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Morphological Assessments and Phylogenetic Relationships of the Seychellean Frogs of the Family Sooglossidae (Amphibia: Anura) Ronald A

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Morphological Assessments and Phylogenetic Relationships of the Seychellean Frogs of the Family Sooglossidae (Amphibia: Anura) Ronald A Zoological Studies 46(3): 322-335 (2007) Morphological Assessments and Phylogenetic Relationships of the Seychellean Frogs of the Family Sooglossidae (Amphibia: Anura) Ronald A. Nussbaum1 and Sheng-Hai Wu2,* 1Division of Amphibians and Reptiles, Museum of Zoology, University of Michigan, Ann Arbor, MI 48109-1079, USA 2Department of Life Sciences, National Chung-Hsing University, 250 Kuo-Kwang Road, Taichung 402, Taiwan (Accepted December 21, 2006) Ronald A. Nussbaum and Sheng-Hai Wu (2007) Morphological assessments and phylogenetic relationships of the Seychellean frogs of the family Sooglossidae (Amphibia: Anura). Zoological Studies 46(3): 322-335. The frog family Sooglossidae is endemic to the Seychelles Is. in the Indian Ocean, and consists of 2 genera and 4 species, the monotypic Nesomantis and Sooglossus (3 species). Many previous studies have suggested that Sooglossus sechellensis is more similar to Nesomantis thomasseti than to its congener, S. gardineri. Based on an extensive dataset of 188 morphological characters, we concluded that Nesomantis and S. sechel- lensis form a monophyletic group with the exclusion of S. gardineri (and S. pipilodryas). We therefore have established a new genus, Sechellophryne, to accommodate the latter 2 species. http://zoolstud.sinica.edu.tw/Journals/46.3/322.pdf Key words: Nesomantis thomasseti, Sooglossus gardineri, Sooglossus pipilodryas, Sooglossus sechellensis, Sechellophyrne gen. nov. The granitic Seychelles Is. lie in the western rious until the late 20th century when continental Indian Ocean between 4 and 5 S latitude and drift gained respectability, and the formation of the 55 and 56 E longitude, °about 1600° km east of western Indian Ocean through the breakup of Africa° (Mombasa),° 926 km northeast of Gondwanaland began to be understood. Madagascar (Antsiranana), and 2900 km south- The amphibian fauna of the Seychelles con- west of India (Mumbai). These islands have fasci- sists of 13 species (6 caecilians and 7 frogs), 12 of nated biogeographers and geologists since the which are endemic. The single non-endemic late 19th century when Darwin (1859) and Wallace species, the ranid frog, Ptychadena mascarenien- (1880) pointed out that they are continental in sis, is widespread in sub-Saharan Africa, character in spite of their great distance from large Madagascar, and the Mascarene Is. It may have continental blocks. been introduced to the Seychelles by humans Among the unusual features of the (Mertens 1934, Vences et al. 2004, but see Seychelles is the presence of endemic amphib- Nussbaum 1984). ians. Wallace (1880) recognized amphibians as a The 6 species of caecilians (in 3 endemic group with very limited means of transoceanic dis- genera) in the Seychelles are monophyletic persal, and their presence in the Seychelles was (Nussbaum and Ducey 1988, Haas et al. 1993, an important factor that prompted him to classify Hedges and Maxson 1993, Nussbaum unpubl. the Seychelles as“continental”as opposed to data), and molecular data (Haas et al. 1993) sug- “oceanic”islands. However, the means by which gest that the lineage has been present in the amphibians and other sedentary groups may have Seychelles for at least 30 million yrs. Wilkinson et populated the granitic Seychelles remained myste- al. (2002 2003) suggested that the nearest relative *To whom correspondence and reprint requests should be addressed. Tel: 886-4-22840319 ext. 711. Fax: 886-4-22874740. E-mail:[email protected] 322 Nussbaum and Wu -- Morphological Phylogeny of Sooglossidae 323 of Seychellean caecilians is the Indian genus The purposes of this paper were to (1) use a Gegeneophis. cladistic methodology applied to a broad morpho- Of the 6 endemic species of frogs, logical database to confirm or refute the mono- Tachycnemis seychellensis belongs to a monotyp- phyletic status of the sooglossids; and (2) deter- ic (Nussbaum and Wu 1995) hyperoliid genus. It mine the cladistic relationships within the family. is placed either in a monotypic subfamily (Tachycneminae) (Channing 1989), sister to the Taxonomic history remaining hyperoliids or to the Malagasy Heterixalus (Frost et al. 2006). Boettger (1896) described the first sooglossid The remaining 4 Seychellean frogs constitute as Arthroleptis sechellensis, thereby assigning the the 2 genera of Nesomantis (thomasseti) and species to a group of African ranids. Boulenger Sooglossus (gardineri, pipilodryas, and sechellen- (1906) transferred Arthroleptis sechellensis to a sis) of the endemic family, the Sooglossidae. At new genus, Sooglossus, on the basis of tongue the time of their discovery at the end of the 19th morphology (entire and elliptical), but retained the century, the sooglossids were assigned to the fam- species among the ranids. Boulenger (1909) ilies Ranidae and Bufonidae. Noble (1926) placed described Nesomantis thomasseti as a new frog them together as a distinctive group of the genus and species from Mahé I., Seychelles. He Pelobatidae, which he later (Noble 1931) formally wrote that Nesomantis is allied to Sooglossus, but named the Sooglossinae; but this was viewed with unlike the latter genus, Nesomantis has vomerine skepticism based on biogeographic considerations teeth and lacks a claw-like, dermal tip to the digits. (e.g., Darlington, 1957). Boulenger viewed S. sechellensis as a dwarf Griffiths (1963) defined the family Nesomantis. He did not comment on its relation- Sooglossidae and placed it among the ranoids, ships to other frogs, although he apparently and this was accepted by most subsequent assumed it was a ranid close to Arthroleptis and authors (e.g., Savage 1973, Duellman 1975). other African ranoids. Shortly afterwards, However, other phylogenetic hypotheses also Boulenger (1911) described a 3rd Seychellean frog abound, mainly as the result of using different as Nectophryne gardineri, believing it to belong to characters and species (as a myobatrachid: Lynch an African bufonid genus, because of webbing at 1973, Nussbaum 1979 1980, Duellman and Trueb the base of the toes. 1986, Ford and Canatella 1993; as a lepto- Noble (1926) noted pelobatid characteristics dactylid: Laurent 1975, Tyson 1987; as a group in all three of these Seychellean frogs, and conse- including dendrobatids and microhyloids: quently, he (1931) established a new pelobatid Blommers-Schlösser 1993; as basal to the subfamily, the Sooglossinae, to accommodate all 3 Hyloidea: Hoegg et al. 2004; as basal in the species. Hyloides: Frost et al. 2006; as basal to the Noble (1926) stated that the differences ranoids: Tyler 1985; or as a sister to the recently between S. sechellensis and Nesomantis“are very described Indian Nasikabatrachidae: Biju and trivial”, but that it is“well to utilize these generic Bossuyt 2003). names until the anatomy of N. thomasseti and S. Although the phylogenetic position of the sechellensis is better known.” He indicated that a , sooglossids is unresolved, Noble s (1926) con- separate genus might be justified for Nec. gar- tention that they are monophyletic has not been dineri, but instead he opted for expanding the defi- refuted (Griffiths 1959 1963, Nussbaum 1979 1980 nition of Sooglossus and placing it in the latter 1982, Tyler 1985, Green et al. 1988). genus as Soo. gardineri. It appears that the classification of the 3 The recently described Nasikabatrachidae species (prior to the description of S. pipilodryas) from the Western Ghats of India is considered to did not accurately reflect their phylogenetic rela- be the sister group of Sooglossidae based on mol- tionships. Comparisons of chromosomes ecular evidence (Biju and Bossuyt 2003). (Nussbaum 1979), advertisement calls (Nussbaum However, based on the limited morphological evi- et al. 1982), and isozymes (Green et al. 1988) all dence from their description, it is more like the suggested that Sooglossus gardineri is the most African Hemisotidae in the unique pectoral girdle divergent species and that it, and not Nesomantis architecture, external morphology, and life history thomasseti, should be placed in a separate genus. (Scott 2005). However, Frost et al. (2006) found This conclusion, based on phenetics, needs to be further support for a sooglossid-nasikabatrachid confirmed using cladistic methods. relationship and considered Nasikabatrachidae to 324 Zoological Studies 46(3): 322-335 (2007) be a junior synonym of the Sooglossidae. 1119 equally parsimonious cladograms had a con- Gerlach and Willi (2002) described Soo. pipi- sistency index of 0.243. Sooglossidae monophyly lodryas from the island of Silhouette as a new was supported by parsimony and bootstrapping cryptic species closely related to S. gardineri. If (Fig. 1). Soo. pipilodryas proves to be a valid species (dis- Twenty-four synapotypies diagnose the family, tinct from S. gardineri), then it is clearly the sister and bootstrap support for the clade was strong. species of S. gardineri, and its existence will not Nesomantis thomasseti and Soo. sechellensis affect the phylogenetic analysis that follows. share 7 synapotypies, and bootstrap support for the grouping was moderately strong, thus support- ing the monophyly of the 2 species with the exclu- MATERIALS AND METHODS sion of Soo. gardineri. The family Sooglossidae is grouped with the We recorded morphological characters from Dendrobatidae and Leptodactylidae, but the boot- 36 frog species representing 12 families (Appendix strapping support was low. Firmsternal families 1). The data set include 188 characters and is part (Arthroleptidae, Microhylidae, and Petropedetidae) of
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