Content uploaded by Biju S D
Author content
All content in this area was uploaded by Biju S D on Feb 24, 2018
Content may be subject to copyright.
Accepted by M. Vences: 4 Dec. 2018; published: 23 Feb. 2018
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
P.O. Box 41-383
Auckland 1346
New Zealand
e-mail: magnolia@mapress.com
http://www.mapress.com/j/zt
© 2018 Magnolia Press
All rights reserved.
No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any
means, without prior written permission from the publisher, to whom all requests to reproduce copyright
material should be directed in writing.
This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose
other than private research use.
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
·
5
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
GARG ET AL.
6
·
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
Zootaxa 4384 (1) © 2018 Magnolia Press
·
7
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
GARG ET AL.
8
·
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
......continued on the next page
Zootaxa 4384 (1) © 2018 Magnolia Press
·
9
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
GARG ET AL.
10
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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
......continued on the next page
Zootaxa 4384 (1) © 2018 Magnolia Press
·
11
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
......continued on the next page
GARG ET AL.
12
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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
Zootaxa 4384 (1) © 2018 Magnolia Press
·
13
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.
GARG ET AL.
14
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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.
Zootaxa 4384 (1) © 2018 Magnolia Press
·
15
TAXONOMIC REVIEW OF GENUS UPERODON
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.
GARG ET AL.
16
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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).
Zootaxa 4384 (1) © 2018 Magnolia Press
·
17
TAXONOMIC REVIEW OF GENUS UPERODON
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).
GARG ET AL.
18
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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
Zootaxa 4384 (1) © 2018 Magnolia Press
·
19
TAXONOMIC REVIEW OF GENUS UPERODON
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)
GARG ET AL.
20
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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,
Zootaxa 4384 (1) © 2018 Magnolia Press
·
21
TAXONOMIC REVIEW OF GENUS UPERODON
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.
GARG ET AL.
22
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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.
Zootaxa 4384 (1) © 2018 Magnolia Press
·
23
TAXONOMIC REVIEW OF GENUS UPERODON
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
GARG ET AL.
24
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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
Zootaxa 4384 (1) © 2018 Magnolia Press
·
25
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.
GARG ET AL.
26
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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.
Zootaxa 4384 (1) © 2018 Magnolia Press
·
27
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).
GARG ET AL.
28
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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).
Zootaxa 4384 (1) © 2018 Magnolia Press
·
29
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
GARG ET AL.
30
·
Zootaxa 4384 (1) © 2018 Magnolia Press
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. Sin