel, 1848 as a replacement

name for Orchestes Tschudi, 1838 (see below). Since in these cases no diagnoses were given

for the supposedly new taxa, and no included species mentioned, Triturus and Philautus should

be considered as nomina nuda, without status in nomenclature. As a result, the valid name

for the genus currently known as Triturus Rafinesque, 1815 should become Molge Merrem,

1820, and that for the genus currently known as Philautus Gistel, 1848 should become Den-

drobatorana Ahl, 1927 (since, as will be shown elsewhere [MORÈRE & DUBOIS, in prepara-

tion], the holotype of Hylambates dorsalis Peters, 1875, type species of Dendrobatorana, be-

longs in fact to the genus Philautus). Numerous similar examples could easily be found.

SAVAGE (1986) is not embarrassed with these problems, since he does not at all consider the

consequences of his refusal to recognize that Asterodactylus was clearly proposed as a replace-

ment name for Pipa.

One may wonder why WAGLER felt necessary to replace some existing generic names

by others, coined by him. Let us note in passing that even if we had no clue to help us to

understand why he did so, that would not in the least allow us to reject the evidence, and

in this respect the discussion given by SAVAGE (1986) about the name Cacopus is completely

irrelevant : no matter if GÜNTHER (1864) was wrong in believing the name Uperodon to be

preoccupied, the fact is that he clearly and “expressly” presented Cacopus as a replacement

name for the latter! However and fortunately in this case, WAGLER himself gave us the ex-

planation. To be sure, to find this it is necessary to read Latin and German in WAGLER’s

text, but I am of the opinion that it is not possible to deal correctly with nomenclatural prob-

lems when one is unable to read, or at least decipher, Greek, Latin, German and French!!

WAGLER (1830 b : 17) gives us a first indication in a footnote concerning the mam-

malian generic name Tapirus, for which he proposes the new replacement name Rhinochoe-

rus. He then quotes LINNAEUS, as follows : “Nomina generica, quae ex graeca vel latina lin-

gua radicem non habent, rejicienda sunt. Linné Philos. bot. stud. Spreng. p. 265.” (Generic

names, which do not have Latin or Greek roots, must be rejected). It is thus clear that he

rejected as invalid all generic (not specific) names for which he did not find a root in Latin

or Greek languages. Careful examination of the whole book of WAGLER bears this interpre-

tation out : in all those cases in which an existing generic name was neither Latin, nor Greek,

1. One of the readers of the manuscript of this paper crossed this paragraph out, writing “Horrors!” in the margin

and adding: “My God, you even demand that people read French!” It may therefore be interesting Lo point out here

that, if English is certainly, at the moment, the most “international” of all languages, French is the second one in this

respect. According 10 MALHERBE (1983), the ten most “international” languages rank as follows: (1) English, official

or “privileged” language in 47 countries; (2) French, 26 countries; (3) Arab, 21 countries; (4) Spanish, 19 countries;

5) Portugese, 7 countries; (6) German and Swahili, 5 countries; (8) Dutch, 4 countries; (9) mandarin Chinese and

talian, 3 countries. The six official languages of the United Nations are English, French, Spanish, Russian, Arab

and mandarin Chinese. If the number of persons in the world who speak a given language as their mother tongue is

considered, English, with 320 millions, is only second, after mandarin Chinese (700 millions) and before Hindi-Urdu

(280 millions), Spanish (190 millions) and Russian (160 millions).

In fact, in one disagrees with my demand that z0ologists should be able to “read, or at least decipher” French,

this simply means that only one language in the world, English, is considered worthy of being known by them! I

would not support such a statement, especially when one deals with disciplines, like systematics and nomenclature,

in which old texts of the XIX century and even before are still in current use: in these times the leadership of English

in science was not established, and many important works were written in French, German, Latin and other lan-

guages. Nowadays, if 67% of the scientific publications in the world are in English, 8% are in French, which

be considered trivial. In recent years, other languages, especially Spanish, have had a growing use in some

like zoology, and there is no reason to believe that this movement will stop. Ignorance of all the non-English s

literature cannot be profitable to any scientist in the world, since ignorance has never been useful to anyone.

Source : MNHN, Paris

42 ALYTES 6 (1-2)

nor based on Latin or Greek roots, WAGLER proposed a new replacement name. In some

cases, he even expressed his perplexity as to the possible etymology of a generic name, which

for this reason he replaced by a nomen novum. Thus, the name Systoma is proposed to re-

place Engystoma, which is presented as follows in a footnote on p. 205: “Gen. Engystoma

(quid?) Fitzing.”.

After WaGLER’s death in 1832, MICHAHELLES (1833) reproduced in sis von Oken an

unpublished manuscript of WAGLER, which had been written before the 1830 book of this

author. In this text we can find additional explanations of the reasons why WAGLER rejected

some generic names as invalid. In a footnote (in MICHAHELLES, 1833 : 888), WAGLER states

that he found no evidence of the use of the names Hyla and Calamita in classical Latin texts,

and that for this reason these names must be rejected. He states that he found the name

Calamites in PLINIUS, as the name of the common European treefrog. He adds that unfor-

tunately FITZINGER had already given the name Calamita, “(das in Calamites umgeändert

werden mu)” (which must be changed into Calamites) to another genus of frogs, and that,

in order not to upset FITZINGER’s work, he refrains from using the name Calamites for the

European treefrog. In consequence he proposes the new remplacement name Discodactylus

for Hyla, the latter not being of classical Latin or Greek origin. This footnote clearly shows :

(1) that, as a general rule, WAGLER did propose new replacement names for generic names

considered by him invalid, because they were not of strictly classical Greek or Latin origin

(i.e. a name used as such in classical Latin or Greek, or a name based on classical Latin or

Greek roots) ; (2) that nevertheless WAGLER made clearly the distinction between new names

for already known taxa, and for new taxa, and that he respected the works of previous au-

thors (his refusal to use the name Calamites for a different genus than that called Calamita

by FITZINGER — after SCHNEIDER, 1799 — is a rare example, for this epoch, of respect of the

works of others) ; (3) more specifically, that SAVAGE’s (1986) analysis of the status of the names

Calamites and Dendrobates is in error.

This leads us now to discuss the case of the unjustified emendations of generic names

proposed by WaGLER (1830 b), such as Bombitator, Megalophrys or Calamites. The name

Calamites is expressly presented by WAGLER (in MICHAHELLES, 1833) as a replacement name

for Calamita, and could either be considered as a nomen novum or as an unjustified emen-

dation of this latter name (see discussion above). Strangely, SAVAGE (1986) considers it as

the name of a new genus, overlooking the fact that WAGLER (1830 b : 200) expressly wrote :

“CALAMITES (...) Fitzing.”, thus clearly acknowledging FITZINGER as the author of the taxon,

even though he emended his name. In reality, in SAVAGE’s logic, if Calamites is not accepted

as a replacement name or an unjustified emendation for Calamita, it should not be consid-

ered as the name of a new genus (since it is credited to a previous author), but as an incor-

rect subsequent spelling, without status in nomenclature. It is true that the spelling Cal-

amita does not appear in WAGLER’s (1830 b) book, but it appears in the manuscript of

WAGLER, anterior to this 1830 book, which was later published by MICHAHELLES (1833). 1

contend that this case is a typical example of type (C3) of evidence for emendations pre-

sented above, which is also acknowledged in Art. 33(b)(i) of the new Code.

SAVAGE’s (1986) too rapid way of working has another, quite funny indeed, conse-

quence. SAVAGE (1986) states that Calamites is not an unjustified emendation of Calamita,

but the name of a new taxon. He adds that FITZINGER’s (1843) designation of Hyla cyanea

as type species of Calamites, rediscovered by Dupois (D-26 : 19), is valid, but he does not

Source : MNHN, Paris

Dugolis 43

discuss the consequences of this fact : these would be that Calamites Wagler, 1830 would be

a senior synonym of the well-known and much used generic name Lütoria Tschudi, 1838!

Fortunately, Calamites Wagler, 1830 is preoccupied by Calamites Guettard, 1770 (see D-26 :

14), so that, even if SAVAGE (1986) was right, no nomenclatural disruption would result.

However, these facts and their consequences should all be considered and discussed when

such question are tackled : this clearly shows that in nomenclature a very slight divergence

of opinion as to the status of a name may have considerable consequences in the long run.

As for the name Bombitator, SAVAGE (1986) is still less excusable to consider it as the

name of a new genus, because WAGLER (1830 b) mentions the name “Bombinator Merr.” in

a footnote on p. 206 and in the index on p. 346. In Latin, the name “bombitator” (bee) only

did exist, while the names “bombina” and “bombinator” did not exist but were coined by XVIII

and XIX century authors on the basis of the name “bombus” (buzzing). That Bombitator can-

not either be considered as a fortuitous incorrect subsequent spelling is also shown by the

fact that this name alone (and not Bombinator) appears on several occasions in WAGLER’s book

(pp. 132, 206, 294, 296, 301, 302, 303, 305, 306, 346). It is thus clear that the spelling Bom-

bitator was purposedly used by WAGLER instead of the spelling Bombinator, but for the same

taxon as MERREM (1820). If this is not an unjustified emendation, I wonder which name of

this epoch will qualify for this category.

To come back finally to the name Dendrobates now, it was clearly presented by WaG-

LER (1830 b) as a replacement name for Hylaplesia because the latter name was not accept-

able according to WAGLER’s conceptions, being based on the root “Hyla” which was rejected

by WAGLER (1830 b ; in MICHAHELLES, 1833) as invalid since it did not exist in classical Latin.

The fact that WAGLER (1830 b) only recognized some of the species originally included in

Hysaplesia (and Hylaplesia) by BoIE (in SCHLEGEL, 1826, 1827) is of no relevance here, be-

cause, as remembered above in the dichotomic key under the heading II, an author may per-

fectly, while accepting a taxon created by a previous author and crediting it to him, modify

in part the diagnosis or content of the taxon : by doing so he does not create a new taxon,

because otherwise any new modification of the diagnosis or content of a taxon would result

in the creation of a new taxon and of a new name, and no stability of taxinomy and nomen-

clature would ever be possible.

As for the status of the name Hylaplesia itself, it is certainly open to discussion. It might

be possible to consider it very formally as an incorrect subsequent spelling. However, as I

have shown (D-18, D-21), this name has an etymological justification (being based on Hyla),

while Hysaplesia has none and is clearly the result of a misprint for Hylaplesia. Since the

spelling Hylaplesia has been used by various authors since its creation, while the spelling

Hysaplesia has remained ignored until the paper by STEJNEGER (1937) where it was resur-

rected, this name, on which is based the family-group name Hylaplesidae, is better consid-

ered as having an independent status in nomenclature (for more detailed discussions of other

similar cases, see D-17). It is therefore justified to consider Hysaplesia Boie in SCHLEGEL,

1826, which was in reality clearly a misprint in the original publication, as the “correct orig-

inal spelling” of the name in the sense of the Code, and Hylaplesia Boie in SCHLEGEL, 1827

as an unjustified emendation of the latter. Finally, Dendrobates Wagler, 1830 is without pos-

sible doubt a new replacement name for Hylaplesia. Therefore the problems raised by Du-

BOIS (D-18) concerning the validity of the names Dendrobates and Dendrobatidae remain,

and must be solved by an action of the ICZN.

Source : MNHN, Paris

44 ALYTES 6 (1-2)

In conclusion the generic names Asterodactylus, Dendrobates, Enydrobius, Systoma,

Bombitator, Calamites and Megalophrys are all new replacement names (or, if one prefers, un-

justified emendations, for the last three ones) for existing generic names which were believed

by WaAGLER (1827, 1830 b) to be invalid because they were not of strict classical Latin or

Greek origin. They have therefore by definition the same type species as the replaced names,

and SAVAGE’S (1986) analysis is in error.

OTHER PROBLEMS RAISED BY SAVAGE (1986) IN GENERIC NAMES

(1) Cacopus Günther, 1864. - As already tackled above, the analysis of this case pre-

sented by SAVAGE (1986 : 261) is clearly in error. He states that Cacopus is a replacement

name for Hyperodon, “an incorrect subsequent spelling (which has no status) as Uperodon is

an available name”. SAVAGE (1986) clearly mingles two very different things : the fact that

an author has intentionally introduced a new spelling (emendation) or a new name (nomen

novum), because he believes, for some reason, that the original name is incorrect or invalid ;

and the fact that this action is or not justified according to our current rules. It is perfectly

true that Uperodon Duméril & Bibron, 1841 is an “available name” and that, according to

the 1985 Code (and also to the preceding editions), it does not have to be emended. But this

has no bearing on the question whether GÜNTHER (1864) intentionally decided to replace it!

As a matter of fact, GÜNTHER (1864 : 415) did clearly and explicitly state that he did so and

why : “The correct spelling of this word would be Hyperodon, a name long previously given

to a genus of Cetaceans. Besides, these frogs have no vomerine teeth.” This means that : (1)

GÜNTHER (1864) considered (in error, but this needs not concern us here) the spelling Uper-

odon to be invalid because incorrectly formed, in his opinion, from the Greek root ; (2) he

stated that the correct spelling of this name should be Hyperodon ; (3) but he immediately

added that this latter name was also invalid, being preoccupied ; (4) furthermore, he added

that this name was inappropriate for the frogs in question, because it did not correspond to

the real biological properties of these animals ; (5) consequently, he proposed the replace-

ment name Cacopus for this genus. Only two interpretations of this case are possible. The

first one, which I had proposed (D-26), is simply to consider Cacopus as a replacement name

for Uperodon. As a matter of fact, (1) GÜNTHER clearly considered this latter name as invalid,

and (2) he only mentioned the spelling Hyperodon as the potential correct spelling of this

name, but not as a valid name. However, if one wishes strictly to follow the very formal

proposal of SAVAGE (1986) that Cacopus be considered a replacement name for Hyperodon,

then the latter name must be considered an unjustified emendation of Uperodon, that must

be credited to GÜNTHER (1864), but cannot in the least be viewed as an incorrect subsequent

spelling! This much more formal solution, although it does not change the final result, is

less logical, because it credits GÜNTHER (1864) with a name which he himself rejected in the

very work where he created it.

(2) Philautus Gistel, 1848. - Concerning this name, SAVAGE (1986 : 261) writes : “I have

not seen the original descriptions (sic) of the latter name but suspect it does not qualify as

a new replacement name.” This statement sounds quite strange to me.? Personally, when I

2. I am even surprised that the journal Copeia should have published this sentence

Source : MNHN, Paris

Duois 45

have not seen a publication, I either try to obtain and see it by myself, or, when this proves

impossible, I use second-hand information. It is true that such information may sometimes

be in error (and this is the reason why I always try my best to obtain the original paper),

but, at least, it is better than no information at all. SAVAGE (1986) seems to act differently.

He apparently does not try seriously to see the original publication (for if he had, he cer-

tainly could have succeeded, for example by writing, as I have done in this case, to the Brit-

ish Museum library), and he a priori mistrusts the information given in a recent paper by a

colleague, who furthermore stated that he had seen the original paper (since, as is easy to

verify, I always state in the bibliographies of my papers when I have not seen a given pub-

lication). This is a strange attitude. Disappointing as it may be for SAVAGE, ÎÏ must precise

that here again he is in error. To avoid repetition of this error, I am obliged to give here

more details about GISTEL’s (1848) work. That may be useful to all batrachologists.

In the introduction of his book, GISTEL (1848 : viii-xi) presents a very long alphabet-

ical list of generic names, for most of which he proposes new replacement names. The way

names are presented may be illustrated by the following example, at the beginning of the

list : “Acanthoderus (Palisot de Beauvois. Wanze : Lygaeus sanctus) bleibt ; aber Acanthoderus

(Serv. Cerambycid.) heifit : Scamillus (Nob.).” This means that the first genus name cited

stands (bleibt), while the second one is replaced by another one, and is (re)named (heifit)

Scamilius. The words “bleibt” and “heifit”, which appear in the first examples, are either

abbreviated (“b1.” for “bleibt”) or omitted (“heift”) in the following of the text ; the word

“Nobis” is abbreviated first in “Nob.”, then in “N.”. Other replacement names are also pro-

posed in a slightly different manner in footnote 19 of p. xi.

As far as the name Philautus is concerned, it appears as follows on p. x of GISTEL

(1848) : “Orchestes (Ilig. Käf.) bl. - Orchestes (Tschudi, Isis 1838. 853.) : Philautus, N. -

Orchestes (Costa Cenni, Crustac.) : Encopis, N.” This is a clear indication that Philautus Gis-

tel, 1848 is proposed as a new replacement name for Orchestes Tschudi, 1838, because the

latter is preoccupied by Orchestes Illiger, 1798 (and by Orchestes Leach, 1830) (see D-11).

In view of the rarity of GISTEL’s (1848) text, I felt it useful to prepare and give here a

complete list of the replacement names proposed in this book for amphibian genera (the same

work should also be done for other groups), most of which have not been recently cited (see

Table 11). These names should be included in the synonymies of their respective genera, as

was done by DuBois (D-11) in Ranoidea. Apart from Philautus, the only one of these names

to have been considered to be valid in the recent years is Hydromantes ; however this name

was recently shown to be invalid (D-24).

(3) Astrodactylus [Hogg, 1838]. - Contrary to SAVAGE (1986), this name is an unjustified

emendation of Asterodactylus Wagler, 1827, not an incorrect subsequent spelling, by virtue

of criteria (C2), (C4) and (C6) above. As a matter of fact, both this spelling and the original

one have an etymological jusitification : Astrodactylus is based on the Latin word astrum, de-

rived from the Greek &orpov, while the original spelling Asterodactylus was based on the Latin

word aster, derived from the Greek &orñp. Both astrum and aster mean “star”, and this re-

fers to the shape of the extremities of the fingers of these animals. Besides, in all his papers,

HoGG (1838, 1839 a, 1839 b, 1841) always used the spelling Astrodactylus (and the derived

spelling Astrodactylidae) for the name of these animals, and never reverted to the original

spelling Asterodactylus. However H0GG was clearly aware of the original spelling, which he

Source : MNHN, Paris

46 ALYTES 6 (1-2)

Table 11. - New replacement names for generic names of amphibians created by GISTEL

(1848).

New replacement name Page in Original generic name Recent reference

GISTEL (1848) to GISTEL'S

(1848) name

Barvboas xi Pelophilus Tschudi, 1838

Borborocoites xi Palaeobatrachus Tschudi, 1838

Bradytes xi Bradybates Tschudi, 1838

Buccinator xi Boophis Tschudi, 1838 D11

Cotobotes xi Hemidactylium Tschudi, 1838

Dendricus vi Buergeria Tschudi, 1838 D-11

Dendromanes xi Microhyla Tschudi, 1838

Dendromedusa xi Hylaplesia Boic in SCHLEGEL, 1827

Epipole ix Eucnemis Tschudi, 1838 D-11

Hydromantes xi Geotrion Bonaparte, 1832 D-24

Hydroscopes xi Pseudosalamandra Tschudi, 1838

Limnarches xi Ambystoma Tschudi, 1838

Pelida xi Hyladactylus Tschudi, 1838

Pelodytes xi Peudorriton Tschudi, 1838

Philautus x Orchestes Tschudi, 1838 D-11, D-26

Phyllodytes xi Comnufer Tschudi, 1838 D-11

Polvphone xi Ranoïdea Tschudi, 1838

Pyleus xi Pseudobufo Tschudi, 1838

Tritogenius xi Andrias Tschudi, 1837

Troglobates xi Palacophrynos Tschudi, 1838

Xiphoctonus xi Xiphonura Tschudi, 1838

Zoodioctes xi Hylarana Tschudi, 1838 D-11

cited when he commented (HOGG, 1839 a : 268) on WAGLER's (1830 b) book where this

spelling appears. There can therefore be no doubt about the intentionally of his act.

(4) Lophiohyla Miranda-Ribeiro, 1926. - Quite funnily, SAVAGE (1986 : 261) writes

concerning this name : “Lophiohila (sic) Miranda-Ribeiro, 1926 is an incorrect subsequent

spelling.” In reality, the spelling which appears in MIRANDA-RIBEIRO’s (1926) work is not

Lophiohila, but Lophiohyla. This name must be considered an unjustified emendation of the

original spelling Lophyohyla (for more detail, see D-26 : 21-22), by virtue of criterion (CS):

this name appears several times in MIRANDA-RIBEIRO’s (1926) work, and again once in a sub-

sequent paper (MIRANDA-RIBEIRO, 1937), while the original spelling of the names does not

appear any more in these texts. This name is clearly based on Hyla, but the etymology of

the element lophio is unclear to me : it seems to be derived from the Greek \ogt& (mane),

which does not make sense to me. However, there can be no doubt about the intention of

MIRANDA-RIBEIRO to use the spelling Lophiohyla for this genus, and the spelling must be

considered an unjustified emendation. As for the spelling Lophiohila, cited by SAVAGE (1986),

itis a new spelling. I admit however perfectly that it is to be considered as an incorrect sub-

sequent spelling, without status in nomenclature, just like the three other new and incorrect

spellings which appear in various parts of his paper (Cycloramphis, Phrynobatracinae, Petro-

pedetine).

(S) Myiobatrachus [Bonaparte, 1850]. - Contrary to SAVAGE (1986), this name must be

considered an unjustified emendation of Myobatrachus Schlegel, 1850, not an incorrect sub-

Source : MNHN, Paris

Dugois 47

sequent spelling, by virtue of criteria (C4), (C5) and (C6). As a matter of fact, both this

spelling and the original one have an etymological justification : Myobatrachus is based on

the root myo-, derived from the Greek ès, prués (rat, mouse ; muscle) ; Myiobatrachus is

based on the root myio-, derived from the Greek puia (fly). Besides, BONAPARTE (1850, 1852

b) used the spellings Myiobatrachidae and Myiobatrachina in two different papers (see D-

26 : 22), and the spelling Myiobatrachus was accepted as valid by the very author of the orig-

inal name Myobatrachus, SCHLEGEL himself (1858).

(6) Cyclorhampus Agassiz, 1846. - This spelling is considered by SAVAGE (1986) as an

incorrect subsequent spelling of Cycloramphus Tschudi, 1838, but it is in fact an unjustified

emendation of this latter name, as shown by types (C2) and (C4) of evidence. In TscHUDrs

(1838) text, this generic name was spelled Cycloramphos on p. 41 and Cycloramphus on p. 81.

AGassiZ (1846 : 110) cites both these spellings, and, in front of each of them, writes “C-

rhamphus” between parentheses. This is a clear indication that he considered Cyclorhamphus

as the only justified spelling for this name, which should in his opinion replace both original

spellings. The reason for that is clearly the recourse to the etymology, this name being based

on the Greek word pépos (beak), which is correctly latinized into rhamphus.

(7) Alytes Wagler, 1829. - For this name, SAVAGE (1986 : 261) most strangely writes :

“the type species of Alytes Wagler, 1829, is Bufo obstetricans Laurent (sic), 1768, by indi-

cation (Art. 12b.5)”. This is perfectly true, but by no means contradictory to what I had

written (D-26 : 18), which SAVAGE does not seem to have understood. I therefore give here

a translation of the relevant part of my discussion on this question : “In the first text where

the generic name Alytes appears (WAGLER, 1829 : 70), this name is accompanied by no de-

scription or diagnosis, but is presented in the combination Alytes obstetricans. The name Bufo

obstetricans Laurenti, 1768 being at this epoch the only existing specific name which could

apply to this taxon (see SHERBORN, 1902 : 682), one may consider that the name Aytes is

available as from 1829, with Bufo obstetricans Laurenti, 1768 for type species by monotypy.

One could also consider this indication as insufficient, the name obstetricans not being de-

fined by the mention of his author and date : in this case, Alytes would be a nomen nudum

in WAGLER (1829), and would only obtain a status in nomenclature with the publication by

WAGLER (1830 a : [53], pl. XXII) of a description and of figures showing the species Alytes

obstetricans (Laurenti, 1768).” (D-26 : 18). My concern here was whether the specific name

obstetricans mentioned by WAGLER (1829) was to be considered as a new name, which would

then be a nomen nudum, or as a subsequent use of the specific name Bufo obstetricans Lau-

renti, 1768. It is true that the subsequent publications of WaAGLER (1830 a, 1830 b) dem-

onstrated without ambiguity that it was this latter name which had been used by WAGLER

(1829), but, had these subsequent works never appeared, there would be no evidence at all

in the original paper itself (WAGLER, 1829) for this interpretation. The only evidence which

could be used at this date was the fact that Bufo obstetricans Laurenti, 1768 was the only spe-

cific name obstetricans which existed then in the whole animal kingdom. These are subtle

matters, I admit, but which should not discourage a truly serious student of nomenclatural

problems.

(8) “Elosia nasuta Tschudi, 1838”. - This is the last of the examples cited by SAVAGE

(1986 : 261) to demonstrate that “it behooves any user of Dubois’ paper to review the orig-

inal literature to determine if a particular name is an emendation or an incorrect spelling.”

While I am not embarrassed by this suggestion, I think this example clearly shows that the

Source : MNHN, Paris

48 ALYTES 6 (1-2)

batrachologists interested in suprageneric nomenclature of amphibians should consult my

work directly, rather than SAVAGE’s (1986) comments upon it, which are based on an ap-

proximate (and sometimes completely erroneous) understanding of its content. As a matter

of fact, SAVAGE (1986 : 261) writes : “Other corrections that need to be mentioned regarding

the genera discussed by Dubois are : (...) d) the name Elosia nasuta Tschudi, 1838, is an

incorrect subsequent spelling.” Now, here is the exact translation of my own comment on

this question : “GORHAM (1966 : 111) and LyNCH (1971 : 166) believe that there exists a

nominal species Elosia nasuta Tschudi, 1838. In fact, the binomial Elosia nasuta which ap-

pears in TscHuprs (1838 : 77) work is clearly a new combination proposed for the nominal

species Hyla nasus Lichtenstein, 1823. The spelling nasuta used by TSCHUDI (1838) for the

specific name cannot be held to be an unjustified emendation of the name nasus Lichten-

stein, 1823, but must be regarded as an incorrect subsequent spelling, devoid of nomencla-

tural status (the same is true for the spelling nasulus, which also appears in TscHUDrs 1838

work).” I think this needs no additional comment.

VALID FAMILY-GROUP NAMES

While SAVAGE (1986) tends to favor a very strict, restrictive and even rigid conception

of the Code regarding the status of new replacement names, he adopts the reverse attitude

concerning the validity of family-group names. In this case he clearly praises a return to pre-

Code rules, since he does not admit the principle or priority for these names. As a matter of

fact, he challenges the use of some family-group names which I consider valid on the basis

of this principle. In particular, he suggests that Dactylethrinae Hogg, 1838 should be re-

placed by Xenopodinae Fitzinger, 1843, and that Tornieriobatinae Miranda-Ribeiro, 1926

should be replaced by Nectophryninae Laurent, 1942. He seems also, though less clearly,

to suggest that Petropedetinae (which he qualifies as “widely used”) should be given priority

over Phrynobatrachinae. Although this point is only slightly tackled by SAVAGE (1986) in this

paper, it is of importance, because it is clear that SAVAGE’s opinion in this respect was ac-

cepted or shared by the editors of ASW, and one may in fact find in SAVAGE’s (1986) paper

the “theoretical” justification for some of the decisions taken in ASW concerning the valid

names of some family-group taxa, which I already commented upon elsewhere (D-41).

As I already pointed out (D-26 : 6), MyYERS & LEVITON (1962 : 290) rightly stressed

that one of the causes of instability of familial nomenclature in zoology stems from the fact

that several conceptions have existed as to the rules which should be followed to determine

the valid name of a family-group taxon. Thus, according to the various “rules” followed in

the past by different authors, the name retained as the valid one may have been : (R1) the

family-group name based on the oldest available genus-group name belonging to the family-

group taxon in question ; (R2) the family-group name based on the oldest valid genus-group

name of the family-group taxon in question ; (R3) the first family-group name to have been

used with a correct spelling (suffix in - idae or - inae) ; (R4) the ofdest family-group name ever

proposed for this group, even when it was then incorrectly formed (suffix in -ae, -ida, -ina,

-ides, etc.). The experience shows that the use of rules (R1) and (R2) leads to catastrophic

consequences as 10 the stability of family-group names. As a matter of fact, under such rules,

very limited taxinomic changes may lead to a change of name for the family-group taxon :

Source : MNHN, Paris

DuBois 49

in case (R1) this may be caused by the simple addition to or substraction from this taxon of

a genus the name of which is older than all the other generic names of the group ; in case

(R2) this may be caused by the simple fact that a generic name, once considered valid, be

rejected as a junior homonym or (subjective or objective) synonym, or on the reverse retired

from synonymy. Rule (R3) would not do justice to the fact that in older times there was no

rule as to the correct spelling of family-groups names and that nevertheless authors of this

epoch, although using spellings which are currently considered incorrect, clearly created

family-group names and should be credited with the authorship of these names. Rule (R4)

is the one that ensures the highest stability possible for family-group names and credits these

names with their proper authors. As I had stressed in my 1984 paper (D-26), the rules then

in force according to the second edition of the Code (ANONYMOUS, 1964, 1974) were excel-

lent, because they allowed a maintenance of the stability of family-group nomenclature even

when minor changes were introduced in the generic content of a family-group taxon or in

the status of some generic names. Unfortunately, as commented above in detail, the changes

brought to Art. 32, 35 and 39 in the 1985 Code are highly open to criticism and should in

my opinion be reconsidered.

Some comments are necessary here concerning Art. 40 of the Code, which is called

upon by SAVAGE (1986) and also in ASW to justify certain decisions taken by these authors.

The relevant parts of this Article are as follows :

“Article 40. Synonymy of the ivpe genus. -

(a) After 1960. - When, after 1960, the generic name on which a valid family-group name is based

is rejected as a junior synonym, that family-group name is not to be replaced (...).

(b) Before 1961. - If a family group name has been replaced before 1961 because of such syn-

onymy, and the replacement name has won general acceptance, it is to be maintained.

@.)

(ii) in the event of divergent interpretations of the expression “general acceptance”, the case is

10 be referred to the Commission for a decision.” (ANONYMOUS, 1985 a : 81).

Some comments on this article seem appropriate here. First of all, it is clear that this

article was devised in order to limit the nomenclatural disturbances which would be entailed

by a too strict application of the principle of priority in the family-group, since in the past

many authors in fact had followed the rule (R2) above. However, this partial suspension of

the principle of priority to family-group names is limited in time, since it applies to actions

prior 10 1961. The aim of setting this limiting date is clear : it is to ensure that, from 1961

on, the principle of priority strictly applies to family-group names, as well as to genus-group

and species-group names. It should therefore be clear that this article only applies to family-

group names which have been proposed and have wvon general acceptance before 1961, and can-

not even be considered as potentially applicable to names either proposed or generally ac-

cepted after that date. Thus for example this article cannot in the least be called upon to

conserve the name Ichthyophiidae Taylor, 1968, which was proposed after 1960 and is a jun-

ior synonym of Epicriidae Fitzinger, 1843. It could not either be called upon to conserve

Bombininae Fejérväry, 1921, which has been used only once before 1961 (D-37) and is a

junior synonym of Bombinatorinae Gray, 1825, a name which has had a large use in the XIX

century, and is the valid name of the subfamily, if one wishes to recognize it as a distinct

taxon (D-19, D-26, D-29, D-35, D-39).

Source : MNHN, Paris

50 ALYTES 6 (1-2)

A second comment is that Art. 40 only applies to the case where a family-group name,

based on a generic name rejected as a junior synonym, is replaced by a family-group name

based on the generic name which is the senior synonym of the rejected generic name (as in

the example given in Art. 40 of the Code). Thus this article could not apply in the case of

Petropedetinae versus Hemimantinae, even if, which is far from being true, the name Pe-

tropedetinae had “won general acceptance” before 1961.

Finally, it is important to note that in the case of disagreements between different au-

thors as to the fact that a name has or not “won general acceptance”, the ICZN alone is

enabled to take a decision. In other words, in such cases it is the responsability of those who

believe, contrary to others, that Art. 40 may be called upon to conserve a more recent name,

to submit an application to the ICZN and give evidence that this recent name has “won gen-

eral acceptance”. Unless they do it, the principle of priority must apply in these cases as in

general.

The proposals of SAVAGE (1986) may now be considered in the light of these com-

ments.

The case of Dactylethrinae versus Xenopodinae was already discussed elsewhere (D-

41). I contend that the name Xenopodinae was only very rarely used before 1961, and even

before 1986, and that it cannot be considered as having “won general acceptance”. The name

Dactylethrinae, which has been more largely used than this latter name and which has prior-

ity, should be conserved for this taxon. I feel it useless to discuss this problem in more detail

here and to give lists of bibliographic references in support of my assertions as long as no-

body deems it justified to submit officially this problem to the ICZN.

The case of Petropedetinae was also discussed elsewhere (D-41). SAVAGE (1986) qual-

ifies this name as “widely used”, but does not produce evidence for this statement : as shown

elsewhere (D-41), I can list 11 references to use of this name, versus 35 references to use of

Phrynobatrachinae. Furthermore, Art. 40 cannot be called upon in this case, for the reasons

developed above, and an action of the ICZN is necessary here if the name Hemimantinae is

to be rejected.

Finally, the most striking case is that of the names Tornieriobatinae and Nectophry-

ninae. SAVAGE (1986 : 261) writes : “Art. 40b is marginally applicable here since the subfam-

ily has been rarely recognized, nevertheless I would favor use of Nectophryninae.” Actually,

it is misleading to say that the subfamily “has been rarely recognized”, since I personally

know only two publications where it was recognized before my own works (D-14, D-19, D-

26, D-29, D-35) : that of MIRANDA-RIBEIRO (1926) where he created the name Tornieriob-

atinae, and that of LAURENT (1942) where he created the name Nectophryninae! This latter

author himself seems to have never used again this name, or recognized this subfamily, in

subsequent publications. Even if I missed a few papers where these names were used, it is

clear that it is impossible to speak in such a case of “general acceptance” before 1961, or

even before 1986. Art. 40 is therefore not applicable at all in this case, not “marginally ap-

plicable”, and the name Tornieriobatinae must remain the valid name of the taxon, if the

latter is to be recognized.

This last example has the merit to clearly tell us what is the philosophy of SAVAGE con-

cerning the validity of family-group names. He in fact favors the rule (R2) above, and con-

siders that, as far as possible, one should suppress family-group names which are based on

Source : MNHN, Paris

Dugois si

generic names currently considered junior synonyms of valid names. He then tries to use

Art. 40 for this purpose, and not for the real purpose of this article, which is to protect the

stability of nomenclature by conserving well-known and widely used names. It is also clear that

itis for this same reason that SAVAGE (1973 : 354) used the name Platymantinae for the taxon

named Cornuferinae by NOBLE (1931 : 521), after the ICZN had decided, in a very disput-

able Opinion, to consider the name Cornufer as a junior synonym of Platymantis (for a more

detailed discussion of these points, see DuBois, D-11).

The exactly same philosophy is adopted in ASW, and I now understand better, in the

light of SAVAGE’s (1986) paper, some of the choices made in this list (concerning e.g. the

names Xenopodinae and Petropedetinae), or the strange statement on p. 105 of ASW, which

had elicited my surprise (see D-41), that Alytidae was the first available name for the family

currently known as Discoglossidae, while I had clearly shown that the first available name

for this taxon was in fact Bombinatoridae : it is because, this latter name being based on a

junior synonym, it was simply considered by the authors of ASW as virtually non-existant!

However, as I discussed in detail elsewhere (D-26, D-37), these matters are not so simple,

and such practices cannot be accepted.

The refusal of respecting the principle of priority for family-group names has impor-

tant consequences. It is almost equivalent to a refusal of accepting that family-group names

be ruled by the /nternational Code of zoological Nomenclature, and is therefore a step back-

ward, contrary to the tendency which has always been increasing to elaborate and use precise

rules for the use of names in systematics. It should therefore not be accepted by taxinomists

who wish to work for a stable system of nomenclature of living beings.

Incidentally, this case also throws a light on another type of errors which had struck

me in ASW, and which is mentioned elsewhere (D-41), but the reasons for which I had not

really understood. Thus for example this list states that the type species of the genus Lep-

todactylus is Rana fusca Schneider, 1799, while it is in fact Rana typhonia Latreille in SON-

NINI & LATREILLE, 1801. The reason of this mistake is the same as that just discussed for

family-group names : since this latter name is currently considered a subjective junior syn-

onym of the former one (HEYER, 1968 ; LyYNCH, 1971), the authors of ASW replaced the

true name of the type species of the genus by the valid name of this species. This clearly

shows a confusion between the concepts of nominal species and of biological species : two

different nominal species may be considered to be the same biological species, they never-

theless remain distinct nominal species, and the type species of genera are nominal species,

not biological species!? À symmetrical case is that e.g. of Necturus, where the reverse mistake

was done : ASW considered FITZINGER’s (1843) designation of “Nectur. lateralis. Wagl.” (in

fact Triton lateralis Say, 1823) as type species of this genus to be valid because this name is

currently considered a subjective synonym of Sirena maculosa Rafinesque, 1818. However,

the nominal species Triton lateralis was not part of the originally included species of Necturus

Rafinesque, 1819, and is not eligible for type fixation in this genus. Therefore, BROWN (1908)

3. This is not understood e.g. by ZHAO (1984), who states that the type species of the genus Liua Zhao & Hu, 1983

Should move from Ranodon wushanensis Liu, Hu & Yang. 1960 to Hynobius shihi Liu, 1950, since the former is à

synonym of the latter: both nominal species may be synonyms (RisCH & THORN, 1982), the former nevertheless re-

mains the type species of the genus Liua, by original designation. Let us note also that the name Liua is mispelled

Liuia in ASW (pp. 565-566).

Source : MNHN, Paris

52 ALYTES 6 (1-2)

is the author of the valid type species designation (see Table VII in D-41). Other mistakes

of this type may exist in ASW, where I have not looked especially for them.

CLASSIFICATION USED

This last point deserves little comment, except that it is in error that, concerning the

few changes that I had proposed (D-19, D-26, D-29) to LAURENT’s (1980 a) classification

scheme, SAVAGE (1986 : 262) writes : “The bases for these decisions are not discussed.” It

should be clear that my paper (D-26) was dealing with nomenclatural problems, not with

classification itself. However, nomenclature is meaningful and can be used only on the basis

of a sification, and I had to choose one, even if I did not ignore that it could not be the

“definitive” one! I chose LAURENT’s (1980 a, 1986 a) scheme and made clearly reference to

it. In the few cases where I introduced (slight) changes, in particular in the recognition of

subfamilies, I clearlÿ gave references to works justifying these choices (Discoglossidae, pp.

24-26 ; Pipidae, p. 27 ; Pelobatidae, pp. 28-29 ; Bufonidae, pp. 33-35). I should stress that

in all these cases I proposed these changes as tentative, but only using already existing names,

in order not to create new taxa or names. In one case I even used voluntarily an artifice in

order not to create a new name. This is the case of the subfamily I suggested to recognize

for East Asian bufonids (including the genera Ansonia, Bufoides, Leptophryne, Pedostibes, Pe-

lophryne, Pseudobufo). No family-group name, based on one of these family-group names or

on a synonym of one of them, exists at present, and I could certainly have created one. How-

ever, since we clearly still lack much information to prove that this group is really homo-

phyletic, I refrained from doing so, and used the following artificial solution. There cur-

rently exists a family-group name Adenomidae Cope, 1860, based on the generic name

Adenomus Cope, 1860. The type species of this nominal genus is Adenomus badioflavus Cope,

1860, à name which currently stands in the synonymy of the name Bufo kelaartii Günther,

1859. Finally, INGER (1972 : 360) has suggested that this latter species, from Ceylon, might

have to be retired from the genus Bufo and placed in a genus of its own. I therefore proposed

to provisionally use the name Adenominae Cope, 1860 for the subfamily of East Asian bu-

fonids, pending further studies on the statuts of Bufo keluartii and of the other species of

this group. This action has the double advantage of avoiding the premature creation of a

family-group name for a poorly known group, but also to draw the attention to the interest

of a study of relationships within this group and between this group and other bufonids, and

I hope it will prompt studies in this field. This is clearly a more economical and prudent

attitude than that of many other authors, not only in the past but also in recent years, like

e.g. SAVAGE (1973), who proposed two new family-group names (Allophrynidae, Platyman-

tinae) for taxa which were not even defined or diagnosed, and the validity of which was not

at all demonstrated in his work (actually, the second of these taxa is clearly heterogenous

and invalid, at least as proposed by SAVAGE ; see D-2, D-11, D-35).

CONCLUSION

After this detailed analysis, [ regret to say that I must reject as wrong all the correc-

tions proposed by SAVAGE (1986) to my list of suprageneric and generic names of anuran

Source : MNHN, Paris

Dugois 53

amphibians (D-26), and that I reject his statement that “the materials (especially the generic

lists) presented [by Dubois] require considerable modification.” This does not mean that this

latter work contained no mistakes or omissions, but these were all missed by SAVAGE (1986).

Since my work was submitted for publication, and apart from the changes made necessary

by the new Code, which were “corrected” elsewhere (D-29), I have found 7 errors and 5

omissions in it. Four of the errors concern the first use of subsequent spellings of family-

group names : the spelling Centroleninae was first used by BRATTSTROM (1957 : 73), not LUTZ

(1969 : 276) ; the spelling Phrynomeridae was first used by PARKER (1932 : 1241), not PAR-

KER (1934 : 9) ; the spelling Ranini was first used by BONAPARTE (1839 : [225]), not BRONN

(1849 : 684) ; the spelling Ranidi was first used by BONAPARTE (1837 : [248]), not ACLOQUE

(1900 : 489). The fifth error concerns the author of the first family-group name based on the

generic name Rana, which proves to be GoLDFuss (1820), not GRAY (1825) (see DuBois, D-

29). The two remaining errors concern the family-group names Allophrynidae and Platy-

mantinae, created by SAVAGE (1973): I had not realized that these names were not accom-

pagnied by any diagnosis and are therefore nomina nuda in this work ; the first one obtained

a status in nomenclature in GOIN, GoIN & ZUG’s (1978) work (see DuBoIs, D-31), and the

second one, which is a junior synonym of Dicroglossidae Anderson, 1871 (see DuBois, D-

26, D-35), obtained a status in nomenclature, under the spelling Platymantini, in LAURENT

(1986 a : 760). As for the omissions, four of them also concern subsequent spellings of fam-

ily-group names : I had missed the spelling Pelobatidea, due to HUXLEY (1871 : 189), and

the spellings Pipini, Bufonini and Hyladini, used by BONAPARTE (1839 : [225]). Finally, I

only recently (D-35) rediscovered the family-group name Colodactyli Tschudi, 1845, which

should now stand in the synonymy of the name Discoglossidae, and entails a change of date

for this name.

The total number of names mentioned in my checklist (D-26) being 572, the total EO

rate for this whole work is therefore of 2.1% (E rate : 1.2% ; O rate : 0.9%), which seems

acceptable according to the a priori criterion proposed elsewhere (D-41). À few additional

mistakes or omissions certainly remain to discover, but this should not significantly alter these

figures.

RÉSUMÉ

L'auteur répond aux critiques diverses récemment émises par SAVAGE (1986) sur ses

propres travaux (notamment DUBOIS, 1984), et montre que celles-ci sont dues à de graves

incompréhensions de plusieurs importantes règles de nomenclature. Une attention particu-

lière est accordée aux problèmes posés par la distinction entre différents types de noms et

d’orthographes (nouveau nom, nouveau nom de remplacement, émendation injustifiée, or-

thographe subséquente incorrecte), et plusieurs exemples sont traités en détail. Il est montré

que certaines des suggestions de SAVAGE (1986) concernant les noms du groupe-famille re-

viennent en réalité à une proposition de retour à des règles antérieures à celles du Code in-

ternational de Nomenclature zoologique pour ces noms, puisque cet auteur n’accepte pas l’ap-

plication du principe de priorité à ces noms. La question des règles suggérées par DuBois

(1984) pour la nomenclature des taxons du groupe-classe est aussi discutée, et les modifi-

cations récemment proposées à cet égard par LESCURE, RENOUS & Gasc (1986) sont criti-

quées.

Source : MNHN, Paris

sa ALYTES 6 (1-2)

LITERATURE CITED

Note . — To save space, the references to the works cited above which are given in the papers

D:8, D-11, D-25, D-26, D-29 and D-41 are not repeated here (the references D-I1 to D-40 are given

in D-41).

BiscHorr, W., 1982. — Die innerartliche Gliederung von Gallotia galloti (Duméril & Bibron 1839)

(Reptilia : Sauria : Lacertidae) auf Tencriffa, Kanarische Inseln. Bonn. zool. Beitr., 33 : 363-

382.

BONAPARTE, C. L., 1837. — Salamandra maculosa. Salamandra pezzata. In : Iconografia della fauna it-

alica per le quauro classi degli animali vertebrati, vol. II, [fasc. XIX], Roma, Salviucci : [245-250],

pl. [52].

ze 1839. — Bufo vulgaris. Rospo comune. /n : Iconografia della fauna italica per le quatro classi degli

animali vertebrati, vol. Il, [fasc. XXIV], Roma, Salviucci : [223-228], pl. [49].

Bou, R. & DuBols, A., 1986. - Nomenclature ordinale et familiale des tortues (Reptilia). Note com-

plémentaire. Bull. Soc. linn. Lyon, 55 : 87-90.

BRATTSTROM, B. H., 1957. - The phylogeny of the Salientia based on skeletal morphology. Syst. Zool.,

6: 70-74.

CLARKE, B. T., 1983. - À morphological re-examination of the frog genus Nannophrys (Anura : Ran-

idae) with comments on its biology, distribution and relationships. Zool. 7. Linn. Soc., 79 : 377-

398.

CUELLAR, H. S., FAWCETT, J. D., FERNER, J. W., MASLIN, P., OLDHAM, J. C., ROTH, J. J., SAVITZKY,

A. & SMITH, H.M., 1972. - Comment on Dendrobates. Z.N.(S.)1930. Bull. zool. Nom., 29 : 24.

Dugois, A., 1987 (D-41). — Living amphibians of the world : a first step towards a comprehensive

checklist. Alytes, 19-149.

1984 (D-42). = Les spécimens-types de Gallotia galloti (Oudart, 1839) (Reptiles, Sauriens). Bull.

Soc. linn. Lyon, 53 : 27-30.

Dupuis, C., 1984. - Explication du vote sur le projet de la troisième édition du Code international de

Nomenclature zoologique. Z.N.(G.)197. Bull. zool. Nom., 41 : 141-148.

Gray, J. E., 1842. - The northern z0ological gallery. Zn : Synopsis of the contents of the British Museum,

44th. ed., London, British Museum : 97-157.

HoG, J., 1839 a. — On the classifications of the Amphibia. Mag. nat. Hist., (n.s.), 3 : 265-274.

---- 1839 b. — On the classifications of the Amphibia (continued). Mag. nat. Hist., (n.s.), 3 : 367-378.

—s 1841. — On the existence of branchiae in the young Caeciliae ; and on a modification and extension

of the branchial classification of the Amphibia. Ann. Mag. nat. Hist., (1), 7 : 353-363.

Horus, L. B., 1983. - Comments on the proposed conservation of Dendrobates Wagler, 1830 and

Dendrobatidae Cope, 1865. Z.N.(S.)1930. Bull. zool. Nom., 40 : 197-198.

HUXLEY, T. H., 1871. — À manual of the anatomy of vertebrated animals. London, Churchill : i-vii + 1-

510.

LESCURE, J., 1982. — L’espèce-type du genre Dendrobates Wagler, 1830 : nouvelles propositions.

Z.N(S.)1930. Bull. zool. Nom., 39 : 264-267.

LESCURE, J., RENOUS, S. & Gasc J.-P., 1986. - Proposition d’une nouvelle classification des Amphi-

biens Gymnophiones. Mém. Soc. zool. France, 43 : 145-177, 1 tab.

MALHERBE, M., 1983. — Les langages de l'humanité. Paris, Seghers : 1-444.

MICHAHELLES, C., 1833. - Waglers Synonymie der Sebaischen Amphibien. Jsis von Oken, 1833 : 884-

905.

Myers, C. W. & DaLv, J. W., 1971. - Comment on the proposed designation of a new type-species

of Dendrobates Wagler, 1830. Z.N.(S.)1930. Bull. zool. Nom., 28 : 141.

PARKER, H. W., 1930 a. - Report on the Amphibia collected by Mr. J. Omer-Cooper in Ethiopia. Proc.

zool. Soc. London, 1930 : 1-6.

— 1930 b. — A collection of frogs from Portuguese East Africa. Proc. z0ol. Soc. London, 1930 : 897-

905, pl. I.

su 1932. — Parallel modifications in the skeleton of the Amphibia Salientia. Arch. zool. ital., 16 : 1239-

1248.

Source : MNHN, Paris

Dugois 55

es 1936 a. - The Amphibians of the Mamfe Division, Cameroons. I. Zoogeography and systematics.

Proc. zool. Soc. London, 1936 : 135-163, pl. I.

_—— 1936 b. — Dr. Karl Jordan’s expedition to South-West Africa and Angola : herpetological collec-

tions. Novit. zool., 40 : 115-146.

ee. 1937. - Reptiles and Amphibians from the Mafia archipelago. Ann. Mag. nat. Hist., (10), 20 :629-

632.

PETERS, J. A., DUELLMAN, W. E., LyNCH, J. D., MYERS, C.W., SAVAGE, J. M. & WALkER, C. F.,

1972. - Comment on the proposed designation of type-species for the Amphibian genus Den-

drobates. Z.N.(S.)1930. Bull. zool. Nom., 29 : 107-108.

RiscH, J.-P. & THORN, R., 1982. - Notes sur Ranodon shihi (Liu, 1950) (Amphibia, Caudata, Hyno-

biidae). I. Taxonomie. Bull. Soc. Hist. nat. Toulouse, 1981 (1982), 117 : 171-174.

SAVAGE, J. M., 1986. — Review of DuBois, 1984 e (D-26). Copeia, 1986 : 259-262.

SCHUCHERT, C. & BUCKMAN, S. S., 1905. — The nomenclature of types in natural history. Ann. Mag.

nat. Hist., (7), 16 : 102-104.

SHERBORN, C. D., 1928. — Index animalium. 1801-1850. Fasc. 15 (M, N). London, British Museum :

3747-4450.

SILVERSTONE, P. À., 1971. - Dendrobates Wagler, 1830 (Amphibia : Anura): proposed designation of

type-species under the plenary powers. Z.N.(S.)1930. Bull. zool. Nom., 27 : 262-264.

STARRETT, P. H., 1973. - Evolutionary patterns in larval morphology. n : J. L. VIAL (ed.), Evolu-

tionary Biology of the Anurans, Columbia, Univ. Missouri Press : 251-271.

STEJNEGER, L., 1937. - Designation of genotype for Hylaplesia Boie. Copeia, 1937 : 139.

Zmao, E., 1984. — The nomenclature of the type species of the genus Liua should be revised. Acta

herpet. sinica, 3 : 40.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2): 56-68.

Discoglossidae Günther, 1858 (Amphibia, Anura):

proposed conservation

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The family-group name Discoglossidae Günther, 1858 is shoun to be a junior

synonym of four other names : Bombinatorina Gray, 1825 : Alytae Fitzinger, 1843 :

Bombitatores Fitzinger, 1843 : and Colodactyli Tschudi, 1845. The Article 40 of the Code

allows to give the name Discoglossidae the date 1845 and priority over Colodactyli, but

such an action cannot be taken as concerns the three other names. The ICZN is the-

refore requested to use its plenary powers to rule that the name Discoglossidae is to be

given precedence over these three names whenever these names are considered to ap-

ply to a single taxon.

Note. — This paper was submitted on 24 September 1982 to the Secretary of the International

Commission on Zoological Nomenclature for publication in the Bulletin of zoological Nomenclature, but,

despite repeated requests since then, has still not been published in this journal. The problem it raised

has therefore remained unresolved, which is unfortunate, especially when one considers that, since then,

several papers discussing the phylogeny, classification and nomenclature of the Discoglossidae, as well

as books and checklists mentioning this and other family-group names, have been published (DUBoOIS,

1983, 1984, 1985, 1987 a ; MAxsON & SZYMURA, 1984 ; SANCHIZ, 1984 ; ALCOVER, SANDERS & SAN-

CHIZ, 1984 ; FROST, 1985 ; DUELLMAN & TRUEB, 1986 ; LAURENT, 1986), and others are in progress.

On 6 May 1986, the editor of the Bulletin of zoological Nomenclature at last proposed me to publish this

paper, but in a drastically reduced version, excluding in particular the list of references, which would

simply have been mentioned as being “held by the Secretariat”. Unfortunately, a list of references held

by a Secretariat cannot compare with a published list of references, which is immediately available to

the whole scientific community, and will remain so in the future. The paper is therefore published here

as it was submitted, except that a few more recent bibliographic references and information have been

incorporated.

() The family-group name Discoglossidae was coined by GÜNTHER (1858 : 346 ; 1859)

for a family including six nominal genera of Amphibia Anura (Chiroleptes, Pelodytes, Dis-

coglossus, Leptobrachium, Megalophrys and Ceratophryne).

In the same works, GÜNTHER (1858, 1859) also recognized a family Bombinatoridae,

with four genera (Pelobates, Bombinator, Alsodes and Telmatobius), and a family Alytidae, with

three genera (Alytes, Scaphiopus and Heleioporus).

(2) Core (1864, 1865) proposed a new classification of the Anura, in which he grouped

in a single family the genera Latonia, Discoglossus, Alytes, Bombinator and later (1866) Za-

Dhrissa. For this family he chose to use the name Discoglossidae.

Source : MNHN, Paris

Dugois 57

In making this choice, CoPE (1864) would have taken a “first reviser action” if the three

family-group names mentioned above had been created in the same paper by GÜNTHER (1858).

Unfortunately, this is not the case.

Indeed, at the time when Core took this action, his choice was particularly ill-foun-

ded. He should have used the name Bombinatoridae for this family : this name had the prio-

rity, having been coined, under the spelling Bombinatorina, by GRAY (1825) ; furthermore

this name had been used regularly by zoologists since its creation and remained in use at

least until 1885 (see references in Appendix).

The second and third family-group names available for the family were those of Alytae

and of Bombitatores, both coined by FITZINGER (1843 : 32). The first one, based on the ge-

neric name Alytes Wagler, 1829, had been used by several authors since its creation (see Ap-

pendix). The second one, based on the generic name Bombitator Wagler, 1830, an unjusti-

fied emendation and hence substitute name for Bombinator Merrem, 1820 (see DuBois, 1984),

had been used only once, and by its author (FITZINGER, 1860 : 415) after its creation. No

first reviser action having ever been taken concerning the relative priority of the names Aly-

tae and Bombitatores, I hereby select Alytae and consider it to have priority over Bombi-

tatores.

The fourth available name for this group is Colodactyli, coined by TsCHUDI (1845) on

the basis of the generic name Colodactylus Tschudi, 1845, a junior subjective synonym of

Discoglossus Otth, 1837 (see DuBois, 1987 : 11).

The name Discoglossidae was only the fifth to have been coined, and had been used

only by GÜNTHER (1858, 1859) when CoPE's (1864) work was published.

(3) Although CopE’s choice was not justified at the time when he made it, it was fol-

lowed by most subsequent authors. This was the case in particular of BOULENGER (1882 :

444), who recognized a family Discoglossidae, including the genera Discoglossus, Bombinator,

Liopelma and Alytes. Since then the name Discoglossidae has been almost universally used

by zoologists (see Appendix) and is now considered the valid name of a family including se-

veral fossil general and four living genera of Anura : Alytes Wagler, 1829 ; Barbourula Tay-

lor & Noble, 1924 ; Bombina Oken, 1816 (Name Number 1064 on the Official List of Generic

Names in Zoology) (a name of which Bombinator Merrem, 1820, and Bombitator Wagler, 1830,

a substitute name for the latter, are junior subjective synonyms) ; and Discoglossus Otth, 1837

(a name of which Colodactylus Tschudi, 1845 is a junior subjective synonym).

Furthermore, this name has also been used, under the spelling Discoglossoidea, for a

superfamily including the Discoglossidae and the Leiopelmatidae, by several recent authors

(DUELLMAN, 1975 ; DOWLING & DUELLMAN, 1978 ; CEI, 1980 ; LAURENT, 1980, 1986 ;

Dugois, 1983, 1984, 1985 ; MAxSON & SzZYMURA, 1984).

(4) In view of the almost universal use of the name Discoglossidae since 1882, conser-

vation of this name and rejection of the names Bombinatoridae, Alytidae, Bombitatoridae

and Colodactylidae is to be commanded.

As concerns the last of these four names, this action may be taken easily and by a simple

recourse 10 Article 40 of the Code (ANONYMOUS, 1985): the generic name Colodactylus having

been rejected as a junior synonym of Discoglossus, the family-group name Colodactyli having

been replaced by Discoglossidae before 1961, and the replacement having won general ac-

Source : MNHN, Paris

58 ALYTES 6 (1-2)

ceptance, this replacement is to be maintained. The name Discoglossidae retains its own au-

thor (GÜNTHER) but takes the precedence of Colodactyli of which it is to be deemed the se-

nior synonym : it must therefore be now written Discoglossidae Günther, 1858 (1845) (see

Dugois, 1987 à).

As concerns the three other names (Bombinatoridae, Alytidae and Bombitatoridae) ho-

wever, simple recourse to the Code does not allow to reject them, and appeal must be made

to the ICZN to solve the problem. Two different actions could be contemplated to achieve

this aim :

— the first one would to the suppression under the plenary powers of the names Bom-

binatorina Gray, 1825, Alytae Fitzinger, 1843 and of all other family-group names based on

the generic names Bombinator, Alytes and Bombitator proposed before 1858 (see Appendix);

— the second one would be a Ruling under the plenary powers stating that the name

Discoglossidae Günther, 1858 (1845) should be given precedence over the name(s) Bombi-

natorina Gray, 1825, Alytae Fitzinger, 1843 and/or Bombitatores Fitzinger, 1843 by any

zoologist who considers that the genera Discoglossus and Bombina and/or Alytes belong to the

same family-group.

The second action is here advocated because it is more economical and also for addi-

tional reasons which will now be discussed.

(5) Recently, LANZA, CEI & CRESPO (1975, 1976) presented evidence for a rather large

immunological distance between the genus Discoglossus on one side and the genera Alytes and

Bombina on the other, and suggested that both groups could be considered as two distinct

families.

For the new family which they proposed to recognize, LANZA, CEI & CRESPO (1975,

1976) used the name Bombinidae, which they credited to FITZINGER (1826). In fact, the name

used by FITZINGER (1826) was Bombinatoroidea, a junior synonym of Bombinatorina Gray,

1825. This family-group name is based on the generic name Bombinator Merrem, 1820, which

is a junior subjective synonym of Bombina Oken, 1816 and has therefore a separate status

in nomenclature. The name Bombinatorina (or Bombinatoroidea) may therefore be emended

into Bombinatoridae or Bombinatorinae, but not into Bombinidae or Bombininae.

The first family-group name based on the generic name Bombina Oken, 1816 is Bom-

bininae Fejérväry, 1921. The existence of this name seems to have been ignored by most

authors after its creation. Thus MERTENS (1955 : 132), in his application for the conservation

of the generic name Bombina, wrote : “The genus Bombina Oken, 1816, is not the type ge-

nus of a taxon belonging to any family-group”. After its creation, the name Bombininae (un-

der this spelling or under the emendation Bombinidae) has been used by TATARINOV (1964),

KUHN (1965, 1967) and LaANZA, CEI & CRESPO (1975, 1976). I have been unable to trace

other uses of this name.

(6) Subsequent authors did not follow LaNZa, CEI & CRESPO’s (1975, 1976) proposal

to recognize two families instead of one, but some authors proposed to divide the family Dis-

coglossidae in two subgroups.

When this application was submitted to the ICZN (24 September 1982), this use of

subfamilial suprageneric groups had not started. Since then, while some authors (e.g. OLMO

et al., 1982 ; MaxsON & SZYMURA, 1984 ; FROST, 1985 ; DUELLMAN & TRUEB, 1986 ; LAU-

Source : MNHN, Paris

DuBois 59

RENT, 1986) maintained all the genera of this group in a single undivided family, other au-

thors proposed to divide the family in two subgroups. These subgroups were given the rank

of “generic groups” (“Discoglossus group” and “Bombina group”: ESTES & SANCHIZ, 1982 a,

1982 b), of tribes (Discoglossini and Alytini : SANCHIZ, 1984 ; ALCOVER, SANDERS & SAN-

cxiz, 1984) or of subfamilies (Discoglossinae and Bombinatorinae : DuBoIs, 1983, 1984, 1985,

1987 a).

Since it is likely that this use of subfamilial suprageneric groups in the Discoglossidae

will be continued, it is important and urgent to stabilize the nomenclature of these groups

and to solve the problems detailed above.

(7) If the family Discoglossidae is to be divided in two subfamilies, which name should

be given to the non-nominotypical one?

In view of the recency of the name Bombininae and of its very scarce use since its crea-

tion (which is far from being a “general acceptance”, in the sense e.g. of Art. 40), conser-

vation of this name in place of its senior subjective synonym Bombinatorina is not justified,

all the more that this latter name has had a period of rather large use in the nineteenth cen-

tury (see Appendix). Furthermore, if the name Bombinatorina was to be suppressed, the

second name available for a family-group including Alytes and Bombina would be Alytae Fit-

zinger, 1843, a name which has priority over Bombininae Fejérväry, 1921 and which has

been used more in the past than this latter name (see Appendix). Finally, a third senior sy-

nonym of Bombininae is Bombitatores Fitzinger, 1843, so that conservation of the name

Bombininae would request the suppression of three family-group names by the Commission.

The simplest and best action appears to be the conservation of the name Bombinato-

rina. This name has priority on the other ones, and was used more, and longer, than them,

in the XIX century (see Appendix). The fact that this family-group name is based on a ge-

nus-group name which is now considered a junior synonym of another name is not a valid

reason for rejecting this name, if one admits that family-group names of taxa are to be re-

gulated by the Code (Art. 1 of the Code, ANONYMOUS, 1985) and in particular by its Prin-

ciple of Priority (Art. 23 of the Code) (for a more detailed discussion of this question, see

Dugois, 1987 b).

The simplest action which can be suggested for solving all the problems discussed above

is therefore to provide for the priority of the name Discoglossidae over the name Bombi-

natorina, but without suppressing any name. The authors who believe that only one family-

group taxon should be recognized for all these genera will therefore keep using the name

Discoglossidae for this taxon, while those who think that Alytes, Bombina and Barbourula

should be separated from Discoglossus at the subfamilial or familial level can use the name

Bombinatorinae in this respect. Finally, for those who may think that even more splitting

is necessary, they could have for example a family Discoglossidae Günther, 1858 (1845) with

two subfamilies, Discoglossinae Günther, 1858 (1845) and Bombinatorinae Gray, 1825, the

latter with two tribes, Bombinatorini Gray, 1825 and Alytini Fitzinger, 1843 (see DUBoIs,

1987 a : 11-12).

(8) Accordingly I ask the International Commission on Zoological Nomenclature :

(1) to use its plenary powers to rule that the family-group name Discoglossidae Gün-

ther, 1858 (1845) is to be given precedence over the name(s) Bombinatorina Gray, 1825,

Alytae Fitzinger, 1843 and/or Bombitatores Fitzinger, 1843 by any zoologist who considers

Source : MNHN, Paris

60 ALYTES 6 (1-2)

that the genera Discoglossus Otth, 1837 and Bombina Oken, 1816 and/or Alytes Wagler, 1829

belong to the same family-group;

(2) to place the following family-group names on the Official List of Family-Group

Names in Zoology:

(a) Discoglossidae Günther, 1858 (1845) (type genus Discoglossus Otth, 1837), with

an endorsement that it is to be given nomenclatural precedence over Bombinatorina Gray,

1825, Alytae Fitzinger, 1843 and/or Bombitatores Fitzinger, 1843 whenever these names are

considered synonyms;

(b) Bombinatorina Gray, 1825 (type genus Bombinator Merrem, 1820, a junior

subjective synonym of Bombina Oken, 1816, Name Number 1064 on the Official List of Ge-

neric Names in Zoology), with an endorsement that it is not to be given priority over Dis-

coglossidae Günther, 1858 (1845) whenever the two names are considered synonyms;

that it is not to be given priority over Discoglossidae Günther, 1858 (1845) whenever the two

names are considered synonyms;

(d) Bombitatores Fitzinger, 1843 (type genus Bombitator Wagler, 1830, a substi-

tute name for Bombinator Merrem, 1820), with an endorsement that it is not to be given

priority over Discoglossidae Günther, 1858 (1845) whenever the two names are considered

synonyms;

(3) to place the following generic names on the Official List of Generic Names in Zoo-

logy:

(a) Discoglossus Otth, 1837 (gender : masculine) (type species, by monotypy, Dis-

coglossus pictus Otth, 1837);

(b) Bombinator Merrem, 1820 (gender : masculine) (type species, by subsequent

designation of DUMÉRIL & BIBRON (1841 : 485), Bufo igneus Laurenti, 1768, a junior sub-

jective synonym of Rana bombina Linnaeus, 1761, Name Number 1141 on the Official List

of Specific Names in Zoology);

obstetricans Laurenti, 1768);

(d) Bombitator Wagler, 1830, a substitute name for Bombinator Merrem, 1820;

(4) to place the following specific names on the Official List of Specific Names in Zoo-

logy:

(a) pictus Otth, 1937, as published in the binomen Discoglossus pictus (specific name

of type species of Discoglossus Otth, 1837);

(b) igneus Laurenti, 1768, as published in the binomen Bufo igneus (specific name

of type species of Bombinator Merrem, 1820 and of Bombitator Wagler, 1830);

cific name of type species of Alytes Wagler, 1829).

Source : MNHN, Paris

Dugois

RÉSUMÉ

Le nom Discoglossidae Günther, 1858 s’avère un synonyme plus récent de quatre autres

noms du groupe-famille : Bombinatorina Gray, 1825 ; Alytae Fitzinger, 1843 ; Bombita-

tores Fitzinger, 1843 ; et Colodactyli Tschudi, 1845. L'article 40 du Code permet d’attribuer

au nom Discoglossidae la date de 1845 et de lui donner la priorité sur Colodactyli, mais une

telle action ne peut être effectuée en ce qui concerne les trois autres noms. En conséquence,

il est demandé à l’ICZN de faire usage de ses pleins pouvoirs pour décider que la priorité

doit être donnée au nom Discoglossidae par rapport aux trois autres noms, lorsque ces noms

sont considérés comme synonymes.

APPENDIX

List of references of works using the family-group names Bombinatorina Gray, 1825,

Alytae Fitzinger, 1843, Bombitatores Fitzinger, 1843, Colodactyli Tschudi, 1845, Disco-

glossidae Günther, 1858 and Bombininae Fejérväry, 1921.

Family-group original name,

author and date

(name of type genus)

Spelling used

References

Bombinatorina Gray, 1825

(Bombinator Merrem, 1820)

Bombinatorina

Bombinatoroidea

Bombinatoridae

Bombinatores

Bombinatorida

Bombinatorinae

Bombinatorini

GRaY, 1825 : 214

BONAPARTE, 1850

BONAPARTE, 1852 : 477

GONTHER, 1858 : 344

GONTHER, 1859 : viii, 40

MivarT, 1869 : 290

FITZINGER, 1826 : 37

Gray, 1831 : 38

BONAPARTE, 1850

BONAPARTE, 1852 : 477

GONTHER, 1858 : 343

GÜNTHER, 1859 : viii, 40

STEINDACHNER, 1867 : 33

MIvarT, 1869 : 286

FaTI0, 1872 : 230

HOFFMANN, 1878 : 633

MULLER, 1878 : 582

HUDI, 1838 : 26

TscHUpI, 1845 : 170

BAYER, 1885 : 22

Dugois, 1983 : 27]

1984 : 26

s, 1985 : 74

Dugois, 1987 a : 12

Source : MNHN, Paris

ALYTES 6 (1-2)

Family-group original name,

author and date

(name of type genus)

Spelling used

References

Alytae Fitzinger, 1843

(Alytes Wagler, 1829)

Bombitatores Fitzinger, 1843

(Bombitator Wagler, 1830)

Colodactyli Tschudi, 1845

(Colodactylus Tschudi, 1845)

Discoglossidae Günther, 1858

(Discoglossus Ouh, 1837)

Alytae

Alytina

Alytidae

Alytini

Alytinae

Bombitatores

Colodactyli

Colodactylidac

Discoglossidae

FITZINGER, 1843 : 32

TscHuDi, 1845 : 168

FITZINGER, 1860 : 415

BONAPARTE, 1850

BONAPARTE, 1852 : 477

MivarT, 1869 : 291

GÜUNTHER, 1858 : 346

GONTHER, 1859 : viii, 37

Corr, 1863 : 51

STEINDACHNER, 1867 : 32

KEFERSTEIN, 1868 : 269

MivarT, 1869 : 286

FATI0, 1872 : 230

HOFFMANN, 1878 : 631

LATASTE, 1878 : 491

MÜLLER, 1878 : 582

LATASTE, 1879 : 761

FROST, 1985 : 105

SaNcHIZ, 1984 : 61

ALCOVER, SANDEI

Duois, 1987 a : 12

FITZINGER, 1843 : 32

FITZINGER, 1860 : 415

TscHUDI, 1845 : 167

Dugois, 1987 a : 11

Dumois, 1987 a: 11

GÜNTHER, 1858 : 346

GÜNTHER, 1859 : vii, 34

Cor, 1864 : 183

STEINDACHNER, 1867 : 28

KEFERSTEIN, 1868 : 267

THEOBALD, 1868 : 82

MivarT, 1869 : 287

FaTI0, 1872 : 230

CorE, 1875 : 10

HOFFMANN, 1878 : 627

LATASTE, 1878 : 491

MULLER, 1878 : 582

LATASTE, 1879 : 761

R, 1882 : 444

CAMERANO, 1884 : 203

S & SANCHIZ, 1984 : 109

Source : MNHN, Paris

Duois 63

Family-group original name, Spelling used References

author and date

(name of type genus)

Corr, 1889 a : 248

CorE, 1889 b : 862

BOULENGER, 1897 : 123

Gapow, 1901 : 152

STEJNEGER, 1907 : 50

BOULENGER, 1910 : 149

SCHREIBER, 1912 : 159

VAN DENBURGH, 1912 : 259

NICHOLLS, 1916 : 81

BoLkay, 1919 : 349

FEJÉRVARY, 1921 : 25

NOBLE, 1922 : 22

NIEDEN, 1923 : 24

MiRANDA-RIBEIRO, 1924 : 143

TayLor & NOBLE, 1924 : 1

STEJNEGER, 1925 : 6

NOBLE, 1927 : 69

ScHmipT, 1927 : 556

MERTENS & MULLER, 1928 : 15

NoBLE, 1931 : 486

Pore, 1931 : 434

Liv, 1935 : 22

MERTENS & MÜLLER, 1940 : 14

PorE & BORING, 1940 : 24

BOURRET, 1942 : 153

INGER, 1954 : 209

RI 1958 : 115

PASTEUR & BONS, 1959 : 100

FUHN, 1960 : 164

MERTENS & WERMUTH, 1960 : 37

Liv & Hu, 1961 : 35

GoIN & GoIN, 1962 : 223

GRIFFITHS, 1963 : 271

TIHEN, 1965 : 315

GoRHAM, 1966 : 9

OKapa, 1966 : 1

INGER, 1967 : 371

KawamURA, NIsHIOKA & UEDA, 1972 : 303

ESTEs & REIG, 1973 : 17

LYNCH, 1973 : 134

SAVAGE, 1973 : 354

STARRETT, 1973 : 251

TRUEB, 1973 : 70

GORHAM, 1974 : 41

DUELLMAN, 197

Lanza, CEI & CRESPO, 1975 : 158

L4 , CET & CRESPO, 1976 : 311

SokoL, 1977 : 505

DOWLING & DUELLMAN, 1978 : 21.1

GOIN, GoiN & ZUG, 1978 : 231

ISKANDAR, 1978 : 564

SANCHIZ & ADROVER, 1979 : 6

Source : MNHN, Paris

64 ALYTES 6 (1-2)

Family-group original name, Spelling used References

author and date

(name of type genus)

LAURENT, 1980 : 399

ESTES & SANCHIZ, 1982 a : 9

EsTEs & SANCHIZ, 1982 b : 33

OLMo et al., 1982 : 283

Dugois, 1983 : 271

Duois, 1984 : 24

MaxsON & SZYMURA, 1984 : 245

VIERTEL, 1984 : 21

SANCHIZ, 1984 : 61

ALCOVER, SANDERS & SANCHIZ, 1984 : 109

Dugois, 1985 : 74

FRosr, 1985 : 105

DUELLMAN & TRUEB, 1986 : 518

LAURENT, 1986 : 661

Dumors, 1987 a : 11

Discoglossina MivarT, 1869 : 294

Discoglossinae FEJÉRVARY, 1921 : 25

Dusois, 1983 : 271

Duois, 1984 : 26

Dugois, 1985 : 74

Dugois, 1987 a : 12

Discoglossoidea DUELLMAN, 1975 : 5

DowLING & DUELLMAN, 1978 : 19.1

Cet, 1980 : 11

LAURENT, 1980 : 398

Dusois, 1983 : 271

Dugois, 1984 : 24

MaxsoN & SZYMURA, 1984 : 245

Dugois, 1985 : 74

LAURENT, 1986 : 658

Bombininae Fejérväry, 1921 Bombininae FEJÉRVARY, 1921 : 25

(Bombina Oken, 1816) KUHN, 1967 : 16

Bombinidae TATARINOV, 1964 : 128

KUHN, 1965 : 89

KUHN, 1967 : 16

Lanza, CEI & CRESPO, 1975 : 158

Lanza, CE1 & CRESPO, 1976 : 311

LITERATURE CITED

ALCOVER, J. A., SANDERS, E. & SANCHIZ, B., 1984. — El registro f6sil de los sapos parteros (Anura,

Discoglossidae) de Baleares. /n : HEMMER & ALCOVER (1984) : 109-121.

BAYER, F., 1885. — O kostie Zab z éeledi pelobatid. Abh. bôhm. Ges. Wiss., (6), 12 (13) : 1-24, pl. 1-

Source : MNHN, Paris

Dugois 65

Boukay, S. J., 1919. - Osnove Uporedne Osteologije Anurskih Batrahija. Glasn. zemalj. Muz. Bosni

Herceg., 31 : 275-357.

BONAPARTE, C. L., 1850. — Conspectus systematum Herpetologiae et Amphibiologiae. Editio altera refor-

mata. Lugduni Batavorum, Brill : 1 pl.

ee 1852. - Conspectus systematum Herpetologiae et Amphibiologiae. Nuovi Ann. Sci. nat. Bologna,

(3), 5 : 477 - 480.

BOULENGER, G. A., 1882. - Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British

Museum. London, Taylor & Francis : i-xvi + 1-503, pl. I-XXX.

ee 1897. — The Tailless Batrachians of Europe. Part I. London, Ray Society : i-äi + 1-210, pl. I-X, 2

maps.

re 1910. — Les Batraciens, et principalement ceux d'Europe. Paris, Doin : ii + 1-305.

BOURRET, R.,; 1942. — Les Batrachiens de l’Indochine. Hanoï, Institut océanographique de l’Indochine :

ix + 1-547, 4 pl.

CaMERANO, L., 1884. - Monografia degli Anfibi Anuri italiani. Mem. real Accad. Sci. Torino, (2), 35 :

187-284, pl. I-II.

Cet, J. M., 1980. — Amphibians of Argentina. Monit. zool. ital., (n.s.), monogr. 2: i-xii + 1-609, pl.

1-16 + 1-24.

Core, E. D., 1863. - On Trachycephalus, Scaphiopus and other American Batrachia. Proc. Acad. nat.

Sci. Phila., 1863 : 43-54.

ee 1864. — On the limits and relations of the Raniformes. Proc. Acad. nat. Sci. Phila., 1864 : 181-

183.

ee 1865. — Sketch of the primary groups of Batrachia Salientia. Nat. Hist. Rev., (n.s.), 5 : 97-120.

ue 1866. — On the structures and distribution of the genera of the Arciferous Anura. 7. Acad. nat.

Sci. Phila., (2), 6 : 67-112.

- 1875. - Check-list of North American Batrachia and Reptilia. Bull. U.S. nat. Mus., 1 : 1-104.

- 1889 a. - The Batrachia of North America. Bull. U. S. nat. Mus., 34 : 1-525, pl. I-LXXXVI.

- 1889 b. — Synopsis of the families of Vertebrata. Am. Nat., 23 : 849-877.

DowLiNG, H. G. & DUELLMAN, W. E., 1978. — Systematic herpetology : a synopsis of families and

higher categories. Publications in Herpetology, Hiss Publications, New York, 7 : i-vii + 1.1-118.3

+ vi.

Dusois, A., 1983. — Classification et nomenclature supragénérique des Amphibiens Anoures. Bull. Soc.

linn. Lyon, 52 : 270-276.

as 1984. — La nomenclature supragénérique des Amphibiens Anoures. Mém. Mus. nat. Hist. nat., (À),

131 : 1-64.

- 1985. - Miscellanea nomenclatorica batrachologica (VIT). Alytes, 4 : 61-78.

- 1987 a. - Miscellanea taxinomica batrachologica (D). Alytes, 5 : 7-95.

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

DuELLMAN, W. E., 1975. - On the classification of frogs. Occ. Pap. Mus. nat. Hist. Univ. Kansas, 42 :

1-14.

DuELLMAN, W. E. & TRUEB, L., 1986. — Biology of Amphibians. New York, McGraw-Hill : i-xix + 1-

670.

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

Tome 8. Paris, Roret : + 1-792.

EsTes, R. & REIG, O. A., 1973. — The early fossil record of frogs : a review of the evidence. /n: VIAL

(973) : 11-63.

EsTEs, R. & SANCHIZ, B., 1982 a. - New discoglossid and palaeobatrachid frogs from the late Creta-

ceous of Wyoming and Montana, and a review of other frogs from the Lance and Hell Creek

formations. 7. Vert. Paleont., 2 : 9-20.

= 1982b. — Early Cretaceous lower Vertebrates from Galve (Teruel), Spain. 7. Vert. Paleont., 2 : 21-

39.

FaTi0, V., 1872. - Faune des Vertébrés de la Suisse. Genève & Bâle, Georg : 1-603 + i-xii, pl. I-V.

FEJéRvARY, G. J. V., 1921. - Kritische Bemerkungen zur Osteologie, Phylogenie und Systematik der

Anuren. Arch. Naturgesch., 87 (3) : 1-30.

FITZINGER, L. L., 1826. - Neue Classification der Reptilien. Wien, Heubner : i-vii + 1-66, 1 tab.

aa 1843. - Systema Reptilium. Fasciculus primus. Amblyglossae. Vindobonae, Braumüller & Seidel :

1-106 + i-ix.

Source : MNHN, Paris

66 ALYTES 6 (1-2)

--- 1860. — Die Ausbeute der ôsterreichischen Naturforscher an Säugethieren und Reptilien während

der Weltumsegelung Sr. Majestät Fregatte Novara, Sber. Akad. Wiss. Wien, 42 : 383-416.

Frost, D. R. (ed.), 1985. - Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst. Coll.:

Liv] + iv + 1.732.

FUHN, I. E., 1960. - Amphibia. Fauna Republicii Populare Romine, Bucuresti, Academiei Republicii

Populare Romine, 14 (1) : 1-288.

Gapow, H., 1901. - Amphibia and Reptiles. London, Macmillan & Co. xiii + 1-668, 1 pl.

Go, & J. & Go, O.B., 1962. — Introduction to herpetology. San Francisco & London, Freeman &

-ix + 1-341.

GoiN, e GoiN, O. B. & ZUG, G. R., 1978. — Introduction 10 herpetology. Third edition. San Fran-

cisco, Freeman & Co. : i-xiii + 1-378.

GorHaAM, S. W., 1966. — Liste der rezenten Amphibien und Reptilien. Ascaphidae, Leiopelmatidea

(sic), Pipidae, Discoglossidae, Pelobatidae, Leptodactylidac, Rhinophrynidae. Das T'ierreich, 85 :

i-xvi + 1-222.

1974. — Checklist of world Amphibians up to January 1, 1970. Saint-John, The New Brunswick Mu-

seum : 1-173.

Gray, J. E., 1825. — À synopsis of the genera of Reptiles and Amphibia, with a description of some

new species. Ann. Philos., (2), 10 : 193-217.

Er 1831. - Description of two new genera of frogs discovered by John Reeves, Esq. in China. /n :

The zoological Miscellany, London, Treuttel, Wurz & Co., 1-40 : 38.

GRIFFITHS, L., 1963. - The phylogeny of the Salientia. Biol. Rev., 38 : 241-292.

GONTHER, AÀ., 1858. — On the systematic arrangement of the Tailless Batrachians and the structure of

Rhinophrynus dorsalis. Proc. zool. Soc. Lond., 1858 : 339-352.

at 1859. Catalogue of the Batrachia Salientia in the collection of the British Museum. London, Taylor

& Francis : i-xvi + 1-160, pl. I-XII.

HEMMER, H. & ALCOVER, J. A. (eds.), 1984. — Historia biologica del ferreret. Ciutat de Mallorca, Mol :

1-252.

HOFFMANN, C. K., 1878. — Klassen und Ordnungen der Amphibien wissenschaftlich dargestelldt in

Wort und Bild. /n: BRONN, H. G., Die Klassen und Ordnungen des Thier-Reichs wissenschaftlich

dargestelldt in Wort und Bild, Leipzig & Heidelberg, Winter, 6 (2) : 1-726, pl. I-LII.

INGER, R. F., 1954. — Systematics and z0ogeography of Philippine Amphibia. Fieldiana : Zool., 33

183-531.

1967. — The development of a phylogeny of frogs. Evolution, 21 : 369-384.

IskANDAR, D. J., 1978. — À new species of Barbourula: first record of a discoglossid Anuran in Bornco.

Copeia, 1978 : 564-566.

KaWAMURA, T., NisH1okA, M. & UEDA, H., 1972. - Reproduction of the oriental fire-bellied toad,

Bombina orientalis, with special reference to the superiority of this species as a laboratory animal.

Sci. Report Lab. amph. Biol. Hiroshima Univ., 03-317.

KEFERSTEIN, W., 1868. — Ucber die Batrachier Australiens. Arch. Naturgesch., 34 (1) : 253-290, pl. V-

VII.

KUHN, O., 1965. — Die Amphibien. System und Stammesgeschichte. Krailling bei München, Ocben : 1-

102.

ir 1967. — Amphibien und Reptilien. Stuttgart, Gustav Fischer : i-vii + 1-124.

Lawza, B., Cet, J. M. & CRESPo, E., 1975. - Immunological evidence for the specific status of Dis-

coglossus pictus Otth, 1837 and D. sardus Tschudi, 1837, with notes on the families Discoglos-

sidae Günther, 1858 and Bombinidae Fitzinger, 1826 (Amphibia Salientia). Monit. zool. ital.,

(n.s.), 9 : 153-162.

eee 1976. — Further immunological evidence for the validity of the family Bombinidae (Amphibia Sal-

ientia). Monit. zool. ital., (n.s.), 10 : 311-314.

LaTasTE, F., 1878. — Division en familles naturelles des Batraciens Anoures d'Europe. Rev. int. Sci.,

2 : 488-499.

2 1879. — Division en familles naturelles des Batraciens Anoures d'Europe. C. r. Assoc. fr. Av. Sci.

7 : 758-765.

LAURENT, R. F., 1980. — Esquisse d'une phylogenèse des Anoures. Bull. Soc. zool. France, 104 : 397-

422

es 1986. — Sous-classe des Lissamphibiens (Lissamphibia). Systématique. Zn : P.-P. GRassé & M.

Source : MNHN, Paris

DuBois 67

DELSOL (eds.), Traité de zoologie, Tome XIV, Batraciens, Fasc. I-B, Paris, Masson : 594-797.

LAURENTI, J. N., 1768. — Specimen medicum, exhibens Synopsin Reptilium emendatam cum experimentis

circa venena et antidota reptilium austriacorum.… Viennae, Joan. Thom. Nob. de Trattnern : i-ii

+ 1-215, pl. I-V.

LINNAEUS, C., 1761. — Fauna Svecica. Editio altera, auctior. Stockholmiae, Laurentii Salvii : i-xlix +

1-79, pl. I-II.

Liv, C. C., 1935. - Types of vocal sac in the Salientia. Proc. Boston Soc. nat. Hist., 41 (3) : 19-40, pl.

4-8.

Liu, C.-C. & Hu, S.-C., 1961. — The Tailless Amphibians of China. (In Chinese). Shanghai : i-xvi + 1-

364, pl. I-VI + I-XXVIII.

LyncH, J. D., 1973. - The transition from archaic to advanced frogs. Jn: VIAL (1973) : 133-182.

MaxsoN, L. R. & SzymuRA, J. M., 1984. — Relationships among discoglossid frogs : an albumin per-

spective. Amphibia-Reptilia, 5 : 245-252.

MERREM, B., 1820. — Versuch eines Systems der Amphibien. Marburg, Johann Christian Krieger : 1-188

(*X 2) + 189-191, 1 pl.

MERTENS, R., 1955. — Proposed use of the plenary powers to conserve the generic name Bombina Oken,

1816 (class Amphibia, order Anura). Bull. zool. Nomencl., 11 : 132-133.

MERTENS, R. & MULLER, L., 1928. — Liste der Amphibien und Reptilien Europas. Abh. senckenberg.

naturf. Ges., 41 : 1-62.

--- 1940. - Die Amphibien und Reptilien Europas. (Zweite Liste, nach dem Stand vom 1. Januar 1940).

Abh. senchenberg. naturf. Ges., 451 : 1-56.

MERTENS, R. & WERMUTH, H., 1960. — Die Amphibien und Reptilien Europas. (Dritte Liste, nach dem

Stand vom 1. Januar 1960). Frankfurt am Main, Waldemar Kramer : i-xi + 1-264.

MIRANDA-RIBEIRO, À. DE, 1924. — Notas batrachologicas. Bol. Mus. nac. Rio de Janeiro, 1 : 137-143.

MivarT, G., 1869. - On the classification of the Anurous Batrachians. Proc. z0ol. Soc. Lond., 1869 :

280-295.

MULLER, F., 1878. — Katalog der im Museum und Universitätskabinet zu Basel aufgestellten Amphi-

bien und Reptilien nebst Anmerkungen. Verhandl. naturf. Ges. Basel, 6 : 559-709, pl. I-III.

NICHOLLS, G. E., 1916. - The structure of the vertebral column in the Anura Phaneroglossa and its

importance as a basis of classification. Proc. Linn. Soc. Lond., 128 : 80-92.

NIEDEN, F., 1923. - Amphibia. Anura I. Subordo Aglossa und Phaneroglossa, sectio 1 Arcifera. Das

Tierreich, 46 : 1-584.

Nosce, G. K., 1922. — The phylogeny of the Salientia. L. The osteology and the thigh musculature :

their bearing on classification and phylogeny. Bull. Am. Mus. nat. Hist., 46 : 1-87, pl. I-XXIII,

1 tab.

ce 1927. - The value of life history data in the study of the evolution of the Amphibia. Annals New

York Acad. Sai., 30 : 31-128, pl. I-IX.

me 1931. - The biology of the Amphibia. New York, Dover : i-xviii + 1-577.

OkaDa, Y., 1966. — Fauna Japonica. Anura (Amphibia). Tokyo, Biogcographical Society of Japan : i-

xii + 1-234, pl. I-XXIV.

OKEN, L. VON, 1816. — Lehrbuch der Naturgeschichte. Dritter Theil, Zoologie, Zweite Abtheilung. Jena,

August Schmid & Comp.: i-xvi + 1-1272.

OLMo, E., MORESCALCHI, À., STINGO, V. & ODIERNA, G., 1982. - Genome characteristics and the sys-

tematics of the Discoglossidae (Amphibia Salientia). Monit. zool. ital., (n.s.), 16 : 283-299.

OrrH, À, 1837. — Beschreibung einer neuen Europäischen Froschgattung, Discoglossus. N. Denkschr.

Schweizer. Ges. Naturwiss., 1 (4) : 1-7, 1 pl.

PasTEUR, G. & Bons, J., 1959. — Les Batraciens du Maroc. Trav. Inst. sci. chérif., (Zool.), 17 :

i-xvi + 1-246, pl. I-II, maps 1-3.

Pope, C. H., 1931. - Notes on Amphibians from Fukien, Hainan, and other parts of China. Bull. Am.

Mus. nat. Hist., 61 : 397-611, pl. XITI-XXII.

Porr, C. H. & BORING, À. M., 1940. — À survey of Chinese Amphibia. Peking nat. Hist. Bull., 15 :

13-86, 1 map.

REIG, O. A., 1958. — Proposiciones para une nueva macrosistematica de los Anuros. Physis, 21 : 109-

118.

SancHiZ, B., 1984. — Anlisis filogenético de la tribu Alytini (Anura, Discoglossidae) mediante el es-

tudio de su morfoestructura 6sea. /n: HEMMER & ALCOVER (1984) : 61-108.

Source : MNHN, Paris

68 ALYTES 6 (1-2)

SancHIZ, F. B. & ADROVER, R., 1979. — Anfibios fosiles del Pleistoceno de Mallorca. Doñana, Acta

vertebr., 4 : 5-25.

Savace, J. M., 1973. — The geographic distribution of frogs : patterns and predictions. /n : VIAL (1973) :

351-445.

SCHMIDT, K. P., 1927. - Notes on Chinese Amphibians. Bull. Am. Mus. nat. Hist., 54 : 553-575, pl.

XXXI-XXXII.

SCHREIBER, E., 1912. — Herpetologia Europaea. Jena, Gustav Fischer

SokoL, O. M., 1977. — A subordinal classification of frogs (Amphibia

505-508.

SrarRETT, P. H., 1973. - Evolutionary patterns in larval morphology. /n : VIAL (1973) : 251-271.

STEINDACHNER, F., 1867. — Reise der ôsterreichischen Fregatte Novara um die Erde in den Jahren 1857,

1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Band

L. Amphibien. Wien, aus der Kaiserlich-Kôniglichen Hof- und Staatsdruckerei : 1-70, pl. I-V.

STEJNEGER, L., 1907. — Herpetology of Japan and adjacent territory. Bull. US. nat. Mus., 58: i-xx +

1-577, pl. I-XXXV.

— 1925. - Chinese Amphibians and Reptiles in the United States National Museum. Proc. U. S. nat.

Mus., 66 (25) : 1-115.

TATARINOV, L. P., 1964. - Nadotryad Salientia. Prygayuséie, ili Beskhvostye. In : Y. A. ORLOV (ed.),

Osnovy Paleontologii, Tome 12, Zemnovodnye, Presmykayuséiesya i Priey. Moskva, Izdatelbstvo

“Nauka” : 125-133.

TAYLOR, E. H. & NOBLe, G. K., 1924. - À new genus of discoglossid frogs from the Philippine islands.

Am. Mus. Novit., 121 : 1-4.

TeoBaLD, W., 1868. — Catalogue of Reptiles in the Museum of the Asiatic Society of Bengal. . asiat.

Soc. Bengal, 1868 : i-iü + 1-88, 4 pl.

TIHEN, J. A., 1965. — Evolutionary trends in frogs. Am. Zool., 5 : 309-318.

TRUEB, L., 1973. - Bones, frogs, and evolution. /n : VIAL (1973) : 65-132.

TscHuDI, J. J., 1838. — Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Abth-

eilung der Reptilien. Neuchâtel, Petitpierre : 1-102, pl. I-VI.

24 1845. — Reptilium conspectus quae in Republica Peruana reperiuntur et pleraque observata vel col-

lecta sunt in itinere. Arch. Naturgesch., 11 : 150-170.

Van DENBURGH, J., 1912. - Notes on Ascaphus, the discoglossoid toad of North America. Proc. Calif.

Acad. Sci., (4), 3 : 259-264. x

VaL, J. L. (ed.), 1973. - Evolutionary biology of the Anurans. Columbia, University of Missouri Press :

i-xii + 1-470.

VIERTEL, B., 1984. - Habit, melanin pigmentation, oral disc, oral cavity and filter apparatus of the

larvae of Baleaphryne muletensis. In : HEMMER & ALCOVER (1984) : 21-43.

WAGLER, J., 1829. — Descriptiones et icones Amphibiorum. Jsis von Oken, 1829 : 68-70.

1830. — Natürliches System der Amphibien, mit vorangehender Classification der Säugethiere und Vôgel.

München, Stuttgart & Tübingen, Cotta : i-vi + 1-354.

+ 1-960.

nura). J. Zool., Lond., 182 :

Source : MNHN, Paris

Alytes, 1987, 6 (1-2): 69-74. 69

Strongylopus Tschudi, 1838 (Amphibia, Anura):

request for the designation under the plenary

powers of a type-species in agreement

with current usage

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

A detailed analysis shows that, as a result of Opinion 713 (1964) of the Inter-

national Commission on Zoological Nomenclature, the nominal genus Strongylopus

Tschudi, 1838 does not have at present a type-species, and that the possible designa-

tions of type-species available under the Rules would result in nomenclatural problems.

The ICZN is therefore asked to use its plenary powers to designate for this genus a type-

species in agreement with current usage.

Note. — This paper was submitted on 8 September 1980 to the Secretary of the International

Commission on Zoological Nomenclature for publication in the Bulletin of zoological Nomenclature, but,

despite repeated requests since then, has still not been published in this journal. The problem it raised

has therefore remained unresolved, which is unfortunate, especially when one considers that, since then,

several papers and checklists mentioning the generic name Strongylopus have been published (CLARKE,

1981 ; Duois, 1981 a, 1981 b, 1987 ; FROST, 1985 ; DUELLMAN & TRUEB, 1986), and others are in

progress. This paper is therefore published here as it was submitted, except that a few more recent

bibliographic references have been incorporated.

(1) After having long been considered a subjective synonym of the Amphibian generic

name Rana Linnaeus, 1758, the name Strongylopus Tschudi, 1838 was resurrected as a valid

generic name by VAN DiyK (1966). Since then it has been used as a valid generic (VAN DIXK,

1971, 1972 ; CHANNING & VAN DK, 1976 ; CHANNING, 1979 ; CLARKE, 1981 ; FROST, 1985 ;

DuELLMAN & TRUEB, 1986) or subgeneric (DUBOIS, 1981 a, 1981 b, 1987) name by some

authors, while others (PASSMORE & CARRUTHERS, 1979) still considered it as a synonym of

Rana. It is likely that this name will remain in use, at least to designate a subgenus of the

genus Rana s.l. However, evidence is below presented that the nominal genus Strongylopus

Tschudi, 1838 does not at present have a type-species, and that the possible designations of

type-species available under the Rules would result in nomenclatural problems. The

Commission is therefore requested to use its plenary powers to designate a type-species for

this nominal genus in agreement with current usage.

(2) The generic name Strongylopus was created by TscHUDI (1838 : 38, 78-79) for a

single nominal species, “Rana fasciata Boie”. As was shown by PARKER & RIDE (1962), BOIE

Source : MNHN, Paris

70 ALYTES 6 (1-2)

(1832 : 62) credited the name Rana fasciata to BURCHELL (1824 : 32). Therefore the type-

species of Strongylopus at its creation was Rana fasciata Burchell, 1824 by monotypy.

(3) As a result of PARKER & RIDE’s (1962) application and after the comments of SMITH

(1963) and PoyNTON (1963) about it, the Commission (Opinion 713 ; ANONYMOUS, 1964)

decided to suppress the name Rana fasciata Burchell, 1824, as well as all other uses of the

combination Rana fasciata prior to that by SMITH (1849), and to place the name Rana fas-

ciata Smith, 1849 on the Official List of Specific Names in Zoology.

(4) It is clear that the generic name Srrongylopus, coined in 1838, cannot have as its

type-species by monotypy a nominal species created in 1849. Furthermore, TsCHUDI (1838)

based his genus Strongylopus on specimens in the Leiden Museum which presumably belon-

ged to the species Rana grayi Smith, 1849 and not to Rana fasciata Smith, 1849 (see PARKER

& RIDE, 1962). In any case, both nominal species Rana fasciata and Rana grayi date from

SmirH’s (1849) paper, and cannot be the type-species of Strongylopus by original monotypy ;

they cannot either be considered as originally included in this genus, at least not as from

TscHuprs (1838) work. The genus Strongylopus is however available as of TSCHUDI (1838),

since based on a diagnosis. The only nominal species originally included in this genus having

been suppressed by the Opinion 713, the result is the same as if this genus had been created

without any nominal species included. Therefore the first author to have associated available

specific names to the name Strongylopus has fixed the originally included species of this ge-

nus.

(5) DUMÉRIL & BIBRON (1841 : 389) mention the combination Strongylopus fasciatus in

the synonymy of Rana fasciata. This work is still anterior to the work of SMITH (1849) and,

following the Opinion 713, the name Rana fasciata is still not available at that time.

As was noted by PARKER & RIDE (1962), SMITH (1849) had relied upon DUMÉRIL &

BiBRON’s (1841) work to define the species Rana fasciata. He had restricted the use of this

name to some of the specimens upon which DUMÉRIL & BIBRON had based their description

of this species, and which were considered by them to constitute a particular variety, “Va-

riété D”, of this species.

While SMITH (1963) and HUBBs (in ANONYMOUS, 1964) are certainly correct in stating

that a specific name, associated with a given species, should not be credited to an author to

whom this given species was unknown, it may be regretted that, when voting on the Opi-

nion 713 (see ANONYMOUS, 1964), the Commission was not proposed as a third alternative

in the Part 2 of the vote to validate the name Rana fasciata as of DUMÉRIL & BIBRON (1841)

and to designate for this nominal species, rather than a neotype, a lectotype, chosen among

the still extant specimens on which DUMÉRIL & BIBRON (1841) had based their description

of Rana fasciata. For this purpose, the specimen No. 396 in the Paris Museum collections,

an adult female collected by DELALANDE in the Cape region, which belongs to DUMÉRIL &

BiBRON’s “Variété D” and which is a typical Rana fasciata, would have been available. If

this had been done, Rana fasciata would be the type-species of Strongylopus by subsequent

monotypy but, unless changes are brought to the Opinion 713, it cannot be the case.

(6) FITZINGER (1843 : 31) mentioned “Strongyl. fasciatus Tschud.” as type-species of

Strongylopus, and GÜNTHER (1859 : 20) quoted “Strongylopus fasciatus, Tschudi” in the sy-

nonymy of “Rana fasciata, Boie”. Both these authors refer to a specific name, “fasciatus

Tschudi” (= fasciata Burchell) which is not available as having been suppressed by the Opi-

Source : MNHN, Paris

Duois 71

nion 713, and therefore cannot be construed as having designated a type-species for Stron-

gylopus or as having included a nominal species in this genus.

(7) The first author to have associated available specific names to Strongylopus is FIT-

ZINGER (1860 : 414), who wrote :

“Strongylopus Delalandii Fitz. (Rana Delalandii Dum. Bibr.) Cap.

Strongylopus fasciatus Tschudi (Rana fasciata Boic. -

Rana fasciata Dum. Bibr.) Cap.

Strongylopus oxyrhynchus Fitz. (Rana oxyrhynchus Sundevall.) Cap.

The first of these three names, Rana delalandii Duméril & Bibron, 1841 (nec Pyxice-

phalus delalandii Tschudi, 1838), is a senior synonym of Rana angolensis Bocage, 1866 (see

e.g. POYNTON, 1964 : 103), and applies to a species of the subgenus or genus Rana s. str.

The third name, Rana oxyrhynchus Smith, 1849, applies to a species of the subgenus or ge-

nus Ptychadena Boulenger, 1917 (see e.g. POYNTON, 1964 : 124). As for the second name,

it refers again to the suppressed name Rana fasciata Burchell, 1824 and is not available, all

the more that SMITH’s Rana fasciata is not even mentioned as a synonym.

Therefore two nominal species only, Rana delalandii Duméril & Bibron, 1841 and Rana

oxyrhynchus Smith, 1849 are to be considered as the species originally included in the genus

Strongylopus Tschudi, 1838, and are the only two species eligible for type-fixation in this ge-

nus. The fact that later STEINDACHNER (1867 : 21-22) mentioned the names Rana grayi Smith,

1849 (as a synonym of “Strongylopus grayi Steind.”) and Rana fasciata Smith, 1849 (as a sy-

nonym of “Strongylopus fasciatus Tschudi”) is of no relevance here, since STEINDACHNER’S

work is largely posterior to FITZINGER’s.

(8) None of the two species originally included in Strongylopus belongs to the group to

which this name is currently applied. If Rana delalandii Duméril & Bibron, 1841 was chosen

as the type-species of Strongylopus, this name would disappear as a subjective junior syno-

nym of Rana Linnaeus, 1758 (see e.g. DuBois, 1987 : 42): this would have no other incon-

venience than the need of coining a new name for the group of frogs including Rana fasciata

Smith, 1849 and Rana grayi Smith, 1849. If, on the other hand, Rana oxyrhynchus Smith,

1849 was chosen as the type-species of Strongylopus, this latter name should replace Prycha-

dena Boulenger, 1917 as the valid name of another subgenus or genus of African ranids. This

latter consequence would be most disturbing for the stability of nomenclature since the name

Ptychadena has been regularly used since its creation.

(9) In order to preserve the stability of nomenclature, an action of the Commission is

necessary. Two possible actions may be contemplated.

The first one, which I strongly advocate, would be to revert to the Opinion 713 and

to make the name Rana fasciata available as of DUMÉRIL & BIBRON (1841) ; the specimen

No. 396 in the Paris Museum would be designated as lectotype of this species and the des-

ignation of a neotype for Rana fasciata made by PARKER & RIDE (1962) would be annulated ;

following this action, the species Rana fasciata Duméril & Bibron, 1841 would automatically

become the type-species of Strongylopus Tschudi, 1838 by subsequent monotypy.

If the Commission refused to change the wording of the Opinion 713, the action to

take would be to suppress all previous designations of type-species for Strongylopus Tschudi,

1838, and to designate a type-species for this genus under the plenary powers. The choice

Source : MNHN, Paris

72 ALYTES 6 (1-2)

of this type-species could again be a matter of discussion : one could advocate the choice of

Rana grayi Smith, 1849, since this species, although under the name “Rana fasciata Boie”,

is likely to be the one on which TscHUDI (1838) had based his genus Strongylopus ; or one

could advocate the choice of Rana fasciata Smith, 1849, in order to follow previous des-

ignations (e.g. FITZINGER, 1843, POYNTON, 1964 and FROST, 1985 mentioned “Rana fas-

ciata” as the type-species of Strongylopus). Since Rana grayi and Rana fasciata are very clo-

sely related species (see e.g. POYNTON, 1964 and PASSMORE & CARRUTHERS, 1979), any genus

containing one of them is bound to contain also the other one, and both possible designa-

tions would be strictly equivalent in terms of generic content.

(10) Accordingly I ask the International Commission on Zoological Nomenclature to

use its plenary powers to settle the matters in this question, by choosing between the three

following alternatives :

(a) Alternative A.

(1) to use its plenary powers to change some parts of the Opinion 713 ; the fol-

lowing new wordings are to replace those which appear in the original Opinion under the

same numbers :

(1) (b) all other uses of the specific name fasciata, in the combination Rana fas-

ciata, prior to that by DUMÉRIL & BIBRON, 1841;

(2) (a) fasciata Duméril & Bibron, 1841, as published in the binomen Rana fas-

ciata, as interpreted by the specimen No. 396, in the Paris Museum collections, which is

hereby designated as lectotype of this species (Name No. 2042);

(3) (b) fasciata, all other uses of, in the combination Rana fasciata prior to that

by DUMÉRIL & BIBRON, 1841 (as suppressed under the plenary powers in (1) (b) above) (Name

No. 807);

(2) to change the wordings of the entry No. 2042 in the Official List of Specific

Names in Zoology and of the entry No. 807 in the Official Index of Rejected and Invalid Spe-

cific Names in Zoology according to (1) above;

(3) to place the generic name Strongylopus Tschudi, 1838 (gender : masculine),

type-species, by subsequent monotypy through DUMÉRIL & BIBRON (1841), Rana fasciata

Duméril & Bibron, 1841, on the Official List of Generic Names in Zoology.

(b) Alternative B.

(1) to use its plenary powers to suppress all previous fixations of type-species for

Strongylopus Tschudi, 1838 and to designate Rana grayi Smith, 1849, as type-species of this

genus;

(2) to place the generic name Strongylopus Tschudi, 1838 (gender : masculine),

type-species, by designation above in (1), Rana grayi Smith, 1849, on the Official List of Ge-

neric Names in Zoology.

Same as Alternative B, but with Rana fasciata Smith, 1849 instead of Rana grayi

Smith, 1849 designated as type-species of Sirongylopus Tschudi, 1838.

Source : MNHN, Paris

Dugois 73

RÉSUMÉ

Une analyse détaillée permet de montrer que, par suite de l’Opinion 713 (1964) de la

Commission Internationale de Nomenclature Zoologique, le genre nominal Strongylopus

Tschudi, 1838 ne possède pas à présent d’espèce-type, et que les désignations d’espèce-type

actuellement possibles en fonction du Code entraîneraient des problèmes nomenclaturaux.

En conséquence il est demandé à l’ICZN de faire usage de ses pleins pouvoirs pour désigner

pour ce genre une espèce-type qui soit en accord avec l’usage actuel.

LITERATURE CITED

ANONYMOUS, 1964. — Opinion 713. Rana fasciata Smith, 1849 (Amphibia): added to the official list

with suppression of Rana fasciata Burchell, 1824, under the plenary powers. Bull. zool. Nom.,

1: 352-354.

Boite, H., 1832. — Briefe von Heinrich Boie ; geschrieben aus Ostindien und auf der Reise dahin. Schleswig,

Kôniglichen Taubstummen Institut : 1-154.

BURCHELL, W. J., 1824. — Travels in the interior of Southern Africa. Vol. II. London, Longman & Co.:

i-viii + 1-648, pl. 1-10.

CHANNING, A., 1979. — Ecological and systematic relationships of Rana and Strongylopus in Southern

Natal (Amphibia : Anura). Ann. Natal Mus.. 23 : 797-831.

CHANNING, À. & Van DK, D. E., 1976. — À guide 10 the frogs of South West Africa. Durban, Univer-

sity of Durban, Westville Press : 1-47.

CLARKE, B. T., 1981. — Comparative osteology and evolutionary relationships in the African Raninae

(Anura Ranidae). Monit. zool. ital., (n. s.), 15, suppl. : 285-331.

Dusois, A., 1981 a. — Deux noms d’espèces préoccupés dans le genre Rana (Amphibiens, Anoures).

Bull. Mus. natn. Hist. nat., (4), 2 (A) : 927-931.

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

avec identification de leurs espèces-types : conséquences nomenclaturales. Monit. zool. ital.,

(n. s.), 15, suppl. : 225-284.

ue 1987. — Miscellanea taxinomica batrachologica (1). Alytes, 5 : 7-95.

DuELLMAN, W. E. & TRUEB, L., 1986. — Biology of Amphibians. New York, McGraw-Hill : i-xix + 1-

670.

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

Tome 8. Paris, Librairie encyclopédique de Roret : i-vii + 1-792.

FITZINGER, L., 1843. - Systema Reptilium. Fasciculus primus. Amblyglossae. Vindobonae, Braumüller

& Seidel : 1-106 + i-ix.

ee 1860. — Die Ausbeute der ôsterreichischen Naturforscher an Säugethieren und Reptilien während

der Weltumsegelung Sr. Majestät Fregatte Novara. Sitz.-ber. kais. Akad. Wiss. Wien, 42 : 383-

416.

FRosT, D. R. (ed.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst. Coll. :

Liv] + Ev + 1732.

GÜNTHER, À., 1859. — Catalogue of the Batrachia Salientia in the collection of the British Museum. Lon-

don, Taylor & Francis : i-xvi + 1-160, pl. I-XII.

PARKER, H. W. & RIpE, W. D. L., 1962. - Rana fasciata Burchell, 1824 (Amphibia) ; proposed des-

ignation of a neotype under the plenary powers. Z.N.(S.) 1253. Bull. zool. Nom., 19 : 290-292.

PASSMORE, N. I. & CARRUTHERS, V. C., 1979. — South African frogs. Johannesburg, Witwatersrand

University Press : i-xviii + 1-270.

PoyNTON, J. C., 1963. - Comment on the proposed use of the plenary powers to designate a neotype

for Rana fasciata Burchell, 1824. Z.N.(S.) 1253. Bull. zool. Nom., 20 : 255.

ee 1964. - The Amphibia of Southern Africa : a faunal study. Ann. Natal Mus., 17 : 1-334.

Source : MNHN, Paris

74 ALYTES 6 (1-2)

SmirH, A., 1849. — Jllustrations of the Zoology of South Africa. Reptlia. London, Smith, Elder & Co.:

pl. 1-78, pp. 1-28 (Appendix).

SurrH, H. M., 1963. - Comment on the proposed use of the plenary powers to designate a neotype for

Rana fasciata Burchell, 1824. Z.N.(S.) 1253. Bull. zool. Nom., 20 : 254.

STEINDACHNER, F., 1867. — Reise der ôsterreichischen Fregatte Novara um die Erde in den Jahren 1857,

1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Bd.

L. Amphibien. Wien, aus der Kaiserlich-Kôniglichen Hof- und Staatsdruckerei : 1-70, pl. 1-V.

TscHuDr, J. J., 1838. - Classification der Batrachier, mit Berücksichtigung der fossilen T'hiere dieser Abthei-

lung der Reptilien. Neuchâtel, Peitpierre : 1-102, pl. 1-VI.

Van Du, D. E., 1966. — Systematic and field keys to the families, genera and described species of

Southern African Anuran tadpoles. Ann. Natal Mus., 18 : 231-286.

ee 1971. - Anuran ecology in relation particularly to oviposition and development out of water. Zoo.

afr., 6 : 119-132.

ee 1972. - The behaviour of Southern African Anuran tadpoles with particular reference to their ccol-

ogy and related external morphological features, Zool. afr., 7 : 49-55.

Source : MNHN, Paris

Alytes, 1987, 6 (1-2) : 75-84. 75

Elachistocleis Parker, 1927 (Amphibia, Anura) :

proposed conservation

Alain DuBois

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

The overlooked fact that DuméRi. & BiBrON (1841) were the first authors to va-

lidly designate a type-species. namely Rana ovalis Schneider, 1799, for the nominal ge-

nus Engystoma Fitzinger, 1826, is stressed. Engystoma is therefore the valid name for

the microhylid genus now universally known as Elachistocleis Parker, 1927. The no-

menclatural consequences of this fact are discussed, both as concerns genus-group and

family-group names, and the ICZN is requested to use its plenary powers to designate

the nominal species Rana gibbosa Linnaeus, 1758 as type-species of Engystoma in or-

der to solve these problems.

Note. — This paper was submitted on 15 October 1982 to the Secretary of the International

Commission on Zoological Nomenclature for publication in the Bulletin of zoological Nomenclature, but,

despite repeated requests since then, has still not been published in this journal. The problems is raised

have therefôre remained unresolved, which is unfortunate, especially when one considers that, since

then, several publications mentioning the names discussed below have appeared, including three check-

lists (HARDING, 1983 ; DuBois, 1984 ; FRosT, 1985) and two important monographs (DUELLMAN &

TRUEB, 1986 ; LAURENT, 1986), and others are in progress. This paper is therefore published here as

it was submitted, except that a few recent bibliographic references have been incorporated, as well as

an additional discussion made necessary by the changes introduced in Art. 32 and 35 of the new Code.

(1) MERREM (1820 : 177) created the Amphibian genus Breviceps for the single no-

minal species Rana gibbosa Linnaeus, 1758, which is therefore the type-species of this genus

by monotypy. This genus, which now includes more than ten species of African frogs, is

now a member of the Brevicipitinae subfamily of the family Microhylidae.

(2) FITZINGER (1826 : 40) created the genus Engystoma. He mentioned the species Rana

gibbosa Linnaeus, 1758 as a “Repräsentant” (an example) of his new genus, but did not clearly,

formally, designate this species as type of his new genus. Beside this “Repräsentant” he also

mentioned other nominal species as being part of the new genus. On page 40 he listed, be-

side Rana gibbosa, the names Rana bufonia, Bufo ventricosus, Bombinator ventricosus, Bom-

binator systoma and Pipa laevis ; on page 65, in the list of specimens in the collection of the

Wien Museum, he quoted the species Rana ovalis, Rana gibbosa and Rana venitricosa as

members of the genus Engystoma. Therefore the following nominal species may be consi-

dered as the originally included species of the genus Engvstoma Fitzinger, 1826, among which

one may be chosen and designated as type-species of this genus : Rana gibbosa Linnaeus,

1758 ; Rana bufonia Merrem, 1820 ; Rana ventricosa Linnaeus, 1758 ; Rana svstoma Schnei-

der, 1799 ; Bufo laevis Daudin, 1802 ; Rana ovalis Schneider, 1799.

Source : MNHN, Paris

76 ALYTES 6 (1-2)

(3) WaGLeR (1830 : 205) proposed the replacement name Systoma for Engystoma Fit-

zinger, 1826. He included in it only one nominal species, “Breviceps gibbosus Merr.” (which

he considered synonym of “Rana Systoma Schn. Daud.”), but did not formally designate it

as type-species of this genus.

(4) TscHupt (1838 : 49, 86) used the generic name Systoma Wagler, 1830, which he

considered synonym of Breviceps Merrem, 1820 and of Engystoma Fitzinger, 1826, for the

nominal species Rana systoma Schneider, 1799, Rana gibbosa Linnaeus, 1758 and “Breviceps

leschenaulti Bibron” (nomen nudum). He did not designate a type-species for the genus.

(5) DuMÉRIL & BIBRON (1841 : 738-746) used the generic name Engystoma Fitzinger,

1826 for five nominal species, and were the first authors to make a designation of type-spe-

cies valid in the light of the present Code, using the word “type” : they chose the species

Rana ovalis Schneider, 1799 (DUMÉRIL & BIBRON, 1841 : 740). This species being one of

the originally included species of the genus, this designation is valid.

(6) FITZINGER (1843 : 33) retained the replacement name Systoma Wagler, 1830 for

the genus he had created under the name Engystoma Fitzinger, 1826. For this genus he des-

ignated the species Rana gibbosa Linnaeus, 1758 as type-species. This designation is not va-

lid, being posterior to that of DUMÉRIL & BIBRON (1841) for Engystoma : Systoma being a

replacement name for Engystoma, both nominal genera have the same type-species, namely

Rana ovalis Schneider, 1799.

(7) Following DUMÉRIL & BIBRON (1841), various authors, including GUNTHER (1859 :

51), BOULENGER (1882 : 160), Core (1875 : 30 ; 1887 : 18 ; 1889 : 385), GaDOW (1901 : 231)

and others, used Engystoma Fitzinger, 1826 for Rana ovalis Schneider, 1799 and related spe-

cies.

(8) STEJNEGER (1910) published a paper entitled : “The Amphibian generic name En-

gystoma untenable.” He argued that Engystoma Fitzinger, 1826 was “based on Linné’s Rana

gibbosa” and was therefore a junior objective synonym of Breviceps Merrem, 1820. For Rana

ovalis Schneider, 1799 and related species, he resurrected the generic name Gastrophryne Fit-

zinger, 1843 (type-species by original designation : Engystoma rugosum Duméril & Bibron,

1841).

(9) PARKER (1927) proposed a generic revision of the frogs which had previously been

referred to the genus Gastrophryne, for which he recognized six distinct genera. In particu-

lar, he created the genus Elachistocleis for the two nominal species Rana ovalis Schneider,

1799 (which PARKER designated as type-species of this new genus) and Oxyrkynchus bicolor

Guérin-Méneville, 1838.

(10) Since PARKER's (1927) work, and except for a few exceptions (ANDERSSON, 1945 :

2 ; BARTH, 1958 : 82 ; and possibly a few others), zoologists have given up using the generic

name Engvstoma Fitzinger, 1826 and h sed the generic name Elachistocleis Parker, 1927

for Rana ovalis Schneider, 1799 and Oxyrhynchus bicolor Guérin-Méneville, 1838 (the latter

being considered synonym of the former by some authors). The following authors used the

name Elachistocleis : MERTENS (1929 : 286 ; 1930 : 163 ; 1957 : 47) ; CRAWFORD (1931 : 38) ;

DuNN (1931 : 416 ; 1949 : 2) ; PARKER (1933 : 3; 1934 : 120) ; FREIBERG (1942: 220 ; 1951 :

1. Contrary to what is stated in FROST {1985 : 379), the author of this name is GUÉRIN-MÉNEVILLE {1838 : 17), not

VALENCIENNES (18381

Source : MNHN, Paris

Dugois 77

330) ; Myers (1942 : 155) ; PENTZ (1943: 182); SCHMIDT & INGER (1951 : 448) ; BOKER-

MANN (1952 : 279 ; 1966 : 34) ; TayLoR (1952 : 594) ; ARAMBURU (1953 : 273) ; CARVALHO

(1954 : 3) ; GRIFFITHS (1954 : 42) ; COCHRAN (1954 : 362 ; 1961 : 62) ; CEI (1955 : 291 ;

1956 : 65 ; 1968 : 211 ; 1980 : 62) ; BARTH (1958 : 82) ; GINÉS (1959 : 140) ; LEGRAND (1959 :

49) ; STEBBINS & HENDRICKSON (1959 : 538) ; Gans (1960 : 290) ; RIVERO (1961 : 177 ;

1966 a : 304 ; 1966 b : 493) ; CEI & RoïG (1961 : 34) ; ACHENBACH (1962 : 265) ; SICK (1965 :

81) ; GALLARDO (1966 : 77 ; 1979 : 301) ; SAVAGE (1966 : 728) ; NELSON (1967 : 3719B ;

1968 : 170 ; 1972 : 897 ; 1973 : 163) ; KENNY (1969 : 12) ; MEDEM (1969 : 160) ; COCHRAN

& Goin (1970 : 77) ; HUTCHISON (1971 : 119) ; NELSON & GUTTMAN (1973 : 424) ; TRUEB

(1973 : 71) ; WALKER (1973 : 6) ; BRAUN & BRAUN (1974 : 35 ; 1976 : 4) ; GORHAM (1974 :

119) ; LESCURE (1975 : 79 ; 1976 : 512) ; BOGART & NELSON (1976 : 199) ; EMERSON (1976 :

547) ; DOWLING & DUELLMAN (1978 : 40.2) ; DUELLMAN (1979 : 18) ; HOOGMOED (1979 :

273) ; LAURENT (1979 : 60 ; 1986 : 745) ; LyNCH (1979 : 193) ; RIVERO-BLANCO & DIxON

(1979 : 290) ; TYLER (1979 : 87) ; HARDING (1983 : 47) ; Dugois (1984 : 15, 20-21, 40) ;

FRoST (1985 : 378) ; DUELLMAN & TRUEB (1986 : 552).

(1) In view of the fact that FITZINGER (1826) did not validly designate a type-species

for his new genus Engystoma, and that DUMÉRIL & BIBRON (1841) validly designated Rana

ovalis as type-species of this genus, Engystoma should be removed from the synonymy of

Breviceps, where it erroneously stands since STEJNEGER’s (1910) paper. Engystoma Fitzinger,

1826 is in fact a senior objective synonym of Elachistocleis Parker, 1927 and should replace

the latter as the valid name of the genus. Such a replacement would be inappropriate, since

the name Elachistocleis has had a large use after its creation, and since the name Engystoma

has been almost universally rejected since the creation of Elachistocleis. An action of the

Commission is therefore needed in order to conserve the name Elachistocleis.

(2) Two different kinds of action could be considered :

(a) The first one would be the suppression, under the plenary powers, of the generic

names Engystoma Fitzinger, 1826, Systoma Wagler, 1830 (a replacement name of the latter)

and also Engistoma Peracca, 1904 (an unjustified emendation of Engystoma), and the place-

ment of the name Elachistocleis Parker, 1927 on the Official List. As shown below in (16),

it would also be advisable in this case, although not absolutely necessary, to suppress the

family-group names Engystomidae Bonaparte, 1850 (and Engystomina Bonaparte, 1850),

Systomata Lichtenstein, 1856 and Engistomatidae Methuen & Hewitt, 1913. Thus the sup-

pression of either three or six names by the Commission would be necessary in this case.

(b) The second possibility would be setting aside by the Commission of all previous

designations of type-series for Engystoma (and its replacement names), including DUMÉRIL

& BIBRON’s (1841) valid designation, and designation by the Commission, under the plenary

powers, of Rana gibbosa Linnaeus, 1758 as type-species of this genus. This action would be

equivalent to a validation of STEJNEGER’S (1910) action considering FITZINGER’s (1826) men-

tion of Rana gibbosa as a “Repräsentant” of his genus Engystoma as a valid type-species des-

ignation for this genus. In this case the name Engystoma and its replacement names would

remain in the synonymy of Breviceps Merrem, 1820, where STEJNEGER (1910) had placed

them, and the name Elachistocleis Parker, 1927 would remain valid. The family-group names

mentioned above in (a) would disappear as junior synonyms, as discussed below in (15), so

that no suppression of name would be needed. This second possibility of action is chosen

here, because it is far more economic than the first one.

Source : MNHN, Paris

78 ALYTES 6 (1-2)

(13) Following STEJNEGER (1910), PARKER (1934) considered Engystoma Fitzinger, 1826

a junior synonym of Breviceps Merrem, 1820. For the subfamily accomodating this genus he

strangely used the name Brevicipitinae, which he credited to CoPE (1867), instead of two

other older family-group names mentioned by him : Systomata Lichtenstein, 1856 and “En-

gystomatidae Günther, 1858”. Since then the name Brevicipitinae has remained in use as the

valid name of the subfamily : see e.g. GOIN & Go (1962 : 234) ; POYNTON (1964 : 69) ;

WAGER (1965 : 40) ; VAN DuyK (1966 : 247) ; LyYNCH (1973 : 117) ; SAVAGE (1973 : 354) ;

BoGarT & NELSON (1976 : 201) ; DOWLING & DUELLMAN (1978 : 40.3, under the incorrect

spelling Brevicipinae) ; GOIN, GoiN & ZUG (1978 : 228) ; LAURENT (1979 : 60 ; 1980 a : 412 ;

1980 b : 86 ; 1986 : 749) ; BOGART & TaNDY (1981 : 59, under the incorrect spelling Bre-

vicepinae) ; MORESCALCHI (1981 : 44) ; DuBois (1983 : 275 ; 1984 : 40 ; 1985 : 75) ; FROST

(1985 : 355) ; DUELLMAN & TRUEB (1986 : 552).

(14) As a matter of fact, all authors until 1979 have overlooked the family-group names

created by BONAPARTE in a table (1850 ; reprinted in article form in 1852) in a work which

was recently rediscovered (DUBoIS, 1981, 1982). BONAPARTE (1850) was the first author to

propose family-group names based on the generic names Breviceps Merrem, 1820 and En-

gystoma Fitzinger, 1826 : he created the name Engystomidae for a family including three

subfamilies, one of which he called Engystomina ; besides, he created the name Brevicipi-

tina for a subfamily of his family Bufonidae.

(5) If the generic name Engystoma Fitzinger, 1826 remains, after action of the

Commission, an objective synonym of Breviceps Merrem, 1820, the family-group names Bre-

vicipitinae and Engystomatinae are also objective synonyms. Since both names were created

in the same publication by BONAPARTE (1850), a first-reviser action is needed to decide upon

their relative priority. I therefore took such an action (DUBoIS, 1983 : 275, 1984 : 40) and

selected the name Brevicipitinae Bonaparte, 1850, in order to maintain the stability of the

nomenclature of this subfamily (see (13) above). The name Engystomatinae Bonaparte, 1850

therefore automatically disappeared as a junior objective synonym of Brevicipitinae and its

existence does not threaten the stability of nomenclature. The same applies to the family-

group names Systomata Lichtenstein, 1856 and Engistomatidae Methuen & Hewitt, 1913,

names which are based upon generic names which are substitute names for Engystoma Fit-

zinger, 1826.

(16) If the Commission refused to follow the suggestion made above, and if Engystoma

Fitzinger, 1826 remained an objective senior synonym of Elachistocleis Parker, 1927, the name

Engystomatinae Bonaparte, 1850 would become a junior subjective synonym of Microhyli-

nae Günther, 1858 (1843), the name traditionally used for this subfamily. Thus its existence

would not cause a threat to the stability of nomenclature. However, if new family-groups

were to be created (for example if the subfamily Microhylinae was to be subdivided into se-

veral tribes), the name Engystomatinae would remain available for the group including Rana

ovalis Schneider, 1799. If, on the other hand, the name Engvstoma Fitzinger, 1826 had been

suppressed by the Commission, it would be unadvisable to use the name Engystomatinae for

the family-group including the genus Elachistocleis Parker, 1927. To avoid such possible dif-

ficulties, it would therefore be justified to ask the Commission to suppress, beside the ge-

neric names Engystoma, Systoma and Engistoma, the familv-group names Engystomidae Bo-

naparte, 1850 (and Engystomina Bonaparte, 1850), Systomata Lichtenstein, 1856 and

Engistomatidae Methuen & Hewitt, 1913. This would make the solution discussed above in

(12a) even heavier, and this reason strengthens the case for choosing the solution (12b).

Source : MNHN, Paris

Dusois 79

(7) A few words must be said about the familial name Microhylidae. The family-group

name Microhylinae was coined by NOBLE (1931 : 537) for a subfamily including the genus

Microhyla Tschudi, 1838 and related genera. For the family he used the name Brevicipiti-

e. PARKER (1934 : 15-16) remarked that the two oldest family-group names available for

mily were Hylaedactyli Fitzinger, 1843 and Gastrophrynae Fitzinger, 1843. PARKER

(1934 : 16) took a first-reviser action concerning these two names, by selecting the latter

against the former. However, both these family-group names were based on generic names

considered by PARKER (1934) as junior synonyms of valid names. Gastrophryne Fitzinger, 1843

was placed by him in the synonymy of Microhyla Tschudi, 1838, and consequently PARKER

(1934 : 16) replaced the name Gastrophrynae Fitzinger, 1843 by the name Microhylidae as

the valid name of the family. He was followed by all authors until now, and the name Mi-

crohylidae is universally used by present-days batrachologists : see e.g., among many others,

BouRRET (1942 : 479) ; Liu (1950 : 231) ; TAYLOR (1952 : 594 ; 1962 : 539) ; COCHRAN

(1954 : 1) ; GRIFFITHS (1954 : 37 ; 1999 : 473 ; 1963 : 273) ; INGER (1954 : 414 ; 1966 : 116 ;

1967 : 370) ; REIG (1958 : 115) ; RIVERO (1961 : 177) ; GoIN & GoIN (1962 : 197) ; PoyN-

TON (1964 : 69) ; OKADA (1966 : 41) ; COCHRAN & GoIN (1970 : 74) ; ESTES & RE1G (1973 :

37) ; LYNCH (1973 : 135 ; 1979 : 192) ; SAVAGE (1973 : 352) ; STARRETT (1973 : 251) ; TRUEB

(973 : 71) ; DuBois (1974 : 352 ; 1983 : 274 ; 1984 : 38 ; 1985 : 75) ; GORHAM (1974 : 115) ;

DUELLMAN (1975 : 5 ; 1979 : 3) ; DOWLING & DUELLMAN (1978 : 40.1) ; GoIN, GoiN & ZUG

(1978 : 69) ; GUIBÉ (1978 : 92) ; HOOGMOED (1979 : 251) ; LAURENT (1979 : 56 ; 1980 a :

410 ; 1980 b : 85 ; 1986 : 736) ; TYLER (1979 : 79) ; CEI (1980 : 152) ; HARDING (1983 :

46) ; FROST (1985 : 349) ; DUELLMAN & TRUEB (1986 : 549). The name Gastrophrynae ha-

ving been changed into Microhylidae on the basis of a subjective synonymy of the generic

names Gastrophryne and Microhvla, and the name Microhylidae having won general accep-

tance since then, the latter name must be maintained as the valid name of the family by vir-

tue of Art. 40(a). This name takes the date of the rejected name (Art. 40b), and should lo-

gically be quoted as Microhylidae Noble, 1931 (1843) ; similarlv, the nominotypical subfamily

should logically be known as Microhylinae Noble, 1931 (1843) (see DUBOIS, 1983, 1984).

However, as a result of the recent wordings of Articles 32 and 35 of the Code (ANONYMOUS,

1985), which I criticized in detail elsewhere (DUBOIS, 1985), the names Microhylidae and

Microhvlinae must now be credited to GÜNTHER (1858), who had created the taxa names

Micrhylina and Micrhylidae, both based on the genus-group name Micrhvla Duméril & Bi-

bron, 1841, an unjustified emendation of Microhyla Tschudi, 1838 (for more details see Du-

BOIS, 1984, 1985, 1987).

(18) To conclude, the International Commission on Zoological Nomenclature is re-

quested :

(D to use its plenary powers to set aside all previous type-species designations for the

genus Engvstoma Fitzinger, 1826, and to designate the nominal species Rana gibbosa Lin-

naeus, 1758 as type-species of this genus :

(2) to place the following names on the Official List of Generic Names in Zoology :

(a) Breviceps Merrem, 1820, type-species, by monotypy, Rana gibbosa Linnaeus,

1758 ;

(b) Microhvla Tschudi, 1838, type-species, by monotvpy, Microhyla achatina

Tschudi, 1838 ;

Source : MNHN, Paris

80 ALYTES 6 (1-2)

Schneider, 1799 ;

(3) to place the following names on the Official List of Specific Names in Zoology :

(a) gibbosa Linnaeus, 1758, as published in the binomen Rana gibbosa (type-species

of Breviceps Merrem, 1820) ;

(b) achatina Tschudi, 1838, as published in the binomen Microhyla achatina (type-

species of Microhyla Tschudi, 1838) ;

Elachistocleis Parker, 1927) ;

(4) to place the following names on the Official List of Family-Group Names in Zoo-

logy :

(a) Microhylidae Günther, 1858 (1843), type-genus Microhyla Tschudi, 1838 ;

(b) Brevicipitina Bonaparte, 1850, type-genus Breviceps Merrem, 1820.

RÉSUMÉ

DUMÉRIL & BIBRON (1841) s’avèrent être les premiers auteurs à avoir effectué une dé-

signation valide d’espèce-type, en l'occurrence Rana ovalis Schneider, 1799, pour le genre

nominal Engystoma Fitzinger, 1826. Il en résulte que le nom Engystoma est en fait le nom

valide pour le genre de Microhylidae actuellement connu sous le nom Elachistocleis Parker,

1927. Les conséquences nomenclaturales de ce fait sont discutées, et il est demandé à l'ICZN

de faire usage de ses pleins pouvoirs pour désigner l’espèce nominale Rana gibbosa Linné,

1758 comme espèce-type du genre Engystoma, de manière à résoudre ces problèmes.

LITERATURE CITED

ANONYMOUS, 1985. — International code of zoological nomenclature. Third edition. London, International

Trust for zoological Nomenclature : i-xx + 1-338.

ACHENBACH, G. M., 1962. - Contribuciôn al conocimiento de Batracios que viven en el departamento

de La Capital de la provincia de Sante Fe (Republica Argentina). /n : Anais do Segundo Congresso

Latino Americano de Zoologia, 2 : 261-285.

ANDERSSON, L. G., 1945. — Batrachians from East Ecuador. Ark. Zool., 37 A (2) : 1-88.

ARSMBURU, À. $. À. DE, 1953. — El género Hypopachus Keferstein en la Argentina (Batrachia : Salien-

ta). Notas del Museo Peron, 16 (Zool. 144) : 271-285.

BaRTH, R., 1958. - À faune do Parque nacional do Itatiaia. Bol. Parg. nac. Hiatiaia, 6 : 1-150.

BoGarT, J. P. & NELSON, C. E., 1976. Evolutionary implications from karvotypic analysis of frogs

of the families Microhylidae and Rhinophrynidac. Herpetologica, 32 : 199-208.

BoGarT, J. P. & TaNDy, M., 1981. - Chromosome lineages in African ranoid frogs. Monit. sool. ital.,

fn: 8), 16, suppl. : 55-91.

BOKERMANN, W. C. A., 1952. — Microhylidae da colecao do departamento de Zoologia (Amphibia-

Anura). Pap. avul. Dep. Zool., 10 : 271-292, 4 pl.

1966. — Lista anotada das localidades tipo de Anfibios brasileir

BONAPARTE, C. L., 1850. — Conspectus svstematum Herpetologiae et

mata. Lugduni Batavorum, Brill : 1 pl.

so Paulo, RUSP : 1-183.

mphibiologiae. Editio altera refor-

Source : MNHN, Paris

Dugois 81

Se 1852. - Conspectus systematum Herpetologiae et Amphibiologiac. Nuovi Ann. Sci. nat. Bologna,

(G), 5 : 477-480.

BOULENGER, G. A., 1882. — Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British

Museum. London, Taylor & Francis : i-xvi + 1-503, pl. I-XXX.

BouRRET, R., 1942. — Les Batraciens de l'Indochine. Hanoï, Institut océanographique de l’Indochine :

ix + 1-547, 4 pl.

BRAUN, P. C. & BRAUN, C. A. S., 1974. — Fauna da fronteira Brasil-Uruguai, lista dos Anfibios dos

departamentos de Artigas, Rivera e Cerro Largo. Zheringia, (Zool.), 45 : 34-49.

1976. - Contribuicäo ao estudo da fauna anfibiolôgica de regiäo metropolitana (Grande Porto Alegre),

Rio Grande do Sul, Brasil. Com. Mus. Ci. PUCRGS, Porto Alegre, 10 : 1-16.

CarvaLHo, A. L. DE, 1954. - A preliminary synopsis of the genera of American microhylid frogs. Occ.

Pap. Mus. Zool. Univ. Michigan, 555 : 1-20, pl. I.

Cet, J. M., 1955. — Chacoan Batrachians in central Argentina. Copeia, 1955 : 291-293, pl. II.

= 1956. - Nueva lista sistematica de los Batracios de Argentina y breves notas sobre su biologia y

ecologia. /nv. Zool. chil., 3 : 35-68, pl. I-IX.

ee 1968. — Distribution et spécialisation des Batraciens sud-américains. Jn : Biologie de l'Amérique

australe, Paris, CNRS, 4 : 199-214.

me 1980. - Amphibians of Argentina. Monit. zool. ttal., (n. s.), monogr. 2 : i-xii + 1-609, pl. 1-16

+ 1-24.

Cet, J. M. & RoiG, V. G., 1961. — Batracios recolectados por la expediciôn biolôgica Erspamer en Cor-

rientes y Selva Oriental de Misiones. Notas biol. F Ci. Corrientes, (Zool.), 1 : 1-40.

Cocra, D. M., 1954. - Frogs of Southeastern Brazil. U. S. nat. Mus. Bull., 206 : i-xvi + 1-423,

pl. 1-34.

= 1961. — Type specimens of Reptiles and Amphibians in the U. S. national Museum. U. $. nat.

Mus. Bull., 220 : i-xv + 1-291.

CocHRAN, D. M. & GoiN, C. J., 1970. — Frogs of Columbia. U. S. nat. Mus. Bull., 288 : i-xii + 1-655.

Cor, E. D., 1867. - On the families of the raniform Anura. 7. Acad. nat. Sci. Phila., (2), 6 : 189-206.

Check-list of North American Batrachia and Reptilia. Bull. U. S. nat. Mus., 104.

. - Catalogue of Batrachians and Reptiles of Central America and Mexico. Bull. U. S. nat.

Mus., 32 : 1-98.

- 1889. - The Batrachia of North America. Bull. U. S. nat. Mus., 34 : 1-525, pl. I-LXXXVI.

CRAWFORD, S. C., 1931. — Field keys to the Lizards and Amphibians of British Guiana. Ann. Carnegie

Mus., 21 : 11-42.

DowLiNG, H. G. & DUELLMAN, W. E., 1978. — Systematic herpetology : a synopsis of families and

higher categories. Publications in Herpetology, Hiss Publications, New York, 7 : i-vii + 1.1-118.3

+ vi.

Dusois, A., 1974. — Liste commentée d'Amphibiens récoltés au Népal. Bull. Mus. nat. Hist. nat., (3),

213 (Zool. 143) : 341-411.

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

avec identification de leurs espèces-types : conséquences nomenclaturales. Monit. zool. ital., (n.

s.), 15, suppl. : 225-284.

- 1982. - Dendrobates Wagler, 1830 and Dendrobatidae Cope, 1865 (Amphibia, Anura) : proposed

conservation. Bull. zool. Nom., 39 : 267-278.

en 1983. — Classification et nomenclature supragénérique des Amphibiens Anoures. Bull. Soc. linn.

Lyon, 52 : 270-276.

La nomenclature supragénérique des Amphibiens Anoures. Mém. Mus. nat. Hit. nat., (A),

1-64.

Miscellanea nomenclatorica batrachologica (VID). Alytes, 4 : 61-78.

- 1987. — Living amphibians of the world : a first step towards a comprehensive checklist. Alyres,

5 : 99-149.

DUELLMAN, W. E., 1975. — On the classification of frogs. Occ. Pap. Mus. nat. Hist. Univ. Kansas, 42 :

1-14.

- 1979. - The South American herpetofauna : a panoramic view. Monogr. Mus. nat. Hist. Univ.

Kansas, 7 : 1-28.

DUuELLMAN, W. E. & TRUEB, L., 1986. — Biology of Amphibians. New York, McGraw-Hill : i-xix +

1-670.

Source : MNHN, Paris

82 ALYTES 6 (1-2)

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

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

Du, E. R., 1931. - The Amphibians of Barro Colorado Island. Occ. Pap. Boston Soc. nat. Hist., 5 :

403-421.

1949. — Notes on South American frogs of the family Microhylidac. Am. Mus. Novit., 1419 : 1-21.

EMERSON, S. B., 1976. — À preliminary report on the superficial throat musculature of the Microhy-

lidae and its possible role in tongue action. Copeia, 1976 : 546-551.

ESTEs, R. & REIG, O. A., 1973. — The early fossil record of frogs : a review of the evidence. /n : Vial.

(1973) : 11-63.

FITzINGER, L. L., 1826. - Neue Classification der Reptilien. Wien, Heubner : i-viii + 1-66, 1 tab.

ystema Reptilium. Fasciculus primus. Amblyglossae. Vindobonae, Braumüller & Seidel :

1-106 + i-ix.

FREIBERG, M. À., 1942. - Enumeraciôn sistemätica y distribuciôn geogräfica de los Batracios argenti-

nos. Physis, 19 : 219-240.

ES 1951. — Sobre la distribuciôn geogräfica de Elachistocleis ovalis bicolor (Valenciennes). Physis, 20 :

330-331.

Frost, D. R. (ed.), 1985. — Amphibian species of the world. Lawrence, Allen Press & Assoc. Syst. Coll. :

Liv] + iv + 1-732.

GaDoW, H., 1901. - Amphibia and Reptiles. London, Macmillan & Co. : i-xiii + 1-668, 1 pl.

GaLLaRDo, J. M., 1966. — Zoogeografia de los Anfibios chaqueños. Physis, 26 : 67-81.

ne 1979. - Composiciôn, distribuciôn y origen de la herpetofauna chaqueña. Monogr. Mus. nat. Hist.

Univ. Kansas, 7 : 299-307.

Gans, C., 1960. — Notes on a herpetological collecting trip through the Southeastern lowlands of Bo-

livia. Ann. Carnegie Mus., 35 : 283-314.

GINÉS, H., 1959. — Familias y géneros de Anfibios (Amphibia) de Venezuela. Mem. Soc. Ci. nat. La

Salle, 19 : 84-146.

Goix, C. J. & GoiN, O. B., 1962. — Introduction 10 herpetology. San Francisco & London, Freeman &

Co. : i-ix + 1-341.

Goiw, C. J., GoiN, O. B. & ZuG, G. R., 1978. — Introduction to herpetology. Third edition. San Fran-

cisco, Freeman & Co. : i-xiii + 1-378.

GorHaM, S. W., 1974. — Checklist of world Amphibians up 10 January 1, 1970. Saint-John, The New

Brunswick Museum : 1-173.

GRIFFITHS, L., 1954. - On the “otic element” in Amphibia Salientia. Proc. zool. Soc. Lond., 124 : 35-50,

pl. 14.

Eur 1959. The phylogeny of Sminthillus limbatus and the status of the Brachycephalidae (Amphibia

Salientia). Proc. zool. Soc. Lond., 132 : 457-487, pl. 14.

ee. 1963. — The phylogeny of the Salientia. Biol. Rev., 38 : 241-292.

GUÉRIN-MÉNEVILLE, F. E., 1838. - Jconographie du Règne Animal de G. Cuvier. Tome II. Texte ex-

tif. Reptiles. Paris, Baillière : 1-24, pl. 1-30.

GuiBé, J., 1978. - Les Batraciens de Madagascar. Bonn. 2ool. Monogr., 11 : 1-144, pl. 1-82.

GUNTHER, À., 1858. — On the systematic arrangement of the tailless Batrachians and the structure of

Rhinophrynus dorsalis. Proc. zool. Soc. Lond., 1858 : 339-352.

ee 1859. - Catalogue of the Batrachia Salientia in the collection of the British Museum. London, Taylor

& Francis : i-xvi + 1-160, pl. I-XII.

HARDING, K. A., 1983. — Catalogue of New World Amphibians. Oxford, Pergamon Press : i-xv + 1-406.

HooGmoEb, M. S., 1979. - The herpetofauna of the Guianan region. Monogr. Mus. nat. Hist. Univ.

Kansas, 7 : 241-279.

Hurcnison, V. H., 1971. — Herpetological expedition to Columbia, National Geographic Society Re-

search Reports, 1965 Projects : 113-122.

INGER, R. F., 1954. - Systematics and z0ogcography of Philippine Amphibia. Fieldiana :

183-531.

1966. — The systematics and zoogeography of the Amphibia of Bornco. Fieldiana : Zool., 52 : 1-402.

1967. - The development of a phylogeny of frogs. Evolution, 21 : 369-384.

KENNY, J. S., 1969. - The Amphibia of Trinidad. Stud. Fauna Curaçao Caribb. Isl., 29 (54) : 1-78,

pl: XV.

LAURENT, R. F., 1979. - Herpetofaunal relationships between African and South America. Monogr.

Mus. nat. Hist. Univ. Kansas, 7 : 55-71.

Zool., 33 :

Source : MNHN, Paris

Dugois 83

me 1980 a. - Esquisse d’une phylogenèse des Anoures. Bull. Soc. zool. France, 104 : 397-422.

- 1980 b. - Géonémie des Anoures. C. r. Soc. Biogéogr., 56 : 81-86.

- 1986. — Sous-classe des Lissamphibiens (Lissamphibia). Systématique. n : P.-P. GRASSÉ & M.

DELsoL (eds.), Traité de zoologie, 14, Batraciens, Fasc. I-B, Paris, Masson : 594-797,

LEGRAND, C. D., 1959. - Comunidades psamofilas de la region de Carrasco (Uruguay). An. Mus. Hist.

nat. Montevideo, (2), 6 (7) : 1-63, pl. I-XXIII.

LESCURE, J., 1975. — Biogéographie et écologie des Amphibiens de Guyane française. C. r. Soc. Bio-

géogr., 50 : 68-82.

---- 1976. - Contribution à l’étude des Amphibiens de Guyane française. VI. Liste préliminaire des

Anoures. Bull. Mus. nat. Hist. nat., (3), 377 (Zool. 265) : 475-524.

Liu, C.-C., 1950. - Amphibians of Western China. Fieldiana : Zool. Mem., 2 : 1-400, pl. 1-10.

Lyc, J. D., 1973. — The transition from archaic to advanced frogs. {n : VIAL (1973) : 133-182.

= 1979. - The Amphibians of the lowland tropical forests. Monogr. Mus. nat. Hist. Univ. Kansas,

7:189-215.

MeveM, F., 1969. — El desarrollo de la herpetologia en Colombia. Rev. Acad. colomb. Ci., 13 : 149-199.

MERREM, B., 1820. — Versuch eines Systems der Amphibien. Marburg, Johann Christian Krieger : 1-188

(x 2) + 189-191, 1 pl.

MERTENS, R., 1929. - Bemerkungen über einige brasilianische Froschlurche. Blätter Aquarien Terra

rienkunde, 40 : 286-287, pl. 37-38.

--- 1930. - Amphibien und Reptilien. Folia zool. hydrobiol., 1 : 161-166.

- 1957. - Froschfang-Exkursionen in Brasilien. 2. Säo Paulo. Agu. Ter. Z., 10 : 47-52.

MORESCALCHI, À., 1981. — Karyology of the main groups of African frogs. Monit. zool. ital., (n. s.),

15, suppl. : 41-53.

Myers, G. $., 1942. — Notes on some frogs from Peru and Ecuador. Proc. biol. Soc. Wash., 55 : 151-155.

NELSON, C. E., 1967. - The evolution of frogs of the family Microhylidae in North America. Diss.

Abstr., 27 : 3719B.

1968. — Ranas de la familia Microhylidae en la America del Norte. ZV Congreso Latinoamericano de

Zoologia, Caracas : 167-180.

er 1972. — Distribution and biology of Chiasmocleis panamensis (Amphibia : Microhylidae). Copeia,

1972 : 895-898.

ue 1973. - Mating calls of the Microhylinac : descriptions and phylogenctic and ecological conside-

rations. Herpetologica, 29 : 163-176.

NELSON, C. E. & GUTTMAN, S. L., 1973. - Serum protein electrophoresis of some Amphibia (Caecili-

dae, Rhinophrynidae, Microhylidae). Com. Biochem. Physiol., 44B : 423-428.

NoBLe, G. K., 1931. — The biology of the Amphibia. New York, Dover : i-xviii + 1-577.

Okapa, Y., 1966. — Fauna Japonica. Anura (Amphibia). Tokyo, Biogeographical Society of Japan : i-

xii + 1-234, pl. I-XXIV.

Parker, H. W., 1927. - The brevicipitid frogs allied to the genus Gastrophryne. Occ. Pap. Mus. Zool.

Univ. Michigan, 187 : 1-6, pl. I.

1933. — A list of the frogs and toads of Trinidad. Tropical Agriculture, 10 : 8-12.

- 1934. — À monograph of the frogs of the family Microhylidæe. London, British Museum : i-vii + 1-208.

PENTZ, K., 1943. - The cranial morphology of the ncotropical microhylid (Anura) Elachistocleis ovalis

(Schneider). South. Afr. J. Sci., 39 : 182-226.

POyNTON, J. C., 1964. - The Amphibia of Southern Africa : a faunal study. Ann. Natal Mus., 17 :

1-334.

REIG, O. A., 1958. — Proposiciones para una nueva macrosistematica de los Anuros. Physis, 21 : 109-118.

RIvERO, J. A., 1961. - Salientia of Venezuela. Bull. Mus. comp. Zool. Harvard Coll., 126 : 1-207, 1 pl.

ee 1966 a. — Salientios (Amphibia) en la coleccién de la Sociedad de Ciencias Naturales La Salle de

Venezuela. Caribb. J. Sci., 4 : 297-305.

_.. 1966 b. — The distribution of Venezuelan frogs. VI. The LLanos and the Delta Region. Caribb.

J. Sci, 4 : 491-495.

RIVERO-BLANCO, C. & DiIxON, J. R., 1979. - Origin and distribution of the herpetofauna of the Dry

Lowland Regions of Northern South America. Monogr. Mus. nat. Hist. Univ. Kansas, 7 : 281-298.

Savace, J. M., 1966. — The origins and history of the Central American herpetofauna. Copeia, 1966 :

719-766.

Source : MNHN, Paris

84 ALYTES 6 (1-2)

1973. - The geographic distribution of frogs : patterns and predictions. /n : ViAL (1973) : 351-445.

ScHMIDT, K. P. & INGER, R. F., 1951. - Amphibians and Reptiles of the Hopkins-Branner expedition

to Brazil. Fieldiana : Zool., 31 : 439-465.

Sick, H., 1965. — À fauna do Cerrado. Argu. Zool. Säo Paulo, 12 : 71-93.

STARRETT, P. H., 1973. - Evolutionary patterns in larval morphology. /n : Via (1973) : 251-271.

STEBBINS, R. C. & HENDRICKSON, J. R., 1959. — Field studies of Amphibians in Colombia, South Ame-

rica. Univ. Calif. Publ. Zool., 56 : 493-540.

STEJNEGER, L., 1910. - The Amphibian generic name Engystoma untenable. Proc. biol. Soc. Washing-

ton, 23 : 165-167.

TayLor, E. H., 1952. - The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull., 35 : 577-942.

2e 1962. - The Amphibian fauna of Thailand. Univ. Kansas Sci. Bull., 43 : 265-599.

TRUEB, L., 1973. - Bones, frogs, and evolution. Zn : VIAL (1973) : 65-132.

TscHuDi, J. J., 1838. — Classification der Batrachier, mit Berücksichtigung der fossilen Thiere dieser Ab-

theilung der Reptilien. Neuchâtel, Petitpierre : 1-102, pl. I-VI.

TYLER, M. J., 1979. - Herpetofaunal relationships of South America with Australia. Monogr. Mus. nat.

Hist. Univ. Kansas, 7 : 73-106.

Van Dix, D. E., 1966. - Systematic and field keys to the families, genera and described species of

Southern African Anuran tadpoles. Ann. Natal Mus., 18 : 231-286.

ViaL, J. L. (ed.), 1973. - Evolutionary biology of the Anurans. Columbia, Univ. Missouri Press : i-xii

+ 1-470.

WAGER, V. A., 1965. — The frogs of South Africa. Cape Town & Johannesburg, Purnell & Sons : i:

+ 1-242.

WaGLeR, J., 1830. — Natürliches System der Amphibien, mit vorangehender Classification des Säugethiere

and Vügel. München, Stuttgart & Tübingen, Cotta : i-vi + 1-354.

WaLker, C. F., 1973. - À new genus and species of microhylid frog from Ecuador. Occ. Pap. Mus.

nat. Hist. Univ. Kansas, 20 : 1-7.

Source : MNHN, Paris

AINTES

Journal International de Batrachologie

International Journal of Batrachology

édité par la Société Batrachologique de France

Rédacteurs : Alain DuBoIs et Jean-Jacques MORÈRE.

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

75005 Paris, France.

Comité de rédaction : Jean-Louis AMIET (Yaoundé), Stephan D. Busack (Urbana), Benedetto LANZA

(Firenze), Raymond F. LAURENT (Tucumän), Richard J. WassERsUG (Halifax).

Recommandations aux auteurs. — Alytes publie des articles originaux en français ou en anglais, consacrés

aux Amphibiens. Les manuscrits doivent être dactylographiés et accompagnés d'un résumé en

anglais (abstract). Les articles en anglais seront suivis d’un résumé assez complet en français (pour

ceux qui le souhaiteraient, les rédacteurs acceptent de revoir les résumés en français à partir d’un

texte en anglais). Tableaux et figures doivent comporter un titre. Les figures, exécutées à l'encre

noire, ne devront pas dépasser le format 16 X 24 cm. Indiquer leur numéro au crayon ; légendes

sur feuille séparée. Présenter les références bibliographiques conformément au dernier numéro

d’Alytes paru (les références de livres doivent comporter la pagination). Adresser les manuscrits

en trois exemplaires aux rédacteurs. L’acceptation d’un article pour publication est décidée par

les rédacteurs après lecture critique de celui-ci par deux lecteurs ou plus.

Instructions to authors. — Alytes publishes original papers in English or in French, dealing with Am-

phibians. Manuscripts should be typewritten, and preceded by an English abstract. Papers in

English should be followed by a detailed French summary (for those who may wish so, the ed-

itors accept to revise such French summaries on the basis of an English text). Tables and figures

should possess titles. Figures should be drawn in black ink and should not exceed 16 X 24 cm

in size. Their numbers should be written in pencil. Figure captions should be assembled on a

Separate sheet. Bibliographic references should be presented as in recent issues of Alytes (book

references should include the pagination). Send the manuscripts in triplicate to the editors (ad-

dress above). Acceptance for publication will be decided by the editors following review by two

referees or more.

Tirés à part. — 25 exemplaires gratuits par article. Au-delà, les tirés à part seront facturés par tranches

de 25 exemplaires.

Publié avec le concours du Muséum national d'Histoire naturelle.

Directeur de la Publication : Alain DUBoIs.

Numéro de Commission Paritaire : 64851. CE NP UUSS

T & JAN, 1998

Alpes, 1987, 6 (1-2) : 1-84.

SOMMAIRE

Alain DuBois

Müscellanea taxinomica batrachologica (IE)... 1

Alain DuBois

Miscellanea nomenclatorica batrachologica (XVI) ..................,,,.,,. 10

Pierre JOLY

Le régime alimentaire des Amphibiens : méthodes d'étude .................... 11

Miloÿ L. KALEZIÉ, Georg DZuKié, Jelka CRNOBRNJA & Nikola TVRTKOVIÉ

On the Triturus vulgaris schreiberi problem : electrophoretic data .......... 18

Manuel POLLS PELAZ

On the identity of the so-called “algae like cells” in tadpole cultures of Euro-

pean green frogs (Rana ridibunda) ..

Alain DUBoIs

Again on the nomenclature of frogs

Alain DUBOIS

Discoglossidae Günther, 1858 (Amphibia, Anura) : proposed conservation .…. 56

Alain DuBors

Strongylopus Tschudi, 1838 (Amphibia, Anura) : request for the designation

under the plenary powers of a twpe-species in agreement with current usage 69

Alain DUBOIs

Elachistocleis Parker, 1927 (Amphibia, Anura) : proposed conservation .….. 75

Imprimerie Fotek, St.-Niklaas, Belgique.

Dépôt légal: 4% trimestre 1987.

Source : MNHN, Paris: