Journal of Threatened Taxa | www.threatenedtaxa.org | 26 June 2017 | 9(6): 10269–10285

 

 

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Identity of Sphaerotheca pluvialis (Jerdon, 1853) and other available names among the burrowing frogs (Anura: Dicroglossidae) of South Asia

 

 

Neelesh Dahanukar 1, Shauri Sulakhe 2 & Anand Padhye 3

 

 

1 Indian Institute of Science Education and Research (IISER), G1 Block, Dr. Homi Bhabha Road, Pashan, Pune, Maharashtra 411008, India

1 Systematics, Ecology and Conservation Laboratory, Zoo Outreach Organization (ZOO), No. 12 Thiruvannamalai Nagar, Saravanampatti - Kalapatti Road, Saravanampatti, Coimbatore, Tamil Nadu 641035, India

2,3 Institute of Natural History Education and Research (INHER), C26/9 Ketan Heights, Kothrud, Pune, Maharashtra 411038, India

3 Department of Zoology, Abasaheb Garware College, Karve Road, Pune, Maharashtra 411004, India.

1 n.dahanukar@iiserpune.ac.in, 2 shaurisulakhe@gmail.com, 3 anand.padhye@mesagc.org (corresponding author)

 

 

 

 

 

 

doi: http://doi.org/10.11609/jott.3358.9.6.10269-10285 | ZooBank: urn:lsid:zoobank.org:pub:AA692CE2-7038-41DF-9F35-DC5E74BCFFFC

 

Editor: Anonymity requested. Date of publication: 26 June 2017 (online & print)

Manuscript details: Ms # 3358 | Received 21 February 2017 | Final received 23 April 2017 | Finally accepted 20 June 2017

 

Citation: Dahanukar, N., S. Sulakhe & A. Padhye (2017). Identity of Sphaerotheca pluvialis (Jerdon, 1853) and other available names among the burrowing frogs (Anura: Dicroglossidae) of South Asia. Journal of Threatened Taxa 9(6): 10269–10285; http://doi.org/10.11609/jott.3358.9.6.10269-

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Copyright: © Dahanukar et al. 2017. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use of this article in any medium, reproduction and distribution by providing adequate credit to the authors and the source of publication.

 

Funding: The study was partially supported by The Mohamed bin Zayed Species Conservation Fund [15254528] to Anand Padhye. Neelesh Dahanukar is supported by DST-INSPIRE Research Grant [IFA12-LSBM-21].

 

Conflict of interest: The authors declare no competing interests. Funding sources had no role in study design, data collection, results interpretation and manuscript writing.

 

Author Details: Neelesh Dahanukar works in ecology and evolution with an emphasis on mathematical and statistical analysis. He is also interested in taxonomy, distribution patterns and molecular phylogeny of fish and frogs. Shauri Sulakhe is a Founder Trustee of Institute of Natural History Education and Research (INHER), Pune. He is interested in wildlife photography and natural history. Anand Padhye is an Associate Professor working on systematics, ecology, diversity, distribution and evolution of amphibians. He is also Founder Trustee of Institute of Natural History Education and Research (INHER), Pune.

 

Author Contribution: ND and AP designed the study; ND and AP diagnosed the species; SS and AP collected specimens; AP and SS studied museum specimens and collected morphometric data; ND collected genetic data; ND performed genetic and morphometric analysis; ND and AP wrote the manuscript.

 

 

Acknowledgements: Anand Padhye is thankful to Head of the Zoology Department and Principal, MES’ Abasaheb Garware College, Pune; and Neelesh Dahanukar is thankful to the Director, Indian Institute of Science Education and Research, Pune, for providing infrastructural facilities. We are thankful to Dr. H.V. Ghate and Dr. Anil Mahabal for helpful discussions and valuable suggestions. Special thanks to Dr. Kailash Chandra, Director, Zoological Survey of India, for necessary permissions; to Dr. Kaushik Deuti, Scientist; and Sudipta Debnath, Zoological Assistant; for their help in the study of specimens in ZSI Kolkata museum collection; and to Dr. K. A. Subramanian, Scientist, ZSI-SRC, Chennai, for helping us variously. We are thankful to Dr. Lauren Scheinberg, collection manager, Herpetology California Academy of Science for providing photographs of holotype of S. swani. We are thankful to Dr. Jefferey Streicher Curator, Amphibians, Natureal History Museum, London, for providing the photographs of S. rolandae holotype. We are thankful to Soham Dixit for providing photographs of syntypes of S. breviceps. We are grateful to Dr. Mark-Oliver Rodel and Dr. Frank Tillack, Museum für Naturkunde (ZMB), Berlin, for helping Soham Dixit in examining the syntypes of S. breviceps. We are grateful to Dr. Deepak Apte, Director; Dr. Rahul Khot, incharge Natural History Collection; and Vithoba Hegde, senior field assistant, for their help in registration of specimens at Bombay Natural History Society (BNHS), Mumbai. We also thank P.S. Bhatnagar, officer-in-charge, and Sameer Pati, Zoological Survey of India, Western Regional Center (ZSI-WRC), Pune, for their help in registering specimens in ZSI-WRC. We are thankful to Dr. Sanjay Molur for help in registration of specimens in the Wildlife Information Liaison Development (WILD), Coimbatore. We thank Dr. Annemarie Ohler (Muséum National d’Histoire Naturelle, Paris), Dr. Lauren Scheinberg, Dr. Jens Vindum, Dr. Darrel Frost (California Academy of Science) and Dr. Frank Tillack for providing important references. We are thankful to Nikhil Dandekar, Shankar Sarma, Satish Pande, Sanjay Khatavkar, Chaitanya Risbud, Makarand Ketkar, Varun Kher, Mayuresh Kulkarni, Srushti Bhave, Nikhil Modak, Amit Vaidya and Rajgopal Patil for their help in the field work. Nikhil Dandekar also helped in morphometry.

 

 

 

 

Abstract: Species diversity of South Asian burrowing frogs within the genus Sphaerotheca is still obscure because of lack of taxonomic review. There are taxonomic issues within the genus Sphaerotheca with several names being synonymized to S. breviceps and names, such as S. pluvialis, being available without proper diagnosis. To resolve the taxonomic confusion, we describe and diagnose S. pluvialis by designating a neotype from Carnatic region of southern India, from where the original species was described, and make S. fodiens as its objective synonym. We designate lectotype of S. breviceps and provide diagnosis for separating the species from its congeners. We clarify the identity of S. dobsonii and resurrect S. swani and S. maskeyi. Based on morphology we define three groups within genus Sphaerotheca and provide key to the groups and eight valid species in the genus. We assign genetic barcodes to S. pluvialis, S. dobsonii and S. breviceps.

 

 

 

Keywords: Amphibia, molecular phylogeny, S. breviceps, S. dobsonii, S. fodiens, S. maskeyi, S. swani, taxonomy.

 

 

 

 

 

Article

 

INTRODUCTION

 

 

South Asian burrowing frogs have a long taxonomic history. Schneider (1799) described Rana breviceps from eastern India, followed by Gravenhorst (1829) who described Rana variegata, without providing the type locality. Tschudi (1838) transferred Rana breviceps to genus Systoma. Subsequently, Jerdon (1853) described Pyxicephalus fodiens, P. rufescens and P. pluvialis from southern India and Sri Lanka. Günther (1859) proposed a new genus Sphaerotheca while describing S. strigata from Madras, southern India, which was later synonymized to the genus Tomopterna by Günther (1860). Günther (1864) provided the first revision of South Asian burrowing frogs and placed the species under Genus Pyxicephalus. He considered two valid species, namely Pyxicephalus breviceps and P. rufescens, with Sphaerotheca striaga, P. fodiens and P. pluvialis as synonyms of P. breviceps. Boulenger (1882) shifted the two valid species to Rana and described a new species Rana dobsonii. Murray (1884) described Tomopterna strachani which was included in Rana by Boulenger (1890). Boulenger (1920) retained all the valid species under Rana subgenus Tomopterna owing to their burrowing habit. Rao (1937) described Rana (Tomopterna) leucorhynchus from southern India; Myers and Leviton In Leviton et al. (1956) described Rana swani from Nepal, and Dubois (1983) described Rana (Tomopterna) breviceps rolandae from Sri Lanka. Dubois (1987) defined tribe Tomopternini under family Ranidae and considered only two species valid from South Asia, namely Tomopterna (Sphaerotheca) breviceps and T. (S.) rolandae while he considered other species, namely Rana dobsonii, Pyxicephalus fodiens, Rana leucorhynchus, Pyxicephalus pluvialis, Tomoptema strachani, Sphaerotheca strigata, Rana swani and Rana variegata, as ‘incertae sedis et synonymes’ without providing any rationale for doing so. He also transferred Rana (Tomopterna) rufescens under Limnonectes (Fejervarya). Subsequently, Schleich & Anders (1998) described Tomopterna maskeyi from Nepal. Marmayou et al. (2000) and Vences et al. (2000) provided genetic support to raise the subgenus Sphaerotheca to genus level. Vences et al. (2000) considered seven species from South Asia valid under Sphaerotheca, namely S. breviceps, S. dobsonii, S. leucorhynchus, S. maskeyi, S. rolandae, S. strachani and S. swani.

Major confusion regarding the valid species under genus Sphaerotheca was due to the following publications—Dubois (1999, 2000, 2004) made the name Sphaerotheca pluvialis available without properly resurrecting the species as he did not provide diagnostic characters to identify it and separate it from other available names. Further, Dubois (1999) synonymized Tomopterna maskeyi and Dubois (2000) synonymized Rana variegata, Pyxicephalus fodiens, Sphaerotheca strigata, Rana dobsonii, Tomopterna strachani, Rana (Tomopterna) leucorhynchus and Rana swani to Sphaerotheca pluvialis without providing any justification. In addition, Marmayou et al. (2000) provided genetic sequence for Sphaerotheca pluvialis, from Myanmar, without clearly defining the species. Moreover, the type locality of S. pluvialis by original description is southern India and not Myanmar, which are biographically well separated areas.

Therefore, in this communication we try to resolve the taxonomic confusions in available names under Sphaerotheca. We validate the resurrection of S. pluvialis by Dubois (1999), by designating a neotype. We designate a lectotype for S. breviceps and clarify the identity of S. dobsonii, S. swani and S. maskeyi. We define morphological groups within the genus Sphaerotheca and provide a key to the groups and species. Further, we assign genetic barcodes to S. pluvialis, S. dobsonii and S. breviceps.

 

 

 

MATERIALS AND METHODS

 

 

 

Specimen studied and museum details

Specimens in the Indian museum collections of Bombay Natural History Society (BNHS, Mumbai), the Zoological Survey of India (ZSI, Kolkata), the Zoological Survey of India, Western Regional Centre (ZSI-WRC, Pune), the Wildlife Information Liaison Development (WILD, Coimbatore), and the Institute of Natural History Education and Research (INHER, Pune), were studied. Photographs of syntypes of Sphaerotheca breviceps were studied from the Museum für Naturkunde (ZMB), Berlin, Germany. Photographs of holotype of Sphaerotheca swani were studied from California Academy of Sciences, Stanford University collection (CAS-SU), San Francisco, USA. Photographs of holotype of Sphaerotheca maskeyi were studied from Zoologische Staatssammlung München, Germany (available online at http://www.biologie.uni-ulm.de/cgi-bin/herbar.pl?herbid=109475&sid=T&lang=d, accessed on 17 February 2017). Photographs of holotype of Sphaerotheca rolandae were studied from the Natural History Museum (BMNH), London, UK.

 

Morphometry

Measurements were taken to the nearest 0.1mm using a digital caliper (Ocean Premium measuring instruments) and include: Length of specimen from snout to the visible tip of urostyle (SUL), head length (HL), head width (HW), nasal snout distance (SN), inter nasal distance (IN, measured between the centre of the nares), diameter of the eye (EL), eye snout distance (SL), eye to nasal distance (EN), shortest distance between eyes (IUE), upper eyelid width (UEW), tympanum diameter (TYD), distance from tympanum to the back of the eye (TYE), length of hand (HAL), F1 to F4 (Finger 1 to Finger 4 length from the base of the sub-articular tubercle), length of forelimb (FLL), length of femur (FL), length of Tibia (TL), foot length (FoL, measured from the base of the inner metatarsal tubercle to the tip of the toe), T1 to T5 (Toe1 to Toe5 length from the base of the respective sub-articular tubercle), and Inner metatarsal tubercle length (IMT). Webbing formula was determined following the method provided by Savage & Heyer (1967) with modifications by Myers & Duellman (1982).

 

Statistical analysis

All characters showed positive linear correlation with SUL. Therefore, to remove size bias, statistical analysis of the morphometric data was performed on size adjusted measurements by taking all measurements as percent of SUL. Multivariate normality of the data was checked using Doornik & Hansen (2008) omnibus. Discriminant Analysis (DA) was performed to understand whether related species form significantly different clusters (Huberty & Olejnik 2006) in the genus Sphaerotheca. Wilks’ Lambda statistic was used to test the null hypothesis that the mean vectors of different clusters are equal (Harris 2001). Mahalanobis distances (Harris 2001) between pair of individuals were calculated and were used for computing Fisher’s distances (distance between the centroids of the clusters, divided by the sum of their standard deviations) between two clusters to check if the clusters were significantly different. Statistical analysis was performed in PAST 3.12 (Hammer et al. 2001).

 

Molecular analysis

Thigh muscle tissue was harvested from two specimens of putative topotypes of S. breviceps (BNHS 6005, WILD-16-AMP-645), two specimens of S. breviceps from Maithon, Jharkhand (BNHS 6006 and WILD-16-AMP-647), three specimen of S. dobsonii from Tamhini, Maharashtra (BNHS 6007, INHER-Amphibi-86, ZSI-WRC A/1548), two specimens of S. dobsonii from Devi-Hasol, Maharashtra (BNHS 6008, WILD-16-AMP-651) and one specimens of S. dobsonii from Bankot, Maharashtra (WILD-16-AMP-653). We could not collect topotypes for Sphaerotheca leucorhynchus for molecular analysis despite our best efforts for three consecutive years (2013 to 2016) to visit the type locality of the species. Further, we could not study topotypes of S. maskeyi, S. rolandae, S. swani and S. strachani for molecular analysis because of logistic and legal reasons due to their presence outside India. Tissues were preserved in absolute ethanol. DNA extraction, PCR amplification of 16S rRNA gene and sequencing protocols followed Padhye et al. (2014). Sequences were analysed by the BLAST tool (Altschul et al. 1990) for similar sequences in NCBI Genbank database (www.ncbi.nlm.nih.gov). Sequences generated in the current study are deposited in GenBank under the accession numbers (KY215969–KY215978). Additional 16S gene sequences were retrieved from the NCBI GenBank database. Gene sequences were aligned using MUSCLE (Edgar 2004). Pair-wise raw phylogenetic distances were calculated in MEGA6 (Tamura et al. 2013). Maximum likelihood phylogenetic tree was built using IQ-TREE software (Nguyen et al. 2015) where the best nucleotide substitution model was analysed based on the minimum Bayesian Information Criterion (BIC) value (Schwarz 1978; Nei & Kumar 2000). Reliability of the phylogenetic tree was estimated with ultrafast bootstrap support (Minh et al. 2013) for 1000 iterations. Phylogenetic tree was edited in FigTree v1.4.2 (Rambaut 2009).

Comparative material and data sources

Sphaerotheca breviceps: Lectotype, ZMB 55005, male (29.9mm SUL) and paralectotype, ZMB 3351 (33.9mm SUL), Indes orientales (only photographs examined); BNHS 6004, male (29.6mm SUL), BNHS 6005, female (33.5mm SUL), WILD-16-AMP-645, male (31.6mm SUL), WILD-16-AMP-646, female (34.8mm SUL), ZSI-WRC A/1546, male (31.7mm SUL), ZSI-WRC A/1547, female (30.0mm SUL), INHER-AMPHIBIA-46, female (34.4mm SUL) and INHER-AMPHIBIA-49, male (27.8mm SUL), 15.xi.2015, India: Tamil Nadu: Nagapattinam District, Tranquebar (=Tharangambadi), Karaikal (11.0620N, 79.8130N, elevation 16m), coll. N. Dandekar and S. Sulakhe; WILD-16-AMP-647, sub adult (26.6mm SUL) and BNHS 6006, sub adult (24.8mm SUL), 26-ix-2014, India: Jharkhand: Dhanbad District, Maithon (23.7760N, 86.8090E, 150m), coll. A.D. Padhye; ZSI 18744, male (48.3mm SUL), Madras, coll. J. Henderson.

Sphaerotheca dobsonii: WILD-16-AMP-648, female (43.0mm SUL), ZSI-WRC A/1548, female (38.0mm SUL) and BNHS 6007, female (45.5mm SUL), 1-vi-2000, India: Maharashtra: Pune District, Tamhini (18.4770N, 73.4270E, 897m), coll. A. D. Padhye; INHER-Amphibia-86, male (55.1mm SUL), 25-vi-2016, India: Maharashtra: Pune District, Tamhini (18.4770N, 73.4270E, 897m), coll. N. Dandekar & M. Ketkar; INHER-Amphibia-151, female (35.0mm SUL), WILD-16-AMP-651, female (34.3mm SUL) and BNHS 6008, female (42.5mm SUL), 31-vii-2016, India: Maharashtra: Ratnagiri District, Devi-Hasol (16.7420N, 73.4320E, 159m), coll. A.D. Padhye; 5 ex., INHER-Amphibia-144, female (37.0mm SUL), WILD-16-AMP-652, female (41.5mm SUL), WILD-16-AMP-653, female (46.0mm SUL), BNHS 6009, female (42.9mm SUL) and BNHS 6010, female (31.7m SUL), 29.vii.2016, India: Maharashtra: Ratnagiri District, Bankot (17.9800N, 73.0650E, 46m), coll. S. Sulakhe et al. Additional data from Boulenger (1920).

Sphaerotheca swani: Holotype, CAS-SU 15371, male (42.3mm SUL), 22.vi.1954, Dharan, eastern Nepal, at an altitude of 1,000 feet, coll. L.W. Swan (only photographs examined). Additional data from Leviton et al. (1956).

Sphaerotheca maskeyi: Holotype, ZSM 106/91-2, 8.vii.1991, Chitwan Jungle Lodge, Royal Chitwan National Park, Central Nepal, at an altitude of approx. 300m, coll. H.H. Schleich and T.M. Maskey (only photographs examined); ZSI 16127, female, 43.7mm SUL, Chinpore (=Chainpur) Nepal. Additional data from Schleich & Anders (1998).

Sphaerotheca rolandae: Holotype, BMNH 1973.3024, Kurunegala, Sri Lanka, altitude 60m, latitude 7029′N, longitude 80022′E (only photographs examined); ZSI 17630 and ZSI 17632, Bangalore, India, coll. C.R. Narayan Rao (only photographs examined). Further details from Dubois (1983) and Dutta & Manamendra-Arachchi (1996).

Data for Sphaerotheca leucorhynchus from Rao (1937) and S. strachani from Murray (1884). Among the current synonyms of S. breviceps, data for Rana variegata was obtained from Gravenhorst (1829) and Sphaerotheca strigata from Günther (1859) and Boulenger (1920).

 

 

RESULTS AND DISCUSSION

 

 

Identity of Sphaerotheca pluvialis (Jerdon, 1853)

Jerdon (1853) made two names available, namely Pyxicephalus fodiens and P. pluvialis, from Carnatic region of southern India based on scanty descriptions. Nevertheless, he acknowledged that the two species are very similar and preferred to call them different just based on the different vernacular names used by the natives. Günther (1859) further mentioned that he has seen the “coloured figures of Mr. Jerdon’s Pyxicephalus fodiens and P. pluvialis, in the possession of Walter Elliot, Esq., from which it is evident that they are identical”. Thus, it can be suggested that Pyxicephalus fodiens and P. pluvialis are indeed the same species. Although, the name P. fodiens appears before P. pluvialis, Dubois (2000) gave preference for the name P. pluvialis over P. fodiens as the first reviser, owing to slightly more detailed original description of P. pluvialis. We designate ZSI A9074 as a neotype of both P. fodiens and P. pluvialis and consider P. pluvialis as a valid species under the genus Spherotheca with P. fodiens as its objective synonym. It is also essential to note that the neotype is collected from within the Carnatic region, the type locality of both the species, which is the part of southern India that includes Tamil Nadu, south-eastern Karnataka, north-eastern Kerala and southern Andhra Pradesh in modern India. We choose this specimen ZSI A9074 because it also has 16S rRNA gene sequence AF215418, which is the genetic barcode for the neotype of Spherotheca pluvialis. Designation of a neotype is necessary for three reasons. First, there are no known types for P. fodiens and P. pluvialis and the types were reported as lost (Jerdon, 1870) and colored figures of the two species, as mentioned by Günther (1859), are also not traceable. Second, the original description is not adequate to identify and diagnose the species. Third, the species has often been confused with other sympatric congeners in the past.

 

 

Sphaerotheca pluvialis (Jerdon, 1853)

(Image 1)

Synonyms:

Pyxicephalus pluvialis Jerdon (1853): p. 534

Pyxicephalus fodiens Jerdon (1853): p. 534 (objective synonym)

Sphaerotheca strigata Günther, 1859: p. 20, Plate II fig. A

Pyxicephalus (Tomopterna) fodiens—Peters (1860, p. 186)

Tomopterna (Sphaerotheca) pluvialis—Dubois (2000, p. 334)

Sphaerotheca dobsonii (non Boulenger, 1882)— Vences et al. (2000, p. 240)

Tomopterna (Sphaerotheca) breviceps (non Schneider, 1799)—Bossuyt & Milinkovitch (2000, p. 6586)

Suggested common name: Jerdon’s Burrowing Frog

Neotype: ZSI A9074, 10.vi.1998, female, 45.1mm SUL, India: Tamil Nadu: Thiruadisoolam near Pattaravakkam (12.6960N, 80.0300E), coll. Romulus Whitaker.

Comparative material: ZSI 18743, female, 51.9mm SUL, Madras, coll. J. Henderson; ZSI 12515, male, 46.5mm SUL, southern India, coll. E. Gerard; ZSI 2681, female, 45.4mm SUL, Sri Lanka, coll. E.F. Kelaart.

Genetic barcode: 16S rRNA sequence AF215418 of the neotype ZSI A9074.

Diagnosis: Sphaerotheca pluvalis differs from all other congeners based on the following combination of characters: medium sized frog with slightly warty dorsum; snout as long as eye diameter; tympanum large, vertically oval, its horizontal diameter less than ½ of the eye diameter; inter-narial width slightly greater than inter orbital distance, which is less than the width of upper eyelid; first finger longer than second and fourth; first finger almost equal to or slightly less than third finger; second finger longer than fourth; outer metatarsal tubercle absent; tibio-tarsal tubercle absent; inner metatarsal tubercle elongated, crescentic, more than 1.5 times but less than two times in length as compared to length of first toe; heels do not touch each other when legs are folded at right angles to the body axis; tibio tarsal articulation barely reaching the tympanum; and webbing formula I12+II1+-3III2+-3½IV3--2V (See species groups and species comparisons section below for species wise comparison.)

Description of the Neotype (ZSI A9074, female, Image 1) (all measurements in mm): Medium sized frog (SUL 45.1); head wider than long (HW 18.8 > HL 15.1); snout shorter than horizontal diameter of eye (SL 6.3 < EL 6.6); outline of snout rounded dorsally, truncated laterally; snout slightly protruding beyond the mouth ventrally; nostrils nearer to eye than to snout (SN 3.6 > EN 3.2); tympanum less than ½ the diameter of eye (TYD = 2.5; EL= 6.6); supra-tympanic fold distinct; upper eyelid width about rd the horizontal diameter of eye(UEW = 4.2; EL= 6.6); upper eyelids warty; Inter narial width slightly greater than shortest distance between eyes (IN 3.7 > IUE 3.6); canthus rostralis distinct; loreal region slightly concave and oblique; buccal cavity shallow, vomerine teeth in two sharply oblique rows at the anterior border of choanae; tongue thin, bifid, without papilla; dorsal skin granulated or slightly warty; ventral and lateral skin smooth.

Length of Forelimb slightly greater than length of hand (FLL 10.8 > HAL 10.5); finger lengths from shortest to longest—F4 (3.4) < F2 (4.1) < F1 (5.9) < F3 (6.1); palmar tubercles present, outer palmar tubercle single, sub-articular tubercles moderate, supernumerary tubercles present, single; fingers without web or fringe of skin.

Hind limbs long; femur longer than tibia (FL 19.1 > TL 17.4); foot longer than tibia (FoL 18.3 > TL 17.4); toe lengths from shortest to longest are—T1 (2.8) < T2 (5.8) < T3 (9.2) < T5 (9.8) < T4 (14.6); inner metatarsal tubercle large, very prominent, compressed, inserted obliquely at the base of the first toe, which it exceeds in length—T1 (2.8) < IMT (4.0); outer metatarsal tubercle absent; supernumerary tubercles absent; sub-articular tubercles moderate; tarsal fold and outer phalangeal fringe absent; webbing formula I1-2II1-3III2-3½IV4-2V.

Coloration: In alcohol preservation (Image 1), creamish-brown above with dark brown patches, dorso-lateral creamish patches on shoulders and above supratympanic fold, mid-dorsal creamish line starting from middle of eyes till vent; inverted V-shaped mark on the back interrupted by mid-dorsal line; creamy white triangular patch starting from snout to middle of the eyes followed by a dark band continuing on the upper eyelids on either side; limbs cross barred, canthal region dark brown with two vertical creamy white bands starting from anterior as well as posterior margins of the eye extending to upper jaw, posterior band covers anterior portion of tympanum, tympanum dark brow in posterior half; supra tympanic fold distinctly marked with dark brown edge; ventral side creamy brown with mottling on throat.

Variation: Morphometric variation is provided in Table 1. Morphological and color variation as per Image 2.

Morphometric analysis: Size corrected morphometric data was not significantly different from multivariate normal (Ep = 55.44, P = 0.3464). Sphaerotheca breviceps, S. dobsonii and S. pluvialis formed distinct clusters in DA (Fig. 1). Sphaerotheca pluvialis formed a distinct cluster from S. brevceps (F = 3824.71, P = 0.004) based on longer toes and smaller femur, inter orbital distance, forelimb and head width (Fig. 1). Sphaerotheca pluvialis formed a distinct cluster from S. dobsonii (F = 1694.35, P = 0.019) based on longer toes and smaller thigh, foot and inter narial distance (Fig. 1).

Genetic analysis: Model test suggested transition model with gamma distribution (TIM2e+G4, -lnL = 1686.798, df = 31, BIC = 3570.69) as the best fit nucleotide substitution model for the data. Sphaerotheca pluvialis formed a distinct clade different from both, the topotypic S. breviceps and S. dobsonii (Fig. 2). Sphaerotheca pluvialis differed from S. breviceps by 7.9–8.3 % raw genetic distance, while it differed from S. dobsonii by 5.6–8.0 % (Table 2). Specimens identified as S. pluvialis by Marmayou (2000) and S. breviceps by Frost et al. (2006) from Myanmar are conspecific to each other but differ from S. pluvialis by 7.1–7.5 % raw genetic distance and from S. breviceps by 9.1–9.6 % indicating that they are neither S. pluvialis nor S. breviceps and they may belong to one of the other available names under Sphaerotheca. Gene sequence AF249042 of a specimen identified as S. breviceps from Sri Lanka by Bossuyt & Milinkovitch (2000) is similar to the sequence of neotype of S. pluvialis. We have not considered five sequences AF249014, DQ346976, AY880442 and GU19112 or specimens identified as Sphaerotheca for genetic analysis because they either did not align properly with other sequences of Sphaerotheca or were not of good quality. Further, we have not considered GU191123 labelled as Sphaerotheca sp. because the sequence is similar to Hoplobatrachus tigerinus and not species of Sphaerotheca.

Distribution: While both Pyxicephalus fodiens and P. pluvialis were described from Carnatic region of India, Jerdon (1853) mentioned that Pyxicephalus fodiens was also present in Ceylon (= Sri Lanka). Interestingly, we found a specimen in the collection of Zoological Survey of India (ZSI 2681) collected from Sri Lanka, which is conspecific with S. pluvialis. Further, 16S rRNA gene sequence AF249042 of a specimen identified as S. breviceps from Sri Lanka by Bossuyt & Milinkovitch (2000) is similar to the sequence of neotype of S. pluvialis, which further vouch for the presence of S. pluvialis from Sri Lanka. Based on the neotype, three comparative specimens and genetic evidence we suggest that S. pluvialis is present in southern India and Sri Lanka.

Remarks: Dubois (2000) used the name Tomopterna (Spherotheca) pluvialis and Dubois (1999; 2004) and Marmayou et al. (2000) used the name S. pluvialis without actually discussing the proper resurrection of the species with detailed diagnosis to identify it. Although Dubois (1999; 2000) referred to a manuscript in preparation that will clarify the identity of S. pluvialis, such a manuscript has not been published till date. Use of a specimen from Myanmar for genetic analysis of S. pluvialis by Marmayou et al. (2000) is certainly wrong, because of two reasons. First, there was no diagnosis for the species and therefore the identity of the species was not clear, and second, S. pluvialis was definitely described from southern India, so a specimen from Myanmar cannot be considered as conspecific with S. pluvialis until it is proven that the species is also distributed in southern India. The neotype of S. pluvialis described in this study is from Carnatic region and is distinctly different from other valid species (see Species groups and species comparisons section below).

It is essential to note that the type locality of S. pluvialis is just south of Madras, the type locality of Sphaerotheca strigata described by Günther (1859). We also found a specimen (ZSI 18743) from Madras that is conspecific to S. pluvialis neotype. We were not able to examine the types of S. striagata; however, the original description and morphometry of types provided by Boulenger (1920) clearly indicate that S. strigata is conspecific with S. pluvialis neotype. We do not use the available name S. strigata and prefer to designate the neotype of S. pluvialis for following reasons. First, S. pluvialis has precedence over S. strigata. Second, Günther (1859) mentions that types if S. strigata were collected by J.C. Jerdon creating the possibility that they were from the same collection that Jerdon (1853) used for describing Pyxicephalus fodiens and P. pluvialis. Third, Günther (1864) himself synonymized Pyxicephalus fodiens, P. pluvialis and S. strigata with S. breviceps, making all three names available for the species that will be resurrected from S. breviceps complex. Thus, we consider S. strigata as a junior subjective synonym of S. pluvialis, maintaining the synonymy suggested by Dubois (2000).

 

 

 

 

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Resurrection of Sphaerotheca maskeyi (Schleich & Anders, 1998)

Dubois (1999) treated S. maskeyi as a synonym of S. pluvialis without providing any rationale for the same. We examined the photographs of the holotype of S. maskeyi, as well as a specimen from Nepal (ZSI 16127, Image 3) that matches the description of the species and was collected just south of the type locality. On the basis of detailed description of the species provided by Schleich & Anders (1998) and study of the holotype and comparative material we suggest that the species is valid and can be diagnosed as follows.

Sphaerotheca maskeyi can be distinguished from all its congeners based on the combination of following characters: tympanum about ½ of the eye diameter; interorbital width less than upper eyelid width; snout to nostril distance more than half of eye diameter; nostril nearer to eye than to snout; finger 2 length more than or equal to finger 4 length; finger 1 length more than finger 3 length; tibio tarsal articulation reaching tympanum; outer metatarsal tubercle absent; tibio tarsal tubercle absent; length of inner metatarsal tubercle more than 2 times the inner toe length; and webbing formula I1--2+II1½-3+III2+-3½IV3½-2+V (See Species groups and species comparisons section below for species wise comparison).

 

 

Resurrection of Sphaerotheca swani (Myers & Leviton in Leviton et al., 1956)

Dubois (2000) treated S. swani as a synonym of S. pluvialis without providing any justification. We examined the photographs of the holotype of S. swani (Image 4). On the basis of detailed description of the species provided by Leviton et al. (1956) and study of the holotype we suggest that the species is valid and can be diagnosed as follows.

Sphaerotheca swani can be distinguished from all its congeners based on the combination of following characters: inter orbital distance about ¾ of upper eyelid width; tympanum large, about rd diameter of eye; first finger equal to third; second finger shorter than first and third; fourth finger shortest; tibio tarsal articulation reaching centre of eye; outer metatarsal tubercle absent; tibio tarsal tubercle absent; length of inner metatarsal tubercle slightly more or equal to inner toe length; and webbing formula I1--2II1+-3III2+-3½IV3½-2+V (See Species groups and species comparisons section below for species wise comparison).

 

 

Identity of Sphaerotheca breviceps

Several species have been considered under the synonymy of Sphaerotheca breviceps because of lack of diagnostic characters that separates S. breviceps from its congeners. Schneider (1799) does not provide a precise type locality for the species; however, he mentions that the specimens were obtained from eastern India. Because the type locality of S. pluvialis is also in eastern coast of southern India, based on the neotype designation, there is a need to clarify the identity of S. breviceps. There are two extant syntypes of S. breviceps, namely ZMB 3351 and ZMB 55005. For clarifying taxonomy we designate the lectotype from among the syntypes; however, as there is no illustration or special mention of one of the syntypes, we designate the male ZMB 55005 (Image 5) as the lectotype of S. breviceps and consider the other specimen ZMB 3351 as the paralectotype. Dubois (1983) suggests that these specimens probably came from Tranquebar (=Tharangambadi), Tamil Nadu, India. We collected several specimens of the species from Tharangambadi and they closely resemble the lectotype based on the diagnostic characters described below. Our specimens from Tharangambadi, therefore, can be considered as putative topotypes of S. breviceps and 16S rRNA sequences from these topotypes, namely KY215977 and KY215978, as the genetic barcode for the species.

Sphaerotheca breviceps can be diagnosed from all its congeners based on the combination of following characters: tympanum small, less than ½ of the eye diameter; interorbital width more than or equal to upper eyelid width; snout to nostril distance less than half of eye diameter; nostril nearer to snout than to eye; finger 2 length more than finger 4 length; finger 1 more than finger 2 length but less than finger 3 length; outer metatarsal tubercle absent; tibio tarsal tubercle absent; length of inner metatarsal tubercle more than 2 times the inner toe length; and webbing formula I1--2-II1½-2½III2½-3½IV4--2+V (See Species groups and species comparisons section below for species wise comparison).

We examined a specimen of S. breviceps (ZSI 18744) collected from Madras (Image 6). This suggests that both S. breviceps and S. pluvialis are probably sympartic. We examined two specimens from Jharkhand (BNHS 6006 and WILD-16-AMP-647), which are morphologically similar to S. breviceps. Although these specimens formed a distinct clade (Fig. 2), their raw genetic distance from topotypic S. breviceps was low (1.8–2.2 %). Therefore, S. breviceps is probably widespread along the eastern coast of India.

We tentatively retain Rana variegata Gravenhorst, 1829 under the synonymy of S. breviceps because we could not trace the type specimens and the figure as well as the description are not enough to diagnose the species. Further, there is no type locality mentioned for Rana variegata. Nevertheless, we think that the type of the species could be present in the museum collection of Natural History Museum (UWZM), Wroclaw University. This is because, for another species described by Gravenhorst (1829), namely Rana gracilis, it was speculated that Gravenhorst did not collect specimens (Biju et al. 2014); however, Padhye et al. (2015) mention a specimen of R. gracilis in the collection of UWZM collected by Gravenhorst. We have already stated that specimens misidentified as S. pluvialis and S. breviceps from Myanmar are not conspecific to either S. breviceps or S. pluvialis. For these specimens, Rana variegata is an available name. It is also essential to note that several of the species described by Gravenhorst (1829) came from Southeast Asia, so a further enquiry into Sphaerotheca from Myanmar is essential to resolve the issue with the identity of Rana variegata Gravenhorst, 1829.

 

 

 

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Identity of Sphaerotheca dobsonii

Dubois (1987) considered Rana dobsonii Boulenger, 1882 as ‘incertae sedis et synonymes’ under the genus Tomopterna subgenus Sphaerotheca without providing any rationale for the same. Similarly, Dubois (2000) placed Rana dobsonii under the synonymy of Tomopterna (Sphaerotheca) pluvialis, a species which he never properly resurrected by providing diagnostic characters to identify it. This makes it important to clarify the identity of Sphaerotheca dobsonii. We were not able to examine the holotype of Sphaerotheca dobsonii; however, we examined several specimens of the species from wide distributional range in Western Ghats of Maharashtra. Our specimens (for example see Image 7) closely resemble the type description and illustration provided by Boulenger (1882) and morphometry of type specimen provided by Boulenger (1920). Further, Kotaki et al. (2008) and Hasan et al. (2014) provided 16S rRNA gene sequences (AB277305 and AB530608 respectively) for topotypic Sphaerotheca dobsonii. These sequences can be considered as genetic barcode for identifying S. dobsonii. Our specimens are genetically closer to these sequences further confirming the species identity. Since S. dobsonii differs from S. breviceps with a raw genetic distance of 7.1–10.9 % and from S. pluvialis with 5.6–8.0 %, it is obvious that S. dobsonii is a genetically distinct species. Morphologically, S. dobsonii can be diagnosed from its congeners as follows.

Sphaerotheca dobsonii differs from all other congeners based on a combination of characters including: snout equal to or longer than eye diameter; tympanum large, vertically oval, its horizontal diameter less than or equal to half of the eye diameter; inter-narial width greater than inter orbital distance; first finger length less than third finger length; second finger longer than fourth; outer metatarsal tubercle absent; tibio-tarsal tubercle absent; inner metatarsal tubercle elongated, crescentic, more than 1.5 times but less than or equal to two times in length as compared to length of first toe; tibio tarsal articulation barely reaches eye; heels just touch each other when legs are folded at right angles to the body axis; and webbing formula I1½-2+II1½-3+III2½-4-IV4--2+V (See Species groups and species comparisons section below for species wise comparison).

 

 

 

 

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Species groups and species comparisons

Based on morphological comparison of the species under Sphaerotheca, we recognize three morphological groups, namely Breviceps group, Dobsonii group and Leucorhynchus group. Diagnostic characters for each group and species comparison within each group are provided below.

Breviceps group: Outer metatarsal tubercle absent and length of inner metatarsal tubercle more than two times the inner toe length. Included species are Sphaerotheca breviceps (Schneider, 1799), S. maskeyi (Schleich & Anders, 1998) and S. rolandae (Dubois, 1983). Sphaerotheca breviceps differs from S. maskeyi in tympanum length less than half of eye diameter (vs. more than or equal to half), interorbital distance more than or equal to upper eyelid width (vs. less than upper eyelid width), snout to nostril less than half of eye diameter (vs. more than half of eye diameter), nostril closer to snout than to eye (vs. closer to eye than snout) and finger 1 length less than finger three length (vs. more than finger three length). Sphaerotheca rolandae differs from both S. breviceps and S. maskeyi in the presence of tibiotarsal tubercle (vs. absent) (see Image 8). Further, all three species have different webbing extent on foot (Fig. 3).

Dobsonii group: Outer metatarsal tubercle absent and length of inner metatarsal tubercle less than or equal to two times the inner toe length. Included species are Sphaerotheca dobsonii (Boulenger, 1882), S. pluvialis (Jerdon, 1853) and S. swani (Myers & Leviton in Leviton et al., 1956). Sphaerotheca dobsonii differs from S. pluvialis in tibio tarsal articulation reaching eye (vs. barely reaching tympanum), heels just touch each other when legs are folded at right angles to the body axis (vs. do not touch); and finger 1 length much lesser than finger 3 length (vs. finger 1 length equal to or slightly lesser than finger 3 length). Further, S. dobsonii is genetically distinct from S. pluvialis (Fig. 2) with raw genetic distance of 5.6–8.0 %. Sphaerotheca swani differs from both S. dobsonii and S. pluvialis in tympanum large about rd of eye diameter (vs. less than or equal to ½), first finger is equal to or longer than third (vs. less than third), inner metatarsal tubercle equal to first toe length (vs. more than 1.5 times first toe length) and tibio tarsal articulation reaching middle of eye (vs. barely reaching tympanum in S. pluvialis and reaching eye in S. dobsonii). Further, all three species have different webbing extent on foot (Fig. 3).

Leucorhynchus group: Can be diagnosed from Breviceps and Dobsonii groups by presence of outer metatarsal tubercles. Included species are Sphaerotheca leucorhynchus (Rao, 1937) and S. strachani (Murray, 1884). Sphaerotheca leucorhynchus differes from S. strachini in outer metatarsal tubercle small (vs. conspicuous), first finger longer than second (vs. first finger nearly equal to second) and tympanum smaller than eye (vs. as large as eye).

Despite extensive surveys in the type locality of S. leucorhynchus in Wattakole, Coorg in Karnataka for three consecutive years (2013–2016), we were unable to record a species of Sphaerotheca resembling the description or the figure provided by Rao (1937). Further surveys to locate this species are essential to clarify its identity. Presence of outer metatarsal tubercles makes this species unique from other species of Sphaerotheca, except S. strachani.

 

 

Key to groups and species of the genus Sphaerotheca

Based on the comparative account provided above, a key to the groups and valid species of Sphaerotheca is provided.

 

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CONCLUSIONS

 

 

We designate, diagnose and describe the neotype of Sphaerotheca pluvialis from Carnatic region, which is a type locality mentioned in the original description of the species. We resurrect S. maskeyi and S. swani with characters to diagnose the species. We clarify the identity of S. breviceps by designating a lectotype, and of S. dobsonii, and provide diagnosis for the two species. Based on our study we suggest that genus Sphaerotheca has three species groups with eight valid species, viz., Breviceps group: S. breviceps (Schneider, 1799), S. maskeyi (Schleich & Anders, 1998), and S. rolandae (Dubois, 1983); Dobsoni group: S. dobsonii (Boulenger, 1882), S. pluvialis (Jerdon, 1853), and S. swani (Myers & Leviton in Leviton et al., 1956); and Leucorhynchus group: S. leucorhynchus (Rao, 1937), and S. strachani (Murray, 1884). We consider Sphaerotheca strigata Günther, 1859 as a junior subjective synonym of S. pluvialis and Rana variegata Gravenhorst, 1829 as a junior subjective synonym of S. breviceps. We provide genetic delimitation of three species, namely S. breviceps, S. dobsonii and S. pluvialis, based on topotypes and type specimen.

 

 

References

 

Altschul, S.F., W. Gish, W. Miller, E.W. Myers & D.J. Lipman (1990). Basic local alignment search tool. Journal of Molecular Biology 215: 403–410; http://doi.org/10.1016/S0022-2836(05)80360-2

Biju, S.D., S. Garg, S. Mahony, N. Wijayathilaka, G. Senevirathne & M. Meegaskumbura (2014). DNA barcoding, phylogeny and systematic of golden-backed frogs (Hylarana, Ranidae) of the Western Ghats-Sri Lanka biodiversity hotspot, with the description of seven new species. Contributions to Zoology 83(4): 269–335.

Bossuyt, F. & M.C. Milinkovitch (2000). Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Science 97(12): 6585–6590; http://doi.org/10.1073/pnas.97.12.6585

Boulenger, G.A. (1882). Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London.

Boulenger, G.A. (1890). The Fauna of British India, including Ceylon and Burma. Reptilia and Batrachia.   Taylor and Francis, London.

Boulenger, G.A. (1920). A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum 20: 1–226.

Doornik, J.A. & H. Hansen (2008). An Omnibus Test for Univariate and Multivariate Normality. Oxford Bulletin of Economics and Statistics 70: 927–939; http://doi.org/10.1111/j.1468-0084.2008.00537.x

Dubois, A. (1983). Note préliminaire sur le groupe de Rana (Tomopterna) breviceps Schneider, 1799 (Amphibiens, Anoures), avec diagnose d’une sous-expèce nouvelle de Ceylan. Alytes 2: 163–170.

Dubois, A. (1999). South Asian Amphibia: a new frontier for taxonomists. Journal of South Asian Natural History 4(1): 1–11.

Dubois, A. (2000). The influence of man on the distribution of amphibians in the Himalayas of Nepal: an example of critical evaluation of biogeographical data. In: Miehe, G. & Y. Zhang (eds.). Environmental changes in high Asia. Marburger Geographische Schriften 135: 326–345.

Dubois, A. (1987). Miscellanea taxinomica batrachologica (I). Alytes 5: 7–95.

Dubois, A. (2004). Amphibians of Nepal: a few words of caution. Alytes 21: 174–180.

Dutta, S.K. & K. Manamendra-Arachchi (1996). The amphibian fauna of Sri Lanka. Wildlife Heritage Trust of Sri Lanka.

Edgar, R.C. (2004). MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797; http://doi.org/10.1093/nar/gkh340

Frost, D.R., T. Grant, J. Faivovich, R.H. Bain, A. Haas, C.F.B. Haddad, R.O. de Sá, A. Channing, M. Wilkinson, S.C. Donnellan, C.J. Raxworthy, J.A. Campbell, B.L. Blotto, P.E. Moler, R.C. Drewes, R.A. Nussbaum, J.D. Lynch, D.M. Green & W.C. Wheeler (2006). The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370.

Gravenhorst, J.L.C. (1829). Deliciae Musei Zoologici Vratislaviensis. Fasciculus primus. Chelonios et Batrachia. Leipzig: Leopold Voss.

Günther, A.C.L.G. (1859). Catalogue of the Batrachia Salientia in the Collection of the British Museum. London: Taylor and Francis.

Günther, A.C.L.G. (1860). Contribution to the knowledge of the reptiles of the Himalaya mountains. Proceedings of the Zoological Society of London 1860: 148–175.

Günther, A.C.L.G. (1864). The Reptiles of British India.  London: Ray Society by R. Hardwicke.

Hammer, Ø., D.A.T. Harper & P.D. Ryan (2001). Past: Paleontological Statistics Software Package for education and data analysis. Paleontología Electrónica 4: 1–9. http://palaeo-electronica.org/2001_1/past/issue1_01.html

Harris, R.J. (2001). A primer for multivariate statistics. Third Edition. Lawrence Erlbaum Associates Publishers, London.

Hasan, M., M.M. Islam, M.R. Khan, T. Igawa, M.S. Alam, H.T. Djong, N. Kurniawan, H. Joshy, Y.H. Sen, D.M. Belabut, A. Kurabayashi, M. Kuramoto & M. Sumida (2014). Genetic divergences of South and Southeast Asian frogs: a case study of several taxa based on 16S ribosomal RNA gene data with notes on the generic name Fejervarya. Turkish Journal of Zoology 38: 389–411.

Huberty, C.J. & S. Olejnik (2006). Applied MANOVA and Discriminant Analysis. 2nd Edition. Hohn Wiley and Sons, N.J., 488pp; http://doi.org/10.1002/047178947X

Jerdon, T.C. (1853). Catalogue of reptiles inhabiting the Peninsula of India. Journal of the Asiatic Society of Bengal 22: 522–534.

Jerdon, T.C. (1870). Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal 1870: 66–85.

Kotaki, M., A. Kurabayashi, M. Matsui, W. Khonsue, T.H. Djong, M. Tandon & M. Sumida (2008). Genetic divergences and phylogenetic relationships among the Fejervarya limnocharis complex in Thailand and neighboring countries revealed by mitochondrial and nuclear genes. Zoological Sciences 25(4): 381–390; http://doi.org/10.2108/zsj.25.381

Leviton, A.E., G.S. Myers & L.W. Swan (1956). Zoological results of the California Himalayan Expedition to Makalu, eastern Nepal. 1. Amphibians and reptiles. Occasional Papers of the Natural History Museum of Stanford University 1: 1–18.

Marmayou, J., A. Dubois, A. Ohler, E. Pasquet & A. Tillier (2000). Phylogenetic relationships in the Ranidae (Amphibia, Anura). Independent origin of direct development in the genera Philautus and Taylorana. Comptes Rendus des Academie des Sciences 323: 287–297.

Minh, B.Q., M.A.T. Nguyen & A. von Haeseler (2013). Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30: 1188–1195; http://doi.org/10.1093/molbev/mst024

Murray, J.A. (1884). The Vertebrate Zoology of Sind. Richardson & Co., London; Education Society’s Press, Bombay.

Myers, C.W. & W.E. Duellman (1982). A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Novitiates 2752: 1–32.

Nei, M. & S. Kumar (2000). Molecular Evolution and Phylogenetics. Oxford University Press, New York.

Nguyen, L.T., H.A. Schmidt, A. von Haeseler & B.Q. Minh (2015). IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268–274; http://doi.org/10.1093/molbev/msu300

Padhye, A.D., A. Jadhav, N. Modak, P.O. Nameer & N. Dahanukar (2015). Hydrophylax bahuvistara, a new species of fungoid frog (Amphibia: Ranidae) from peninsular India. Journal of Threatened Taxa 7(11): 7744–7760; http://doi.org/10.11609/JoTT.o4252.7744-60

Padhye, A.D., N. Modak & N. Dahanukar (2014). Indirana chiravasi, a new species of leaping frog (Anura: Ranixalidae) from Western Ghats of India. Journal of Threatened Taxa 6(10): 6293–6312; http://doi.org/10.11609/JoTT.o4068.6293-312

Peters, W.C.H. (1860). Über einige interessante Amphibien, welche von dem durch seine zoologischen Schriften rühmlichst bekannten österreichischen Naturforscher Professor Schmarda während seiner auf mehrere Welttheile ausgedehnten, besonders auf wirbellose Thiere geichteten, naturwissenschaftlichen Reise, mit deren Veröffentlichung Hr. Schmarda gegenwartig in Berlin beschafigt ist, auf der Insel Ceylon gesammelt wurden. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin 1860: 182–186.

Rambaut, A. (2009). FigTree, ver. 1.4.2. [Online]. Available: http://tree.bio.ed.ac.uk/software/figtree/ Accessed on 28 February 2015.

Rao, C.R.N. (1937). On some new forms of Batrachia from S. India. Proceedings of the Indian Academy of Sciences Section B 6: 387–427; http://doi.org/10.1007/BF03051434

Savage, J.M. & W.R. Heyer (1967). Variation and distribution in the tree‐frog genus Phyllomedusa in Costa Rica, central America: With 6 figures. Studies on Neotropical Fauna and Environment 5(2): 111–131; http://doi.org/10.1080/01650526709360400

Schneider, J.G. (1799). Historia Amphibiorum Naturalis et Literarariae. Fasciculus Primus. Continens Ranas, Calamitas, Bufones, Salamandras et Hydros in Genera et Species Descriptos Notisque suis Distinctos. Jena: Friederici Frommanni.

Schleich, H.H. & C.C. Anders (1998). Tompterna maskeyi spec. nov. from Nepal (Amphibia, Anura). Volume 4: 57–72. In: Schleich, H.H. & W. Kästle (eds.), Contributions to the Herpetology of South Asia (Nepal, India). Veröffentlichungen aus dem Fuhlrott-Museum. Fuhlrott-Museum.

Schwarz, G. (1978). Estimating the dimension of a model. Annals of Statistics 6: 461–464.

Tamura, K., G. Stecher, D. Peterson, A. Filipski & S. Kumar (2013). MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30: 2725–2729; http://doi.org/10.1093/molbev/mst197

Tschudi, J.J. v. (1838). Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien.  Neuchâtel: Petitpierre.

Vences, M., F. Glaw, J. Kosuch, I. Das & M. Veith (2000). Polyphyly of Tomopterna (Amphibia: Ranidae) based on sequences of the mitochondrial 16S and 12S rRNA genes, and ecological biogeography of Malagasy relict amphibian groups, pp. 229–242. In: Lourenço, W.R. & S.M. Goodman (eds.). Memoires de la Société de Biogéographie. Paris, France, Société de Biogéographie.