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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4384 (1): 001
–
088
http://www.mapress.com/j/zt/
Monograph
https://doi.org/10.11646/zootaxa.4384.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:18DD1320-7914-4E09-A46C-707069DC69F5
ZOOTAXA
An integrative taxonomic review of the South
Asian microhylid genus Uperodon
SONALI GARG
1,9
, GAYANI SENEVIRATHNE
2,3
, NAYANA WIJAYATHILAKA
4
,
SAMADHAN PHUGE
5
, KAUSHIK DEUTI
6
, KELUM MANAMENDRA-ARACHCHI
7
,
MADHAVA MEEGASKUMBURA
8,2,9
& SD BIJU
1,9
1
Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India
2
Department of Molecular Biology & Biotechnology, Faculty of Science, University of Peradeniya, Peradeniya, Sri Lanka
3
Department of Organismal Biology & Anatomy, University of Chicago, IL, USA
4
Department of Zoology, Faculty of Applied Sciences, University of Sri Jayewardenepura, Sri Lanka
5
Department of Zoology, Savitribai Phule Pune University, Ganeshkhind, Pune, India
6
Herpetology Division, Zoological Survey of India, 27 JL Nehru Road, Kolkata, West Bengal, India
7
Postgraduate Institute of Archaeology, University of Kelaniya, 407 Bauddhaloka Mawatha, Colombo, Sri Lanka
8
Guangxi Key Laboratory of Forest Ecology and Conservation, College of Forestry, Guangxi University, Nanning, China
9
Corresponding authors. E-mail: sdbiju@es.du.ac.in, sonali1909@gmail.com, madhava_m@mac.com
Magnolia Press
Auckland, New Zealand
4384
GARG ET AL.
2
·
Zootaxa 4384 (1) © 2018 Magnolia Press
SONALI GARG, GAYANI SENEVIRATHNE, NAYANA WIJAYATHILAKA, SAMADHAN PHUGE,
KAUSHIK DEUTI, KELUM MANAMENDRA-ARACHCHI, MADHAVA MEEGASKUMBURA & SD
BIJU
An integrative taxonomic review of the South Asian microhylid genus Uperodon
(Zootaxa 4384)
88 pp.; 30 cm.
22 Feb. 2018
ISBN 978-1-77670-315-9 (paperback)
ISBN 978-1-77670-316-6 (Online edition)
F
IRST
P
UBLISHED I
N
2018 B
Y
Magnolia Press
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ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
Zootaxa 4384 (1) © 2018 Magnolia Press
·
3
TAXONOMIC REVIEW OF GENUS UPERODON
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
DNA analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Order Anura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Family Microhylidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Subfamily Microhylinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Genus Uperodon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Uperodon systoma (Schneider, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Uperodon globulosus (Günther, 1864 “1863”). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Uperodon taprobanicus (Parker, 1934) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Uperodon anamalaiensis (Rao, 1937) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Uperodon montanus (Jerdon, 1854 “1853”). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Uperodon mormorata (Rao, 1937) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Uperodon nagaoi (Manamendra-Arachchi and Pethiyagoda, 2001) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Uperodon obscurus (Günther, 1864 “1863”) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Uperodon palmatus (Parker, 1934) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Uperodon triangularis (Günther, 1876 “1875”) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Uperodon variegatus (Stoliczka, 1872) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Uperodon rohani sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Natural history and reproductive behaviour . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .45
Comparative morphology of tadpoles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .58
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Supplementary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Abstract
Based on a recent molecular phylogenetic study, the South Asian microhylid genus Uperodon (subfamily Microhylinae)
currently comprises of 12 valid species that are largely restricted to India and Sri Lanka. Considering the revised generic-
level status of its various members, here we review the taxonomy of all known species in this genus and clarify their no-
menclatural status and geographical distribution, by integrating evidence from genetics, adult and tadpole morphology,
breeding ecology, and bioacoustics. Our molecular analyses of a mitochondrial 16S rRNA gene fragment combined with
external and internal morphological studies also revealed a distinct new species in the genus. This species, formally de-
scribed as Uperodon rohani sp. nov., is endemic to Sri Lanka and widely distributed at lower elevations in the island. For
nomenclatural stability of various previously known members, the following actions are also undertaken: (1) redescription
of the poorly-defined species Ramanella anamalaiensis Rao (= Uperodon anamalaiensis) and Hylaedactylus montanus
Jerdon (= Uperodon montanus); (2) neotype designation for Ramanella anamalaiensis Rao (= Uperodon anamalaiensis),
Ramanella minor Rao, Ramanella mormorata Rao (= Uperodon mormorata), and Ramanella triangularis rufeventris
Rao; (3) lectotype designation for Callula variegata Stoliczka (= Uperodon variegatus); and (4) synonymization of Ra-
manella minor Rao with Uperodon anamalaiensis.
Key words: Amphibians, bioacoustics, endemism, mitochondrial DNA, natural history, neotype, lectotype, new species,
tadpoles, Western Ghats-Sri Lanka biodiversity hotspot
Introduction
The genus Uperodon Duméril and Bibron is classified in subfamily Microhylinae of the narrow-mouthed frog
family Microhylidae. This genus was originally described to accommodate Engystoma marmoratum Guérin-
Méneville, subsequently considered as a junior subjective synonym of Rana systoma Schneider, which was among
the first frog species described from India. Another species, Cacopus globulosus Günther was transferred to
Uperodon and the genus continued to comprise of two species—Uperodon systoma and Uperodon globulosus, until
GARG ET AL.
4
·
Zootaxa 4384 (1) © 2018 Magnolia Press
recently. A molecular study by Peloso et al. (2016) revised the contents of this genus mainly to resolve the
paraphyly of genus Kaloula Gray that resulted from the phylogenetic position of Kaloula taprobanica Parker as
being sister to the clade containing Ramanella Rao and Ramanna (currently unrecognized) + Uperodon sensu
stricto (e.g., Van Bocxlaer et al. 2006; Pyron & Wiens 2011). As a result, all the members of genus Ramanella (=
Uperodon) and Kaloula taprobanica (= Uperodon taprobanicus) are now included in genus Uperodon (sensu lato),
which together comprises of 12 recognised species (Frost 2017).
Originally, Ramanella was diagnosed as a distinct genus on the basis of external and internal morphological
characters (Rao & Ramanna 1925; Parker 1934), and Uperodon taprobanicus (Parker 1934) was described as a
subspecies—Kaloula pulchra taprobanica. There are two notable studies concerning the morphological
relationship between the three ‘genera’—Kaloula, Ramanella and Uperodon (sensu stricto). Parker’s monograph
of the family Microhylidae (1934) discussed various morphological similarities and dissimilarities useful for
genus-level diagnosis. He also transferred four species—Uperodon montanus Jerdon, U. obscurus Günther, U.
triangularis Günther and U. variegatus Stoliczka—originally described in other genera to Ramanella. Ramaswami
(1936), however, considered Ramanella not to be distinct from Kaloula (Peloso et al. 2016). Nevertheless,
subsequent authors continued to recognize Ramanella, and several new species were added to this genus (Parker
1934; Rao 1937; Manamendra-Arachchi & Pethiyagoda 2001).
Over the years, considerable natural history information has accumulated for various Uperodon species (see
‘results’ for detailed discussion). However, several members of the genus are still poorly studied and known either
only from their original descriptions and type localities, or from invalidated reports (e.g., U. anamalaiensis Rao, U.
montanus, U. minor Rao, U. nagaoi Manamendra-Arachchi and Pethiyagoda, U. palmatus Parker). Another
species, U. mormorata Rao was only recently rediscovered (Das & Whitaker 1997). Although numerous additional
distribution records are available for Uperodon species in literature (see Supplementary Table S1), many lack
vouchers or photographs, and therefore require further confirmation. This is also one of the major problems
currently plaguing the genus since identities of many Uperodon species remain obscure in the absence of type
material. The type specimens of species described by Rao (1937) are considered lost (Dubois 1984; Biju 2001)
resulting in several uncertainties concerning species identity. Two species, U. anamalaiensis and U. minor, had not
been reported for several decades since their original description. While the rediscovery of U. anamalaiensis from
its type locality was also recently announced by Vijaykumar et al. (2011), no voucher-based observations have
been made for U. minor. Further, there are several available names currently in the synonymy of U. variegatus, of
which Hylaedactylus carnaticus Jerdon is suggested to be a senior synonym (Parker 1934) but the type material for
this species is also reported to be lost (Jerdon 1870). Another taxon, Ramanella triangularis rufeventris Rao,
currently in the synonymy of Uperodon triangularis, was described as a subspecies (Rao 1937) but its type is
considered lost (Dubois 1984; Biju 2001). The type locality of Uperodon triangularis itself has been noted to be
imprecise (Biju 2001). With such essential information currently unavailable for various nominal species, the
taxonomy of Uperodon is in need of a comprehensive revision to ensure proper species identification, which is also
necessary for any meaningful studies on this group. Additionally, several known taxa require designation of
neotypes and lectotypes in order to establish nomenclatural stability.
In this work, we studied over 70 Uperodon populations from across their range in India and Sri Lanka to
understand the existing diversity, variations, geographical ranges and distribution patterns, by integrating evidence
from genetics and morphology. For the first time, we present a comprehensive taxonomic revision of all known
members of the genus Uperodon, along with the description of a new species. Additional information on tadpole
morphology, advertisement call structure, breeding ecology, and natural history is summarized with previously
available knowledge of various members. Morphological comparisons and affinities are also provided for all the
species to facilitate a better working taxonomy for this group.
Materials and methods
Field surveys and specimen collection. Field surveys were conducted in India and Sri Lanka between 2002 and
2015. A total of 44 localities were surveyed in India (33) and Sri Lanka (11) (Table 1). Our sampling effort was
focused largely on the Western Ghats–Sri Lanka biodiversity hotspot because of the high diversity and endemism
of Uperodon frogs in this region. Sampling was usually done through opportunistic searches during both day and
Zootaxa 4384 (1) © 2018 Magnolia Press
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TAXONOMIC REVIEW OF GENUS UPERODON
night. Males were also located by their calls after sunset, mostly during the breeding season (June–August in the
Western Ghats, October–December in the Eastern Ghats and Sri Lanka) or immediately after sporadic rains.
Suitable habitats such as temporary ponds and tree holes were specifically searched for the presence of tadpoles
and eggs. Observations on breeding behaviour and ecology were also made during the monsoon seasons in India
(June–August/October–November) and Sri Lanka (October–December). Sampled individuals were euthanized in
MS-222 (Tricaine methanesulfonate). A portion of the thigh muscle or liver tissue (adult) or tail tissue (tadpole)
was taken in absolute ethanol and stored at a temperature of -20°C for molecular studies. Adult specimens were
fixed in 4% formalin and preserved in 70% ethanol. Eggs and tadpoles were stored in 3.5% neutral buffered
formalin. Type specimens of the new species are deposited in Department of Molecular Biology & Biotechnology,
University of Peradeniya (DZ), Sri Lanka; neotypes are deposited in the Zoological Survey of India–Western Ghats
Regional Centre (ZSI-WGRC), Calicut, India. Referred specimens are available at Systematics Lab, University of
Delhi (SDBDU) and Department of Molecular Biology & Biotechnology, University of Peradeniya (DZ), for the
Indian and Sri Lankan species, respectively. All authors were involved in sampling and/or collection of field data.
Molecular study. Genomic DNA was extracted from 67 tissue samples (Table 2) using Qiagen DNeasy blood
and tissue kit (Qiagen, Valencia, CA, USA), following the manufacturer’s protocol. A fragment of the
mitochondrial 16S rRNA gene (≈ 540 bp) was PCR-amplified using the previously published primer set: 16Sar and
16Sbr (Simon et al. 1994). Purified PCR products were sequenced on both strands using BigDye Terminator v3.1
Cycle Sequencing Kit on ABI 3730 automated DNA sequencer (Applied Biosystems). DNA sequences were
assembled and edited in ChromasPro v1.34 (Technelysium Pty Ltd.). The alignment was created using ClustalW in
MEGA 6.0 (Tamura et al. 2013) and manually optimized. New sequences are deposited in the GenBank under
accession numbers MG557906–MG557972 (Table 2). Uncorrected pairwise genetic distances were computed in
PAUP* (Swofford 2002) and intra- and interspecific comparisons between the sequences were made for
preliminary genetic delineation of species. SG (Indian material), GS and NW (Sri Lankan material) generated
DNA sequences.
Maximum Likelihood (ML) and Bayesian analyses were performed to reconstruct phylogenetic relationships.
Ambiguously aligned regions were excluded from the analyses. Trees were rooted using Kaloula conjuncta, K.
pulchra and K. mediolineata as outgroups (Van Bocxlaer et al. 2006; Pyron & Wiens 2011; Peloso et al. 2016).
Akaike Information Criteria were implemented in ModelTest 3.4 (Posada & Crandall 1998) to estimate the
appropriate model of sequence evolution. Analyses were performed using the best-fit General time reversible
model with a proportion of invariant sites and gamma-distributed rate variation among sites (GTR+I+G).
Maximum Likelihood searches were executed in PAUP* (Swofford 2002), with all parameters estimated. Bayesian
analysis was implemented in MrBayes (Ronquist & Huelsenbeck 2003) using uniform priors and four Metropolis-
Coupled Markov Chain Monte Carlo (MCMCMC) chains run for 10 million
generations. Trees were sampled every
1000 generations and based on a conservative approach samples from the first five million
generations were
discarded as burn-in (Huelsenbeck et al. 2001). Bayesian posterior probabilities (BPP) for the clades were
estimated by summarizing the remaining trees with 50% majority rule in PAUP* (Swofford 2002). Maximum
Likelihood clade support (BS) was also assessed by 10,000 rapid bootstrap replicates executed using RAxML 7.3.0
(Stamatakis et al. 2008) as implemented in raxmlGUI 1.1 (Silvestro & Michalak 2012). SG and SDB performed the
molecular analyses.
TABLE 1. Collection localities for the genetically and/or morphologically studied Uperodon samples in the present
study.
Locality Altitude Coordinates Species
(metres) Latitude (°N) Longitude (°E)
INDIA
Tam i l Na du
Coimbatore district
Coimbatore 400 11.0825 77.0617 U. systoma
1
Kanchipuram district
......continued on the next page
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Zootaxa 4384 (1) © 2018 Magnolia Press
TAB LE 1. (Continued)
Locality Altitude Coordinates Species
(metres) Latitude (°N) Longitude (°E)
Chengalpattu 60 12.6902 80.0258 U. variegatus
1
Kunnapattu 20 12.6303 80.1521 U. systoma
1
Mamallapuram 23 12.7412 80.2372 U. taprobanicus
1
Nilgiris district
Udhagamandalam (Ooty) 2215 11.4347 76.6692 U. triangularis
1
Tirunelve li district
Kakkachi 1266 08.5359 77.3952 U. montanus
1
Sengaltheri 1000 08.5371 77.4544 U. montanus
2
Kerala
Idukki district
Devikulam 1670 10.0626 77.1040 U. montanus
2
Methooty 446 09.8423 78.8712 U. anamalaiensis
1
Munnar 1410 10.0833 77.1333 U. montanus
1
Upper Manalar, Periyar TR 1564 09.5748 77.3362 U. montanus
1
Kollam district
Pandimotta, Shendurney WLS 1221 08.8266 77.2122 U. montanus
1
Palakkad district
Kesavapara, Nelliyampathy 923 10.5243 76.6673 U. montanus
1
Palakkad 86 10.8026 76.6361 U. variegatus
1
Parambikulam TR 544 10.4453 76.8130 U. anamalaiensis
1
Siruvani 880 10.9789 76.6154 U. montanus
2
Thiruvananthapuram district
Chathankod 140 08.6593 77.1494 U. anamalaiensis
1
Kallar, Ponmudi 180 08.8490 77.1322 U. anamalaiensis
1
Karamama 18 08.4780 76.9683 U. variegatus
2
Makki 227 08.6723 77.1301 U. anamalaiensis
2
Thrissur district
Mannuthy 40 10.5236 76.2940 U. taprobanicus
2
Meladoor 150 10.3167 76.3500 U. variegatus
2
Wayanad district
Kalpetta 950 11.6000 76.0833 U. triangularis
1
,
U. taprobanicus
2
Kurichiyarmala 1210 11.6006 75.9664 U. montanus
1
Mananthavady 1100 11.8333 76.0833 U. triangularis
1
Pozhuthana 731 11.5861 76.0309 U. triangularis
1
Settukunnu 823 11.6172 75.9913 U. montanus
1
Suganthagiri 790 11.5407 76.0242 U. triangularis
1
Sulthanbathery 908 11.6642 76.2653 U. taprobanicus
1
Thirunelli 850 11.9106 76.0002 U. montanus
1
......continued on the next page
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TAXONOMIC REVIEW OF GENUS UPERODON
TAB LE 1. (Continued)
Locality Altitude Coordinates Species
(metres) Latitude (°N) Longitude (°E)
Karnataka
Bangalore Urban district
Bangalore 900 13.064253 77.620233 U. variegatus
1
,
U. taprobanicus
1
Bannerghatta 910 12.7215 77.5680 U. systoma
1
Belgaum district
Londa 660 15.4433 74.5200 U. mormorata
2
Chikmagalur district
Bygoor 1042 13.3111 75.6128 U. mormorata
1
,
U. triangularis
1
Mudigere 900 13.1258 75.5979 U. triangularis
1
Hassan district
Kempholey 649 12.8566 75.6955 U. anamalaiensis
2
Sakleshpur 948 12.9356 75.7697 U. mormorata
2*
Sakleshpur 782 12.8680 75.7029 U. anamalaiensis
1
Kodagu district
Madikeri 1102 12.4528 75.7027 U. triangularis
1
Wat tak oll i, Coorg 67 5 12. 1266 75.78 60 U. triangularis
1
Shimoga district
Kachigebailu, Shimoga 639 13.9835 75.1099 U. mormorata
1
Uttara Kannada district
Castle Rock 622 15.4019 74.3299 U. mormorata
1
Dandeli 494 15.1669 74.6323 U. mormorata
1
Maharashtra
Kolhapur district
Amba 590 16.9167 73.7833 U. mormorata
1
Pune district
Pune 585 18.5641 73.8256 U. globulosus
1
Tam hini G hat 600 18.5278 73.5744 U. globulosus
2
Sindhudurg district
Amboli 733 15.9691 73.9882 U. mormorata
1
Goa
North Goa district
Anjuna 23 15.5742 73.7482 U. mormorata
1
South Goa district
Netravali 73 15.0891 74.2331 U. mormorata
1
Gujarat
Narmada district
Samot 527 21.6481 73.8566 U. mormorata
2
......continued on the next page
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Zootaxa 4384 (1) © 2018 Magnolia Press
TAB LE 1. (Continued)
Locality Altitude Coordinates Species
(metres) Latitude (°N) Longitude (°E)
Andhra Pradesh
West Godavari district
Ellore (= Eluru) 22 16.7205 81.0903 U. variegatus
2*
Koruturu, Srikakulam 170 17.4297 81.5397 U. variegatus
1
Odisha
Odisha (unspecified) – – – U. systoma
1
, U. taprobanicus
1
Ganjam district
“Russelconda” (=Bhanjanagar) 83 19.9358 84.5824 U. globulosus
2*
Khurdha district
Bhubaneswar, Utkal University
campus
43 20.3034 85.8401 U. variegatus
2
SRI LANKA
Ampara district
Ampara 30 07.3004 81.6688 U. rohani sp. nov.
2
Anuradhapura district
Anuradhapura 90 08.3333 80.4000 U. rohani sp. nov.
2
Mihintale 117 08.3549 80.5054 U. rohani sp. nov.
1
Nochchiyagama 70 08.2667 80.2167 U. rohani sp. nov.
2
Badulla district
Badulla 830 06.9953 81.0456 U. rohani sp. nov.
2
Mahiyanganaya 90 07.3167 80.9833 U. rohani sp. nov.
2
Galle district
Hiyare 125 06.0609 80.3240 U. nagaoi
1
Kanneliya Forest Reserve 150 06.2500 80.3300 U. nagaoi
2*
Hambantota district
Bundala 8 06.1990 81.2108 U. rohani sp. nov.
1
Palatupana, Kirinda 5 06.2667 81.4000 U. rohani sp. nov.
2
Kandy district
Kandy 530 07.2908 80.6369 U. obscurus
2
Panwila 652 07.3726 80.7326 U. obscurus
1
Peradeniya 480 07.2575 80.5963 U. obscurus
1
Pussellawa 900 07.0833 80.5000 U. obscurus
2
Udawatta Kele 465 07.3000 80.5833 U. obscurus
2
Kegalle district
Babilapitiya estate 120 07.2407 80.2140 U. obscurus
2
Kilinochchi district
Paranthan, Jaffna 6 09.4333 80.4000 U. rohani sp. nov.
2
Kurunegala district
Karawaddana 114 07.5115 80.4270 U. rohani sp. nov.
1
,
U. taprobanicus
1
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TAXONOMIC REVIEW OF GENUS UPERODON
TAB LE 1. (Continued)
Locality Altitude Coordinates Species
(metres) Latitude (°N) Longitude (°E)
Kurunegala 1 230 07.4704 80.3801 U. rohani sp. nov.
2
Kurunegala 2 400 07.4448 80.4080 U. obscurus
2
Makandura 25 07.3245 79.9887 U. rohani sp. nov.
2
Matale district
Dambulla 188 07.8601 80.6746 U. rohani sp. nov.
1
,
U. systoma
1
,
U. taprobanicus
1
Gammaduwa 1120 07.5885 80.7088 U. obscurus
2
Sigiriya 175 07.9500 80.7500 U. rohani sp. nov.
2
Nuwara Eliya district
Bogawantalawa 1160 06.8250 80.6300 U. obscurus
2
Diyagama estate, Agrapatana 1830 06.8289 80.7797 U. palmatus
2
Laxapana 1220 06.8333 80.5333 U. obscurus
2
Pattipola 1860 06.8500 80.8333 U. palmatus
2
Nuwara-Eliya 1907 06.9717 80.7866 U. palmatus
1
Morey estate 1424 06.7951 80.5240 U. palmatus
1
Polonnaruwa district
Polonnaruwa 55 07.9333 81.0000 U. rohani sp. nov.
2
Ratnapura district
Koskulana, Panapola, Sinharaja 460 06.4167 80.4500 U. obscurus
2
Morningside, Rakwana 1060 06.4000 80.6333 U. obscurus
2
Colonial locations
“Base of Anamalai Hills, Coimbatore
District”, India
U. anamalaiensis
*
“Bangalore (Mysore), S. India” U. variegatus
2*
“Ceylon” (=Sri Lanka) U. obscurus
2*
“Gadavery Valey” or “Gadavere”,
India
U. variegatus
2*
“India orientali” U. systoma
2*
“Malabar”, India U. montanus
2
,
U. triangularis
2*
“Newera Ellya”, Sri Lanka U. palmatus
2*
“Nilgiri Hills” or “Nilgheries”, India U. triangularis
2*
“Punduloya, Ceylon”, Sri Lanka U. taprobanicus
2*
“the Carnatic”, India U. variegatus
*
“Wynaad”, India U. montanus
2*
“Yellagherry Hills”, India U. variegatus
2*
1
Genetically and morphologically studied collections from the present study
2
Other morphologically studied collections and museum specimens
*
Typ e lo ca lity
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External morphology. The new collections were morphologically compared with all the available types and
other museum specimens. In cases where the type specimens are either lost or destroyed, morphological
identification of species was based on the original descriptions. Gonads were examined by a small lateral or ventral
incision to determine the sex and maturity of specimens. Only mature (adult) animals were used for morphological
studies. Measurements and associated terminologies follow Biju & Bossuyt (2006) and Biju et al. (2014a). The
following measurements were taken to the nearest 0.1 mm using a digital vernier caliper or a binocular microscope
with an ocular micrometre: SVL (snout-vent length), HW (head width, at the angle of the jaws), HL (head length,
from the rear of the mandible to the tip of the snout), MN (distance from the rear of the mandible to the nostril), SL
(snout length, from the tip of the snout to the anterior orbital border), EL (eye length, horizontal distance between
the bony orbital borders), IUE (inter upper eyelid width, shortest distance between the upper eyelids), UEW
(maximum upper eyelid width), IFE (internal front of the eyes, shortest distance between the anterior orbital
borders), IBE (internal back of the eyes, shortest distance between the posterior orbital borders), FAL (forearm
length, from the flexed elbow to the base of the outer palmar tubercle), HAL (hand length, from the base of the
outer palmar tubercle to the tip of the third finger), SHL (shank length), TL (thigh length, from the vent to the
knee), FOL (foot length, from the base of the inner metatarsal tubercle to the tip of the fourth toe), FD (disc width
of finger), FW (width of finger, measured at the base of the disc), TD (disc width of toe), TW (width of toe,
measured at the base of the disc), digit number is represented by roman numerals I–V. All measurements provided
in the taxonomy section are in millimetres. Webbing is described as by Biju et al. (2014b). Measurements and
photographs were taken for the right side of the specimen, except when a character was damaged, in which case the
measurement was taken on the left side. SDB and SG performed all the morphological studies.
Principal Component Analyses (PCA) were performed on 17 morphometric variables from adult male and
female specimens (separately) of Uperodon variegatus and the new species, using the software Statistica v7.1
(StatSoftInc). PCA factor scores for Principal Components (PC) with eigenvalues >1.0 were reported. Factor
scores of the first two components were visualized on scatterplots to assess the degree of morphological
differentiation between males and females of the two species.
TABLE 2. Samples used for the molecular study.
Species Collection Locality Voucher No./Source Accession No.
Present study
Uperodon anamalaiensis India: Karnataka: Sakleshpur ZSI/WGRC/V/A/957 MG557906
Uperodon anamalaiensis India: Kerala: Chathankod SDBDU 2013.2302 MG557907
Uperodon anamalaiensis India: Kerala: Chathankod SDBDU 2002.929 MG557908
Uperodon anamalaiensis India: Kerala: Methooty SDBDU 2012.1994 MG557909
Uperodon anamalaiensis India: Kerala: Methooty SDBDU 2012.1995 MG557910
Uperodon anamalaiensis India: Kerala: Parambikulam TR SDBDU 2011.548 MG557911
Uperodon globulosus India: Maharashtra: Pune SDBDU 2014.2672 MG557912
Uperodon globulosus India: Maharashtra: Pune SDBDU 2014.2682 MG557913
Uperodon globulosus India: Maharashtra: Pune SDBDU 2014.2685 MG557914
Uperodon montanus India: Kerala: Kesavapara SDBDU 2011.1197 MG557915
Uperodon montanus India: Kerala: Kurichiyarmala SDBDU 2006.4877 MG557916
Uperodon montanus India: Kerala: Munnar SDBDU 2005.30 MG557917
Uperodon montanus India: Kerala: Pandimotta SDBDU 2011.905 MG557918
Uperodon montanus India: Kerala: Kurichiyarmala SDBDU 2006.4846 MG557919
Uperodon montanus India: Kerala: Settukunnu SDBDU 2011.849 MG557920
Uperodon montanus India: Kerala: Thirunelli SDBDU 2011.228 MG557921
Uperodon montanus India: Tamil Nadu: Kakkachi SDBDU 2003.40116 MG557922
Uperodon montanus India: Tamil Nadu: Kakkachi SDBDU 2002.1925 MG557923
Uperodon montanus India: Kerala: Upper Manalar SDBDU 2015.3001 MG557924
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TAXONOMIC REVIEW OF GENUS UPERODON
TAB LE 2. (Continued)
Species Collection Locality Voucher No./Source Accession No.
Uperodon mormorata India: Goa: Anjuna SDBDU 2015.3082 MG557925
Uperodon mormorata India: Karnataka: Bygoor SDBDU 2011.507 MG557926
Uperodon mormorata India: Karnataka: Castle Rock SDBDU 2011.835 MG557927
Uperodon mormorata India: Karnataka: Dandeli SDBDU 2011.1331 MG557928
Uperodon mormorata India: Karnataka: Shimoga SDBDU 2012.43 MG557929
Uperodon mormorata India: Maharashtra: Amba SDBDU 2005.1763 MG557930
Uperodon mormorata India: Maharashtra: Amba SDBDU 2005.2520 MG557931
Uperodon mormorata India: Maharashtra: Amboli SDBDU 2004.4513 MG557932
Uperodon mormorata India: Maharashtra: Amboli SDBDU 2005.4757 MG557933
Uperodon mormorata India: Maharashtra: Amboli SDBDU 2012.2127 MG557934
Uperodon nagaoi Sri Lanka: Southern: Hiyare DZ 1356 MG557935
Uperodon nagaoi Sri Lanka: Southern: Hiyare DZ 1367 MG557936
Uperodon obscurus Sri Lanka: Central: Peradeniya DZ 1364 MG557937
Uperodon obscurus Sri Lanka: Central: Peradeniya DZ 1365 MG557938
Uperodon obscurus Sri Lanka: Central: Panwila DZ 1408 MG557939
Uperodon palmatus Sri Lanka: Central: Moray estate DZ 1275 MG557940
Uperodon palmatus Sri Lanka: Central: Nuwara-Eliya DZ 1331 MG557941
Uperodon palmatus Sri Lanka: Central: Nuwara-Eliya DZ 1371 MG557942
Uperodon rohani sp. nov. Sri Lanka: North Western: Karawaddana DZ 1112 MG557943
Uperodon rohani sp. nov. Sri Lanka: Central: Dambulla DZ 1129 MG557944
Uperodon rohani sp. nov. Sri Lanka: Central: Dambulla DZ 1130 MG557945
Uperodon rohani sp. nov. Sri Lanka: Southern: Bundala NP DZ 1336 MG557946
Uperodon rohani sp. nov. Sri Lanka: North Central: Mihintale DZ 1448 MG557947
Uperodon rohani sp. nov. Sri Lanka: North Central: Mihintale DZ 1449 MG557948
Uperodon systoma India: Tamil Nadu: Kunnapattu SDBDU 2005.4723 MG557949
Uperodon systoma India: Tamil Nadu: Coimbatore SDBDU 2005.4750 MG557950
Uperodon systoma India: Karnataka: Bannerghatta SDBDU 2014.2522 MG557951
Uperodon systoma India: Odisha SDBDU 2002.1151 MG557952
Uperodon systoma Sri Lanka: Central: Dambulla DZ 1128 MG557953
Uperodon triangularis India: Karnataka: Bygoor SDBDU 2010.332 MG557954
Uperodon triangularis India: Karnataka: Bygoor SDBDU 2011.503 MG557955
Uperodon triangularis India: Karnataka: Madikeri SDBDU 2012.2220 MG557956
Uperodon triangularis India: Karnataka: Madikeri SDBDU 2012.2221 MG557957
Uperodon triangularis India: Karnataka: Mudigere ZSI/WGRC/V/A/959 MG557958
Uperodon triangularis India: Kerala: Kalpetta SDBDU 2012.2038 MG557959
Uperodon triangularis India: Kerala: Mananthavady SDBDU 2003.40240 MG557960
Uperodon triangularis India: Kerala: Pozhuthana SDBDU 2014.2627 MG557961
Uperodon triangularis India: Kerala: Suganthagiri SDBDU 2014.2595 MG557962
Uperodon variegatus India: Andhra Pradesh: Srikakulam SDBDU 2007.4994 MG557963
Uperodon variegatus India: Karnataka: Bangalore SDBDU 2002.524 MG557964
Uperodon variegatus India: Tamil Nadu: Chengalpattu SDBDU 2005.4727 MG557965
Uperodon taprobanicus India: Karnataka: Bangalore SDBDU 2014.2567 MG557966
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Osteology. Adult specimens (N = 3) kept in 10% neutral buffered formalin were cleared and stained using
Alcian blue and Alizarin Red following Hanken & Wassersug (1981) and Taylor & Van Dyke (1985). Stained
specimens were scored for bone and cartilage within two days of completion of the procedure. Chondrocranial and
osteological terminologies follow Cannatella (1999) and Duellman & Trueb (1986), respectively. For comparisons,
the term “vomer” (Duellman & Trueb 1986) is used instead of “prevomer” (Trueb 1993), following previous
osteological literature on microhylids (e.g., Parker 1934; Ramaswami 1936; Peloso et al. 2016). Photographs were
taken either using a digital camera or captured with the aid of a QImaging camera mounted on Nikon SMZ 1500
stereomicroscope. GS performed the osteological study.
Larval study. The identity of the tadpoles was confirmed genetically using a 16S rRNA gene fragment.
Tadpoles were staged according to Gosner (1960). External morphology of the larvae was described following the
terminologies of Altig & McDiarmid (1999a) and Grillitsch et al. (1993). Tadpoles were measured to the nearest
0.01 mm using a digital vernier caliper. The following measurements (Altig & McDiarmid 1999a; Bowatte &
Meegaskumbura 2012) were taken for each tadpole stage: maximum height of body (bh), maximum width of body
(bw), maximum diameter of eye (ed), internarial distance (nn), naro-pupular distance (np), interpupular distance
(pp), rostro-narial distance (rn), distance from tip of snout to opening of spiracle (ss), distance from tip of snout to
insertion of upper tail fin (su), snout-vent length (svl), total length (tl), distance from vent to tip of tail (vt),
maximum height of tail (ht), tail-muscle height (tmh) and tail-muscle width (tmw). SG performed DNA barcoding
and GS described the larval morphology.
Bioacoustics. Male calls were recorded for three species using a Marantz PMD620 solid-state digital recorder
(44.1 kHz sampling rate, 16-bit resolution) and a Sennheiser ME 66 unidirectional microphone. The gain level of
the recorder was set at the beginning of each recording and the microphone was positioned at about 30–50 cm from
the calling individual. One specimen from each of the recorded populations was sampled for morphological and
genetic identification. Dry and wet bulb air temperatures were noted at each recording site. Five temporal
properties (call duration, call rise time, call fall time, number of pulses per call and pulse rate) and one spectral
TAB LE 2. (Continued)
Species Collection Locality Voucher No./Source Accession No.
Uperodon taprobanicus India: Kerala: Sulthanbathery SDBDU 2005.4732A MG557967
Uperodon taprobanicus India: Odisha SDBDU 2002.3107 MG557968
Uperodon taprobanicus India: Tamil Nadu: Mamallapuram SDBDU 2010.011 MG557969
Uperodon taprobanicus Sri Lanka DZ 1157 MG557970
Uperodon taprobanicus Sri Lanka: Central: Dambulla DZ 1133 MG557971
Uperodon taprobanicus Sri Lanka: North Western: Karawaddana DZ 1110 MG557972
GenBank
Uperodon mormorata
(as U. montanus)
––
AB612000
1
Uperodon palmatus
(as U. obscurus)
Sri Lanka MM 5980 KC180006
2
Uperodon systoma Indian subcontinent VUB1151 EF017960
3
Uperodon variegatus India VUB1342 EF017963
3
Uperodon taprobanicus Sri Lanka VUB0102 AF249057
4
Uperodon taprobanicus Sri Lanka KUHE: 37252 AB634691
5
Outgroups
Kaloula pulchra China NMNS 3208 KC822614
6
Kaloula mediolineata Thailand: Wang Nam Kieow: Sakaerat KU 328280 KC822509
6
Kaloula conjuncta Philippines: Luzon Island: Palola ACD 769 KC822537
6
1
Kurabayashi et al. 2011;
2
de Sá et al. 2012;
3
Va n B oc x la e r et al. 2006;
4
Bossuyt & Milincovitch 2000;
5
Matsui et al. 2011;
6
Blackburn et al. 2013
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TAXONOMIC REVIEW OF GENUS UPERODON
property (dominant frequency) for a single call of each species were measured using Raven Pro 1.4 (Charif et al.
2010). Spectrograms were prepared for graphical representation of the call structure. Terminology and call property
definitions follow Bee et al. (2013). SG and SDB performed the acoustic study.
Museum and other abbreviations. For museums and frequently used terms, abbreviations are as follows:
SDBDU (Systematics Lab, University of Delhi, India), DZ (Department of Molecular Biology & Biotechnology,
University of Peradeniya, Sri Lanka), NHM (Natural History Museum, London, United Kingdom) formerly
BMNH (British Museum (Natural History), London, United Kingdom), ZSIC (Zoological Survey of India,
Kolkata, India), ZSI/WGRC (Zoological Survey of India–Western Ghats Regional Centre, formerly Zoological
Survey of India–Western Ghats Field Research Station, Calicut, India), BNHS (Bombay Natural History Society,
Mumbai, India), WHT (Wildlife Heritage Trust of Sri Lanka, Colombo), HT (Holotype), NT (Neotype), PT
(Paratype), LT (Lectotype), PLT (Paralectotype), ST (Syntype), TT (Topotype), NP (National Park), WLS (Wildlife
Sanctuary), TR (Tiger Reserve), SDB (SD Biju), SG (Sonali Garg), MM (Madhava Meegaskumbura), GS (Gayani
Senevirathne) and NW (Nayana Wijayathilaka).
Results
DNA analyses. Genetic differentiation among all the Uperodon species was assessed based on mitochondrial 16S
rRNA gene. The resultant alignment comprised of 511 bp (including gaps) of which 501 unambiguously aligned
positions with 108 variable sites and 97 parsimony-informative sites were used for phylogenetic analyses. The ML
and Bayesian analyses recovered 12 major lineages with significant support (BPP 0.99–1.0, BS 80–100%) (Fig. 1).
These corresponded to 11 previously established Uperodon species (except U. minor) and one additional divergent
lineage. The candidate species from Sri Lanka was delineated based on considerable genetic distance of 2.9%
(range 2.4–3.5%) from its closest relative U. variegatus known from India (Table 3). This new lineage was highly
supported (BPP 1.0, BS 99%). Interspecific uncorrected genetic divergence between the previously known
Uperodon species ranged from 2.4–9.1% (Table 3), which are consistent or slightly lower than the species-level
distances observed in other anuran groups from the region (e.g., Biju et al. 2014a; Biju et al. 2014b; Garg & Biju
2016; Wijayathilaka et al. 2016; Garg et al. 2017; Garg & Biju 2017). The observed intraspecific distances reached
a maximum of 2.2% and the following major sub-lineages (>1%) were observed: (1) Uperodon triangularis
comprised of two sub-lineages, one identified as the typical form from the ‘Malabar’ region of Kerala, and the
second with populations from Karnataka that most likely correspond to the subspecies Ramanella triangularis
rufeventris (for detailed discussion see the taxonomic account for Uperodon triangularis); (2) Uperodon
mormorata also showed two sub-lineages that correspond to populations from two different states, Karnataka
(typical) and Maharashtra, respectively; (3) Uperodon variegatus populations from the Eastern Ghats (Andhra
Pradesh and Tamil Nadu) and Western Ghats (Karnataka and Kerala) formed separate sub-lineages; (4) Uperodon
systoma sample from Bannerghata (Karnataka) showed a divergence of 2.2% from the other populations; (5)
Uperodon taprobanicus population from the Eastern Ghats (Odisha) differed genetically from all other Western
Ghats and Sri Lankan populations by 1.0–1.2%; and (6) the candidate species also showed intraspecific divergence
of up to 1.4% (see taxonomic description of the new species for discussion). In addition, U. montanus populations
found north and south of the Palghat gap were divergent by up to 1.2%. For U. anamalaiensis, a single population
from ‘Sakleshpur’ in Karnataka that is likely to correspond with U. minor showed shallow divergence (0.8%) from
populations found south of the Palghat gap in Kerala. For intra- and interspecific pairwise comparisons among all
the members see Table 3, and for detailed taxonomic notes and implications see the respective species accounts in
the ‘Taxonomy’ section.
Taxonomy
Detailed morphological comparison with available types, museum specimens and original descriptions, shows that
the newly identified lineage from Sri Lanka represents a distinct species. This species is formally described as
Uperodon rohani sp. nov. based on molecular and morphological evidence. In anticipation of this outcome, we
refer to the species under its new name in the following sections, prior to its formal taxonomic description, which is
provided subsequently.
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FIGURE 1. Maximum Likelihood phylogram based on 511 bp of mitochondrial 16S rRNA gene sequences from 73
individuals representing all known members of the genus Uperodon. Three samples representing taxon under synonymy are
shown in quotation marks. Outgroup not shown for graphical purposes. Bayesian Posterior Probabilities and bootstrap values of
>50 are shown above and below the branches, respectively. Values towards the species labels represent mean uncorrected
genetic distances between the species with range values in parentheses.
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FIGURE 2. A. Geographic distribution of the genus Uperodon in Bangladesh, Bhutan, India, Nepal, Pakistan and Sri Lanka;
B. Distribution of Uperodon globulosus, U. systoma and U. taprobanicus; C. Distribution of four endemic Uperodon species of
the Western Ghats (excluding the widely distributed Indian member U. variegatus, shown in Fig. 12A); D. Distribution of four
endemic Uperdodon species of Sri Lanka. Details of localities reported in the present study are provided in Table 1. Records
from literature are listed in supplementary Table S1 and indicated here based on approximate coordinates.
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Order Anura
Family Microhylidae
Subfamily Microhylinae
Genus Uperodon
Type species Uperodon systoma
Uperodon Duméril and Bibron, 1841. Erpétologie Genérale ou Histoire Naturelle Complète des Reptiles. Volume 8. Paris:
Librarie Enclyclopedique de Roret. Type species. Engystoma marmoratum Guérin-Méneville, 1838 (= Rana systoma
Schneider, 1799) by monotypy.
Hyperodon Agassiz, 1846. Nomenclator Zoologicus Continens Nomina Systematica Generum Animalium tam Viventium quam
Fossilium. Fasicle 12 (Indicem Universalem). Soloduri: Jent et Gassmann.
Cacopus Günther, 1864 “1863”. The Reptiles of British India. London: Ray Society by R. Hardwicke.
Ramanella Rao and Ramanna, 1925. On a new genus of the family Engystomatidae. Proceedings of the Zoological Society of
London, 1925: 587–597.
FIGURE 3. Preserved type specimens of Uperodon species in dorsal and ventral view. A. Uperodon systoma, Syntype, ZMB
3551 (female); B. Uperodon globulosus, Syntype, NHM 62.12.29.4 (ex. BMNH 1947.2.10.97) (female); C. Kaloula pulchra
taprobanica, Holotype, NHM 1905.3.25.138 (ex. BMNH 1947.2.10.62) (female); D. Ramanella anamalaiensis, Neotype, ZSI/
WGRC/V/A/956 (male); E. Hylaedactylus montanus, Holotype, NHM 72.4.17.212 (ex. BMNH 1947.2.11.19) (female); F.
Ramanella mormorata, Neotype, ZSI/WGRC/V/A/958, (female); G. Ramanella nagaoi, Holotype, WHT 2826 (male); H.
Callula obscura, Holotype, NHM 58.11.28.42 (ex. BMNH 1947.2.11.20) (female); I. Ramanella palmata, Holotype, NHM
90.11.8.62 (ex. BMNH 1947.2.11.21); J. Callula triangularis, Lectotype, NHM 74.4.29.897 (ex. BMNH 1947.2.11.15) (male);
K. Hylaedactylus carnaticus, Lectotype, ZSI 2761 (female); L. Uperodon rohani sp. nov., Holotype, DZ 1449 (male). (Images:
SD Biju and Sonali Garg).
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Common name. Globular Frogs (Frank & Ramus 1995)
Diagnosis. Small to large-sized frogs (male SVL 21–56 mm, female SVL 28–72 mm) with globular
appearance; head wider than long; pupil oval; skin shagreened to glandular; tympanum hidden; tongue emarginate
posteriorly, with or without lingual papillae; fingers free, dermal fringes present, tips of fingers with truncate discs
(except in Uperodon globulosus and U. systoma: rounded, slightly enlarged without discs), without dorsoterminal
grooves; webbing between fingers absent; webbing between toes present or absent; prevomers divided or
undivided, clavicles absent, and procoracoids rudimentary (Parker 1934; Ramaswami 1936).
Distribution. Genus Uperodon is restricted to the Indian subcontinent. It is known to occur widely in India and
Sri Lanka, with some reports from adjacent countries, namely, Bangladesh, Bhutan, Nepal and Pakistan. The
distribution of the genus extends from 05.9°–33.7° N and 70.5°–95.4° E (Fig. 2). Of the 12 nominal species, eight
are restricted to the Western Ghats–Sri Lanka biodiversity hotspot region, with one Indian member (U. variegatus)
more widely distributed outside the Western Ghats, while three species occur across the entire known range of the
genus.
Uperodon systoma (Schneider, 1799)
Marbled Globular Frog
(Figs. 1, 2A–B, 3A, 4A–F, 5A; Tables 1–4)
Original name and description. Rana systoma Schneider, 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, 144–146. Syntype. ZMB 3551, original description mentions
two specimens, one likely to be lost. Type locality. “India orientali”, India. Current status of specific name. Valid
name, as Uperodon systoma (Schneider, 1799). Synonyms. Engystoma marmoratum Cuvier, 1829 (Duméril &
Bibron 1841; Peters 1864 “1863”) and Systoma leschenaultii Tschudi, 1838 (Duméril & Bibron 1841).
Note on doubtful synonym. Pachybatrachus Petersii Keferstein, 1868 described from “Neu-Süd-Wales”
Australia was considered as a synonym of Uperodon systoma (Schneider 1799) by Boulenger (1882) and followed
by Parker (1934). However, if this taxon was indeed described from Australia (although it might be erroneous
according to Frost 2017) it cannot be homogenous with Uperodon systoma or any other member of the genus
Uperodon, which is clearly restricted to South Asia. Although, we could not examine the type of Pachybatrachus
petersii Keferstein, 1868, its synonymy with Uperodon systoma is considered doubtful.
Comments. The original description of Rana systoma (Schneider 1799) mentions two specimens (= syntypes)
from “India orientali” but the description is likely to have been based on a single specimen (Bauer 1998). Currently
a single specimen is available in the ZMB (Berlin) collection (Peters 1864 “1863”; Bauer 1998). We examined the
available syntype (an adult female) and found it to clearly match the original description of the species. However,
since at least two specimens were originally catalogued for this taxon and the original description was brief, we
herein provide a detailed description of the available type specimen, ZMB 3551 (an adult female), as a reference of
this species for future taxonomic works.
Comparison. Uperodon systoma differs from all other known members of the genus, except U. globulosus and
U. taprobanicus, by its larger snout-vent size, male SVL 51–56 mm, female SVL 47–64 mm (vs. smaller, male
SVL 21–40 mm, female SVL 28–43 mm); finger and toe tips rounded, without discs (vs. finger tips enlarged in to
discs, except in U. globulosus); and presence of two dermal projections on the neopalatinal ridges on posterior side
of each choana, widely separated from the vomerine odontophores (vs. prominent dermal projections on the
neopalatinal ridges on posterior side of each choana, narrowly separated from the vomerine odontophores in U.
globulosus; indistinct or weakly to well-developed neopalatinal ridges on posterior side of each choana, fused with
the vomerine odontophores in all other species) (Fig. 5A). Uperodon systoma differs from U. globulosus and U.
taprobanicus in having divided prevomers, as in the rest of Uperodon species (vs. undivided prevomers in U.
globulosus and U. taprobanicus) (Parker 1934); and by its dorsum being light or dark brown with prominent
yellow markings (vs. light brown to greyish-brown without any prominent markings in U. globulosus; light to dark
grey with prominent red or reddish-orange blotches in U. taprobanicus). Specifically it also differs from U.
globulosus by its snout length nearly equal to eye diameter, male EL/SL ratio 95.3–104%, N = 4, female EL/SL
ratio 98–106%, N = 1 (vs. snout nearly twice as long as diameter of eye, male EL/SL ratio 40.5–50.0%, N = 3,
female EL/SL ratio 44.4–54.8%, N = 1) (Table 4).
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FIGURE 4. A–D. Uperodon systoma (Syntype, ZMB 3551) in preservation: A. lateral view of head; B. ventral view of hand;
C. ventral view of foot; D. schematic illustration of foot webbing; E–F. Uperodon systoma (SDBDU 2005.2732, female) in
life: E. dorsolateral view; F. anterior view; G–J. Uperodon globulosus (Syntype, NHM 62.12.29.4 ex. BMNH 1947.2.10.97) in
preservation: G. lateral view of head; H. ventral view of hand; I. ventral view of foot; J. schematic illustration of foot webbing;
K–L. Uperodon globulosus (SDBDU 2014.2672, male) in life: K. anterior view (not preserved); L. dorsolateral view; M–P.
Uperodon taprobanicus (Holotype, NHM 1905.3.25.138 ex. BMNH 1947.2.10.62) in preservation: M. lateral view of head; N.
ventral view of hand; O. ventral view of foot; P. schematic illustration of foot webbing. (Images: SD Biju).
FIGURE 5. Ventral surface of the palate in Uperodon species showing vomerine odontophores separated or fused with dermal
projections and/or neopalatinal ridges on posterior sides of the choanae (indicated with arrows). A. Uperodon systoma, SDBDU
2005.2787 (female) and SDBDU 2005.2806 (male); B. Uperodon globulosus, SDBDU 2014.2673 (female) and SDBDU
2005.4762 (female); C. Uperodon taprobanicus, SDBDU 2007.6364 (female) and SDBDU 2005.4732A (male); D. Uperodon
anamalaiensis, SDBDU 2013.2302 (female) and SDBDU 2002.3050 (male); E. Uperodon montanus, SDBDU 2002.24B
(female) and SDBDU 2003.40115 (male); F. Uperodon mormorata, SDBDU 2011.507 (male) and SDBDU 2012.43 (male); G.
Uperodon obscurus, DZ 1366 (male); H. Uperodon palmatus, DZ 1331 (male); I. Uperodon triangularis, SDBDU 2007.6400
(female) and ZSI/WGRC/V/A/959 (male); J. Uperodon variegatus, SDBDU 2005.4726 (female) and SDBDU 2005.4727
(male); K. Uperodon rohani sp. nov., DZ 1521 (male). (Images: SD Biju).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon systoma
showed an average intraspecific distance of 0.9% (range 0–2.2%, N = 6). The sample from Bannerghatta
(Karnataka) showed considerable divergence of up to 2.2% from the typical populations in Tamil Nadu, excluding
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which, U. systoma showed an average intraspecific distance of 0.5% (range 0–0.8%, N = 5). Genetically, U.
systoma is closely related to U. globulosus, from which it was found to differ by an average uncorrected genetic
distance of 3.5% (range 3.2–4.0%, N = 18). For interspecific genetic distances with all other members of the genus,
see Table 3.
Description of syntype, ZMB 3551 (measurements in mm) (Figs. 3A, 4A–D). Large-sized (SVL 49.7),
globular and robust adult female; head small (HW 11.7, HL 7.9, IFE 6.6, IBE 12.4), less than one-sixth (15.9%) of
body length, wider than long (HW/HL ratio 1.5); snout semi-circular to nearly rounded in dorsal and ventral view,
rounded in lateral view, its length (SL 4.8) nearly equal to horizontal diameter of eye (EL 4.9); loreal region obtuse
with rounded canthus rostralis; interorbital space (IUE 4.8) nearly twice as wide as upper eyelid width (UEW 2.6);
nostril oval without lateral flap of skin, nearly equidistant to tip of the snout (NS 1.6) and the eye (EN 1.4);
tympanum not visible externally, distinct supratympanic fold extending from posterior corner of upper eyelid near
to insertion of forelimb at axilla; eye diameter (EL 4.9); vomerine odontophores present on the palate, prominent,
without teeth; presence of two prominent dermal projections on the neopalatinal ridges on posterior side of each
choana, widely separated from the vomerine odontophores; tongue moderately large, rounded, shallowly
emarginated. Forelimbs moderately short and strong; forearm (FAL 13.2) shorter than hand length (HAL 23.8);
finger length formula I<IV<II<III (FI 3.5, FII 5.1, FIII 6.7, FIV 3.6); tips of all fingers rounded, without discs;
webbing absent between fingers; subarticular tubercles weakly-developed, oval, all present; two well-developed
palmar tubercles (inner, oval, 1.3 mm long; outer, bilobed, 1.7 mm long). Hind limbs relatively short and strong,
thigh length (TL 18.3) longer than shank (SHL 15.4) and shorter than foot (FOL 18.6); tips of all toes rounded,
without discs; webbing present, small: I2–2II1
3
/
4
–3III3–4
–
IV4–2
3
/
4
V; well-developed dermal fringes present on all
toes; subarticular tubercles weakly-developed, oval, all present; two shovel-shaped metatarsal tubercles (inner, 2.3
mm long; outer, 4.3 mm long).
Skin of dorsum, sides of head, and dorsal parts of forelimb, thigh, tibia and tarsus, shagreened (Fig. 3A);
posterior parts of thigh, especially cloacal region, granular. Ventral surfaces shagreened and wrinkled (Fig. 3A).
Colouration. In preservation (ZMB 3551): Dorsum brown with prominent light grey elongate blotches, flanks
lighter in colour than dorsum; forelimbs, dorsal surfaces of thigh, shank and foot, light creamish-brown with darker
brown markings; chest, belly, and ventral surfaces of thigh, tibia and foot grey with creamish-brown (Figs. 3A, 4A–
D). In life (SDBDU 2005.2732): Dorsum brown with prominent yellow blotches, dorsal surface of head yellowish
brown, upper eyelids yellow; lateral surfaces of head and flank lighter in colour compared to dorsum; forelimbs,
dorsal surfaces of thigh, shank and foot, light brown with yellow blotches; throat, chest, and belly, greyish-white
without any marking, ventral surfaces of forelimb, thigh, tibia and foot, white (Figs. 4E–F).
Variations. Morphometric measurements for six specimens are given in Table 4. SDBDU 2005.2806:
subarticular tubercles on fingers weakly-developed, throat greyish-white with light brown marbling; SDBDU
2005.2787: Skin of dorsum and limbs coarsely glandular, posterior parts of thigh especially the cloacal region
prominently glandular, and throat coarsely granular.
Secondary sexual characters. Male (SDBDU 2005.2787): vocal sac externally visible on the lower jaw;
female (SDBDU 2005.2806): ova white, pigmented on pole (diameter 0.9–1.3 mm, N = 10).
Geographical distribution and habitat. Uperodon systoma is widely distributed in India and Sri Lanka, and
also reported from adjoining regions in Nepal and Pakistan (Fig. 2B). In India, it has been reported from the states
of Tamil Nadu, Kerala, Karnataka, Maharashtra, Gujarat, Andhra Pradesh, Telangana, Odisha, Madhya Pradesh,
Chhattisgarh, Rajasthan, Uttar Pradesh, Himachal Pradesh, West Bengal, Assam and Mizoram. In Sri Lanka, the
species is known from Central, Eastern, Northern, North Western, Sabaragamuwa, Southern and Uva Provinces.
For detailed list of localities and references thereto see Supplementary Table S1. In the present study, this species
was found in Tamil Nadu (Coimbatore and Kunnapattu), Karnataka (Bannerghatta) and Odisha in India, and
Dambulla (Central Province) in Sri Lanka (Table 1). Individuals were collected from soil surface close to
temporary pools or dry burrows in agricultural fields. See ‘Natural history and breeding’ observations in the
respective section, after the species accounts.
Uperodon globulosus (Günther, 1864 “1863”)
Indian Globular Frog (Frank & Ramus 1995)
(Figs. 1, 2A–B, 3B, 4G–L, 5B; Tables 1–4)
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Original name and description. Cacopus globulosus Günther, 1864 “1863”. The Reptiles of British India.
London: Ray Society by R. Hardwicke, p. 416. Syntypes. NHM 62.12.29.4 (ex. BMNH 1947.2.10.97) and NHM
62.12.29.3 (ex. BMNH 1947.2.10.77). Type loc ali ty. “Russelconda (Madras Presidency)”, India. Current status
of specific name. Valid name, as Uperodon globulosus (Günther, 1864 “1863”).
Comments. The original description (Günther 1864 “1863”) mentions that “two specimens” (= syntypes) from
“Russelconda (Madras Presidency)” collected by Dr. Trail were used for the description of this species. We
examined the two syntypes available at NHM, London: (1) a gravid female, NHM 62.12.29.4 (ex. BMNH
1947.2.10.97), SVL 67.4 mm, and (2) a sub-adult, NHM 62.12.29.3 (ex. BMNH 1947.2.10.77), SVL 26.9 mm. The
female specimen that is also illustrated with the original description (Plate XXVI, Fig. K) was found to be in
relatively good condition (Fig. 3B). Since the original description was brief, we herein provide a detailed
description of the adult female specimen, NHM 62.12.29.4 (ex. BMNH 1947.2.10.97) that may serve as a reference
for Uperodon globulosus (Günther, 1864 “1863”) in future taxonomic works.
Comparison. Uperodon globulosus differs from all other known members of the genus, except U. systoma and
U. taprobanicus, by its larger snout-vent size, male SVL 51–56 mm, female SVL 66–72 mm (vs. smaller, male
SVL 21–40 mm, female SVL 28–43 mm); tips of fingers rounded, without discs (vs. finger tips enlarged into discs,
except in U. systoma); and presence of two prominent dermal projections on the neopalatinal ridges on posterior
side of each choana, narrowly separated from the vomerine odontophores (vs. dermal projections widely separated
from the vomerine odontophores in U. systoma; indistinct or weakly to well-developed neopalatinal ridges on
posterior side of each choana, fused with the vomerine odontophores in all other species) (Fig. 5B). Uperodon
globulosus also differs from all other known Uperodon species, except U. taprobanicus, in having undivided
prevomers (vs. divided prevomers), and differs from U. taprobanicus by its dorsum being light brown to greyish-
brown without any prominent markings (vs. light to dark grey with prominent red or reddish-orange blotches). For
more differences with U. systoma see the ‘Comparison’ section for that species.
Genetic divergence. For 16S mitochondrial gene sequences, intraspecific variation was not observed between
the sampled populations of Uperodon globulosus (N = 3). Genetically, U. globulosus is closely related to U.
systoma, from which it was found to differ by an average uncorrected genetic distance of 3.5% (range 3.2–4.0%, N
= 18). For interspecific genetic distances with all other members of the genus, see Table 3.
Description of syntype, NHM 62.12.29.4 (measurements in mm) (Figs. 3B, 4G–J). Large-sized (SVL 67.4),
globular and robust adult female; head small (HW 19.2, HL 13.0, IFE 9.7, IBE 13.9), about one-fifth (19.3%) of
body length, wider than long (HW/HL ratio 1.5); snout nearly truncate in dorsal and ventral view, obtuse in lateral
view, its length (SL 7.7) more than twice as long as horizontal diameter of eye (EL 3.6); loreal region obtuse, with
rounded canthus rostralis; interorbital space over three times wider (IUE 8.8) than upper eyelid width (UEW 2.7);
nostril oval without lateral flap of skin, closer to tip of snout (NS 2.8) than to eye (EN 3.3); tympanum not visible
externally, distinct supratympanic fold extending from posterior corner of upper eyelid near to insertion of forelimb
at axilla; eye diameter (EL 3.6); vomerine odontophores present on the palate, prominent, without teeth; presence
of two prominent dermal projections on the neopalatinal ridges on posterior side of each choana, narrowly
separated from the vomerine odontophores (Fig. 5B); tongue moderately large, rounded, shallowly emarginated.
Forelimbs moderately short and strong, forearm (FAL 14.5) shorter than hand length (HAL 18.0); finger length
formula IV<I<II<III; tips of all fingers rounded, without discs; webbing absent between fingers; subarticular
tubercles weakly-developed, oval, all present; two well-developed palmar tubercles (inner, oval, 3 mm long; outer,
bilobed, 4 mm long). Hind limbs relatively short and strong, thigh length (TL 29.1) longer than shank (SHL 21.7)
and foot (FOL 26.5); tips of all toes rounded, without discs; webbing present, small: I1
1
/
2
–2
–
II2
–
–3
–
III2
1
/
2
–3
1
/
2
IV4
–
–
2
1
/
2
V; well-developed dermal fringes present on all toes; subarticular tubercles weakly-developed, oval, all present;
two shovel-shaped metatarsal tubercles (inner, 8.6 mm; outer, 4.8 mm).
Skin of dorsum, sides of head, and dorsal parts of forelimb, thigh, tibia and tarsus, shagreened (Fig. 3B);
posterior parts of thigh, especially cloacal region, distinctly granular. Ventral surfaces shagreened and wrinkled
(Fig. 3B).
Colouration. In preservation (NHM 62.12.29.4): Dorsum brown, flanks light greyish-brown; forelimbs,
dorsal surfaces of thigh, shank and foot, dark greyish-brown; chest, belly, and ventral surfaces of thigh, tibia and
foot, light greyish-brown (Figs. 3B, 4G–I). In life (SDBDU 2014.2672): Dorsum greyish-brown, dorsal surface of
head grey with bluish tinge, lateral surfaces of head and flank lighter in colour compared to dorsum, with yellow
tinge; forelimbs, dorsal surfaces of thigh, shank and foot, greyish-brown with yellow tinge; throat, chest, and belly,
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greyish-white without any markings; ventral surfaces of forelimb, thigh, tibia and foot, light grey with cream
coloured blotches (Figs. 4K–L).
Variations. Morphometric measurements for five specimens are given in Table 4. SDBDU 2014.2673 (female)
has prominent neopalatinal ridges on posterior side of each choana (Fig. 5B).
Secondary sexual characters. Male (SDBDU 2014.2672): single vocal sac externally visible on the lower
jaw; female (syntype): ova white, pigmented on pole (diameter 0.9–1.4 mm, N = 20).
Tad pol e morpholog y. In dorsal view, tadpoles of Uperodon globulosus (Gosner stage 32, N = 2) have an
elliptical body with narrower and shorter anterior region, wider and longer posterior region; snout appears rounded
in dorsal and lateral view (from eyes to base of upper labium); eyes small, bulbous, positioned laterally;
umbraculum absent; nasolacrimal duct absent; narial depressions clearly visible in lateral view, located much
closer to snout than to eyes, pigmentation not visible around nasal depressions; sinistral spiracle present, opening
located at ventral tail-musculature margin; tubular vent tube present posteriorly, with aperture opening medially in
line with plane of the ventral fin; tail ends bluntly, composed of bilateral myotomic muscle masses divided by V-
shaped septa, with unequal membranes on either side of tail musculature, tail musculature extends up to the end of
tail tip; dorsal fins originate after tail-body junction, margin of lower fin not parallel to the margin of tail muscle;
mouth (oral disc) absent, keratodonts and jaw sheaths not seen, papillae absent.
Measurements (mm) (N = 1): Lateral measurements include, maximum height of body (bh) = 3.66, maximum
width of body (bw) = 5.98, maximum diameter of eye (ed) = 0.40, internarial distance (nn) = 2.07, naro-pupular
distance (np) = 2.58, rostro-narial distance (rn) = 0.02, interpupular distance (pp) = 5.78, distance from tip of snout
to opening of spiracle (ss) = 4.55, distance from tip of snout to insertion of upper tail fin (su) = 6.60, snout-vent
length (svl) = 7.74, total length (tl) = 19.15, distance from vent to tip of tail (vt) = 11.60, maximum height of tail
(ht) = 4.95, tail muscle height (tmh) = 1.61, tail muscle width (tmw) = 1.35.
Geographical distribution and habitat. Uperodon globulosus is widely distributed in India and neighboring
regions in Bangladesh, Bhutan and Nepal (Fig. 2B). In India, this species has been reported from the states of
Andhra Pradesh, Maharashtra, Gujarat, Odisha, Madhya Pradesh, Chhattisgarh, Uttar Pradesh, Jharkhand, Odisha,
West Bengal, Assam and Arunachal Pradesh (for detailed list of localities and references thereto see supplementary
Table S1). In the present study, we collected specimens from temporary pools and puddles inside the Pune city
(Table 1). At Tamhini Ghats, a large number of males were observed on the road passing through open or
agricultural areas, immediately after the first monsoon showers. See ‘Natural history and breeding’ observations in
the respective section, after the species accounts.
Uperodon taprobanicus (Parker, 1934)
Painted Globular Frog
(Figs. 1, 2A–B, 3C, 4M–P, 5C, 6A–C; Tables 1–4)
Original name and description. Kaloula pulchra taprobanica Parker, 1934. A Monograph of the frogs of the
family Microhylidae. London: Trustees of the British Museum, 86–87. Holotype. By original designation, NHM
1905.3.25.138 (ex. BMNH 1947.2.10.62), an adult female, SVL 54.9 mm. Type lo cali ty. “Punduloya”, Sri Lanka.
Current status of specific name. Valid name, as Uperodon taprobanicus (Parker, 1934).
Comment. Even though Parker (1934) provided a description for this nominal taxon, considering its recent
taxonomic placement in the genus Uperodon, here we provide a detailed description of the holotype and
comparison of the species with all its presently recognized congeners.
Comparison. Uperodon taprobanicus differs from all other known members of the genus by presence of two
well-developed neopalatinal ridges on posterior side of each choana, fused with the vomerine odontophores (vs.
prominent dermal projections on the neopalatinal ridges on posterior side of each choana, separated from the
vomerine odontophores in U. globulosus and U. systoma; indistinct or weakly-developed neopalatinal ridges in all
other species), and its dorsum being light to dark grey with prominent red or reddish-orange blotches (vs. absent).
Specifically, it differs from U. globulosus and U. systoma by its smaller snout-vent size, male SVL 42–52 mm,
female SVL 53–55 mm (vs. larger, U. globulosus: male SVL 51–56 mm, female SVL 66–72 mm; U. systoma: male
SVL 51–56 mm, female SVL 47–64 mm), and tips of fingers with discs (vs. rounded, without discs); and from
other members of the genus by its larger snout-vent size, male SVL 42–52 mm, female SVL 53–55 mm (vs.
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smaller, male SVL 21–40 mm, female SVL 28–43 mm). Uperodon taprobanicus also differs from all other known
Uperodon species, except U. globulosus, in having undivided prevomers (vs. divided prevomers).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon
taprobanicus showed an average intraspecific distance of 0.3% (range 0–1.2%, N = 9). The sample from Odisha
showed a divergence of up to 1.2% from the typical populations. Genetically, U. taprobanicus is the most divergent
member of the genus and shows high interspecific genetic distances with all the other known members (see Table 3).
Redescription of holotype (measurements in mm) (Figs. 3C, 4M–P). Large-sized (SVL 54.9), globular and
robust adult female; head small (HW 16.5, HL 12.4, IFE 8.0, IBE 14.9), less than one-fourth (22.6%) of body
length, wider than long (HW/HL ratio 1.3); snout truncate in dorsal and ventral view, vertical in lateral view, its
length (SL 5.2) longer than horizontal diameter of eye (EL 4.7); loreal region obtuse with rounded canthus
rostralis; interorbital space wider (IUE 5.7) than upper eyelid (UEW 3.8); nostril oval without lateral flap of skin,
closer to tip of snout (NS 1.6) than to eye (EN 2.9); tympanum not visible externally, distinct supratympanic fold
extending from posterior corner of upper eyelid near to insertion of forelimb at axilla; eye diameter (EL 4.7);
vomerine odontophores present on the palate, without teeth; presence of two prominent neopalatinal ridges on
posterior side of each choana, fused with the vomerine odontophores (Fig. 5C); tongue moderately large, rounded,
shallowly emarginated. Forearms (FAL 16.8) shorter than hand length (HAL 24.5); finger length formula
I<II<IV<III; tips of all fingers with truncate discs; webbing absent between fingers; subarticular tubercles rather
well-developed, oval, all present; two well-developed palmar tubercles, inner one oval and smaller, and outer one
bilobed. Hind limbs relatively short and strong, thigh length (TL 20.8) longer than shank (SHL 17.2) and foot (FOL
19.9); tips of all toes rounded, slightly enlarged, without discs; webbing present, small: I2–2II2
–
–3III2–3
1
/
3
IV3
1
/
3
–
2V; weakly-developed dermal fringes present on all toes; subarticular tubercles well-developed, oval, all present;
two shovel-shaped metatarsal tubercles (inner, 1.4 mm long; outer, 3.1 mm long).
Skin of dorsum, sides of head, and dorsal parts of forelimb, thigh, tibia and tarsus, shagreened with scattered
glandular projections (Figs. 3C, 4M–P); posterior parts of thigh, especially cloacal region, distinctly granular.
Ventral surface of throat granular, chest and belly, sparsely granular (Fig. 3C).
Colouration. In preservation: Dorsum light brown with large irregular greyish-brown blotches, a dark grey
stripe between eyes, flanks and lateral sides of abdomen lighter brown in colour compared to dorsum; chest, belly
and ventral surfaces of thigh, tibia and foot, light greyish-brown with dark brown mottling (Fig. 3C). Colour in life
(SDBDU 2014.2567): Dorsal surface of head, anterior and posterior parts of back, brown and having brick-red
blotches with dark brown margins, brick-red stripe with dark brown margin between eyes, lateral surfaces of head
light brick-red, and flanks light brown with grey reticulations; forelimbs, dorsal surfaces of thigh, shank and foot,
light brown with dark blackish-brown spots and brick red patches; fingers and toes light brown with grey spots;
anterior part of thigh and groin brown with light grey reticulations; throat, chest and belly, light reddish-brown with
uniform light grey reticulations except on fingers and toes (Figs. 6A–C).
Variations. Morphometric measurements for five specimens, including the holotype, are given in Table 4.
SDBDU 2005.3107: dorsal skin sparsely granular; throat prominently granular, chest and belly sparsely granular;
anterior parts of thigh prominently granular. SDBDU 2005.4732A: dorsal skin coarsely granular; ventral surface
prominently granular, especially throat region; anterior parts of thigh, shank and groin, light greyish-brown with
creamy white mottling.
Secondary sexual characters. Male (SDBDU 2005.4732A): vocal sac externally visible on the lower jaw;
female (Holotype): ova white, pigmented on pole (diameter 0.8–1.2 mm, N = 10).
Geographical distribution and habitat. Uperodon taprobanicus is widely distributed in India and Sri Lanka,
and also reported from Bangladesh and Nepal (Fig. 2B). In India, it has been reported from the states of Tamil
Nadu, Kerala, Karnataka, Maharashtra, Gujarat, Andhra Pradesh, Telangana, Odisha, Madhya Pradesh,
Chhattisgarh, West Bengal, Bihar and Assam. In Sri Lanka, it is known from Central, Eastern, North Western,
Western, Sabaragamuwa, Southern and Uva Provinces. For detailed list of localities and references thereto see
Supplementary Table S1. In the present study, we collected and genetically confirmed the presence of this species
in Tamil Nadu (Mamallapuram), Kerala (Mannuthy, Kalpetta and Sulthanbathery), Karnataka (Bangalore) and
Odisha in India, and Central (Dambulla) and North Western (Karawaddana) regions of Sri Lanka. It was observed
in roadside puddles, wall crevices, cultivated fields and tree holes in both primary and secondary forests, from sea
level up to 950 m asl. See ‘Natural history and breeding’ observations in the respective section, after the species
accounts.
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FIGURE 6. Uperodon taprobanicus, U. anamalaiensis and U. montanus in life. A–C. Uperodon taprobanicus (SDBDU
2014.2567, male): A. dorsolateral view; B. dorsal view; C. ventral view; D–F. Uperodon anamalaiensis: D. dorsolateral view
(SDBDU 2013.2302, female); E. dorsal view (SDBDU 2017.3507, female); F. ventral view (SDBDU 2017.3507, female); G–
L. Uperodon montanus: G. dorsolateral view (SDBDU 2011.849, male); H. dorsolateral view (SDBDU 2011.1197, male); I.
ventral view (SDBDU 2002.1925, male); J. dorsolateral view (SDBDU 2002.1925, male); K. dorsal view (SDBDU 2015.3001,
male); L. dorsal view (SDBDU 2011.905, male). (Images: SD Biju and Sonali Garg).
Uperodon anamalaiensis (Rao, 1937)
Anamalai Globular Frog
(Figs. 1, 2A, 2C, 3D, 5D, 6D–F, 7A–E, 8A–D; Tables 1–4)
Original name and description. Ramanella anamalaiensis Rao, 1937. On some new forms of Batrachia from S.
India. Proceedings of the Indian Academy of Sciences. Section B, 6: 387–427. Neotype. By present designation,
ZSI/WGRC/V/A/956, an adult male, SVL 30.9 mm, from Parambikulam, Kerala state, India, collected by SD Biju
and team on 15 June 2011. Ty pe l ocality. “Base of Anamalai Hills, Coimbatore District”, Tamil Na du, India.
Current status of specific name. Valid name, as Uperodon anamalaiensis (Rao, 1937). Synonym. Ramanella
minor Rao, 1937 syn. nov. [= Uperodon minor (Rao, 1937)], Neotype by present designation, ZSI/WGRC/V/A/
957, an adult male, from Sakleshpur, Karnataka state, India, collected by SD Biju and team on 29 September 2012.
Neotypification of Ramanella anamalaiensis Rao, 1937. This species was described from the “Base of
Anamalai Hills, Coimbatore District” based on a single specimen (“total length 28.00 mm”), which was deposited
in the Central College, Bangalore (CCB). The original name bearing type for this taxon is considered lost (Dubois
1984; Biju 2001) and the species was known only from its original description, hence also presumed to be “lost”,
until it was rediscovered recently from the wet evergreen forests of Parambikulam Tiger Reserve (Vijayakumar et
al. 2011). In the original description, Rao (1937) stated several morphological characters based on which he
recognized this taxon to be new, but did not provide any comparison with the other closely known members. Our
collections from Parambikulam, which is part of the Anamalai Hills (type locality), are comparable with the
original description of Uperodon anamalaiensis (Rao 1937) in their general size, morphology, and dorsal and
ventral markings. The primary inconsistency between our recent collections and the original description is in the
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degree of webbing on foot. Rao (1937) mentions “toes perfectly free” but the accompanying illustration (Rao 1937,
plate XXIX, fig. 20) suggests presence of at least small foot webbing. However, since we found variation in foot
webbing among the various examined populations of this species, we do not consider this as a reliable character for
diagnosing the species. Further, since the taxonomic status of Ramanella anamalaiensis Rao, 1937 has remained
doubtful (Dutta 1997) primarily due to unavailability of its name bearing type, in accordance with Article 75 of
The Code, designation of a neotype is considered necessary for this taxon, in order to clarify its taxonomic status
and to define it objectively. Hence, based on diagnostic characters found useful for differentiating Ramanella
anamalaiensis Rao, 1937 (= Uperodon anamalaiensis) from its congeners (as stated in the ‘comparison’ section
below), we herein formally designate ZSI/WGRC/V/A/956, an adult male (SVL 30.9 mm) from Parambikulam, as
the neotype of Ramanella anamalaiensis Rao, 1937. The neotype description provided subsequently also shows
that the neotype is largely consistent with what is known of the former name-bearing type.
Synonymization and neotypification of Ramanella minor Rao, 1937. Rao (1937) described another taxon,
Ramanella minor, from Sakleshpur in Karnataka state along with Ramanella (= Uperodon) anamalaiensis. During
our surveys in the Sakleshpur region (type locality) we collected an Uperodon specimen that was comparable with
Rao’s description of Ramanella minor. Since the type specimen of this taxon that was originally deposited in the
Central College Bangalore (CCB), is also considered lost (Dubois 1984; Biju 2001), a critical study of the original
description of Ramanella minor Rao suggests that the majority of the mentioned characters are nearly the same as
for Ramanella anamalaiensis Rao and not helpful in distinguishing them from each other. The major diagnostic
characters stated for Ramanella minor by Rao (1937), such as ‘snout broadly truncated, distance between nostrils
equals width of the upper eyelid, interorbital space nearly twice the width of the upper eyelid, no occipital fold,’
and the overall colouration, also match with our collections of Uperodon anamalaiensis. The major differences
between these two taxa as per Rao’s descriptions seem to be ‘postnarial ridges strongly developed’ (vs. ‘ridges
incompletely developed’ in U. anamalaiensis), ‘diameter of the eye less than the length of the snout’ (vs. ‘diameter
of the eye greater than the length of the snout’ in U. anamalaiensis), ‘canthus rostralis obtuse’ (vs. ‘rostralis
rounded’ in U. anamalaiensis) and few other minor characters, which are all found to be variable among the
various Uperodon anamalaiensis populations examined by us (Table 4) and therefore may not be sufficient to
distinguish U. minor and U. anamalaiensis from each other. We also compared the 16S rRNA gene sequences of
our collection from Sakleshpur and the Uperodon anamalaiensis population from Parambikulam, and found them
to show a low genetic difference of 0.8%. Hence, in accordance with Articles 24.2.1 (Statement of the Principle of
the First Reviser) and 24.2.2 (Determination of precedence of names or acts by the First Reviser) of The Code, and
Recommendation 24A of The Code, we act as the First Revisers and consider Ramanella minor Rao, 1937 as a
junior subjective synonym of Ramanella anamalaiensis Rao, 1937 (= Uperodon anamalaiensis). Furthermore,
since the taxonomic status of Ramanella minor Rao, 1937 syn. nov. has been considered doubtful (Dutta 1997)
primarily due to unavailability of name bearing types, in accordance with Article 75 of The Code, we herein
designate ZSI/WGRC/V/A/957 (an adult male, SVL 31.7 mm, from the original type locality ‘Sakleshpur’) as the
neotype of Ramanella minor Rao, 1937, in order to define this taxon objectively and to establish taxonomic
stability. The neotype description of Ramanella minor Rao, 1937 syn. nov. provided along with figures (Figs.
S1A–E) in the supplementary file S1, also shows that the neotype is largely consistent with what is known of the
former name-bearing type.
Comparison. Uperodon anamalaiensis differs from U. globulosus, U. systoma and U. taprobanicus by its
smaller snout-vent size, male SVL 25–37 mm, N = 14, female SVL 35–39 mm, N = 4 (vs. larger, U. globulosus:
male SVL 51–56 mm, N = 2, female SVL 66–72 mm, N = 3; U. systoma: male SVL 51–56 mm, N = 2, female SVL
47–64 mm, N = 4; U. taprobanicus: male SVL 42–52 mm, N = 3, female SVL 53–55 mm, N = 2). It also differs
from the former two by its finger tips with truncate discs (vs. finger tips rounded without discs), ventral surface
with large spots or blotches (vs. without any markings), and presence of two indistinct neopalatinal ridges on
posterior side of each choana, fused with the vomerine odontophores (vs. two prominent dermal projections on the
neopalatinal ridges on posterior side of each choana, separated from the vomerine odontophores); differs from U.
taprobanicus by presence of two indistinct neopalatinal ridges on posterior side of each choana, fused with the
vomerine odontophores (vs. well-developed neopalatinal ridges). Further, U. anamalaiensis differs from U.
nagaoi, U. rohani sp. nov. and U. variegatus by its ventral surface in having large spots or blotches (vs. absent) and
presence of webbing between toes, I1
+
–2
–
II1–2
–
III1
+
–2
–
IV2–1V (vs. absent). This species could be confused with
U. montanus, U. mormorata, U. obscurus, U. palmatus and U. triangularis due to presence of large spots or
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TAXONOMIC REVIEW OF GENUS UPERODON
blotches on ventral skin. However, U. anamalaiensis differs from all these species by presence of continuous broad
median band extending from behind the eye up to near the vent (vs. absent or discontinuous patches). Specifically,
U. anamalaiensis differs from three Indian congeners by the following characters: from U. montanus by its
relatively larger snout-vent size, male SVL 25–37 mm, N = 14, female SVL 35–39 mm, N = 4 (vs. relatively
smaller, male SVL 21–28 mm, N = 12, female SVL 28–34 mm, N = 8); shank shorter than thigh, male: SHL/TL
ratio 0.8–0.9, N = 14, female: SHL/TL ratio 0.9, N = 4 (vs. shank equal to thigh, male: SHL/TL ratio 1.0, N = 12,
female: SHL/TL ratio 1.0, N = 8); and thigh almost equal to foot length, male: TL/FOL ratio 1.0–1.1, N = 14,
female: TL/FOL ratio 1.0, N = 4 (vs. thigh shorter than foot, male: TL/FOL ratio 0.8–0.9, N = 14, female: TL/FOL
ratio 0.8–1.0, N = 8) (Table 4). It differs from U. mormorata by its relatively smaller snout-vent size, male SVL
25–37 mm, N = 14, female SVL 35–39 mm, N = 4 (vs. relatively larger, male SVL 31–40 mm, N = 9, female SVL
32–43 mm, N = 3) and ventral skin surface with scattered spots or blotches (vs. densely marbled); and from U.
triangularis by presence of webbing between toes (Figs. 7C–E) (vs. absent) and presence of dermal fringes on
fingers and toes (vs. absent). Further, U. anamalaiensis differs from the Sri Lankan congeners by the following
characters: from U. palmatus by its snout longer than eye diameter, male: EL/SL ratio 0.5–0.7, N = 14, female: EL/
SL ratio 0.6–0.8, N = 4 (vs. equal, male: EL/SL ratio 1.0, N = 4, female: EL/SL ratio 1.0, N =2); and from U.
obscurus by its shank shorter than thigh, male: SHL/TL ratio 0.8–0.9, N = 14, female: SHL/TL ratio 0.9, N = 4 (vs.
nearly equal, male: SHL/TL ratio 0.9–1.0, N = 17, female: SHL/TL ratio 0.9–1.1, N = 5) (Table 4).
FIGURE 7. Head, hand, foot and foot webbing in Uperodon anamalaiensis, U. montanus, U. mormorata and U. nagaoi. A–E.
Uperodon anamalaiensis: A–D. Neotype of Uperodon anamalaiensis (ZSI/WGRC/V/A/956, male): A. lateral view of head; B.
ventral view of hand; C. ventral view of foot; D. schematic illustration of foot webbing; E. schematic illustration of foot
webbing in referred specimen of U. anamalaiensis (SDBDU 2013.928, female); F–L. Uperodon montanus: F–H. Topotype of
Uperodon montanus (SDBDU 2011.849, male): F. lateral view of head; G. ventral view of hand; H. ventral view of foot; I.
schematic illustration of foot webbing; J–L. schematic illustration of foot webbing in referred specimens of Uperodon
montanus: J. SDBDU 2015.3001, male; K. SDBDU 2003.40115, female; L. SDBDU 2005.30, female; M–S. Uperodon
mormorata: M–P. Neotype of Uperodon mormorata (ZSI/WGRC/V/A/958, female): M. lateral view of head; N. ventral view
of hand; O. ventral view of foot; P. schematic illustration of foot webbing; Q–S. schematic illustration of foot webbing in
referred specimens of Uperodon mormorata: Q. SDBDU 2002.567, female; R. SDBDU 2012.43, male; S. SDBDU 2015.3082,
male. T–W. Holotype of Uperodon nagaoi (WHT 2826, male): T. lateral view of head; U. ventral view of hand; V. ventral view
of foot; W. schematic illustration of foot webbing. (Illustrations: SD Biju).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon
anamalaiensis showed an average intraspecific distance of 0.4% (range 0–0.8%, N = 6). Genetically, U.
anamalaiensis is closely related to U. montanus, from which it was found to differ by an average uncorrected
genetic distance of 3.2% (range 2.6–4.0%, N = 60). For interspecific genetic distances with all other members of
the genus, see Table 3.
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Description of neotype (measurements in mm) (Figs. 3D, 7A–D). Small-sized (SVL 30.9), slender adult male;
head small (HW 8.5, HL 6.3, IFE 3.9, IBE 7.0), about one-fifth (20.3%) of body length, wider than long (HW/HL
ratio 1.3); snout truncate in dorsal and ventral view, vertical in lateral view, protruding, its length (SL 3.0) longer
than horizontal diameter of eye (EL 2.2); loreal region acute with rounded canthus rostralis; interorbital space
about twice as wide (IUE 3.6) as upper eyelid width (UEW 1.7); nostril oval without lateral flap of skin, closer to
tip of snout (NS 0.7) than to eye (EN 1.7); tympanum not visible externally, distinct supratympanic fold extending
from posterior corner of upper eyelid near to insertion of forelimb at axilla; eye diameter (EL 2.2); vomerine
odontophores present on the palate, prominent, with or without teeth; presence of two indistinct neopalatinal ridges
on posterior side of each choana, fused with the vomerine odontophores; tongue moderately large, emarginate.
Forelimbs moderately long and thin; forearm (FAL 6.9) shorter than hand length (HAL 8.5); finger length formula
I<II<IV<III; tips of all fingers with truncate discs, without grooves, moderately wide compared to finger width
(FD
I
1.5, FW
I
0.3; FD
II
1.5, FW
II
0.4; FD
III
1.8, FW
III
0.5; FD
IV
1.5, FW
IV
0.4); subarticular tubercles prominent,
oval, all present; two well-developed palmar tubercles, inner oval, outer bilobed. Hind limbs relatively long and
thin, thigh length (TL 13.9) longer than shank (SHL 12.4) and foot (FOL 13.5); relative digit lengths
I<II<V<III<IV; tips of all toes with small truncate to rounded discs, rather wide compared to toe width (TD
I
0.7,
TW
I
0.4; TD
II
0.9, TW
II
0.5; TD
III
1.0, TW
III
0.5; TD
IV
1.1, TW
IV
0.5; TD
V
0.8, TW
V
0.4); foot webbing present,
moderate: I1
+
–2
–
II1–2
–
III1
+
–2
–
IV2–1V; well-developed dermal fringes present on all toes; subarticular tubercles
prominent, oval, all present; two smooth metatarsal tubercles, oval, outer tubercle slightly larger than the inner.
Skin of snout, between eyes, and sides of head, shagreened to granular; anterior and posterior parts of back,
and upper and lower parts of flank, granular (Fig. 3D); dorsal surfaces of forelimb, thigh, tibia and tarsus,
shagreened to sparsely granular; throat, chest, abdomen, and ventral surfaces of limbs, shagreened (Fig. 3D).
Colouration. In preservation: Dorsum dark greyish-brown with faint dark grey band extending from posterior
end of upper eyelid on either side, along the side of dorsum, and approaching the vent; a faint dark grey stripe
between eyes; throat, chest and belly, brown with scattered light grey blotches; ventral surfaces of forelimb, thigh,
tibia and foot, dark grey with light grey blotches (Fig. 3D). Colour in life: Dorsum brown with irregular yellowish-
brown spots and prominent yellowish-brown band extending from posterior end of upper eyelid near to vent;
yellowish-brown stripe between eyes; lower part of flank brown with yellow speckles; dorsal surfaces of forelimbs,
thigh, shank and foot, brown with yellowish-brown cross bands; throat, chest and belly, greyish-brown with
scattered light grey blotches; ventral surfaces of forelimb, thigh, tibia and foot, dark grey with light yellow
blotches.
Variations. Morphometric measurements for 18 specimens, including the neotype, are given in Table 4. There
is not much variation observed in the overall morphology, except for skin colour and markings, sex-based webbing
differences (Figs. 7D–E), and presence of vomerine teeth in SDBDU 2013.2302 (Fig. 5D). Dorsal colour and skin
granulation varies slightly between individuals even from the same populations. SDBDU 2011.233A, SDBDU
2011.234, SDBDU 2011.548, SDBDU 2012.2000 and SDBDU 2012.1998: dorsum dark grey with a light grey
dorsal band extending over the back and a light grey stripe between the eyes, dorsal skin shagreened to sparsely
granular, ventral surface (including limbs) uniformly coffee brown with yellowish-grey spots and blotches;
SDBDU 2013.2302: anterior parts of thigh sparsely granular with scattered spinular projections; SDBDU
2013.928: dorsum light greyish-brown with a straw coloured band extending over the back, foot webbing I2–2II2
–
–3
–
III2–3IV3
–
–2
–
V (Fig. 7E); SDBDU 2012.1995: anterior parts of thigh prominently granular with scattered
spinular projections.
Secondary sexual characters. Male: vocal sac externally visible on the lower jaw; female (SDBDU
2017.3507): ova white, pigmented on pole (diameter 0.6–0.9 mm, N = 10).
Tad pol e morphology. In dorsal view, tadpoles of Uperodon anamalaiensis (Gosner stage 34, N = 2) have an
elliptical body with narrower and shorter anterior region, and wider and longer posterior region; snout appears
rounded in dorsal view and depressed in lateral view; eyes small, bulbous, positioned laterally; nasolacrimal duct
absent; narial depressions visible clearly in lateral view, located much closer to snout than to eyes, pigmentation
not visible around nasal depressions; spiracle opens near the vent, flap present; vent tubular, positioned mid-
ventrally with aperture opening medially in line with the plane of ventral fin; tail ends with round tip, composed of
bilateral myotomic muscle masses divided by V-shaped septa, with unequal membranes on either side of tail
musculature; dorsal fins originate after tail-body junction, margin of lower fin not parallel to margin of tail muscle;
mouth terminal and appears as a slit opening, keratinized structures absent, papillae absent.
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TAXONOMIC REVIEW OF GENUS UPERODON
FIGURE 8. Male advertisement calls of Uperodon anamalaiensis, U. mormorata and U. triangularis. A–D. Uperodon
anamalaiensis: A. ten second call segment; B. one second call segment; C. 0.1 second segment showing a single non-pulsatile
call; D. spectrogram of the call shown in Figure 8B; E–H. Uperodon mormorata: E. ten second call segment; F. one second call
segment; G. 0.1 second segment showing a single non-pulsatile call; H. spectrogram of the call shown in Figure 8G; I–L.
Uperodon triangularis: I. ten second call segment; J. one second call segment; K. 0.1 second segment showing a single non-
pulsatile call; L. spectrogram of the call shown in Figure 8J.
Measurements (mm) (N = 1): Lateral measurements include, maximum height of body (bh) = 3.41, maximum
width of body (bw) = 6.87, maximum diameter of eye (ed) = 0.58, internarial distance (nn) = 1.65, naro-pupular
distance (np) = 2.86, rostro-narial distance (rn) = 0.02, interpupular distance (pp) = 5.42, distance from tip of snout
to opening of spiracle (ss) = 6.44, distance from tip of snout to insertion of upper tail fin (su) = 9.72, snout-vent
length (svl) = 8.86, total length (tl) = 21.78, distance from vent to tip of tail (vt) = 13.51, maximum height of tail
(ht) = 3.83, tail muscle height (tmh) = 1.97, tail muscle width (tmw) = 1.97.
Vocalization. A male of Uperodon anamalaiensis (SDBDU 2012.1994) from Methooty produced a single type
of call with pulsatile temporal structure. Calls were not delivered in groups. A typical male call had a duration of
208.9 ms, a short rise time of 4 ms, and fall time of 202.3 ms. The call comprised of 35 pulses that were delivered
at a rate of 175.8 pulses/s. Spectrum was characterized by an overall dominant frequency of 1.2 kHz (Figs. 8A–D).
Geographical distribution and habitat. Uperodon anamalaiensis was only known from its original
description until its recent rediscovery from Parambikulam in the southern Western Ghats of India. The present
study provides further morphological and genetically confirmed reports of this species from Kerala (Chathankod
and Methooty) and Karnataka (Sakleshpur) (Fig. 2C; Table 1), at elevations between 100–650 m asl. It is also
likely to be present in the Anamalai region of Tamil Nadu. Individuals were collected from tree holes, ground
puddles and shallow water channels inside disturbed secondary forests (Chathankod and Methooty) as well as
primary forest areas, either protected (Parambikulam) or unprotected (Ponmudi and Sakleshpur).
Notes on previous reports. This species was previously misidentified as Uperodon triangularis from
Ponmudi hills (Inger et al. 1984) and Thiruvananthapuram district (Andrew et al. 2005) of Kerala state. The report
of U. anamalaiensis from Munnar in Kerala (Harpalani et al. 2015) most likely refers to U. montanus (see
Supplementary Table S1 for list of localities and references thereto).
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Uperodon montanus (Jerdon, 1854 “1853”)
Mountain Globular Frog
(Figs. 1, 2A, 2C, 3E, 5E, 6G–L, 7F–L; Tables 1–4)
Original name and description. Hylaedactylus montanus Jerdon, 1854 “1853”. Catalogue of reptiles inhabiting
the Peninsula of India. Journal of the Asiatic Society of Bengal, 22:522–534. Holotype. By monotypy, NHM
72.4.17.212 (ex. BMNH 1947.2.11.19), an adult female, SVL 28.3 mm. Type locality. “Mountain streams in
Wynaad”, Kerala, India. Current status of specific name. Valid name, as Uperodon montanus (Jerdon, 1854
“1853”).
Comments. The original description by Jerdon (1854 “1853”) was brief and subsequently elaborated by
Parker (1934). This species is largely known only from its original collection and some scanty reports or checklists
(e.g., Dutta 1997; Subramanian et al. 2013). Further, considering the recent taxonomic placement of this taxon in
genus Uperodon and morphological variations observed among populations (particularly in foot webbing), here we
provide a detailed redescription of the holotype and comparison of this species with all its presently recognized
congeners, to facilitate proper identification.
Comparison. For comparison of Uperodon montanus with U. globulosus, U. systoma, U. taprobanicus and U.
anamalaiensis see ‘comparison’ section of those species. Uperodon montanus differs from U. nagaoi, U. rohani
sp. nov. and U. variegatus by presence of prominent spots and blotches on its ventral surface (vs. inconspicuous
spots and speckles present or absent in U. nagaoi, and completely absent in U. rohani sp. nov. and U. variegatus)
and relatively smaller snout-vent size, male SVL 21–28 mm, N = 12; female SVL 28–34 mm, N = 8 (vs. relatively
larger, U. nagaoi: male SVL 25–32 mm, N = 8; U. rohani sp. nov.: male SVL 26–34 mm, N = 16, female SVL 28–
35 mm, N = 8; U. variegatus: male SVL 25–33 mm, N = 12, female SVL 29–36 mm, N = 7). Uperodon montanus
could be confused with U. anamalaiensis, U. mormorata, U. obscurus, U. palmatus and U. triangularis due to
presence of prominent spots and blotches on its ventral skin surface. However, U. montanus differs from all these
species by its relatively smaller snout-vent size, male SVL 21–28 mm, N = 12; female SVL 28–34 mm, N = 8 (vs.
relatively larger, U. anamalaiensis: male SVL 25–37 mm, N = 14, female SVL 35–39 mm, N = 4; U. mormorata:
male SVL 31–40 mm, N = 9, female SVL 32–43 mm, N = 3; U. obscurus: male SVL 22–31 mm, N = 17, female
SVL 30–38 mm, N = 5; U. palmatus: male SVL 27–31 mm, N = 4, female SVL 34–36 mm, N = 2; U. triangularis:
male SVL 25–37 mm, N = 20, female SVL 33–42 mm, N = 4). Specifically, U. montanus also differs from U.
mormorata by its ventral skin being light to dark brown with scattered grey spots or blotches (vs. dark brown with
dense marbling); from U. palmatus by its snout longer than eye diameter, male: EL/SL ratio 0.6–0.8, N = 12,
female: EL/SL ratio 0.6–0.8, N = 8 (vs. equal, male: EL/SL ratio 1.0, N = 4, female: EL/SL ratio 1.0, N =2); and
from U. triangularis by presence of webbing between toes (vs. absent).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon montanus
showed an average intraspecific distance of 0.7% (range 0–1.2%, N = 10). Genetically, U. montanus is closely
related to U. anamalaiensis, from which it was found to differ by an average uncorrected genetic distance of 3.2%
(range 2.6–4.0%, N = 60). For interspecific genetic distances with all other members of the genus, see Table 3.
Redescription of holotype (measurements in mm) (Fig. 3E). Small-sized (SVL 28.3), slender adult female;
head small (HW 10.1, HL 7.0, IFE 3.9, IBE 7.5), one-fourth (24.7%) of body length, wider than long (HW/HL ratio
1.4); snout nearly rounded in dorsal and ventral view, nearly acute in lateral view, its length (SL 3.3) longer than
horizontal diameter of eye (EL 2.1); loreal region obtuse with rounded canthus rostralis; interorbital space nearly
twice as wide (IUE 3.1) as upper eyelid width (UEW 1.6); nostril closer to tip of snout (NS 0.8) than to eye (EN
2.1); supratympanic fold distinct, extending from posterior corner of upper eyelid to insertion of forelimb at axilla;
eye diameter (EL 2.1); vomerine odontophores present on the palate; indistinct neopalatinal ridges on posterior side
of each choana, appear fused with the vomerine odontophores; tongue moderately large, circular, shallowly
emarginated. Forelimbs moderately long and thin; forearm (FAL 6.8) shorter than hand length (HAL 8.7); finger
length formula I<II<IV<III. Hind limbs relatively long and thin, thigh length (TL 12.3) shorter than shank (SHL
12.4) and foot (FOL 13.9).
Skin of snout, between eyes, sides of head, and anterior part of dorsum, shagreened to granular; posterior part
of back, and upper and lower parts of flank, granular (Figs. 3E); dorsal surfaces of forelimb, thigh, tibia and tarsus,
shagreened to sparsely granular; ventral surface uniformly shagreened (Fig. 3E).
Since the hand and foot morphology could not be reliably determined due to poor preservation of the holotype
specimen, we are describing these characters based on a topotype specimen (SDBDU 2011.849).
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TAXONOMIC REVIEW OF GENUS UPERODON
Topotype specimen (SDBDU 2011.849). Hand (Fig. 7G): tips of all fingers with truncate discs, moderately
wide compared to finger width (FD
I
0.6, FW
I
0.3; FD
II
1.0, FW
II
0.7; FD
III
1.0, FW
III
0.7; FD
IV
0.9, FW
IV
0.4); well-
developed dermal fringes present on all fingers; subarticular tubercles rather prominent, oval, all present; two well-
developed palmar tubercles (inner, oval, 0.8 mm; outer, bilobed, 1.1 mm). Foot (Figs. 7H–I): tips of all toes with
small rounded discs, rather wide compared to toe width (TD
I
0.5, TW
I
0.4; TD
II
0.6, TW
II
0.4; TD
III
0.6, TW
III
0.5;
TD
IV
0.6, TW
IV
0.4; TD
V
0.5, TW
V
0.4); well-developed dermal fringes present on all toes; subarticular tubercles
rather prominent, oval; two metatarsal tubercles, oval, outer (0.5 mm long) slightly larger than the inner (0.9 mm
long); webbing present, basal: I2–2II2–3III3–4IV4–3V.
Colouration. In preservation (SDBDU 2011.849): Dorsum greyish-brown; flanks, forelimbs, dorsal surfaces
of thigh, shank and foot, light greyish-brown; chest and belly dark grey with light grey spots and patches, ventral
surfaces of thigh, tibia and foot, light grey with lighter grey spots. Colour in life (SDBDU 2011.849): Dorsum and
lateral sides of head uniformly dark brown with pale brown median patch on the back and few irregular patches
near groin and thigh; dorsal surface of limbs dark brown; ventral surface dark brown with irregular white spots and
blotches (Fig. 6G).
Variations. Morphometric measurements of 20 specimens, including the holotype, are given in Table 4. Dorsal
colouration and webbing is highly variable in this species. Foot webbing: basal in SDBDU 2011.849 (male), I2–
2II2–3III3–4IV4–3V; medium in SDBDU 2015.3001 (male), I2
–
–2II2
–
–3III2–3IV3–2
1
/
2
V; medium in SDBDU
2003.40115 (female), I2–2II2
–
–3
–
III2–3
+
IV3
–
–2V; and medium in SDBDU 2005.30 (female), I2–2II2
–
–3III2–
3IV3–2V (Figs. 7I–L). SDBDU 2011.849 (male): Dorsum uniformly brown with faint grey markings; SDBDU
2015.3001 (male): colour in life: dorsum orangish-brown with dark brown median patch on the back and few
irregular patches near groin and thigh, forelimbs orangish-brown, and hind limbs orangish-yellow with dark brown
cross bands (Fig. 6K); colour in preservation: dorsum grey with brown median patch and irregular brown spots,
brown stripe between eyes, posterior part of dorsum (near groin) with two brown blotches on either side, dorsal
surface of hand and foot grey with dark brown cross bands; SDBDU 2015.3002 (female): dorsum prominently
granular, dorsal markings almost similar to SDBDU 2015.3001.
Note. Parker (1934) illustrated the foot webbing for this species (figure 38, p. 91) and showed relatively more
webbing in the male compared to the female. He also stated toes to be “half webbed in males” and “1/3 webbed in
females”. In our study (N = 20), we found webbing to be highly variable, and since females were also often found
to have more extensive webbing than males, this character should not be considered reliable for diagnosing this
species.
Secondary sexual characters. Male: Vocal sac externally visible on the lower jaw; female (SDBDU
2002.24B): ova white, pigmented on pole (diameter 0.9–1.5mm, N = 20).
Geographical distribution and habitat. Uperodon montanus was originally described from the Wa ya nad
region. In the present study, it is confirmed to occur in the Western Ghats states of Kerala (Devikulam, Munnar,
Upper Manalar, Pandimotta, Nelliyampathy, Siruvani, Kurichiyarmala, Settukunnu and Thirunelli) and Tamil Nadu
(Kakkachi and Sengaltheri), predominantly from mid to high elevations ranging from 800–1700 m asl (Fig. 2C;
Table 1). Individuals were collected either from water-filled tree holes, damp leaf litter on forest floor or ground
puddles. This species is likely to be distributed in southern parts of Karnataka. However, reports of U. montanus
from Gujarat, Maharashtra and northern Karnataka require confirmation and may refer to U. mormorata (see
Supplementary Table S1 and references thereto).
Uperodon mormorata (Rao, 1937)
Mottled Globular Frog
(Figs. 1, 2A, 2C, 3F, 5F, 7M–S, 8E–H, 9A–D; Tables 1–4)
Original name and description. Ramanella mormorata Rao, 1937. On some new forms of Batrachia from S.
India. Proceedings of the Indian Academy of Sciences. Section B, 6: 387–427. Neotype. By present designation,
ZSI/WGRC/V/A/958, an adult female, SVL 32.4 mm, from Sakleshpur, Karnataka state, India, collected by SD
Biju on 26 June 2003. Type lo cal ity. “Saklespur (= Sakleshpur), Hassan District, Mysore”, India. Current status
of specific name. Valid name, as Uperodon mormorata (Rao, 1937).
Neotypification of Ramanella mormorata Rao, 1937. This species was described from “Saklespur” based on
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a single specimen “total length 25.00 mm”, which was deposited in the Central College, Bangalore. The original
name bearing type (or types) of this species are considered lost (Dubois 1984; Das & Whitaker 1997; Biju 2001),
and this nominal taxon was also presumed to be “lost” until its rediscovery from “Canacona (15°01’N 74°04’E),
adjacent to Cotigao Wildlife Sanctuary (Goa, south-Western India)” based on an available museum specimen from
“Malabar” at MCZ, USA (Das & Whitaker 1997). As per the original description, Rao had several available
specimens including “young”, “immature”, “mature males” and “mature females”, however, the description was
based on a single specimen for which absolute measurements were included. Das & Whitaker (1997) provided a
detailed description of this species based on examined specimens, however, without designating a neotype. Our
new collection from the type locality Sakleshpur is comparable with the original description of Ramanella
mormorata Rao, 1937, in overall general morphology, body size and dorsal and ventral markings. Since the
original name-bearing type of this species is lost, we consider designation of a neotype necessary to define this
nominal taxon objectively for clarification of its taxonomic status and to establish nomenclatural stability. Hence,
in accordance with Article 75 of The Code, and based on diagnostic characters found useful for differentiating
Ramanella mormorata Rao, 1937 (= Uperodon mormorata) from its congeners (as stated in the ‘comparison’
section below), we herein formally designate ZSI/WGRC/V/A/958 (an adult female, SVL 32.4 mm) from the
original type locality Sakleshpur as the neotype of Ramanella mormorata Rao, 1937. The neotype description
provided subsequently also shows that the neotype is largely consistent with what is known of the former name-
bearing type.
Comparison. For comparison of Uperodon mormorata with U. globulosus, U. systoma, U. taprobanicus, U.
anamalaiensis and U. montanus see ‘comparison’ section of those species. Uperodon mormorata differs from U.
nagaoi, U. rohani sp. nov. and U. variegatus by its densely mottled ventral surface (vs. inconspicuous spots and
speckles present or absent in U. nagaoi; and complete absence of spots, blotches or mottling in U. rohani sp. nov.
and U. variegatus) and relatively larger snout-vent size, male SVL 31–40 mm, N = 9, female SVL 32–43 mm, N =
3 (vs. relatively smaller, U. nagaoi: male SVL 25–32 mm, N = 8; U. rohani sp. nov.: male SVL 26–34 mm, N = 16,
female SVL 28–35 mm, N = 8; U. variegatus: male SVL 25–33 mm, N = 12, female SVL 29–36 mm, N = 7). This
species could be confused with U. anamalaiensis, U. obscurus, U. palmatus and U. triangularis due to its mottled
ventral skin. However, U. mormorata differs from all these species by its relatively larger snout-vent size, male
SVL 31–40 mm, N = 9, female SVL 32–43 mm, N = 3 (vs. smaller, U. obscurus: male SVL 22–31 mm, N = 17,
female SVL 30–38 mm, N = 5; U. palmatus: male SVL 27–31 mm, N = 4, female SVL 34–36 mm, N = 2; U.
triangularis: male SVL 25–37 mm, N = 20, female SVL 33–42 mm, N = 4), its ventral surface (including throat
and limbs) being densely marbled (vs. scattered spots or blotches, largely on chest and belly), and shank relatively
shorter than thigh, male: SHL/TL ratio 0.8–1.1, N = 9, female: SHL/TL ratio 0.8–0.9, N = 3 (vs. nearly equal, U.
obscurus: male: SHL/TL ratio 0.9–1.0, N = 17, female: SHL/TL ratio 0.9–1.1, N = 5; U. palmatus: male: SHL/TL
ratio 1.0, N = 4, female: SHL/TL ratio 1.0, N = 2; U. triangularis: male: SHL/TL ratio 0.9–1.1, N = 20, female:
SHL/TL ratio 0.9–1.0, N = 4).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon mormorata
showed an average intraspecific distance of 0.6% (range 0–1.2%, N = 11). Populations from Maharashtra and
Karnataka were observed to be divergent by up to 1.2%. Genetically, U. mormorata is closely related to U.
triangularis, from which it was found to differ by an average uncorrected genetic distance of 2.9% (range 2.4–
3.6%, N = 99). For interspecific genetic distances with all other members of the genus, see Table 3.
Description of neotype (measurements in mm) (Figs. 3F, 7M–P). Small-sized (SVL 32.4), relatively robust
adult female; head small (HW 9.6, HL 8.8, IFE 4.4, IBE 8.6), less than one-third (27.2%) of body length, wider
than long (HW/HL ratio 1.1); snout truncate in dorsal and ventral view, vertical in lateral view, its length (SL 3.1)
shorter than horizontal diameter of eye (EL 2.6); loreal region acute with rounded canthus rostralis; interorbital
space nearly twice as wide (IUE 3.0) as upper eyelid width (UEW 1.6); nostril closer to tip of snout (NS 0.6) than
to eye (EN 1.5); supratympanic fold distinct, extending from posterior corner of upper eyelid to insertion of
forelimb at axilla; eye diameter (EL 2.6); vomerine odontophores present on the palate, prominent, without teeth;
weakly-developed neopalatinal ridges on posterior side of each choana, appear fused with the vomerine
odontophores; tongue moderately large, emarginate. Forelimbs moderately long and thin; forearm (FAL 6.5)
shorter than hand length (HAL 9.1); finger length formula I<II<IV<III; tips of all fingers with truncate discs,
without grooves, moderately wide compared to finger width (FD
I
1.5, FW
I
0.3; FD
II
1.5, FW
II
0.4; FD
III
1.8, FW
III
0.5; FD
IV
1.5, FW
IV
0.4); subarticular tubercles prominent, oval, all present; two well-developed palmar tubercles
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TAXONOMIC REVIEW OF GENUS UPERODON
(inner, oval, 1.6 mm long; outer, bilobed, 1.9 mm long). Hind limbs relatively long and thin, thigh length (TL 13.5)
longer than shank (SHL 11.1) and foot (FOL 12.9); tips of all toes with small truncate to rounded discs, discs rather
wide compared to toe width (TD
I
0.7, TW
I
0.4; TD
II
0.8, TW
II
0.6; TD
III
1.0, TW
III
0.7; TD
IV
1.0, TW
IV
0.6; TD
V
0.8,
TW
V
0.5); foot webbing present: I2
–
–2II2–3III2–3IV3–2V; well-developed dermal fringe present on all toes;
subarticular tubercles prominent, oval; two smooth metatarsal tubercles, oval, outer slightly longer than the inner.
Skin of snout, between eyes, sides of head, anterior part of the dorsum, posterior parts of back, and upper and
lower parts of flank, granular; dorsal surfaces of forelimb, thigh, tibia and tarsus, shagreened to sparsely granular
(Figs. 3F, 7M); ventral surfaces shagreened.
Colouration. In preservation: Dorsum olive grey with scattered dark brown spots, dark brown inverted ‘V’ at
the level of shoulder (directed downwards), a dark brown line between eyes, another two lines: one on snout, and
another between inverted ‘V’ and line connecting the eyes; dorsal surfaces of forelimb, thigh and shank (including
fingers and toes), olive grey with dark brown cross bands; groin and anterior parts of thigh, light grey with light
brown mottling, and a light brown patch near cloacal region; throat, chest, belly, and ventral surfaces of forelimbs
and hind limbs, light brown with prominent light grey reticulations (Fig. 3F). Colour in life (SDBDU 2016.3404):
Dorsum dark olive grey with scattered dark brown spots especially on posterior part of dorsum, thigh and shank,
faint dark brown inverted ‘V’ at the level of shoulder (directed downwards); dorsal surfaces of forelimb, thigh and
shank (including fingers and toes), olive grey with dark brown cross bands; groin and anterior parts of thigh, light
brown with light grey and white mottling; throat, chest, belly, and ventral surfaces of forelimb and hind limb, dark
brown with prominent light greyish-blue reticulations (Figs. 9A–C).
FIGURE 9. Uperodon mormorata, U. nagaoi and U. obscurus in life. A–C. Uperodon mormorata (SDBDU 2016.3404, male):
A. Dorsolateral view; B. dorsal view; C. ventral view; D. Uperodon mormorata (SDBDU 2011.835, male), dorsolateral view;
E–G. Uperodon nagaoi (DZ 1367, male): E. Dorsolateral view; F. dorsal view; G. frontal view; H–K. Uperodon obscurus: H.
Dorsolateral view (DZ 1366, male); I. Dorsal view (DZ 1366, male); J. dorsolateral view (DZ 1408, male); K. dorsolateral
view (DZ 1369, male); L. ventral view (DZ 1369, male). (Images: SD Biju and E–G Madhava Meegaskumbura).
Variations. Morphometric measurements for 12 specimens, including the holotype, are given in Table 4.
Overall, the skin colouration and texture (except markings), are nearly similar in all the examined specimens.
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However, webbing between toes is observed to be highly variable, without being consistent even between the same
sexes. Foot webbing: SDBDU 2002.567 (female): medium, I2
–
–2II2–3III2
+
–3IV3–2V; SDBDU 2012.43 (male):
large, I1–2II1–1III1–2IV2–1V; SDBDU 2015.3082 (male): medium, I1–2II1–2III2–3IV3–2V (Figs. 7Q–S).
SDBDU 2015.3085: dorsum with more prominent blackish-brown spots.
Note. In the original description, Rao (1937) stated this species to have “No occipital fold”. However, Das &
Whitaker (1997) discussed its presence based on their observation of occipital folds in both their examined samples
of Uperodon mormorata (MCZ A. 116283 and MCZ A. 15421). Variable occurrence of occipital fold was earlier
also reported by Parker (1934). We closely examined the presence and absence of occipital fold in members of the
genus Uperodon, and observe that its presence may be an artifact of preservation caused due to stretching of loose
skin of the snout towards the head.
Secondary sexual characters. Male (SDBDU 2016.3404): Vocal sac externally visible on the lower jaw;
female (ZSI/WGRC/V/A/958) ova white, pigmented on pole (diameter 0.6–0.8 mm, N = 20).
Vocalization. Male of Uperodon mormorata (SDBDU 2016.3404) from Goa produced a single type of call
with pulsatile temporal structure. The calls had uniform intervals and were not delivered in groups. A typical male
call had a duration of 182.6 ms, rise time of 108.3 ms, and fall time of 57.4 ms. The call comprised of 28 pulses that
were delivered at a rate of 154.7 pulses/s. The spectrum was characterized by single broad peak with mean
dominant frequency of 2.6 kHz (Figs. 8E–H).
Geographical distribution and habitat. In the present study, the presence of Uperodon mormorata is
confirmed in the Western Ghats states of Karnataka, Maharashtra, Goa and Gujarat (Fig. 2C; Tables 1). For
additional previous records and references thereto see supplementary Table S1. Report of this species from Assam
(Dey 2001) is doubtful. We collected this species predominantly from terrestrial habitats such as rock crevices,
fallen logs and under stones, and occasionally from tree barks about two metres above the ground. It was found in
and around secondary forest areas (Amba, Amboli, Bygoor, Dandeli, Londa, Netravali, Sakleshpur, Samot and
Castle rock), primary forests (Shimoga), as well as marshes (coastal Goa), at elevations from sea level up to 1050
m asl.
Uperodon nagaoi (Manamendra-Arachchi and Pethiyagoda, 2001)
Nagao’s Globular Frog
(Figs. 1, 2A, 2D, 3G, 7T–W, 9E–G; Tables 1–4)
Original name and description. Ramanella nagaoi Manamendra-Arachchi and Pethiyagoda, 2001. Ramanella
nagaoi, a new tree-hole frog (Microhylidae) from southern Sri Lanka. Journal of South Asian Natural History, 5:
121–133. Holotype. WHT 2826, by original designation, an adult male, SVL 25.6 mm. Type locality. “Kanneliya
Forest Reserve (near Galle), Sri Lanka”. Current status of specific name. Valid name, as Uperodon nagaoi
(Manamendra-Arachchi and Pethiyagoda, 2001).
Comparison. For comparison of Uperodon nagaoi with U. globulosus, U. systoma, U. taprobanicus, U.
anamalaiensis, U. montanus and U. mormorata see ‘comparison’ section of those species. Uperodon nagaoi differs
from U. rohani sp. nov. and U. variegatus by its dorsum being light grey to dark brownish-grey without prominent
markings (vs. light to dark brown with light grey or olive markings) and nearly uniform brown ventral skin without
or with inconspicuous spots and speckles (vs. uniformly white, without any spots, blotches or mottling).
Specifically, it differs from U. variegatus by its shank relatively longer than thigh, SHL/TL ratio 1.0–1.2, N = 8 (vs.
shorter, male: SHL/TL ratio 0.9–1.0, N = 16, female: SHL/TL ratio 0.8–0.9, N = 7) (Table 4). Further, U. nagaoi
differs from U. obscurus and U. palmatus by its shank relatively longer than thigh, male: SHL/TL ratio 1.0–1.2, N
= 8 (vs. nearly equal, U. obscurus: male: SHL/TL ratio 0.9–1.0, N = 17, female: SHL/TL ratio 0.9–1.1, N = 5; U.
palmatus: male: SHL/TL ratio 1.0, N = 4, female: SHL/TL ratio 1.0, N = 2), rudimentary webbing between toes,
I1–2II2–3
+
III3–4IV4
+
–3V (vs. more, U. obscurus: I1–2II1
1
/
2
–3
–
III2–3
+
IV3
+
–2V; U. palmatus: I1–2II1–3
–
III1–
3IV2
2
/
3
–1V), and ventral skin without or with inconspicuous spots and speckles (vs. prominent spots or blotches).
It differs from U. triangularis by its dorsal skin in having faint brown to dark brown coloured markings (vs.
prominent bright orange coloured markings), ventral skin without or with inconspicuous spots and speckles (vs.
prominent spots or blotches), and tadpoles with funnel-shaped mouth and papillae (vs. terminal mouth lacking
papillae).
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TAXONOMIC REVIEW OF GENUS UPERODON
Genetic divergence. For 16S mitochondrial gene sequences, intraspecific variation was not observed between
the sampled populations of Uperodon nagaoi (N = 2). Genetically, U. nagaoi is closely related to U. obscurus and
U. palmatus; from these it was found to differ by an average uncorrected genetic distance of 5.6% (N = 6) and 4.0%
(range 4.0–4.2%, N = 8), respectively. For interspecific genetic distances with all other members of the genus, see
Table 3.
Description of holotype. A detailed general description was published by Manamendra-Arachchi &
Pethiyagoda (2001).
Variations. Morphometric measurements for eight specimens, including the type series, are given in Table 4.
Secondary sexual character. Male (DZ 1367): Vocal sac externally visible on the lower jaw.
Geographical distribution and habitat. This species is endemic to Sri Lanka, where it is known to occur in
the Central, Sabaragamuwa, Southern and Western Provinces (Fig. 2D). For list of localities and references thereto
see Supplementary Table S1. In the present study, we collected this species from tree holes at Hiyare Forest
Reserve (Galle district) from elevation of 125 m asl, in the vicinity of its type locality Kanneliya Forest Reserve.
See ‘Natural history and breeding’ observations in the respective section, after the species accounts.
Uperodon obscurus (Günther, 1864 “1863”)
Sri Lankan Globular Frog
(Figs. 1, 2A, 2D, 3H, 5G, 9H–K, 10A–D; Tables 1–4)
Original name and description. Callula obscura Günther, 1864 “1863”. The Reptiles of British India. London:
Ray Society by R. Hardwicke: 438. Holotype. By monotypy, NHM 58.11.28.42 (ex. BMNH 1947.2.11.20), an
adult female, SVL 32.2 mm. Type locality. “Ceylon” (Sri Lanka). Current status of specific name. Valid name,
as Uperodon obscurus (Günther, 1864 “1863”).
Comparison. For comparison of Uperodon obscurus with U. globulosus, U. systoma, U. taprobanicus, U.
anamalaiensis, U. montanus, U. mormorata and U. nagaoi see ‘comparison’ section of those species. Uperodon
obscurus is closely related to U. palmatus, both genetically and morphologically. However, it differs from the latter
by its snout length longer than horizontal diameter of eye, male: EL/SL ratio 0.6–0.9, N = 17; female: EL/SL ratio
0.6–0.9, N = 5 (vs. equal, male: EL/SL ratio 1.0, N = 4, female: EL/SL ratio 1.0, N = 2) (Table 4), snout nearly
acute in lateral view (vs. vertical), relatively less webbing, I1–2II1
1
/
2
–3
–
III2–3
+
IV3
+
–2V (vs. more, I1–2II1–3
–
III1–
3IV2
2
/
3
–1V) (Fig. 10D), and dorsal skin shagreened to sparsely granular (vs. distinctly granular with warty
projections). It differs from U. triangularis by its fourth toe webbing extending close up to second subarticular
tubercle: I1–2II1
1
/
2
–3
–
III2–3
+
IV3
+
–2V (vs. toes completely free of webbing). Uperodon obscurus cannot be
confused with three other species, U. nagaoi, U. rohani sp. nov. and U. variegatus, due to its relatively extensive
foot webbing, I1–2II1
1
/
2
–3
–
III2–3
+
IV3
+
–2V (vs. rudimentary or absent), and ventral skin being light to dark brown
with distinct blotches or spots (vs. brown without prominent mottling in U. nagaoi, and white without any blotches
or spots in U. rohani sp. nov. and U. variegatus).
Genetic divergence. For 16S mitochondrial gene sequences, intraspecific variation was not observed between
the sampled populations of Uperodon obscurus (N = 3). Genetically, U. obscurus is closely related to U. palmatus,
from which it was found to differ by an average uncorrected genetic distance of 2.4% (range 2.4–2.6%, N = 12).
For interspecific genetic distances with all other members of the genus, see Table 3.
Remarks. The original description by Günther (1864 “1863”) was very brief. Subsequently Parker (1934)
provided a general description based on the types, without a comparison with other closely related members.
Hence, a detailed description and comparison of this species with all its congeners is provided here.
Redescription of holotype (measurements in mm) (Figs. 3H, 10A–D). Small-sized (SVL 32.2), slender adult
female; head small (HW 9.3, HL 8.8, IFE 4.9, IBE 8.6), less than one-third (27.3%) of body length, wider than long
(HW/HL ratio 1.1); snout truncate in dorsal and ventral view, nearly acute in lateral view, its length (SL 4.2) longer
than horizontal diameter of eye (EL 3.9); loreal region acute with rounded canthus rostralis; interorbital space more
than twice as wide (IUE 3.3) as upper eyelid width (UEW 1.4), and wider than internarial distance (IN 2.7); nostril
much closer to tip of snout (NS 0.7) than to eye (EN 2.4); supratympanic fold distinct, extending from posterior
corner of upper eyelid to insertion of forelimb at axilla; eye diameter (EL 3.9); vomerine odontophores present on
the palate, without teeth; weakly-developed neopalatinal ridges on posterior side of each choana, appear fused with
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the vomerine odontophores; tongue moderately large, oval, slightly emarginate. Forelimbs (FAL 8.8) shorter than
hand length (HAL 9.5); finger length formula I<II<IV<III, tips of all fingers with oval discs, moderately wide
compared to finger width (FD
I
0.8, FW
I
0.4; FD
II
1.1, FW
II
0.5; FD
III
1.0, FW
III
0.5; FD
IV
0.9, FW
IV
0.4); subarticular
tubercles prominent, oval, all present; two well-developed palmar tubercles, oval. Hind limbs relatively long and
thin, thigh length (TL 13.5) longer than shank (SHL 13.3), and shorter than foot (FOL 14.9); relative digit lengths
I<II<V<III<IV, tips of all toes with small truncate to rounded discs, rather wide compared to toe width (TD
I
0.7,
TW
I
0.5; TD
II
0.8, TW
II
0.5; TD
III
0.7, TW
III
0.5; TD
IV
0.7, TW
IV
0.5; TD
V
0.5, TW
V
0.5); foot webbing present, I1–
2II1
1
/
2
–3
–
III2–3
+
IV3
+
–2V; well-developed dermal fringes present on all toes; subarticular tubercles prominent,
oval; two smooth metatarsal tubercles, oval, outer one slightly larger than the inner.
FIGURE 10. Head, hand, foot and foot webbing in Uperodon obscurus, U. palmatus, U. triangularis and U. variegatus. A–D.
Holotype of Uperodon obscurus (NHM 58.11.28.42 ex. BMNH 1947.2.11.20, female): A. Lateral view of head; B. ventra l view
of hand; C. ventral view of foot; D. schematic illustration of foot webbing; E–H. Holotype of Uperodon palmatus (NHM
90.11.8.62 ex. BMNH 1947.2.11.21, female): E. Lateral view of head; F. ventral view of hand; G. ventral view of foot; H.
schematic illustration of foot webbing; I–L. Lectotype of Uperodon triangularis (NHM 74.4.29.897 ex. BMNH 1947.2.11.15,
male): I. Lateral view of head; J. ventral view of hand; K. ventral view of foot; L. schematic illustration of foot webbing; M.
Lectotype of Uperodon variegatus (ZSI 2761, female), lateral view of head; N–P. Referred specimen of Uperodon variegatus
(ZSI 4726, female): N. ventral view of hand; O. ventral view of foot; P. schematic illustration of foot webbing. (Illustrations:
SD Biju).
Skin of snout, between eyes, sides of head, and anterior part of dorsum, shagreened to granular; posterior parts
of back, and upper and lower parts of flank, granular; dorsal surfaces of forelimb, thigh, tibia and tarsus,
shagreened to sparsely granular; throat, chest and abdomen, shagreened; ventral surfaces of forelimb and hind
limb, shagreened (Figs. 3H, 10A).
Colouration. In preservation: Dorsum greyish-brown with dark brown median patch; flanks light greyish-
brown; dorsal surfaces of forelimbs, thigh, shank and foot, light brown; chest and belly, dark grey with faint grey
spots; ventral surfaces of thigh, tibia and foot, greyish-brown with faint grey spots (Fig. 3H). Colour in life (DZ
1366): Dorsum light orangish-brown with irregular blackish-brown spots, prominent blackish-brown band
extending from behind the eye up to vent where it becomes wider; lower parts of flank brown with light bluish-grey
spots; dorsal surfaces of forelimb, thigh, shank and foot, orangish-brown with dark brown cross bands; throat light
brown; chest greyish-brown; belly and ventral surfaces of forelimb, thigh, tibia and foot, brown with light bluish-
grey spots (Figs. 9H–I).
Variations. Morphometric measurements of 17 males and five females, including the holotype, are given in
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Table 4. DZ 1408: dorsal skin bright orange and more prominently granular compared to DZ 1366; DZ 1369:
dorsum orangish-grey with a dark grey median band, and ventral surfaces coffee brown with light bluish-grey
spots, except on the throat.
Secondary sexual characters. Male (DZ 1366): vocal sac externally visible on the lower jaw; female (NHM
1982.1290): ova white, pigmented on pole (diameter 0.7–1.0 mm, N = 20).
Geographical distribution and habitat. Uperodon obscurus is endemic to Sri Lanka and known to occur in
Central, North Western, Sabaragamuwa, Southern and Uva Provinces (Fig. 2D; Tables 1, S1). In the present study,
we confirmed the occurrence of this species in Kandy district (Kandy, Peradeniya, Panwila, Pussellawa and
Udawatta Kele), Kegalle district (Babilapitiya estate), Kurunegala district (Kurunegala), Matale district
(Gammaduwa), Nuwara Eliya district (Bogawantalawa and Laxapana) and Ratnapura district (Sinharaja and
Morningside), from elevations of 100 m up to 1220 m asl. It was collected either from tree hollows in disturbed
forests adjacent to urban areas (Peradeniya and Panwila), or temporary ponds surrounded with vegetation
(Laxapana).
Uperodon palmatus (Parker, 1934)
Parker’s Globular Frog
(Figs. 1, 2A, 2D, 3I, 5H, 10E–H, 11A–E; Tables 1–4)
Original name and description. Ramanella palmata Parker, 1934. A Monograph of the Frogs of the Family
Microhylidae. London: Trustees of the British Museum: 93. Holotype. NHM 90.11.8.62 (ex. BMNH
1947.2.11.21), by original designation. Ty pe loca lity. “Newera Ellya, Ceylon, 7,000 ft” (= Nuwara-Eliya), Sri
Lanka. Current status of specific name. Valid name, as Uperodon palmatus (Parker, 1934).
Comparison. For comparison of Uperodon palmatus with U. globulosus, U. systoma, U. taprobanicus, U.
anamalaiensis, U. montanus, U. mormorata, U. nagaoi and U. obscurus see ‘comparison’ section of those species.
Uperodon palmatus differs from U. triangularis by presence of webbing between toes, I1–2II1–3
–
III1–3IV2
2
/
3
–1V
(vs. toes without webbing); and snout equal to eye diameter, male: EL/SL ratio 1.0, N = 4, female: EL/SL ratio 1.0,
N = 2 (vs. longer, male: EL/SL ratio 0.6–0.9, N = 20, female: EL/SL ratio 0.7–0.9, N = 4) (Table 4). Uperodon
palmatus cannot be confused with three other species, U. nagaoi, U. rohani sp. nov. and U. variegatus, due to
presence of more extensive webbing between toes, I1–2II1–3
–
III1–3IV2
2
/
3
–1V (vs. rudimentary or absent); and
ventral skin light to dark brown with distinct blotches or spots (vs. brown without prominent blotches or spots in U.
nagaoi, and white without blotches or spots in U. rohani sp. nov. and U. variegatus).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon palmatus
showed an average intraspecific distance of 0.1% (range 0–0.2%, N = 4). Genetically, U. palmatus is closely
related to U. obscurus, from which it was found to differ by an average uncorrected genetic distance of 2.4% (range
2.4–2.6%, N = 12). For interspecific genetic distances with all other members of the genus, see Table 3.
Remark. A short general description was published by Parker (1934), though without a comparison with other
closely related members. Hence, a detailed description and comparison of this species with all its congeners is
provided here.
Redescription of holotype (measurements in mm) (Figs. 3I, 10E–H). Small-sized (SVL 34.6), slender adult
female; head small (HW 11.8, HL 9.1, IFE 4.8, IBE 9.4), less than one-third (26.3%) of body length, wider than
long (HW/HL ratio 1.3); snout truncate in dorsal view, truncate to rounded in ventral view, and vertical in lateral
view, its length (SL 3.6) slightly longer than horizontal diameter of eye (EL 3.5); loreal region acute with rounded
canthus rostralis; interorbital space twice as wide (IUE 3.8) as upper eyelid width (UEW 1.9), and internarial
distance (IN 1.9); nostril much closer to tip of snout (NS 0.5) than to eye (EN 1.9); supratympanic fold distinct,
extending from posterior corner of upper eyelid to insertion of forelimb at axilla; eye diameter (EL 3.5); vomerine
odontophores present on the palate, without teeth; indistinct neopalatinal ridges on posterior side of each choana,
appear fused with the vomerine odontophores; tongue moderately large, oval, slightly emarginate. Forelimbs (FAL
7.4) shorter than hand length (HAL 10.4); finger length formula I<II<IV<III, tips of all fingers with oval discs,
moderately wide compared to finger width (FD
I
1.0, FW
I
0.3; FD
II
1.7, FW
II
0.5; FD
III
1.7, FW
III
0.5; FD
IV
1.3, FW
IV
0.5); subarticular tubercles prominent, oval, all present; two well-developed palmar tubercles. Hind limbs
relatively long and thin, thigh length (TL 14.8) longer than shank (SHL 14.6), and shorter than foot (FOL 15.4);
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tips of all toes with small truncate to rounded discs, rather wide compared to toe width (TD
I
0.8, TW
I
0.4; TD
II
0.8,
TW
II
0.5; TD
III
0.9, TW
III
0.5; TD
IV
1.1, TW
IV
0.5; TD
V
0.8, TW
V
0.4); foot webbing present, I1–2II1–3
–
III1–3IV2
2
/
3
–1V; well-developed dermal fringes present on all toes; subarticular tubercles prominent, oval; two oval smooth
metatarsal tubercles.
Skin of snout, between eyes, sides of head, anterior part of dorsum, posterior parts of back, and upper and
lower parts of flank, granular (Fig. 3I); dorsal surfaces of forelimb, thigh, tibia and tarsus, shagreened to sparsely
granular; anterior parts of thigh with spinular projections.
FIGURE 11. Uperodon palmatus, U. triangularis and U. variegatus in life. A–E. Uperodon palmatus: A. Dorsolateral view
(DZ 1372, male); B. dorsolateral view (DZ 1372, male); C. ventral view (DZ 1371, male); D. dorsal view (DZ 1331, male); E.
dorsolateral view (DZ 1331, male); F–K. Uperodon triangularis: F. Dorsolateral view (ZSI/WGRC/V/A/959, male); G. dorsal
view (SDBDU 2015.2857, male); H. ventral view (SDBDU 2015.2857, male); I. dorsolateral view (SDBDU 2011.502, male);
J. dorsolateral view (SDBDU 2012.2200, subadult male); K. dorsofrontal view (SDBDU 2012.2038, male); L–P. Uperodon
variegatus: L. Dorsolateral view (not preserved); M. ventral view (SDBDU 2007.4994, male); N. dorsolateral view (not
preserved); O. dorsolateral view of an albino (or possibly leucistic) form (not preserved); P. dorsolateral view (not preserved).
(Images: SD Biju, Sonali Garg, N & P - Shyamal L, O - Romulus Whitaker).
Colouration. In preservation: Dorsum reddish-brown with light brown spots, prominent light brown median
band starting well behind the eye at the level of shoulders and extending up to the vent; flanks light greyish-brown;
dorsal surfaces of forelimbs, thigh, shank and foot, light greyish-brown; throat reddish brown; chest light brown;
belly light brown with grey spots; ventral surfaces of thigh, tibia and foot, light brown with faint grey spots (Fig.
3I). Colour in life (DZ 1372). Dorsum greyish-brown with irregular blackish-brown spots, prominent dark brown
band starting well behind the eye at the level of shoulders and extending up to vent; lower parts of flank brown with
faint greyish-white spots; dorsal surfaces of forelimb, thigh, shank and foot, greyish-brown with dark brown cross
bands; throat light greyish-brown; chest greyish-brown; belly and ventral surfaces of forelimb, thigh, tibia and foot,
brown with scattered greyish-white spots (Fig. 11A).
Variations. Morphometric measurements of six specimens, including the holotype, are given in Table 4. DZ
1331: dorsum light greyish-brown with a blackish-brown dorsal band; DZ 1371: dorsum orangish-brown with dark
brown dorsal band.
Secondary sexual characters. Male (DZ 1371): vocal sac externally visible on the lower jaw; female
(holotype): ova white, pigmented on pole (diameter 0.7–1.0 mm, N = 10).
Geographical distribution and habitat. Uperodon palmatus is endemic to Sri Lanka with confirmed
distribution only in the high elevation regions (1400–2000 m asl) of the Central Province: Agrapatana,
Bogawantalawa, Hakgala, Horton plains, Pattipola, Nuwara-Eliya and Morey estate (Fig. 2D; Tables 1, S1). See
‘Natural history and breeding’ observations in the respective section, after the species accounts.
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Uperodon triangularis (Günther, 1876 “1875”)
Triangle-spotted Globular Frog
(Figs. 1, 2A, 2C, 3J, 5I, 8I–L, 10I–L, 11F–K; Tables 1–4)
Original name and description. Callula triangularis Günther, 1876 “1875”. Third report on collections of Indian
reptiles obtained by the British Museum. Proceedings of the Zoological Society of London, 1875: 567–577.
Syntypes. NHM 74.4.29.891–900 and NHM 74.4.29.1010, according to Parker (1934). Typ e loca lity. “Malabar”,
India. Current status of specific name. Val i d n am e , a s Uperodon triangularis (Günther, 1876 “1875”). Synonym.
Ramanella triangularis rufeventris Rao, 1937 (Dutta 1997), Neotype by present designation, ZSI/WGRC/V/A/959,
an adult male, from Mudigere, Karnataka state, India, collected by SD Biju and team on 28 September 2012.
Comments. The original description (Günther, 1876 “1875”) mentions 11 syntypes from Malabar, all from
Col. Beddome’s collection. We examined all the syntypes (nine males and two females) available at NHM, and
found NHM 74.4.29.897 (ex. BMNH 1947.2.11.15) to be in agreement with the original description. This specimen
was also found to be in well-preserved condition (Fig. 3J). Since the original description for this nominal taxon was
brief and based on several specimens from an imprecise locality “Malabar” (Biju 2001), and because this species
has often been misidentified by researchers (e.g., Rao 1937; Inger et al. 1984), we herein provide a detailed
description of an adult male specimen from the type series, NHM 74.4.29.897 (ex. BMNH 1947.2.11.15), that may
serve as a reference of this species for future taxonomic works.
Notes on synonymy and neotypification of Ramanella triangularis rufeventris Rao, 1937. A new variety of
this species was described from “Mudigere, Kadur, Saklespur, Hassan” in Karnataka as Ramanella triangularis
rufeventris n. var Rao, 1937. This taxon was considered to be a synonym of Ramanella triangularis (= Uperodon
triangularis) by implication (Dutta 1997), however, without any justification or comparison. In order to verify this
action, we compared the two taxa based on the original description. While describing Ramanella triangularis
rufeventris, Rao stated two major differences between the new variety “of the plains” and the typical forms of
Uperodon triangularis from “Malnad areas” (= Malenadu region of Karnataka): (1) “the snout is pointed, tips
obliquely truncated, prominent”, and (2) “Tibio-tarsal articulation stands well behind the shoulder”, and also
included illustrations (Pl. XXIX. Figs. 18 and 18a). Apart from these characters, he used colour and marking
differences to distinguish the new variety. During our surveys in regions spanning the type locality of this name
bearing taxon (“Mudigere”), we collected Uperodon specimens that were comparable with Rao’s description of
“Ramanella triangularis rufeventris” particularly in having a more reddish dorsal colour. Due to unavailability of
type specimens, considered lost (Dubois 1984), we relied on the original description to compare our new
collections. We found majority of the characters to be same as in Uperodon triangularis, with only minor
differences in colour and markings that were not reliable for distinguishing the two taxa from each other. Further,
since the snout shape mentioned in the original description of Ramanella triangularis rufeventris is contradicting,
we relied on the illustration that clearly shows the snout as “truncate” and not “pointed”, which is also similar to
Uperodon triangularis. The mitochondrial 16S rRNA gene sequences of our new collections comparable with
Rao’s new variety, and those of Uperodon triangularis from the Malabar region of northern Kerala, were also
found to be shallowly divergent (1.5%). Hence, based on findings in this study we confirm Ramanella triangularis
rufeventris Rao, 1937 to be a junior subjective synonym of Callula triangularis Günther, 1876 “1875” [=
Uperodon triangularis (Günther, 1876 “1875”)].
Additionally, since the type material for Ramanella triangularis rufeventris Rao, 1937 is lost (Dubois 1984),
we herein also designate ZSI/WGRC/V/A/959 (an adult male from Mudigere) as the neotype of Ramanella
triangularis rufeventris Rao, 1937, in order to define this taxon objectively and to establish taxonomic stability.
The description provided along with figures (Figs. S1F–J) in the supplementary file S1 also shows that the neotype
is largely consistent with what is known of the former name-bearing type.
Comparison. For comparison of Uperodon triangularis with U. globulosus, U. systoma, U. taprobanicus, U.
anamalaiensis, U. montanus, U. mormorata, U. nagaoi, U. obscurus and U. palmatus see ‘comparison’ section of
those species. Uperodon triangularis cannot be confused with the two other species, U. rohani sp. nov. and U.
variegatus, because of its ventral skin being brown to dark blackish-brown with prominent light grey spots and
blotches (vs. uniformly white with complete absence of spots and blotches).
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon
triangularis showed an average intraspecific distance of 0.9% (range 0–1.6%, N = 9). Populations from Karnataka
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and Kerala were observed to be divergent by up to 1.6%. Genetically, U. triangularis is closely related to U.
mormorata, from which it was found to differ by an average uncorrected genetic distance of 2.9% (range 2.4–3.6%,
N = 99). For interspecific genetic distances with all other members of the genus, see Table 3.
Description of syntype, NHM 74.4.29.897 (measurements in mm) (Figs. 3J, 10I–L). Small-sized (SVL 31.6),
slender adult female; head small (HW 9.6, HL 7.7, IFE 3.6, IBE 8.4), almost one-fourth (24.4%) of body length,
wider than long (HW/HL ratio 1.2); snout rounded to nearly truncate in dorsal and ventral view, nearly acute in
lateral view, its length (SL 3.7) longer than horizontal diameter of eye (EL 2.5); loreal region obtuse with rounded
canthus rostralis; interorbital space wider (IUE 3.4) than upper eyelid width (UEW 1.9); nostril much closer to tip
of snout (NS 0.9) than to eye (EN 2.1); supratympanic fold distinct, extending from posterior corner of upper
eyelid to insertion of forelimb at axilla; eye diameter (EL 2.5); vomerine odontophores present on the palate,
without teeth; presence of two weakly-developed neopalatinal ridges on posterior side of each choana, fused with
the vomerine odontophores; tongue emarginated. Forelimb (FAL 7.1) shorter than hand length (HAL 8.7); finger
length formula I<II<IV<III, tips of all fingers with truncate discs, discs moderately wide compared to finger width
(FD
I
0.8, FW
I
0.6; FD
II
1.0, FW
II
0.7; FD
III
0.9, FW
III
0.6; FD
IV
0.8, FW
IV
0.5); subarticular tubercles prominent,
oval, all present; two well-developed palmar tubercles (inner, oval, 0.7 mm; outer, shallowly lobed, 1.5 mm). Hind
limbs relatively long and thin, thigh length (TL 11.4) longer than shank (SHL 10.9), and shorter than foot (FOL
12.1); tips of all toes with small truncate to rounded discs, discs rather wide compared to toe width; webbing absent
between toes; dermal fringes absent on all toes; subarticular tubercles prominent, oval; two smooth metatarsal
tubercles, oval, outer one longer (1.6 mm) than the inner (0.9 mm).
Skin of snout, between eyes, sides of head, anterior parts of dorsum, posterior parts of back, and upper and
lower parts of flank, shagreened (Fig. 3J); dorsal surfaces of forelimb, thigh, tibia and tarsus, shagreened to
sparsely granular; ventral surface uniformly shagreened.
Colouration. In preservation (NHM 74.4.29.897): Dorsum orangish-brown with solid darker brown median
band on the back starting just above the level of shoulders and extending up to vent, where it becomes wider;
brown stripe between eyes; flanks with orangish-brown patches; lateral sides of head brown, anterior and posterior
parts orangish-brown; ventral surfaces (including limbs) brown with scattered light brown spots (Fig. 3J). Colour
in life (SDBDU 2015.2857): Dorsum orange with solid dark brown median band on the back starting just above the
level of shoulders and extending close to vent, where it becomes wider; dark brown stripe between eyes and
another streak over snout; lateral sides of head and flanks dark brown; dorsal surfaces of limbs (including fingers
and toes) dark brown, elbow and knee orange; ventral surfaces (including limbs) dark greyish-brown with scattered
light grey spots (Figs. 11F–K).
Variations. Morphometric measurements for 19 males and four females are given in Table 4. Dorsal
colouration and markings are highly variable among the examined specimens. SDBDU 2011.503: dorsal skin
shagreened to sparsely granular, orange-coloured dorsolateral band discontinuous towards vent; SDBDU 2011.502:
dorsal skin shagreened to sparsely granular, orange-coloured dorsolateral band patchy and discontinuous; SDBDU
2012.2038: dorsum largely olive grey in colour with dark brown median patch. SDBDU 2008.409: dorsum light
brown with narrow band between upper eyelids.
Secondary sexual characters. Male (SDBDU 2011.504): vocal sac externally visible on the lower jaw;
female (SDBDU 2007.6400): ova white, pigmented on pole (diameter 1.3–1.6 mm, N = 10).
Tad pol e m or ph ol og y. In dorsal view, tadpoles of Uperodon triangularis (Gosner stage 35, N = 2) have an
elliptical body with narrower anterior region, and wider and longer posterior region; snout appears rounded in
dorsal view and depressed in lateral view; eyes small, bulbous, positioned laterally; nasolacrimal duct absent;
narial depressions clearly visible in lateral view, located much closer to snout than eyes, pigmentation not visible
around nasal depressions; spiracle opens near vent, flap absent; vent tubular, positioned mid-ventrally with aperture
opening medially in line with the plane of ventral fin; tail ends bluntly, composed of bilateral myotomic muscle
masses divided by V-shaped septa, with unequal membranes on either side of tail musculature; dorsal fins originate
after tail-body junction, margin of lower fin not parallel to margin of tail muscle; mouth terminal and appears as slit
opening, keratinized structures absent, papillae absent (Figs. 16K–M).
Measurements (mm) (N = 1): Lateral measurements include; maximum height of body (bh) = 3.14, maximum
width of body (bw) = 6.67, maximum diameter of eye (ed) = 0.56, internarial distance (nn) = 1.63, naro-pupular
distance (np) = 2.78, rostro-narial distance (rn) = 0.02, interpupular distance (pp) = 5.43, distance from tip of snout
to opening of spiracle (ss) = 6.24, distance from tip of snout to insertion of upper tail fin (su) = 9.82, snout-vent
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length (svl) = 8.56, total length (tl) = 21.18, distance from vent to tip of tail (vt) = 12.51, maximum height of tail
(ht) = 3.85, tail muscle height (tmh) = 1.87, and tail muscle width (tmw) = 1.89.
Vocalization. Male of Uperodon triangularis (SDBDU 2015.2857) from Suganthagiri produced a single type
of call with pulsatile temporal structure. The calls had uniform intervals and were not delivered in groups. A typical
male call had a duration of 228.9 ms, rise time of 132.3 ms, and fall time of 97.4 ms. The call comprised of 29
pulses that were delivered at a rate of 133.1 pulses/s. The spectrum was characterized by single broad peak with
mean dominant frequency of 2.4 kHz (Figs. 8I–L).
Geographical distribution and habitat. Uperodon triangularis is known to occur in the South Indian states
of Kerala, Karnataka and Tamil Nadu (Fig. 2C; Tables 1, S1). In the present study, we confirmed the presence of
this species in following regions north of Palghat gap in the Western Ghats: Tamil Nadu (Ooty), northern Kerala
(Kalpetta, Mananthavady, Pozhuthana and Suganthagiri) and southern Karnataka (Bygoor, Madikeri, Mudigere,
and Wattakoli). It is a mid to high elevation species (about 675 m up to 2200 m asl). Individuals were found under
fallen logs and stones during the non-breeding season, and observed around temporary puddles and permanent
water bodies during the breeding season. At Bygoor, this species was found about four feet above ground on tree
bark inside coffee plantations. For misidentified records from south of Palghat gap see ‘Notes’ under U.
anamalaiensis.
Uperodon variegatus (Stoliczka, 1872)
Variegated Globular Frog
(Figs. 1, 2A, 3K, 5J, 10M–P, 11L–P, 12A, 13, 14; Tables 1–4)
Original name and description. Callula variegata Stoliczka, 1872. Observations on Indian Batrachia.
Proceedings of the Asiatic Society of Bengal, 1872: 101–113. Lectotype. By present designation, ZSI 2761, an
adult female, SVL 29.8 mm. Type locality. “Ellore” (=Eluru, Andhra Pradesh) India. Current status of specific
name. Va li d n a m e , a s Uperodon variegatus (Stoliczka, 1872). Synonyms. Hylaedactylus carnaticus Jerdon, 1854
“1853” (by Parker 1934), Callula olivacea Günther, 1876 “1875” (by Boulenger 1890), and Ramanella symbioitca
Rao and Ramanna, 1925 (by Parker 1934).
Lectotypification of Callula variegata St olicz ka, 1872. The original description of this species (Stoliczka
1872) mentions two syntypes from “Ellore”. ZSI 2761 was indicated as one of the syntypes (Sclater 1892), and for
the second specimen NHMW 4019 and BMNH 74.11.12.2 have been suggested as possible candidates (Tiedemann
et al. 1994; Dutta & Manamendra-Arachchi 1996; Dutta 1997; Chanda et al. 2000). In the present study, the
available samples at ZSI, Kolkata (ZSI 2761) and Natural History Museum, London (BMNH 74.11.12.2) were
examined. Both these specimens were found to be identical in species status and were in agreement with the
original description. However, since this species was described based on two specimens, and there have been
confusions related to the status of syntypes (Chanda et al. 2000) as well as species identification (e.g., Rao 1937;
Günther 1876 “1875”), apart from the availability of several poorly-defined nominal taxa currently in the
synonymy of Uperodon variegatus, in accordance with Article 74 of The Code, designation of a lectotype from
syntypes is advisable for it “to become the unique bearer of the name of a nominal species-group taxon and the
standard for its application” in order to clarify the taxonomic status of this species. Hence, based on diagnostic
characters found useful for differentiating Uperodon variegatus from its congeners (as stated in the ‘comparison’
section below), we herein designate ZSI 2761 (an adult female, SVL 29.8 mm) from “Ellore”, as the lectotype of
Callula variegata Stoliczka, 1872. The lectotype description provided subsequently is consistent with the original
description.
Comparison. Uperodon variegatus cannot be confused with any of the known member of this genus, except
Uperodon rohani sp. nov., because of its dorsal skin that has prominent spots or blotches, and ventral skin white to
off-white without any spots, blotches or mottling. Uperodon variegatus differs from Uperodon rohani sp. nov. by a
suite of morphological characters (including osteology) and considerable genetic divergence. For comparison with
U. globulosus, U. systoma, U. taprobanicus, U. anamalaiensis, U. montanus, U. mormorata, U. nagaoi, U.
obscurus, U. palmatus, U. triangularis and U. rohani sp. nov. see ‘comparison’ sections of those species.
Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon variegatus
showed an average intraspecific distance of 0.7% (range 0–1.2%, N = 4). The samples from Karnataka and Kerala
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(Western Ghats) showed a divergence of up to 1.2% with the populations from Andhra Pradesh and Tamil Nadu
(Eastern Ghats). Genetically, U. variegatus is closely related to U. rohani sp. nov., from which it was found to
differ by an average uncorrected genetic distance of 2.8% (range 2.4–3.4%, N = 24). For interspecific genetic
distances with all other members of the genus, see Table 3.
Description of lectotype (measurements in mm) (Figs. 3K, 10M). Small-sized (SVL 29.8), slender adult
female; head small (HW 8.7, HL 7.1, IFE 4.3, IBE 8.3), less than one-fourth (23.8%) of body length, wider than
long (HW/HL ratio 1.2); snout rounded to nearly truncate in dorsal and ventral view, acute in lateral view, its length
(SL 3.2) longer than horizontal diameter of eye (EL 2.7); loreal region obtuse with rounded canthus rostralis;
interorbital space more than twice as wide (IUE 3.6) as upper eyelid (UEW 1.4); nostril closer to tip of snout (NS
0.9) than to eye (EN 1.9); supratympanic fold distinct, extending from posterior corner of upper eyelid to insertion
of forelimb at axilla; eye diameter (EL 2.7); vomerine odontophores present on the palate, without teeth; presence
of two weakly-developed neopalatinal ridges on posterior side of each choana, fused with the vomerine
odontophores; tongue emarginate. Forearm (FAL 5.0) shorter than hand length (HAL 7.8); finger length formula
I<II<IV<III, tips of all fingers with truncate discs, discs moderately wide compared to finger width; subarticular
tubercles prominent, oval, all present; two well-developed palmar tubercles (inner, oval, 1.1 mm long; outer,
shallowly lobed, 1.0 mm long). Hind limbs relatively long, thigh length (TL 11.3) longer than shank (SHL 9.9) and
shorter than foot (FOL 12.7); tips of all toes with small truncate to rounded discs; webbing absent between toes;
dermal fringes absent on all toes; subarticular tubercles prominent, oval; two smooth metatarsal tubercles, oval,
outer one longer (1.3 mm long) than the inner (0.7 mm long).
Skin of snout, between eyes, sides of head, anterior part of dorsum, posterior parts of back, and upper and
lower parts of flank, shagreened (Fig. 3K); dorsal surfaces of forelimb, thigh, tibia and tarsus, shagreened to
sparsely granular; ventral surfaces shagreened.
FIGURE 12. A. Distribution of Uperodon variegatus and Uperodon rohani sp. nov. Details of localities reported in the prese nt
study are provided in Table 1. Records from literature are listed in supplementary Table S1 and indicated here based on
approximate coordinates; B–E. Head, hand, foot and schematic illustration of foot webbing in the holotype of Uperodon rohani
sp. nov. (DZ 1449, male): B. lateral view of head; C. ventral view of hand; D. ventral view of foot; E. foot webbing; F–J.
Uperodon rohani sp. nov. (DZ 1449, male) in life: F. dorsolateral view; G. dorsal view; H. ventral view; I. Uperodon rohani sp.
nov. (Paratype, DZ 1485, female), dorsal view; J. Uperodon rohani sp. nov. (DZ 1130, female), dorsolateral view. (Images: SD
Biju).
Colouration. In preservation: Dorsum uniformly light greyish-brown; flanks greyish-brown; lateral sides of
head light brown; ventral surface uniformly light grey (Figs. 3K). Colour in life (SDBDU 2007.4994, male). Dorsal
surfaces (including limbs) dark brown with scattered olive yellow spots throughout; lateral sides of head dark
greyish-brown with scattered olive coloured spots; throat dark grey with calling patch; chest, belly and thigh, flesh-
white without markings; ventral surfaces of forelimb, tibia and foot, light grey (Fig. 11L–M).
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FIGURE 13. Principal Component Analyses showing morphometric differentiation between Uperodon rohani sp. nov. and U.
variegatus based on 17 morphological characters. Left: Plot for the first two factor planes explaining 54.4% of the variations
among 28 adult male specimens. Right: Plot for the first two factor planes explaining 73% of the variations among 15 adult
female specimens. Factor loadings for each component are provided in Table S2.
FIGURE 14. Cleared and double stained specimens of Uperodon rohani sp. nov. (above) and U. variegatus (below). From left
to right: Dorsal view of the crania composed of nasals (NS), frontoparietals (FP), prootics (PR), exoccipitals (EX) and
squamosals (SQ); anteroventral view of the crania showing components of the upper jaw, premaxillae (PM) and maxillae
(MA); ventral view of the lower jaw comprising of dentaries (DN), angulosplenials (AG) and mentomeckelians (MM). Bones
are indicated in red, cartilage in blue. (Images: Gayani Senevirathne).
Variations. Morphometric measurements for 12 males and seven females, including the lectotype, are given in
Table 4. Dorsal colouration and markings were highly variable among the examined specimens. SDBDU
2007.6347: dorsum brown with yellowish-grey blotches in life (that turns light brown with grey blotches in
preservation); SDBDU 2005.4726: light brown with olive grey blotches in life. Albino (or possibly leucistic) forms
of this species are known from Mamallapuram in Tamil Nadu (Fig. 11O).
Secondary sexual characters. Male: vocal sac externally visible on the lower jaw; females (SDBDU
2005.4726): ova white, pigmented on pole (diameter 0.7–1.0 mm, N = 20).
Geographical distribution and habitat. Uperodon variegatus is a widely distributed species in India. It is
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known to occur right from Tamil Nadu and Kerala in the south, northwards up to Karnataka, Andhra Pradesh,
Telangana, Odisha, Jharkhand and West Bengal (Fig. 12A; Tables 1, S1). In the present study, genetically identified
individuals of this species were largely collected from urban landscapes and disturbed secondary forests (Tamil
Nadu: Chengalpattu and Mamallapuram; Kerala: Karamana, Meladoor and Palakkad; Karnataka: Bangalore; and
Andhra Pradesh: Srikakulam), from sea level up to 900 m elevations. This species is predominantly found in drier
regions towards the eastern coast. At Mamallapuram, it was found in brackish waters, just a few metres away from
the seacoast.
Uperodon rohani sp. nov.
Rohan’s Globular Frog
(Figs. 1, 2A, 2D, 3L, 5K, 12, 13, 14; Tables 1–4)
http://zoobank.org/urn:lsid:zoobank.org:act:D612A25A-E09E-4193-8B89-3DC62746FB55
Holotype. DZ 1449, an adult male, from Mihintale, Anuradhapura district, Sri Lanka, collected by Madhava
Meegaskumbura and team, on 02 October 2014. Paratypes. DZ 1448, DZ 1450 and DZ 1374, three adult males,
collected along with the holotype; DZ 1336, an adult male, from Bundala NP, Hambantota district, Sri Lanka,
collected by MM and team, on 04 November 2013; DZ 1520 and DZ 1521, two adult males, and DZ 1425, an adult
female, from Makandura, Kurunegala district, Sri Lanka, collected by MM and team, on 11 September 2014; DZ
1486, an adult male, and DZ 1485, an adult female, from Ampara, Ampara district, Sri Lanka, collected by MM
and team, on 10 October 2012; DZ 1129–DZ 1132, four adult females, from Dambulla, Matale district, Sri Lanka,
collected by MM and team, on 04 January 2013. Referred specimens. NHM 1931.2.1.21, an adult male, from
“Ceylon”, Sri Lanka, collected by GH Herry; NHM 1973.3015, an adult male, from Kurunegala, “Kurunegala
dist.,” Sri Lanka, collected by R Sananayake; NMH 1976.264–NHM 1976.265, two adult males, from Badulla,
“Hambantota dist.,” Sri Lanka, collected by JC Naggs; WHT 5320, an adult male, from Nochchiyagama,
Anuradhapura district, Sri Lanka, collected by MM Bahir, C Gans and C Yahalmulla, on 18 November 2000; WHT
01188, an adult male, from Sigiriya, Near Rest house, Matale district, Sri Lanka, collected by D Gabadge, on 08
February 1996; WHT 1749, an adult male, from Polonnaruwa, Polonnaruwa district, Sri Lanka, collected by A
Nollert, on 28 July 1997; WHT 01189, an adult male, from Anuradhapura, Anuradhapura district, Sri Lanka,
collected by D Gabadage and MM Bahir, on 11 August 1995; NHM 1973.3014, an adult female, from Kurunegala:
Ceylon, “Kurunegala dist.,” Sri Lanka, collected by R Sananayake; WHT 01191, an adult female, from Palatupana,
Kirinda, Hambantota district, Sri Lanka, collected by U. Premachandra, in August 1995.
Etymology. This species is named after Rohan Pethiyagoda, in recognition of his contribution to biodiversity
studies in India and Sri Lanka, especially on frogs and fishes. The species epithet is formed as a noun in the
genitive case.
Comparison. Uperodon rohani sp. nov. cannot be confused with U. globulosus, U. systoma and U.
taprobanicus due to its smaller snout-vent size, male SVL 26–34 mm, N = 16, female SVL 28–35 mm, N = 8 (vs.
larger, U. globulosus male SVL 51–56 mm, N = 2, female SVL 66–72 mm, N = 3; U. systoma male SVL 51–56 mm,
N = 2, female SVL 47–64 mm, N = 4; and U. taprobanicus male SVL 42–52 mm, N = 3, female SVL 53–55 mm, N
= 2). Uperodon rohani sp. nov. specifically also differs from the former two by its finger tips having truncate discs
(vs. rounded without discs), and presence of two weakly-developed neopalatinal ridges on posterior side of each
choana, fused with the vomerine odontophores (vs. two prominent dermal projections on the neopalatinal ridges on
posterior side of each choana, separated from the vomerine odontophores); and from U. taprobanicus by presence of
two weakly-developed neopalatinal ridges on posterior side of each choana, fused with the vomerine odontophores
(vs. well-developed neopalatinal ridges). Further, U. rohani sp. nov. differs from U. anamalaiensis, U. montanus, U.
obscurus and U. palmatus by its ventral skin being white to off-white in life (that turns greyish-white in
preservation) without any spots or blotches (vs. light or dark brown with prominent spots or blotches), and absence
of webbing between toes in both the sexes (vs. present). Uperodon rohani sp. nov. differs from U. nagaoi by its
ventral skin being white to off-white in life (that turns greyish-white in preservation) without any spots or speckles
(vs. brown to dark brown in life that turns brown or dark greyish-brown in preservation, with or without
inconspicuous spots or speckles). For detailed comparison of U. rohani sp. nov. with U. globulosus, U. systoma, U.
taprobanicus, U. anamalaiensis, U. montanus, U. mormorata, U. nagaoi, U. obscurus, U. palmatus and U.
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triangularis see ‘comparison’ section of those species. Uperodon rohani sp. nov. is morphologically and genetically
close to U. variegatus, from which it differs by its snout nearly equal to eye diameter, male SL/EL ratio 1.0–1.2, N =
16, female SL/EL ratio 0.8–1.0, N = 8 (vs. snout relatively longer than eye, male SL/EL ratio 1.2–1.6, N = 12, female
SL/EL ratio 1.2–1.4, N = 7); head relatively narrower, male HW/HL 1.0–1.1, N = 16, female HW/HL 1.0–1.2, N = 8
(vs. head relatively wider, male HW/HL 1.1–1.4, N = 12, female HW/HL 1.1–1.4, N = 7); and shank relatively
longer than thigh, male SHL/TL ratio 1.0–1.1, N = 8, female SHL/TL ratio 1.1, N = 8 (vs. relatively shorter, male
SHL/TL ratio 0.9–1.0, N = 12, female SHL/TL ratio 0.8–0.9, N = 7) (Table 4).
The new species cannot be confused with previously available names, Engystoma marmoratum Cuvier, 1829
and Systoma leschenaultii Tschudi, 1838, currently under the synonymy of Uperodon systoma Schneider, 1799.
The former was described from “l’Inde” (= India) and as stated by Peters (1864 “1863”)—“bereits richtig erkannt
batten = Uperodon marmoratum D. B. und miifste daher den Species namen “systoma” behalten”, was already
correctly recognized as Uperodon marmoratum (= Uperodon systoma) by Duméril and Bibron (1841). The latter,
also stated by Duméril and Bibron (1841)—“peut-être de la même espèce que notre Upérodonte marbré”, is
perhaps the same as Uperodon systoma, from which the new species cannot be confused based on several
morphological characters such as smaller body size (vs. larger), finger tips with truncate discs (vs. rounded without
discs) and vomerine odontophores fused with neopalatinal ridges (vs. separated). Furthermore, three other names—
Hylaedactylus carnaticus Jerdon, 1854 “1853”, Callula olivacea Günther, 1876 “1875” and Ramanella symbioitca
Rao and Ramanna, 1925, currently under the synonymy of Uperodon variegatus Stoliczka, 1872, were also
described from regions in India (“Carnatic”, “Yellagherry hills” and “Ellore”, respectively), while the distribution
of U. rohani sp. nov. is restricted to Sri Lanka. For specific differences with U. systoma and U. variegatus, see the
detailed comparison with those species.
Morphometric differentiation. Uperodon rohani sp. nov. showed morphological differentiation from its
closely related species, U. variegatus. Principal Component Analysis (PCA) for the adult male specimens (U.
rohani sp. nov.: N = 12; U. variegatus: N = 16) recovered five Principal Components (PC) with eigenvalues >1.0
that accounted for 77.2% of the total variance; PC 1 explained 32.3% of the variations, with highest loadings for
SVL, HW, IUE, IFE, MN, HAL and TL; PC 2 explained 22.1% of the variations, with highest loadings for SL,
IBE, EN, SHL and FOL; PC 3 explained 10% of the variations, with highest loadings for UEW and NS; PC 4
explained 6.9% of the variations, with highest loadings for EL; PC 5 explained 6% of the variations; and the
remaining factors altogether explained 22.8% of the variations (Table S2). Principal Component Analysis (PCA)
for the adult female specimens (U. rohani sp. nov.: N = 7; U. variegatus: N = 8) recovered three Principal
Components (PC) with eigenvalues >1.0 that accounted for 82.3% of the total variance; PC 1 explained 46.2% of
the variations, with highest loadings for SVL, HW, SL, IUE, UEW, IFE, IBE, MN, EN and TL; PC 2 explained
26.9% of the variations, with highest loadings for HL, FAL, HAL and SHL; PC3 explained 9.2% of the variations,
with highest loadings for EN; and the remaining factors altogether explained 17.7% of the variations (Table S2).
Projections of factor planes 1 and 2 showed distinct clusters for the two species (Fig. 13).
Osteological comparison. Uperodon rohani sp. nov. and its closely related species, U. variegatus, have the
following skeletal features: Anteriorly slender, basally wider crania, with widest at anterior margins of tympanic
annulus in both species. Cranium length is 93% of its width in U. rohani sp. nov. and 84.2% in U. variegatus.
Frontoparietals are largest, covering more than half of crania, slender and quadrangular shaped in both species.
Nasals, lying dorsal to nasal capsules, triangular-shaped, medial edge 20.6% of the mid-cranial width in U.
variegatus and 20.9% in U. rohani sp. nov. Posterolateral margins of frontoparietals slightly overlay anterolateral
margins of prootics. Major differences between the two species concentrate mainly on three bones of the lower and
upper jaws—premaxillae, mentomeckelians and dentaries. Premaxillae usually composed of alary, dentary and
maxillary processes. In both species, premaxillae are edentate. However, shape of dentary process differs
immensely, wherein U. rohani sp. nov. has well-bifurcated dentary process with 74.9% height compared to width
of premaxillae. Comparatively, dentary process of U. variegatus is rectangular in shape, with a slight median
depression, and 27.4% height compared to premaxillary width. Mentomeckelians are observed as cylindrical
endochondral elements, anterior to the anterior ends of Meckel’s cartilages, only in U. variegatus.
Mentomeckelians are absent in U. rohani sp. nov.; however, the infrarostrals, which are the precursors of
Mentomeckelians are still present and can be seen clearly (Fig. 14). Development of dentaries also shows
differences between the two species. Dentaries are well-developed, extend for about one-third and two-third of the
length of the mandible arc in U. rohani sp. nov. and U. variegatus, respectively (Fig. 14).
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Genetic divergence. For 16S mitochondrial gene sequences, the sampled populations of Uperodon rohani sp.
nov. showed an average intraspecific distance of 0.5% (range 0–1.4%, N = 6). The sample from Bundala showed a
divergence of up to 1.4% from other populations. Genetically, U. rohani sp. nov. is closely related to U. variegatus,
from which it was found to differ by an average uncorrected genetic distance of 2.8% (range 2.4–3.4%, N = 24).
For interspecific genetic distances with all other members of the genus, see Table 3.
Description of holotype (measurements in mm) (3L, 5K, 12B–J). Small-sized (SVL 26.0), relatively slender
adult male; head small (HW 6.7, HL 6.2, IFE 3.5, IBE 5.5), less than one-fourth (23.4%) of body length, wider than
long (HW/HL ratio 1.1); snout subovoid in dorsal, subovoid to nearly truncate in ventral view, rounded in lateral
view, its length (SL 2.6) slightly longer than horizontal diameter of eye (EL 2.5); loreal region rounded with
vertical canthus rostralis; interorbital space wider (IUE 2.9) than upper eyelid (UEW 1.6) and internarial distance
(IN 1.9); nostril closer to tip of snout (NS 0.7) than to eye (EN 1.4); supratympanic fold distinct, extending from
posterior corner of upper eyelid near to insertion of forelimb at axilla; eye diameter (EL 2.5); vomerine
odontophores present on the palate, without teeth; presence of two weakly-developed neopalatinal ridges on
posterior side of each choana, fused with the vomerine odontophores; tongue emarginate. Forelimbs moderately
long and thin; forearm (FAL 5.2) shorter than hand length (HAL 7.2); finger length formula I<II<IV<III, tips of all
fingers with oval discs, discs moderately wide compared to finger width (FD
I
0.9, FW
I
0.4; FD
II
1.3, FW
II
0.4; FD
III
1.2, FW
III
0.4; FD
IV
1.2, FW
IV
0.5); subarticular tubercles prominent, oval, all present; two well-developed palmar
tubercles, oval. Hind limbs relatively long and thin, thigh length (TL 9.2) slightly shorter than shank (SHL 9.4) and
shorter than foot (FOL 11.1); relative digit lengths I<II<V<III<IV; tips of all toes slightly enlarged with truncate to
rounded discs, discs rather wide compared to toe width (TD
I
0.7, TW
I
0.6; TD
II
0.8, TW
II
0.6; TD
III
0.7, TW
III
0.5;
TD
IV
0.7, TW
IV
0.5; TD
V
0.5, TW
V
0.5); foot webbing absent; subarticular tubercles prominent, oval; two smooth
metatarsal tubercles, oval, outer one slightly longer than the inner.
Skin of dorsum, snout, and between eyes, shagreened; sides of head, posterior part of back, and upper and
lower parts of flank, smooth; dorsal surfaces of forelimb, thigh, tibia and tarsus, smooth to shagreened; ventral
surfaces smooth.
Colouration. In preservation: Dorsum dark brown with irregular olive grey coloured blotches, pair of faint
and irregular olive grey longitudinal streaks extending from upper eyelids up to vent; dorsal surfaces of forelimbs
and hind limbs lighter in colour compared to dorsum and with faint yellowish-grey spots or blotches; sides of head
brown; ventral surfaces (including limbs) greyish-white, throat with blackish-brown calling patch (Fig. 3L). Colour
in life: Dorsum reddish brown with pair of irregular olive yellow coloured longitudinal streaks extending from
upper eyelids up to vent, scattered olive yellow coloured spots and blotches especially towards dorsolateral
surfaces and posterior part of back, larger olive yellow blotch in center of dorsum, few scattered spots on head,
olive yellow patch between nostrils and upper eyelids; lateral sides of head brown with scattered olive yellow spots
near margins of mouth; anterior and posterior parts of groin light brown; dorsal surfaces of limbs brown to light
brown with scattered olive yellow spots; throat light brown with bluish-black calling patch and irregular light grey
speckles; chest, belly and thighs, greyish-white with light brown margins; limbs light brown with irregular light
grey speckles (Figs. 12F–J).
Variations. Morphometric measurements for 16 males and eight females, including the type series, are given
in Table 4. DZ 1374 and DZ 1336: dorsal skin smooth; DZ 1129: lateral surfaces slightly granular; DZ 1374: lateral
surfaces of head slightly granular. DZ 1374: dorsum coffee brown in colour with prominent irregular olive grey
spots or patches; DZ 1521: dorsum dark brown with olive grey blotches; DZ 1425: dorsum dark brown with small
olive grey spots scattered throughout, scattered spots also seen on surfaces of hand, foot, sides of head and groin;
DZ 1485: dorsum darker brown with fewer blotches.
Secondary sexual characters. Male (DZ 1449): vocal sac externally visible on the lower jaw; female (DZ
1425): ova white, pigmented on poles (diameter 0.6–1.0 mm).
Geographical distribution and habitat. Uperodon rohani sp. nov. is endemic to Sri Lanka, where it is widely
distributed throughout the island (Fig. 2D). In the present study, we genetically and/or morphologically confirmed
the occurrence of this species at several localities in the Central, Eastern, Northern, North Central, North Western,
Southern, and Uva provinces (Table 1). Previous records of U. variegatus from Sri Lanka refer to this species (see
supplementary Table S1 and references thereto).
Comment. This species was historically considered to be conspecific with the Indian species Uperodon
variegatus and widely reported as Ramanella variegata (see Table 1 and supplementary Table S1).
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Natural history and reproductive behaviour
Like most microhylids, Uperodon are explosive breeders (e.g., Altig & McDiarmid 1999b; Manamendra-Arachchi
& Pethiyagoda 2001). Nevertheless, an extraordinary diversity has been observed within the genus in terms of
reproductive behaviour, breeding ecology and larval development. The Sri Lankan species U. nagaoi, U. obscurus
and U. palmatus are reported to be phytotelm breeders (Meegaskumbura 1999; Meegaskumbura 2001;
Manamendra-Arachchi & Pethiyagoda 2001; Chandrasegaran & Ranawana 2002; Karunarathna & Amarasinghe
2009; Bowatte & Meegaskumbura 2012). Of these three species, U. nagaoi is suggested to obligately inhabit tree
holes, making it ecologically specialized (Manamendra-Arachchi & Pethiyagoda 2001). We observed a female of
U. nagaoi submerged in a water-filled tree hole with a clutch of gel-encapsulated eggs adhered to the inner walls
(about 4 to 5 cm above the water level) at Hiyare forest reserve in Sri Lanka (Figs. 15I–J), suggesting that this
species might exhibit parental care behavior, similar to that observed in some tree hole breeding frog species (e.g.,
Biju et al. 2016). On the other hand, U. obscurus and U. palmatus are more adaptive, commonly found on forest
floor, and also breed in small terrestrial ponds, rock pools and puddles (Meegaskumbura 1999; Meegaskumbura
2001; Manamendra-Arachchi & Pethiyagoda 2001; Ukuwela et al. 2010; Janzen & Bopage 2011). Chandrasegaran
& Ranawana (2002) noticed various-staged tadpoles of U. obscurus inside a tree hole at the same time and pointed
out that this species may breed multiples times in a season. Conversely, this also suggests the possibility of sharing
of phytotelm by multiple females during a single season.
As for the Indian members, Uperodon montanus is known to inhabit and breed inside tree holes (e.g., Krishna
et al. 2004; Krishna & Bosch 2007). In this study, we observed U. montanus inside a water-filled tree hole about
six metres above ground at Kesavapara (Nelliyampathy, Kerala) and a damp tree hole about three metres above
ground at Upper Manalar (Kerala). However, at Pandimotta (Shendurney WLS, Kerala) calling males of U.
montanus were collected from a temporary puddle. We also collected U. anamalaiensis from tree holes about 8–10
metres above the ground at Chathankod (Kerala) in the southern Western Ghats. This species, however, was
observed to breed in shallow-water channels on the ground immediately after the first rains. At Methooty in Kerala,
U. anamalaiensis was found breeding in rock puddles on exposed streams during the month of May. Males of this
species exhibit a single vocal sac while calling and form axillary amplexus with the females. Pigmented eggs of U.
anamalaiensis, surrounded with a thick jelly layer, can be found as surface films on the breeding sites. Inger et al.
(1984) observed microhylid tadpoles inside a tree hole at Ponmudi (Kerala) and suspected them to belong to U.
triangularis. However, since Inger et al.’s (1984) descriptions match closely with U. anamalaiensis adult and
tadpoles reported in our study (see ‘Comparative morphology of tadpoles’ section), and the presence of U.
anamalaiensis at Ponmudi is also genetically confirmed (Fig. 1), we consider their observations to be of this
species. Although U. anamalaiensis and U. montanus are reported from tree holes, these are not necessarily
phytotelm inhabitants or breeders, as evident from available information and our observations. Uperodon
triangularis has occasionally been reported from tree holes (Abraham et al. 2001) but all collections of this species
in the present study were from the ground, usually around puddles, tree bases or from fallen logs. Uperodon
mormorata may also occasionally be arboreal (Raju Vyas personal communication). We found this species on a
tree trunk at Shimoga (Karnataka) 4–5 metres above ground.
Another species, Uperodon taprobanicus is also a semi arboreal species widely reported from tree holes both
in India and Sri Lanka (e.g., Daniels 2005; Siliwal & Ravichandran 2008; Ganesh 2012). In the present study, this
species was collected from tree holes about 4–5 metres above ground at Kalpetta (Wayanad, Kerala) and Odisha,
but also from roads (Bangalore town, Karnataka) and agricultural fields (Wayanad) in India. In West Bengal this
species has been found inside tree holes about one and a half metres above the ground, and usually descends from
tree holes and hollow trunks climbing down backwards with its legs first (Deuti & Goswami 1995). In Sri Lanka it
was observed in several habitats such as coconut trunks at Makandura (1–6 individuals), tree trunks of Terminalia
arjuna adjacent to permanent water holes at Ampara and Kurunegala, termite mounds at Dambulla and on the
roadside at Bundala, Katharagama (Hambanthota district) and Anuradhapura.
The wide-ranging Indian species Uperodon variegatus is found closest to human-dominated landscapes such
as agricultural fields, wall crevices, and often from damp areas of houses such as bathrooms and water closets. This
species was observed to breed in stagnant pools and water tanks. Rao & Ramanna (1925) also reported U.
variegatus from scorpion burrows. The breeding behavior of the new species U. rohani sp. nov. is close to its
congener U. variegatus. Amplexus in U. rohani sp. nov. is axillary as in the case of most species in this genus.
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FIGURE 15. Breeding behavior in Uperodon species. A–C. Uperodon globulosus: A. Male calling from a temporary water
puddle (not preserved); B. Male and female in axillary amplexus; C. Eggs floating on water surface; D–G. Uperodon
anamalaiensis: D. Male calling from a temporary pool of water on a hill stream (SDBDU 2012.1998); E. Male and female in
axillary amplexus; F. jelly-covered eggs floating on water surface; G. individual eggs after removal of the jelly layer; H. Male
of Uperodon palmatus calling from a small permanent water pool adjacent to a forest; I–J. Uperodon nagaoi: I. early
developmental stages inside a gel-encapsulated egg mass on inner wall of a tree hole; J. oviposition site with gel-encapsulated
eggs adhered on inner wall of a tree hole (about 4 to 5 cm above the water level on a tree at the edge of a stream in a lowland
canopy covered forest) along with an adult male found submerged about 1 cm below the water surface. (Images: A–C.
Samadhan Phuge, D–G. SD Biju, H–J. Nayana Wijayathilaka).
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Among all the members of this genus, Uperodon globulosus and U. systoma are unique because of their
fossorial habit that also makes it difficult to sight them. In India, our collections of U. systoma were made either
from small burrows in dry agricultural fields (Coimbatore and Kunnapattu in Tamil Nadu) or tilled soil in
uncultivated fields (Bannerghatta, Karnataka). We did not observe the breeding of U. systoma, however, though on
one occasion erratic calls were heard from the ground adjacent to a temporary pond in Coimbatore town (Tamil
Nadu) during the month of October. Individuals were located close to soil burrows. In Sri Lanka, specimens were
collected from Makandura (Kurunegala district), Dambulla (Matale district) and Anuradhapura (Anuradhapura
district). At Makandura, more than 12 animals were found feeding on termites in and around a termite mound. One
of the individuals quickly escaped into a ground burrow upon being disturbed. During a repeat visit to this site, an
animal was seen emerging out of the same burrow suggesting that this species may use permanent tunnels.
However, so far no information is available on the breeding behavior of these frogs. The second fossorial species,
U. globulosus, is currently known to occur only in India. In the present study, large aggregations of males were
either observed in and around temporary rainwater pools and puddles (Pune, Maharashtra, India; Jaldapara, West
Bengal, India) or on roads (Tamhini Ghat, Maharashtra, India) within a few days after the first monsoon showers.
Breeding activity was observed between 18:00–02:00 hrs. Male calls are loud and produced by inflating a single
gular pouch (Fig. 15A). Calling individuals and amplecting pairs (in axillary amplexus) were found floating inside
or at the edges of stagnant water, with the male’s head and throat rested over the female’s back. During amplexus, a
thick cream-coloured sticky substance is observed and it seems to help the pair stick together while mating since
pairs were also seen struggling to detach themselves after completion of egg-laying. Before egg laying the female
bends its body in the form of an arch. Eggs were seen as surface films in puddles (Fig. 15C). Upon being disturbed,
U. globulosus frogs are observed to inhale air and swell the two sides of their abdomen like a balloon, while also
secreting a sticky white fluid.
The breeding behavior of U. taprobanicus, which is also a semi-fossorial species (Dutta & Manamendra-
Arachchi 1996), is also likely to be similar to that observed in U. globulosus. In India, its breeding activities are
usually observed to start with the first rains in June and extend up to the end of July. After a heavy shower, they
emerge from their hiding places and mostly breed in pools of dirty water. Males inflate single vocal sacs while
calling and use them as floats in water. During amplexus, the mating pairs lay eggs while floating passively in
water. At Pondicherry, Tamil Nadu (India), a male and a female of U. taprobanicus were found in amplexus while
floating in a roadside ditch. Upon handling the pair, we noticed a sticky substance with strong odour over the
female’s dorsal body surface, especially near the head region, where the male’s throat was pressed against the
female during axillary amplexus. When disturbed, this species is also reported to exhibit a defense behavior by
inhaling air, swelling up like a ball while keeping the head down (Somaweera 2006) and secreting a slimy white
fluid through skin pores.
Male advertisement calls have so far been described for Uperodon taprobanicus (reported as ‘Kaloula pulchra’
from “Ceylon”, Nelson 1973), U. triangularis (Kuramoto & Dubois 2009), U. montanus (reported as U.
‘anamalaiensis’, Harpalani et al. 2015), U. ‘montanus’ (Kadadevaru et al. 1998; Krishna & Bosch 2007), U.
variegatus (Kanamadi et al. 1993), U. nagaoi (Manamendra-Arachchi & Pethiyagoda 2001), and U. obscurus
(Nelson 1973). In the present study, we have further described the call structure in U. anamalaiensis, U. mormorata
and U. triangularis, based on genetically confirmed identities (see ‘Vocalization’ sections under the individual
species accounts).
Comparative morphology of tadpoles
Free-swimming exotrophic tadpoles (Altig & Johnston 1989) are one of the three developmental modes observed
in the family Microhylidae (Parker 1934). Many microhylids are known to be suspension feeders of small particles
while swimming (Wells 2007); tadpoles of Uperodon also fall under this category. Microhylid tadpoles are also
known to exhibit variations in terms of tail morphology and shape of the spiracular opening as well as the location
of the mouth and spiracle (Donnelly et al. 1990; Bowatte & Meegaskumbura 2012). However, only a few studies
are available on tadpole morphology of species in this group (Rao 1918; Kirtisinghe 1958; Ukuwela et al. 2010;
Manamendra-Arachchi & Pethiyagoda 2001; Bowatte & Meegaskumbura 2012). The location of mouth in U.
montanus (Parker 1934), U. triangularis (Rao 1918; present study), U. obscurus (Bowatte & Meegaskumbura 2012),
GARG ET AL.
48
·
Zootaxa 4384 (1) © 2018 Magnolia Press
TABLE 3. Inter and intraspecific uncorrected p-distances (in percent) for the mitochondrial 16S rRNA gene sequences. Values in bold indicate intraspecific distances.
Species U.
anamalaiensis
U.
montanus
U.
mormorata
U.
nagaoi
U.
obscurus
U.
palmatus
U. rohani
sp. nov.
U.
triangularis
U.
variegatus
U.
globulosus
U.
systoma
U.
taprobanicus
U. anamalaiensis 0.4
(0–0.8)
U. montanus 3.2
(2.6–4.0)
0.7
(0–1.2)
U. mormorata 5.7
(5.3–6.3)
5.6
(4.8–6.1)
0.6
(0–1.2)
U. nagaoi 7.3
(7.2–7.4)
6.6
(6.4–6.8)
7.2
(7.0–7.4)
0
U. obscurus 7.8
(7.6–8.0)
6.7
(6.6–7.0)
6.7
(6.3–6.9)
5.6
(5.6–5.6)
0
U. palmatus 6.2
(6.0–6.4)
5.5
(5.4–5.8)
5.8
(5.5–6.1)
4.0
(4.0–4.2)
2.4
(2.4–2.6)
0.1
(0–0.2)
U. rohani
sp. nov.
5.2
(4.6–5.6)
5.3
(5.0–5.8)
6.6
(6.3–7.1)
7.2
(7.0–7.4)
7.7
(7.5–7.9)
6.5
(6.3–6.7)
0.5
(0–1.4)
U. triangularis 6.6
(6.3–7.1)
5.9
(5.1–6.5)
2.9
(2.4–3.6)
8.0
(7.5–8.5)
6.7
(6.5–6.7)
6.0
(5.7–6.5)
7.3
(6.7–7.9)
0.9
(0–1.6)
U. variegatus 6.3
(5.8–6.7)
5.8
(5.4–6.4)
7.0
(6.5–7.5)
7.1
(7.0–7.4)
7.7
(7.3–8.0)
7.0
(6.9–7.1)
2.8
(2.4–3.4)
7.2
(6.7–7.9)
0.7
(0–1.2)
U. globulosus 8.1
(7.7–8.4)
7.7
(7.2–8.0)
7.7
(7.5–7.9)
8.1
(8.1–8.1)
8.2
(8.2–8.2)
7.6
(7.5–7.7)
8.2
(8.1–8.5)
8.3
(7.9–8.9)
8.8
(8.5–9.1)
0
U. systoma 8.0
(7.5–8.6)
7.5
(6.8–8.4)
7.6
(7.1–8.3)
8.9
(8.8–9.0)
8.4
(8.2–8.8)
8.0
(7.8–8.2)
9.3
(8.9–10.1)
8.2
(7.5–8.9)
9.4
(8.7–9.9)
3.5
(3.2–4.0)
0.9
(0–2.2)
U. taprobanicus 9.6
(9.1–10.3)
9.7
(8.9–10.3)
8.7
(8.1–9.3)
10.8
(10.5–11.1)
10.2
(9.9–10.5)
10.4
(10.1–10.9)
10.1
(9.5–10.7)
8.7
(8.3–9.1)
10.7
(10.5–11.5)
9.9
(9.7–10.1)
9.1
(8.3–9.7)
0.3
(0–1.2)
Zootaxa 4384 (1) © 2018 Magnolia Press
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49
TAXONOMIC REVIEW OF GENUS UPERODON
TABLE 4. Morphometric measurements of Uperodon species. All measurements are in millimetres (mm). The status of each specimen is given after the museum number: Holotype (HT), Paratype (PT),
Neotype (NT), Lectotype (LT), Syntype (ST).
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
Uperodon systoma (Schneider, 1799)
F "India orientali" ZMB 3551 (ST)
49.7 11.7 7.9 4.8 4.9 4.8 2.6 6.6 12.4 7.1 1.4 1.6 13.2 23.8 18.3 15.4 18.6
F Kunnapattu SDBDU 2005.2806
63.3 15.5 10.2 5.0 5.2 6.5 3.3 7.9 14.8 9.3 1.5 1.9 17.1 27.3 23.2 19.4 23.6
F Kunnapattu SDBDU 2005.4724
48.2 14.1 8.8 4.5 4.4 5.6 3.1 6.5 13.0 7.4 1.7 2.1 13.9 23.9 19.4 16.4 20.5
F Coimbatore SDBDU 2005.2732
47.0 12.5 8.9 4.3 4.1 5.1 2.6 6.8 11.9 7.3 1.9 2.0 13.3 21.3 17.3 15.2 19.0
Average 52.1 13.5 9.0 4.7 4.7 5.5 2.9 7.0 13.0 7.8 1.6 1.9 14.4 24.1 19.6 16.6 20.4
Standard deviation 7.6 1.7 0.9 0.3 0.5 0.7 0.4 0.6 1.3 1.0 0.2 0.2 1.8 2.5 2.6 1.9 2.3
M Kunnapattu SDBDU 2005.2787
55.2 13.6 8.7 5.0 4.9 5.8 3.1 7.3 13.2 8.8 2.0 2.2 15.6 24.5 22.1 17.4 21.1
M Dambulla DZ 1128
51.0 14.8 11.5 4.8 5.1 5.8 3.3 6.5 13.1 9.3 1.8 2.1 12.2 20.9 21.6 17.6 22.0
Average 53.1 14.2 10.1 4.9 5.0 5.8 3.2 6.9 13.2 9.1 1.9 2.2 13.9 22.7 21.9 17.5 21.6
Standard deviation 3.0 0.8 2.0 0.1 0.1 0.0 0.1 0.6 0.1 0.4 0.1 0.1 2.4 2.5 0.4 0.1 0.6
Uperodon globulosus (Günther, 1864 “1863”)
F Russelconda
NHH 62.12.29.4 (ex.
BMNH 1947.2.10.97) (ST)
67.4 19.2 13.0 7.7 3.6 8.8 2.7 9.7 13.9 10.2 3.3 2.8 14.5 18.0 29.1 21.7 26.5
F Pune SDBDU 2014.2673
71.7 18.7 12.7 7.4 3.7 8.7 2.6 9.6 15.3 11.3 3.4 2.9 20.5 32.0 27.4 23.3 28.1
F Odisha SDBDU 2005.4762
66.5 17.0 12.0 7.4 3.0 8.1 2.3 9.4 13.8 10.0 3.2 2.8 17.5 28.9 24.0 21.6 25.0
Average 68.5 18.3 12.6 7.5 3.4 8.5 2.5 9.6 14.3 10.5 3.3 2.8 17.5 26.3 26.8 22.2 26.5
Standard deviation 2.8 1.2 0.5 0.2 0.4 0.4 0.2 0.2 0.8 0.7 0.1 0.1 3.0 7.4 2.6 1.0 1.6
M Bihar ZSI A9575
55.9 15.9 10.6 6.3 2.8 6.7 1.5 7.7 11.9 7.8 2.5 2.5 11.3 13.7 21.1 15.6 20.3
M Dhanbad ZSI A4478
51.7 14.2 11.8 6.2 3.4 7.5 1.9 8.1 12.9 7.9 2.6 2.4 11.2 14.2 19.1 17.2 21.8
Average
53.8 15.1 11.2 6.3 3.1 7.1 1.7 7.9 12.4 7.9 2.6 2.5 11.3 14.0 20.1 16.4 21.1
Standard deviation 3.0 1.2 0.8 0.1 0.4 0.6 0.3 0.3 0.7 0.1 0.1 0.1 0.1 0.4 1.4 1.1 1.1
Uperodon taprobanicus (Parker, 1934)
F “Punduloya”
NHM 1905.3.25.138
(ex. 1947.2.10.62) (HT)
54.9 16.5 12.4 5.2 4.7 5.7 3.8 8.0 14.9 9.9 2.9 1.6 16.8 24.5 20.8 17.2 19.9
F Mannuthy SDBDU 2007.6364
53.7 16.0 12.1 5.3 4.5 5.2 3.3 7.8 14.2 10.3 2.9 1.5 16.3 26.5 21.1 17.8 20.2
Average 54.3 16.3 12.3 5.3 4.6 5.5 3.6 7.9 14.6 10.1 2.9 1.6 16.6 25.5 21.0 17.5 20.1
Standard deviation 0.8 0.4 0.2 0.1 0.1 0.4 0.4 0.1 0.5 0.3 0.0 0.1 0.4 1.4 0.2 0.4 0.2
M Bangalore SDBDU 2014.2567
42.5 12.7 11.0 4.5 3.7 4.1 2.9 5.7 11.6 9.3 2.5 1.3 12.5 19.0 15.2 12.9 16.7
M Odisha SDBDU 2002.3107
51.9 14.4 11.2 5.0 4.2 5.2 3.6 7.6 12.9 9.6 3.0 1.6 14.9 23.9 19.2 16.6 20.2
M Kalpetta SDBDU 2005.4732A
51.1 13.8 10.5 4.7 3.8 5.4 2.5 7.3 12.5 8.5 2.8 1.5 15.0 23.6 19.5 16.5 19.4
……continued on the next page
GARG ET AL.
50
·
Zootaxa 4384 (1) © 2018 Magnolia Press
TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
Average 48.5 13.6 10.9 4.7 3.9 4.9 3.0 6.9 12.3 9.1 2.8 1.5 14.1 22.2 18.0 15.3 18.8
Standard deviation 5.2 0.9 0.4 0.3 0.3 0.7 0.6 1.0 0.7 0.6 0.3 0.2 1.4 2.7 2.4 2.1 1.8
Uperodon anamalaiensis (Rao, 1937)
M Parambikulam ZSI/WGRC/V/A/956 (NT)
30.9 8.5 6.3 3.0 2.2 3.6 1.7 3.9 7.0 6.2 1.7 0.7 6.9 8.5 13.9 12.4 13.5
M Parambikulam SDBDU 2011.233A
36.7 10.2 8.2 4.3 2.3 3.9 1.6 4.3 8.4 6.3 2.3 0.9 7.1 10.4 14.4 12.9 14.6
M Parambikulam SDBDU 2011.234
35.2 8.9 6.7 3.4 2.3 3.6 1.7 4.5 8.3 6.7 2.1 0.8 7.5 9.3 14.5 13.1 14.2
M Parambikulam SDBDU 2011.548
28.7 8.8 6.5 3.0 2.3 2.7 1.3 3.4 6.2 6.2 1.7 0.5 6.5 8.3 11.2 9.3 10.4
M Methooty SDBDU 2012.2000
32.1 9.0 7.5 3.2 2.2 3.6 1.8 4.2 7.8 6.6 2.1 0.9 6.4 8.5 13.3 11.7 13.0
M Methooty SDBDU 2012.1994
32.2 9.8 7.6 3.6 2.2 3.5 1.8 4.2 9.1 6.7 2.0 1.0 8.0 10.3 13.1 12.1 13.7
M Methooty SDBDU 2012.1998
31.8 8.5 6.2 3.0 2.1 3.2 1.6 3.9 7.4 6.0 2.2 0.8 6.1 8.9 13.1 11.6 12.5
M Methooty SDBDU 2012.1996
30.6 8.3 6.9 3.2 2.1 4.1 1.7 3.8 7.7 6.0 1.9 0.8 7.1 8.5 13.0 11.2 12.7
M Methooty SDBDU 2012.1999
31.7 9.4 7.2 3.5 2.3 3.3 1.5 4.8 8.2 6.1 2.3 0.8 6.6 9.1 13.0 11.7 13.5
M Methooty SDBDU 2011.121
25.4 7.4 6.1 3.1 2.1 2.9 1.2 3.5 6.1 5.4 2.3 0.7 6.1 7.3 10.8 9.1 10.6
M Ponmudi SDBDU 2002.929
30.5 8.6 6.4 3.5 2.0 3.7 1.7 4.4 7.5 5.3 1.8 0.9 7.0 8.3 12.1 10.4 12.5
M Chathankod SDBDU 2002.3050
31.8 9.3 7.7 3.1 2.2 3.4 1.8 4.4 8.1 6.5 2.0 0.6 6.8 9.1 13.7 12.3 13.5
M Chathankod SDBDU 2002.3051
28.9 8.2 6.1 3.0 1.9 3.4 1.5 3.8 8.0 5.8 2.3 0.7 7.1 8.8 12.1 11.3 12.6
M Sakleshpur ZSI/WGRC/V/A/957 (NT)
31.7 9.1 7.1 3.1 2.1 3.4 1.3 4.2 7.4 5.9 2.2 0.9 6.2 8.6 13.8 11.8 13.3
Average 31.3 8.9 6.9 3.3 2.2 3.5 1.6 4.1 7.7 6.1 2.1 0.8 6.8 8.9 13.0 11.5 12.9
Standard deviation 2.7 0.7 0.7 0.4 0.1 0.4 0.2 0.4 0.8 0.4 0.2 0.1 0.5 0.8 1.1 1.2 1.2
F Chathankod SDBDU 2013.2302
38.6 10.6 8.6 3.7 2.3 4.4 1.9 4.7 8.3 7.7 2.4 0.9 9.2 11.9 15.4 14.2 15.7
F Chathankod SDBDU 2013.928
38.5 10.6 8.6 3.2 2.6 3.7 1.7 5.3 8.4 7.5 2.3 0.9 8.4 10.0 14.9 13.1 15.2
F Methooty SDBDU 2012.1995
37.4 9.8 8.2 3.9 2.9 4.3 1.7 4.9 9.2 6.4 2.2 1.0 8.2 10.7 15.0 12.8 15.7
F Chathankod SDBDU 2017.3507
35.3 8.9 6.8 3.3 2.8 3.3 1.7 4.7 7.6 6.3 1.9 0.9 6.9 10.0 14.9 12.9 14.3
Average 37.5 10.0 8.1 3.5 2.7 3.9 1.8 4.9 8.4 7.0 2.2 0.9 8.2 10.7 15.1 13.3 15.2
Standard deviation 1.5 0.8 0.9 0.3 0.3 0.5 0.1 0.3 0.7 0.7 0.2 0.1 1.0 0.9 0.2 0.6 0.7
Uperodon montanus (Jerdon, 1854 “1853”)
M Settukunnu SDBDU 2011.849
27.1 8.1 6.9 2.9 1.9 3.1 1.0 3.6 6.6 5.8 1.6 1.1 5.6 7.2 11.1 11.2 12.4
M Pandimotta SDBDU 2011.905
25.6 7.6 6.6 2.7 2.0 3.0 1.1 4.1 5.6 5.1 1.4 0.8 5.6 7.4 11.1 11.1 12.5
M Pandimotta SDBDU 2011.906
22.5 7.5 6.0 2.8 2.0 3.1 1.1 4.0 6.1 5.5 1.9 0.8 4.9 6.5 10.5 10.6 11.6
M Kesavapara SDBDU 2011.1197
23.6 8.0 6.3 2.6 1.6 2.6 1.3 3.4 5.8 5.1 2.0 0.7 5.5 6.6 10.1 10.2 11.7
M Kakkachi SDBDU 2003.40116
22.1 6.7 5.8 2.8 1.9 2.9 1.1 3.1 6.0 5.1 1.4 0.7 5.4 6.1 9.8 9.4 11.2
M Sengaltheri SDBDU 2002.24A
21.2 6.5 5.5 2.8 2.0 3.1 1.0 3.3 6.3 4.8 1.6 0.6 5.1 6.4 9.2 9.1 10.1
M Kakkachi SDBDU 2002.1925
22.2 7.1 5.6 2.7 1.8 3.2 1.2 3.1 6.0 4.1 1.7 0.6 5.7 6.1 9.5 9.2 10.1
……continued on the next page
Zootaxa 4384 (1) © 2018 Magnolia Press
·
51
TAXONOMIC REVIEW OF GENUS UPERODON
TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
M Upper Manalar SDBDU 2015.3001
25.2 7.2 6.4 2.8 2.1 2.7 1.2 3.9 6.9 4.9 1.4 0.7 5.4 6.7 10.8 10.7 11.7
M “Malabar” NHM 74.4.29.999
26.6 7.9 6.5 2.8 1.9 3.1 1.2 4.3 6.3 5.6 1.8 0.7 5.7 7.2 11.1 11.1 12.0
M “Malabar” NHM 74.4.29.1000
22.4 7.1 6.0 2.8 2.1 3.1 1.1 4.0 6.1 5.2 1.7 0.8 5.7 7.3 9.1 9.3 11.4
M “Malabar” NHM 74.4.29.1002
23.4 7.6 5.9 2.7 2.2 3.0 1.2 3.5 6.2 5.7 1.4 0.6 5.6 7.2 10.1 10.4 11.2
M Devikulam NHM 97.10.1.35
22.7 7.2 5.6 2.6 1.8 3.0 1.2 3.5 6.0 4.9 1.6 0.6 5.7 6.8 9.8 10.1 11.2
Average 23.7 7.4 6.1 2.8 1.9 3.0 1.1 3.7 6.2 5.2 1.6 0.7 5.5 6.8 10.2 10.2 11.4
Standard deviation 1.9 0.5 0.4 0.1 0.2 0.2 0.1 0.4 0.3 0.5 0.2 0.1 0.3 0.5 0.7 0.8 0.8
F “Wynad”
NHM 72.4.17.212
(ex. 1947.2.11.19) (HT)
28.3 10.1 7.0 3.3 2.1 3.1 1.6 3.9 7.5 5.9 2.1 0.8 6.8 8.7 12.3 12.4 13.9
F Kakkachi SDBDU 2003.40115
28.1 8.9 7.2 3.1 2.3 3.2 1.4 3.8 7.2 5.7 1.7 0.5 6.6 8.4 12.0 12.2 13.8
F Sengaltheri SDBDU 2002.24B
31.2 10.9 8.5 3.2 2.2 4.3 1.6 4.8 8.3 6.5 1.9 0.8 7.5 9.1 13.8 13.9 14.5
F Upper Manalar SDBDU 2015.3002
31.8 11.0 8.1 3.5 2.4 3.5 1.7 4.6 8.0 6.2 2.0 0.9 7.2 9.5 13.7 13.9 15.2
F Munnar SDBDU 2005.30
29.8 10.5 7.0 3.2 2.0 3.0 1.7 4.0 7.5 6.0 1.9 0.9 7.0 8.8 12.2 12.4 14.0
F Malabar NHM 74.4.29.993
33.6 11.1 8.1 3.5 2.6 3.6 1.7 4.8 8.6 7.1 2.6 1.1 7.9 9.8 13.9 13.9 14.9
F Devikulam NHM 97.10.1.34
30.2 10.0 7.1 3.3 2.5 3.1 1.6 4.7 7.9 6.9 2.5 1.0 7.8 9.1 13.1 13.3 14.9
F Devikulam NHM 97.10.1.37
30.0 10.0 7.0 3.2 2.1 3.2 1.5 4.5 8.1 6.9 2.6 1.0 7.8 9.2 13.0 13.1 14.3
Average 30.4 10.3 7.5 3.3 2.3 3.4 1.6 4.4 7.9 6.4 2.2 0.9 7.3 9.1 13.0 13.1 14.4
Standard deviation 1.8 0.7 0.6 0.1 0.2 0.4 0.1 0.4 0.5 0.5 0.4 0.2 0.5 0.4 0.8 0.7 0.5
Uperodon mormorata (Rao, 1937)
M
Shimoga SDBDU 2012.43
31.2 9.9 8.0 2.6 2.0 3.9 1.5 4.5 7.9 7.2 2.5 0.7 6.2 8.9 12.4 11.1 13.1
M
Bygoor SDBDU 2011.507
38.9 11.5 9.2 3.3 2.7 4.5 2.0 6.0 8.8 7.6 2.0 1.1 8.6 10.5 15.5 13.3 15.4
M
Castle Rock SDBDU 2011.835
37.2 11.4 9.5 2.2 2.9 3.7 1.7 5.3 8.7 7.3 2.3 1.1 7.9 10.2 15.5 12.6 15.7
M
Castle Rock SDBDU 2003.40197
35.9 10.5 7.1 3.2 2.9 3.9 1.9 5.1 9.3 7.3 1.9 0.8 7.3 10 14.9 12.5 14.3
M
Amboli SDBDU 2002.2018B
33.2 10.5 7.3 3.4 2.7 3.7 1.9 5.1 8.6 7.0 2.1 1.0 7.0 9.0 12.7 11.9 13.6
M
Amboli SDBDU 2002.2017
38.3 12.2 7.6 3.6 3.4 4.7 1.9 5.0 9.4 7.4 1.8 1.0 9.1 10.3 13.0 14 15.3
M
Amboli SDBDU 2002.2018A
39.3 12.9 9.6 3.0 3.4 4.7 2.1 6.2 9.7 9.5 2.2 1.4 8.7 10.5 15.2 14.1 15.5
M
Goa SDBDU 2015.3082
32.8 10.1 8.4 3.5 3.0 3.8 1.7 4.2 8.2 7.5 2.0 0.7 6.6 9.2 13.4 12.2 13.9
M
Netravali SDBDU 2016.3404
34.2 11.0 9.1 3.6 2.5 4.0 1.7 5.0 9.0 7.8 1.8 1.0 7.3 9.3 13.0 12.0 14.1
Average 35.7 11.1 8.4 3.2 2.8 4.1 1.8 5.2 8.8 7.6 2.1 1.0 7.6 9.8 14.0 12.6 14.5
Standard deviation 2.9 1.0 1.0 0.5 0.4 0.4 0.2 0.6 0.6 0.7 0.2 0.2 1.0 0.7 1.3 1.0 0.9
F Sakleshpur ZSI/WGRC/V/A/958 (NT)
32.4 9.6 8.8 3.1 2.6 3.0 1.6 4.4 8.6 6.8 1.5 0.6 6.5 9.1 13.5 11.1 12.9
F Samot SDBDU 2002.567
38.6 11.8 9.8 3.1 2.8 3.9 1.5 5.3 8.9 8.6 2.8 1.1 7.7 9.6 14.3 12.9 14.7
F Amboli SDBDU 2015.3085
42.5 13.2 9.8 4.1 2.7 4.6 2.4 5.2 10.4 8.3 2.2 1.2 8.9 11.5 16.4 14.5 16.7
……continued on the next page
GARG ET AL.
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·
Zootaxa 4384 (1) © 2018 Magnolia Press
TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
Average 37.8 11.5 9.5 3.4 2.7 3.8 1.8 5.0 9.3 7.9 2.2 1.0 7.7 10.1 14.7 12.8 14.8
Standard deviation 5.1 1.8 0.6 0.6 0.1 0.8 0.5 0.5 1.0 1.0 0.7 0.3 1.2 1.3 1.5 1.7 1.9
Uperodon nagaoi (Manamendra-Arachchi and Pethiyagoda, 2001)
M
Kanneliya FR WHT 2826 (HT)
25.6 6.8 6.0 3.3 2.0 2.9 1.3 4.2 6.5 4.7 1.6 0.9 6.2 6.5 11.5 11 11.2
M Kanneliya FR WHT 2867 (PT)
27.4 8.5 7.1 3.5 2.2 3.2 1.4 4.2 7.9 4.5 2.0 0.7 6.4 8.0 10.7 12.6 12.7
M
Kanneliya FR WHT 2865 (PT)
27.3 7.4 6.7 3.2 2.1 3.2 1.5 3.7 6.9 4.8 1.6 0.7 6.2 8.1 11.7 12.3 12.3
M
Kanneliya FR WHT 2868 (PT)
26.5 7.8 6.9 3.3 2.0 3.1 1.6 4.6 6.7 5.1 1.7 0.7 6.6 7.8 10.5 11.3 12.5
M Kanneliya FR WHT 2866 (PT)
27.7 7.4 6.5 3.4 2.1 3.3 1.7 4.0 6.7 4.3 1.6 0.6 7.5 7.6 10.8 12.3 12.4
M
Kanneliya FR WHT 2863 (PT)
27.2 7.6 6.8 3.3 2.1 3.4 1.3 4.0 6.7 4.5 1.7 0.7 6.5 7.4 10.4 12.1 12.2
M
Kanneliya FR WHT 2864 (PT)
26.7 8.2 7.2 3.5 2.1 2.8 1.7 3.9 6.9 4.4 1.6 0.7 6.2 7.8 11.0 11.7 11.8
M Hiyare DZ 1367
30.1 8.1 7.1 3.6 2.3 3.7 1.4 4.4 7.0 5.9 2.0 0.8 7.4 8.0 12.4 13.4 13.3
Average 27.3 7.7 6.8 3.4 2.1 3.2 1.5 4.1 6.9 4.8 1.7 0.7 6.6 7.7 11.1 12.1 12.3
Standard deviation 1.3 0.5 0.4 0.1 0.1 0.3 0.2 0.3 0.4 0.5 0.2 0.1 0.5 0.5 0.7 0.8 0.6
Uperodon obscurus (Günther, 1864 “1863”)
M
Morning side SL WHT 3510
25.4 7.9 6.9 2.8 2.1 3.2 1.5 4.0 6.6 5.7 1.3 0.4 5.6 6.5 11.1 10.9 11.1
M
Laxapana, SL WHT 7500
27.8 8.7 6.8 3.0 2.1 2.8 1.7 3.8 6.9 6.1 1.5 0.5 5.5 7.4 10.5 10.3 11.8
M
Pussellawa, SL WHT 07697
26.2 7.7 6.5 2.7 2.2 2.4 1.6 3.6 7.2 5.6 1.3 0.6 5.8 7.5 12.0 12.1 11.1
M
Udawattekelle SL WHT 00463A
22.8 6.9 6.5 2.3 2.1 2.7 1.7 3.5 6.3 4.4 1.4 0.4 5.1 6.2 9.9 10 9.9
M
Koskulana WHT 1194
24.5 7.6 6.7 2.5 1.8 3.0 1.2 3.7 6.3 5.7 1.5 0.6 5.4 7.1 11.5 10.6 10.4
M Paradeniya Campus DZ 1369
26.3 7.7 6.9 2.8 1.8 3.6 1.3 4.2 6.8 6.2 1.7 0.6 5.9 7.0 10.4 10.6 10.3
M
Paradeniya Campus DZ 1370
24.9 7.4 6.6 2.5 2.0 3.0 1.5 4.1 6.7 6.1 1.5 0.5 5.5 7.0 10 10.0 9.5
M
Paradeniya Campus DZ 1366
24.2 7.5 6.8 2.6 2.0 3.2 1.7 4.1 7 5.9 1.5 0.5 5.9 7.2 10.9 11.0 12.7
M Panwila, SL DZ 1408
24.8 7.6 6.8 2.8 2.0 3.3 1.7 3.7 6.5 5.5 1.8 0.5 5.5 7.2 10.2 9.9 10.9
M
Paradeniya Campus DZ 1365
27.8 8.0 7.2 3.1 2.1 3.6 1.6 4.3 6.9 5.9 1.8 0.7 6.4 7.3 10.9 11.1 11.1
M
Kandy: Sri Lanka NHM 1982.1291
24.5 7.7 6.8 2.7 2.0 3.1 1,4 3.9 7 5.8 1.9 0.6 6.3 6.7 10.2 10.6 10.8
M
Kandy: Sri Lanka NHM 1982.1292
25.9 7.7 5.9 2.7 2.1 3.0 1.3 3.9 6.6 5.2 2.4 0.4 4.6 7.6 10.1 10.6 12.0
M Babilapitiya Estate NHM 1973.3308
23.5 7.5 6.5 2.6 2.1 2.8 1.6 3.7 6.8 6.1 1.7 0.6 4.5 5.7 10.4 10.0 9.5
M
Babilapitiya Estate NHM 1973.3309
22.7 7.9 6.6 2.4 1.7 3.1 1.7 3.4 7.0 6.1 2.0 0.6 5.2 6.3 10.4 10.2 9.7
M Bogawantalowa NHM 1973.3010
21.9 8.6 7.3 2.2 1.6 3.2 1.2 3.8 7.0 6.4 1.9 0.7 5.5 7.0 10.6 10.5 10.4
M
Krunegala: Ceylon NHM 1973.3012
30.1 8.9 7.8 3.3 2.6 2.8 1.9 4.5 7.8 6.7 1.3 0.6 7.6 8.1 12.4 12.1 13.4
M
Krunegala: Ceylon NHM 1973.3013
30.1 8.7 7.6 3.3 2.5 3.1 1.5 4.4 8.0 7.1 1.7 0.6 7.2 7.3 12.7 12.7 13.8
……continued on the next page
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·
53
TAXONOMIC REVIEW OF GENUS UPERODON
TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
Average 25.5 7.9 6.8 2.7 2.0 3.1 1.5 3.9 6.9 5.9 1.7 0.6 5.7 7.0 10.8 10.8 11.1
Standard deviation 2.4 0.5 0.4 0.3 0.3 0.3 0.2 0.3 0.5 0.6 0.3 0.1 0.8 0.6 0.8 0.8 1.3
F
“Ceylon”
NHM 58.11.28.42 (ex.
1947.2.11.20) (HT)
32.2 9.3 8.8 4.2 3.9 3.3 1.4 4.9 8.6 6.5 2.4 0.7 8.8 9.5 13.5 13.3 14.9
F
Kandy: Sri Lanka NHM 1982.1290
32.3 8.5 7.4 3.6 2.1 3.4 1.7 4.5 7.7 6 1.9 0.7 6.1 9.3 11.7 12.7 13.3
F
Gammaduwa NHM 1955.1.10.77
30.1 8.9 7.7 3.5 2.1 3.6 2.1 4.7 8.5 6.5 1.9 0.7 7.1 6.9 12.6 13.3 12.6
F
Morning side, SL WHT 05982
30.9 9.9 8.9 3.4 2.1 4 2 4.1 8.6 7.2 1.9 0.7 7.1 8.6 14.3 13.1 14.8
F
Morning side, SL WHT 5980
37.1 10.4 9.1 3.9 3.1 4 1.9 4.5 8.7 7.4 2.2 1.0 8.5 9.7 15.4 14.9 15.6
Average 32.5 9.4 8.4 3.7 2.7 3.7 1.8 4.5 8.4 6.7 2.1 0.8 7.5 8.8 13.5 13.5 14.2
Standard deviation 2.7 0.8 0.8 0.3 0.8 0.3 0.3 0.3 0.4 0.6 0.2 0.1 1.1 1.1 1.4 0.8 1.2
Uperodon palmatus (Parker, 1934)
M
“Newera Ellya”
NHM 90.11.8.63
(ex. 1947.2.11.22)
27.9 8.5 7 2.5 2.4 3.1 1.7 4.2 7.5 6.4 1.7 0.4 7.4 7.9 11.2 11.5 12.4
M
Nuwara Eliya DZ 1371
28.9 7.9 6.9 2.8 2.7 3.1 1.6 4.1 7.4 6.9 1.3 0.8 7.1 8.5 11.4 11.3 13.2
M
Nuwara Eliya DZ 1331
30.7 8.8 7.4 2.7 2.8 2.8 1.9 4.5 8.0 7.5 1.7 1.0 7.6 9.0 12.1 12.2 14.5
M
Nuwara Eliya DZ 1372
30.5 8.7 7.6 2.7 2.8 3.4 2 4.4 8.4 7.2 1.8 0.6 7.7 8.9 12.4 12.4 14.1
Average 29.5 8.5 7.2 2.7 2.7 3.1 1.8 4.3 7.8 7.0 1.6 0.7 7.5 8.6 11.8 11.9 13.6
Standard deviation 1.3 0.4 0.3 0.1 0.3 0.2 0.2 0.2 0.5 0.5 0.2 0.3 0.3 0.5 0.6 0.5 0.9
F “Newera Ellya”
NHM 90.11.8.62
(ex. 1947.2.11.21) (HT)
34.6 11.8 9.1 3.6 3.5 3.8 1.9 4.8 9.4 5.7 1.9 0.5 7.4 10.4 14.8 14.6 15.4
F
Diyagama estate WHT 5424
35.3 10.1 8.5 3.3 3.4 3.5 2.2 4.5 8.3 6.8 1.9 0.8 8.2 9.7 13.5 13.6 15.6
Average 35.0 11.0 8.8 3.5 3.5 3.7 2.1 4.7 8.9 6.3 1.9 0.7 7.8 10.1 14.2 14.1 15.5
Standard deviation 0.5 1.2 0.4 0.2 0.1 0.2 0.2 0.2 0.8 0.8 0 0.2 0.6 0.5 0.9 0.7 0.1
Uperodon triangularis (Günther, 1876)
M
“Malabar” NHM 74.4.29.897
(ex. 1947.2.11.15) (ST)
31.6 9.6 7.7 3.7 2.5 3.4 1.9 3.6 8.4 5.7 2.1 0.9 7.1 8.7 11.4 10.9 12.1
M
“Malabar” NMH 74.4.29.892
(ex. 1947.2.11.10)
32.3 10.7 8.3 3.3 2.9 2.1 1.9 4.9 9.4 6.8 1.9 1.7 7.7 9.5 13.1 12.0 13.8
M
“Malabar” NHM 74.4.29.894
(ex. 1947.2.11.12)
32.2 2.0 10 3.8 2.8 3.5 2.0 5 9.4 8 1.7 0.8 7.3 9.8 11.8 12.9 14.8
M
“Malabar” NHM 74.4.29.895
(ex. 1947.2.11.13)
32.7 9.3 9.4 3.6 2.3 3.9 1.8 4.7 8.7 8.1 1.9 1.1 6.8 8.6 11.2 11.3 13.3
M
“Malabar” NHM 74.4.29.896
(ex. 1947.2.11.14)
31.9 10 7.7 3.6 2.7 3.4 2.0 4.4 9.2 6.1 2.0 0.7 5.6 8.9 12.4 11.5 13.3
……continued on the next page
GARG ET AL.
54
·
Zootaxa 4384 (1) © 2018 Magnolia Press
TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
M
“Malabar” NHM 74.4.29.898
(ex. 1947.2.11.16)
30.5 9.1 7.2 3.6 2.5 2.9 1.7 4.1 8.9 5.7 2.0 1.0 6.5 9 11.4 11.6 11.5
M
“Malabar” NHM 74.4.29.899
(ex. 1947.2.11.17)
29.5 9.3 7.4 3.0 2.6 4.1 1.7 4.0 8.4 6.1 1.9 0.9 6.9 8.9 12.2 11.7 13.7
M
“Malabar”
NHM 74.4.29.900
(ex. 1947.2.11.18)
25.6 7.8 5.8 3.0 2.4 2.5 1.5 3.6 7.2 6.5 1.6 0.9 5.8 6.9 8.8 10.0 10.5
M
Nilgheries NHM 87.8.26.2
29.4 8.9 6.9 3.6 2.3 2.7 1.7 3.9 8.1 6 1.6 0.9 6.1 8.2 11.7 10.3 12.6
M
Bygoor SDBDU 2011.504
32.7 10.3 8.2 3.5 2.8 3.6 2.0 4.3 8.7 6.8 1.6 0.9 6.5 8.8 12.4 11.9 13.0
M
Kalpetta SDBDU 2012.2038
27.2 8.9 7.8 2.9 2.4 3.5 1.4 4.2 7.9 6.6 1.4 0.9 6.4 8.2 11.1 11.0 12.3
M
Bygoor SDBDU 2011.503
36.0 11.1 8.6 3.4 2.4 3.9 2.1 5.1 8.9 6.9 1.9 0.9 8.0 10.3 13.2 12.6 14.6
M
Bygoor SDBDU 2010.332
35.2 10.7 7.3 3.6 2.7 3.7 2.0 5.3 9.6 6.5 2.2 0.8 8.1 10.1 13.7 12.7 13.9
M
Kalpetta SDBDU 2008.409
28.8 9.6 7.3 3.3 2.2 3.1 2.1 4.9 8.4 6.4 1.8 0.9 6.9 8.6 12.9 11.0 12.5
M
Bygoor SDBDU 2011.502
34.2 10.1 7.6 3.8 2.4 3.9 2.1 4.9 8.8 6.3 1.8 0.9 7.4 9.7 12.6 12.7 13.4
M
Bygoor SDBDU 2011.506
36.8 10.7 7.9 3.6 2.9 3.1 2.2 4.2 9.1 6.7 2.1 0.9 6.4 9.3 13.6 12.9 13.1
M
Mudigere ZSI/WGRC/V/A/959 (NT)
29.7 9.4 6.7 2.7 2.2 3.0 1.9 4.0 7.9 6.0 1.6 0.9 6.0 8.8 11.0 10.1 12.4
M
Kalpetta SDBDU 2007.6358
28.9 9.3 7.1 2.9 2.5 3.1 2.1 3.7 8.5 6.0 1.8 0.9 6.2 8.1 11.1 11.4 12.6
M
Suganthagiri SDBDU 2015.2857
31.0 10.7 8.2 3.5 2.6 3.2 2.1 4.5 8.8 6.3 1.8 0.9 7.0 9.1 11.8 11.8 13.5
Average 31.4 9.3 7.7 3.4 2.5 3.3 1.9 4.4 8.6 6.5 1.8 0.9 6.8 8.9 12.0 11.6 13.0
Standard deviation 2.9 2.0 0.9 0.3 0.2 0.5 0.2 0.5 0.6 0.7 0.2 0.2 0.7 0.8 1.2 0.9 1.0
F
Ooty SDBDU 2007.6400
40.8 11.5 8.4 3.7 2.7 4.6 2 4.9 9.5 8.1 2.1 1.1 8.1 10.8 13.8 14.1 14.2
F
Kalpetta SDBDU 2003.40240
33.0 9.4 8 3.5 2.5 3.7 2 4.1 8.5 6.4 2 1.2 7.3 9 12.9 11.5 13.4
F
“Malabar”
NHM 74.4.29.891
(ex. 1947.2.11.9)
35.8 11.1 9.3 3.3 2.9 3.5 1.7 5.7 9.9 7.1 1.6 1 7.8 9.8 13.9 12.9 14.2
F
“Malabar”
NHM 74.4.29.893
(ex. 1947.2.11.11)
41.1 12.6 9.2 3.4 3 4.9 1.6 5.2 10.1 8.9 1.9 0.9 7.9 11.4 15.4 14.2 15.7
Average 37.7 11.2 8.7 3.5 2.8 4.2 1.8 5.0 9.5 7.6 1.9 1.1 7.8 10.3 14.0 13.2 14.4
Standard deviation 4.0 1.3 0.6 0.2 0.2 0.7 0.2 0.7 0.7 1.1 0.2 0.1 0.3 1.1 1.0 1.3 1.0
Uperodon variegatus (Stoliczka, 1872)
M
Yellagherry Hills
NHM 72.4.29.1351
(ex. 1947.2.11.23)
26.8 7.4 6.2 3.1 2.4 3.1 1.4 3.9 6.6 5.9 1.6 0.8 5.8 7.5 10.6 9.8 11.5
M
Yellagherry Hills
NHM 72.4.29.1352
(ex. 1947.2.11.24)
25.6 7.0 6.3 3.2 2.6 2.9 1.3 3.4 6.5 5.3 1.6 0.8 5.6 7.1 10.2 9.9 11.0
M
Bangalore
NHM 1925.9.18.1
(ex. 1947.2.11.35)
30.1 8.8 7.3 3.4 2.4 3.3 1.3 4.1 6.9 6.2 1.7 0.6 5.6 7.4 10.8 9.9 10.8
……continued on the next page
Zootaxa 4384 (1) © 2018 Magnolia Press
·
55
TAXONOMIC REVIEW OF GENUS UPERODON
TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
M
Koruturu SDBDU 2007.4994
32.1 8.9 6.5 3.3 2.6 3.3 1.4 4.2 7.5 6.3 1.6 0.8 5.7 8.8 11.1 9.5 12.1
M
Karamana SDBDU 2007.6348
28.8 8.4 7.2 3.0 2.3 4.0 1.5 4.3 7.4 7.0 1.8 0.9 6.2 8.5 10.5 9.8 12.4
M
Palakkad SDBDU 2002.524
29.8 8.5 6.8 3.2 2.4 3.6 1.6 4.2 7.2 6.2 1.3 0.7 6.0 7.1 11.0 9.4 10.4
M
Palakkad SDBDU 2002.523
27.7 7.1 6.6 3.0 2.4 3.2 1.3 3.5 6.5 5.9 1.5 0.9 4.9 7.0 9.6 8.8 10.2
M
Meladoor SDBDU 2003.40222
27.5 7.9 6.7 3.2 2.6 2.9 1.3 3.5 6.7 5.0 1.0 0.9 5.5 7.5 9.5 8.9 10.3
M
Chengalpattu SDBDU 2005.4727
28.9 7.8 6.7 3.4 2.2 3.4 1.5 3.8 7.0 6.0 1.7 0.7 5.6 7.8 10.9 9.8 12.6
M
Utkal Uni. Campus NHM 1984.132
27.4 6.9 6.3 3.3 2.1 3.2 1.4 3.7 6.7 5.8 1.5 0.8 5.5 7.3 9.5 9.1 10.3
M
Utkal Uni. Campus NHM 1984.133
28.8 7.5 6.3 3.5 2.5 3.4 1.5 3.9 6.8 5.2 1.6 0.8 5.6 7.2 9.8 9.6 10.4
M
RR Lab area NHM 1984.135
26.9 7.8 6.3 3.4 2.6 3.2 1.4 3.6 6.8 4.8 1.5 0.7 5.1 7.1 9.9 9.0 10.1
Average 28.4 7.8 6.6 3.3 2.4 3.3 1.4 3.8 6.9 5.8 1.5 0.9 5.6 7.5 10.3 9.5 11.0
Standard deviation 1.8 0.7 0.4 0.2 0.2 0.3 0.2 0.3 0.3 0.6 0.2 0.2 0.3 0.6 0.6 0.4 0.9
F
Ellore ZSI 2761 (LT)
29.8 8.7 7.1 3.2 2.7 3.6 1.4 4.3 8.3 6.2 1.9 0.9 5.0 7.8 11.3 9.9 12.7
F
Gadavery Valey NHM 1908.1.29.26
30.6 8.9 7.4 3.3 2.6 3.9 1.5 4.0 8.2 6.1 1.9 0.9 6.1 8.0 11.3 10.1 12.9
F
Gadavere NHM 77.8.6.12
34.2 9.6 7.4 3.8 2.9 4.1 1.6 4.6 8.1 6.9 1.8 0.8 6.3 8.2 12.1 9.9 12.8
F
Bangalore
NHM1925.9.18.1
(ex. 1947.2.11.34)
35.6 9.9 7.7 3.8 2.8 3.8 1.8 4.7 8.3 7.3 1.9 0.8 6.4 8.3 12.9 10.5 13.1
F
Chengalpattu
SDBDU 2005.4726
33.8 9.6 7.1 3.4 2.9 3.8 1.7 4.5 8.0 7.0 1.8 0.9 6.3 8.6 12.1 10.4 13.1
F
Karamana
SDBDU 2007.6347
32.9 9.5 7.4 3.4 2.5 4.0 1.6 4.5 8.1 7.2 1.7 0.9 5.8 8.9 12.1 10.6 13.9
F
RR Lab area
NHM 1984.134
30.7 8.0 7.1 3.3 2.6 3.6 1.6 4.2 7.7 6.2 1.9 0.6 6.0 8.4 11.9 9.9 12.7
Average 32.5 9.2 7.3 3.5 2.7 3.8 1.7 4.4 8.1 6.7 1.9 0.8 6.0 8.3 12.0 10.2 13.0
Standard deviation 2.2 0.7 0.2 0.2 0.2 0.2 0.2 0.2 0.2 0.5 0.1 0.1 0.5 0.4 0.6 0.3 0.4
Uperodon rohani sp. nov.
M
Mihintale DZ 1449 (HT)
26.0 6.7 6.2 2.6 2.5 2.9 1.6 3.5 5.5 5.2 1.4 0.7 5.2 7.2 9.2 9.4 11.1
M
Mihintale
DZ 1448 (PT)
28.0 7.4 7.2 2.7 2.6 3.0 1.4 3.4 5.9 6.4 1.9 0.9 6.0 7.7 9.4 9.9 12.4
M
Mihintale
DZ 1450 (PT)
30.7 7.8 7.1 2.8 2.4 3.6 1.5 3.6 6.2 5.8 1.9 0.9 5.6 8.0 10.0 10.4 12.1
M
Mihintale
DZ 1374 (PT)
26.8 7.3 7.0 2.8 2.6 3.2 1.5 3.5 5.4 5.3 1.7 1.1 5.7 7.7 9.9 10.3 11.8
M
Bundala NP DZ 1336 (PT)
26.4 7.4 7.0 2.6 2.4 2.9 1.5 3.9 6.1 5.8 1.8 0.5 5.7 7.5 9.3 9.6 11.3
M
Makandura DZ 1520 (PT)
26.2 6.6 6.3 2.6 2.5 3.1 1.2 3.8 5.4 5.2 1.6 0.8 4.8 7.2 9.9 10.2 11.1
M
Makandura DZ 1521 (PT)
28.9 7.7 7.2 2.8 2.4 2.8 1.5 3.7 5.7 5.8 1.9 0.9 5.8 7.7 11.0 10.9 12.2
M
Ampara DZ 1486 (PT)
33.1 7.2 7.0 2.4 2.3 3.3 1.4 3.5 5.8 5.3 1.8 0.9 6.0 8.0 10.0 10.4 12.6
M
Ceylon NHM 1931.2.1.21
28.6 7.5 7.0 2.6 2.4 3.2 1.2 3.3 6.6 5.7 1.8 0.9 6.1 7.9 10.1 10.7 11.6
M
Kurunegala: Ceylon NHM 1973.3015
27.6 7.2 6.8 2.8 2.5 2.9 1.2 3.6 6.3 5.5 1.7 0.8 5.9 7.2 9.7 10.3 10.2
……continued on the next page
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TABLE 4. (Continued)
Sex Locality Museum Number
SVL
HW HL SL EL IUE UEW IFE IBE MN EN NS FAL
HAL
TL
SHL FOL
M
Badulla, Sri Lanka NMH 1976.264
26.3 7.0 6.7 2.7 2.5 3.0 1.3 3.7 6.3 5.9 1.8 0.6 6.4 7.5 10.0 11.0 11.6
M
Badulla, Sri Lanka NHM 1976.265
27.4 7.1 6.9 2.8 2.5 2.9 1.2 3.7 6.1 5.8 1.9 0.6 5.6 7.1 9.9 10.5 11.2
M
Nachchiyagama WHT 5320
30.1 8.0 7.7 2.6 2.4 2.9 1.1 3.4 6.5 5.8 1.6 0.6 5.6 7.4 10.2 10.6 12.5
M
Sigiriya, Near RH WHT 01188
28.0 7.8 7.3 2.6 2.6 3.0 1.2 3.5 6.6 5.9 1.7 0.5 5.6 7.6 10.0 10.5 10.9
M
Polonnaruwa WHT 1749
26.3 6.6 6.4 2.9 2.6 2.7 1.2 3.5 6.2 5.2 1.5 0.6 5.7 7.5 9.4 9.8 10.1
M
Anuradhapura WHT 01189
27.7 7.6 7.1 2.7 2.3 2.9 1.3 3.5 6.3 5.5 1.5 0.6 5.4 7.1 9.6 9.9 11.2
Average 28.0 7.3 6.9 2.7 2.5 3.0 1.3 3.6 6.1 5.6 1.7 0.7 5.7 7.5 9.9 10.3 11.5
Standard deviation 1.9 0.4 0.4 0.1 0.1 0.2 0.2 0.2 0.4 0.3 0.2 0.2 0.4 0.3 0.4 0.5 0.8
F
Kurunegala: Ceylon NHM 1973.3014
28.7 7.4 6.8 2.6 2.7 3.4 1.3 3.5 6.3 5.1 2.0 0.6 5.9 7.8 10.3 11.4 11.0
F
Palatupana, Kirinda WHT 01191
30.0 7.6 6.8 2.4 2.6 3.5 1.5 3.4 6.7 5.4 2.1 0.5 6.1 7.9 9.9 10.9 11.2
F
Dambulla DZ 1129 (PT)
34.5 9.1 8.8 2.5 2.6 3.7 1.5 4.5 6.8 6.4 2.0 0.6 7.8 10.0 11.9 12.9 14.4
F
Dambulla DZ 1130 (PT)
33.3 8.6 7.9 2.7 2.6 3.6 1.3 4.2 6.6 6.4 2.0 0.5 6.9 9.0 11.0 12.2 13.8
F
Dambulla DZ 1132 (PT)
34.2 8.5 7.2 2.8 2.7 3.6 1.4 4.5 7.1 6.2 2.1 0.6 6.6 8.4 12.0 13.1 13.9
F
Dambulla DZ 1131 (PT)
31.9 8.3 7.3 2.5 2.5 3.4 1.3 4.1 6.7 6.4 2.1 0.6 6.9 8.8 11.2 12.4 13.3
F
Makandura DZ 1425 (PT)
31.1 8.4 7.8 2.6 2.7 3.5 1.6 4.1 6.8 6.9 2.3 0.9 6.3 7.3 10.8 11.8 12.1
F
Ampara DZ 1485 (PT)
31.3 8.3 7.4 2.4 2.9 3.6 1.4 4.5 7.0 6.7 2.3 0.9 6.5 8.8 11.1 12.1 12.7
Average
31.9 8.3 7.5 2.6 2.7 3.5 1.4 4.1 6.8 6.2 2.0 0.7 6.6 8.5 11.0 12.1 12.8
Standard deviation
2.0 0.5 0.7 0.1 0.1 0.1 0.1 0.4 0.2 0.6 0.2 0.2 0.6 0.8 0.7 0.7 1.3
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U. anamalaiensis (present study), U. variegatus (Rao 1918), U. systoma (Kirtisinghe 1957), and U. globulosus
(present study) is terminal, whereas U. palmatus has a dorsoterminal mouth (Kirtisinghe 1958) and U. nagaoi has a
funnel-shaped mouth (Manamendra-Arachchi & Pethiyagoda 2001). Tail morphology does not differ substantially
among these species and almost all are known to have rounded tail tips as studied in U. palmatus (Kirtisinghe 1958),
U. triangularis, U. variegatus (Rao 1918), U. obscurus (Bowatte & Meegaskumbura 2012) and U. nagaoi
(Manamendra-Arachchi & Pethiyagoda 2001). Location of spiracle tends to show variation among the studied
species, with spiracles in U. variegatus, U. triangularis (Rao 1918), U. anamalaiensis and U. globulosus (present
study) located ventral to the gut; spiracle openings in U. palmatus (Kirtisinghe 1958) and U. montanus (Parker
1934) near tubular vent; and the opening slightly beyond posterior end of body in U. obscurus and U. nagaoi
(Manamendra-Arachchi & Pethiyagoda 2001; Bowatte & Meegaskumbura 2012) (Fig. 16).
FIGURE 16. Tadpoles of Uperodon globulosus, U. anamalaiensis, U. nagaoi, U. obscurus and U. triangularis. A–C.
Uperodon globulosus in life, Gosner stage 32: A. Lateral view; B. dorsal view; C. ventral view; D–F. Uperodon anamalaiensis
in preservation, Gosner stage 34; D. Lateral view; E. dorsal view; F. ventral view; G–H. Uperodon nagaoi in preservation,
Gosner stage 37: G. Lateral view; H. lateral view of head; I–J. Uperodon obscurus in life: I. Lateral view, Gosner stage 30; J.
Lateral view of head, Gosner stage 38; K–M. Uperodon triangularis in life, Gosner stage 35: K. Lateral view; L. ventral view;
M. dorsal view. (Images: SD Biju, I–J & G–H. Madhava Meegaskumbura).
Discussion
The genus Uperodon is morphologically one of the most diverse anuran groups of the Western Ghats–Sri Lanka
biodiversity hotspot. Its members exhibit immense variations with respect to body shape (slender vs. robust and
globular), body size (21–72 mm), lifestyle (burrowing vs. arboreal), reproductive behaviour (arboreal vs.
terrestrial), and tadpole morphology. Species within the group also form three highly divergent evolutionary
lineages, which along with striking phenotypic differences could be the major reason for acceptance of the previous
generic-level classification despite the advent of molecular systematics (e.g., Van Bocxlaer et al. 2006; Pyron &
Wiens 2011). Recently, based on molecular evidence and earlier observations of Ramaswami (1936), Peloso et al.
(2016) considered the synonymy of Ramanella (sensu stricto) and Kaloula pulchra taprobanica with Uperodon
(sensu stricto) as the most parsimonious option for stabilizing the taxonomy of this group. Our taxonomic revision
using an integrative approach, which includes all members of this morphologically variable group, will aid future
studies on Uperodon (sensu lato) frogs. Furthermore, this study delineates the geographical ranges of all known
Uperodon species, clarifying their patterns of distribution. Of the 12 nominal species, eight are restricted to the
Western Ghats–Sri Lanka biodiversity hotspot (U. anamalaiensis, U. montanus, U. mormorata, U. nagaoi, U.
obscurus, U. palmatus, U. rohani sp. nov. and U. triangularis), accounting for 67% endemism at the species level.
Among these species, four each are endemic to the Western Ghats and Sri Lanka, respectively. The remaining
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species—U. globulosus, U. systoma, U. taprobanicus and U. variegatus, are more widely distributed outside the
Western Ghats–Sri Lanka hotspot region, with distributions of the former three extending up to adjoining countries
such as Bangladesh, Bhutan, Nepal and Pakistan (Fig. 2A; Tables 1, S1).
Although knowledge on the natural history of Uperodon species is sparse, some interesting insights can be
gained regarding the reproductive requirements of these frogs. Several species are known to be arboreal and may
therefore possess dilated finger and toe tips as an adaptation for climbing (Blackburn et al. 2013). The non-dilated
digit tips in some members (U. globulosus and U. systoma), however, are suggestive of terrestrial lifestyles.
Further, only a few species breed inside tree holes; most others are opportunistic, breeding in ground puddles. It
would thus be interesting to investigate the various reproductive modes in Uperodon frogs, how they have evolved
to utilize different niches, and the possible factors responsible for diversification. In the future, based on
comprehensive phylogenies, morphological and ecological traits can be analyzed to address various evolutionary
questions. Such studies could also shed light on the adaptability of Uperodon frogs, especially in the absence of
suitable habitats such as tree holes, which is likely to be their most immediate threat with increasing habitat
destruction and degradation in the Western Ghats–Sri Lanka biodiversity hotspot.
Acknowledgements
We are grateful to Rohan Pethiyagoda and Mark D Scherz for their valuable comments and suggestions that were
helpful in improving the earlier draft of this paper; Robin Suyesh for help in the acoustic and statistical analyses;
Mahadev Mathpati for support in compilation and preparation of Table S1; K Jayaram for field and logistic support
to SDB and SG in Tamil Nadu; Aravind Madhyasthya, KV Gururaja and KP Dinesh for field support to SDB and
SG in Karnataka; Past members of the Systematics Lab (University of Delhi) for field support; Romulus Whitaker
and Janaki Lenin for logistic support to SDB and SG during fieldwork at Mamallapuram (Tamil Nadu) and for
permission to use their photographs of Uperodon variegatus; L. Shyamal for permission to use the image of
Uperodon variegatus; Mark-Oliver Rödel and Frank Tillack (ZMB, Berlin) for providing access to specimens in
their care and their hospitality to SDB and SG; Kailash Chandra and G Maheswaran (ZSI, Kolkata), PM Sureshan
(ZSI, Calicut) and Rahul Khot (BNHS, Mumbai), Manori Goonatilake and Chandrika Munasinghe (NHM,
Colombo) for access to specimens in their care and museum support to SDB and SG; Barry Clarke and David
Gower (NHM, London) for access to specimens in their care and museum support to SDB; the state forest
departments in India (Kerala, Karnataka, Tamil Nadu, Maharashtra) for study permits and field support; the
Department of Wildlife Conservation and Forest Department of Sri Lanka for research permits. We also thank the
editor (Miguel Vences) and the two reviewers (Mark D Scherz and KV Gururaja) for valuable suggestions. This
study was partially supported by the following grants to SDB: Critical Ecosystem Partnership Fund, Conservation
International, USA (Project 55918/ 2009); University of Delhi Research and Development Grants 2013–14 and
2015–16; DST Purse Grant Phase I (2009–2013) and Phase II (2015–16), Department of Science and Technology,
Ministry of Science and Technology, Government of India. SG received research fellowships from University of
Delhi, University Teaching Assistantship Scheme (2010/56562), and Council for Scientific and Industrial Research
(CSIR No. 9/45(1381)/2015-EMR-I). GS received a Rufford Small Grant (No. 192721) and MM received a grant
from National Research Council of Sri Lanka (No. 11-124).
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TAXONOMIC REVIEW OF GENUS UPERODON
Supplementary Material
An integrative taxonomic review of the South Asian microhylid genus Uperodon
Sonali Garg, Gayani Senevirathne, Nayana Wijayathilaka, Samadhan Phuge, Kaushik Deuti, Kelum Manamendra-
Arachchi, Madhava Meegaskumbura and SD Biju
TABLE S1. Distribution of Uperodon species. 1. Genetically and morphologically studied collections from the present
study, 2. Other morphologically studied collections and museum specimens in the present study, 3. Previously published
literature and other available records (including reports that require further confirmation). Asterisk symbol denotes the
type locality.
State /
Province
Locality Reference
Uperodon anamalaiensis
INDIA
Tamil Nadu “Base of Anamalai Hills, Coimbatore District”* Rao 1937
Kerala Parambikulam TR, Palakkad district Vijaykumar et al. 2011
2
;
Present Study
1
Munnar, Idukki district Harpalani et al. 2015
3
Chathankod, Thiruvananthapuram district Present Study
1
Kallar, Ponmudi, Thiruvananthapuram district Present Study
1
Makki, Thiruvananthapuram district Present Study
2
Methooty, Idukki district Present Study
1
Karnataka Kempholey, Hassan district Present Study
1
Sakleshpur, Hassan district Present Study
1
Uperodon globulosus
INDIA
Kerala Manimala River, Erumely, near Koratty bridge, Kottayam
district
George & Alex 1995
3
; Dutta 1997
3
Karnataka Khanapur, Belgaum district Abdulali 1962
3
“Mysore” Dutta 1997
3
Maharashtra “Kanheri Caves (ca. 1,300 ft.), near Borivli, Salsette Island,
Bombay”
Abdulali & Daniel 1954
3
Kanheri Caves, Mumbai Bhaduri & Daniel 1956
3
“Bombay” Daniel 1963a
3
Balewadi, Pune district Padhye et al. 2002
3
Pashan, Pune district Padhye et al. 2002
3
Navegaon National Park, Gondia district Bhandarkar et al. 2012
3
Supegaon, Phansad Wildlife Santuary Katwate et al. 2013
3
“Satara Tehsil” Prasad et al. 2013
3
Radhanagari Wildlife Santuary Yadav & Yankanchi 2014
3
Shivaji University, Kolhapur district Yadav et al. 2014
3
Pune, Pune district Present Study
1
Tamhini Ghat, Pune district Present Study
2
Andhra
Pradesh
Sundipenta/Sikharam, Nallamalai Hills Rao et al. 2005
3
...Continued on next page
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TABLE S1. (Continued)
State /
Province
Locality Reference
Mannanur – Farahabad, Nallamalai Hills Srinivasulu et al. 2006
3
;
Srinivasulu et al. 2007
3
Mukhadwaram – Srisailam, Nallamalai Hills Srinivasulu et al. 2007
3
“Nallamala hills” Srinivasulu & Das 2008
3
Seshachalam Biosphere Reserve Guptha et al. 2012
3
Thummalapalle uranium mining area, Thummalapalle, YSR
Kadapa district
Reddy et al. 2013
3
Gujarat Ahwa, Dang district Daniel 1963a
3
Waghai, Dang district Daniel 1963a
3
Shoolpaneshwar WLS, Bharuch district Naik & Vinod 1993
3
;
Sarkar & Ray 2000
3
Surat Dangs, Narmada district Dutta 1997
3
Vansda National Park Vyas 2004
3
Madhya
Pradesh
“Jabalpur” Dutta 1992
3
; Chandra & Gajbe 2005
3
“Berar” Dutta 1997
3
Kachpura Ward, Jabalpur Chandra & Ray 2007
3
Tropical Forest Research Institute campus, Jabalpur Paunikar 2017
3
Chhattisgarh “Bastar” Sarkar 1993
3
; Chanda 1995
3
; Chandra & Gajbe
2005
3
; Dutta 2015
3
Indravati Tiger Reserve Chanda 1995
3
;
Chandra & Gajbe 2005
3
Odisha “Russelconda”* Günther, 1864 “1863”
2
Sambalpur University Campus, Sambalpur district Sarkar 1993
3
“Dhenkanal” Dutta 1997
3
Bedkot, Karlapat WLS Deuti & Raha 2010
3
Karlapat, Karlapat WLS Deuti & Raha 2010
3
;
Deuti et al. 2014
“Mundguda” (= Mundraguda), Karlapat WLS Deuti & Raha 2010
3
“Rayagada” Deuti et al. 2014
3
North Orissa University Campus, Mayurbhanj district Boruah et al. 2016
3
Jharkhand Tundi, Dhanbad district Sarkar 1991
3
West Bengal “Calcutta” Bhaduri 1945
3
“Jalpaiguri” Bhaduri & Basu 1956
3
; Dutta 1997
3
24 Pargonas district Sarkar et al. 1992
3
Haora (=Howrah) district Sarkar et al. 1992
3
Hugli (=Hooghly)
district Sarkar et al. 1992
3
Dhupguri, Jalpaiguri district Sarkar et al. 1992
3
Sulkapara, Jalpaiguri district Sarkar et al. 1992
3
Mandalpushkarini, Medinipur district Sarkar et al. 1992
3
“40 miles north of Calcutta: Camp Kanchrapara” Dutta 1997
3
Midnapore district Pratihar & Deuti 2011
3
...Continued on next page
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TAXONOMIC REVIEW OF GENUS UPERODON
TABLE S1. (Continued)
State /
Province
Locality Reference
Durgapur city, Paschim Bardhaman district Pal et al. 2012
3
Amarabati pond, Durgapur, Paschim Bardhaman district Dutta & Mukhopadhyay 2013
3
Durgapur Government College campus, Paschim Bardhaman
district
Dutta & Mukhopadhyay 2013
3
Uttar Pradesh Dudhwa National Park, Kamrup district Hegde 2013
3
Assam Kamrup district Bhaduri & Saha 1980
3
; Dutta 1997
3
Mandakata, Kamrup district Choudhury et al. 1999
3
Sonitpur district Choudhury et al. 1999
3
Dibru Saikhowa National Park Choudhury et al. 2001
3
Barak Valley Dey 2001
3
Sivasagar district Bortamuli & Bordoloi 2008
3
Arunachal
Pradesh
“Pakhui Wildlife Sanctuary, Bhalukpung, Tippi and Near
Khari River”
Borah & Bordoloi 2001
3
Itanagar, Papumpare Borah & Bordoloi 2003
3
BANGLADESH
Chittagong Bandarban Hill district Reza & Perry 2015
3
Kaptai National Park, Rangamati district Shihan & Kabir 2015
3
Dhaka Madhupur National Park, Tangail district Reza & Perry 2015
3
Rasulpur, Madhupur NP, Tangail district Mahony et al. 2009
3
Savar, Dhaka district Mahony et al. 2009
3
Jahangirnagar University, Dhaka district Reza & Perry 2015
3
Rangpur Berakuthi, Barua, Nilphamari district Sarker et al. 2012
3
BHUTAN
– Royal Manas National Park Das et al. 2016a
3
NEPAL
(n.a.) Shrestha 2001
3
; Anders 2002
3
;
Shah & Tiwari 2004
3
; Frost 2017
Uperodon montanus
INDIA
Colonial
location “Anamallays” Parker 1934
3
“Malabar” Parker 1934
2
“Travancore” Parker 1934
3
Tamil Nadu Kalakkad Mundunthurai Tiger Reserve Cherian et al. 1999
3
; Johnsingh 2001
3
;
Vasudevan et al. 2001
3
Kakkachi, Tirunelveli district Present Study
1
Sengaltheri, Tirunelveli district Present Study
2
Kerala “Wynaad”* Jerdon 1854 “1853”
2
“Devicolum, Travancore (4,000-7,000 ft)” Parker 1934
3
“Trevandrum, Travancore” Parker 1934
3
“Kochi” Dutta 1997
3
...Continued on next page
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TABLE S1. (Continued)
State /
Province
Locality Reference
Wayanad Abraham et al. 2001
3
Amballoore, Chimmini Wildlife Sanctuary Andrews et al. 2005a
3
Silent Valley, Poochappara Andrews et al. 2005a
3
Thiruvananthapuram district Andrews et al. 2005b
3
Devikulam, Idukki district Present Study
2
Kesavapara, Nelliyampathy, Palakkad district Present Study
1
Kurichiyarmala, Wayanad district Present Study
1
Munnar, Idukki district Present Study
1
Pandimotta, Shendurney WLS, Kollam district Present Study
1
Settukunnu, Wayanad district Present Study
1
Siruvani, Palakkad district Present Study
2
Upper Manalar, Periyar TR, Idukki district Present Study
1
Thirunelli, Wayanad district Present Study
1
Karnataka Dandeli, Uttara Kannada district Adbulali 1962
3
Sringeri, Chikkamagaluru district Kadadevaru et al. 1998; Krishnamurthy 1996
3
Kudremukh National Park Krishnamurthy 2003
3
Hosagadde (Cardamom estate), Bisale Reserve forest Krishna et al. 2004
3
;
Krishna & Bosch 2007
3
Agumbe, Shimoga district Ganesh et al. 2007
3
Tirthahalli, Shimoga district Ganesh et al. 2007
3
Sharavathi, Uttara Kannada district Gururaja et al. 2008
3
;
Ramachandra et al. 2012
3
Talagini, Shimoga district Kuramoto & Joshy 2009
Thomra, southern Coorg, Kodagu district Ganesh & Asokan 2010
3
Sagar, Shimoga district Joshy & Kuramoto 2011
3
Mirjan, Uttara Kannada district Kumar et al. 2012
3
Kali river basin, Uttara Kannada district Ramachandra et al. 2012
3
Maharashtra Sanjay Gandhi National Park Yazdani et al. 1992
3
Kanari Caves, Salsette Island Dutta 1997
3
Khandala, Pune district Dutta 1997
3
; Padhye & Ghate 2012
3
Sawantwadi, Sindhudurg district Sohani & Samant 2009
3
Amboli, Sindhudurg district Sohani & Samant 2009
3
;
Padhye & Ghate 2012
3
Koyna, Satara district Padhye & Ghate 2012
3
Tamhini, Pune district Dahanukar & Padhye 2005
3
;
Padhye & Ghate 2012
3
“Satara Tehsil” Prasad et al. 2013
3
Gujarat Vansda National Park Vyas 2004
3
Uperodon mormorata
INDIA
Karnataka Sakleshpur, Hassan district* Rao 1937; Present Study
2
...Continued on next page
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TAXONOMIC REVIEW OF GENUS UPERODON
TABLE S1. (Continued)
State /
Province
Locality Reference
Gundia river Basin Gururaja et al. 2007
3
;
Dudani et al. 2010
3
Agumbe, Shimoga district Purushotham & Tapley 2011
3
Baba Budan Hills, Chikmagalur district Gururaja 2012
3
Kempholey, Hassan district Gururaja 2012
3
Dandeli Anshi Tiger Reserve, Uttara Kannada district Gururaja & Ramachandra 2012
3
Kuvempu University Campus, Bhadra Wildife Sanctuary
range, Shimoga district
Narayan et al. 2017
3
Bygoor, Chikmagalur district Present Study
1
Castle Rock, Uttara Kannada district Present Study
1
Dandeli, Uttara Kannada district Present Study
1
Kachigebailu, Shimoga district Present Study
1
Londa, Belgaum district Present Study
2
Maharashtra Barshiv, Phansad Wildlife Sanctuary Katwate et al. 2013
3
Supegaon, Phansad Wildlife Sanctuary Katwate et al. 2013
3
Amboli, Sindhudurga district Sayyed 2014
3
; Present Study
1
Radhanagari Wildlife Sanctuary Yadav & Yankanchi 2014
3
Shivaji University, Kolhapur Yadav et al. 2014
3
Ajoba fort, Thane district Pandirkar et al. 2016
3
Amba, Kolhapur district Present Study
1
Goa Canacona, Cotigao Wildlife Sanctuary Das & Whitaker 1997
3
Anjuna, North Goa district Present Study
1
Netravali, South Goa district Present Study
1
Gujarat Samot, Shoolpaneshwar Wildlife Sanctuary, Narmada district Vyas 2012
3
; Present Study
2
Assam Barak Valley Dey 2001
3
Uperodon nagaoi
SRI LANKA
Central
Province Sri Pada Nature Reserve Karunarathna & Amarasinghe 2009
3
Sabaragamuw
a Province Kithulgala Pethiyagoda et al. 2006
3
; Karunarathna &
Amarasinghe 2009
3
Bambarabotuwa-Ratnapura Karunarathna & Amarasinghe 2009
3
Gilimale Karunarathna & Amarasinghe 2009
3
Koskulana-Panapola Karunarathna & Amarasinghe 2009
3
Pompekale-Ratnapura Karunarathna & Amarasinghe 2009
3
Uda Mäliboda, Samanala Nature Reserve Samarasinghe et al. 2012
3
Southern
Province
Kanneliya Forest Reserve* Manamendra-Arachchi & Pethiyagoda 2001;
Present Study
2
Kudawa, Sinharaja Pethiyagoda et al. 2006
3
Hiyare, Galle district Karunarathna & Amarasinghe 2009
3
; Present
Study
1
Beraliya- Elpitiya, Galle district Karunarathna & Amarasinghe 2009
3
...Continued on next page
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TABLE S1. (Continued)
State /
Province
Locality Reference
Dediyagala Karunarathna & Amarasinghe 2009
3
Hiniduma Karunarathna & Amarasinghe 2009
3
Kanneliya Karunarathna & Amarasinghe 2009
3
Kottawa-Kombala, Hiyare Karunarathna & Amarasinghe 2009
3
Nakiyadeniya Karunarathna & Amarasinghe 2009
3
Matara district (Dediyagala, Wilpita and Sinharaja) Wildlife Conservation Society Report 2009
3
Western
Province Athwelthota, Kalutara district
Karunarathna & Amarasinghe 2009
3
Uperodon obscurus
SRI LANKA
Colonial
location “Ceylon” (=Sri Lanka)* Günther 1864 “1863”
Central
Province Knuckles Mountain Forest Range Dutta & Manamendra-Arachchi 1996
3
;
Weerawardhena & Russell 2012
3
Sarasavigama, Kandy district Lokuhewage & Amerasinghe 2001
3
Gannoruwa, Kandy district Chandrasegaran & Ranawana 2002
3
;
Pethiyagoda et al. 2006
3
Illukkumbura, Matale district Pethiyagoda et al. 2006
3
Kotagala, Nuwara Eliya district Pethiyagoda et al. 2006
3
Gannoruwa Forest Reserve Ukuwela & Ranawana 2007
3
Ambagamuwa, Nuwara Eliya district Kumara & Ukuwela 2009
3
Peradeniya, Kandy district Bowatte & Meegaskumbura 2011
3
; Present
Study
2
Samanala Nature Reserve Karunarathna et al. 2012
3
Bogawantalawa, Nuwara Eliya district Present Study
2
Gammaduwa, Matale district Present Study
2
Laxapana, Nuwara Eliya district Present Study
2
Panwila, Kandy district Present Study
2
Pussellawa, Kandy district Present Study
2
Udawatta Kele, Kandy district Present Study
2
North Western
Province Kurunegala, Kurunegala district Present Study
2
Sabaragamuw
a Province
Morningside, Rakwana, Ratnapura district Janzen & Bopage 2011
3
;
Present Study
2
Uda Mäliboda, Samanala Nature Reserve Samarasinghe et al. 2012
3
Babilapitiya estate, Kegalle district Present Study
2
Koskulana, Panapola, Sinharaja, Ratnapura district Present Study
2
Sabaragamuw
a & Southern
Provinces
Sinharaja, World Heritage Site Dutta & Manamendra-Arachchi 1996
3
;
Bambaradeniya et al. 2003
3
; Meegasakumbura
et al. 2012
3
Uva Province Namunukula, Badulla district Dutta & Manamendra-Arachchi 1996
3
;
Pethiyagoda et al. 2006
3
...Continued on next page
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TAXONOMIC REVIEW OF GENUS UPERODON
TABLE S1. (Continued)
State /
Province
Locality Reference
Uperodon palmatus
SRI LANKA
Central
Province “Newera Ellya”, Sri Lanka* Parker 1934
2
Nuwara-Eliya, Nuwara Eliya district Dutta & Manamendra-Arachchi 1996
3
; Present
Study
1
Bogawantalawa Pethiyagoda et al. 2006
3
Horton plains Pethiyagoda et al. 2006
3
Diyagama estate, Agrapatana Present Study
2
Morey estate Present Study
1
Pattipola, Nuwara Eliya district Present Study
2
Central and
Uva Provinces Hakgala
Pethiyagoda et al. 2006
3
Southern and
Uva Provinces Yala National Park Karunarathna et al. 2017
3
Uperodon rohani sp. nov.
SRI LANKA
Central
Province
Sigiriya, Matale district Dutta & Manamendra-Arachchi 1996
3
; Present
Study
2
Menikdena (Dambulla), Matale district Somaweera et al. 2001
3
Kalugala Proposed Forest Reserve (KPFR), Kaluthara district Botejue & Wattavidanage 2012
3
Dambulla, Matale district Present Study
1
Eastern
Province
Trincomalee Dutta & Manamendra-Arachchi 1996
3
Panam a, Ampara district Somaweera et al. 2004
3
Ampara, Ampara district Present Study
2
Northern
Province
Paranthan, Jaffna, Kilinochchi district Dutta & Manamendra-Arachchi 1996
3
; Present
Study
2
Jaffna, Jaffna district Dutta & Manamendra-Arachchi 1996
3
Opata, Kotugoda Dutta & Manamendra-Arachchi 1996
3
North Central
Province
Anuradhapura, Anuradhapura district Dutta & Manamendra-Arachchi 1996
3
; Present
Study
2
Mihinthale, Anuradhapura district* Dutta & Manamendra-Arachchi 1996
3
; de
Silva 2011
3
; Present Study
1
Mihinthale Sanctuary, Anuradhapura district Sarathchandra et al. 2017
3
Nochchiyagama, Anuradhapura district Present Study
2
Polonnaruwa, Polonnaruwa district Present Study
2
North Western
Province
Anawilundawa Wetland Santuary Rodrigo et al. 2008
3
Puttalam city, Puttalam district Karunarathna & Amaras inghe 2011
3
Karawaddana, Kurunegala district Present Study
1
Kurunegala, Kurunegala district Present Study
2
Makandura, Kurunegala district Present Study
2
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TABLE S1. (Continued)
State /
Province
Locality Reference
Western
Province
National Zoological Garden, Dehiwala Karunarathna et al. 2008
3
Bellanwila - Attidiya Sanctuary (BAS) Karunarathna et al. 2010
3
Uva Province Nilgala Forest Area, Monaragala district Sarathchandra et al. 2017
3
Badulla, Badulla district Present Study
2
Mahiyanganaya, Badulla district Present Study
2
Southern
Province
Palatupana, Kirinda, Hambantota district Dutta & Manamendra-Arachchi 1996
3
; Present
Study
2
Bundala, Hambantota district Present Study
1
Uperodon systoma
INDIA
Colonial
location “India orientali”* Schneider 1799
Tamil Nadu Chengai-MGR (formerly Chengleput) district Das 1996
3
Mamallapuram, Kanchipuram district Dutta 1997
3
Madras Khan 2002
3
; Ganesh & Asokan 2010
3
Mannampandal, Nagapattinam district Ganesh & Chandramouli 2007
3
AVC College, Mannampandal, Nagapattinam district Nath et al. 2012
3
Kalpakkam Nuclear Campus, Kanchipuram district Ramesh et al. 2013
3
Meghamalai, Theni district Srinivas & Bhupathy 2013
3
Ilangi, Tirunelveli district Vasanthi et al. 2014
3
Kudirapppu, Tirunelveli district Vasanthi et al. 2014
3
Nannagaram, Tirunelveli district Vasanthi et al. 2014
3
Coimbatore, Coimbatore district Present Study
1
Kunnapattu, Kanchipuram district Present Study
1
Kerala Kottayam, Kottayam district Dutta 1997
3
Trivandrum, Thiruvananthapuram district Ferguson 1904
3
; Dutta 1997
3
; Saidapur 2001
3
Nilambur, Malappuram district Andrews et al. 2005b
3
Karnataka “Dharwar” (= Dharwad) Uttangi 1951
3
“Biligiriranga Hills” Khan 2002
3
“Mysore” Khan 2002
3
Agumbe, Shimoga district Ganesh & Mouli 2006
3
Kadatoka village, Uttara Kannada district Hegde 2012
3
Bannerghatta, Bangalore Urban district Present Study
1
Maharashra University Campus, Aurangabad, Aurangabad district Jamdar et al. 2010
3
Navegaon National Park, Gondia district Bhandarkar et al. 2012
3
Solapur district Padhye & Ghate 2012
3
“Krishna & Koyna Rivers” in Satara district Sathe & Patil 2016
3
Andhra
Pradesh Chinnarutla, Nallamalai Hills Rao et al. 2005
3
Mannanur, Nallamalai Hills Rao et al. 2005
3
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TAXONOMIC REVIEW OF GENUS UPERODON
TABLE S1. (Continued)
State /
Province
Locality Reference
Sundipenta/Sikharam, Nallamalai Hills Rao et al. 2005
3
Perumukkal, Kaliveli watershed region Ramanujan & Anbarasan 2007
3
Pitchandikulam, Kaliveli watershed region Ramanujan & Anbarasan 2007
3
Success canyon, Kaliveli watershed region Ramanujan & Anbarasan 2007
3
Mannanur – Farahabad, Mahboobnagar district Srinivasulu et al. 2007
3
Mukhadwaram – Srisailam, Kurnool district Srinivasulu et al. 2007
3
Shikaram, Kurnool district Srinivasulu et al. 2007
3
Sunnipenta, Kurnool district Srinivasulu et al. 2007
3
“Nallamala hills” Srinivasulu & Das 2008
3
Seshachalam Biosphere Reserve Guptha et al. 2012
3
Srikakulam district Sebastian et al. 2012
3
Thummalapalle uranium mining area, Thummalapalle, YSR
Kadapa district
Reddy et al. 2013
3
Telangana Ameenpur Lake, Medak district Sailu et al. 2017
3
Gujarat Polo Forest Vanaj, Sabarkantha district Sharma 2005
3
; Jangid et al. 2017
3
Mahi River, Kotna (Vadodara) – Amrol (Anand) Vyas 2013
3
Mini Nadi, Sherkhi (Vadodara) – Angadh (Vadodara) Vyas 2013
3
Vartak River, Pipariya+Koshiyal – Vasanha Buzarg (Both in
Kheda)
Vyas 2013
3
Vishwamitri River, Khalipur (Vadodara) – Maretha/Maneja
(Vadodara)
Vyas 2013
3
Ahwa, Dang district Vyas & Parasharya 2016
3
Bagodara, Ahmedabad district Vyas & Parasharya 2016
3
Bharuch City, Bharuch district Vyas & Parasharya 2016
3
Chhatral, Gandhinagar district Vyas & Parasharya 2016
3
Karamsad, Anand district Vyas & Parasharya 2016
3
Rajkot City, Rajkot district Vyas & Parasharya 2016
3
Shivran, Talala, Junagadh district Vyas & Parasharya 2016
3
Vasna Village, Ahmedabad district Vyas & Parasharya 2016
3
Waghai, Dang district Vyas & Parasharya 2016
3
Rajasthan Phulwari Ki Nal Wildlife Sanctuary Dutta 1997
3
Jhalan Hills, Jaipur Sharma 1992
3
; Sharma et al. 2007
3
“Udaipur” Sharma et al. 2007
3
Merwara Hills, Aravalli range, Ajmer district Sharma & Sharma 2009
3
Odisha “Bubaneswar”, Khordha district Mohanty-Hejmadi et al. 1979; Saidapur 2001
3
Sambalpur district Dash & Mahanta 1993
3
Balighai, Konark-Balukhand WLS Deuti & Raha 2010
3
Bedkot, Karlapat WLS Deuti & Raha 2010
3
Chandraprabha, Konark-Balukhand WLS Deuti & Raha 2010
3
“Mundguda” (= Mundraguda), Karlapat WLS Deuti & Raha 2010
3
Ramchandi, Konark-Balukhand WLS Deuti & Raha 2010
3
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TABLE S1. (Continued)
State /
Province
Locality Reference
“Mendiabiri”, Karlapat WLS Deuti & Raha 2010
3
;
Deuti et al. 2014
3
“Sunabeda”, Koraput district Deuti et al. 2014
3
Gandhamardan Hills
Sahu et al. 2014
3
;
Pradhan et al. 2015
3
Madhya
Pradesh
“Gwalior” Saksena et al. 1988
3
;
Sarkar & Ray 2000
3
; Chandra & Gajbe 2005
3
“Turiya, Seoni district & Pyorthadi” Chandra & Ray 2007
3
Chhattisgarh “Bastar” Dutta 2015
3
Uttar Pradesh Shiwalik Hills, near Badshahibag, Saharanpur district Tilak & Husain 1976
3
Agra (St. John’s College Staff Club grounds – Hariparbat
Police Station; Nala Burhan Sayyad)
Mahendra 1939
3
; Dutta 1997
3
“Allahabad” Dutta 1997
3
“Kanpur” Dutta 1997
3
Dudhwa National Park, Kamrup district Hegde 2013
3
Uttarakhand Rajaji National Park Ray & Tilak 1994
3
Dehradun Husain 2015
3
Himachal
Pradesh Paonta Saheb, Sirmour district Singh 1982
3
Simbalbara Wildlife Sanctuary Mehta & Sharma 2009
3
Mizoram Ngenpui Wildlife Sanctuary Pawar 1999
3
Assam Barak Valley Dey 2001
3
SRI LANKA
Central
Province
Menikdena (Dambulla) Somaweera et al. 2001
3
Dambulla, Matale district Present Study
1
Eastern
Province
Panam a, Ampara district Som aweera et al. 2004
3
Northern
Province
Jaffna Kirtisinghe 1957
3
Marichchukkaddi Dutta & Manamendra-Arachchi 1996
3
North Western
Province
Nawadankulama Dutta & Manamendra-Arachchi 1996
3
Anawilundawa Wetland Santuary Rodrigo et al. 2008
3
Eluwankulama Aruwakkalu Forest Kumarsinghe et al. 2013
3
Sabaragamuw
a Province
Aluthnuwara Kirtisinghe 1957
3
; Dutta & Manamendra-
Arachchi 1996
3
Southern
Province
Lunama-Kalametiya Wetland Sanctuary Ekanayake et al. 2005
3
Uva Province Nilgala Forest Area Sarathchandra et al. 2017
3
NEPAL
(n.a.) Shrestha 2001
3
; Anders 2002
3
; Schleich et al.
2002
3
; Shah & Tiwari 2004
3
; Frost 2017
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TAXONOMIC REVIEW OF GENUS UPERODON
TABLE S1. (Continued)
State /
Province
Locality Reference
Ghodaghodi Lake Bista et al. 2010
3
“15 kms east of Mahendranagar” Frost 2017
PAKISTA N
– Thal and Cholistan Desert Khan 1999
3
; Khan 2004
3
– Margalla Hills, Islamabad Masroor 2011
3
Uperodon taprobanicus
INDIA
Colonial
location
“Cauvery River, Madras”
Parker 1934
3
Tamil Nadu Mamallapuram (= Mahabalipuram), Kanchipuram district Dutta 1997
3
; Present Study
1
Point Calimere, Nagapattinam district Giri et al. 2001
3
Mannampandal, Nagapattinam district Ganesh & Chandramouli 2007
3
Chennai snake park campus, Guindy, Chennai Ganesh 2012
3
AVC College, Mannampandal, Nagapattinam district Nath et al. 2012
3
Kanakaneri, Puducherry Seshadri et al. 2012
3
Korkad aeri, Puducherry Seshadri et al. 2012
3
Kalpakkam Nuclear Campus, Matale district Ramesh et al. 2013
3
Kerala Wayanad Abraham et al. 2001
3
Kuttanad, Pathanamthitta district Andrews et al. 2005b
3
Madras Ganesh & Asokan 2010
3
Kulathupuzha forest range Jose et al. 2014
3
Shendurney Wildlife Sanctuary Jose et al. 2014
3
Kalpetta, Wayanad district Present Study
2
Mannuthy, Thrissur district Present Study
2
Sulthanbathery, Wayanad district Present Study
2
Karnataka Dandeli, Uttara Kannada district Adbulali 1962
3
Dharwad, Dharwad district Dutta 1997
3
; Pattar et al. 2010
3
“Mysore” Dutta 1997
3
Lukunda, Kodagu district Sengupta et al. 2009
3
Devarabetta, Hosur district Chandramouli & Baskaran 2012
3
Devbagh, Karwar, Uttara Kannada district Lewis et al. 2010
3
Uttara Kannada district Ramachandra et al. 2012
3
Taralu, Bangalore Rural district Alexander et al. 2014
3
Gulbarga University Campus, Gulbarga district Jadesh et al. 2014
3
Tumakuru, Tumkur district Kumar & Asiyanuzhat 2016
3
Kuvempu University Campus, Bhadra Wildife Sanctuary
range, Shimoga district
Narayan et al. 2017
3
Bangalore, Bangalore Urban district Present Study
1
Maharashtra Navegaon Bandh Wildife Sanctuary Padhye et al. 2002
3
Arnala Beach, Virar, Mumbai Nandvikar & Shaikh 2009
3
Navegaon National Park, Gondia district Bhandarkar et al. 2012
3
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TABLE S1. (Continued)
State /
Province
Locality Reference
Daryapur taluka, Amravati district Dhande & Khandare 2013
3
Paratwada city, Amravati district Wagh et al. 2017
3
Andhra
Pradesh
Sriharikota Island Sivakumar et al. 2003
3
Chinnarutla, Nallamalai Hills Rao et al. 2005
3
Mallelatheertham, Nallamalai Hills Rao et al. 2005
3
Mannanur, Nallamalai Hills Rao et al. 2005
3
Pulicheruvu, Nallamalai Hills Rao et al. 2005
3
Sundipenta/Sikharam, Nallamalai Hills Rao et al. 2005
3
Pitchandikulam, Kaliveli watershed region Ramanujan & Anbarasan 2007
3
Mukhadwaram, Sunnipenta Srinivasulu et al. 2007
3
Seshachalam Biosphere Reserve Guptha et al. 2012
3
Telangana Ameenpur Lake, Medak district Sailu et al. 2017
3
Gujarat Mal-Samot, Shoolpaneswar Wildlife Sanctuary, Bharuch
district
Naik et al. 1993
3
Ratanmahal Wildlife Sanctuary Vyas & Patel 1994
3
;
Singh et al. 2002
3
Agricultural Universty Campus, Anand, Anand district Vyas & Parasharya 2004
3
Surat city, Surat district Vyas & Parasharya 2004
3
Vartak River, Pipariya+Koshiyal – Vasanha Buzarg (Both in
Kheda)
Vyas 2013
3
Ankleshwar Town, Bharuch district Vyas & Parasharya 2016
Odisha Konarak, Puri district Dutta 1997
3
Bhitarkanika Sanctuary, Behrempur district Sengupta et al. 2009
3
;
Jena et al. 2013
3
Baleshwar, Konark-Balukhand WLS Deuti & Raha 2010
3
Balighai, Konark-Balukhand WLS Deuti & Raha 2010
3
Mendiabiri, Karlapat WLS Deuti & Raha 2010
3
“Mundguda” (= Mundraguda), Karlapat WLS Deuti & Raha 2010
3
Bolangir district Mishra & Guru 2013
3
Gandhamardan Hills Sahu et al. 2014
3
;
Pradhan et al. 2015
3
North Orissa University Campus, Mayurbhanj district Boruah et al. 2016
3
Madhya
Pradesh “Jabalpur” Chandra & Gajbe 2005
3
Pench Tiger Reserve Chandra & Gajbe 2005
3
“Seoni” Chandra & Gajbe 2005
3
Shahdol district Gajbe 2003
3
; Chandra & Gajbe 2005
3
Pench and Satpura Ramakrishna et al. 2006
3
Tropical Forest Research Institute (TFRI) campus, Jabalpur Paunikar 2017
3
Chhattisgarh Jagdalpur, Bastar district Daniel & Verma 1964
3
“Jabalpur” (= Probably Jagdalpur), Indravati Tiger Reserve Sengupta et al. 2009
3
...Continued on next page
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TABLE S1. (Continued)
State /
Province
Locality Reference
“Bastar” Dutta 2015
3
Uttar Pradesh Katerniaghat Wildlife Sanctuary, Bahraich district Das et al. 2012
3
Dudhwa National Park, Kamrup district Hegde 2013
3
West Bengal “Calcutta” (= Kolkata) Parker 1934
3
Camp Kanchrapara (40 kms north of Kolkata) Dutta 1997
3
“Howrah” Sengupta et al. 2009
3
Midnapore district Pratihar & Deuti 2011
3
Durgapur, Paschim Bardhaman district Pal et al. 2012
3
Durgapur Government College campus, Paschim Bardhaman
district
Dutta & Mukhopadhyay 2013
3
Kanthi, Purba Medinipur district Hazra et al. 2014
3
University of Burdwan, Golapbag, Burdwan Aditya & Hazra 2016
3
Bihar –Dutta 1997
3
Assam North of Tinsukia Dutta 1997
3
Santinagar, Kokrajhar town Sengupta et al. 2009
3
SRI LANKA
Colonial
locations
“Ceylon” (= Sri Lanka) Parker 1934
3
“Punduloya, Ceylon, 4,000 ft.”, Sri Lanka* Parker 1934
2
“Kurumgalla, Ceylon”, Sri Lanka Parker 1934
3
Central
Province
Menikdena (Dambulla) Somaweera et al. 2001
3
Kalugala Proposed Forest Reserve, Kaluthara district Botejue & Wattavidanage 2012
3
Knuckles Mountain Forest Range Weerawardhena & Russell 2012
3
Dambulla, Matale district Present Study
1
Eastern
Province
Panam a, Ampara district Som aweera et al. 2004
3
North Western
Province
Kurunegala Kirtisinghe 1957
3
Chilaw Dutta & Manamendra-Arachchi 1996
3
Anawilundawa Wetland Santuary Rodrigo et al. 2008
3
Eluwankulama Aruwakkalu Forest Kumarsinghe et al. 2013
3
Mihinthale Sanctuary Sarathchandra et al. 2017
3
Karawaddana, Kurunegala district Present Study
1
Western
Province
Colombo Kirtisinghe 1957
3
Panadura Kirtisinghe 1957
3
National Zoological Garden, Dehiwala Karunarathna et al. 2008
3
Bellanwila - Attidiya Sanctuary (BAS) Karunarathna et al. 2010
3
Sabaragamuw
a Province
Warakapola Dutta & Manamendra-Arachchi 1996
3
Uda Mäliboda, Samanala Nature Reserve Samarasinghe et al. 2012
3
...Continued on next page
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TABLE S1. (Continued)
State /
Province
Locality Reference
Sabaragamuw
a and Southern
Provinces
Sinharaja World Heritage Site Bambaradeniya et al. 2003
3
Southern
Province
Deniyaya, Sinharaja Forest Reserve Dutta & Manamendra-Arachchi 1996
3
Lunama-Kalametiya Wetland Sanctuary Ekanayake et al. 2005
3
Matara district Wildlife Conservation Society Report 2009
3
Uva Province Nilgala Forest Area Sarathchandra et al. 2017
3
BANGLADESH
Dhaka Madhupur National Park Sengupta et al. 2009
3
;
Reza & Perry 2015
3
NEPAL
(n.a.) Dubois 1987 “1986”
3
; Anders 2002
3
; Shah &
Tiwari 2004
3
; Frost 2017
Province No. 3 Chitwan National Park Bhattarai et al. 2017
3
Uperodon triangularis
INDIA
Colonial
locations
“Malabar”* Günther 1876 “1875”
2
; Parker 1934
3
“Nilgiri Hills” or “Nilgheries” Parker 1934
3
Tamil Nadu “Pycara and Kotagiri”, Nilgiris district Thurston 1888
3
Srivilliputhur Reserve Forests Malhotra & Davis 1991
3
Shevaroy hills Dutta 1997
3
Udhagamandalam (Ooty), Nilgiris district Kuramoto & Dubois 2009
3
;
Present Study
1
Emerald Area, Avalanche Reserve Forest Santhoshkumar et al. 2017
3
Kerala “Chimanji”, Travancore Ferguson 1904
3
Kallar, Ponmudi Inger et al. 1984
3
Silent Valley, Palakkad district Dutta 1997
3
; Andrews et al. 2005a
3
Wayanad Abraham et al. 2001
3
Thiruvananthapuram district Andrews et al. 2005b
3
Kulathupuzha forest range Jose et al. 2014
3
Kalpetta, Wayanad district Present Study
1
Mananthavady, Wayanad district Present Study
1
Pozhuthana, Wayanad district Present Study
1
Suganthagiri, Wayanad district Present Study
1
Karnataka “Hassan” Rao 1937
3
; Dutta 1997
3
“Kadur”, Chikmagalur district Rao 1937
3
“Malnad areas” (= Malenadu region) Rao 1937
3
Mudigere, Chikkamagaluru district Rao 1937
3
“Sakleshpur”, Hassan district Rao 1937
3
Lackunda Estate, Coorg Hampson & Bennett 2002
3
...Continued on next page
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TABLE S1. (Continued)
State /
Province
Locality Reference
Gundya catchment Gururaja et al. 2007
3
Gundia Basin, Mallahlli Waterfalls Dudani et al. 2010
3
Brahmagiri Wildlife Sanctuary, Kodagu district Lewis et al.2010
3
Kempholey, Hassan district Gururaja 2012
3
Rajiv Gandhi National Park, Nagarhole Krishna & Sreepada 2012
3
Kodagu district Rathod & Rathod 2013
3
Madikeri, Kodagu district Present Study
1
Mudigere, Chikmagalur district Present Study
1
Wattakolli, Coorg, Kodagu district Present Study
1
Uperodon variegatus
INDIA
Colonial
locations “Carnatic”
Jerdon “1854” 1853
3
“Yellagherry Hills”
Günther 1876 “1875”
2
;
Boulenger 1882
3
“Bangalore (Mysore), S. India” Rao & Ramanna 1925
2
“Gadavery Valey” (= Godavari Valley) or “Gadavere, near
Bhadrachalam”
Parker 1934
2
; Boulenger 1882
3
“Chanda” Parker 1934
3
“S.E. of Bhadrachalam” Parker 1934
3
Tamil Nadu Tambaram, Kanchipuram district Chakko 1968
3
St. Thomas Mount, Madras Murthy 1973
3
Mannampandal, Nagapattinam district Ganesh & Chandramouli 2007
3
Chennai snake park campus, Guindy, Chennai Ganesh 2012
3
AVC College, Mannampandal, Nagapattinam district Nath et al. 2012
3
Korkad aeri, Puducherry Seshadri et al. 2012
3
Velrampetai, Puducherry Seshadri et al. 2012
3
Courtallam, Tirunelveli District Vasanthi et al. 2014
3
Kudirapppu, Tirunelveli district Vasanthi et al. 2014
3
Melagaram, Tirunelveli district Vasanthi et al. 2014
3
Minnagar, Tirunelveli district Vasanthi et al. 2014
3
Nannagaram, Tirunelveli district Vasanthi et al. 2014
3
Chengalpattu, Kanchipuram district Present Study
1
Kerala Thiruvananthapuram district Andrews et al. 2005b
3
Karamama, Thiruvananthapuram district Present Study
2
Meladoor, Thrissur district Present Study
2
Palakkad, Palakkad district Present Study
1
Karnataka Bangalore Rao & Ramanna 1925
2
; Bai 1956
3
Dharwad, Dharwad district Kanamadi et al. 1993
3
; Dutta 1997
3
; Pattar et
al. 2010
3
Agumbe, Shimoga district Ganesh et al. 2007
3
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References
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Abdulali, H. & Daniel, J.C. (1954) Extension of range of the frog Uperodon globulosum Gunth. Journal of the Bombay Natural
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TABLE S1. (Continued)
State /
Province
Locality Reference
Tirthahalli, Shimoga district Ganesh et al. 2007
3
Devarabetta, Hosur district Chandramouli & Baskaran 2012
3
Brahmagiri Wildlife Sanctuary, Kodagu district Lewis et al. 2010
3
Taralu, Bangalore Rural district Alexander et al. 2014
3
Maharashtra Navegaon Bandh Wildife Sanctuary Padhye et al. 2002
3
Navegaon National Park, Gondia district Bhandarkar et al. 2012
3
Andhra
Pradesh
Ellore (= Eluru), West Godavri district* Stoliczka 1872
2
Potharajupenta–Naguluty, Nagarjunasagar-Srisailam Tiger
Reserve
Srinivasulu et al. 2007
3
“Nallamala hills” Srinivasulu & Das 2008
3
Koruturu, Srikakulam, West Godavri district Present Study
1
Telangana Dongala Dharmaram, Medak district Narayana et al. 2014
3
Ameenpur Lake, Medak district Sailu et al. 2017
3
Madhya
Pradesh
“Chanda in Madhya Pradesh” Daniel 1963b
3
–Dutta 1992
3
; Chandra & Gajbe 2005
3
Odisha Bhubaneswar, Utkal University campus Dutta et al. 1991
3
; Saidapur 2001
3
; Present
Study
2
Sambalpur district Dash & Mahanta 1993
3
Barapali, Bargarh district Dutta 1997
3
Bedkot, Karlapat WLS Deuti & Raha 2010
3
Jaccum, Karlapat WLS Deuti & Raha 2010
3
Mendiabiri, Karlapat WLS Deuti & Raha 2010
3
“Mundguda” (= Mundraguda), Karlapat WLS Deuti & Raha 2010
3
Gandhamardan Hills Sahu et al. 2014;
Pradhan et al. 2015
3
Buguda forest, Ganjam district Das et al. 2016b
3
Jharkhand Chakradharpur (Chota Nagpur), Singhbhum district Rao 1918
3
; Sarkar 1991
3
West Bengal Banspahari, West Midnapur district Deuti 1998
3
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TAXONOMIC REVIEW OF GENUS UPERODON
1831.
Andrews, M.I., George, S. & Joseph, J. (2005b) A survey of the amphibian fauna of Kerala–Distribution and Status. Zoos' Print
Journal, 20 (1), 1723–1735.
Bai, A.K. (1956) Analysis of chromosomes in two genera of Microhylidae (Amphibia; Anura). Proceedings of the Indian
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Supplementary File S1
Neotype description of Ramanella minor Rao, 1937 [= Uperodon minor (Rao, 1937)], ZSI/WGRC/V/A/957, an
adult male, from Sakleshpur (12.8680°N 75.7029°E, 782 m asl), Hassan district, Karnataka state, India, collected
by SD Biju and team on 29 September 2012.
Description of neotype (measurements in mm) (Figs. S1A–E). Small-sized, slender adult male (SVL 31.7;
head small (HW 9.1, HL 7.1, IFE 4.2, IBE 7.4), more than four times (22.4%) smaller than the body length, wider
than long (HW/HL ratio 1.3); snout truncate in dorsal and ventral view, vertical in lateral view, protruding, its
length (SL 3.1) longer than the horizontal diameter of the eye (EL 2.1); loreal region acute with rounded canthus
rostralis; interorbital space more than twice as wide (IUE 3.4) as the upper eyelid width (UEW 1.3); nostril closer
to the tip of snout (NS 0.9) than the eye (EN 2.2); tympanum not visible externally; eye diameter (EL 2.1);
vomerine odontophores present on the palate; presence of two indistinct neopalatinal ridges on posterior side of
each choana, fused with the vomerine odontophores; tongue moderately large, emarginated. Forelimbs moderately
long and thin; forearm (FAL 6.2) shorter than hand length (HAL 8.6); finger length formula I<II<IV<III; tips of all
fingers with truncate discs, without grooves, moderately wide compared to finger width; subarticular tubercles
prominent, oval, all present; two well-developed palmar tubercles, inner oval, outer bilobed. Hindlimbs relatively
long and thin, thigh length (TL 13.8) longer than shank (SHL 11.8) and foot (FOL 13.3); foot webbing present,
moderate: I1
+
–2II1–2III1
+
–2
1
/
2
IV2
1
/
2
–1V; well-developed dermal fringes present on all toes; subarticular tubercles
prominent, oval, all present; two smooth metatarsal tubercles, oval, outer tubercle slightly larger than the inner.
Skin of snout, between eyes, and sides of head, shagreened to granular; anterior and posterior parts of the back,
and upper and lower parts of flank, granular; dorsal surfaces of forelimb, thigh, tibia and tarsus, sparsely granular;
throat granular, chest, abdomen, and ventral surfaces of limbs, shagreened.
Colouration. In preservation: Dorsum dark greyish-brown with a faint grey band; limbs lighter than dorsum;
throat black, chest and belly, light brown with scattered light grey blotches.
Neotype description of Ramanella triangularis rufeventris Rao, 1937, ZSI/WGRC/V/A/959, an adult male, from
Mudigere (13.1258°N 75.5979°E, 900 m asl), Chikkamagaluru district, Karnataka state, India, collected by SD
Biju and team on 28 September 2012.
Description of neotype (measurements in mm) (Figs. S1F–J). Small-sized, slender adult male (SVL 29.7);
head small (HW 9.4, HL 6.7, IFE 4.0, IBE 7.9), more than four times smaller (22.6%) than the body length, wider
than long (HW/HL ratio 1.4); snout rounded to nearly truncate in dorsal and ventral view, nearly acute in lateral
view, its length (SL 2.7) longer than the horizontal diameter of eye (EL 2.2); loreal region obtuse with rounded
canthus rostralis; interorbital space wider (IUE 3.0) than upper eyelid width (UEW 1.9); nostril much closer to the
tip of snout (NS 0.9) than the eye (EN 1.6); eye diameter (EL 2.2); vomerine odontophores present on the palate;
presence of two weakly-developed neopalatinal ridges on posterior side of each choana, fused with the vomerine
odontophores. Forelimb (FAL 6.0) shorter than hand length (HAL 8.8); finger length formula I<II<IV<III, tips of
all fingers with truncate discs, discs moderately wide compared to finger width (FD
I
0.9, FW
I
0.7; FD
II
1.1, FW
II
0.8; FD
III
1.0, FW
III
0.7; FD
IV
1.0, FW
IV
0.6); subarticular tubercles prominent, oval, all present; two well-developed
palmar tubercles (inner oval, 1.2 mm long; outer 1.4 mm long). Hindlimbs relatively long and thin, thigh length
(TL 11.0) longer than shank (SHL 10.1), and shorter than foot (FOL 12.4); tips of all toes with small truncate to
rounded discs, discs rather wide compared to toe width; webbing absent between toes; dermal fringes absent on all
toes; subarticular tubercles prominent, oval; two metatarsal tubercles, oval, outer one longer (1.3 mm) than the
inner (0.7 mm).
Skin of snout, between eyes, sides of head, and anterior parts of the dorsum, shagreened to sparsely granular;
ventral surface uniformly shagreened.
Colouration. In preservation: Dorsum olive grey with a interrupted dark brown median band on the back
starting just above the level of the shoulders and extending up to the vent; a brown stripe between the eyes; flanks
and lateral sides of the head light brown; ventral surfaces (including limbs) coffee brown with scattered cream
coloured spots.
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FIGURE S1. A–E. Neotype of Ramanella minor Rao, 1937 (ZSI/WGRC/V/A/957, male) in preservation: A. dorsal view; B. ventral
view; C. lateral view of head; D. ventral view of hand; E. ventral view of foot; F–J. Neotype of Ramanella triangularis rufeventris
Rao, 1937 (ZSI/WGRC/V/A/959, male) in preservation: A. dorsal view; B. ventral view; C. lateral view of head; D. ventral view of
hand; E. ventral view of foot.
TABLE S2. Factor loadings and variance explained by Principal Components (PC) with eigenvalues of >1.0 based
on Principal component analysis of 17 morphometric variables from adult male and female specimens of Uperodon
rohani sp. nov. and U. variegatus. Values in bold indicate variables with the highest loadings. Abbreviations:
Eigenvalue (Eig), Cumulative (Cum), Variance (Var).
...
Variable Males Females
PC 1 PC 2 PC 3 PC 4 PC 5 PC 1 PC 2 PC 3
SVL -0.726174 0.083894 -0.006431 0.269998 0.429796 -0.740246 0.547413 0.055544
HW -0.786735 -0.285862 0.313254 0.290593 0.132505 -0.961223 0.096277 0.026117
HL -0.450758 0.587954 0.337607 0.068704 0.360313 -0.330804 0.753251 0.068523
SL -0.218642 -0.843790 0.044857 0.081008 -0.144538 -0.826575 -0.463668 -0.178072
EL 0.365862 0.042291 0.146692 0.672302 -0.490739 -0.345379 -0.223589 0.704103
IUE -0.724689 -0.325178 -0.321521 -0.099136 0.100847 -0.850903 -0.159707 -0.162859
UEW -0.348858 -0.317688 -0.633666 -0.232398 -0.140563 -0.722297 -0.263169 0.172053
IFE -0.644302 -0.478635 0.068625 -0.265918 -0.279931 -0.858687 0.322445 0.214330
IBE -0.573113 -0.609823 0.382376 0.067136 -0.008183 -0.810170 -0.498851 -0.120936
MN -0.782379 0.082459 0.160830 -0.286269 -0.091218 -0.839213 0.173740 0.316211
EN -0.369134 0.698583 -0.075633 -0.246473 -0.229042 0.632036 0.325283 0.640278
NS -0.154895 0.029135 -0.776577 0.403826 0.053515 -0.555151 -0.425809 0.448797
FAL -0.543871 0.376210 0.069565 -0.022183 -0.406813 -0.022655 0.918307 0.058434
HAL -0.732483 0.220303 -0.230874 0.305230 -0.102551 -0.279739 0.790193 -0.314204
TL -0.755158 -0.133946 0.153317 0.116153 -0.157851 -0.912485 0.163434 -0.059741
SHL -0.177198 0.856745 0.166785 -0.005020 -0.205022 0.387974 0.844937 0.208278
...Continued on next page
GARG ET AL.
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Zootaxa 4384 (1) © 2018 Magnolia Press
TABLE S2. (Continued)
Variable Males Females
PC 1 PC 2 PC 3 PC 4 PC 5 PC 1 PC 2 PC 3
FOL -0.597825 0.601743 -0.252702 0.020813 0.056578 -0.594520 0.672472 -0.243202
Eig 5.485337 3.755638 1.699489 1.173053 1.011991 7.850641 4.566733 1.572220
Cum Eig 5.48534 9.24097 10.94046 12.11352 13.12551 7.85064 12.41737 13.98959
%Total Var 32.26669 22.09199 9.99700 6.90031 5.95289 46.18024 26.86313 9.24835
% Cum Var 32.2667 54.3587 64.3557 71.2560 77.2089 46.1802 73.0434 82.2917
