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Systematics of the Genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae): Some Historical and Metataxonomic Comments

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Systematics of the Genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae): Some Historical and Metataxonomic Comments J. South Asian Nat. Hist., ISSN 1022-0828. May, 2001. Vol. 5, No. 2, pp. 173-186,8 figs., 9 tabs. © 2001, Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka. Systematics of the genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae): some historical and metataxonomic comments Alain Dubois & Annemarie Ohler Laboratoire des Reptiles et Amphibiens, Museum national d'Histoire naturelle, 25 rue Cuvier, 75005 Paris, France. Abstract We present some historical and metataxonomic comments on the data presented by Bossuyt & Dubois (2001) in their review of the genus Philautus. This stresses the existence of some historical and geographical patterns in the descriptions of species once referred to this genus, and in their changes of generic allocation. These data suggest that changes in taxonomies are not only linked to theoretical evolution of the dominant ideas in the field of taxonomy, but are also largely influenced by the ideas and skills of individual taxonomists. Some comments are also offered regarding the replacement in the scientific literature of the name Ixalus by the name Philautus, and the fate of the name-bearing types of species referred to this genus. We suggest that the methodology here proposed for historical and metataxonomic analysis of systematic data could be used profitably for studying the taxonomies of other zoological groups. Introduction Taxonomy is an old, but still very lively and recent research field sometimes also known as promising, discipline of biology. In zoology, the "comparative systematics" (Mayr & Short, 1970; taxonomic and nomenclatural system currently in Dubois, 1977, 1988, 1998; Bock & Farrand, 1980; force, used by the vast majority of specialists, is Dubois & Ohler, 1995) or "metataxonomy" (Van Valen, derived from that first used by Linnaeus (1758). Since 1973; Dubois, 1988). that time, methods and concepts have undergone Bossuyt & Dubois (2001) provided a review of the considerable evolution, and the current concepts in information dealing with the descriptive taxonomy taxonomy are quite different from what they were then of the frog genus Philautus Gistel, 1848 (Ranidae, (see e.g. Mayr, 1982, 1999; Mayr & Ashlock, 1991). Rhacophorinae). They proposed a working taxonomy However, in order for taxonomy and nomenclature to for this group and a provisional status for the 177 be able properly to play their role of storage and species-group names published since 1758 for frogs retrieval of information, they must show both a allocated by previous authors to this group. The data historical continuity and robustness, and a capacity presented in this work attract some interesting for adaptation and evolution. Thus, the taxonomy of historical and metataxonomic comments. any zoological group continuously evolves through We tried to measure the "robustness" of the time, both because of the evolution of methods and taxonomies proposed by all authors who described concepts, and because of the addition of new basic new species once referred to the genus Philautus data, in particular through field surveys and during the long history (1822-1999) of study of this collection and study of specimens (see e.g. Dubois, genus. For the purpose of this evaluation, 1998). A historical and quantitative analysis of this "robustness" is not exactly equivalent to "stability", evolution could serve to enlighten the way taxonomy which would mean long-term persistence of a works and evolves, independently of the ideas and taxonomy without change: we consider that a theories of taxonomists themselves. Such analyses taxonomy proposed by an author is "robust" if it is belong in the domain of "comparative taxonomy", a currently in use, even if it had been abandoned for D u b o i s & O h l e r some time between its initial proposal and the present species referred by them to the genus Philautus, while time. We used three indexes to measure the robustness no less than 56 (i.e. 31.6 %) apply to species referred of a taxonomy as just defined: the "species-name to other genera. As shown here in Table 1, among the robustness index", the "generic allocation robustness 121 names that refer to Philautus as here understood, index" and the "lower taxonomy robustness index". 84 (i.e. 69.4 %) are here considered provisionally valid, Other indexes could also be computed, dealing e.g. while 37 (i.e. 30.6 %) are invalid, being either junior with higher taxonomy (above that of genus) or with subjective synonyms (25 names, i.e. 20.7 %), or junior the whole scheme of classification used by an author objective synonyms, junior homonyms or (rather than only its original new species names and nomenclaturally unavailable names. The percentage combinations) or at a given date, but we do not explore of 69.4 % of valid names is somewhat higher, but of them here. the same order of magnitude, as that (64 %) obtained The "species name robustness index" (SNRI) is for all living Amphibia of the world at the time of the ratio (expressed in percent) of the number of Gorham's (1974) checklist, as analysed by Dubois species-group names created by an author or in a (1977: 235-248). given period that are currently considered valid If we consider the geographic distribution over 11 (irrespective of their having possibly been considered geographic regions of the type localities of these 121 as synonyms for some time) to the total number of nominal species (Table 2), it is clear that this genus species-group names created by this author or in this has been unevenly studied over its wide range. The period. best studied areas appear to be Sri Lanka (16 valid The "generic allocation robustness index" (GARI) species, i.e. 55.2 % of names), Borneo (14 valid species, is the ratio (expressed in percent) of the number of i.e. 93.3 % of names), southern and central India (12 species-group names that are currently allocated to the same genus as they were at their creation (irrespective of possible changes in their generic Table 1. Current taxonomic status of the 121 nominal spec­ allocation in the meanwhile) to the total number of ies described from 1822 to 1999 and here referred to the species-group names created by this author or in this genus Philautus (excluding nominal species now placed in period. other genera). TN, total number of nominal species creat­ ed during period; VN, number of nominal species cur­ Finally, the "lower taxonomy robustness index" rently considered valid; JSS, number of junior subjective (LTRI) is the mean (also expressed in percent) of SNRI synonyms; JOS, number of junior objective synonyms and GARI. (including new replacement names and unjustified Of course, robustness as measured by these indexes emendations); JH, junior (primary and secondary) homo­ is highly dependent on the current taxonomy, which nyms; UN, unavailable names (nomina nuda, incorrect is taken as a comparative base line: the values of these spellings); IN, total number of nominal species currently indexes will change every time the "current considered invalid (JSS + JOS + JH + UN); SNRI, species taxonomy" changes. This will no doubt be the case name robustness index = VN / TN %. with the taxonomy proposed by Bossuyt & Dubois (2001) for the 177 species studied, and the conclusions Period TN VN JSS JOS JH UN IN SNRI below will probably have to be modified accordingly. However, we present them here for two major reasons: 1820-1829 1 0 0 0 0 1 1 0 1830-1839 1 1 0 0 0 0 0 100 (1) despite these predictable changes we think that 1840-1849 0 0 0 0 0 0 0 0 the major trends described below will probably 1850-1859 7 7 0 0 0 0 0 100 remain true; (2) the new methodology here proposed 1860-1869 9 8 1 0 0 0 1 88.9 for such analyses can be used again to study the 1870-1879 15 8 5 1 1 0 7 53.3 forthcoming new taxonomies of this group, as well as 1880-1889 6 3 1 1 1 0 3 50.0 of other animal groups. 1890-1899 9 8 1 0 0 0 1 88.9 All calculations below were made using the SPSS 1900-1909 11 6 5 0 0 0 5 54.5 Base 9.0 software (Anonymous, 1999). 1910-1919 7 5 2 0 0 0 2 71.4 1920-1929 13 5 6 2 0 0 8 38.5 1930-1939 7 5 1 1 0 0 2 71.4 Historical and geographical patterns 1940-1949 3 2 1 0 0 0 1 66.7 in the description of species of the genus Philautus 1950-1959 2 0 2 0 0 0 2 0 1960-1969 2 2 0 0 0 0 0 100 Bossuyt & Dubois (2001) proposed a provisional 1970-1979 4 4 0 0 0 0 0 100 status for the 177 species-group names studied in 1980-1989 11 9 0 2 0 0 2 81.8 detail by them. Of these 177 names (listed in Table 1 1990-1999 13 11 0 1 0 1 2 84.6 of Bossuyt & Dubois, 2001), 121 (i.e. 68.4 %) apply to Total 121 84 25 8 2 2 37 69.4 174 J. South Asian Nat. Hist. H istory and metataxonomy of P h il a u t u s Table 2. Location of the type-localities of the 121 nominal species here referred to the genus Philautus. Southern and central India: Andhra Pradesh, Karnataka, Kerala, Madhya Pradesh, Tamil Nadu. Northeastern India: Arunachal Pradesh, Meghalaya, West Bengal. TN, total number of names; VN, number of names here considered valid; IN, number of names here considered invalid; SNRI, species name robustness index = VN / TN %. Region and area TNVNINSNRI Sri Lanka 29161355.2 Southern and central India 21 12 9 57.1 Subtotal south Asia 50 28 22 56.0 Northeastern India 9 8 1 88.9 China 9 8 1 88.9 Figure 1.
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