Zoosyst. Evol. 97 (2) 2021, 451-470 | DO! 10.3897/zse.97.61770 yee BERLIN Lost, forgotten, and overlooked: systematic reassessment of two lesser-known toad species (Anura, Bufonidae) from Peninsular India and another wide-ranging northern species Karan Bisht', Sonali Garg’, A. N. D. Akalabya Sarmah!, Saibal Sengupta’, S. D. Biju! 1 Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India 2 School of life sciences, Assam Don Bosco University, Tapesia, Assam, India http://zoobank.org/C09D919D-FB05-4D9F-9F BC-D453F83FACBC Corresponding authors: S. D. Biju (Sdbiju.es@gmail.com); Sonali Garg (sgarg.du@gmail.com) Academic editor: Rafe Brown @ Received 9 December 2020 @ Accepted 30 August 2021 Published 21 September 2021 Abstract We rediscovered two species of toads, Bufo stomaticus peninsularis and Bufo brevirostris, which were described from Peninsular India 84 and 101 years ago, respectively, but have not been reported since. Because the name-bearing types of both species are either damaged or lost, we provide detailed redescriptions, morphological comparisons, and insights into phylogenetic relationships with closely related members of the genus Duttaphrynus sensu lato, based on new material from the type locality of each species. We clar- ify and validate the identity of D. brevirostris, which was rediscovered from multiple localities in the Malenadu and adjoining coastal regions of Karnataka. We also demonstrate that Bufo stomaticus peninsularis, which was considered a synonym of Duttaphrynus scaber, is a distinct species. Bufo stomaticus peninsularis differs from Duttaphrynus scaber morphologically and genetically, and is more closely related to members of the Duttaphrynus stomaticus group. We also clarify the identity of the namesake species of the Duttaphrynus stomaticus group, which is reported widely in India and neighbouring countries, but lacks sufficient taxonomic information due to its brief original description and reportedly untraceable type material. We located and studied the complete syn- type series of D. stomaticus, probably for the first time in over a century, and we report on the status of available specimens, provide detailed description of a potential type, compare it to related species, and clarify the species’ geographical range. Our molecular analyses suggest that D. stomaticus is minimally divergent from, and possibly conspecific with, D. olivaceus. Our analyses also clarify its relationship to the closely-related D. peninsularis comb. nov., with which it was previously confused. Finally, our study provides other insights into the phylogenetic relationships and genetic differentiation among various species of Duttaphrynus toads. Key Words Amphibia, Bufo stomaticus peninsularis, distribution, Duttaphrynus brevirostris, Duttaphrynus stomaticus group, Firouzophrynus, molecular phylogeny, redescription, rediscovery, taxonomy Introduction The genus Duttaphrynus sensu lato, comprising 26 rec- ognised Asian species, is a widely-distributed and com- monly-occurring group of toads, found at elevations from sea level up to 2500 m asl (Frost et al. 2006; Van Bocxlaer et al. 2009; Portik and Papenfuss 2015). The genus is represented by 19 species in India, 16 of which were described with type localities designated in the country. Among the Indian Duttaphrynus species, nine occur in Peninsular India and of these, six are endem- ic to the region. Although the wide-ranging species (D. melanostictus, D. stomaticus, D. hololius, and D. scaber) are frequently studied and reported from Peninsular India (Sarkar et al. 1993; Dutta 1997; Biju 2001; Chanda 2002; Van Bocxlaer et al. 2009; Dinesh et al. 2009; Srinivasulu et al. 2013; Ganesh et al. 2020), the taxonomic status of the endemic species has not been thoroughly investigated Copyright Bisht K. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 452 subsequent to their original descriptions (Dubois and Ohler 1999; Biju 2001). These include five recognised species—D. beddomii (Gunther, 1876), D. brevirostris (Rao, 1937), D. microtympanum (Boulenger, 1882), D. parietalis (Boulenger, 1882), and D. silentvalleyensis (Pillai, 1981). Their identities remain somewhat doubtful, due to reasons such as either brief or cursory original de- scriptions, unavailability of type specimens, or absence of new topotypic collections (Dubois and Ohler 1999; Biju 2001). In addition, identification of Duttaphrynus species is challenging, due to their overall phenotypic similari- ties and substantial intraspecific morphological variabil- ity (Inger 1972; Dubois and Ohler 1999; Biju 2001; Van Bocxlaer et al. 2010; Wogan et al. 2016; Jayawardena et al. 2017). Another four available names from Penin- sular Indian regions exist as junior subjective synonyms (Dubois and Ohler 1999). Given such complex nomen- clatural histories, misidentifications of Duttaphrynus spe- cies in museum specimens (S.D.B., personal observation) and regional biodiversity reports (Ray and Deuti 2008; Gururaja 2012; Hegde 2012; Seshadri et al. 2012; Ganesh et al. 2020) are frequent. Two Duttaphrynus toads were described by C. R. Narayan Rao (15 August 1882-2 January 1960), who was among the most notable amphibian taxonomist in south- ern India during the colonial and post-colonial periods of the twentieth century. He described a total of 27 new spe- cies of frogs, including subspecies and varieties, largely from the states of Karnataka and Tamil Nadu (Rao 1920, 1922, 1937). However, a large number of his types (19 species; deposited in the Central College, Bangalore) are lost (Dubois 1984; Biyu 2001). Seventeen of Rao’s spe- cies currently are recognised as valid; nine of these have had their name-bearing type status stabilised through des- ignation of neotype specimens (e.g., Bossuyt and Dubois 2001; Biyu et al. 2011, 2014a, 2014b; Garg et al. 2018). Similarly, the fate of Rao’s bufonid species has remained precarious: (1) Bufo brevirostris Rao, 1937 was described based on a single specimen from “Kempholey, Hassan District, Mysore State,” which subsequently was reported to be lost (Dubois 1984; Biyu 2001). Hence, this species is known only from its original description. Dubois and Ohler (1999) discussed the problematic taxonomic status of this taxon, and, later Van Bocxlaer et al. (2009) trans- ferred it to Duttaphrynus based on DNA sequences from a single specimen, without further information or discus- sion. The species continues to be recognised in the litera- ture, albeit in the absence of new reliable records, photo- graphs, or voucher specimens (Dutta 1997, Chanda 2002; Dinesh et al. 2009; Subramanian et al. 2013; Jayawardena et al. 2017). Additionally, (2) Bufo stomaticus peninsu- laris Rao, 1920 was described as a new variety of “Bufo stomaticus” from “Mavkote and Watekolle, Coorg,” based on a specimen (ZSIC 19176) designated as the holotype by Chanda et al. (2001 “2000”). This taxon was considered a synonym of Duttaphrynus stomaticus (Daniel 1963; Dan- iels 2005), until Srinivasulu et al.’s (2013) correction of some photograph-based misidentifications of “D. scaber” zse.pensoft.net Bisht K. et al.: Duttaphrynus toads from India (not Duttaphrynus stomaticus peninsularis) as “D. stomat- icus,” which was implicitly considered as the transfer of Bufo stomaticus peninsularis into the synonymy of Dut- taphrynus scaber (Schneider, 1799) by Frost (2021). However, most recently Ganesh et al. (2020) made a cur- sory statement referring to the identity of this taxon as “status: incertae sedis” without any clarification. The confusing taxonomic status of Rao’s variety Bufo stomaticus peninsularis is also undeniably linked to its originally assigned species—Duttaphrynus stomaticus (Litken, 1864). Although Srinivasulu et al. (2013) re- ported on misidentifications of D. stomaticus from Pen- insular India, no studies to date have provided direct and conclusive evidence for either resolving the identity of Bufo stomaticus peninsularis or clarifying the occurrence of Duttaphrynus stomaticus in Peninsular India. The latter is considered as a widely distributed species in south and southwest Asia, with its range encompassing nearly the whole of India and the neighbouring Bangladesh, Nepal, Pakistan, Afghanistan, and Iran (Stéck et al. 2006; Ras- tegar et al. 2008; Van Bocxlaer et al. 2009; Shaikh et al. 2014; Portik and Papenfuss 2015; Nepali and Singh 2018; Frost 2021) (Suppl. materal 1: Table S1). However, Dut- taphrynus stomaticus was originally described from “ostin- diske” (= East Indies or East India) (Lutken 1864), where its type locality was subsequently restricted to “Assam” (Boulenger 1891). Since type specimens were reported as untraceable (Dutta 1997), the identification of this species in recent literature is apparently based only on its brief orig- inal description, rather than examination of name-bearing types, or detailed redescription of topotypic material. The present study was undertaken to conclusively resolve the taxonomic identity and stabilise the nomen- clatural status of the two lesser-known Duttaphrynus toads from Peninsular India (Bufo brevirostris Rao, 1937 and Bufo stomaticus peninsularis Rao, 1920) and another wide-ranging northern species (Bufo stomaticus Litken, 1864). We do so based on morphological comparison with original descriptions and available type specimens (except for D. brevirostris), as well as molecular and morphological insights gathered from new topotypic ma- terial, arguably rediscovered for the first time since both species’ original descriptions. We also aimed to infer phylogenetic relationships of the focal species, as well as gather insights on patterns of genetic differentiation among all known members of the genus Duttaphrynus that are characterised by known localities, and represent- ed by accompanying vouchered molecular data. Materials and methods Field study Surveys were carried out for sampling the target spe- cies from regions encompassing their type localities in the Indian states of Karnataka, Andhra Pradesh, Tam- il Nadu, and Assam. Additionally, some populations of Zoosyst. Evol. 97 (2) 2021, 451-470 ‘Duttaphrynus stomaticus’ were randomly sampled from regions across India to understand intra and interspecific relationships. A total of 15 newly sampled populations are included in the study (Suppl. materal 1: Tables S2 and S3). Surveys and sampling were conducted both during day and night hours, mostly during the pre-monsoon and monsoon months (April—August), but occasionally also at other times of the year (March and October). The sampled individuals were photographed to document colouration and characters in life, followed by euthanisation using Tricaine methanesulphonate (MS-222). Tissue samples were taken from the thigh muscle or liver, preserved in absolute ethanol, and stored at -20 °C for molecular stud- ies. Locality information was recorded using a GPS with the WGS84 datum system. Distribution maps were pre- pared in QGIS version 2.6.1 (http://www.qgis.org). Morphological study Sex and maturity were determined by examining the go- nads through a small lateral or ventral incision, or by the presence of secondary sexual characters (such as nup- tial pads and vocal sacs in males). The following mea- surements were taken to the nearest 0.1 mm with digital slide-calipers: SVL (snout-vent length), HW (head width, at the angle of the jaws), HL (head length, from rear of mandible to tip of snout), SL (snout length, from tip of snout to anterior orbital border), EL (eye length, horizon- tal distance between bony orbital borders), IFE (internal front of the eye, shortest distance between the anterior orbital borders), IBE (internal back of the eyes, shortest distance between the posterior orbital borders), IUE (in- ter upper eyelid width, the shortest distance between the upper eyelids), VUEW (maximum upper eyelid width), IN (internarial distance), NS (distance from the nostril to the tip of the snout), EN (distance from the front of the eye to the nostril), PD (minimum distance between parotoids), PL (maximum parotoid length), PW (maximum parotoid width), TYD (greatest tympanum diameter), TYE (dis- tance from the tympanum to the back of the eye), FAL (forearm length, from flexed elbow to base of outer pal- mar tubercle), HAL (hand length, from base of outer pal- mar tubercle to tip of third finger), TL (thigh length, from the vent to the knee), SHL (shank length, from knee to heel), FOL (foot length, from base of inner metatarsal tu- bercle to tip of fourth toe), TFOL (total foot length, from heel to tip of fourth toe), ITL (inner toe length), OMTL (length of outer metatarsal tubercle), and IMTL (length of inner metatarsal tubercle). Digit number is represented by roman numerals I—V in subscript. All measurements provided in the taxonomy section are in millimetres (mm). Measurements and associated terminology follow Dubois and Ohler (1999) and Biju and Bossuyt (2009). The webbing formulae follow Savage and Heyer (1967) as modified by Myers and Duellman (1982). The amount of webbing relative to subarticular tubercles is described by numbering the tubercles 1—3, starting from the base. 453 For the convenience of discussion, webbing is addition- ally defined as basal, small, medium, or large, following Garg and Biju (2017). To ascertain the degree of morphometric differentia- tion among the three Indian members of the Duttaphry- nus stomaticus group, a multivariate analysis was per- formed using 21 morphometric characters from male specimens. The data for each character was expressed as the ratio of the respective SVL so as to reduce the im- pact of allometry, and subjected to Principal Component Analysis (PCA), a dimensionality reduction technique. Furthermore, Box and Whiskers plots were created for a univariate analysis of SVL and five morphometric char- acters that yielded the most significant contribution to the PCA, in order to visualise differences among the species. The analyses were performed in R (R Development Core Team 2008) using the package MASS and the plots were made using the ggplot2 and ggfortify packages. Molecular study Genomic DNA was extracted from the new samples us- ing Qiagen DNeasy Blood and Tissue Kit (Qiagen, Va- lencia, CA, USA) following the manufacturer’s proto- cols. A short fragment of the mitochondrial 16S rRNA (~540 bp) was PCR-amplified using previously published primer sets 16Sar and 16Sbr (Simon et al. 1994). Puri- fied PCR products were sequenced with the same prim- ers using BigDye Terminator v3.1 Cycle Sequencing Kit on ABI 3730 automated DNA sequencer (Applied Bio- systems). Raw sequences were checked and assembled in ChromasPro v1.34 (Technelysium Pty Ltd.) and de- posited in the NCBI GenBank under accession numbers MZ816170—-MZ816184. We reconstructed phylogenetic relationships among major distinct evolutionary lineages representing known or putative Duttaphrynus species (Van Bocxlaer et al. 2009; Portik and Papenfuss 2015). DNA sequences for nine mitochondrial gene regions (12S ribosomal RNA, tRNA“, 16S ribosomal RNA, tRNA‘, NADH dehy- drogenase subunit 1, tRNA"*, tRNAS™, tRNA™*, and NADH dehydrogenase subunit 2) and two nuclear genes (NCX1 and CXCR4) from previously published stud- ies (Byyu and Bossuyt 2003; Van Bocxlaer et al. 2009; Portik and Papenfuss 2015; Liedtke et al. 2016) were retrieved from the GenBank and assembled along with selected new sequence data (Suppl. materal 1: Table S2). Sequences were aligned using ClustalW in MEGA 6.0 (Tamura et al. 2013). Alignments for coding DNA were checked by comparison with amino acid sequenc- es, whereas the alignment for non-coding sequences was visually optimised and the ambiguously aligned regions were subsequently excluded from phylogenetic analyses. A character set of total 5,737 bp assembled for 18 taxa was used for the Maximum Likelihood (ML) and Bayes- ian Inference (BI). Appropriate models of sequence evo- lution were determined for each gene by implementing zse.pensoft.net 454 Akaike Information Criteria in ModelTest 3.4 (Posada and Crandall 1998). Maximum Likelihood (ML) search- es were performed on a partitioned dataset using the GTRGAMMA model with 2,000 independent runs ex- ecuted alongside 10,000 rapid bootstrap replicates in RAXxML 7.3.0 (Stamatakis et al. 2008) as implemented in raxmlGUI 1.1 (Silvestro and Michalak 2012). Bayes- ian analyses were performed using the best-fit General Time Reversible (GTR) model with a proportion of in- variant sites (+1) and gamma-distributed rate variation among sites (+G) independently for each gene partition, with all parameters estimated. Bayesian searches were executed in MrBayes (Ronquist and Huelsenbeck 2003) with two parallel runs of four Metropolis-Coupled Mar- kov Chain Monte Carlo (MCMCMC) chains executed for 10 million generations using uniform priors and sam- pling frequency of trees after every 1,000 generations. Convergence of the parallel runs was determined by split frequency standard deviations of less than 0.01 and ~1.0 potential scale reduction factors for all model pa- rameters. Bayesian posterior probabilities (BPP) for the clades were summarised after discarding the first 2,500 trees (25 percent) as burn-in from each run (Huelsen- beck et al. 2001). We further assessed relationships using available ho- mologous mitochondrial 16S rRNA sequences from GenBank and our new samples (Suppl. materal 1: Table S3). Sequences were aligned using ClustalW in MEGA 6.0 (Tamura et al. 2013) and the alignment was manually checked for the presence of any ambiguous or doubtful sites. Certain short GenBank sequences and sequences or positions that showed low confidence for homology were excluded from phylogenetic analyses. A character set of 524 bp from 137 taxa, including an outgroup, was subjected to ML and BI analyses. The ML search was ex- ecuted in RAXML based on 500 independent runs using the GTRGAMMA model and clade support was assessed through 1,000 rapid bootstrap replicates. The Bayesian analysis was performed with two parallel runs of four MCMCMC chains executed for 10 million generations using the GTR+I+G model, with a sampling frequency of 1,000 and 25 percent burn-in. The resultant ~15,000 trees were summarised to determine clade support (BPP). The details of the analyses were as described above for the multi-gene dataset. Additionally, the ML phylogram was used as input for performing species delimitation analyses by Bayesian implementation of the Poisson Tree Processor (PTP) method (Zhang et al. 2013) on the bPTP webserver (https://species.h-its.org). Intra- and interspe- cific uncorrected pairwise genetic distances for the 16S tRNA were computed in PAUP* (Swofford 2002). A Me- dian-Joining (MJ) network was further constructed using the software Network 4.6.1.0 (www.fluxus-engineering. com) to evaluate relationships and possible mutation steps among 42 haplotypes recovered from 133 sequenc- es of the 16S rRNA after performing the PHASE algo- rithm (Stephens et al. 2001) in DnaSP version 5 (Librado and Rozas 2009). zse.pensoft.net Bisht K. et al.: Duttaphrynus toads from India Abbreviations Museum acronyms and other abbreviations used herein are as follows: BNHS (Bombay Natural History Society, Mumbai); CCB (Central College, Bangalore); CSPT (Chennai Snake Park Trust, Chennai); ICZN (The In- ternational Code of Zoological Nomenclature); SDBDU (Systematics Lab, University of Delhi, India); ZMUC (Universitets Kobenhavn, Zoologisk Museum, Denmark); ZSIC (Zoological Survey of India, Kolkata, India). Results and discussion Taxonomic accounts Duttaphrynus brevirostris (Rao, 1937) Figs 1-4; Table 1; Suppl. materal 1: Tables S1-S4 Kempholey Toad Original name and description. Bufo brevirostris Rao, 1937. Rao, C. R. N. 1937. On some new forms of Ba- trachia from S. India. Proceedings of the Indian Acad- emy of Sciences. Section B 6: 387-427. Type locality. “Kempholey, Hassan District, Mysore State,” Karnataka, India. Current status of specific name. Valid name, as Duttaphrynus brevirostris (Rao, 1937). Material studied. Jopotype. An adult male, BNHS 6126 (SVL 45 mm), from Kempholey Ghat region in Sakleshpur taluk, Hassan district, Karnataka State, India, collected by S. D. Biyyu and Sonali Garg in June 2013. Oth- er referred specimens. An adult male, SDBDU 2008.410 (SVL 48.6 mm), from Bhagamandala, Kodagu district, Karnataka State; an adult male, SDBDU 2015.3075 (SVL 46 mm), from Manipal, Udupi district, Karnataka State; and a subadult, SDBDU 4714 (SVL 25 mm), from Someshwara, Udupi district, Karnataka State. Rediscovery and validation of taxonomic status. This species was described based on a single specimen (“snout to vent, 27.00 mm’) deposited in the Central Col- lege, Bangalore (CCB). This original name-bearing type specimen is considered lost (Dubois 1984; Biju 2001) and the species currently is known only from its original description. Rao (1937) enumerated several morphologi- cal character states to describe this taxon, but did not pro- vide comparisons with other species. Our collection from a region of Kempholey Ghat in Sakleshpur taluk, that is part of the type locality (Rao 1937), is comparable with the original description with respect to several mentioned characters such as “canthus rostralis angular,” “nostril nearer to the end of the snout than to the eye,” “first fin- ger equal to the second,” “parotoids elongate, moderate- ly prominent,” and “upper surface of the skin covered with small uniformly distributed tubercles; with a small row of larger warts on the median line of the back.” The primary inconsistencies between Rao’s described speci- men and our new collection involve snout-vent length, SVL 45 mm (vs. “27.00 mm”) and weakly developed or Zoosyst. Evol. 97 (2) 2021, 451-470 455 P Figure 1. Morphological characters for topotype of Duttaphrynus brevirostris (Rao, 1937), topotype of D. peninsularis (Rao, 1920), and syntype of D. stomaticus (Ltitken, 1864) in preservation. A-G. Duttaphrynus brevirostris, BNHS 6126: A. Dorsal view; B. Ven- tral view; C. Lateral view of head; D. Dorsal view of hand showing brown nuptial pad on fingers I, II, and II; E. Ventral view of hand; F. Ventral view of foot; G. Schematic illustration of webbing on foot. H-N. Duttaphrynus peninsularis: H. Holotype, ZSIC 19176; IN. Topotype, SDBDU 6370: I. Dorsal view; J. Ventral view; K. Lateral view of head; L. Ventral view of hand; M. Ventral view of foot; N. Schematic illustration of webbing on foot. O-T. Duttaphrynus stomaticus, ZMUC 131137 (ex 196): O. Dorsal view; P. Ven- tral view; Q. Lateral view of head; R. Ventral view of hand; S. Ventral view of foot; T. Schematic illustration of webbing on foot. inconspicuous cephalic ridges (vs. “crown without bony ridge”). The cephalic ridges in our new collection are relatively smooth, depressed, or less conspicuous (Figs 1A, C, 2A) when compared to other species of the Dut- taphrynus melanostictus group from Peninsular India. Hence, presence or absence of this character may be con- sidered a matter of interpretation depending on degree of its prominence. Furthermore, the body size disparity be- tween our collection and that of Rao (1937) also suggests that the type specimen he described could have been a subadult. We examined another subadult specimen from Someshwar (SDBDU 4714; SVL 25 mm), previously zse.pensoft.net Duttaphrynus toads from India Bisht K. et al.: 456 HO Se oe Ce acter eae = Oo e?e ILWO Le 8c vt ET 8l GL Te ILWO HO ae a a Ss oe Fm —l S44 ALI Se oe Ce | ca oT ILI 0? LY G9¢ GE? ECS 692 EVE 1OsL 0? VCE UTE O-9€ 6 Ce OTE SOE 1OsL GG 6 GC Bid T'9¢ T'9¢ VL¢e €8e 1OsL vt 6 8 c 8e SOE T'8¢ 1OsL Lea 89T 98T T9l 1-91 89T Ee OT 104 LI, £1¢ 9°02 Sve vie Goo Vol 104 £1 OLT Tvl ZT co Ll est VSl 104 30 ¢ ol ¢ 6L Og GST 104 90 LL1 SLi SLT c Ll 3881 VLT AHS 8C COC c O¢ OLE SECC € 02 €0¢ HS vt VET LS TEL 391 vol SLl JHS ime) 881 681 8 8l 388i AHS £0 U'st 681 LST LL1 cy a! 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Stl VOL LLY it Sc vor SST c Sv sfels BC OVI O°EL O'¢S it Te OVI Ost 80S hl GAL dH MH TAS SN}RYS 20 60 vO 3T ds BC vVvt LOT VOY = Ueel| 6? Sel EO Lv SY 6? VST 6 9T V SV Su O'S OVI 6 9T OST LL GAL dH MH TAS SM}e4S SZLOeE STO?e NdEds OTvV 800¢ NdEds 9¢T9 SHNG ON Jayono, (eyejeusey Wo ‘sajew |je) sUIsolAelg snuAiydeyng zse.pensoft.net Zoosyst. Evol. 97 (2) 2021, 451-470 457 eo. ee ee ee Sarees oath jet Sie Figure 2. Topotype of Duttaphrynus brevirostris (Rao, 1937), topotype of D. peninsularis (Rao, 1920), and referred specimens of D. stomaticus (Litken, 1864) in life. A. Duttaphrynus brevirostris (BNHS 6126) from Kempholey Ghat region in Sakleshpur taluk. B. Duttaphrynus peninsularis (SDBDU 6370) from Wattakolli. C-F, Duttaphrynus stomaticus: C. SDBDU 2015.2909 from Assam; D. SDBDU 2012.2170 from Rajasthan; E. SDBDU 2012.2172 from Delhi; and F. SDBDU 2012.2268 from Bihar. reported along with DNA sequence data (Van Bocxlaer et al. 2009), and found some comparable characters such as “a small row of larger warts on the median line of the back,” “a network of dark lines,” and “a dark temporal line extending to the sides,” which can usually also be ob- served in subadults of Duttaphrynus melanostictus group species (S.D.B., personal observations). The Someshwar specimen is genetically identical to our Sakleshpur col- lection. Together, these two populations are also morpho- logically and genetically similar to our additional collec- tions from other localities within the Malenadu (Malnad) and adjoining coastal regions of Karnataka (see ‘Material studied’). Altogether, we consider the available morpho- logical and molecular evidence reliable for assigning all the mentioned populations to D. brevirostris (Rao, 1937). Since the absence of a name-bearing type has contrib- uted towards poor knowledge and uncertainty regarding the taxonomic identity of this taxon, as evident from the absence of new records, below we provide a detailed de- scription of a newly-collected voucher specimen from the original type locality (Kempholey Ghat region in Saklesh- pur taluk, Hassan district, Karnataka State, India: BNHS 6126), which is largely consistent with what is known of the former name-bearing type (Rao 1937). The topotype description provided below, augmented by a range of variation observed in vouchered specimens and genetic data from additional localities (Table 1; Suppl. materal 1: Tables S3, S4), validate the identity of D. brevirostris and also serve as a redescription of this poorly known species for the benefit of future taxonomic work. Description of topotype, BNHS 6126 (measurements in mm). A medium-sized, robust adult male (SVL 45.0); head of moderate size, wider (HW 16.9) than long (HL 14.0); snout subovoid in dorsal and ventral view, not pro- zse.pensoft.net 458 jecting, its length (SL 6.1) longer than horizontal diameter of eye (EL 5.9); loreal region obtuse with sharp canthus rostralis; distance between posterior borders of the eyes (IBE 13.9) 2.2 times the distance between the anteri- or borders (IFE 6.3); interorbital space 1.2 times wider (TUE 5.1) than upper eyelid width (UEW 4.1); nostril oval without lateral flap of skin, closer to tip of snout (NS 1.7) than to eye (EN 3.2); tympanum distinct (TYD 2.6), verti- cally oval, 44.1% of eye diameter (EL 5.9), tympanum to eye distance (TYE 0.7); pineal ocellus absent; vomerine ridge and teeth absent; tongue small, oval, entire, median lingual projection absent; parotoid glands present, oval, flat, without spines and warts, longer (PL 6.2) than wide (PW 3.4), shorter than distance between them (PD 8.7); supraorbital and postorbital ridges weakly developed. Forelimbs short; forearm length (FAL 10.8) short- er than hand length (HAL 11.3); fingers rather thin, FL, nearly equal to FL, FL,,, longest (6.3); relative length of fingers: I=II