H GMS mm

AILNTTES

Septembre 1986 ua ; Volume 5, fascicule 3

Source : MNHN, Paris

SOCIÉTÉ BATRACHOLOGIQUE DE FRANCE

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

SIÈGE SOCIAL

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

25 rue Cuvier, 75005 Paris, France.

CONSEIL D'ADMINISTRATION POUR 1986

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 et Luck MARTIN-BOUYER.

ADHÉSION

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

écrire au Secrétaire général. La cotisation inclut le service du Bulletin d’information Circalytes.

TARIFS 1986

LIEU DE RÉSIDENCE

France Europe Hors-Europe

Membres de la S.B.F. : É

Cotisation seule .

Cotisation + abonnement à Alyres

100 F 100 F 100F

160F 160 F 26$ US

Abonnement à Alyres pour les non-membres : 5

Individus 90F 1I0F 20$US

Intitutions étrangères . _ 220F 36$ US

Supplément pour expédition d’Abpres par avion (membres et non-membres) = = 8$ Us

Achats au numéro et rachats d’anciennes séries d’ Alyres :

écrire au Secrétaire général pour information.

MODALITÉS DE RÈGLEMENT

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

sorière, 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 Chèque Postal : “Société Batrachologique de France”, C.C.P.7976 90 K, Paris.

OUTSIDE EUROPE. — Payments should be made in US Dollars by checks (payable to “Société Batrachologique

de France”) sent to our Treasurer (address above), or by credit cards (contact the General Secretary for

instructions).

Source : MNHN, Paris.

ALVTES

Bulletin trimestriel Volume 5

Septembre 1986 Fascicule 3

Alytes, 1986, 5 (3) : 97-98. 97

Miscellanea nomenclatorica batrachologica (XII)

Alain DUBOIS

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

A replacement name is proposed for Glandula Tian & Hu, 1985, which is preoc-

cupied by Glandula Stimpson, 1852 (Tunicata).

TIAN & HU (1985) ont récemment suggéré de subdiviser le genre Bombina Oken,

1816 en deux sous-genres, le sous-genre nominatif et un nouveau sous-genre, pour lequel

ils ont proposé le nom Gandula. Malheureusement, ce nom ne peut être conservé pour

ce sous-genre, car le même nom avait été donné par STIMPSON (1852) à un genre de Tuni-

ciers. Nous proposons ci-dessous un nom de remplacement pour le sous-genre d’Amphi-

biens. Les deux sous-genres du genre Bombina peuvent désormais être caractérisés comme

suit :

Sous-genre BOMBINA Oken, 1816

Espèce-type, par désignation subséquente de la Commission Internationale de Nomen-

clature Zoologique (ANONYME, 1957): Rana bombina Linné, 1761.

Diagnose : voir TIAN & HU (1985).

Espèces incluses : Bombina (Bombina) bombina (Linné, 1761) ; Bombina (Bombina) orien-

talis (Boulenger, 1890) ; Bombina (Bombina) variegata (Linné, 1758).

Sous-genre GROBINA nom. nov.

Nomen novum pro Glandula Tian & Hu, 1985 (nec Glandula Stimpson, 1852).

Espèce-type, par désignation originale sous Glandula Tian & Hu, 1985 : Bombinator maxi-

D LE

mus Boulenger, 1905. x] Bibliothèque Centrale Mus

LU

lsource : MNHN, Paris

98 ALYTES 5 (3)

Diagnose : voir TIAN & HU (1985).

Espèces incluses : Bombina (Grobina) fortinuptialis Tian & Wu, 1981 ; Bombina (Gro-

bina) maxima (Boulenger, 1905) ; Bombina (Grobina) microdeladigitora Liu, Hu & Yang,

1960.

Etymologie du nom subgénérique : du nom générique Bombina et de l’adjectif latin gros-

sus (gros, épais).

RÉFÉRENCES BIBLIOGRAPHIQUES

ANONYME, 1957. — Opinion 453. Validation under the plenary powers of the generic name Bom-

bina Oken, 1816 (class Amphibia, order Anura). Opin. Declar. intern. Comm. zool. Nom.,

15 : 347-356.

STIMPSON, Mr, 1852. — [Several new Ascidians from the coast of the United States]. Proc. Bos-

ton Soc. nat. Hist., 4 : 228-232.

Tian, W. & HU, Q., 1985. — Taxonomical studies on the primitive Anurans of the Hengduan

Mountains, with descriptions of a new subfamily and subdivision of Bombina. Acta herpet.

sin., 4 : 219-224.

Source : MNHN, Paris

Alytes, 1986, 5 (3) : 99-149.

Living amphibians of the world :

a first step towards a comprehensive checkl

©

Alain DUBOIS

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

CONTENTS

Abstract 99

Introduction ... 100

General comments 102

Choice of the information given . 102

Conditions of preparation .. 110

Choice of the classification .. 113

Specific comments ...... 115

Valid names of the taxa, with their correct authors, spellings, dates, references

‘and: pages of publication, 40e meer ele ln einen autel L ne 117

In the class-group .. 117

In the family-group . 117

In the genus-group . 126

In the species-group 129

Type species of genera and subgenera . 132

Type specimens of species … 137

Other types of information . 139

Final miscellaneous comments . 142

Conclusion . 143

Résumé ..... z . . 144

Literature cited .. perse ... 145

The checklist Amphibian species of the world, result of a collective work

by 59 authors, gives the names of 4015 living species of amphibians, along with

various types of information concerning them. Although such a list is potentially

of great interest, both to systematists and to all other biologists working on amphi-

bians, this first edition raises several important problems. The choice of the infor-

mation given in the book seems open to criticism : in particular, the absence

of synonyms and of subspecies names greatly reduces the usefulness of such a

list. Besides, even for the data which appear in the book, the rate of errors and

of omissions, such as it may be estimated for a variety of types of information,

is much too high for this list to be considered a valuable standard in amphibian

taxinomy, nomenclature and bibliography. This is clearly due to the fact that

this first edition was prepared and published much too quickly. It therefore seems

advisable to await at least the second, revised, edition of this book.

! or

99

ist

Source : MNHN, Paris

100 ALYTES 5 (3)

FROST, Darrel R. (ed.). — Amphibian species of the world. À taxonomic and

geographical reference. First edition. Lawrence, Kansas, Allen Press and the Associa-

tion of Systematics Collections, 23 August 1985 : [i-iv] + i-v + 1-732. Price (cloth-bound) :

US $ 85. — Cited below as ASW.

Note. — This paper was written at the request of the editors of Copeia for the “Reviews and

comments” section of this journal, but was finally considered too long and detailed for this journal.

It is therefore published integrally here, while a brief summary of it will appear in Copeia (1987).

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

code numbers, which are made explicit in the bibliography. D-41 refers to the present paper.

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

INTRODUCTION

The checklist Amphibian Species of the World recently published by the Association

of Systematics Collection and Allen Press is an impressive hard-bound volume of more

than 700 pages, the aim of which is “to serve as a standard reference to amphibian nomen-

clature” (ASW : [ii]. Although this does not appear in its title, the list concerns only

the living species of this class. The list was edited under the direction of Darrel R. FROST,

under the auspices of the World Congress of Herpetology and its Checklist Committee,

composed of William E. DUELLMAN (Chairman), Robert C. DREWES, Carl GANS, Alice

G. C. GRANDISON and Marinus S. HOOGMOED. It was compiled by 59 batrachologists

working in 20 countries (not 21 as stated in ASW : iii), who clearly devoted a lot of their

time to the patient collection of the information which is finally presented.

The book poses a genuine bibliographic problem. How should it be cited ? The

simplest solution would be to quote it in all cases as “FROST, 19857, or “FROST et al.,

1985”. But in many cases this would not do justice to the fact that most sections of the

book have been entrusted to specific ‘“‘contributors”, and modified by specific “review-

ers” (some of whom made specific comments which are signed with their initials). À com-

ment like the one on p. 14 signed “JDL” should certainly be quoted as “LYNCH in

FROST, 1985”. A section like the genus Arshroleptis (pp. 14-16), contributed by Raymond

LAURENT alone, should certainly be quoted as ‘LAURENT in FROST, 1985”. Now, how

should be cited an information which appears in a section for which there was no specific

contributor, and which was “completed in cooperation between the editor and the review-

ers” (p. 2} The section Brachycephalidae for example (p. 24), which only contains two

monotypic genera, has no specific contributor, but is credited with 10 reviewers : should

this section be cited as ““FROST, BOKERMANN, CANNATELLA, CARAMASCHI, CROMBIE,

DUELLMAN, GUDYNAS, HOOGMOED, MCDIARMID, SCOTT & VANZOLINI in FROST,

1985” (!), as “FROST et al. in FROST, 1985”, or as “FROST, 1985°”? The same problem

arises for the sections which are credited to several contributors, even when the checklist

mentions the geographic region of their respective contributions. Thus, the text on the

genus Rana is credited to 6 contributors, but the geographic regions covered by three

of them partially overlap : Sushil DUTTA (Tropical Asia), Shuqin HU (China) and Masa-

fumi MATSUI (East Asia). How should be quoted any given information dea-

Source : MNHN, Paris

DUBOIS 101

ling for example with a species of tropical China ? Furthermore, the contributors mentio-

ned at a given taxinomic' level (e.g. the subfamily Raninae : 5 names) may not include

those mentioned at a lower level for included taxa (e.g. the genera Batrachylodes, Cerato-

batrachus, Discodeles, Palmatorappia, Platymantis, Rana, Strongylopus and Tomopterna,

which were partially or totally contributed by authors not included among the first 5 ones).

For this reason, correct citation of the checklist will be a difficult task in many cases where

the attribution of a given information to a given author is ambiguous. To simplify, in

the following discussion the checklist will be quoted in all cases as ASW(Amphibian Spe-

cies of the World), but this should not obscure the fact that the various parts of it were

contributed by different authors.

In this list, “complete to 1 June 1984” (ASW : 2), I counted 4015 nominal species

considered valid (not 4014, as stated on p. 1), arranged in 395 genera and 37 families.

The last checklist of the amphibians of the world, supposedly complete “to January 1,

1970”, that of GORHAM (1974), contained 3343 nominal species (D-4), arranged in 347

genera and 28 families. Even though both checklists are incomplete and contain mistak-

es, these figures show that an important increase (by 20.1 %) in the number of known

or recognized species of amphibians has taken place in the last 15 years period, thus indi-

cating that the field of “microtaxinomy” (MAYR, 1982) in still lively in this group of

vertebrates, and will remain so still long (see also D-4). The increase is also high for the

number of recognized genera and families (respectively 13.8 % and 24.3 %), which also

points to the fact that the ‘“‘macrotaxinomy” of this group is not yet stabilized. In fact,

even if many genera are named and formally described, their phylogenetic relationships

are often poorly known and still not known at all for some of them. In the future it will

be as necessary to work on the higher classification of amphibians as to continue descri-

bing new species and revising difficult and poorly understood groups.

Every batrachologist can only be enthusiastic with the idea of such a book. À com-

prehensive list of names of living Amphibia in the world would be most useful not only

to taxinomists but also to all batrachologists and to all biologists of all disciplines (ecol-

ogy, biogeography, nature conservancy, evolution, genetics, biochemistry, development-

al biology, physiology, etc.) who may have, at a given stage of their research, to know

the valid name of any amphibian species, its phylogenetic relationships, its place within

the current classification, its geographical range and its possible status as an officially

endangered species. Like many colleagues, I had applauded this project and thought it

would happily replace the Das Tierreich volumes, which cover only certain groups and

are now outdated.

My hope was only very partially fulfilled. Certainly, the book exists, which is bet-

ter than nothing ; it will certainly also be useful, at least to those who will use it knowing

its strengths and weaknesses. However, this book is not what one could expect. What

constitutes, in my opinion, its weaknesses, is detailed below. Before doing so, however,

I wish to make clear that I had been contacted by William E. DUELLMAN and Darrel

R. FROST, who had proposed me to participate in the elaboration of this book, but that

I had finally resigned from this work. In a letter dated 25 January 1983, they proposed

1. I use the correct spelling “taxinomy” instead of “taxonomy”, following PASTEUR (1976) and FISCHER & REY

(1983).

Source : MNHN, Paris

102 ALYTES 5 (3)

me to act as contributor for the sections Megophryinae and European and Asian Raninae

of the book. In a letter dated 8 March 1983, I answered that I would accept this work

under certain conditions. FROST sent me the first computer printout of most of the Rani-

dae and of the Megophryinae on 1st April 1983 and asked me for my corrected manu-

script in June 1983. After having started this work, I realized that it could not be ready

for this date and answered I was giving up the project because the time provided was

too short. Beside that, I stated that I was ready to care for the suprageneric nomenclature

part of the whole project. To this last proposal, FROST answered on 14 June 1983 that

he would send me a printout of the citations for higher taxinomic taxa, but I never receiv-

ed it.

The above comments are given in order to show that the following review is not

the reaction of a frustrated batrochologist angry not to have been asked to participate

in this collective work : actually, I had been asked to do it, but refused, for the very rea-

sons which lead me now to criticize the work, i.e., that it suffers mainly from having

been produced in haste. The following comments shall be divided into two parts : gene-

ral comments dealing with the conception and realization of the book ; analytical com-

ments dealing with precise examples scattered throughout the book and dealing with various

types of information.

GENERAL COMMENTS

CHOICE OF THE INFORMATION GIVEN

The quantity of information which can be included or indexed in a checklist is vir-

tually unlimited. It would certainly be nice to have a book giving the names of all taxa

with information on their history, nomenclature, classification, phylogeny, distribution

and all other biological attributes. However, generation of such a list would take a very

long time, and its publication would occupy several volumes the size of ASW. It is there-

fore necessary, to save time and space, to make a choice among the possible information

which such a checklist can contain. The choice of this information should be based on

the public to whom the book is addressed, and in this respect it is not quite clear to me

for whom this book is specially intended. From the various forewords, il would seem that

this list is not meant to be useful only to professional batrachologists, but also to all other

biologists. But one may wonder if the choice made concerning the types of information

given in the checklist is the best one in this respect. These types of information are of

12 sorts.

(1) Valid (or supposed valid) names of all taxa of Amphibia of the following ranks :

class, order, family, subfamily, tribe, genus, subgenus and species. In addition, a few names

of taxa of the ranks of suborder, superfamily and subspecies are given incidentally in the

book, but there is no consistency.

(2) Authors, dates and bibliographic references of the works where the names of

all taxa of the ranks class, order, family, subfamily, genus and species were created (or

supposedly created). In addition, a few references are also given for some taxa at the sub-

generic and subspecific levels, but again without consistency. No reference is given for

taxa of the ranks suborder, superfamily and tribe.

Source : MNHN, Paris

DuBolIs 103

(3) “Original names” of species (when they are different from the current names).

Under this title are given in fact two rather different types of information. In some cases

the “original name” is the original combination, when a species was first described in

a different genus from that where it is currently placed ; the author of the current combi-

nation is generally not mentioned. In some other cases the “original name” is only the

original spelling, when this spelling was an “incorrect original spelling” in the sense of

the Code (ANONYMOUS, 1985 a), and has to be replaced by a “justified emendation” ;

the first author to have used the justified emendation is generally not given. In a few

cases, finally, the “original name” differs from the name considered valid in the checklist

both in combination and spelling.

(4) Original spellings of names of families and subfamilies. This information appears

incidentally in the “comments” following a few names, but is not given in all cases.

(5) Authors, dates and references of the works where first appeared (or supposedly

appeared) the justified emendations of familial or subfamilial names which had been created

under incorrect original spellings. As in the preceding section, this information is given

incidentally in some cases but not in all.

(6) Type species of genera. This information is given for all genera of Amphibia

recognized as valid in the book, but not for the few generic names currently considered

synonyms which are mentioned. It is not given, except for one exception (Echinotriton),

for the subgenera recognized as valid in the checklist.

(7) Kind of type fixation of genera. This information is given in only a part of the

cases (see below for more detail).

(8) Type specimens (holotypes, syntypes, lectotypes or neotypes) of nominal spe-

cies : location (identification of the collections where they are kept, in general Museums)

and collection numbers. This information is given only for a part of the species, and is

sometimes complete, sometimes incomplete (e.g. the location may be known but not the

collection number, or only a part of the syntypes of a species may have been traced ; see

below for more detail).

(9) Type locality. This information is given for all species, but usually not for sub-

species and synonyms.

(10) Distribution. This information is given for all taxa of the ranks class, order,

family, subfamily, genus and species, and exceptionally for a few taxa of other rank.

(11) “Comments”. This section gives various types of information, including : di-

scussions on phylogeny and classification ; nomenclatural discussions ; in a few cases,

mention of synonyms (for some taxa of the ranks genus, subgenus and species) ; in a few

cases, mention of subspecies (with or without complete bibliographic reference to the works

where their names were created, with or without mention of their type specimens, with

or without their type localities and distributions) ; and references to relevant works of

all kinds dealing with these taxa.

(12) Protected status. This information is given for the few species of Amphibia

governed by the USA Endangered Species Act (USA ESA) and listed in the Appendices

of the Convention on International trade in Endangered Species of Wild Flora and Fauna

(CITES).

Source : MNHN, Paris

104 ALYTES 5 (3)

Of the above types of information, most (i.e. types 1 to 9) give nomenclatural data

on names, while only 3 (i.e. types 10 to 12) give biological information on taxa.

Among all these types of information, which ones are likely to be useful to the

various kinds of biologists who are not primarily concerned with systematics and nomen-

clature and to whom the book is supposedly intended ? I would suggest such biologists

will mainly seek in such a book : (a) information about the relationships of the taxa and

the supraspecific classification ; (b) exact names of the taxa with dates and authors (but

not necessarily details such as the original name or the detailed citation of the original

reference) ; (c) complete synonymies of all taxa names ; (d) type species of genera ; (e)

information on the geographical distribution of taxa, including data on geographical varia-

tion within species and the existence, name and distribution of subspecies ; (f) a list of

key references allowing to trace all relevant works dealing with a given taxon (taxinomic

revisions, faunistic works, other biological works of importance).

The importance of some of this information, maybe not evident at first sight, should

be underlined. The need for complete synonymies, for example, may be underestimated.

They are necessary in order to allow the users of the list to trace the current names of

species which were given other names in the past. In Amphibia, such cases are nume-

rous. Many older experimental works deal for example with species which were then known

as Bufo vulgaris, Rana fusca, Triton palmatus, etc. AI the information that may have been

gathered then on these animals (through field work, observations or experimental research),

and which had then been attached to these older names, may be lost, or improperly allo-

cated, if nonsystematists cannot easily identify the biological species to which both older

and newer names apply. This in my opinion is one of the major objectives one should

keep in mind when preparing such a checklist.

Besides, it is important to clearly make the distinction between two kinds of names :

those which are nomenclaturally available, and those which are considered valid at a given

moment of the nomenclatural history of a biological group. It should be stressed that such

an history is extremely dynamic, because a name that has been once considered invalid

as being a junior synonym of another name may be resurrected as a valid name later if

it is shown that both apply to different biological species : as a matter of fact this happen-

ed already for many names of Amphibia ; similarly, in the species-group some second-

ary homonyms may in some cases be resurrected when such names are transferred from

one genus to another (see Art. 59 ofthe Code). The fact of giving in a checklist only those

names which are considered valid at the time of completion of this list supports a static

conception of nomenclatural history which is not born out by the facts. The absence of

synonyms and homonyms greatly reduces the usefulness of the book as a “standard refe-

rence to amphibian nomenclature”. It has also another disadvantage : since it is not com-

plete, this checklist (or its index) cannot be used to know whether a given specific name

has already been given to a species of a given genus, an information which is useful in

order to avoid creating homonyms. In its present state, the checklist and its index will

only allow to find such names if these are currently considered valid specific names, but

not, except for some exceptions, if these names are currently considered synonyms, nomina

dubia, or valid subspecific names (see below).

Knowledge of the type species of genera is also useful : for example, if an experi-

mental biologist wants to compare different genera for given characteristics, he may wish

Source : MNHN, Paris

Dugois 105

to use only one species by genus, and in such a case it would seem best to use, if possible,

the type species of all compared genera : even if the limits of genera are liable to change

with future taxinomic works, the type species of nominal genera won’t change and will

remain stable reference marks for future comparative works.

Information on geographic variation and on subspecies is also important, for all

workers interested in evolution at lower levels. Precise data on the geographic distribu-

tion of recognized subspecies of polytypic species, or bibliographic reference to such data,

should appear in any such checklist. It should be stressed furthermore that the taxinomic

research of the last period has shown that many taxa considered in the past as subspecies

were either strict synonyms or, on the contrary, good species ; symmetrically, cases of

downgrading of specific names to subspecific level are also known, although less abun-

dant (see D-4). À good knowledge of the present state of this question in any given group

is most important in many types of works, and should be given without restriction in

a list like ASW.

If we now compare these needs with ASW, we can make the following statements :

(a) the information about relationships and classification is generally satisfying, except

at higher levels (suborders and subfamilies of Anura) and at lower levels (subgenera and

species groups), and although the classification in its whole is not very homogeneous (see

below) — but this is quite unavoidable in the present state of the taxinomy of the Amphib-

ia ; (b) the list gives the names, authors and dates of the taxa, but furthermore is made

cumbersome by the mention of the original names and the heavy mode of notation of

the original references in the text ; (c) synonymies are rarely given (sometimes, some

synonyms are mentioned in the “comments”, but there is no real rule in this respect :

a species name that has recently been synonymized or resurrected may often be mention-

ed as such, but most older synonyms are not ; generic names are almost never given) ;

(d) type species of genera are given, although in many cases without details on their mode

of fixation ; (e) some information about the geographical distribution is given, but the

subspecies are usually not listed (there is also no real rule about it : when a subspecies

has recently been described, or elevated to species rank, or the contrary, it may be men-

tioned ; but most subspecies are not) ; (f) from one group to another, the completeness

ofthe comments and of the relevant bibliography given varies exceedingly : in a few cases

it is excellent (e.g. genus Xenopus), in other ones it is really bad (e.g. Rana esculenta synklept-

on) ; in general, it is very brief, rather insufficient.

Besides, it should be stressed that the checklist devoids great deal of space to a type

of information which cannot be considered as important for nonsystematists as some of

the lacking information mentioned above : the information on type specimens and type

localities of species. My opinion is that this information will be useful mostly, if not only,

to taxinomists, and should not have been given priority over information on subspecies

and synonyms in a checklist intended to be useful to biologists of various kinds.

What is the use of type specimens ? In the older days of pre-evolutionary, typologi-

cal thinking (see e.g. MAYR, 1969, 1982), types were believed to be “‘typical”? specimens,

exhibiting “typical” characters of species, and specimens showing different characters

were attributed to different species. Al specimens of a given species were believed to

be identical or almost identical to the type, and were often treated, in older Museums,

as “‘duplicates”, which were often disposed of (exchanged, given away or destroyed). The

Source : MNHN, Paris

106 ALYTES 5 (3)

situation changed with the emergence and development of the evolutionary and popula-

tional thinking : the “type” was reduced to being only one of the specimens of a variable

species, and could well not exhibit some of the characters considered diagnostic of a spe-

cies (see e.g. D-4). The rôle of type specimens in systematics has considerably changed,

to a point that has perhaps been under-estimated by some taxinomists. Type specimens

are not used any more for supposedly exhibiting diagnostic characters (although they may

often also do so), but primarily for a nomenclatural purpose : a type specimen is an objec-

tive reference to a natural population of animals, and a way to objectively attach a name

to this population. In recent revisionary works for example, taxinomists study species

limits, diagnoses, distributions, etc., on the basis of as many specimens of various origins

as possible. During this work, they do not (or should not) give a particular importance

or significance to type specimens, which are only specimens among others. Only once

the revision is finished, the biological species and their intra- and interpopulational varia-

bility understood, should the type specimens of the nominal species included in the revi-

sion be examined, in order to know which names should be applied to the various biolo-

gical species defined by the revision. The type specimens have now a purely nomenclatu-

ral rôle in systematics, for defining names, but they do not any more have the special

rôle which they used to have in the past for defining raxa.

Two conclusions may be derived from this assertion : (1) except in the case of the

now relatively rare species of Amphibia which are still known only by their types, type

specimens are not any more important than any other Museum specimens referred to

a given nominal species after a recent revision, or than any other freshly collected speci-

men determined on the basis of such a recent work ; (2) on the other hand type specimens

still play an important rôle in systematics in that they are the only “bridge” linking the

real world (biological species) with the world of language (nomenclature). Type speci-

mens remain thus most useful to systematists, but to systematists alone, in allowing them

to objectively tie scientific names to biological species. But in this respect all type speci-

mens of existing nominal taxa play the same rôle : type specimens of nominal species

the names of which are homonyms or are currently considered synonyms or valid names

of subspecies are as important for systematists as type specimens of nominal species the

names of which are currently considered valid specific names. Therefore, if a list like

ASW is to give the references of type specimens, it should give all of them, irrespective

of the fact that the names to which they are attached are currently considered valid speci-

fic names or not. Doing differently reminds me of the true story of a Museum worker

who told me that, when a specific names is placed in synonymy, its type specimens lose

their status of type and should not be listed in a catalogue of types ! Unfortunately, by

listing only the type specimens of the species currently considered valid, the authors of

ASW seem to adopt the same philosophy. To list type specimens of valid species but

not of homonyms, synonyms and subspecies seems to imply that these types are more

important than other specimens to know and characterize biologically a species, which

is completely wrong but shows that typological thinking is still alive among contempora-

neous taxinomists. I therefore consider the choice of mentioning type specimens rather

than listing synonyms and subspecies in such a book as a scientific error due to a survival

of typological thinking.

Source : MNHN, Paris

DuUBoIs 107

Mention of type localities of species meets with the same criticism. Since the checklist

gives the distributions of species, mention of type localities has only a nomenclatural in-

terest, and particularly in the case of species which can be subdivided into several subs-

pecies : but here again, to make this information useful, il would be necessary to give

also the distributions and type localities of all subspecies.

Thus the information concerning type specimens and type localities is of little in-

terest to nonsystematists. It is true however that this information will be of interest to

taxinomists, although its incompleteness will greatly limit this interest. It should be stressed

however that systematists have other ways to seek this information : once one knows the

name of the species, its author and date, it is easy to go back to the original description

and see whether the type locality and the type specimens are given there ; if not, in many

cases it will be possible to find this information by writing to the curator of the institu-

tion where the author was working. In the remaining cases, it is true that a long inquiry

may be necessary to find out the present location of the types, and if this information

was available for all taxa in a checklist like ASW, this would save much time to Museum

systematists all over the world. However, careful examination of ASW/shows that it usually

gives this information only in two cases : (1) when it has already been searched for by

the author of a revision, of a previous checklist or of a type catalogue : this is for example

the case of all the types traced earlier by DUELLMAN (1977) in the families Hylidae, Cen-

trolenidae and Pseudidae ; in this case, direct use of DUELLMAN's checklist would pro-

vide the same information as ASW — and even more, because it includes the types of

synonyms and subspecies ; (2) for species described by well-known authors who used to

deposit their types in a given institution ; in this case, ASW often gives the location of

the types without giving their numbers : the same information could have been found

by any taxinomist without the help of the checklist.

A checklist should not contain, as far as possible, original data. These should be

published independently in papers cited in the checklist. This means that a checklist is

unavoidably rather heterogeneous, according to whether any given group has or not been

the matter of a recent revisionary work. In the case of the type specimen section, a pecu-

liar problem arises. As far as in most ancient publications the types, and their Museums

of deposition and numbers, are not mentioned (if it was not the case, it would be comple-

tely irrelevant to search for them and list them in a checklist), searching this information

is a genuine scientific work, sometimes long and unrewarding. To include such an infor-

mation in a checklist is contrary to the principle just set forth. When this research has

been done on the occasion of another revision or list, like for example DUELLMAN's (1977)

checklist mentioned above, reference to this list should be enough. In the other cases,

if genuine research is involved, who will be the author(s) of the original information first

published in the checklist ? The editor ? The contributor(s) ? The reviewer(s) ? In most

cases, this information appears anonymously in ASW. This raises additional problems.

For example, when type specimens are mentioned, for the first time in ASW/, as being

“lost”? or “destroyed”, this list should give us the evidence on which such an information

is based : either a bibliographical reference, or the author of this statement (letter from

a curator, personal observation of a contributor, etc.). In the absence of such statements,

it is difficult to know the seriousness and validity of such types of information. Actually,

in several cases I found the mentions “not traced”’ or even “lost” for type specimens

Source : MNHN, Paris

108 ALYTES 5 (3)

which are still perfectly extant, and easy to trace, in major Museums (details on these

cases will be published elsewhere).

The preparation of this book could have been a good occasion to make an interna-

tional inquiry among all natural history Museums (which are listed in Appendix II of

ASW, pp. 665-669). This could have been done for example in taking advantage of the

fact that all the data on which the list is based are stored on computer and in using the

latter to prepare a list of species names for which every Museum collection is supposed

to have the type specimens and to ask these Museum confirmation of the existence of

these specimens at this date and their proper collection numbers. However, no such inquiry

was done by the editors of ASW— or, at least, the Paris Museum, the third largest collec-

tion in the world for the types of amphibians (see Table XI), was never contacted in this

respect.

There is at least one kind of type specimens which could have been quite useful

for taxinomists to list in a book like ASW : the types which were fixed by an action poste-

rior to the original publication where the name had been created, either by a selection

among various syntypes and choice of a lectotype, or by designation of a neotype when

the original type(s) is (are) no longer extant. It is true that these types are much more

difficult to find than original holotypes or syntypes, because any author acting after the

original description may potentially designate such types, and it would be useful to have

a reference to such actions in the “comments”. In this respect, it is disappointing to find

that the information given in ASW on such types is incomplete or false in several cases

(see below), thus reducing again the interest of giving the information on type specimens

in the book.

À good decision of the authors of the checklist has been to list only holotypes, but

not paratypes, which have no particular meaning or importance in nomenclature. Then,

why have some paralectotypes been listed in the book ? There is no rule about it : in

some cases, only the lectotype is listed, which is correct ; in other ones, all syntypes are

listed, then the lectotype designation is added, which is much too heavy : after a lectotype

has been designated, all other syntypes lose their status of “types”, and become “paralec-

totypes”, which have as little nomenclature meaning or importance as paratypes.

Another type of information which could have been useful to amphibian taxino-

mists is the one bearing on the nomenclatural decisions taken by the ICZN (International

Commission on Zoological Nomenclature) with respect to family-group, genus-group and

species-group names of Amphibia. Some of these decisions are mentioned incidentally

in ASW, but most of them are not. In particular it would have been useful (and not espe-

cially time and space consuming) to list in this book all the amphibian names which have

been placed on Official Lists of Conserved Names in Zoology and on Official Indexes

of Rejected and Invalid Names in Zoology, with their numbers on these Lists and Indexes.

This has already been done for all familly-group names and some genus-group and species-

group names by DUBOIS (D-26), and should in my opinion be added in future editions

of ASW.

À strange situation is met with as concerns the status of nomina dubia (names which

are nomenclaturally valid but cannot be associated, at the moment, with given biological

taxa). Such names can have some importance for systematists, as potential senior or junior

homonyms, and as potential senior or junior synonyms (if their status happens to be elu-

Source : MNHN, Paris

DuBois 109

cidated). However, they are less important than names the status of which is clear (either

as valid specific or subspecific names or as synonyms), and it does not seem logical to

include them in a checklist which ignores subspecies and synonyms. While most of these

names have in fact been deleted from the checklist, it is thus surprising to note that some

of them have been listed, for example in the genera Hyla, Hyperolius or Leptodactylus :

in these few genera we have therefore a strange situation where some names are listed

but almost nothing is known about them, while other well-known names are not even

mentioned simply because they apply to subspecies or are currently considered junior

synonyms of other names. It would have seemed justified to homogeneize the checklist

by suppressing all nomina dubia — or including all nomenclaturally available names of

Amphibia without exception.

In the Preface of ASW (p. ïüi), the World Congress of Herpetology Amphibian Check-

list Committee writes : “Originally it was hoped that the list would contain synonyms

and subspecies. However, it soon became obvious that the inclusion of synonyms (and

their citations and type localities) would not only increase the size of the volume beyond

the reasonable limitation that had been set for it, but would require more effort by the

contributors and reviewers.” It is thus clear that this information was not included in

ASW to save space and time. However, one may wonder if space and time could not

have been spared more efficiently if a different choice had been made as to the type of

information to be included in the checklist and its mode of presentation.

A first remark can be made about the way bibliographic citations are printed. Refer-

ences are given in the text itself, in an abbreviated form including only the abbreviated

title of the periodical or book and the relevant pages, but not the complete title of the

work (as in traditional synonymies from the end of the XIX century and the first part

of ours), and are not given in full in an alphabetical bibliography at the end of the book

(as in most recent works). A selected bibliography appears at the end of the volume, giving

the titles of some of the books cited in the checklist (but not all) and the full titles of

the periodicals cited. This presentation has several disadvantages. It is for example impos-

sible to find out immediately, by looking at the bibliography, if a given publication has

been known to the authors of the checklist and used by them, or not. It is also impossible

to find a list of references of a given author cited in the book. This system of presentation

is probably intended to save some space (and also perhaps to remain in the tradition of

presentation of synonymies in older works). It may save some space at the end of the

volume, but it certainly expands the text itself, and makes its reading much more diffi-

cult and unpleasant. Furthermore, on the whole it may even not really save space, when

one considers that many papers (catalogues, checklists, faunas, revisions, etc.), the abbre-

viated reference of which may be almost one line long, are cited several times in the book,

and some of them many times : if these papers were quoted in full only once in the bibli-

ography, and referred to only by the author’s name and a date (with or witout an addition-

al letter) in the text, this would save much space. It is impossible to know a priori which

system would occupy more space, but the modern system of short references in the text

and complete references in bibliography at the end is much more efficient to use and

provides more information (complete titles of publications, number of pages, etc.), and

for this reason should have been preferred in this book. But there are several other ways

to save space in such a checklist.

Source : MNHN, Paris

110 ALYTES 5 (3)

In some checklists, species are only indicated by their names, with their authors

and dates, and, sometimes, the original generic names under which they had been describ-

ed if they have changed later (this is the case for example in GORHAM’s 1974 checklist).

This présentation has the advantage of saving space (the space for the original name, when

relevant, and for the original reference, be it in the text or in a final bibliography) and

time (for the collection of the above information). Besides, it has no real disadvantage :

with the help of the Zoological Record, it only takes a few minutes to find out the original

reference, starting with the name of the author, the date and the name of the taxon ; for

references prior to 1864, date when the Zoological Record was founded, this work may

be a little longer, but with the help of the previous checklists (British Museum’s Catalo-

gues, Das Tierreich, etc.), it usually remains rapid.

What space would save the suppression of this information ? I had the curiosity

(and patience) to count the number of lines which would have been saved if only author

and date had been given for all scientific names, without the original reference, and also

if the sections “original name”, ‘“‘type(s)” and “type locality” had been deleted : I found

respectively 606, 1626, 4448 and 5823 lines, i.e. a total of 12503 lines. Altogether, there-

fore, “suppression” of this information would allow to save much more space than would

be needed to allow adding synonyms and subspecies without increasing the size of the

book, provided each of these names is given only with its author and date, without refe-

rence, nor original name in full (a different generic attribution could be mentioned on

the same line, like in GORHAM’s 1974 checklist), thus occupying only one line each. As

a matter of fact, in GORHAM's (1974) book, DUBOIS (D-4) counted 1009 subspecies (includ-

ing 323 nominative ones and 686 others) and 2252 names considered invalid (synonyms,

homonyms, replacement names), which would occupy 2938 lines altogether. From 1970

to 1984, the number of subspecies and synonyms has probably not increased very much,

because if it is true that new synonymies have been discovered and new subspecies de-

scribed (or downgraded from species rank), it is also true that in the same period a num-

ber of names were resurrected from synonymy or elevated from subspecific to specific

rank. However, even if we assume, probably incorrectly, that the rate of increase of these

two categories of names has been the same during this period as the rate of increase of

specific names considered valid, i.e. 20.1 %, the total estimate would be 3529 names,

which would still occupy less space than the information on type specimens alone (4448

lines) or on type localities alone (5823 lines).

It is thus clear that with a different editorial policy as to the type of information

to be given in the book, inclusion of data on synonyms, homonyms and subspecies could

have been possible without increasing the size of the book. The problem “space” could

then have been solved, but this would not have suppressed the problem “time” and

“effort”, and I think that here lies the true reason for not including synonyms and sub-

species in ASW : for some reason, the editors wanted to publish the list quickly, and could

not afford for the large increase of working time which would have been necessary to

add this information. We are therefore now led to the study of the conditions of prepara-

tion of the checklist, especially as concerns the quickness of preparation.

CONDITIONS OF PREPARATION

As explained by FROST in the Introduction of ASW (p. 2), the procedure of compi-

lation of the book was an “iterative” one : a ‘‘skeleton checklist” was compiled by FROST,

Source : MNHN, Paris

DuBoIs 111

and entered on computer files ; its various sections were sent to various contributors for

completion, and returned to the editor for addition to the computerized data base ; after

correction, they were sent back again to the contributors, and the completed sections were

also sent to one or more reviewers for augmentation and correction ; the final copy was

read and approved by the contributors and the checklist committee ; final standardiza-

tion work was done by the editor. This work was carried over from 1982 to mid-1984.

The data base for the checklist is being maintained on computer files, which will be updated

“so that a new up-to-date version of the list can be produced at any time in the future”

(ASW : ii).

While the process of compilation described above may sound very nice, being a

collective one in which any one’s work was supposedly commented upon and corrected

by others, an important point must be stressed here : the production time for the work

was very short. Having myself worked quite a bit on similar projects, I know by expe-

rience that they cannot be seriously completed within a prefixed amount of time, espe-

cially when this time is short, and even when one devotes an important part of one’s time

to this work. Serious compilation of checklists of this kind really demands an enormous

amount of time, which is clearly underestimated by the “users” of these lists. It requires

in particular considerable bibliographic research, which is extremely time-consuming as

all original works must be traced and directly consulted ; the experience proves that, when

one avoids this work and when one uses second-hand references and information, one

often repeats the mistakes of previous authors (see below for more detail). Furthermore,

it is clear that most of the contributors and reviewers agreed to devote a lot of their time

to the project to build up a list which seemed to them the best one, but that it was not

their primary job at this time ; only the editor was “hired” by the ASC and paid for the

completion of the book. As many contributors and reviewers were unable, for reasons

of time and sometimes of geographic location, to have access to all the relevant original

literature, a lot of verification work rested on the shoulders of the editor at the end of

the work. For a really serious work, he should then have checked himself in the original

publications all the information given by the contributors and reviewers. That he did

not do so will be made clear in the detailed analysis that will be presented below. The

main reason for that is clearly the lack of time, due to the stringent time constraints which

were apparently imposed by the publishers.

It is true that we are now in a period where rapidity of work and publication is

considered one of the qualities a scientific researcher must have. The reasons for that

are not scientific at all, they are rather sociological, or more precisely economical. It is

a rule that many scientific works are now prepared and published very rapidly, for rea-

sons of financing, contracts, careers, etc. Careful study often reveals that these works were

prepared and completed 100 quickly. I strongly urge my colleagues to consider that, con-

trary to the current dominant ideas in this field, rapidity of execution is nor a criterion

of good quality of a scientific work. The relevant criterion should be seriousness, which

should assure the work to have a long life, and to be still useful to workers in the distant

future.

The ASW checklist falls fully into this trap. Furthermore, there seems to have been

a certain fascination of the publishers with the computer. It is true that, as mentioned

in the Foreword of ASW, the computer allows a considerable saving of time for the treat-

ment of data. But the computer only gives back to you what you have put in it. If you

Source : MNHN, Paris

112 ALYTES 5 (3)

introduce false or incomplete data, they will remain so even after the most sophisticated

treatment by the computer.

It is also perfectly true that any checklist is provisional and subject to revision in

the future. But it may be so for three distinct reasons : (1) subsequent progress of science,

which leads to a modification of what was known when the checklist was compiled ; (2)

the inevitable errors in works of this magnitude ; (3) incomplete, too rapid work during

the preparation of the checklist, which is a bad or incomplete reflection of the state of

science at the time of its publication.

Unfortunately, whereas the first two reasons are unavoidable, the third one is not.

As will be shown in detail below, the ASW checklist cannot be confidently used as a pri-

mary source by taxinomists and other biologists, because of its high rate of mistakes, in

all fields. It will merely be useful as a source of references, but any information that it

contains should be checked against the original publications. It will therefore be impossi-

ble to use this book alone, but its use will have to be combined with the use of other

checklists, sometimes partial and older, some of which are devoid of bibliography, like

that of GORHAM (1974). I wish here to insist on a point. GORHAM's (1974) checklist has

been much criticized by several authors. Some of these criticisms were well-founded, and

it is true that GORHAM'S list is neither perfect nor complete. However it was the result

of the work of a single man, without computer, and was nevertheless extremely useful.

It is true that it contains mistakes and omissions, that it does not give citations of original

descriptions, nor type specimens references, type localities or detailed distributions ; an

important weakness of this book is that it has no index. However, experience has shown

me that, with the help of GORHAM's list and of the Zoological Record, it is usually extre-

mely easy and rapid to trace all the available information in these fields. Furthermore,

GORHAM's book gives some important pieces of information (syÿnonyms, homonyms and

subspecies) which are not given in ASW, and it will therefore be necessary to combine

the use of both books to have a comprehensive view of “the summary of the state of the

literature of amphibian taxonomy” (4SW : 1). A book that would contain the informa-

tion of both GORHAM's and FROST’s books still remains to be written.

It is clear to me that all or most of the authors who have worked on the establish-

ment of the ASW checklist made use of GORHAM’s checklist, at least as a “starting point”

for their compilation. If they have not, they have certainly lost time. For this reason,

I feel that the introductions to ASW do not do enough justice to GORHAM’s work, which

is only mentioned incidentally and rather contemptuously on p. ii as being incomplete.

Incomplete as it is, GORHAM'S list has certainly rendered many services to amphibian

taxinomists since its publication. It has also allowed the realization of quantitative analy-

ses of the characteristics of the classification of the amphibians at the level of higher taxa

(D-30) and of species and subspecies (D-4) ; the latter type of analysis, which takes into

account the dates and current status (as valid species or subspecies, or as invalid synonyms,

homonyms or replacement names) of all species-group names created for amphibians since

1758, would have been impossible on the basis of ASW.

GORHAM's checklist has also been criticized for having introduced new combina-

tions, “none of which is supported by evidence” (DUELLMAN, 1977 : x). However, this

is unavoidable in any checklist, and neither DUELLMAN's (1977) nor FROST’s (1985)

checklists are beyond this criticism. In ASW, new combinations which are not “support-

Source : MNHN, Paris

DuBoIs 113

ed by evidence” appear for example in the genera Rana (African species of the subgenus

Hylarana), Tomopterna (Asian species) and Scutiger (species transferred from the nominal

genus Oreolalax). There is no way to avoid such actions in a checklist if this list is to

be somewhat heterogeneous ; what is problematic here is the author who should be credi-

ted with these new combinations (see above). More open to criticism is the introduction

in the ASW checklist of unjustified emendation, some of which, like Rana quadrana and

Rana unculuana, are truly barbarian (see below).

CHOICE OF THE CLASSIFICATION

Systematics is not yet a unified discipline of biology. There still exist several schools

of systematics, the most important ones of which are the phenetic, cladist and evolution-

ary ones (see e.g. MAYR, 1969, 1981, 1982). Taxinomists belonging to these three schools

have different conceptions of how a classification should be constructed, of what is a “natu-

ral” taxon, etc. These three different schools each have supporters among current amphib-

ian taxinomists. As these authors work on different groups of amphibians, they produce

subclassifications of various subgroups of this class which, being based on different ratio-

nales, are not “compatible” and cannot be truly combined into a single classification.

The building up of a checklist like ASW/is thus difficult as concerns the choice of a clas-

sification. À possible approach would be to choose a single philosophy of classification

and to produce a whole classification of the class under phenetic, cladist or evolutionary

principles. Unfortunately, no such classification exists for amphibians (from the class cate-

gory to the subspecies category), because a lot of relevant basic information on taxa is

still lacking. At this point, any checklist of the amphibians of the world cannot but adopt

an'eclectic frame of classification. While the classification at the higher level may, at least

partially (because of the gaps in the information), follow one of the three philosophies

of classification, subclassifications of lower taxa will have to relÿ mainly on the most recent

revisionary works, which may have been made according to different principles. This

is unavoidable, but should be made clear in the introduction to the checklist.

In the Introduction of ASW, FROST (pp. 1-2) states that the philosophy retained

in this respect has been to follow the most recent révisions. This is only partially true.

In several cases, the most evident of which is the subfamily Raninae, the classification

used in the book is a compromise between various conceptions of the classification of

the group which have been defended by different authors in the recent years. It is not

quite clear to me which principles have guided the authors in making these choices and

compromises, since the philosophy of classification to which they adhere is not explicitly

stated. However, from several remarks scattered throughout the book, I seem to conclude

that they favor the cladist philosophy. What is extremely misleading however is that this

philosophical choice is never explicited, but that in various parts of the book some of

the cladist ideas are presented as universally agreed upon ideas. For example, the terms

“monophyly” and ‘“‘monophyletic”? are used several times in the book, but without sta-

ting if this word is used in its traditional sense, as a qualification of taxa (ASHLOCK, 1971 ;

MAYR, 1974, 1981 ; DUBOIS, D-30, D-33), or in its Hennigian sense, as a qualification

of descent (WILEY, 1981). Confusion would have been spared if the authors had abando-

ned the equivocal term of “monophyletic” and used the well-defined term “holophyle-

tic” (ASHLOCK, 1971) ; for “monophyletic” in the traditional sense, I recently proposed

the new term ‘“‘homophyletic” (D-33), in the hope to stop a long period of confusion.

Source : MNHN, Paris

114 ALYTES 5 (3)

Other examples of the “cryptic” use of a cladist philosophy in this book can be

found : e.g. the assumption that sister groups should automatically be given the same

rank in taxinomic hierarchies (see e.g. SIBLEY & AHLQUIST, 1982 : 13), or the idea that

paraphyletic groups are not “natural” taxa. This later idea is even pushed extremely far

in a sentence like : “This ‘group’ is likely paraphyletic with respect to the Mantellinae

and through that group to the Rhacophoridae.” (4SW : 451). By putting the word group

between quotation marks in this sentence, the authors seem to imply that a paraphyletic

group is not a group, which is philosophically inadmissible ! À group may be natural

or not, homophyletic, holophyletic, polyphyletic or paraphyletic, it is nevertheless a group :

even a group consisting of “all black animals” would be a real group, even if it does not

correspond to a phylogenetic unit.

The author of these lines adheres to the evolutionary school of MAYR (1969, 1974,

1981), as explained in several papers (D-11, D-12, D-14, D-30, D-33), and he considers

that sister groups should not automatically be given the same rank in classification, that

paraphyletic groups may, in some cases, be perfectly natural groups, and that the build-

ing up of a classification should take into account not only the cladistic relationships bet-

ween taxa but also the amount of genetic, phenetic and ecological divergence which has

occurred between taxa after the cladogenesis which separated the lineages from which

they are issued. Such a conception, which relies both on phylogeny and on divergence,

is “synthetic” or “evolutionary”, not “gradist”, as stated in ASW (p. 452), because a

purely gradist approach to classification would take into account only ecological differen-

ces between taxa, without considering their phylogeny.

Divergences of opinion on the “best” system of classification are normal in the

present state of the theory of systematics, and it is the full right of any taxinomist to adhere

to a philosophy of classification of his choice, but such choices should be stated clearly.

Otherwise, the risk is to present as universal truths ideas which are only the matter of

personal opinions, and this is contrary to science.

Not all groups of Amphibia have been subject to recent revisionary works using

modern concepts and methods of analysis, and this makes unavoidable the presence of

important heterogeneities in such a list. Some of these heterogeneities, however, are due

to the compromise which has been tempted in some cases between quite different con-

cepts of the classification of a group. In such cases it would have been better to choose

one of these options (and to clearly mention this choice) rather than to try an impossible

compromise. À few examples may be mentioned in this respect. Among the subfamily

Megophryinae of the Pelobatidae, if full generic rank is to be given to Arympanophrys

and Brachytarsophrys, it should certainly also be given to Ophryophryne, which is much

more divergent than the latter two groups from Megophrys ; if Ophryophryne is to be main-

tained as a subgenus of Megophrys, then these two former groups should be considered

as simple synonyms of Megophrys (see e.g. D-35). The classification retained for the Raninae

is extremely heterogeneous. If full generic rank is given to taxa like Hildebrandtia, Prycha-

dena, Lanzarana, Pyxicephalus, Aubria, Tomopterna, Strongylopus, Altirana and Nano-

rana, it should also be given to several of the subgenera or even species groups main-

tained in the genus Rana in ASW (see e.g. D-35). There is a striking discrepancy in the

checklist between the few cases where small, sometimes monotypic, genera have recently

been established, often as a result of a strict application of the cladist principles, such:

Source : MNHN, Paris

DuBoIs 115

as the “genera”’ Baleaphryne, Lanzarana, Strongylopus, Atympanophrys, Brachytarsophrys,

Kassinula, Tornierella, and other cases where large and clearly heterogeneous ‘‘genera”

are maintained as such, sometimes with a subdivision in subgenera or species groups (Bufo,

Hyla, Litoria, Eleutherodactylus, Rana, Bolitoglossa, etc.), but even sometimes without

such a subdivision : the most extreme example in the latter case is the “‘genus” Manti-

dactylus, where the 10 species groups recognized by BLOMMERS-SCHLÔSSER (1979) are

not even mentioned, although this genus is clearly heterogeneous and corresponds to a

number of “genera” of the kind of Baleaphryne or Lanzarana.

Such problems clearly show that we are still far from having a unified, homoge-

neous frame of classification for all amphibians of the world, from the lowest to the high-

est taxa. This is rather exciting for the future, and it is likely that the classification which

appears in the first edition of ASW will undergo severe modifications in the years to come.

SPECIFIC COMMENTS

Before starting with these detailed comments, I wish to make a few preliminary

remarks. In what follows, I shall discuss at some length various points which may be

considered as relatively minor, if not fully trivial, such as the publication dates of some

names, the authors of these names, their first page of appearance, etc. This should not

be interpreted as an evidence that I consider these details of utmost importance in syste-

matics. The most important part of taxinomic research is by far the biological study of

organisms and of all aspects of their relationships, in order to build up a classification

in agreement with our knowledge of the evolution of these organisms. The nomenclatu-

ral aspect of classification, which is not biology, is only a minor complement to this scientific

work. Nomenclature bears to biological classification the same relationship as grammar

and orthography to literature : the literary value of a book does not lie in its absence of

grammatical or orthographic mistakes, but the presence of such mistakes usually makes

the publication of a book simply impossible ! The reluctance of most systematists to take

care of nomenclatural rules in their work is to my mind of the same kind as would be

the reluctance of a novelist or a poet to follow the rules of ianguage, which in many re-

spects are sometimes as arbitrary and disputable as the rules of zoological nomenclature,

but which impose their strength of law to all humans who want to communicate by print-

ed ways.

Many systematists claim not to be interested in nomenclature, and prefer to work

only on “biological problems”, leaving to other, supposedly less creative, taxinomists,

the care of nomenclatural problems. This is their full right. But when systematists work

on a checklist which is proposed to “serve as a standard reference to amphibian nomen-

clature”, one may expect that they dont have such contempt for nomenclatural problems.

Furthermore, in a checklist, various degrees of precision may be chosen, but, at the level

retained, information must be carefully checked before being published. For example,

in most works dealing with amphibians systematics and faunistics published until now,

the family-group names are given without their authors and dates, and even more without

the precise reference of their first publication (including the page and the original name);

books and papers giving such details are extremely rare (see D-26 : 5). As long as the

family-group names given are the valid ones however, such absence of details is not a

major deficiency ; but insofar as a book gives authors, dates, bibliographic references,

Source : MNHN, Paris

116 ALYTES 5 (3)

pages, original spellings of names, references of first uses of emended spellings, modes

of designation of type species of genera, etc., these problems should be treated seriously.

It is also perfectly possible, e.g. to save space, to build up a very useful checklist without

including bibliographic references to original publications of scientific names : with the

authors and dates of the names, it is a relatively easy task to find the original references,

as outlined above. But if citations, and first pages of publication of names, are to be given

in a checklist, the references should be given in full (including title of the paper or book

and its complete pagination), and the first pages of publications of names should be

checked carefully.

The ASW checklist is an ambitious book. It attempts to give in a limited space

many different kinds of information (taxinomic, nomenclatural, biogeographic, evolutio-

nary). This makes the “venture”, as it calls itself, quite exciting. However, this calls also

for more stringent criticism. The ASW list aims at giving not only the valid names of

taxa, their authors and dates, but also the authors and dates of the justified emendations,

and all the relevant bibliographic references. If all this information is to be added, this

needs a considerable amount of additional work to produce a list with an acceptable rate

oferrors. On the other hand, if adding all this information increases significantly the rate

of errors, the overall quality and usefulness of the book won’t be higher. This is unfortu-

nately the case of ASW. Although it has included among it aims to give all these see-

mingly minor details, the verification work has not been carefully done. This is shown

by the presence in the book of numerous mistakès of various kinds, concerning namely :

valid names of the taxa (in the class-group, family-group, genus-group and species-group);

authors, dates, references and pages of first publications of names ; original spellings of

names ; authors, dates, references and pages of first publications of emended spellings

of names ; type species of genera and subgenera ; modes of designation of type species

of genera and subgenera ; type specimens of species ; type localities of species ; distribu-

tions of species ; and comments. These types of errors will be analyzed in detail below.

I shall also attempt to estimate the quantitative weight of such mistakes in the book,

with the following ideas in mind. It is certainly impossible for anyone, or any group of

persons, to build up a completely perfect checklist. Mistakes and omissions will inevi-

tably persist in any similar monumental work, although successive reeditions of such a

book, taking advantage of the various comments received from colleagues worldwide, are

liable to reduce significantly the rate of mistakes and omissions. The question one can

ask is : what rate of mistakes and omissions is acceptable in the first edition of such a

list ? By “acceptable”, I mean, wich makes the publication of the list worthwhile, useful

to the international scientific community, and worthy to “serve as a standard reference

to amphibian nomenclature”. I would suggest that the acceptable rate should be fixed

at one of the usual levels of mistakes (in measurement, for example) in biological works,

i.e. 0.05, 0.01 or 0.001. Considering the difficulty of the establishment of such a check-

list, I suggest the highest level, 0.05, should be considered as the maximum rate of errors

and omissions (“EO rate”) “acceptable” in such a work. In what follows, however, it

will be shown that this “acceptable” rate is exceeded in several cases in the ASW check-

list, thus making the value of its publication, in its present state, questionable.

Obviously, the following mistakes and omissions are considered in relation to the

status of amphibian taxinomic literature in early June 1984, the date of completion of

Source : MNHN, Paris

DuBoIs 117

Table I. — Correct names of class-group (sensu DUBOIS, D-26) and family-group taxa of Amphib-

ia for which the names given in ASW are incorrect.

Name, author and date Correct information

which appear in ASW

Correct name, author and date References

Class-group taxa :

Caudata Oppel, 1811 (order) Caudata Scopoli, 1777 (super-order) Da de

Urodela Rafinesque, 1815 (order) :

Lissamphibia * (subclass) Batrachia Brongniart, 1800 (subclass) D-19, D-26, D-29

Family group taxa :

Caeciliidae / -inae Gray, 1825 Cecilidae / -inae Rafinesque-Schmaltz, 1814 D-25, D-29

Dermophidae / -inae Taylor, 1969 Siphonopidae / -inae Bonaparte, 1850 D-25, D-29

Grypiscini * Cycloramphini Bonaparte, 1850 D-i9, D-25, D-29

Hemidactylini * Mycetoglossini Bonaparte, 1850 D-25, D-29

Ichthyophidae / sinae Taylor, 1968 Epicriidae / -inae Fitzinger, 1843 D-25, D-29

Petropedetinae Noble, 1931 Phrynobatrachinae Laurent, 1940 (1878) D-l6, D-19, D-26, D-29

Xenopodinae Fitzinger, 1843 Dactylethrinae Hogg, 1838 D-19, D-26, D-29

% ASW provides neither author nor date for this name.

the manuscript. Since then, new species and genera have been described, new data publish-

ed on already known taxa and on nomenclatural problems, but these will hopefully be inclu-

ded in the next edition of ASW.

VALID NAMES OF THE TAXA, WITH THEIR CORRECT AUTHORS, SPELLINGS, DATES,

REFERENCES AND PAGES OF PUBLICATION

In the class-group

Fourteen names of the class-group (as defined by DÜBOIS, D-26) are mentioned

in ASW, 11 of which are considered valid in this book. Of these, 2 are credited with incor-

rect names (see Table I), 12 are either credited with incorrect authors and dates or with

no author and date (see Table II), and 6 with incorrect spellings (see Table IT). The

EO rates for these different types of information are thus respectively 14.3 %, 85.7 %

and 42.9 %. For more detailed discussions of the valid names of these taxa, see D-25,

D-26 and D-29.

In the family-group

For the names of this group, we shall consider independently the information con-

cerning the names themselves, and concerning the various spellings taken by these names.

Table I gives a list of names of the family-group mentioned in ASW which are

incorrect, with the valid corresponding names. Out of a total of 97 family-group names

mentioned in the book, 7 are wrong (EO rate 7.2 %). Table II gives a list of the family-

group names which are credited with incorrect authors and dates in ASW or for which

Source : MNHN, Paris

118 ALYTES 5 (3)

Table 11. — Correct authors and dates of the class-group (sensu DUBOIS, D-26) and family-group

names of Amphibia mentioned in ASW but for which this information is either incorrect

or lacking. The names as given in the first column appear under the spelling which they

bear in ASW, and which in some cases is incorrect (see Table III).

Name which Author and date Correct information

appears in ASW mentioned in ASW

Correct author and date References

Class-group names :

Ambystomatoidea . Noble, 1931 D25, D-29

Amphiumoidea . Cope, 1888 D-29

Apoda © . Merrem, 1820 D25

Caudata Oppel, 1811 Scopoli, 1777 D-26, D-29

Cryptobranchoidea * Bonaparte, 1832 D25, D-29

Gymnophiona Müller, 1831 Rafinesque-Schmaltz, 1814 D-25, D-29

Lissamphibia . Haeckel, 1866 D-19, D-26

Meantes © . Linnaeus, 1767 D41

Plethodontoidea . Smith & Taylor, 1948 D-29

Proteoidea * Müller, 1831 D-25, D-29

Salamandroidea # Müller, 1831 D-25, D-29

Trachystomata . Cope, 1866 DAi

Familÿ-group names :

Allophrynidae “Nomen nudum” Goin, Goin & Zug, 1978 D:31

Alsodini ® . Mivar, 1869 D-19, D-26

Ambystomatidae Hallowell, 1858 Gray, 1850 D-34

Ascaphidae ® . Fejérvéry, 1923 D-19, D-26

Asterophryinae Günther, 1859 1858" Günther, 1858 D-19, D-26, D-29

Atelopodidae © . Fitinger, 1843 D-19, D-26, D-29

Batrachyl Û Gallardo, 1965 D-19, D-26, D-29

Bolitoglossini . Hallowell, 1856 D-29

Bombinatoridae ® Û Gray, 1825 D-19, D-26, D-29

Bombinidae . Fejérväry, 1921 D19, D-26

Brachycephalidae Günther, 1859 “1858” Günther, 1858 D-19, D-26, D-29

Calyptocephalellini . Reig, 1960 D-19, D-26, D-29

Cycloraninae © . Parker, 1940 D-19, D-26

Dendrobatidae Cope, 1865 Cope, 1865 (1850) D-18, D-19, D-26,

D-2%

Dermophiinae Taylor, 1969 b Taylor, 1969 a D41

Desmognathidae / -inae Cope, 1859 Gray, 1850 D-4

Discoglossidae Günther, 1859 1858" Günther, 1858 (1845) D35

Eleutherodactylini Fe Lutz, 1954 D-19, D-26, D-29

Elosiinae © Mirando-Ribeiro, 1926 Miranda-Ribeiro, 1923 D19, D-26

Gripiscini + Mivart, 1869 D-19, D-26

Hemidactylini Q Hallowell, 1856 D25

Herpelinae . Laurent, 1984 D-29

Hylidae / -inae Gray, 1825 (1815) Rafinesque, 1815 D-29

Source : MNHN, Paris

DuBoIs 119

Hylodinae Günther, 1859 “1858 Günther, 1858 D-19, D-26, D-29

Hynobiidae Cope, 1860 1859" Cope, 1859 (1856) D-25, D-29

Leptobrachiinae © * Dubois, 1980 D-19, D-26, D-29

Limnodynastinae Lynch, 1971 Lynch, 1969 D-19, D-26, D-29

Megophryinae Noble, 1931 Bonaparte, 1850 D-19, D-26, D-29

Microhylidae / -inae Günther, 1859 “1858” (1843) Günther, 1858 (1843) D-19, D-26, D-29

Odontophrynini . Lynch, 1969 D-19, D-26, D-29

Pelodryadidae / -inae Günther, 1859 1858" Günther, 1858 D-19, D-26, D-29

Phyllomedusidae / -inae Günther, 1859 “1858 Günther, 1858 D-19, D-26, D-29

Polypedatidae ® Günther, 1859 1858" Günther, 1858 D-19, D-26

Ranidae / -inae Gray, 1825 Rafinesque-Schmaltz, 1814 D-19, D-26, D-29

Rhacophoridae / sine Hoffman, 1932 (1859) Hoffinan, 1932 (1858) D-19, D-26, D-29

Rhinophrynidae Günther, 1859 “1858 Günther, 1858 D-19, D-26, D-29

Salamandridae Gray, 1825 Goldfuss, 1820 D29

Scaphiopodidae / -inae © . Cope, 1865 D-19, D-26, D-29

Scolecomorphidae Taylor, 1969 b Taylor, 1969 a D41

%* ASW provides neither author nor date for this name.

° name mentioned in ASW, but as an invalid name

this information is lacking, with the correct corresponding information. These number

39 (EO rate 40.2 %).

Some of the corrections here proposed deserve a more detailed discussion.

A few mistakes only bear upon the authors and publication dates of names. Among

them, particulary noticeable are the family-group names due to Albert GÜNTHER, and

which are credited in ASW to Günther, 1859 “1858”, because the Catalogue of the Batra-

chia Salientia in the Collection of the British Museum, although dated 1858, was in fact

published on 12 February 1859 (SHERBORN, 1934). However, as I had already noted

(D-18, D-19, D-26, D-29), prior to the publication of his Catalogue, GÜNTHER had written

a paper entitled “On the systematic arrangement of the tailless Batrachians and the struc-

ture of Rhinophrynus dorsalis”, which had been read at the meeting of 22 June 1858 of

the Zoological Society of London, and published on pages 339-352 of the 1858 volume

ofthe Proceedings of this Society, which were published on 9 November 1858 (SCLATER,

1893). In this paper, GÜNTHER gave a preliminary account of the classification of the

anurans to be used in his Catalogue , with the names of all the suprageneric taxa to appear

there ; all these names being accompanied in this paper by clear diagnoses, they are nomen-

claturally valid as of 9 November 1858, and all these names should now be quoted as

of Günther, 1858, not Günther, 1859 “1858”.

In Urodela, it should be noted that I recently discovered that several family-group

names created by GRAY (1850) have been overlooked by all subsequent authors (see D-34).

A few other problems arise from the recent rediscovery of previously overlooked

senior synonyms of family-group names currently considered valid (D-11, D-16, D-18,

D-19, D-25, D-26, D-29, D-32, D-35, D-39). À preliminary remark is necessary here.

When my bibliographic research led me to the rediscovery of all these older names, I

had the choice between two attitudes. The first one, certainly the simpliest and less time-

consuming of both, would have been to follow a single rule in all of these cases (either

Source : MNHN, Paris

120 ALYTES 5 (3)

Table III. — Correct spellings of the class-group (sensu DUBOIS, D-26) and family-group names

mentioned in ASW under incorrect spellings.

Spelling which appears in ASW Correct information

Correct spelling References

Class-group names :

Ambystomatoidea Ambystomatoidei D-25, D-29

Amphiumoidea Amphiumoidei D-29

Cryptobranchoidea Cryptobranchoidei D25, D-29

Plethodontoidea Plethodontoidei D4i

Proteoidea Proteoidei D-25, D-29

Salamandroidea Salamandroidei D-25, D-29

Family-group names :

Hemidacylini Hemidactylini D25

to decide to simply follow the Principle of Priority ; or to decide to ask the ICZN to sup-

press collectively all these older synonyms — as was for example done by MERTENS &

WERMUTH (1960) for eleven specific names of Amphibia and Reptilia). The second atti-

tude, which was chosen, was to study all these cases individually, and to treat them diffe-

rently according to the precise situation encountered in each of them. Thus, in several

cases, it was simply decided to follow the Principle of Priority, because the nomenclatu-

ral disturbance was appreciated as minor ; this has led to the replacement of Bombininae

by Bombinatorinae, of Xenopodinae by Dactylethrinae, of Grypiscini by Cycloramphini,

of Elosiinae by Hylodinae, of Sphenophryninae by Genyophryninae, of Hemidactylini

by Mycetoglossini, of Triturinae by Molginae, of Ichthyophiidae by Epicriidae and of

Dermophiidae by Siphonopidae. In a few other cases, application of Art. 40 of the Code

has allowed the conservation of some junior synonyms having “‘won general acceptance” :

Leptodactylidae, Microhylidae, Rhacophoridae, Hynobiidae. Following certain changes

introduced in the new edition of the Code (ANONYMOUS, 1985 a) in its Art. 32, 35 and

39, changes which are open to criticism (see D-29), some names are conserved but take

the date of an older name, based on an unjustified emendation of a generic name : Pipi-

dae, Megophryinae, Cycloramphini, Hylidae, Microhylidae, Ranidae (see DUBOIS, D-29,

who had overlooked Microhylidae as being in this case). Finally, a few names raise spe-

cial, more important problems, and need special discussions.

1 wish first to underline the fact that I consider the Code as having a strong legisla-

tive value in nomenclature, not only an indicative meaning. In other words, I think excep-

tions to the Rules should be accepted only exceptionally, as seems obvious. Appeal to

the ICZN to use its Plenary Powers to suspend such or such Rule of the Code should

be restricted to the few cases when the nomenclatural disturbance entailed by the simple

application of the Rules would really be important, and should not be made in every

case where a name, which has been used for only a few dozen years and in only a few

Source : MNHN, Paris

DuBois 121

dozen publications, should be replaced by an older one. In this respect, I consider that

the conditions given in Art. 79 (c) (2) of the Code, which requires that a name has been

used “by at least 5 different authors and in at least 10 publications” during the immedi-

ately preceding fifty years, in order to show that “stability is threatened”, are not strin-

gent enough. I would personally favor higher values, say 10 or 20 different authors and

at least 50 or even 100 publications. These principles have guided me in the actions I

have taken regarding family-group nomenclature of Amphibia.

The first name to raise important nomenclatural problems in this class is that of

Discoglossidae : it was discussed in detail elsewhere (D-35).

The second important problem is that raised by the name Dendrobatidae, which

was also discussed in detail elsewhere (D-18, D-21).

The third problem is related to the use of the names Hemimantinae Hoffmann,

1878, Petropedetinae Noble, 1931 and Phrynobatrachinae Laurent, 1940. Strict applica-

tion of the Rules in this case would require retention of the first of these names for this

subfamily. Since this name is based on a generic name (Hemimantis Peters, 1863) which

is a junior subjective synonym of a widely used generic name (Phrynobatrachus Günther,

1862), I considered an action of the ICZN was necessary. For several reasons, in particu-

lar the fact that the name Phrynobatrachinae had been a little more used than the name

Petropedetinae, I suggested that the Commission should make use of its Plenary Powers

to rule that the family-group name Phrynobatrachinae is to be cited as of “Laurent, 1940

(1878)”, and that it has priority over the family-group name Hemimantidae Hoffmann,

1878. This application, although published several years ago (D-16), has still not yet been

voted upon. Despite this application, the choice made in ASW was to use the name Petro-

pedetinae for this subfamily. It is stated in this list that FROST and LYNCH, responsible

for this choice, “disagree with this petition and see the situation as identical to that with

Genyophryninae Boulenger, 1890, vs. Sphenophryninae Noble, 1931, one of simple prior-

ity” (ASW : 439). This is simply not true, because if “simple priority” is to be followed,

the subfamily should be called Hemimantinae Hoffmann, 1878. Contrary to what is sta-

ted in ASW, Art. 40 of the Code cannot be called upon in this case, because Hemimantis

is not a synonym of Perropedetes but of Phrynobatrachus, because LAURENT (1940) was

not aware of the existence of the name Hemimantidae Hoffmann, 1878, and finally because

neither the name Petropedetinae nor the name Phrynobatrachinae can be considered as

having “‘won general acceptance” (see D-16). In the original application, I had mentio-

ned 21 uses of the first name against 10 uses of the second one after 1940. To these fig-

ures, I can now add 14 references to uses of the name Phrynobatrachinae (BOGART &

TANDY,1981 ; DuBois, D-11, D-19, D-26, D-29, D-35 ; MORESCALCHI, 1981 ; NUSS-

BAUM, 1982 ; DREWES, 1984 ; LAURENT, 1980 b, 1984 b, 1986 a ; PERRET, 1984 ;

POYNTON & BROADLEY, 1985 ; LAURENT & FABREZI, 1986), but only one to use of the

name Petropedetinae (BLOMMERS-SCHLÔSSER, 1981). This argument strengthens the case

for the selection of the first of these names rather than the second one for this subfamily,

which has been presented in more detail in my previous paper (D-16 : 137-138). At any

rate this case must be voted upon by the ICZN, and no individual taxinomist can solve

this problem by himself.

The next problem is that raised by the names Dactylethridae Hogg, 1838 and Xeno-

poda Fitzinger, 1843. DUBOIS (D-19, D-26, D-29) pointed to the priority of the first of

Source : MNHN, Paris

122 ALYTES 5 (3)

these names over the second, and estimated that, the second one having been used only

very little, it was not justified in this case to call upon Art. 40 to conserve it (D-26 : 27).

The authors of the ASW list took a different position, and wrote that “the incorrect group

name Xenopinae has had widespread use” (4SW : 425), but they provided no evidence

for that statement (no list of references, not even a number of references), and they pre-

sented “Xenopodinae Fitzinger, 1843” as the valid name for the subfamily. First of all,

it should be noted that if this name was to be conserved by virtue of Art. 40, it would

take the date of the replaced name, and thus be written “Xenopodinae Fitzinger, 1843

(1838)”. Furthermore, this case is clearly one which corresponds with the situation de-

scribed as follows in Art. 40 (b) (ii) of the Code : “In the event of divergent interpreta-

tions of the expression ‘general acceptance’, the case is to be referred to the Commission

for a decision.” In the present case, I consider the name Xenopodinae (or its unjustified

emendations) to have had only a very limited use, not wider than the use of the name

Dactylethrinae, which has priority. Most authors until now have only used the family-

group name Pipidae, and only a minority of authors have divided this family into subfa-

milies. Strict application of the Code requires in this case to follow the principle of prio-

rity. In the case where one author or several would believe that the name Xenopodinae

should be given priority over Dactylethrinae, he or they should submit an application

to the ICZN for a decision, and provide evidence that the stability of nomenclature is

threatened by the simple respect of the Code. Unless, or until, such an action is taken,

and followed by a decision of the Commission, the valid name for this subfamily remains

Dactylethrinae Hogg, 1838. The same holds true for all the other cases listed above where

the nomenclatural disturbance is, in my opinion, minor, and where I replaced recent names

by older synonyms. In particular, the family-group name Hylodinae was resurrected by

Dugois (D-19, D-26, D-29), who gave (D-26 : 32) an extensive nomenclatural discus-

sion of the names Hylodes Fitzinger, 1826, “Hylodes Fitzinger, 1843”, Hylodidae Gün-

ther, 1858, and Elosiidae Miranda-Ribeiro, 1923 (not 1926, as erroneously stated in ASW),

which should be consulted in addition to the quite incomplete discussion given in ASW

(pp. 236-237).

The case of the names Caeciliidae Gray, 1825 and Ceciliidae Rafinesque-Schmaltz,

1814 was discussed in detail elsewhere (D-25, D-29, D-32).

À few words must be said about the name Allophrynidae, which the ASW rejects

as having no status in nomenclature : as shown elsewhere (D-31), it is true that this name

is a nomen nudum in the paper of SAVAGE (1973), but not in the book of GOIN, GOIN

& ZUG (1978), who are therefore the authors, in the nomenclatural sense of the term,

of this family-group name. LAURENT (1980 a, 1986 a) gave good arguments for the inclu-

sion of the genus Alophryne in the family Bufonidae, where it seems best to place it in

its own subfamily Allophryninae (D-19, D-26, D-29, D-31 ; LAURENT, 1986 b).

Besides these mistakes, some omissions bear on the actions of first revisors who

chose between two family-group names published contemporaneously. One of these first-

revisor actions is mentioned in ASW (p. 439, for the names Petropedetinae Noble, 1931

and Cacosterninae Noble, 1931), but the following ones are not (see D-26) : the priority

of Leptodactylidae Werner, 1896 (1838) over Ceratophryidae Tschudi, 1838 rests on the

first revisor action of COPE (1866), who gave priority to Cystignathi Tschudi, 1838 over

Ceratophrydes Tschudi, 1838 ; the priority of Microhylidae Günther, 1858 (1843) over

Kaloulidae Noble, 1931 (1843) rests on the first revisor action of PARKER (1934), who

Source : MNHN, Paris

Dugois 123

gave priority to Gastrophrynae Fitzinger, 1843 over Hylaedactyli Fitzinger, 1843 ; the

priority of Brevicipitinae Bonaparte, 1850 over Engystomatinae Bonaparte, 1850 rests

on the first revisor action of DUBOIS (D-19, D-26) ; the priority of Melanobatrachinae

Noble, 1931 over Hoplophryninae Noble, 1931 was fixed by the first revisor action of

PARKER (1934) ; the priority of Pelobatidae Bonaparte, 1850 over Megophryidae Bona-

parte, 1850 rests on the first revisor action of DUBOIS (D-19, D-26), who gave priority

to Pelobatidae Bonaparte, 1850 over Megalophreidina Bonaparte, 1850 ; the priority of

Dicamptodontidae Tihen, 1958 over Rhyacotritonidae Tihen, 1958 was fixed by the first

revisor action of REGAL (1966).

Let us now look at the information concerning the spellings of family-group names

given in ASW. 131 such spellings are mentioned in the book. Except for one name (Table

TT), the spelling used in the checklist is correct (EO rate 0.8 %). The rate of errors and

omissions is much higher however as concerns the reference to the first use of the justi-

fied emendation of family-group names currently in use. There is no general rule about

giving or not the authors and references to these first uses of justified emendations in ASW.

In some cases no information is given, in some others the correct information is given,

in other ones an incorrect information appears. Table IV gives the correct information

in this respect for the spellings for which ASW provides either no information or an incor-

rect one. The total number of spellings in this case being 102, for a total of 131 spellings,

the total EO rate for this information is 77.9 %.

Let us note in passing that the presentation chosen for this information in ASW

is particularly cumbersome and space inefficient. This is clearly shown, for example, by

the case of the names Arthroleptidae and Arthroleptinae, for which ASW (p. 14) writes :

“FAMILY : Arthroleptidae Mivart, 1869. (...) Comment : As originally formed,

the group name was Arthroleptina ; the first use of the group name Arthroleptidae

was by Dubois, 1981, Monit. Zool. Ital., N.S., Suppl. 15 : 259. (...)

SUBFAMILY : Arthroleptinae Mivart, 1869. (...) Comment : The original spelling

of the group name was Arthroleptina ; the first use of the justified emendation Ar-

throleptinae was by Noble, 1931, Biol. Amph. : 515.”

This mode of notation is much heavier than that adopted e.g. by DUBOIS (D-11,

D-26), which gives exactly the same type of information in three lines, under the form

of a synonymy :

“Arthroleptina Mivart, 1869 : 294. - Genre-type : Arthrolepris Smith, 1949.

Arthroleptinae : NOBLE, 1931 : 515.

Arthroleptidae : LAURENT, 1972 : 200.”

To save space, the same information could also be presented as follows : ‘“‘Arthro-

leptina Mivart, 1869 : 294. / Arthroleptinae : NOBLE, 1931 : 515. / Arthroleptidae : LAU-

RENT, 1972 : 200.”

This type of information (on the first authors to have used the justified emenda-

tion of a given name) may be considered trivial, but is necessary if one wants to present

family-group names, with their correct spellings, under the mode of presentation sugges-

ted and used by SMITH & SMITH (1980) and modified by DUBOIS (D-35). Under such

a mode of notation, the two family-group names here in question should be written as

follows : Arthroleptidae Mivart, 1869 (Laurent, 1972) ; Arthroleptinae Mivart, 1869 (Noble,

Source : MNHN, Paris

124 ALYTES 5 (3)

Table IV. — Correct reference to the first use of the correct spelling of the family-group names

mentioned in ASW but for which this information is either incorrect or lacking.

Correct spelling of the First use of the correct spelling Correct information

family-group name as given in ASW”

Real first use of the correct spelling References

Allophrynidae © # Goin, Goin & Zug, 1978:224 D-31

Alsodini ® ÿ Lynch, 1969 : 3 D-26

Alytidae © Günther, 1859 “1858" : 37 Günther, 1858 : 346 D-26

Ambystomatidae Tihen, 1958 : 20 Steineger, 1907 : 24 D41

Amphignathodontinae ® “ Gadow, 1901 : 139 D-26

Amphiumidae . Gray, 1825 : 216 DAI

Anthroleptidae Dubois, D-11 : 259 Laurent, 1972 : 200 DA, D26

Ascaphidae © . Fejérväry, 1923 : 178 D-26

Asterophryinae . Fejérvéry, 1923 : 181 D26

Astylosterninae * Noble, 1927 : 110 Di, D26

Atelopodidae © . Parker, 1934 : 8 D26

Batrachylini * Lynch, 1971 : 123 D-26

Bolitoglossini * Wake, 1966 D41

Bombinatoridae © . Gray, 1831 : 38 D-26

Bombinidae . Tatarinov, 1964 : 126 D-26

Brachycephalidae * Günther, 1858 : 346 D-26

Brevicipitinae . Van Kampen, 1923 : x D-26

Cacosterninae © . Noble, 1931 : 527 DA, D-16, D-26

Cacciliidae + Bonaparte, 1850 D41

Cacciliinae Taylor, 1969 b : 604 Taylor, 1969 a : 303 D41

Calyptocephalellini G Lynch, 1978 : 42 D-26

Centrolenidae . Taylor, 1951 : 36 D-26

Ceratophryidae © , Parker, 1933 : 12 D-26

Ceratophryinae . Parker, 1935 : 511 D-26

Cryprobranchidae * Cope, 1889 : 18 D41

Ccloraninae © * Parker, 1940 : 2 D-26

Cystignathidae © * Günther, 1858 : 346 D-26

Dactylethrinae ® ji Metcalf, 1923 : 391 D26

Dendrobatidae . Cope, 1865 : 100 DI8, D-26

Dermophidae ® . Laurent, 1984 a : 199 D4i

Dermophiinae . Laurent, 1984 a : 199 DAi

Desmognathinae , Boulenger, 1882 b : vi D4i

Discoglossidae Günther, 1859 “1858” : 37 Günther, 1858 : 346 D-26

Dyscophinae . Gadow, 1901 : 139 D-26

Eleutherodactylini . Lynch, 1969 : 3 D26

Elosiinae © . Lur, 1930 : 195 D-26

Genyophryninac Dubois, D-19 : 274 Gadow, 1901 : 139 D-26

Grypiscini s Lynch, 1969 : 3 D-26

Heleophrynidae Gi Hoffman, 1935 : 2 D-26

Hemidactyliini ° Wake, 1966 : vi D4i

Hemisidae Laurent, “1979” [1980 a] : 417 Cope, 1867 : 198 D:11, D26

Herpelinae ® . Laurent, 1984 a : 199 D41

Source : MNHN, Paris

DuBois 125

Hoplophryninae © . Noble, 1931 : 539 D-26

Hylidae . Bonaparte, 1850 D-26

Hylinae . Gadow, 1901 : 139 D-26

Hylodinae . Savage, 1973 : 354 D-26

Hynobiidae . Cope, 1866 : 103 D41

Hyperolini ® . Laurent, 1972 : 201 D-26

Ichthyophiidae . Taylor, 1968 : x D41

Ichthyophiinae . Nussbaum, 1979 : 13 D41

Kassininae . Dubois, D-11 : 227 D-11, D-26

Leptobrachiinae © . Dubois, D-19 : 272 D-26

Leptodactylinae Noble, 1931 : 504 Metcalf, 1923 : 272 D-26

Leptopelinae . Dubois, D-11 : 227 DA, D-26

Limnodynastinae . Heyer & Liem, 1976 : 5 D-26

Mantellinae . Laurent, 1946 : 336 D-26

Megophryinae . Noble, 1931 : 492 D-26

Melanobatrachinae . Noble, 1931 : 538 D-26

Micrhylidae © Günther, 1859 “1858" : 121 Günther, 1858 : 346 D-26

Myobatrachidae . Schlegel, 1850 : 10 D-26

Odontophrynini . Lynch, 1973: 497 D-2

Pelobatidae Lataste, 1879 : 761 Bonaparte, 1850 D-2

Pelodryadidae © Günther, 1859 “1858" : 119 Günther, 1858 : 346 D-26

Pelodryadinae Duellman, 1977 : 1-25 Dowling & Duellman, 1978 : 37.1 D-26

Pelodytidae . Bonaparte, 1850 D-26

Pelodytinae ® . Fejérväry, 1923 : 181 D-26

Petropedetinae . Noble, 1931 : 520 D-11, D-16, D-26

Philautinae . Dubois, D-11 : 227 D-11, D-26

Phrynobatrachinae © . Laurent, 1940 : 79 D-11, D-16, D-26

Phrynomeridae . Parker, 1934 : 9 D-26

Phrynomerinae . Noble, 1931 : 538 D-26

Phyllobatidae © . Parker, 1933 : 12 D-18, D-26

Phyllomedusidae ° Günther, 1859 “1858” : 120 Günther, 1858 : 346 D-26

Phyllomedusinae * Miranda-Ribeiro, 1926 : 64 D-26

Pipidae * Swainson, 1839 : 88 D-26

Pipinae Noble, 1931 : 491 Metcalf, 1923 : 3 D-26

Platymantinae ® * Savage, 1973 : 354 D-26

Plethodontidae . Gray, 1850 : 5 D41

Plethodontinae . Boulenger, 1882 b : vit D41

Plethodontini . Wake, 1966 : vi D41

Polypedatidae ® . Günther, 1858 : 346 D-26

Proteidae . Hogg, 1838 : 152 D41

Pseudidae . Savage & Carvalho, 1953 : 198 D-26

Ranidae Bonaparte, 1831 : 66 Boie, 1828 : 363 D-26

Raninae . Boulenger, 1888 : 205 D-26

Rhacophoridae * Hoffinan, 1932 : 562 DA, D-26

Rhinatrematidae . Nussbaum, 1977 : 1 D41

Rhinodermatidae + Günther, 1858 : 346 D-26

Rhinophrynidae . Günther, 1858 : 348 D-26

Rhyacotritoninae . Tihen, 1958 : 1 D41

Salamandridae * Gray, 1825 : 215 D41

Source : MNHN, Paris

126 ALYTES 5 (3)

Table IV. — (continued) Correct reference to the first use of the correct spelling of the family-group

names mentioned in ASW but for which this information is either incorrect or lacking.

Scaphiophryninae . Laurent, 1946 : 337 D-26

Scaphiopodidae ® . Cope, 1865 : 104 D-26

Scaphiopodinae ® + Dubois, D-19 : 271 D-26

Scolecomorphidae . Taylor, 1969 a : 297 D41

Sirenidae . Gray, 1825 : 215 D41

Sphenophryninae ® . Noble, 1931 : 531 D-26

Telmatobidae © Miranda-Ribeiro, 1926 : 12 Miranda-Ribeiro, 1920 : 320 D-26

Telmatobisai . Lynch, 1969 : 3 D-26

Typhlonectidae + Taylor, 1968 : xi D41

Uraeotyphlinae * Nussbaum, 1979 : 14 D41

Xenopodinae * Metcalf, 1923 : 3 D-26

+ ASW provides neither author nor date for this spelling

* name mentioned in ASW, but as an invalid name.

1931). Therefore, if the checklist is to give this kind of information, it should give it

correctly.

Finally, some even smaller errors must be mentioned, bearing on the first page

of publication of new names or spellings, even when the reference to the work where

these names or spellings appear is correct (see Table V). It is striking to note that even

in these smaller details the EO rate is high (35.7 %), since out of 70 first pages cited,

25 are wrong. À funny example in this respect is the case of the name Leptodactylidae

Werner, 1896, which is credited with the page 15 in ASW: this is the number of the

page of the renumbered reprint of the paper (which starts on p. 1), but not the real num-

ber in the periodical where the paper was published. To save space, this type of small

mistakes won’t be mentioned again below for generic and specific names, where they also

occur and at a similar rate.

A few additional comments : the first spelling given the name Brevicipitinae by

BONAPARTE (1850) was Brevicipitina, not Brevicipina, as stated in ASW (pp. 355, 677) ;

the first spelling given the name Pelobatidae by BONAPARTE (1850) was Pelobatidae, not

Pelobatina, as stated in ASW (p. 409).

In the genus-group

The number of genus-group and species-group names mentioned in ASW is much

too high to allow for a rapid quantitative survey of the type presented above for class-

group and family-group names. I shall limit myself here to a few examples which struck

me while reading the book. These examples are somewhat biased in that they mostly belong

to the Asian and European groups of Amphibia with which I am personally best acquain-

ted, but there is no reason to believe that they are not representative of the whole class.

An example which supports the latter assumption is a trivial but illustrative one :

it concerns the references to the first pages of publication of the generic names of Amphi-

Source : MNHN, Paris

DUBoIs 127

Table V. — Correct first page of appearance of new names or new spellings of names of the family-

group mentioned in ASW/, for which the bibliographic reference given is correct but not the

page.

Name or spelling mentioned in First page of publication Correct information

ASW with its author and date given in ASW

Real first page of publication References

Ceratophrydes Tschudi, 1838 44 26 D-26

Cophylidae Cope, 1889 390 248 D-26

Cophylinae : Parker, 1934 10 v D-26

Dendrobatidae Cope, 1865 103 100 D-26

Dicamptodontinae Then, 1958 2 1 D41

Dyscophidae Boulenger, 1882 a 179 x D41

Genyophryninae Boulenger, 1890 326 327 D-26

Hemiphractinae : Gadow, 1901 210 139 D-26

Hyperolidae : Laurent, 1951 119 né D-26

Ichthyophidae Taylor, 1968 46 x D41

Leptodactylidae Werner, 1896 15 357 D-26

Leptopelinae Laurent, 1972 198 201 D-26

Microhylidæe : Parker, 1934 9 i D-26

Microhylinae Noble, 1931 537 451 D-26

Myobatrachidae Schlegel, 1850 9 10 D-26

Myobatrachinae : Parker, 1940 6 2 D-26

Philautinae Dubois, 1981 258 221 © D26

Piprina Gray, 1825 213 214 D-26

Plethodontidae Gray, 1850 3 5 D41

Rhinatrematidae Nussbaum, 1977 3 1 D41

Rhyacotritoninae Tihen, 1958 25 1 D41

Sooglossinae Noble, 1931 494 492 D-26

Telmatobii Fitzinger, 1843 43 32 D-26

Telmatobiinae : Vellard, 1951 21 3 D-26

Typhlonectidae Taylor, 1968 231 x D41

bia created by TSCHUDI (1838). After the introduction, TSCHUD/s work is divided into

two parts : in the first part (pp. 27-69), the various genera recognized by this author are

diagnosed and discussed ; in the second part (pp. 70-98), the same genera are listed again,

with their specific contents, the synonymies of the specific names and the distributions

of the species. The new generic names created by TSCHUDI (1838) are nomenclaturally

valid from the first part of this text, since they are associated there to diagnoses. Thus,

if a checklist is to give the precise pages of publications of names (which, of course, is

not absolutely necessary), the pages cited should be those of the first part of this work.

The ASW checklist mentions 24 generic names of TSCHUDI (1838) : for 12 (50.0 %) of

these names (Boophis, Buergeria, Crinia, Hemidactylium, Litoria, Onychodactylus, Pleuro-

dema, Polypedates, Pseudotriton, Pyxicephalus, Strongylopus, Theloderma), the correct page,

in the first part of the work, is given ; for 2 (8.3 %) of these names (Cycloramphus, Lepto-

brachium), the ASW list correctly mentions the page of the second part of the work (this

Source : MNHN, Paris

128 ALYTES 5 (3)

is the correct page there because both these generic names appear with different spellings

in the two parts of the work, and the “correct original spellings” fixed later by first revi-

sor actions are those of the second part) ; for 9 (37.5 %) of these names (Ambystoma, Aste-

rophrys, Cynops, Kalophrynus, Microhyla, Plethodon, Pseudobufo, Sphaenorhynchus, Trachy-

cephalus), the page of the second part of the work is incorrectly given as the first page

of appearance of the name ; finally, for 1 (4.2 %) name (Hynobius), the page 56 (in the

first part) is cited, but the correct page is 60. Thus, for this small information, the EO

rate is 41.7 %. One cannot help from thinking that, if such small mistakes may in some

cases be made by the various contributors of the book, it would have been the work of

the editor and/or the editorial committee to check for homogeneity and precision of such

type of information. If this was not done for well-known and important works like

TSCHUDP's one, it is even less likely to have been done for smaller, less important ones.

It is therefore likely that ASW'is filled up with similar such small mistakes or omissions,

which however cannot be discovered collectively but must be traced individually : in other

words, in any given case, it seems reasonable not to directly use the information given

in ASW , but rather to go back to the original publications for checking. The following

examples taken at random in the book will further illustrate this assertion.

BUFONIDAE. — The generic name Pedostibes must be credited to Günther, 1876, not 1875

(see DUNCAN, 1937).

DENDROBATIDAE. — The authors of the generic name Phyllobates are DUMÉRIL & BIBRON

(1841), not BIBRON in SAGRA (1843), the latter work having appeared later (see SMITH & GRANT,

1958).

DISCOGLOSSIDAE . — Baleaphryne is not a valid genus name : it is at least a subgenus (see

HEMMER & ALCOVER, 1984) and more probably a simple synonym of Alytes (see DUBOIS, D-26).

This latter generic name was created by WAGLER in 1829, not 1830 (see D-26).

HYLIDAE. — The name Hemiphractus was created by WAGLER (1828 a) in a first paper which

appears a few pages before that (1828 b) cited in ASW.

HYPEROLIDAE. — DUBOIS (D-11) recently noted that the name Eremiophilus Fitzinger, 1843

had priority over Kassina Girard, 1853, and discussed the problems raised by the conservation of

this latter name. This question was the matter of an application by DUBOIS et al. (1983) and of

a recent Opinion of the ICZN (ANONYMOUS, 1985 b). — The generic name Megalixalus must be

credited to Günther, 1869, not 1868 (see DUNCAN, 1937).

LEPTODACTYLIDAE. — The author of the names Adenomera and À. marmorata is STEINDACH-

NER (1867), not FITZINGER in STEINDACHNER (1867). — The choice of the name A/sodes rather

than Hammatodactylus as the valid name of the genus containing both species Cystignathus nodosus

and Alsodes monticola implies that the authors of ASW possess some evidence that BELL's (1843)

paper was published earlier than FITZINGER's (1843) book. I do not know of such an evidence,

and would have been interested to know if it exists. If not, both names must be considered pub-

lished on 31 December 1843 (see SMITH & GRANT, 1958), and the choice between them must be

the matter of a first revisor action, but here again we would like to have a reference, LYNCH

(1971 :124) mentioned the problem, but did not solve it, because he only supposed that FITZIN-

GER's (1843) work might antedate BELL's (1843), and indicated that if it was true Hammatodactylus

would have the priority.

MICROHYLIDAE. — DUBOIS (D-26 : 20-21, D-40) showed that, following the designation by

DUMÉRIL & BIBRON (1841 : 740) of Rana ovalis Schneider, 1799 as type species of the nominal

genus Engystoma Fitzinger, 1826, this latter name has priority over Elachistocleis Parker, 1927. He

asked the ICZN to suppress DUMÉRIL & BIBRON's (1841) designation and to designate Rana gib-

bosa Linnaeus, 1758 as type species of Engystoma, following the incorrect actions of FITZINGER

Source : MNHN, Paris

DuBois 129

(1843) (under Systoma) and STEJNEGER (1910). Although this application was sent to the ICZN

on 15 October 1982, it has still not yet been published in the Bulletin of Zoological Nomenclature,

and much less voted upon. — The generic name G/yphoglossus must be credited to Günther, 1869,

not 1868 (see DUNCAN, 1937).

PELOBATIDAE. — It is stated in ASW that the name Carpophrys is a nomen nudum because

it is unsupported by a diagnosis. This is simply not true : this name was clearly associated with

a diagnosis (although in the form of a dichotomic key) in the original publication (ANONYMOUS,

1977). The reasons that make this name nomenclaturally unavailable are different (see D-10) : the

name was published anonymously, and without designation of a type species. Under the provisions

of the current Code, a new generic name, to be available, after 1950 must not have been published

anonymously (Art. 14), and after 1930 must be accompanied by the fixation of a type species (Art.

13b). None of these conditions were met with in the original publication (ANONYMOUS, 1977), but,

quite funnily, they are now met with in ASW since the checklist and its sections are not anony-

mous (the section on Megophryinae-was contributed by Masafumi MATSUI), and since the follow-

ing statement appears in ASW : “Leprolalax oshanensis was placed in the monotypic genus Car-

pophrys..”. By doing so, ASW makes the generic name Carpophrys “almost” nomenclaturally avai-

lable : it now has an author (MATSUI in FROST, 1985), a type species (Megophrys oshanensis Liu,

1950), and an “indication”? (reference to the diagnosis given in ANONYMOUS, 1977) ; fortunately,

this name is published in ASW as a junior synonym and for this reason cannot be made available

(rt. 11 (e) (ii) of the Code).— The generic name Leptobrachella was created by SMITH (1925), not

SMITH (1931) (see D-8).

RANIDAE. + The author of the mantelline generic name Trachymantis is METHUEN (1920),

not “METHUEN & HEWITT (1920)” (see D-7).

RHACOPHORIDAE. - The name Chiromantis was created by PETERS in 1854, not 1855 (see

D:-11).

Altogether, and apart from the problems surrounding TSCHUDPs names, 15 errors

or omissions were mentioned above, for 395 generic names listed in the book, but this

analysis is not at all exhaustive and other mistakes certainly exist : the EO rate for this

item is thus ar least 3.8%, certainly much higher.

Besides the 395 generic names considered valid in ASW/ this list mentions 11 sub-

generic names considered valid, but gives the author and date for only one of them (Echi-

notriton). Authors and dates of the other ones are given here in Table VI. The EO rate

for this information is thus 90.9 %.

Before leaving this chapter let us mention an important information which was

published only after June 1984 and could therefore not be included in ASW but should

appear in the next edition of the book. All authors until now have incorrectly considered

Salamandra genei to be the type species of the genus Hydromantes Gistel, 1848. As shown

elsewhere in detail (D-24), this is incorrect, and the name Hydromantes is a synonym of

Triturus. Fortunately, this name can be replaced without problem by the very similar

name Hydromantoides Lanza & Vanni, 1981, which causes virtually no nomenclatural dis-

turbance (see D-24 for additional comments).

In the species-group

As for the genus-group names, I shall limit myself here to a few examples, and

the following list of mistakes is certainly very incomplete. For simple reasons of time

economy, it was mainly based on my personal knowledge of certain taxinomic groups

or geographical regions. À simple extrapolation from these groups to other ones may not

Source : MNHN, Paris

130 ALYTES 5 (3)

Table VI. — Authors, dates, and type species of the subgenera considered valid in ASW but for

which this information is not given there.

Genus Subgenus Author and date Type species and its mode of designation References

Ceratophrys Stombus Gravenhorst, 1825 Rana comuta Linnaeus, 1758, by subsequent designation

of GRAVENHORST, 1829 D-26

Megophrys … Ophryophryne Boulenger, 1903 Ophryophryme microstoma Boulenger, 1903, by monotypy D-8

Prychadena Parkerana Dubois, 1984 Nomen novum for Abrana Parker, 1931 ; type species is

therefore Abrana cotti Parker, 1931, by monotypy Dil, D-22

Rana Babina Van Denburgh, 1912 Rana holsti Boulenger, 1892, by original designation D-11

Euphiyeris — Fitzinger, 1843 Rana leschenaultit Duméril & Bibron, 1841, by original

designation D-11

Hylarana Tschudi, 1838 Hyla erythraea Schlegel, 1837, by monotypy DA

Limmonectes — Fitzinger, 1843 Rana kuhli Duméril & Bibron, 1841, by

designation D-11

Paa Dubois, 1975 Rana liebigi Günther, 1860, by original designation D-11

Scaphiopus … Spea Cope, 1866 Scaphiopus bombifrons Cope, 1863, by original designation D-41

Seutiger Oreolalax Myers & Leviton, 1962 Seuriger pingif Liu, 1943, by original designation D8

be warranted, since different groups were clearly submitted to different treatments, but

at least this biased sample shows that the book is still rich in mistakes.

BUFONIDAE. — If the name Bufo arabicus is to be considered a synonym of the name Bufo

orientalis, it should replace it as the valid name of the species. — If the name Bufo sulphureus is a

synonym of Bufo koynayensis, it should not be cited as a valid name. — The specific name Hylaplesia

borbonica which appears in SCHLEGEL (1827) is a nomen nudum ; this name became nomenclatu-

rally available only with the publication by TSCHUDI (1838) of a diagnosis of this species (see D-15).

— The nominal species Bufo glaberrimus should be credited to Günther, 1869, not 1868, and the

nominal species Ansonia ornata, Bufo beddomii, Bufo hololius and Pedostibes tuberculosus to Günther,

1876, not 1875 (see DUNCAN, 1937).

DENDROBATIDAE. — The original name of the type species of Dendrobates is Rana tinctoria

Cuvier, 1797, not Calamita tinctorius Schneider, 1799. — The specific name Phyllobates bicolor must

be credited to Duméril & Bibron, 1841, not Bibron in SAGRA 1843 (see SMITH & GRANT, 1958).

DISCOGLOSSIDAE. — The correct name of the type species of Barbourula is busuangensis, not

busangensis.

HYLIDAE. — The authors of the name Hyla albovittata (and of other names published in the

same work) are LICHTENSTEIN, WEINLAND & VON MARTENS (1856), not LICHTENSTEIN & MAR-

TENS (1856) (see D-26). — The correct reference to the name Hyla cinerea is Schneider, 1799, not

1792, and that to the name Hyla leucotaenia is Günther, 1869, not 1868 (see DUNCAN, 1937). —

Bour & DuBOIS (1984) recently gave a discussion about the nomenclatural availability of the name

Rana bicolor Boddaert, 1772.

HYPEROLIIDAE. — The nominal species Hylambates viridis should be credited to Günther, 1869,

not 1868 (see DUNCAN, 1937).

LEPTODACTYLIDAE. — Although FITZINGER (1843) cited “Hylod. lineatus. Dum. Bibr.” as type

species of his new genus Lithodytes, the valid name of this species is Rana lineata Schneider, 1799,

a name mentioned by DUMÉRIL & BIBRON (1841) in the synonymy of their Hylodes lineatus.

Source : MNHN, Paris

DuUBoIs 131

There exists therefore no nominal species “Hylodes lineatus Duméril and Bibron, 1841”. This is

fortunate, because otherwise the name Eleutherodactylus lineatus (Brocchi, 1879) would be a pri-

mary homonym and would have to be replaced. — The correct name of the type species of Cyclo-

ramphus is C. fulginosus, not C. fuliginosus. — The original reference to the name Ceratophrys boiei

is WIED-NEUWIED (1824 : 673), not WIED-NEUWIED (1825 : 592) (see D-26). — The emendation

of the specific name Eeutherodactylus gaigei (Dunn, 1931) into gaigeae used in ASW is unjustified.

As a matter of fact, Art. 31 (a) (i) of the Code reads as follows : “A species-group name, if a noun

in the genitive case formed (...) from a modern personal name that is or has been latinized, is to

be formed in accordance with the rules of Latin grammar”’. The spelling gaigei, dedicated to Helen

T. GAIGE, must be considered as the genitive of the name gaigeus. It is true that most Latin names

of the second declension are of masculine gender, but some are feminine, in particular names of

trees (like fagus), countries (like Aegyprus) or cities (like Corinthus) — just like some names of the

first declension are of masculine gender. Therefore, names like Eleutherodactylus gaigei or similarly

Leptobrachium boringii (dedicated to Alice M. BORING , but not emended in ASW/), are correct and

should not be emended. — The nominal species Cystignathus rhodonotus should be credited to Gün-

ther, 1869, not 1868 (see DUNCAN , 1937).

MICROHYLIDAE . — The original name of the type species, by monotypy, of the genus Rama-

nella, was R. simbiotica, not symbioitica “(in error for symbiorica) ”! The author of the unjustified

emendation symbioitica is PARKER (1934 : 90). — The author of the name Engystoma marmora-

tum, type species by monotypy of Uperodon Duméril & Bibron 1841, is GUÉRIN-MÉNEVILLE (1838),

not CUVIER (1829) (see D-26 : 23). — The nominal species Glyphoglossus molossus should be cred-

ited to Günther, 1869, not 1868, and the nominal species Callula triangularis to Günther, 1876,

not 1875 (see DUNCAN, 1937).

PELOBATIDAE. — The correct spelling of the name of the species called Leprobrachium pul-

lus in ASW is L. pullum. — The author of the specific name Megophrys monticola, synonymous

of Megophrys montana Kulh & Van Hasselt, 1822 a, is SMITH (1931), not KUHL & VAN HASSELT

(1822 b), for the reasons explained elsewhere (D-17). — The specific name Megophrys pachyproctus

is based on the Greek name proctos (vent), of masculine gender, and should not be emended into

pachyprocta. — 1 showed elsewhere (D-17) that the valid name of the species currently known as

Megophrys parva (Boulenger, 1893) is in fact Megophrys monticola (Günther, 1864). To avoid nomen-

clatural disturbance, I addressed on 27 May 1981 an application to the ICZN (D-37), which has

yet not been plublished.

PIPIDAE. — If the synonymy of Xenopus wittei includes Xenopus (laevis) bunyoniensis, this lat-

ter name has priority and the species should be known as Xenopus bunyoniensis.

RANIDAE. — BROWN (in ASW : 459) suggests to use the name Discodeles ventricosus for the

species described as Rana ventricosus by VOGT (1912). However, according to the Code (Art. 52), “a

junior primary homonym is permanently invalid”, and this name is preoccupied by Rana ventricosa

Linnaeus, 1758. The valid name of this species is therefore Discodeles vogti (Hediger, 1934) (not Hei-

diger, as printed in ASW). — The following original names are omitted in ASW/: Rana (Paa) arnoldi,

Rana (Chaparana) fansipani, Rana (Paa) hazarensis, Rana (Euphiyctis) keralensis, Rana (Paa) minica

and Rana (Strongylopus) bonaespei. This latter name (see D-9) is mispelled bonaspei in ASW (p. 522).

— The author of the name Rana dalmatina is Fitzinger in BONAPARTE, 1838, not Bonaparte, 1840

(see D-27). — The valid name of the species called Rana dauchina in ASW (p. 488) is R. daunchina.

— The author of the name Rana limnocharis is not “Boie in WIEGMANN (1835)”, but Gravenhorst,

1829 (see D-28). This case is exactly similar to that of the name Polypedares leucomystax, correctly

analyzed by HOOGMOED in ASW (p. 541), and should be treated in the

same way. — The name Rana mortenseni was synonymized with Rana nigrovittata by SMITH (1922),

and Ï am not aware that it has been resurrected since . — The names Rana quadranus and Rana

unculuanus are based on the Latin word “anus” (vent), and must therefore be treated as invariable

names standing in apposition to the generic name Rana : the emendations guadrana and unculuana

Source : MNHN, Paris

132 ALYTES 5 (3)

which appear in ASW/are thus totally unjustified and unacceptable ! — There exists no such name

as “Rana pullus Stoliczka, 1870”, but a name Rana gracilis var. pulla Stoliczka, 1870 (see D-28 :

155). — The nominal species Ixa/us opisthorhodus should be credited to Gunther, 1869, not 1868,

and the nominal species Ixalus diplostictus, Polypedates beddomii, Polypedates brachytarsus, Polype-

dates formosus, Rana pygmaea and Rana verrucosa to Günther, 1876, not 1875 (see DUNCAN, 1937).

RHACOPHORIDAE. — As mentioned in ASW, the name Rhacophorus moschatus Kuhl & Van

Hasselt, 1822 has priority over the name Hyla reinwardtit Schlegel, 1840. To solve this problem,

T'addressed on 27 May 1981 to the ICZN an application (D-38) which has not yet been published.

— The nominal species Ixalus nasutus, Polypedates cavirostris, Polypedates nasutus and Polypedates

rufescens should be credited to Gunther, 1869, not 1868, and the nominal species Ixalus beddomii,

Ixalus chalazodes and Polypedates jerdonii to Gunther, 1876, not 1875 (see DUNCAN, 1937).

The above mistakes are only examples and certainly much below the total number

of mistakes for this type of information, which could be known precisely only by a de-

tailed checking of the 4015 nominal species’ names, authors, dates and references. They

bear on 60 names, and thus indicate a certainly much underestimated EO rate of at least

1.5 %.

TYPE SPECIES OF GENERA AND SUBGENERA

In the ASW checklist, 406 genus-group names are cited as valid (395 for genera,

11 for non-nominative subgenera). The type species are given for all genera, and for 1

of the subgenera ; the omission rate for this information on the 406 names is therefore

2.5 %. Table VI gives the lacking information for the 10 subgeneric names.

Among the 396 genus-group names for which the list provides the name of the

type species, this information is incorrect for at least 10 of them (at least, because I did

not systematically check the validity of the information for all names, but only noticed

some mistakes while looking through the book). Table VII gives the correct information

in these cases. The error rate for this information on the 406 names is therefore of at

least 2.5 %, and the total EO rate for this information is at least 4.9 %.

As far as the mode of designation of the type species of genera is concerned, this

information is given for only 297 of the 395 genera listed in the book (omission rate

24.8 %). As shown in Table VIII, a detailed analysis shows that the quality of the infor-

mation in this respect greatly depends on the group studied, and hence on the contribu-

tors which were in charge of the various groups (apparently the editor and the checklist

committee did not improve contributors’ works in this respect). Thus the information

in this field is null for 5 families (Discoglossidae, Heleophrynidae, Leiopelmatidae, Pelody-

tidae, Pseudidae), it is between 25 and 90 % for 12 families, between 90 and 100 % for

5 families, and it is 100 % for 15 families. The information is more complete in the Uro-

dela (95.1 %) than in the Gymnophiona (85.3 %) or Anura (70.0 %).

However, even among the 297 genera for which this information is given, many

mistakes remain. Table IX gives a list of the genera for which I noted that the mode

of designation given in ASW was incorrect : they are at least 26 in number (again, this

is a lower estimate, since I did not check the validity of this information for all genera,

far from this). The error rate for this information is at least 8.8 % of the 297 names,

or 6.6 % of the 395 genera listed in the book. The total EO rate for the mode of designa-

tion of type species of genera is thus at least 124 out of 395, i.e. at least 31.4 %.

Source : MNHN, Paris

Dugois 133

Table VII. — Correct information concerning the type species of some genera for which ASW pro-

vides incorrect information.

Correct information

Genus Type species and its mode of designation

as given in ASW Type species and its mode of designation Ref.

Afrisalus — Megalixalus fornasinit congieus Euchnemis fornasinii Bianconi, 1849, by DA1

Laurent, 1944 Laurent, 1941 original designation (LAURENT, 1944 : 111)

Dendrobates Calamita tinctorius Schneider, 1799, by subsequent Nomen novum for Hylaplsia Boie in ScuLece, D-18,

Wagler, 1830 designation of DUMÉRIL & BIBRON, 1841 1827, which is an unjustified emendation of D-21,

Hysaplesia Boie in ScHLeGEL, 1826 ; type spe- D-26

cies is Rana tinctoria Cuvier, 1797, by subse-

quent designation of DUMÉRIL & BiBRON, 1841

Lchthyophis — Ichthyophis hasseli Fitznger, 1826 (nomen nudum) _ Caecilia glutinosa Linnaeus, 1758, by monotypy D-41

Fitzinger, 1826 (= Caecilia hypocyanea Boie, 1827)

Leprodactylus … Rana fusca Schneider, 1799 Rana typhonia Latreille in Sonnint & D-2%6

Fitzinger, 1826 LarreiLLe, 1801 (non Rana syphonia Linnaeus,

1758), by subsequent designation of FITZINGER,

1843

Lithodyres — Hylodes lineatus Duméril & Bibron, 1841 Rana lineata Schneider, 1799, by original D41

Fitinger, 1843 designation

Mertensiella Not designated Nomen novum for Exaeretus Waga, 1876 ; type DA1

WolterstorfT, species is therefore Exaererus caucasieus Waga,

1925 1876, by monotypy

Necturus Necturus lateralis Wagler, 1830 (= Sirena maculosa … Sirena maculosa Rafinesque, 1818, by subsequent D-41

Rafinesque, Rafinesque, 1818), by subsequent designation of designation of Browx, 1908

1819 TANGER, 1843

Pseudophryne … Pseudophryne australis Duméril & Bibron, 1841, by Bombinator australis Gray, 1835, by original D-41

Fitinger, 1843 original designation designation

Rhinatrema Caecilia bivitrata Cuvier, 1829 (nomen nudum) Caerilia bivitrata Guérin-Méneville, 1838, by D-41

Duméril & (= Caecilia bivittatum Guérin-Méneville, 1829) monotypy

Bibron, 1841

Strongylopus … Rana fasciata Tschudi, 1838 No type species at present ; case submitted on 8

Tschudi, 1838 September 1980 by Dusors to the ICZN inan D-I1,

application which has still not been published D-36

Since the various families and genera have been entrusted to different contribu-

tors, a certain heterogeneity was inescapable in their treatment. Homogeneization of these

different contributions, which could have been done by the editors, is clearly wanting.

As in the case of TSCHUD/’s names discussed above, an instructive example in this re-

spect is given by the genus-group names created by FITZINGER (1843) and considered

valid in ASW, which are 16 in number. As remarked by DUBOIS (D-15), all these names

are perfectly valid according to the Code, being associated with clear type species desig-

nations (in a column bearing the title “Typus”?). For 3 (18.8 %) of these names (Limnody-

nastes, Pseudophryne, Tachycnemis), the designation of the type species is correctly stated

in ASW to be “by original designation” ; for 4 (25.0 %) other names (Gastrophryne,

Gastrotheca, Ololygon, Pseudacris), this designation is erroneously stated to be “by mono-

typy” ; for 1 (6.3 %) name (Phrynohyas), it is erroneously stated to be “by monotypy

Source : MNHN, Paris

134 ALYTES 5 (3)

Table VIII. — Quantitative analysis of the information given in ASW on the mode of designation

of the type species of the genera considered valid in ASW. The mode of designation

stated in ASW is sometimes in error : see text and Tables VII and IX.

Order, family Number of genera considered Mode of designation of the type species stated in ASW

valid in ASW

N %

Anura 300 210 700

Arthroleptidae 8 8 100

Brachycephalidae 2 1 50.0

Bufonidae 2 20 80.0

Centrolenidae 2 1 50.0

Dendrobatidae 4 3 750

Discoglossidae 5 ° 0

Heleophrynidae î 0 0

Hemisidae 1 1 100

Hylidae 37 3 622

Hyperoliidae 4 15 92.9

Leiopelmatidae 2 ° 0

Leptodactylidae si 2 431

Microhylidae 59 4 69.5

Myobatrachidae 20 20 100

Pelobatidae 9 4 444

Pelodyt 1 0 0

Pipidae 4 1 25.0

Pseudidae 2 0 0

Ranidae 39 38 974

Rhacophoridae 10 10 100

Rhinodermatidae 1 1 100

Rhinophrynidae 1 1 100

Sooglossidae 2 2 100

Urodela GI 58 95.1

Ambystomatidae 2 2 100

Amphiumidae 1 1 100

Cryptobranchidae 2 2 100

Dicamptodontidae 2 2 100

Hynobiidae 9 8 88.9

Plethodontidae 21 26 2 96.3

Proteidae 2 2 100

Salamandridae 14 E 92.9

Sirenidae 2 2 100

Gymnophiona # 2 853

Ceciliidae 24 2 91.7

Epicriidae 3 1 33.3

Rhinatrematidae 2 1 50.0

Scolecomorphidae 1 1 100

Typhlonectidae 4 4 100

Total Amphibia 395 297 752

and subsequent designation” (!) ; and finally for 8 (50.0 %) other names (Euphlycris, Eup-

sophus, Leptophryne, Limnomedusa, Limnonectes, Lithodytes, Osteopilus, Peltophryne), the

mode of designation of the type species is not stated. The total EO rate for this informa-

tion is therefore of 13 out of 16, i.e. 81.3 %.

Source : MNHN, Paris

DuUBoIs 135

Beside this, it should be noted that FITZINGER (1843) also designated type species

for some genera created before him by other authors. Some of these designations are clearly

acknowledged in ASW (e.g. for the genera Plethodon, Polypedates, Rana) while others are

not. Thus, FITZINGER (1843: 31) clearly designated Rana gryllus LeConte, 1825 as type

species of the genus Acris Duméril & Bibron, 1841, but, following DUELLMAN (1970),

ASW writes that Rana gryllus is the type species of Acris “by fiat”.

Some comments are necessary for some of the information which appears in Tables

VII and IX.

A first comment concerns the type species of Zchthyophis Fitzinger, 1826. FITZIN-

GER (1826 : 36) referred two nominal species to this new genus : Caecilia glutinosa Lin-

naeus, 1758, and “a new species from Java” which he called Zchthyophis hasseltii but for

which he proposed no diagnosis. The name Jchrhyophis hasselti is therefore a nomen nudum

in this work, and Caecilia glutinosa, the only available name associated with the generic

name Jchthyophis in the original description, is the name of the type species of this genus

by monotypy. À similar situation is met with for the genus Pseudorriton Tschudi, 1838,

which was created for the nominal species Sa/amandra subfusca Green, 1818, and for

a second species which is called Triton major on p. 60 and Pseudotriton nigra on p. 95

of TSCHUDPs (1838) work. Both these later names are nomina nuda in this work, and

S. subfusca is therefore the type species of Pseudotriton by monotypy.

A different case is that of the type species of the genus Pseudophryne Fitzinger, 1843,

which is stated in ASW to be : “Phryniscus australis Duméril and Bibron, 1841 [not of

Gray, 1835] (= Pseudophryne semimarmorata Lucas, 1892, according to Parker, 1940, Novit.

Zool., 42 : 101), by original designation”. As a matter of fact, there exists no nominal

species “Phryniscus australis Duméril and Bibron, 1841”. DUMÉRIL & BIBRON (1841 :

725) clearly attributed this species name to GRAY (1835), and it is irrelevant in this re-

spect that the specimens to which they applied this name were in fact members of a biolo-

gical species distinct from that described by GRAY (1835) as Bombinator australis : any

author who misidentifies specimens and applies in error an existing name to them does

not create a new specific name, otherwise the taxinomic literature would be extremely

overcongested (in this respect see D-24) ! Incidentally, the same applies to the problem

of the type species of the genus Synapruranus Carvalho, 1954, and I personally disagree

with the petition presented by LESCURE & NELSON (1977) concerning the type species

of this genus.

Another problem is raised by the type species of the genus Srrongylopus Tschudi,

1838. In a detailed paper (D-36) submitted for publication to the ICZN on 8 September

1980 but which has not yet been published, I presented evidence that this nominal genus

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. I therefore asked the

Commission to use its Plenary Powers to designate for it a type species in agreement

with current usage.

The type species of Caudiverbera Laurenti, 1768 cannot be “Caudiverbera peru-

viana Laurenti, 1768, by tautonymy”” (ASW : 262) ! STEJNEGER (1936) considered C.

Peruviana as the type species of Caudiverbera because this species was “‘the same as Lacerta

caudiverbera Linnaeus”. Had this latter name been cited by LAURENTI (1768) among the

included species of his genus Caudiverbera, it would actually become the type species

Source : MNHN, Paris

136 ALYTES 5 (3)

Table IX. — Correct information concerning the mode of designation of the type species of some

genera for which ASW provides incorrect information.

Genus Type species Mode of designation Correct information

2 pvenin 7 AS Mode of designation Ref.

Acris Duméril & Bibron, Rana gryllus LeConte, 1825 “By fiat” Subsequent designation of D-26

1841 FITANGER, 1843

Ambystoma Tschudi, 1838 Lacerta subviolacea Subsequent designation of Monotypy D41

Barton, 1804 FITanceR, 1843

Caudiverbera Laurenti, 1768 Caudiverbera peruviana Tautonymy Subsequent designation of D-41

Laurenti, 1768 STEMNEGER, 1936

Chrysobatrachus Chrysobatrachus aupreonitens Original designation Monotypy DA1

Laurent, 1951 Laurent, 1951

Colostethus Cope, 1866 Phyllobates latinasus Monotypy Original designation D-26

Cope, 1863

Dpscophus Grandidier, 1872. Dyscophus insularis Original designation Monotypy D-26

Grandidier, 1872

Gastrophryne Fitzinger, 1843 Engystoma rugosum Duméril Monotypy Original designation D-26

& Bibron, 1841

Gastrotheca Fitzinger, 1843 Hyla marsupiata Duméril … Monotypy Original designation D-26

& Bibron, 1841

Hemiphractus Wagler, 1828 Hemiphractus spixit Subsequent designation of Monotypy D-26

Wagler, 1828 Perens, 1862

Hymenochirus Xenopus boetgeri Original designation and Monotypy D-26

Boulenger, 1896 Tornier, 1896 monotypy (!)

Kassinula Laurent, 1940 Kassinula wittei Original designation Monotypy D41

Laurent, 1940 x

Laurentomantis Microphryne malagasia Monotypy Monotypy under D7

Dubois, 1980 Methuen & Hewitt, 1913 Microphryne Methuen &

Hewitt, 1913

Nyctymystes Stejneger, 1916 Nyctimantis papua Original designation Monotypy D-2%6

Boulenger, 1897

Obolygon Fitzinger, 1843 Hyla strigilata Spix, 1824 Monotypy Original designation D41

Osteocephalus Osteocephalus taurinus Original designation Subsequent designation of D-41

Steindachner, 1862 Steindachner, 1862 KELLOGG, 1932

Phrynohyas Fitzinger, 1843 Hyla zonata Spix, 1824 Monotypy and subsequent Original designation D41

designation of the ICZN,

1958, Opin. 520 ()

Phrynomerus Noble, 1926 Brachymerus bifasciatus Monotypy Original designation D-26

Smith, 1849

Proteus Laurenti, 1768 Proteus anguinus Monotypy (!) Subsequent designation of D-41

Laurenti, 1768 STENEGER, 1936

Pseudacris Fitzinger, 1843. Rana nigrita LeConte, 1825 Monotypy Original designation D41

Pseudohemisus Hemisus obseurus Original designation Monotypy D-26

Mocquard, 1895 Grandidier, 1872

Pseudotriton Tschudi, 1838 Salamandra subfusca Subsequent designation of Monotypy D41

Green, 1818 FITZINGER, 1843

Salamandra Laurenti, 1768 Salamandra maculosa Tautonymy Subsequent designation of D-41

Laurenti, 1768 FITZINGER, 1843

Scaphiophryne Seaphiophryne marmorata … Original designation Monotypy D-26

Boulenger, 1882 Boulenger, 1882

Somuncuria Lynch, 1978 Telmatobius somuncurensis Monotypy Original designation D41

Cei, 1969

Source : MNHN, Paris

DuBols 137

Tornierella Ahl, 1924 Tornierella pulehra Monotypy Original designation D41

Ahl, 1924

Triturus Rafinesque, 1815 Triton cristatus Subsequent designation Subsequent designation of D-41

Laurenti, 1768 Gimplied) of Duxw, 1918 FrrziNGER, 1843, under

Triton Laurenti, 1768

by tautonymy of this genus, but, since it was not, it is not eligible for type fixation, and

STEJNEGER‘S action results in a subsequent designation of type species for this genus.

The same situation is encountered in the case of the generic name Sa/amandra. The spe-

cific name Lacerta salamandra Linnaeus, 1758 was not part of the originally included

species of this genus and is not eligible for type fixation. The author of the subsequent

designation in this case is FITZINGER (1843).

On the whole, for only three quarters of the generic names considered valid in the

list, information is given on the mode of designation of the type species : one may sup-

pose that in the remaining quarter of cases, the primary literature (original description,

revision papers) has not been consulted by the authors of the list (if not, why has this

information been omitted in these cases and given in the other ones ?). Furthermore, when

one considers that, even when some information in this field is provided in the list, this

information is incorrect in at least 8.8 % of the cases, it is clear that the primary literature

has not been consulted in more than one quarter of the cases, perhaps as much as in half

of the cases. If the same figures hold also for all other types of information given, espe-

cially at the specific level (where I have not made such a detailed analysis), the situation

is quite unsatisfactory.

TYPE SPECIMENS OF SPECIES

The type specimens section is one of the poorest of all (see Table X). The book

reports about 4015 nominal species. Of these, only 3216 (80.1 %) are acknowledged with

the proper collection number(s) of their type(s) ; for 539 other species (13.4 %), only the

Museum(s) where the types are deposited are listed, or, in the case of syntypes, only a

part of them have been traced ; finally, for 260 names (6.5 %), the types either have not

been traced, or are “‘supposed lost”. Lectotypes and neotypes, which are of particular

interest and more important than original types to list in such a book (since they are more

difficult to trace, not being mentioned in the original publication), are not all listed. As

mentioned above in the “General comments”, even when information is given on type

specimens, it may be wrong, in particular as concerns the types considered as “lost” in

the list (details on this will be published elsewhere).

The quantity and quality of the information on type specimens is highly depen-

dent on the Museum in which these types are deposited. It is clear that a particular effort

of location of types has been done by some, but not all, of the contributors or reviewers

of the checklist, working in some Museums, but this effort has been unequal. Table XI

presents a quantitative analysis of the completeness of the information presented in ASW

on type specimens (irrespective of the correctness of this information, which I did not

check) in the 20 major collections ofthe world. The information, is stated to be complete

(100 %) for only four of these Museums (Lawrence, Ann Arbor, Chengdu, Säo Paulo) ;

it is between 90 and 100 % for 9 Museums, between 50 and 90 % for 5 Museums, and

Source : MNHN, Paris

138

ALYTES 5 (3)

Table X. — Quantitative analysis of the information given in ASW on the type specimens of the

species considered valid in ASW. Signification of the categories used to describe the iden-

tification of the type specimens :

Complete

ie. either all types identified and indicated by their Museum numbers, or types defi-

nitely indicated as lost or destroyed (this information may sometimes be in error ; see

text).

Partial : i.e. Museum only definitely or tentatively identified, but no specimen numbers given, or

partial list of syntypes, or types indicated as “probably lost”, etc.

. types indicated as “unknown”, “not traced”, “not located”, ‘not known to exist”,

Absent :

etc., or also “not stated” or “not designated” (which is misleading, because many of the

type specimens for which ASW gives collection numbers were also not stated or designa-

ted in the original publications).

Order, family

Number of species listed

Identification of the type specimens

as valid in ASW

Partial Absent

N L N N

Anura 3495 2781 76 495 219

Arthroleptidae Le) 44 603 2 °

Brachycephalidae 2 2 100 0 0

Bufonidae 339 266 785 si 2

Centrolenidae &4 &4 100 0 0

Dendrobatidae u6 106 914 7 3

Discoglossidae 14 6 428 4 4

Heleophrynidae 4 2 50.0 2 0

Hemisidae 8 6 15.0 1 1

Hylidae 637 593 93.1 16 2

Hyperoliidae 219 138 60 68 15

Leiopelmatidae 4 3 15.0 1 0

Leptodactylidae 72 624 864 50 48

Microhylidae 281 234 83 31 16

Myobatrachidae 100 94 94.0 2 4

Pelobatidae sa 67 m8 10 7

Pelodytidae 2 0 0 1 1

Pipidae 2% 20 769 1 5

Pseudidae 4 4 100 0 0

Ranidae 603 387 642 168 48

Rhacophoridae 187 ns 615 5 19

Rhinodermatidae 2 2 100 0 0

Rhinophrynidae 1 1 100 0 0

Sooglossidae 3 3 100 0 0

Urodela 357 287 804 32 38

Ambystomatidae 31 2 645 7 4

Amphiumidae 3 2 66.7 0 1

Cryptobranchidae 3 66.7 1 0

Dicamptodontidae 4 15.0 0 1

Hynobiidae 3 25 758 4 4

Plethodontidae 21 200 905 u 10

Source : MNHN, Paris

DuBois 139

Proteidae 6 4 667 ° 2

Salamandridae 53 31 58.5 9 13

Sirenidae 3 ° 0 0 3

Gymnophiona 163 148 90.8 12 3

Ceciliidae 88 8 92.0 4 3

Epicriidae 40 36 90.0 4 0

Rhinatrematidae 9 8 88.9 1 0

Scolecomorphidae 7 7 100 0 0

Typhlonectidae 19 16 #42 3 0

Total Amphibia 4015 3216 80.1 539 260

less than 50 % for 2 Museums. For all other collections altogether, it is a little less than

80 %, and for all Museums altogether a little more than 80 %. The completeness of the

information is a bit higher on the whole for the Museums or Universities in which work

members of the Checklist Committee under the auspices of which the ASW list was com-

piled : Lawrence (100 %), San Francisco (96.3 % ; not among those listed in Table XI),

Ann Arbor (100 %), London (73.6 %) and Leiden (96.8 %) — thus 93.3 % on average.

Some major Museums like Paris or Berlin Museums were clearly not contacted for a veri-

fication of the correctness or a completion of the information on type specimens given

in ASW. Among the 255 type specimens (or type series) listed as being deposited in the

Paris Museum, only 229 (89.8 %) are definitely identified with their collection numbers.

However, among those, I found at least 22 (9.6 %) wrong sets of information (wrong num-

bers, incomplete series of syntypes, lectotypes not mentioned as such, etc.) (more details

on this question will be given in my forthcoming catalogue of the type specimens of anu-

rans in the Paris Museum collection). Thus the total EO rate for the information concern-

ing type specimens in the Paris Museum is at least 18.8 %.

The authors of ASW do not seem to be aware of Art. 74 (b) of the Code, which

states that any author who has published the inference that a specimen, which was part

of the original syntypes of a nominal species, is the “holotype’”’ or the “type”, is deemed

to have validly designated a lectotype. This “designation of lectotype by inference of

holotype” is not that rare, and should be carefully traced in a checklist which has the

ambition to list all type specimens of a group. Among all the specimens which are listed

as “holotypes” in ASW, at least those of the following species (and certainly others) are

in fact lectotypes designated “by inference of holotype” : Hyla boans, Hyla chinensis, Lim-

nodynastes peronii, Rana delacouri, Rana fuscigula, Rana kuhli, Rana moluccana and Rana

sanguinea.

Among the lectotypes and neotypes which are not mentioned in ASW/, are at least

the lectotype of Laurentomantis ventrimaculata (D-7) and the neotype of Nyctixalus mar-

garitifer (D-11).

OTHER TYPES OF INFORMATION

For all other types of information (type localities, distributions, comments, biblio-

graphic references), I did not attempt an exhaustive-analysis, but only picked up various

types of errors and omissions, in the groups which I know better, during a rapid survey

of the book.

Source : MNHN, Paris

140 ALYTES 5 (3)

Table XI. — Quantitative analysis of the information given on the type specimens listed in ASW,

according to their Museum of deposition.

This information is given for the 20 largest collections of Amphibians in the world (more

than 40 types or type series mentioned in ASW). The information is considered complete

when the collection numbers are given in full in ASW/and when the status of type is consi-

dered definite, not doubtful, in ASW/ (this information may sometimes be in error ; see text).

N : number of species for which ASW indicates the presence, at least doubtful, of a type

specimen or a type series in this Museum.

€ : number of species for which the information on these type specimens is complete.

% : ratio of C to N.

Anura Urodela Gymnophiona Amphibia

Museum

NERCOER N OC % N € % N C %

London 659 472 716 24 22 917 3%4 4 100 717 528 736

Washington 193 188 974 73 69 945 12 10 83.3 278 267 96.0

Paris 225 205 91.1 19 14 737 11 10 909 255 229 898

Harvard 221 216 97.7 12 12 10 14 4 100 247 242 980

Lawrence 181 181 100 12 12 10 4. 4 10 197 197 100

Berlin 179 87 486 5 4 800 1922865; 50.0 194 96 49.5

New York 133 128 962 10 10 100 13 12 923 156 150 96.2

Chicago 9 90 90.9 46 45 978 10 9 900 155 144 929

Ann Arbor 78 78 100 16 16 100 4. 4 100 98 98 100

Philadelphia 54 43 796 18 14 778 20 #20 100 T4 59 797

Wien 57 54 947 TT 100 Nul T1 68 958

Tervuren 71 35 493 0 0 0 0 - 71 35 493

Frankfurt-am-

Main 66 62 93.9 10 0 2% 0:20 "100 69 64 928

Chengdu 59 59 100 TAMCT. 100 0 0 - 66 66 100

So Paulo* 66 64 97.0 00 CE 66 64 970

Leiden 52 50 962 6 6 100 5 15 100 63 61 968

Genova 56 44 78.6 0 0 11. «100! 57 45 789

Genève 47 45 95.7 CE 0 0 = AT AS ne 9527

Rio de Janciro 43 35 814 0 0 1. 1 10 44 36 818

Säo Paulo** 41 41 100 9 oo 1. 1 10 42 42 100

Other collections 848 663 78.2 84 71 845 36 28 718 968 762 18.7

Total 3428 2840 828 340 309 90.9 167 149 89.2 3935 3298 838

* Werner C. A. Bokermann, Parque Zoolégico de Säo Paulo

** Universidade de So Paulo, Museu de Zoologia

BUFONIDAE. — On the basis of various works, DUBOIS (D-19, D-26, D-29) tentatively re-

cognized 5 subfamilies in this family, which is not even mentioned in ASW. — Bufo himalayanus

is also present in the western part of Himalayas (DUBOIS & MARTENS, 1977 ; DUBOIS, D-6, D-13).

— The name Bufo bufo formosus, often cited in the literature, should be mentioned as a synonym

or subspecies of Bufo japonicus (see MATSUI, 1984). — The validity of the placement of the species

Source : MNHN, Paris

DuBoIs 141

Bufo kelaartii in the genus Bufo has been questioned (see INGER, 1972 ; DUBOIS, D-26). — DuBOIS

(D-1) gave an extensive discussion of the systematics of the species Bufo stomaticus and of related

forms. — The current status of the name rondoensis, type species of the genus Mertensophryne, should

be given. — The recent work by WAKE (1980) should be cited in the comments of Nectophrynoides.

DENDROBATIDAE. — The history of this family-group name is incomplete, not mentioning

the name Eubaphidae Bonaparte, 1850 and the possible use of Art. 40 in particular (see D-18).

Besides, as shown in detail elsewhere (D-18, D-21), the generic name Dendrobates Wagler, 1830

is a substitute name for Hylaplesia Boie in Schlegel, 1827, which is itself an unjustified emendation

of Hysaplesia Boie in Schlegel, 1826.

HYLiDAE. — Dugois (D-14, D-23) suggested to consider Pseudacris as a subgenus of Hyla,

and consequently proposed the replacement name Hyla (Pseudacris) nigrita floridensis for the sub-

species previously known as Pseudacris nigrita verrucosa.

LEPTODACTYLIDAE. — In ASW/, the species Physalaemus nattereri is placed in the P. biligoni-

gerus group, but P. biligonigerus is placed in the P. cuvieri group ! — The nominal species Batra-

chophrynus patagonicus could not have been transferred to Arelognathus previous to the description

of this genus, of which furthermore it is the type species !

MICROHYLIDAE. — The distributions of Kaloula pulchra, Microhyla ornata and Uperodon glo-

bulosus include Nepal (D-1, D-3, D-6, D-13).

MYOBATRACHIDAE. — Since the original names of the species Crinia bilingua, C. deserticola,

C. remota and C. signifera contained the genus-group name Ranidella, it is clear that these species

were “formerly in Ranidella” (ASW : 402)!

PELOBATIDAE. — DUBOIS (D-19, D-26, D-29) recognized 4 subfamilies in this family on the

basis of the works by DUBOIS (D-8), ROËEK (1981) and SOKOL (1982). — The range of Megophrys

parva includes eastern and central nepal (D-1, D-3, D-6, D-13).

RANIDAE. — Laurentomantis Dubois, 1980 is a replacement name for Microphryne Methuen

& Hewitt, 1913, not for Trachymanris Methuen, 1920. — Without any explanation, the genus Phryno-

batrachus is said to “include” the genera Stenorhynchus, Hemimantis, Hylarthroleptis, Pararthrolep-

tis (part) and Pseudarthroleptis. In fact, although this is not stated in ASW, this means that the

synonymy first proposed by DUBOIS (D-11 : 253) for this genus is followed — except that the name

Micrarthrolepris should be added to the above list. The specification “part”? (after Pararthrolepris)

is not relevant here : a synonymy of generic names means that the type species of the various nomi-

nal genera are considered congeneric, irrespective of the fact that the authors of these generic names

may or not have included other species in these genera. In this case, the species Pararthrolepris

nanus, type species of Pararthroleptis, is included in Phrynobatrachus, thus agreeing with DUBOIS

(D-11) in considering Pararthrolepris as a synonym of Phrynobatrachus.

Dusois (D-1) suggested that Amolops kaulbacki was possibly a subspecies of À. formosus, not

of À. afghanus. — Lanzarana was explicitly (and not “by implication”) regarded as a subgenus of

Rana by DUBOIS (D-22). — The problems associated with the conservation of the name Plaryman-

tis in the place of Cornufer were discussed by DUBOIS (D-11). — The genera related to Euphlyctis

mentioned by DUBOIS (D-11) cannot correspond to all the platymantines of SAVAGE (1973), as sug-

gested in ASW (p. 478), since DUBOIS considered the latter heterogeneous ! The genera Amolops,

Micrixalus and Staurois at least are certainly not closely related to Euphlyctis (see D-35).

It is not true that CLARKE (1981) rejected the view that Euphlyctis and Limnonectes are“close”

to Tomopterna (ASW : 478). He only rejected the possibility to consider them as sister groups. The

use of the vague term “close” (which does not mean “sister group”) by DUBOIS (D-11) was pur-

poseful, because (1) we still lack a lot of information on the phylogenetic relationships within the

ranines ; and (2) this term conveys the idea that, whatever the phylogenetic relationships between

the taxa, they have not experienced considerable divergence since the cladogenesis which has sepa-

rated the lineages which led to them ; this latter type of information is considered by cladists of.

Source : MNHN, Paris

142 ALYTES 5 (3)

no value for the construction of a classification, but evolutionary systematists do not agree with

them in this respect. — The use of the terms “species group” and ”’species complex” is not consis-

tent within the genus Rana : the Rana pipiens “complex” includes several “groups”, while the

“Rana macrodon complex” is included in the “Rana grunniens group”. DUBOIS (D-4, D-14, D-30)

suggested some rules for the use of these categories in zoology, in particular in Amphibia, which

are not discussed or taken into account in ASW. — The evolutionary and systematic problems posed

by the Rana pipiens group were discussed in some detail by DUBOIS (D-4). — The references given

in ASW on the problems posed by the Rana esculenta group are most insufficient. In particular,

the paper by DUBOIS & GÜNTHER (1982) should at least be cited and discussed, even if their pro-

posals were not adopted. These authors argued that forms like Rana esculenta or, in salamanders,

Ambystoma platineum, which arose from the hybridization of “good species” but do not behave

genetically and evolutionarily like “normal hybrids”, should not be considered like normal species

or hybrids, but as belonging to a new category of the species-group, different from both the catego-

ries species and subspecies, and which they proposed to call “klepton”. Any klepton may be grouped

with the “good species” from which it arose by hybridization in a “synklepton”. Following these

proposals, amphibians kleptons should have been listed in ASW as Rana kl. esculenta or Ambys-

toma KI. platineum, such a way of notation indicating their peculiar characteristics. — Rana blanfor-

di is absent from Uttar Pradesh and all the central and western Himalayas (D-2, D-3, D-5, D-6,

D-13). — The name Rana fusca Blyth is preoccupied by several senior primary homonyms, not

only by “Rana fusca Meyer”. — The status of the species Rana crassa was discussed in some detail

by DUBOIS (D-1). — The ranges of Amolops monticola, Rana humeralis, Rana nigrovittata and Rana

taipehensis extend to the west to eastern Nepal (D-1, D-3, D-6, D-13). — The range of Rana polu-

nini was recently discussed by DUBOIS (D-5). — It is not GORHAM (1974), but LIU (1935) who

synonymized Rana shini whith Rana spinosa. — The status of the types of Rana sikimensis was

discussed in detail by DUBOIS (D-3). — Dugois (D-20) placed the species Rana leucorhynchus in

the Rana breviceps group of the subgenus Tomopterna, not in the Rana rufescens group (transferred

later in the subgenus Féjervarya by DUBOIS, D-28). — The species Rana sauriceps belongs to this

latter subgenus, not to Hylarana (see D-28).

RHACOPHORIDAE. — The name Philautus was not proposed as the name of a new genus,

but as a replacement name for Orchestes. — The ranges of the species Philautus annandalii, Polype-

dates leucomystax, Polypedates maculatus and Rhacophorus maximus include Nepal (D-1, D-3, D-6,

D-13).

AMBYSTOMATIDAE. — Some “forms” of the genus Ambystoma, such as Ambystoma kl. plati-

neum and Ambystoma K1. tremblayi, are not “true species” but belong in fact to the category klep-

ton, as defined by DUBOIS & GÜNTHER (1982).

SALAMANDRIDAE. — DUBOIS (D-14, D-30) suggested that Tylorotriton and Echinorriton be con-

sidered as subgenera of Pleurodeles.

Again, the above list of mistakes and omissions is certainly very incomplete, since

I did not, far from this, check all the information provided for the 4015 species of the

list. However the total number of mistakes is 52, for a total of 4488 taxa (4015 species,

395 genera, 78 families and subfamilies), i.e. an estimated EO rate of at least 1.2 % for

all types of miscellaneous information together.

FINAL MISCELLANEOUS COMMENTS

Thanks to the use of a computer, the index of this book has been carefully pre-

pared. AIl the names which I have looked for and which are mentioned in the book appear

in the index. This is a strong superiority of this book over GORHAM's (1974) cheklist,

for which the absence of any index is a major weakness. The existence of a finely pre-

pared index in ASW is another reason for regretting the absence in this cheklist of the

Source : MNHN, Paris

DuBols 143

synonyms and of the subspecies (except for arbitrary exceptions). As mentioned above,

this will limit the usefulness of this index in allowing to avoid creating homonyms within

a given genus.

I only found one misprint in the index of ASW : the name “iris, Hyla” appears

on p. 693 between “houyi, Hyperolius” and “hubeiensis, Rana”, and is lacking in its prop-

er place on p. 699.

In the book itself, the following mistake may be noted : on p.598, under Plethodon

serratus, the information on the type locality appears under the title “type species”, and

several lines are out of line in relation to the margin.

Recommendation 28A of the Code reads : “A species-group name should not be

put as the first word of a sentence, to avoid its being printed with a capital initial letter.”

Actually, the editors of ASW ignored this recommendation, at least in the case of the

name Rana ventricosus (p. 459), and possibly in others (I did not look systematically for

this type of error).

The binding of the book is solid, and has resisted my repeated inquiries in the

volume. However, the specimen I received for review had a defect in the back cover.

CONCLUSION

Table XII summarizes the data concerning the EO rates estimated above. The ave-

rage EO rate for all these 18 types of data is 33.3 %, i.e. much higher than the 5 % thresh-

old which we had fixed a priori as an “acceptable” rate of error or omission for such

a checklist. However this average value is of little interest, because the types of informa-

tion concerned are most unequal in importance and interest, because the precision of the

estimate of the EO rate is variable from one case to another, and finally because no EO

rate was calculated for some types of information. In particular, no EO rate was estima-

ted for the completeness of the list as far as the species are concerned, but such a rate

would certainly be very low. On the other hand, a high rate of error may be expected

for real dates of publications of papers or books, since, as exemplified several times above,

no systematic research of this information was carried out (as shown e.g. by the fact that

papers like SCLATER, 1893 and DUNCAN, 1937 are never cited). What is relevant how-

ever is the fact that, out of 18 different types of information studied, only five (Nos. 6,

10, 12, 13, 18) have an EO rate below the 5 % threshold. Furthermore, for 4 of these

5 items, the EO rate is clearly underestimated, for reasons explained above. For the 13

other items, the rate is between 7.2 % and 90.9 %, which is much too high to be “‘accep-

table” according to our a priori criterion.

This and the detailed discussion above clearly point to the fact that this checklist

has been prepared and published much too quickly and that it does not fit the require-

ments which it should fit to be fully useful to the international batrachological commu-

nity. What may be feared now is that, despite its statement to the contrary (4SW : 1),

this checklist might tend “to standardize or institutionalize amphibian taxonomy”. If the

numerous mistakes which appear in the book are uncritically repeated by many authors,

they will become more difficult to rectify. My hope in working on this detailed review

has been to limit at least partially this negative impact by providing corrections to some

of the mistakes of the book. Other mistakes certainly remain. It will be necessary to take

Source : MNHN, Paris

144 ALYTES 5 (3)

Table XII. — Rates of errors (E rate) and of omissions (O rate), and cumulated rate of errors and

omissions (EO rate) for various types of information given in ASW/ (see text for expla-

nation). Rates are given in percent.

Type of information Table E rate Orae EO rate

1. Valid names of taxa of tHe class-group I 43 - 143

2. Correct authors and names of taxa of the class-group ul 143 714 85.7

3. Correct spellings of names of taxa of the class-group ml 429 - 429

4. Valid names of taxa of the family-group 1 72 - 72

5. Correct authors and names of taxa of the family-group nu 231 165 402

6. Correct spellings of names of taxa the family-group ml 08 - 08

7. Correct references 10 the first uses of the correct spellings of

names of taxa of the family-group w 122 656 779

8. Correct first page or appearance of new names or new spellings of

names of taxa of the family-group v 35.7 - 357

9. Correct first page of appearance of generic names created by

TscHut (1838) Text a17 - A7

10. Information on valid names, authors, dates and references of

genus-group taxa (excluding TscHUDIs names) Text 30* 08* 38°

11, Correct authors and dates of names of taxa of the rank subgenus vI - 90.9 90.9

12. Information on valid names, authors, dates and references of

species-group taxa Text 153 02* 1,5%

13. Correct type species of nominal genera and subgenera VI - VIT 25* 25 49*

14. Correct mode of designation of type species of nominal genera VIII - IX 6.6* 248 314*

15. Correct mode of designation of type species of nominal genera

created by FITZINGER (1843) Text 313 50.0 813

16. Identification of the type specimens of nominal species x - 199 19.9

17. Identification of the type specimens of nominal species deposited

in the Paris Museum Text 10,2 g6* 18.8*

18. Various informations on taxa Text 0.8* 04 12*

* this rate is certainly underestimaded (see text).

advantage of these and other comments to correct the list and prepare an improved second

edition of this book. The sooner this revised edition appears, the better, since it will limit

the spread of some mistakes in batrachological publications. I would therefore suggest

that, except for the major institutions which cannot do without buying this expensive

volume for their libraries, individual batrachologists rather await the second, revised, edi-

tion of the book to buy it. Hopefully, this will include synonyms and subspecific names,

which would make it a most useful tool for all biologists interested in Amphibia.

RÉSUMÉ

Le volume Amphibian species of the world publié en 1985 sous la direction de D.R.

FROST est le résultat d’un travail collectif dû à 59 auteurs. Il donne les noms de 4015

espèces actuelles d’Amphibiens, ainsi que diverses informations les concernant. Bien qu’une

telle liste soit d’un grand intérêt potentiel, aussi bien pour les systématiciens que pour

Source : MNHN, Paris

DUBOIS 145

tous les autres biologistes travaillant sur les Amphibiens, cette première édition pose plu-

sieurs problèmes importants. Le choix des informations données dans cet ouvrage, notam-

ment, est fort discutable : en particulier, l’absence des synonymes et des noms subspéci-

ques réduit grandement l’utilité d’une telle liste. De plus, même pour les données qui

y figurent, le taux d’erreurs et d’omissions, tel qu’on peut l’estimer pour différents types

d'informations, est bien trop élevé pour que cette liste puisse être considérée comme un

document de base fiable dans les domaines de la taxinomie, de la nomenclature et de la

bibliographie batrachologiques. De telles faiblesses proviennent manifestement de ce que

cette première édition a été préparée et publiée bien trop vite, et il paraît en conséquence

indiqué d’attendre au moins la deuxième édition, révisée, de cet ouvrage.

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 and D-29 are not repeated here.

ANONYMOUS, 1977. — Systematic keys to the Amphibians of China. (In Chinese). Beijing, Kexue

Chupanche : [i=iv] + iv + 1-93, pl. I-XVIL.

— 1985 a. — International code of zoological nomenclature. Third edition. London, International

Trust for zoological Nomenclature : i-xx + 1-338.

— 1985 b. — Opinion 1364. Kassina Girard, 1853 (Amphibia, Anura) : conserved. Bull. zool. Nom.,

42 : 355-356.

ASHLOCK, P. D., 1971. — Monophyly and associated terms. Syst. Zool., 20 : 63-69.

BLOMMERS-SCHLÔSSER, R. M. A., 1981. — On endemic Malagasy frogs (Ranidae, Rhacophoridae

and Hyperoliidae). Monit. zool. ital., (n. s.), suppl. 15 : 217-224.

BOULENGER, G. A., 1882 a. — Catalogue of the Batrachia Salientia s. Ecaudata in the collection of

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

Er 1882 b. — Catalogue of the Batrachia Gradientia s. Caudata and Batrachia Apoda in the collec-

tion of the British Museum. London, Taylor and Francis : i-viii + 1—127, pl. I-IX.

BoUR, R. & DUBOIS, A., 1984. — Nomenclatural availability of Testudo coriacea Vandelli, 1761:

à case against a rigid application of the Rules to old, well-known zoological works. 3. Her-

per, 17 : 356-361.

CLARKE, B. T., 1981. — Comparative osteology and evolutionary relationships in the African Rani-

nae (Anura Ranidae). Monit. zool. ital., (n. s.), suppl. 15 : 285-331.

Copr, E. D., 1866. — On the structures and distribution of the genera of the arciferous Anura.

J. Acad. nat. Sci, Phila., (2), 6 : 67-112.

DREWES, R. C., 1984. — A phylogenetic analysis of the Hyperoliidae (Anura) : treefrogs of Africa,

Madagascar, and the Seychelles Islands. Occ. Pap. Calif. Acad. Sci., 139 : i-x + 1-70.

DuBois, A., 1974 (D-1). — Liste commentée d’Amphibiens récoltés au Népal. Bull. Mus. natn.

Hist. nat., (3), 213 (Zool. 143) : 341-411.

- 1975 (D-2). — Un nouveau sous-genre (Paa) et trois nouvelles espèces du genre Rana. Remar-

ques sur la phylogénie des Ranidés (Amphibiens, Anoures). Bull. Mus. natn. Hist. nat., (3),

324 (Zool. 231) : 1093-1115.

-— 1976 (D-3). — Les Grenouilles du sous-genre Paa du Népal (famille Ranidae, genre Rana).

Cahiers népalais - Documents, Paris, CNRS, 6 : i-vi + 1-275.

- 1977 (D-4). — Les problèmes de l’espèce chez les Amphibiens Anoures. Mém. Soc. zool. France,

39 : 161-284.

Source : MNHN, Paris

146 ALYTES 5 (3)

= 1979 (D-5). — Notes sur la systématique et la répartition des Amphibiens Anoures de Chine

et des régions avoisinantes. II. Rana blanfordii Boulenger, 1882, Rana polunini Smith, 1951

et Rana yadongensis Wu, 1977. Bull. Soc. linn. Lyon, 48 : 657-661.

ER 1980 a (D-6). — L'influence de l’homme sur la répartition des Amphibiens dans l'Himalaya

central et occidental. C. R. Soc. Biogéogr., 55 : 155-178.

= 1980 b (D-7). — Un nom de remplacement pour un genre de Ranidés de Madagascar (Amphi-

biens, Anoures). Bull. Mus. nan. Hist. nat., (4), 2 (A) : 349-351.

- 1980 c (D-8). — Notes sur la systématique et la répartition des Amphibiens Anoures de Chine

et des régions avoisinantes. IV. Classification générique et subgénérique des Pelobatidae

Megophryinae. Bull. Soc. linn. Lyon, 49 : 469-482.

— 1981 a (D-9). — Deux noms d’éspèces préoccupés dans les genre Rana (Amphibiens, Anoures).

Bull. Mus. nat. Hist. nat., (4), 2 (A): 927-931.

— 1981 b (D-10). — Notes sur la systématique et la répartition des Amphibiens Anoures de Chine

et des régions avoisinantes. V. Megophrys oshanensis Liu, 1950 et Leprobrachium minimum

Taylor, 1962. Bull. Soc. linn. Lyon, 50 : 182-192.

-— 1981 c (D-11). — 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.), suppl. 15 : 225-284. .

= 1981 d (D-12). — Quelques réflexions sur la notion de genre en zoologie. Bull. Soc. zool. France,

106 : 503-513.

— 1981 e (D-13). — Biogéographie des Amphibiens de l'Himalaya : état actuel des connaissances.

In : Paléogéographie et biogéographie de l'Himalaya et du sous-continent indien, Paris, CNRS,

Cahiers népalais : 63-74.

= 1982 a (D-14). — Les notions de genre, sous-genre et groupe d'espèces en zoologie à la lumière

de la systématique évolutive. Monir. zool. ital., (n. s.), 16 : 9-65.

re 1982 b (D-15). — Leprophryne Fitzinger, 1843, a senior synonym of Cacophryne Davis, 1935

(Bufonidae). J. Herper., 16 : 173-174.

= 1982 c (D-16). — Phrynobatrachinae Laurent, 1940 (Amphibia, Anura) : proposed conserva-

tion. Z.N. (S.) 2362. Bull. zool. Nom., 39 : 134-140.

-—— 1982 d (D-17). — Le statut nomenclatural des noms génériques d'Amphibiens créés par Kuhl

& Van Hasselt (1822) : Megophrys, Occidozyga et Rhacophorus. Bull. Mus. natn. Hist. nat.,

(4), 4 (A) : 261-280.

= 1982 e (D-18). — Dendrobates Wagler, 1830 and Dendrobatidae Cope, 1865 (Amphibia, Anura) :

proposed conservation. Bull. zool. Nom., 39 : 267-278.

ee 1983 a (D-19). — Classification et nomenclature supragénérique des Amphibiens Anoures. Bull.

Soc. linn. Lyon, 52 : 270-276.

= 1983 b (D-20). — Note préliminaire sur le groupe de Rana (Tomopterna) breviceps Schneider,

1799 (Amphibiens, Anoures), avec diagnose d’une sous-espèce nouvelle de Ceylan. Alyres,

2: 163-170.

1983 c (D-21). — Comments on the proposed conservation of Dendrobates Wagler, 1830 and

Dendrobatidae Cope, 1865. Z.N. (S.) 1930. (2). Bull. zool. Nom., 40 : 198-199.

= 1984 a (D-22). — Miscellanea nomenclatorica batrachologica (1). Alytes, 3 : 39-43.

-— 1984 b (D-23). — Miscellanea nomenclatorica batrachologica (III). Alytes, 3 : 85-89.

- 1984 c (D-24). — Miscellanea nomenclatorica batrachologica (IV). Alyres, 3 : 103-110.

1984 d (D-25). — Miscellanea nomenclatorica batrachologica (V). Alyres, 3 : 111-116.

- 1984 e (D-26). — La nomenclature supragénérique des Amphibiens Anoures. Mém. Mus. natn.

Hist. nat., (A), 131 : 1-64.

= 1984 F(D-27). — Notes sur les Grenouilles brunes (groupe de Rana temporaria Linné, 1758).

III. Un critère méconnu pour distinguer Rana dalmatina de Rana temporaria. Alytes, 3

117-124.

Source : MNHN, Paris

DUBOIS 147

-— 1984 g (D-28). — Note préliminaire sur le groupe de Rana limnocharis Gravenhorst, 1829

(Amphibiens, Anoures). Alytes, 3 : 143-159.

-— 1985 a (D-29). — Miscellanea nomenclatorica batrachologica (VII). Alytes, 4 : 61-78.

— 1985 b (D-30). — Le genre en zoologie : essai de systématique théorique. Thèse, Montpellier :

i-xiv+ 1-167.

—— 1986 a (D-31). — Miscellanea nomenclatorica batrachologica (VIII). Alytes, 4 : 94-96.

— 1986 b (D-32). — Commentaire sur Caeciliidae chez les Amphibiens et chez les Psocoptères :

nouveaux éléments et nouvelle proposition. Z.N. (S.) 2333. Bull. zool. Nom., 43 : 6.

Du 1986 c (D-33). — A propos de l'emploi controversé du terme “monophylétique” : nouvelles

propositions. Bull. Soc. linn. Lyon, 55 : 248-254.

-— 1987 a (D-34). — Miscellanea nomenclatorica batrachologica (XI). Alyres, 4 : 155-156.

-— 1987 b (D-35). — Miscellanea taxinomica batrachologica (1). Alyres, 5 :7-95.

- unpublished (D-36). — Strongylopus Tschudi, 1838 (Amphibia, Anura) : request for the desi-

gnation under the plenary powers of a type-species in agreement with current usage. Sub-

mitted on 8 September 1980 to Bull. zool. Nom.

2 unpublished (D-37). — Leptobrachium parvum Boulenger, 1893 (Amphibia, Anura) : proposed

conservation. Submitted on 27 May 1981 to Bull. zool. Nom.

ss unpublished (D-38). — Hyla reinwardtii Schlegel, 1840 (?) (Amphibia, Anura) : proposed con-

servation. Submitted on 27 May 1981 to Bull. zool. Nom.

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Source : MNHN, Paris

Alytes, 1986, 5 (3): 150.

Miscellanea nomenclatorica batrachologica (XIII)

Alain DUBOIS

Laboratoire des Reptiles et Amphibiens,

Muséum national d'Histoire naturelle,

25 rue Cuvier, 75005 Paris, France

A replacement name is proposed for Rana muta Su & Li, 1986, a primary

homonym of Rana muta Laurenti, 1768.

Su & Li (1986 : 152) ont récemment décrit sous le nom de Rana muta une espèce

nouvelle du groupe de Rana (Paa) yunnanensis Anderson, 1878 (sensu DUBOIS, 1987).

Ce nom ne peut être conservé pour cette espèce, étant un homonyme primaire du nom

Rana muta Laurenti, 1768, actuellement considéré comme un synonyme de Rana tempo-

raria Linné, 1758. Nous proposons le nom de remplacement suivant pour cette espèce,

que nous dédions à la mémoire du Professeur Cheng Chao LIU de Chengdu :

Rana (Paa) liui nom. nov.

Nomen novum pro Rana muta Su & Li, 1986 (nec Rana muta Laurenti, 1768).

RÉFÉRENCES BIBLIOGRAPHIQUES

Dugois, A., 1987. — Miscellanea taxinomica batrachologica (1). Alytes, 5 : 7-95.

Su, C. & Li, S., 1986. — À new anuran species from Yunnan - Rana muta. Acta herpet. sin., 5 :

152-154.

Source : MNHN, Paris

151

NOTES DE LECTURE

Alpes, 1986, 5 (3): 151-152.

Un livre sur les Amphibiens d'Australie occidentale

Jean-Louis AMIET

Laboratoire de Zoologie, Faculté des Sciences,

Université de Yaoundé, B.P. 812, Yaoundé, Cameroun

TYLER, SMITH & JOHNSTONE’s book on frogs of Western Australia is reviewed.

TYLER, M. J., SMITH, L. A. & JOHNSTONE, R. E., 1984. — Frogs of Western Australia.

Perth, Western Australian Museum : i-xii + 1-109. Prix : 16.00 $.

Pour le batrachologue les régions australes partagent le double intérêt de posséder à la fois

des peuplements d’Anoures très originaux et des spécialistes attachés à en faire connaître les riches-

ses à travers des Faunes, des Catalogues et des Guides. Ainsi, en Afrique du Sud comme en Austra-

lie, il existe à présent des ouvrages sur les Anoures qui s’adressent non seulement aux batracholo-

gues confirmés mais visent aussi, par leur présentation et leur illustration, un public d'amateurs

désireux de mieux connaître la faune de leur pays. Le fait que de tels ouvrages, coûteux en raison

de leur iconographie, puissent cependant être édités montre combien le “marché” représenté par

les “batrachophiles” doit être important dans ces régions : on peut regretter que le phénomène ne

revête pas la même ampleur chez nous.

“Frogs of Western Australia”, paru en 1984, est le cinquième ouvrage sur les Anoures d’Aus-

tralie, ce qui confirme notre propos. Il est dû à la collaboration de M. J. TYLER, un des Maîtres

de la batrachologie australienne, de L. À. SMITH et de R. E. JOHNSTONE. Couvrant la totalité de

la faune amphibienne de la moitié occidentale du continent australien, il comprend 171 pages et

12 planches réunissant 72 photos en couleurs, auxquelles s’ajoutent 18 dessins d'excellente facture

et 49 cartes de répartition.

Dans un premier chapitre, les auteurs comparent la faune de l'Australie occidentale à celle

de l’ensemble du continent : 70 espèces sur les 177 actuellement répertoriées, mais une quarantaine

sont propres au territoire considéré.

Un second chapitres permet de définir les différents termes descriptifs employés en Morpho-

logie des Anoures ; l'utilisateur non spécialiste verra sa tâche considérablement simplifiée grâce à

la qualité des dessins au trait.

Deux courts chapitres donnent ensuite quelques précisions sur les œufs, les pontes et les têtards.

Le début de la partie proprement faunistique est marqué non pas par une mais par trois clés

de détermination. Nul n’ignore combien il est difficile d'élaborer de telles clés quand elles concer-

Source : MNHN, Paris

152 ALYTES 5 (3)

nent des régions à batrachofaune riche, où certains genres peuvent compter plus d’une dizaine d’espè-

ces. Les auteurs n’ont pas refusé l'obstacle, mais en ont diminué l'importance en proposant des

clés distinctes pour chacun des territoires biogéographiques reconnus en Australie occidentale. Il

est certain que cette méthode facilite l'établissement des clés en restreignant le nombre des espèces

qui doivent y être incorporées — avantage particulièrement sensible pour les genres multispécifi-

ques dont certaines espèces, allopatriques, ne risquent pas d’être trouvées ensemble. Nous doutons

toutefois que ce procédé puisse être appliqué sans inconvénient à d’autres parties du globe, comme

l’Afrique centrale par exemple, où les zones biogéographiques montrent d’amples gradients inter-

médiaires, quand elles ne s’interpénètrent pas.

Lorsqu'il tente de construire des clés dichotomiques, le batrachologue se heurte à une autre

difficulté : trouver de bons caractères diagnostiques morphologiques, tâche épineuse dans un groupe

où les espèces “jumelles” ou “sub-jumelles” ne sont pas rares. Ici, les auteurs ont préféré contour-

ner l'obstacle, en introduisant dans leurs clés des caractères biologiques. C’est ainsi que le lecteur

a le choix (p. 19), pour séparer deux groupes d'espèces de Ranidella, entre “Call resembles a squelch”

et “Call does not resembles a squelch”. Un pareil critère sera sûrement apprécié du naturaliste

de terrain — auquel s’adresse surtout cet ouvrage — encore qu’une femelle ou un mâle en dehors

de la période de reproduction puissent le laisser perplexe, mais l'identification au laboratoire posera

un problème insoluble.

De nombreuses photos en couleurs complétées par de courtes descriptions axées sur les carac-

tères discriminants permettront cependant à l’utilisateur d’arriver à ses fins. Pour chaque espèce,

l'exposé suit un plan standard comprenant les points suivants : description, distribution, appel nuptial,

reproduction, habitat, auxquels s’ajoutent des remarques de nature diverse. La partie traitant des

espèces comprend 75 pages. Une bibliographie de 70 références termine l’ouvrage.

Grâce aux photos, les amis des Batraciens y découvriront une galerie de portraits particuliè-

rement dépaysante en raison du faciès étrange de certaines espèces. L’écologiste y trouvera quel-

ques beaux exemples d'adaptation à la vie fouisseuse, fréquente dans une faune riche en espèces

déserticoles. Quant aux batrachologues, ceux d’entre eux qui ont l’expérience de la faune africaine

ne manqueront pas d'admirer certains cas de convergence morphologique : la palme revient ici au

genre Litoria, où des espèces ressemblent, même dans leurs attitudes, à des Hyperolius (L. bicolor,

L. microbelos) ou à des Lepropelis (L. rothit), similitude concevable pour des “‘rainettes”, mais aussi,

ce qui est plus extraordinaire, à des Prychadena (L. nasuta, L. tornieri).

1

FAR

* /

Source : MNHN, Paris

ALVTES

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), Michael J.TYLER (Adelaide), Richard J.

WASSERSUG (Halifax).

Recommandations aux auteurs. — Alytes publie des articles originaux en français ou en anglais, consa-

crés 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 fran-

ais (pour ceux qui le souhaiteraient, les rédacteurs acceptent de revoir les résumés en fran-

çais à partir d’un texte en anglais). Tableaux et figures doivent comporter un titre. Les figu-

res, 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’ A/ytes 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. Manuscrits 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

editors 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 Alyres (book references should include the pagination). Send the manuscripts in triplicate

to the editors (address above). Acceptance for publication will be decided by the editors fol-

lowing review by two referees or more.

Türés à part. — 25 exemplaires gratuits par article. Au-delà, les tirés à part seront facturés par tran-

ches de 25 exemplaires.

Publié avec le concours du Muséum national d'Histoire naturelle.

Directeur de la Publication : Alain DUBOIS.

Numéro de Commission Paritaire : 64851.

Source : MNHN, Paris

a

Abjtes, 1986, 5 (3) : 97-152. 47 SEP, 1901

SOMMAIRE

Alain DUBOIS

Miscellanea nomenclatorica batrachologica (XII)

Alain DUBOIS

Living amphibians of the world : a first step towards a comprehen-

sive checklist

Alain DUBOIS

Miscellanea nomenclatorica batrachologica (XIII) ............

NOTES DE LECTURE

Jean-Louis AMIET

Un livre sur les Amphibiens d'Australie occidentale ..........

Photocomposition/Photogravure : Alexandre, Paris. 42 46 17 57.

Imprimé aux Ateliers de la Couronnerie, 45750 Saint-Pryvé Saint-Mesmin, France.

Dépôt légal : 2 trimestre 1987.

97

99

150

151

Source : MNHN, Paris: