J. South Asian Nat. Hist., ISSN 1022-0828. May, 1999. Vol.4, No. 1, pp. 1-11. © Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka.

INVITED EDITORIAL / BOOK REVIEW South Asian Amphibia: a new frontier for taxonomists

Alain Dubois*

* Laboratoire des Reptiles et Amphibiens, Museum national d'Histoire naturelle, 25 rue Cuvier, 75005 Paris, France.

Books reviewed

Dutta, Sushil K. 1997. Amphibians of India and Sri Lanka (Checklist and bibliography). Bhubaneswar, India, Odyssey Publishing House, [i- iii] + i-xiii + 1-342 + i-xxii. [Cited below as AISL].

Dutta, Sushil K. & Manamendra-Arachchi, Kelum. 1996. The amphib­ ian fauna of Sri Lanka. Colombo, Sri Lanka, Wildlife Heritage Trust of Sri Lanka, 1-232. [Cited below as AFSL].

South Asia (here understood in a biogeographical were among the first published monographs deal­ sense, i.e. from Pakistan to Myanmar and from Ne­ ing with the amphibians of a tropical region of the pal to Sri Lanka) is a fascinating area for all evolu­ world. tionary biologists: at the meeting of several major However, after the important works of N. biogeographic zones, it harbours composite faunas Annandale and G. A. Boulenger at the beginning of and floras of various origins and affinities, that oc­ our century, which culminated in Boulenger's (1920) cupy a wealth of habitat types at all altitudes from still unequalled review of Asian ranids, taxonomic sea level to snow level in the Himalayas. This tropi­ study of the amphibians of this region slowed down cal region has been an important "natural labora­ considerably. For about 50 years, most authors tory" where many variants of the "experience of evo­ merely copied Boulenger's taxonomy, which some lution" have developed under a variety of different of them seemed to consider the "final word". An in­ conditions, thus offering an infinity of possible sub­ teresting exception was Kirtisinghe's (1957) book on jects of interest for scientific research. the amphibians of Ceylon, which was based on origi­ Concerning the group of amphibians, this region nal observations and a personal knowledge of these was one of the first tropical areas of the world where animals both in the field and in collections (although taxonomic research was undertaken, mostly under he did not examine or refer to type material). Be­ the leadership of British zoologists. From the begin­ sides this, a few species were described in isolation, ning to the end of the 19th century, descriptive works but no comprehensive revision of a South Asian by various authors, including J. Anderson, R. H. amphibian group was published during this period. Beddome, E. Blyth, A.-M.-C. Dumeril & G. Bibron, In 1970-1973, A. Dubois and collaborators col­ T. C. Jerdon, R. P. Lesson, W. L. Sclater, F. Stoliczka lected more than 15,000 amphibian specimens in and W. Theobald, gave zoologists the first rough idea most parts of Nepal, probably one of the largest of the diversity of amphibian species in "British In­ amphibian collections from South Asia ever gath­ dia". This led to the writing of two books (Gunther, ered. Instead of using these data merely to write a 1864; Boulenger, 1890) presenting a complete survey checklist or a field guide to the amphibians of Ne­ of the then-available information in this field; these pal, I considered that this material should be used D ubois

for the completion of revisions of groups taken one long "sleeping" subject is very promising, but here by one, as problems were evident in all the amphib­ some words of warning are in order. At the verge of ian taxa of Nepal (Dubois, 1974,1975a-b, 1976,1980, the 21th century, taxonomy can no longer be consid­ 1983,1984b, 1987a, in press). To be serious and mean­ ered a young science, and what was permissible at ingful, such revisions should not only take into ac­ the end of the 19th century is no longer so. It was count all available specimens from Nepal and the then usual to describe a "new species" without ex­ immediately neighbouring countries (India and Ti­ tensive comparison with related species, and in par­ bet), but should also include re-examination of the ticular without actual comparison with specimens status of all related taxa from South and South-East in other collections: evaluation of a specimen against Asia, including Indochina and China, which makes written descriptions was then believed to be suffi­ such a project formidable. In particular, owing to the cient to decide if the same or a different taxon was long lack of communication between batrachologists involved. Nowadays, any serious alpha-taxonomic in China and in other countries, "parallel" work must rely not only upon the study of descrip­ taxonomies have tended to develop north and south tions and other bibliographic data, but also of of the southern border of China. Such a situation is voucher specimens of related species, particularly of quite unsatisfactory, as it does not allow taxonomic, type-specimens of all concerned nominal species. phylogenetic and biogeographic generalizations to The taxonomy of the amphibian fauna of the Ori­ be made: recent opening of scientific communication ental Region, and particularly of South Asia, is still with Chinese zoologists will allow the homogeniza­ very unsatisfactory, especially when compared with tion of the taxonomy of amphibians in all of Eastern that of most other regions of the world. Many spe- Asia, but this will still require time, since travel, loans cies-groups or even genera of this region have not and examination of specimens will be necessary to been the subject of recent revisions, and the solve many doubts and problems. Because of these nomenclatural status of many names, including very difficulties, up to now only the revision of the "common" ones, for which no types are identified Himalayan ranids now referred to the genera and no recent descriptions available, is unclear: in Chaparana and Paa could be completed (Dubois, such circumstances, adding further new names to 1975a, 1976, 1979,1980,1992; Dubois & Khan, 1980; the already existing ones, without proper compari­ Dubois & Matsui, 1983), and work on the other sons, will only complicate further the work of fu­ groups is still in progress. Despite this, progress in ture serious workers, who will have to take all these our knowledge of the Nepalese fauna has been enor­ new names into account in their revisions, to look at mous: while Swan & Leviton's (1962) checklist re­ their type-specimens and to deal with the resulting ported about 16 amphibian species in this country, nomenclatural problems. A further problem is posed Dubois's (in press) checklist, submitted for publica­ by authors not preserving type-specimens in a way tion in 1997, mentioned 42 species and subspecies, that will ensure their long-term survival, i.e. in com­ to which, according to Anders et al. (1998), petent institutions as defined by the Code (Anony­ Rhacophorus (Polypedates) taeniatus Boulenger, 1906 mous, 1985: Recommendations 72.D and 72.G, Arti­ should be added: this represents an increase by 169 cle 75.d.6), and publishing their descriptions in jour­ % over 35 years (43 against 16), including 11 species nals which do not practice a high quality of peer re­ first described after 1962 (26 % of the total fauna). view, or too often, no peer review at all. Editors of An important finding of this research is that, for 43 zoological journals would perform a much appreci­ valid names, this checklist reports about 50 syno­ ated service to the scientific community if they refuse nyms (54 % of the total number of 93 names consid­ to publish such isolated descriptions of so-called ered altogether), so that it is clear that even in a "new species" in the absence of comprehensive di­ poorly-explored country with a little-studied fauna, agnosis against all available possible subjective syno­ systematic and nomenclatural work must be carried nyms through the examination of type-specimens: out carefully if one wishes to build up a serious and in most cases, descriptions of new species is war­ useful taxonomy. ranted only in groups which have been subject to However, the finding that Boulenger's taxonomy recent revision. Rather than publish such papers, edi­ was not the "final" one, even for the ranids, had tors should encourage the authors to carry out com­ important consequences, as it seems to have "freed" prehensive nomenclatural and taxonomic revisions, some taxonomists from the dreadful respect they had to examine and describe old type-specimens, to des­ shown for this taxonomy for more than half a cen­ ignate and describe neotypes when necessary, and tury: this resulted in the recent description of a to review critically the species-group taxa: only when number of "new species" of Indian frogs by several this is done will it be meaningful to describe new authors in various journals. This reborn interest in a species.

2 ]. South Asian Nat. Hist. S o uth A sian A m phibia

Actually, and although this is not understood by describes a specimen of Amolops as a new species of some taxonomists (and has led to some serious mis­ Rana or, more "precisely", of Rana (Sylvirana), a understandings on the nature of zoological nomen­ young Bufo as a new species of Ansonia, or, even clature: see e.g. De Queiroz & Gauthier, 1994), allo­ worse, a specimen of Paa as a new species of Scutiger cation of zoological names to taxa is not at all based (none of these examples is imaginary), one will find on definitions, diagnoses or descriptions, but on the many distinctive characters that will purportedly allocation to the taxa of name-bearing type specimens justify the creation of a new nominal species! As the or onomatophores, which constitute an objective, ma­ Code (Anonymous, 1985: Article 13) only requires, terial connection between the real world of animal for the nomenclatural availability of a name, that it populations and the world of language (for more be "accompanied by a description or definition", but details, see Dubois & Ohler, 1997). Any zoological not that the description be correct or complete (for rea­ taxon name therefore rests on its actual or potential sons explained in detail by Dubois & Ohler, 1997), onomatophore, and creation of new names should the new name, if published, will be available for the never been made before the nomenclatural status of taxon represented by its type-specimen, but it will all existing names of the same taxonomic group has remain concealed in the wrong genus until, perhaps been ascertained. much later, a subsequent author re-examines the Before describing any "new species", the sup­ specimen and correctly allocates the name: of course, posed new taxon must be tentatively assigned to a in many cases this is liable to result in disturbing genus, subgenus or species-group; then, it must be consequences on the then-current nomenclature. compared with type-specimens of all or most nomi­ It is difficult to avoid the suspicion that authors nal taxa already recognized in the group, including in search of quick and easy publications, nowadays those of current junior subjective synonyms, in or­ necessary for progress through institutional aca­ der to ascertain whether a name may not be already demic hierarchies, resort to "single species" publi­ available for the taxon. Failure to do so entails the cations as an easy means of augmenting their publi­ risk of creating unnecessary new names, that will cations lists. Such hasty and superficial publications ultimately have to be invalidated as junior synonyms are likely to cause more problems to future taxono­ through much unnecessary work by later taxono­ mists than actually aiding the progress of taxonomy, mists (for recent examples in the amphibians, see e.g.: especially as type-specimens of the "new species", Dubois & Ohler, 1995,1998; Dubois, 1995,1998b). As sometimes deposited in inaccessible and often tem­ most South Asian amphibian groups have not been porary local collections, will all have to be examined properly revised since Boulenger, any description of to ascertain the status of the new names. As a referee a new frog species from this region should first be frequently solicited by various journals to review preceded by a revision of the group concerned, and papers dealing with Asian or Oriental frog taxonomy, all possible efforts should be made to clarify the I have encountered many surprising manuscripts nomenclatural status of all other names available in and often had occasion to point out these problems this group. Of course, such a study takes longer to to the concerned authors and editors, but with only complete and is more involved and expensive than limited effect. Interestingly, several recent authors merely "describing" isolated "new species", but this and editors provided a "philosophical" justification is the only way to produce work that is useful in the for rejecting my advice: they seemed to consider long term. This process has been followed in recent these comments as constituting a "limitation of the decades by almost all authors who have undertaken freedom of taxonomists". They also pointed out that, the systematic re-evaluation of amphibian groups as South Asian scientists find it difficult (for finan­ that had not for long been revised in various regions cial reasons) to travel, and as many institutions refuse of the world (see e.g. Duellman, 1968, 1971; Perret, to send type specimens for study in South Asia, re­ 1976; Lavilla, 1988). quiring these zoologists to examine types would be Unfortunately, this is not the process that has been a way of impeding them in their study of their own followed by several recent workers on the frogs from fauna — a kind of "colonialist" attitude that would the South Asian region. This has resulted in mistakes, result in allowing only scientists from developed such as the creation of synonyms or the placement countries to study this fauna: in consequence, it was of species in wrong genera, and therefore based on suggested that South Asian scientists should not be irrelevant and incorrect diagnoses: several of these required to adopt the same "standards" for modern errors were briefly mentioned by Dubois (in press), taxonomic research as are currently required of say and others will be cited elsewhere. Particularly per­ European or American taxonomists. I take strong nicious are descriptions of "new species" that rely exception to this view, which I consider, in fact, to be on an incorrect generic allocation: of course, if one the real "colonialist" way of tackling this problem,

Vol. 4, No. 1. 3 D ubois

and according to which there should be two kinds knowledge available as at July 1996 on the amphib­ of taxonomies: a "local", "second-class" one, based ians of Sri Lanka. It recognizes 53 valid species dis­ purely on local surveys, description and naming of tributed in five families. Efforts have been made to nominal species without proper comparisons with trace and examine as many type-specimens of nomi­ already existing type material; and an "interna­ nal species as possible, and photographs of many of tional", "first-class" level, where scientists from rich them are provided. For each family, a key to the Sri countries, having the possibility to travel, visit mu­ Lankan species is given. Then, for each species, the seums, obtain specimens in loan, and use modern following information is provided: a genuine syn­ methods of analysis, would propose revisionary onymy (i.e., containing only once each different name works, phylogenetic analyses, comprehensive once given to the species, not the various "uses" or taxonomies and evolutionary and biogeographic "new combinations" of these names); a diagnosis; a syntheses. On the contrary, I believe taxonomists in description; data on colour in life; comments; data this region, like in all other countries in the world, on distribution; etymology of valid specific name of should be given every opportunity to work accord­ species (but, curiously, not of synonyms); and a list ing to modern standards of taxonomic research, but of material examined. A useful companion to this should also be subjected to the same stringent rules volume is the booklet by Anslem De Silva (1996), as others for publication of their results: this is the which provides a list of 248 references of publica­ only way these taxonomists could enter the world tions dealing with the amphibians of Sri Lanka. of professional taxonomy at a level of full equality The second book, AISL, by Sushil K. Dutta, was with their colleagues everywhere, which will never published in India. It contains no illustrations or be the case if "the bar is lowered" for them, as was maps, and consists of two roughly equal parts: a 156- clearly the case in the recent publication by some so- page checklist of the 216 amphibian species recog­ called major international journals of papers from nized as valid by this author in India and Sri Lanka; South Asia that would never have been accepted had and a 168-page bibliography of works dealing with they come from America or Europe. Of course, the the amphibians of these two countries. This enor­ financial and material limitations cited above are real, mous bibliography will certainly prove useful in the but then it should be the duty of the colleagues work­ future, although the fact that taxonomic papers are ing in developed countries to help South Asian col­ intermingled with other kinds of papers (e.g., deal­ leagues address them. This can be done in many ing with parasitology, physiology, developmental ways, for example by inviting and funding these biology), some of which merely use Indian amphib­ colleagues to conduct collaborative research in their ians as research material but show no genuine inter­ own institutions and through sending information est for these animals by themselves, will complicate (including descriptions, drawings or photographs) its use. As for the taxonomic checklist, for each spe­ on type-specimens. Besides, South Asian zoologists cies, the following information is provided: a chreso- would be greatly aided if they developed the prac­ synonymy, i.e. a list of genuine synonyms and of tice of submitting their manuscripts to colleagues for their various uses and combinations (see Dubois, "friendly review" before submitting them formally 1997b: 184-185); data on distribution; a list of refer­ to periodicals for publication. Such a collaborative ences; and also, but for some species only, system­ attitude between colleagues from different parts of atic notes and a list of specimens examined. Here the world would seem to be a better and sounder also, attempts have been made to include as much way of developing batrachology in this region and information as possible about name-bearing types improving its standards, rather than through the con­ of the nominal species, considered either valid or temptuous practice of setting different rules for au­ invalid (synonyms and homonyms), listed in the thors according to their origin. book. In this respect, this checklist is more complete The two books reviewed here auger well for the and useful than Frost's (1985) checklist, a major future (see above for the abbreviations of their ti­ weakness of which was the absence of data on syno­ tles). The first published one, AFSL, is the result of a nyms and subspecies (see Dubois, 1987fc-c). collaborative work between Sushil K. Dutta of India Both these books are useful contributions to the and Kelum Manamendra-Arachchi of Sri Lanka. batrachology of India and Sri Lanka. Their authors Published in Sri Lanka, this is a nicely printed, care­ clearly made a real effort to provide a reasonably fully edited and produced book, with numerous col­ complete coverage of the published information, our and black-and-white photographs (including the thus giving a picture of the "state of the art" regard­ first-ever published photographs in life of many Sri ing the taxonomy of the amphibians of this region. Lankan amphibian species), good drawings and dis­ Quite correctly, the authors of both books considered tribution maps. This book presents a review of the this review of existing data, including search for type-

4 J. South Asian Nat. Hist. S outh A sian A m phibia specimens, as a necessary prerequisite for any genu­ cies Polypedates microtympanum Gunther, 1859 and ine revision work dealing with these groups. It is other closely related species from Sri Lanka striking that none of these two books contains any (Polypedates cavirostris Gunther, 1869; Rhacophorus new name, although in one of the books (AFSL) sev­ fergusonianus Ahl, 1927; Polypedates nanus Gunther, eral new taxa are recognized, but using old names 1869; Rhacophorus pleurotaenia Boulenger, 1904; previously buried in synonymies. This careful and Polypedates reticulatus Gunther, 1864) probably all prudent attitude strongly contrasts with the reckless have a peculiar reproductive biology, which should attitude criticized above, and should be taken as a be recognized taxonomically: for this purpose, the model by future workers on the taxonomy of the genus-group name Kirtixalus, proposed by Dubois amphibians of South Asia (and of other regions as (1987a) as a subgeneric name within Philautus, is well, including Europe: see Dubois, 1998b). available. Quite possibly Kirtixalus will have to be A few weaknesses may be pointed out in both raised to the genus rank, but anyhow this name, at books, that do not however diminish their interest least at subgeneric level within either Philautus or and usefulness. Rhacophorus, should be used for the group recognized The first point is that it is always hazardous to under the latter name in AFSL. More work is needed base a taxonomy on information about taxa from a on the reproductive biology of species of Kirtixalus. restricted area: taxonomic decisions should be based Direct aerial development in these frogs has been on comprehensive, whole-taxon revisions, not on described on several occasions (Gunther, 1876; geographical surveys of limited taxonomic scope (see Ferguson, 1876; Kirtisinghe, 1946,1957), but for the Dubois, 1981,1992). This can be exemplified here by time being, as stressed in AFSL (p. 201-202), it is still the taxonomic treatment of the Rhacophorinae in unclear which species were really involved: further AFSL. First of all, recent molecular data (Marmayou field work on these frogs is badly needed (see also et al., in preparation) strongly suggest that this group, Bossuyt & Dubois, in preparation). if holophyletic (which is not certain), is a member As for the distinction between the genera of the Ranidae and should at best be treated as a Rhacophorus and Polypedates, proposed by Liem subfamily of the latter, as suggested by Laurent (1970), it is tenuous, and was not accepted by all (1951, 1986), rather than as a family of its own. authors (see e.g. Dubois, 1987a). Fixation of a stable Within this group, AFSL recognizes four distinct status for these names will have to await a compre­ genera in Sri Lanka: Philautus Gistel, 1848; hensive generic revision of the Rhacophorinae (a very Polypedates Tschudi, 1838; Rhacophorus Kuhl & Van heavy task indeed), but, for the time being, a con­ Hasselt, 1822; and Theloderma Tschudi, 1838. Treat­ servative attitude might be to treat Polypedates as a ment of Polypedates as a genus distinct from subgenus of Rhacophorus (see e.g. Dubois, in press). Rhacophorus is based on the following rationale: "Sri Finally, within the nominative subgenus of the ge­ Lankan Polypedates and Rhacophorus differ in the size nus Philautus, several species-groups were defined of their vomerine teeth and body coloration, but by Dring (1987), and it would have been useful to otherwise are morphologically similar. The only study whether the Sri Lankan species belong in one striking difference between these two genera in Sri or several of these groups or should be placed in dif­ Lanka concerns their reproductive biology: all Sri ferent groups altogether. In particular, the nominal Lankan Rhacophorus have direct development on species Ixalus temporalis Gunther, 1864 and related land, whereas Polypedates lays eggs in foam nests species (Ixalus leucorhinus Lichtenstein, Weinland & and have aquatic larvae." (AFSL: 140). Such a tax­ Von Martens, 1856, Ixalus hypomelas Gunther, 1876 onomy makes sense in Sri Lanka where it is clear and Ixalus nasutus Gunther, 1869 from Sri Lanka, as that these two groups of frogs should be placed in well as Phyllomedusa wynaadensis Jerdon, 1853 from different genera, but the nomenclatural treatment India) share several external morphological charac­ used is not correct. What seems true of Sri Lankan teristics suggesting that they might belong in a par­ "Rhacophorus" (direct development on land) does ticular group, for which the genus-group name not apply to all the other numerous species currently Pseudophilautus Laurent, 1943 would be available allocated to the genus Rhacophorus (see e.g. Dubois, should this group need to be given a separate iden­ 1987a), and in particular to its type-species tity (see Bossuyt & Dubois, in preparation). Rhacophorus moschatus Kuhl & Van Hasselt, 1822, A refreshing feature of AFSL is that it recognizes currently known as Rhacophorus reinwardtii Kuhl & several new taxa but resurrects for them available Van Hasselt, 1822 (see Dubois, 1982), which is names which had until then been considered as jun­ known to lay eggs in a foam nest and to have aquatic ior subjective synonyms. This attitude is much more larvae (Siedlecki, 1909; Van Kampen, 1909, 1923). "professional" than that of many recent taxonomists On the other hand, it is clear that the nominal spe­ who coined unwarranted new names in similar situ­

Vol. 4, No. 1. 5 D ubois

ations (see references above and below). However, through these neotype designations, for these names in order to be beyond criticism, resurrection of old to disappear as junior synonyms of older names. names must be made with some precautions, in par- Actually, as will be shown elsewhere (Bossuyt & ticular regarding possible first-reviser actions carried Dubois, in preparation), Rao's (1937) names must be out by previous authors in the past. Thus, AFSL res­ used with the greatest caution, as careful examina­ urrects the name Ixalus macropus Gunther, 1869 from tion of his descriptions and figures very clearly show the synonymy of the nominal species Polypedates that most of his species are not correctly allocated to microtympanum Gunther, 1859 where it had been genera: thus, over a total of eight species described placed (Wolf, 1936:173; Gorham, 1974:170). Besides, as new Philautus in this paper, only three are likely AFSL also considers the names Polypedates nanus to have been genuine members of Philautus, while Gunther, 1869 and Ixalus sarasinorum Muller, 1887 to four were probably members of the genus Micrixalus apply to the same species. While choosing the name and one a member of the genus Indirana. In such a Ixalus macropus as the valid one for the species, AFSL situation, the greatest care should be taken to avoid ignored the first-reviser action of Boulenger (1882: these names playing a disturbing nomenclatural role 81), who also considered the names I. macropus and in the future, at least until neotypes have been des­ P. nanus to be synonyms, and afforded priority ignated for them. Quite luckily, two of these names among them to the latter. If Kirtixalus is recognized had already been "eliminated by themselves", be­ as a subgenus of Philautus, the valid specific name ing junior primary homonyms in the genus Philautus, of this species is therefore Philautus (Kirtixalus) nanus but, most unfortunately, Dutta (1985) proposed new (Gunther, 1869). replacement names for them, without trying to at­ Despite a clear effort to do so, AFSL does not pro­ tach them to actual specimens: such thoughtless prac­ vide a complete review of the taxonomic literature tices cannot but add new nomenclatural problems concerning Sri Lankan frogs, nor a complete list of to the already existing ones. Great caution should be names available for these frogs. At least the follow­ exerted before using any of the names of Rao (1937) ing names, based on type-specimens from Sri Lanka, mentioned as valid in AISL: thus for example the are not addressed in this book, and possibly others nominal species Rana parambikulamana Rao, 1937 is are lacking as well: Ixalus poecilopleurus Lichtenstein, not a Tomopterna (or a Sphaerotheca), but a Fejervarya, Weinland & Von Martens, 1856; Ixalus adspersus in the same way as Pyxicephalus rufescens Jerdon, 1853 Gunther, 1872; Ixalus oxyrhynchus Gunther, 1872; (see Dubois, 1984b). Ixalus semiruber Annandale, 1913; Rhacophorus rugatus Several other problems linked to improper study Ahl, 1927; Rhacophorus malcolmsmithi Ahl, 1927. The of the status of old names can be found in AISL. As status of these names will be discussed in a forth­ an example, another irritating case must be discussed coming paper (Bossuyt & Dubois, in preparation). here once again, as it continuously reappears, al­ The scope of AISL being wider than that of AFSL, though it was solved some 80 years ago: this is the it is not surprising if this book should contain more problem raised by the name Ixalus argus Annandale, mistakes of the same kind, a few of which are men­ 1912, a subjective synonym (just like the name tioned below. Polypedates afghanus Gunther, 1859) of Amolops A major problem of Indian's batrachology is Rao's marmoratus (Blyth, 1855) (see Boulenger, 1920; (1937) paper, where this author described and fig­ Dubois, 1974, 1992; Chanda & Sarkar, 1997). With­ ured 20 new frog species and subspecies from south­ out explanation, Inger (in Frost, 1985: 527), Chanda ern India. Most descriptions were based on holotypes & Ghosh (1988, 1989) and Dutta (AISL: 74) recog­ alone, generally in poor condition (desiccated, hav­ nized a species Philautus argus (Annandale, 1912), ing probably not been properly fixed in formalin), probably by simple ignorance of the synonymy as can be seen from the drawings of the specimens. above. Even more strangely, Bourret (1942: 390,451) Dubois (1984b: 157) reported that all these specimens, and Gorham (1974:127,166) cited this nominal spe­ which had been deposited in the Central College in cies twice, first as a Staurois or as an Amolops and Bangalore, had been lost and were no longer avail­ second as a Philautus. Before discussing this prob­ able for study. He suggested that, to definitely re­ lem in some detail, a few preliminary comments are solve the nomenclatural problems posed by Rao's justified. (1937) names, neotypes should be designated for the Science is an international activity, but, unfortu­ nominal taxa concerned, which requires to have nately, several languages are spoken in the world and specimens coming from a locality as close as possi­ scientists from different countries use them to com­ ble to the original type-locality, and corresponding municate. Two centuries ago, at the beginning of reasonably well with the original descriptions; the western biological science, universality of commu­ best solution in several cases will be to manage, nication between scientists was maintained by the

6 J. South Asian Nat. Hist. So uth A sian A m phibia use of Latin as the scientific lingua franca. In the 19th another case. Chanda & Sarkar (1997) devoted a century, scientific publications were written in a va­ whole paper to the refutation of an imaginary state­ riety of European languages, mostly English, French ment wrongly attributed to me. They wrote: "Dubois and German, but also Italian, Portuguese, Russian, (1992) failed to find differences between P. Spanish and sometimes others. During the 20th cen­ shyamrupus and Ixalus argus Annandale, 1912". They tury, a clear trend has developed towards an increase compared the types of these two nominal species, in the proportion of scientific papers published in showed that they were widely different, and con­ English across the entire planet, but this does not cluded: "Relegation of P. shyamrupus to the syn­ mean that English has become the only scientific lan­ onymy of A. afghanus by Dubois (1992), without ex­ guage. In parallel with this trend, another trend is amination of the relevant types, is therefore an er­ clearly visible, that of an increase of scientific publi­ ror, and we formally revive Philautus shyamrupus cations in languages, like Chinese, Japanese, Russian Chanda and Ghosh, 1989 as a distinct species of or Spanish, that were not among the dominant sci­ rhacophorid, which is known only from the type lo­ entific languages of the 19th century (see also in this cality." However, here is a complete translation of respect Dubois, 1987b: 41). Should these publications the part of my paper (Dubois, 1992: 340) referred to be considered "local" publications which can be ig­ by these authors: "Another undeniable synonym of nored, or grossly misunderstood, by English-speak­ the name Rana afghana is Ixalus argus Annandale, ing scientists? Certainly not. It should be realized by 1912. Strangely, and without explanation, Inger (in the latter that the current trend in the increase of Frost, 1985: 527) took this name off this synonymy, English as an international scientific language gives where it was standing since the work of Boulenger scientists whose native language is English a strong (1920: 217), and created the combination Philautus advantage over all other scientists worldwide: when argus. He was followed by Chanda & Ghosh (1988, writing a scientific paper, they only have to concen­ 1989), who compared their new species Philautus trate on the effort of the scientific presentation of re­ shyamrupus to Philautus argus. In fact, the descrip­ sults and discussion, while non-native English- tion and the excellent figure of Annandale (1912) speaking scientists must produce a greater effort, leave no doubt on the synonymy proposed by both scientific and linguistic. Language corrections Boulenger (1920) and adopted since by all other au­ often imposed on them by editors, especially when thors (Bourret, 1942; Dubois, 1974; Gorham, 1974)." an American journal feels entitled to correct the Brit­ It is therefore clear that I never proposed to ish way of writing English, or the reverse, may be synonymize Philautus shyamrupus with Amolops extremely time and energy consuming, and place afghanus and that the whole "refutation" of this syn­ non-English scientists at a disadvantage when com­ onymy is completely irrelevant. pared with their native English-speaking colleagues. My intention in citing the name Philautus The minimum expectation scientists speaking other shyamrupus in this discussion, however, was a dif­ languages can have is that, in the minority of remain­ ferent one. The original description of this species is ing cases of scientific papers written in languages only one among several published in recent years other than English, English-speaking scientists make where a new amphibian species was proposed with­ a corresponding effort to understand correctly their out relevant comparisons with closely related nomi­ contents: of course, no one can be expected to read nal taxa, and without a revision of the group that fluently Chinese, French, German and Russian, but includes this species. As explained above, such iso­ when a scientist encounters a paper in a language lated descriptions of "new species" will only add to he /she cannot personally decipher, he/she should the existing taxonomic and nomenclatural confusion make the effort of asking for a translation from a bi­ and will not help to increase our knowledge of these lingual colleague or some other source. Otherwise, groups. The nominal genus Philautus Gistel, 1848 is the risk exists to have progressively on this planet a good example of this situation. More than 160 several "parallel sciences" with no or only minimal nominal species have been or are currently referred communication between them, which of course to this genus (Bossuyt & Dubois, in preparation), would have unfortunate consequences on the unity which has not been the subject of a general revision, and universality of science. although partial, local revisions have been recently Some years ago (Dubois, 1987b), I had to publish provided (Dring, 1987; Inger, 1989; Brown & Alcala, an English translation of some parts of a paper of 1994; Malkmus & Riede, 1996a-b). In such circum­ mine in French (Dubois, 1984a) which had been stances, it is neither justified to propose new replace­ grossly misunderstood, and consequently submitted ment names for preoccupied names which have not to irrelevant criticisms, by Savage (1986). Unfortu­ been shown to apply to valid species (as discussed nately, I now have to make a similar correction in above), nor to describe new species without relevant

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comparisons. Chanda & Ghosh (1989) compared meet in the future when dealing with many recently their "new species" Philautus shyamrupus only with described "new species" from South Asia. two other nominal species, namely Ixalus argus The comments above should by no means be Annandale, 1912, which is an Amolops, and Philautus taken as suggesting that I consider that South Asian cherrapunjiae Roonwal & Kripalani, 1961, a doubtful taxonomists always publish bad-quality works, species whose name is based on a just-metamor- while their colleagues from America or Europe phosed frog, but they failed to compare it with the would always produce good publications. Quite to other nominal species from the Himalayas and sur­ the contrary, and as I have already stressed it on sev­ rounding regions currently referred (although clearly eral occasions (e.g., Dubois, 1987b-d, 1995, 1998a-b; in error for several of them) to the genus Philautus Dubois & Ohler, 1995,1997), many "western" publi­ (Frost, 1985; Dubois, 1987a). More than 20 nominal cations contain mistakes, particularly in the currently species, several of which are probably synonyms, much neglected field of zoological nomenclature, but would deserve consideration in this context: also as concerns basic taxonomic procedures. A re­ Rhacophorus andersoni Ahl, 1927; Philautus cent example, concerning South Asian amphibians, albopunctatus Liu & Hu, 1962; Ixalus annandalii is Schleich & Anders's (1998) description of Boulenger, 1906; Ixalus carinensis Boulenger, 1893; Tomopterna maskeyi, a member of a group which Ixalus cinerascens Stoliczka, 1870; Rhacophorus dubius Marmayou et al. (in preparation) suggested to re­ Boulenger, 1882; Ixalus garo Boulenger, 1919; Philautus gard as a distinct genus Sphaerotheca Gunther, 1859. gracilipes Bourret, 1937; Polypedates jerdonii Gunther, The new specific name was proposed without proper 1876; Philautus jinxiuensis Hu, 1978; Ixalus kempiae study of intra- and interspecific variation (including Boulenger, 1919; Philautus longchuanensis Yang & Li, polychromatism) among South Asian species of 1979; Philautus maosonensis Bourret, 1937; Philautus Sphaerotheca, nor bioacoustic study of the mating medogensis Ye & Hu, 1984; Philautus menglaensis Kou, calls, nor critical evaluation of the status of all spe­ 1990; Rhacophorus microdiscus Annandale, 1912; cific names already available in this group (see Philautus namdaphaensis Sarkar & Sanyal, 1985; Dubois, 1983), nor examination of the type-speci- Philautus odontotarsus Ye & Fei, 1993; Ixalus parvulus mens still existing for some of these names. Dubois Boulenger, 1893; Philautus rhododiscus Liu & Hu, 1962; (in press) examined two syntypes of Rana breviceps Philautus shillongensis Pillai & Chanda, 1973; Philautus Schneider, 1799 (Zoologisch Museum Berlin 3351) tytthus Smith, 1940. Probably only part of these nomi­ and stated that they represent a small-sized species nal species are genuine members of the genus of this group, Sphaerotheca breviceps, similar to Philautus: several of them (including Philautus Sphaerotheca rolandae but possibly distinct. On the cherrapunjiae Roonwal & Kripalani, 1961) were de­ other hand, he considered that the currently avail­ scribed as having a free tadpole stage, which ex­ able information is for the time being insufficient to cludes them from this genus, at least as defined by support the existence of more than one species of Dubois (1987a). Chanda & Sarkar (1997) just repeated large-sized Sphaerotheca in South Asia, for which the the irrelevant comparison with Ixalus argus and pro­ following names are available: Rana variegata vided no new information to support the validity of Gravenhorst, 1829 (nec Linnaeus, 1758); Pyxicephalus Philautus shyamrupus Chanda and Ghosh, 1989, fodiens Jerdon, 1853; Pyxicephalus pluvialis Jerdon, which remains therefore unsupported by scientific 1853; Sphaerotheca strigata Gunther, 1859; Rana arguments. Unfortunately, AISL once more repeats dobsonii Boulenger, 1882; Tomopterna strachani this meaningless statement, as it writes again that Murray, 1884; Rana leucorhynchus Rao, 1937; Rana Philautus shyamrupus is "close to P. argus and P. swani Myers & Leviton, 1956. Acting as first-reviser, aurifasciatus" (AISL: 86), which has little meaning as he chose Sphaerotheca pluvialis as the valid name of the former is an Amolops and the latter a Philautus. this species. Pending a serious revision of this group, In fact, several characters in the descriptive notes of the name Tomopterna maskeyi will just have to provi­ Chanda & Ghosh (1989) and in the photograph of sionally join the other seven names above in the syn­ Fig. 1 of Chanda & Sarkar (1997) are incompatible onymy of this species. Redescriptions of type-speci­ with the genus Philautus, such as the presence of lon­ mens of this group will soon be available (Dubois & gitudinal bands or lines on its body or flanks, its Ohler, in preparation), but a complete revision of the smooth chest and belly, its large webbing, and its taxonomy of these variable frogs will not be possi­ lack of both inner and outer metatarsal tubercles. ble until field data, particularly on mating calls, and Final stabilization of the status of this name will re­ detailed comparisons of large series of specimens quire re-examination of the holotype and relevant from various localities, can be completed. Taxonomy, comparisons with truly related species. This is just being an ancient scientific discipline, is sometimes one example of the kind of situation we will often seen by some as old-fashioned and outdated. Some

8 J. South Asian Nat. Hist. S o uth A sian A mphibia researchers in other fields of biology find it difficult 1983; Dubois, 1987a) and some others (e.g., femoral to understand the need of our work of inventory, glands in males) reminiscent of some groups of description and classification of biodiversity Of Madagascan frogs: phylogenetic relationships be­ course, we have much to reply from a theoretical tween these and other Southern Indian groups (e.g., point of view, and also on the grounds of conserva­ Sphaerotheca and Hydrophylax) and frogs from Mada­ tion biology and of various aspects of applied biol­ gascar and Africa are therefore a highly promising ogy. But it becomes more difficult to promote our field of research. Another promising region for am­ research field when our scientific productions are of phibian diversity is Myanmar, which has never been a low, "nonprofessional" quality Bad taxonomic seriously surveyed in this respect: the few collections works do not only do harm to their authors, but may ever made in this country by naturalists of the past also contribute to a poor rating of taxonomic activ­ (J. Anderson, E. Blyth, L. Fea, R. Kaulback, M. Stuart, ity outside this research field. Hence the particular K. Ward, etc.) are enough to show that a high rate of responsibility of editors of journals and books to endemicity is to be expected in several parts of this avoid publication of such "second-quality" works. country. Finally, even in countries whose amphibian What will be the future of amphibian taxonomy fauna is less rich than in the preceding ones, i.e., in South Asia? Predictions can be made of burgeon­ Nepal, Bhutan, Bangladesh and Pakistan, several ing activity and outstanding results in the coming new species can be expected to be found in the forth­ decades. All evaluations that can be made suggest coming years, as is suggested by the fact that some that more amphibian species remain to be discov­ of the species of these countries are known just by ered in this region than those already described. The one series of specimens, if not by a single specimen most precise such evaluation concerns Sri Lanka. At (see Dubois, 1980). the end of AFSL, an 8-page appendix presented 16 All these data suggest that exploration of amphib­ colour photos of "unidentified species". In a later ian diversity in South Asia will no doubt become a work by the same team, Pethiyagoda & fascinating "new frontier" for taxonomists at the Manamendra-Arachchi (1998: 5-8) showed 69 pho­ beginning of the 21th century. This task is an urgent tos of "undescribed species of tree frogs" from this one, as the threats to the habitats of these animals in island. These authors estimated the total number of this region are currently very serious and may lead amphibian species in Sri Lanka to about 250, versus to a rapid impoverishment of this unique fauna be­ 53 recognized in AFSL. If this is true, with about 3.9 fore it has even been inventoried and described. Let species per 1000 km2 (versus e.g. 2.75 in Costa Rica us hope that the priority of this exploration will be and 1.3 in Ecuador), Sri Lanka is one of the amphib­ understood by administrative and financial decision­ ian hotspots of the world, with a speciation level rare makers, so that the human and material tools neces­ in vertebrates and reminiscent of the radiation of sary for a thorough and professional treatment of cichlid fishes in the African lakes Malawi, Tangan­ this unique fauna be quickly made available, before yika and Victoria. Forthcoming works on the tax­ the damage has reached much higher levels than onomy of these frogs can therefore be expected to be now. In some respects, it is already too late, as exem­ rich in findings that may lead to partial or total re- plified by Sri Lanka where, according to Pethiyagoda evaluation of the taxonomy of some groups of & Manamendra-Arachchi (1998:10), more than 80 % anurans. of the rain forest has been lost already, and for which Although probably much less rich in species than several amphibian specimens in European museums Sri Lanka, neighbouring countries probably also har­ seem to be no longer represented by known living bour many more species than are currently known. populations. In view of this, collecting of specimens, Southern India in particular can be expected to pro­ tissues, and various kinds of biological information, vide many new species of the family Microhylidae in all possible regions and habitats appears a major and, in the family Ranidae, of the subfamilies priority In a few cases, quick "provisional" naming Ceratobatrachinae (as defined by Marmayou et al., of species (before proper taxonomic work as advised in preparation), Rhacophorinae and Ranixalinae. The above) may be justified by the need to bring sup­ last subfamily (first recognized by Dubois, 1987a and port to projects of natural parks or reserves, or other 1992, and later incorrectly renamed twice, as conservation-oriented programs. But, even for con­ Indiraninae and as Nyctibatrachinae, by Blommers- servation purposes, really important targets will be Schlosser, 1993) is particularly interesting as it is a difficult to ascertain until a proper, professional tax­ geographically very circumscribed taxon, with two onomy of the organisms is available. Supporting genera in India (Indirana and Nyctibatrachus) and one high quality works in this domain should be a ma­ in Sri Lanka (Nannophrys), showing some peculiar jor aim of biological research in the 21th century derived characters in adults and tadpoles (see Clarke, (see also Dubois, 1997a).

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Literature cited Dubois, A., 1975a. Un nouveau sous-genre (Paa) et trois nouvelles especes du genre Rana. Remarques sur la Anonymous [International Commission on Zoological phylogenie des Ranides (Amphibiens, Anoures). Bull. Nomenclature], 1985. International code of zoological Mus. natn. Hist, nat., (3), 324 (Zool. 231): 1093-1115. nomenclature. Third edition. London, International Dubois, A., 1975b. Un nouveau complexe d'especes Trust for zoological Nomenclature: i-xx + 1-338. jumelles distinguees par le chant: les grenouilles du Anders, C. C., Diener, A. & H. H. Schleich, 1998. First Nepal voisines de Rana limnocharis Boie (Amphibiens, record of Polypedates taeniatus (Boulenger, 1906) from Anoures). C. r. Acad. Sci., (D), 281: 1717-1720. Nepal (Amphibia, Anura: Rhacophoridae). Dubois, A., 1976. Les grenouilles du sous-genre Paa du Veroffentlichungen Fulhrott-Mus., 4: 73-86. Nepal (famille Ranidae, genre Rana). Cahiers Nepalais Annandale, N., 1912. Zoological results of the Abor - Documents, Paris, CNRS, 6: i-vi + 1-275. expedition 1911-1912. I. Batrachia. Rec. Indian Mus., Dubois, A., 1979. Notes sur la systematique et la 8: 7-36, pi. 2-4. repartition des Amphibiens Anoures de Chine et des Blommers-Schlosser, R. M. A., 1993. Systematic regions avoisinantes. II. Rana blanfordii Boulenger, relationships of the Mantellinae Laurent 1946 (Anura 1882, Rana polunini Smith, 1951 et Rana yadongensis Wu, Ranoidea). Ethol. Ecol. Evol., 5: 199-218. 1977. Bull. mens. Soc. Linn. Lyon, 48: 657-661. Bossuyt, F. & A. Dubois, in preparation. A review of the Dubois, A.,1980. L'influence de l'homme sur la repartition frog genus Philautus Gistel, 1848 (Amphibia, Anura, des amphibiens dans l'Himalaya central et occidental. Ranidae, Rhacophorinae). C. r. Soc. Biogeogr., 55: 155-178. Boulenger, G. A., 1882. Catalogue of the Batrachia Salientia Dubois, A.,1981. Liste des genres et sous-genres nominaux s. Ecaudata in the collection of the British Museum. de Ranoidea (Amphibiens Anoures) du monde, avec London, Taylor & Francis: i-xvi + 1-503, pi. 1-30. identification de leurs especes-types: consequences Boulenger, G. A., 1890. The fauna of the British India, nomenclaturales. Monit. zool. ital., (n.s.), 15, suppl.: including Ceylon and Burma. Reptilia and Batrachia. 225-284. London, Taylor & Francis: i-xviii + 1-541. Dubois, A., 1982. Le statut nomenclatural des noms Boulenger, G. A., 1920. A monograph of the South Asian, generiques d'amphibiens crees par Kuhl & Van Hasselt Papuan, Melanesian, and Australian frogs of the genus (1822): Megophrys, Occidozyga et Rhacophorus. Bull. Rana. Rec. indian Mus., 20: 1-126. Mus. Natn. Hist, nat., (4), 4 (A): 261-280. Bourret, R., 1942. Les Batraciens de lTndochine. Hanoi, Dubois, A., 1983. Note preliminaire sur le groupe de Rana Institut oceanographique de lTndochine: i-x + 1-547, (Tomopterna) breviceps Schneider, 1799 (Amphibiens, 4 pi. Anoures), avec diagnose d'une sous-espece nouvelle Brown, W. C. & A. C. Alcala, 1994. Philippine frogs of the de Ceylan. Alytes, 2 (4): 163-170. family Rhacophoridae. Proc. Calif. Acad. Sci., 48 (10): Dubois, A., 1984a. La nomenclature supragenerique des 185-220. Amphibiens Anoures. Mem. Mus. natn. Hist, nat., (A), Chanda, S. K. & A. K. Ghosh, 1988. Addenda to the 131: 1-64. amphibian fauna of India. J. Bombay nat. Hist. Soc., Dubois, A., 1984b. Note preliminaire sur le groupe de Rana 85: 626-627. limnocharis Gravenhorst, 1829 (Amphibiens, Anoures). Chanda, S. K. & A. K. Ghosh,1989. A new frog of the genus Alytes, 3: 143-159. Philautus Gistel, from the proposed Namdapha Dubois, A.,1987a. Miscellanea taxinomica batrachologica Biosphere Reserve, Arunachal Pradesh, Northeast (I). Alytes, «1986», 5 (1-2): 7-95. India. J. Bombay nat. Hist. Soc., 86: 215-217. Dubois, A., 1987b. [Review of Frost, 1985]. Copeia, 1987: Chanda, S. K. & A. K. Sarkar, 1997. The validity of Philautus 830-833. shyam rupus Chanda and Ghosh, 1989 (Anura: Dubois, A., 1987c. Living amphibians of the world: a first Rhacophoridae). Hamadryad, 22 (1): 46-47. step towards a comprehensive checklist. Alytes, 5: 99- Clarke, B. T., 1983. A morphological re-examination of the 149. frog genus N annophrys (Anura: Ranidae) with Dubois, A., 1987d. Again on the nomenclature of frogs. comments on its biology, distribution and Alytes, 6 (1-2): 27-55. relationships. Zool. J. Linn. Soc., 79: 377-398. Dubois, A., 1992. Notes sur la classification des Ranidae De Queiroz, K. & J. Gauthier, 1994. Toward a phylogenetic (Amphibiens Anoures). Bull. mens. Soc. linn. Lyon, 61 system of biological nomenclature. Trends Ecol. Evol., (10): 305-352. 9: 27-31. Dubois, A., 1995. The valid scientific name of the Italian De Silva, A., 1996. The Amphibia of Sri Lanka: a checklist treefrog, with comments on the status of some early and an annotated bibliography. Kandy, Sri Lanka, scientific names of Amphibia Anura, and on some Department of Wildlife Conservation, GEF, UNDP & articles of the Code concerning secondary homonyms. FAO, 1996: [i-ii] + 1-48. Dumerilia, 2: 55-71. Dring, J., 1987. Bornean treefrogs of the genus Philautus Dubois, A., 1997a. Editorial: 15 years of Alytes. Alytes, 14 (Rhacophoridae). Amphibia-Reptilia, 8 (1): 19-47. (4): 129. Dubois, A., 1974. Liste commentee d'amphibiens recoltes Dubois, A., 1997b. Instructions to authors of papers au Nepal. Bull. Mus. natn. Hist, nat., (3), 213 (Zool. submitted to Alytes. Alytes, 14 (4): 175-200. 143): 341-411. Dubois, A., 1998a. Mapping European amphibians and

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reptiles: collective inquiry and scientific methodology. Kirtisinghe, P., 1957. The Amphibia of Ceylon. Colombo: Alytes, 15 (4): 176-204. i-xiii + 1-112, 1 pi. Dubois, A., 1998b. Lists of European species of amphibians Laurent, R., 1951. Sur la necessite de supprimer la famille and reptiles: will we soon be reaching "stability"? des Rhacophoridae mais de creer celle des Amphibia-Reptilia, 19 (1): 1-28. Hyperoliidae. Rev. Zool. Bot. afr., 45: 116-122. Dubois, A., in press. The influence of man on the Laurent, R. F., 1986. Sous-classe des Lissamphibiens distribution of amphibians in the Himalayas of Nepal: (Lissamphibia). Systematique. In: P.-P. Grasse & M. an example of critical evaluation of biogeographical Delsol (ed.), Traite de Zoologie, 14, Amphibiens, fasc. data. Marburger geogr. Schriften, in press since 1997. I-B, Paris, Masson: 594-796. Dubois, A. & M. S. Khan, 1980. A new species of frog Lavilla, E. O., 1988. Telmatobius (Anura: Leptodactylidae): (genus Rana, subgenus Paa) from northern Pakistan the name-bearing types of five Vellard's taxa. Alytes, (Amphia, Anura). J. Herp., 13: 403-410. 7 (1): 6-18. Dubois, A. & M. Matsui, 1983. A new species of frog (genus Liem, S. S., 1970. The morphology, systematics, and Rana, subgenus Paa) from western Nepal (Amphibia: evolution of the Old World treefrogs (Rhacophoridae Anura). Copeia, 1983 (4): 895-901. and Hyperoliidae). Fieldiana: Zool., 57: i-vii + 1-145. Dubois, A. & A. Ohler, 1995. Frogs of the subgenus Malkmus, R. & K. Riede, 1996a. Die Baumfrosche der Pelophylax (Amphibia, Anura, genus Rana): a catalogue Gattung Philautus vom Mount Kinabalu. Teil I. of available and valid scientific names, with comments Uberblick und die aurifasciatus-G ruppe mit on name-bearing types, complete synonymies, Beschreibung einer neuen Art (Philautus saueri n. sp.). proposed common names, and maps showing all type Sauria, 18 (1): 27-37. localities. Zool. Polon., (1994), 39 (3-4): 139-204. Malkmus, R. & K. Riede, 1996b. Die Baumfrosche der Dubois, A. & A. Ohler, 1997. Early scientific names of Gattung Philautus vom Mount Kinabalu. Teil II. Die Amphibia Anura. I. Introduction. Bull. Mus. natn. Hist, vermiculatus-Gruppe mit Beschreibung einer neuen nat., (4), 18 (3-4): 297-320. Unterart (Philautus aurantium gunungensis n. ssp.) und Dubois, A. & A. Ohler, 1998. A new species of die /zosd-Gruppe. Sauria, 18 (2): 21-28. Leptobrachium (Vibrissaphora) from northern Vietnam, Marmayou, J., Dubois, A., Ohler, A., Pasquet, E. & A. with a review of the taxonomy of the genus Tillier, in preparation. Phylogenetic relationships in the Leptobrachium (Pelobatidae, Megophryinae). Ranidae (Amphibia, Anura): independent origin of Dumerilia, 4 (1): 1-32. direct development in the genera Philautus and Dubois, A. & A. Ohler, in preparation. Frogs of the genera Taylor ana. Tomopterna, Sphaerotheca and Laliostoma (Amphibia, Perret, J.-L., 1976. 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Hist., (4), 18: 356-357. Schleich, H. H. & C. C. Anders, 1998. Tomopterna maskeyi Frost, D. R. (ed.), 1985. Amphibian species of the world. spec. nov. from Nepal (Amphibia, Anura). Lawrence, Allen Press & Assoc. Syst. Coll.: [i-iv] + i-v Veroffentlichungen Fulhrott-Mus., 4: 57-72. + 1-732. Siedlecki, M., 1909. Zur Kenntnis des javanischen Gorham, S. W., 1974. Checklist of world amphibians up Flugfrosches. Biol. Centralblatt, 29: 704-714, pi. 7-8. to January 1, 1970. Saint-John, The New Brunswick Swan, L. W. & A. E. Leviton, 1962. The herpetology of Museum: 1-173. Nepal: a history, check list, and zoogeographical Gunther, A., 1864. The reptiles of British India. London, analysis of the herpetofauna. Proc. Calif. Acad. Sci., Ray Society: i-xxvii + 1-452, pi. 1-26. (4), 32: 103-147. Gunther, A., 1876. Notes on the mode of propagation of Van Kampen, P. N., 1909. Beitrag zur Kenntnis der some Ceylonese tree-frogs, with description of two Amphibienlarven des indischen Archipels. new species. Ann. Mag. nat. Hist., (4), 17: 377-380, pi. Natuurkundig Tijdschrift Ned.-Indie, 69: 25-48, pi. 2. 20 fig. C. Van Kampen, P. N., 1923. The Amphibia of the Indo- Inger, R. F., 1989. Four new species of frogs from Borneo. Australian archipelago. Leiden, Brill: i-xii + 1-304. Malayan Nature J., 42: 229-243. Wolf, S. 1936. Revision der Untergattung Rhacophorus Kirtisinghe, P., 1946. The presence in Ceylon of a frog with (ausschliesslich der Madagaskar-Formen). Bull. Raffles direct development on land. Ceylon J. Sci., (B), 23:109- Mus., 12: 137-217. 112.

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