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Leaping Frogs (Anura: Ranixalidae) of the Western Ghats of India: an Integrated Taxonomic Review

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Leaping Frogs (Anura: Ranixalidae) of the Western Ghats of India: an Integrated Taxonomic Review OPEN ACCESS All articles published in the Journal of Threatened Taxa are registered under Creative Commons Attribution 4.0 Interna- tional License unless otherwise mentioned. JoTT allows unrestricted use of articles in any medium, reproduction and distribution by providing adequate credit to the authors and the source of publication. Journal of Threatened Taxa The international journal of conservation and taxonomy www.threatenedtaxa.org ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print) Monograph Leaping frogs (Anura: Ranixalidae) of the Western Ghats of India: An integrated taxonomic review Neelesh Dahanukar, Nikhil Modak, Keerthi Krutha, P.O. Nameer, Anand D. Padhye & Sanjay Molur 15 September 2016 | Vol. 8 | No. 10 | Pp. 9221–9288 10.11609/jott.2532.8.10.9221-9288 For Focus, Scope, Aims, Policies and Guidelines visit http://threatenedtaxa.org/About_JoTT.asp For Article Submission Guidelines visit http://threatenedtaxa.org/Submission_Guidelines.asp For Policies against Scientific Misconduct visit http://threatenedtaxa.org/JoTT_Policy_against_Scientific_Misconduct.asp For reprints contact <[email protected]> Publisher/Host Partner Threatened Taxa Journal of Threatened Taxa | www.threatenedtaxa.org | 15 September 2016 | 8(10): 9221–9288 Leaping frogs (Anura: Ranixalidae) of the Western Ghats of Monograph India: An integrated taxonomic review Neelesh Dahanukar 1,*, Nikhil Modak 2,*, Keerthi Krutha 3, P.O. Nameer 4, Anand D. Padhye 5 & ISSN 0974-7907 (Online) ISSN 0974-7893 (Print) Sanjay Molur 6 *Joint first authors OPEN ACCESS 1 Indian Institute of Science Education and Research (IISER), G1 Block, Dr. Homi Bhabha Road, Pashan, Pune, Maharashtra 411008, India 1,6 Systematics, Ecology and Conservation Laboratory, Zoo Outreach Organization (ZOO), 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India 2 Department of Biodiversity, MES Abasaheb Garware College, Karve Road, Pune, Maharashtra 411004, India 3,6 Wildlife Information Liaison Development (WILD) Society, 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India 4 Centre for Wildlife Studies, College of Forestry, Kerala Agricultural University, Thrissur, Kerala 680656, India 5 Department of Zoology, MES Abasaheb Garware College, Karve Road, Pune, Maharashtra 411004, India 5 Institute of Natural History Education and Research (INHER), B1/602 Kumar Parisar, Kothrud, Pune, Maharashtra 411038, India 1 [email protected] (corresponding author), 2 [email protected], 3 [email protected], 4 [email protected], 5 [email protected], 6 [email protected] Abstract: Leaping frogs of the family Ranixalidae are endemic to the Western Ghats of India and are currently placed in a single genus, Indirana. Based on specimens collected from their entire range and a comprehensive study of type material defining all known species, we propose a revised taxonomy for the leaping frogs using an integrative approach including an analysis of the mitochondrial 16S rRNA and nuclear rhodopsin genes, as well as multivariate morphometrics. Both genetic and morphological analyses suggest that the genus Indirana is paraphyletic and a distinct monophyletic group, Walkerana gen. nov., is described herein. The new genus is separated from Indirana sensu stricto by an apomorphic character state of reduced webbing, with one phalange free on the first and second toe (vs. no free phalanges), two phalanges free on the third and fifth toe (vs. one free phalange), and three phalanges free on the fourth toe (vs. 2–2½ phalanges free). This review includes (i) identification of lectotypes and redescription of three species of the genus Walkerana; (ii) identification of lectotypes for Indirana beddomii and I. semipalmata and their redescription; (iii) redescription of I. brachytarsus and I. gundia; and (iv) descriptions of four new species, namely, I. duboisi and I. tysoni from north of the Palghat gap, and I. yadera and I. sarojamma from south of the Palghat gap; and (iv) a key to the genera and species in the family Ranixalidae. Keywords: Discriminant analysis, genetic gap analysis, Indirana, molecular phylogeny, new combinations, new genus, new species, taxonomy, Walkerana gen. nov. DOI: http://dx.doi.org/10.11609/jott.2532.8.10.9221-9288 | ZooBank: urn:lsid:zoobank.org:pub:0AD3BF63-ECF2-433F-A47F-5F96B15D3725 Editor: Anonymity requested. Date of publication: 15 September 2016 (online & print) Manuscript details: Ms # 2532 | Received 09 February 2016 | Final received 15 August 2016 | Finally accepted 10 September 2016 Citation: Dahanukar, N., N. Modak, K. Krutha, P.O. Nameer, A.D. Padhye & S. Molur (2016). Leaping frogs (Anura: Ranixalidae) of the Western Ghats of India: An integrated taxonomic review. Journal of Threatened Taxa 8(10): 9221–9288; http://dx.doi.org/10.11609/jott.2532.8.10.9221-9288 Copyright: © Dahanukar et al. 2016. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use of this article in any medium, reproduc- tion and distribution by providing adequate credit to the authors and the source of publication. Funding: This paper is an outcome of the work conducted by the ‘amphibian chytrid project’ partially funded by the U.S. Fish and Wildlife Service ‘Amphibians in Decline’ Fund no. F11AP00334 to Sanjay Molur and Mohammed bin Zayed Species Conservation Fund (Grant nos. 12254621, 14254361) to Keerthi Krutha. Neelesh Dahanukar is supported by DST-INSPIRE Research Grant [IFA12-LSBM-21]. Nikhil Modak is supported by DST-INSPIRE Student Fellowship [IF 120398]. Nikhil Modak was supported by Ernst Mayr Travel Grant, MCZ, Harvard University, US, for examination of specimens at NHM, London and MNHN, Paris. P.O. Nameer’s biodiversity monitoring projects are funded by the Kerala Agricultural University. Conflict of Interest: The authors declare no competing interests. Funding sources had no role in study design, data collection, results interpretation and manuscript writing. For Author Details, Author Contributionand Acknowledgements see end of this article. LOGOs 9221 Leaping frogs of the Western Ghats Dahanukar et al. INTRODUCTION comprises 12 species, including: I. beddomii (Günther, 1876), I. brachytarsus (Günther, 1876), I. chiravasi The taxonomic history of anurans currently Padhye et al., 2014, I. diplosticta (Günther, 1876), I. grouped into the genus Indirana, in the monotypic gundia (Dubois, 1986), I. leithii (Boulenger, 1888), I. family Ranixalidae, dates back to Günther (1876), leptodactyla (Boulenger, 1882), I. longicrus (Rao, 1937), who described four species, Polypedates beddomii, P. I. phrynoderma (Boulenger, 1882), I. salelkari Modak brachytarsus, P. brevipalmatus and Ixalus diplostictus et al., 2015, I. semipalmata (Boulenger, 1882), and I. from the Malabar Coast of southwestern India. tenuilingua (Rao, 1937). Boulenger (1882) transferred these species to the Dubois (1987a) proposed the tribe Ranixalini, later genus Rana; however, as the name brevipalmata was treated as the subfamily Ranixalinae by Dubois (1992), preoccupied by R. brevipalmata Peters, 1871, Boulenger with the type genus Ranixalus (now a synonym of (1882) provided a replacement name R. leptodactyla for Indirana) and included two other genera, Nannophrys P. brevipalmatus. Boulenger (1882) also synonymized P. and Nyctibatrachus, based on the presence of femoral brachytarsus with R. beddomii and described two other glands in males. On the basis of priority, the subfamily related species, R. phrynoderma and R. semipalmata. Indiraninae Blommers-Schlösser, 1993, is a junior Subsequently, Boulenger (1888) described another synonym of Ranixalinae Dubois, 1987. Van Bocxlaer et member of the same group, R. leithii from Matheran in al. (2006) in their taxonomic analysis using three nuclear the northern Western Ghats. The combination of these and one mitochondrial genetic markers elevated the species into a single group was a result of Boulenger’s taxon to the family Ranixalidae with the genus Indirana (1920) monograph on Rana, in which he defined the as its sole member. subgenus Discodeles by using characters including “toes, More recent molecular studies (Nair et al. 2012a; often also fingers, dilated at the end, the dilation, or disc, Modak et al. 2014) have revealed that several species bearing a crescentic or horseshoe-shaped horizontal in the genus Indirana are undescribed. Although some groove, outer metatarsals united or separated only in the studies have been published that attempt to resolve distal third; omosternum forked at the base”. Although these issues by the study of topotypic material to assist Boulenger (1920) included three other species from the in delineating geographic distribution boundaries of Solomon Islands, Oceania, in the same subgenus, he species (e.g., Modak et al. 2014), and by defining new specifically considered a distinct ‘Ranae beddomianae’ species using an integrated taxonomic approach (Padhye group to consolidate the six species, namely, R. et al. 2014; Modak et al. 2015), a detailed taxonomic (Discodeles) beddomii, R. (D.) diplosticta, R. (D.) leithii, review of leaping frogs is still pending. R. (D.) leptodactyla, R. (D.) phrynoderma, and R. (D.) In the current study, using type material of known semipalmata. Rao (1937) later added R. (D.) tenuilingua species, and with freshly collected material from from Kemphole Forest, Karnataka, India to the same throughout the Western Ghats, emphasizing type group, elevating the number of species to seven. localities of known species, we present genetic barcodes Laurent
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