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New insights on the systematics and reproductive behaviour in tree frogs of the genus Feihyla, with description of a new related genus from Asia (Anura, Rhacophoridae)

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New insights on the systematics and reproductive behaviour in tree frogs of the genus Feihyla, with description of a new related genus from Asia (Anura, Rhacophoridae)

Abstract

The taxonomy of the Asian tree frog genus Feihyla has been in a state of flux ever since its proposal in 2006. Allocation of species to Feihyla remains confusing, particularly with respect to the closely related genus Chirixalus (formerly Chiromantis sensu lato). At the same time, several Chirixalus species are known only from cursory descriptions and remain poorly studied. In this study, we review the systematics of the genus Feihyla and clarify the generic placement of its members along with all other species currently assigned to Chirixalus. Based on integrative evidence gathered from new collections, examination of types and original descriptions, morphological comparisons, phylogenetic relationships inferred from a multi-gene (three mitochondrial + two nuclear) 1,937 bp dataset, as well as reproductive modes including egg-laying, nesting behaviour, and clutch morphology, our results show that the six species formerly attributed to Feihyla represent three morphologically and phylogenetically distinct groups: (1) the type species of the genus, F. palpebralis, along with F. fuhua constitute the Feihyla palpebralis group or Feihyla sensu stricto; (2) F. inexpectata and F. kajau represent the Feihyla vittiger group and are closely related to the Feihyla palpebralis group; (3) ‘F. hansenae’ and ‘F. vittata’ constitute another distinct and reciprocally monophyletic lineage, more closely related to Chirixalus + Chiromantis. In light of long-standing taxonomic confusions and unresolved phylogenetic relationships, we propose recognition of a new genus to accommodate ‘F. hansenae’ and ‘F. vittata’ in order to stabilise the classification of several Asian rhacophorid species that have been frequently confused and transferred within the Chirixalus–Chiromantis–Feihyla complex. Based on integrative evidence, eight species previously attributed to the genus Chirixalus or Feihyla are formally transferred to Rohanixalus gen. nov., two Chirixalus members are allocated to Feihyla, and one synonymised with Rhacophorus bipunctatus. In addition, we report the first member of the tree frog family Rhacophoridae from the Andaman Islands of India—Rohanixalus vittatus, along with description of its male advertisement call, reproductive behaviour including parental care by the female, and larval morphology. Extended distributions are also provided for Rohanixalus species across Northeast India. The study further reveals the presence of potentially undescribed diversity in the new genus. Altogether, the revised classification and novel insights presented herein will facilitate a better working taxonomy for four phylogenetically distinct but morphologically related groups of Old World tree frogs.
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by M. Vences: 12 Oct. 2020; published: 12 Nov. 2020 1
Zootaxa 4878 (1): 001–055
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4878.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:34C96340-F0F5-440F-AEEB-6AC50F175950
New insights on the systematics and reproductive behaviour in tree frogs of
the genus Feihyla, with description of a new related genus from Asia
(Anura, Rhacophoridae)
S. D. BIJU1,2,13*, SONALI GARG1,3,13*, G. GOKULAKRISHNAN6,7, SIVAPERUMAN CHANDRAKASAN6,8,
PANUPONG THAMMACHOTI9, JINLONG REN10, C. GOPIKA1,4, KARAN BISHT1,5,
AMIR HAMIDY11 & YOGESH SHOUCHE12
1Systematics Lab, Department of Environmental Studies, University of Delhi, Delhi, India
2
sdbiju@es.du.ac.in; https://orcid.org/0000-0003-1039-4421
3
sgarg.du@gmail.com; https://orcid.org/0000-0002-0048-4346
4
gopikacjnv33@gmail.com; https://orcid.org/0000-0002-0047-0438
5
karranbisht5@gmail.com; https://orcid.org/0000-0001-9595-3450
6Andaman and Nicobar Regional Centre, Zoological Survey of India, Port Blair, Andaman and Nicobar Islands, India
7
gokul7701@gmail.com; https://orcid.org/0000-0002-3574-1891
8
c_sivaperuman1@rediffmail.com; https://orcid.org/0000-0002-3582-7767
9Department of Biology, Faculty of Science, Chulalongkorn University, Phyathai Road, Pathumwan, Bangkok, Thailand.
tatsuya_th@hotmail.com; https://orcid.org/0000-0003-3477-8964
10Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu, Sichuan, China.
renjl@cib.ac.cn; https://orcid.org/0000-0003-4896-7600
11Museum Zoologicum Bogoriense, Research Center for Biology, Indonesian Institute of Sciences, Cibinong, West Java, Indonesia.
hamidyamir@gmail.com; https://orcid.org/0000-0001-6426-7458
12National Centre for Cell Science, Savitribai Phule University of Pune Campus, Ganeshkhind, Pune, Maharashtra, India.
yogesh@nccs.res.in; https://orcid.org/0000-0002-5055-5931
13Equally contributing authors
*Corresponding authors.
sdbiju.es@gmail.com;
sgarg.du@gmail.com
Table of contents
Abstract ...................................................................................................2
Introduction ................................................................................................2
Material and methods ........................................................................................4
Results ..................................................................................................16
Phylogenetic relationships .............................................................................16
DNA barcoding .....................................................................................17
TaxonomictreatmentTaxonomic treatment .................................................................................18
I. Systematic review of genus Feihyla ..................................................................18
Genus Feihyla .................................................................................18
Feihyla palpebralis group .....................................................................21
Feihyla fuhua Fei, Ye, and Jiang, 2010 .........................................................21
Feihyla palpebralis (Smith, 1924) .............................................................24
Feihyla vittiger group ........................................................................ 24
Feihyla inexpectata (Matsui, Shimada, and Sudin, 2014). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Feihyla kajau (Dring, 1983) .................................................................26
Feihyla samkosensis (Grismer, Thy, Chav, and Holden, 2007) comb. nov. .............................26
Feihyla vittiger (Boulenger, 1897) comb. nov. ...................................................27
II. Description of a new genus ........................................................................27
Genus Rohanixalus gen. nov. ....................................................................27
Rohanixalus vittatus (Boulenger, 1887) comb. nov. ................................................31
Rohanixalus baladika (Riyanto and Kurniati, 2014) comb. nov. .......................................37
Rohanixalus hansenae (Cochran, 1927) comb. nov. ................................................39
Rohanixalus marginis (Chan, Grismer, Anuar, Quah, Grismer, Wood, Muin, and Ahmad, 2011) comb. nov. . . . . 40
Rohanixalus nauli (Riyanto and Kurniati, 2014) comb. nov. .........................................41
Rohanixalus punctatus (Wilkinson, Win, Thin, Lwin, Shein, and Tun, 2003) comb. nov. ...................43
BIJU ET AL.
2 · Zootaxa 4878 (1) © 2020 Magnolia Press
Rohanixalus senapatiensis (Mathew and Sen, 2009) comb. nov. ......................................44
Rohanixalus shyamrupus (Chanda and Ghosh, 1989) comb. nov. .....................................45
III. Taxonomic identity of Chirixalus cherrapunjiae (Roonwal and Kripalani, 1966 “1961”) . . . . . . . . . . . . . . . . . . . . . . 47
Discussion ...............................................................................................47
Acknowledgments ..........................................................................................50
References ................................................................................................50
Abstract
The taxonomy of the Asian tree frog genus Feihyla has been in a state of flux ever since its proposal in 2006. Allocation
of species to Feihyla remains confusing, particularly with respect to the closely related genus Chirixalus (formerly
Chiromantis sensu lato). At the same time, several Chirixalus species are known only from cursory descriptions and
remain poorly studied. In this study, we review the systematics of the genus Feihyla and clarify the generic placement of
its members along with all other species currently assigned to Chirixalus. Based on integrative evidence gathered from
new collections, examination of types and original descriptions, morphological comparisons, phylogenetic relationships
inferred from a multi-gene (three mitochondrial + two nuclear) 1,937 bp dataset, as well as reproductive modes including
egg-laying, nesting behaviour, and clutch morphology, our results show that the six species formerly attributed to Feihyla
represent three morphologically and phylogenetically distinct groups: (1) the type species of the genus, F. palpebralis, along
with F. fuhua constitute the Feihyla palpebralis group or Feihyla sensu stricto; (2) F. inexpectata and F. kajau represent
the Feihyla vittiger group and are closely related to the Feihyla palpebralis group; (3) F. hansenaeand F. vittata
constitute another distinct and reciprocally monophyletic lineage, more closely related to Chirixalus + Chiromantis. In
light of long-standing taxonomic confusions and unresolved phylogenetic relationships, we propose recognition of a new
genus to accommodate F. hansenae and ‘F. vittata in order to stabilise the classification of several Asian rhacophorid
species that have been frequently confused and transferred within the ChirixalusChiromantisFeihyla complex. Based
on integrative evidence, eight species previously attributed to the genus Chirixalus or Feihyla are formally transferred
to Rohanixalus gen. nov., two Chirixalus members are allocated to Feihyla, and one synonymised with Rhacophorus
bipunctatus. In addition, we report the first member of the tree frog family Rhacophoridae from the Andaman Islands
of India—Rohanixalus vittatus, along with description of its male advertisement call, reproductive behaviour including
parental care by the female, and larval morphology. Extended distributions are also provided for Rohanixalus species
across Northeast India. The study further reveals the presence of potentially undescribed diversity in the new genus.
Altogether, the revised classification and novel insights presented herein will facilitate a better working taxonomy for four
phylogenetically distinct but morphologically related groups of Old World tree frogs.
Key words: Chiromantis, Chirixalus, egg attendance, gel-nesting, generic reallocation, integrative taxonomy, larval
morphology, molecular phylogeny, Rohanixalus gen. nov., species groups
Introduction
The Old World tree frog subfamily Rhacophorinae Hoffman, 1932 (1858) is one of the most speciose anuran groups,
with 19 recognised genera and 422 species (Frost 2020). Generic allocation among rhacophorids has been difficult
due to the rapid and high level of diversification within the family and paucity of morphological synapomorphies,
as well as often unresolved phylogenetic relationships (e.g., Bossuyt & Dubois 2001; Wilkinson et al. 2002; Delo-
rme et al. 2005; Frost et al. 2006; Bossuyt et al. 2006; Biju et al. 2008; Biju & Bossuyt 2009; Li et al. 2008, 2009,
2013; Yu et al. 2008). One such taxonomically challenging group is an Afro-Asian radiation of tree frogs, first
described as Chiromantis Peters, 1854, with two African members—Chiromantis xerampelina Peters, 1854 (the
type species) and Chiromantis rufescens (Günther, 1869). The Asian members of this radiation were originally as-
signed to another genus, Chirixalus Boulenger, 1893, with Chirixalus doriae Boulenger, 1893 as the type species.
The latter was synonymised with Chiromantis based on close phylogenetic relationship, though the monophyly of
all previously attributed Chirixalus members (such as Chirixalus vittatus’) remained weakly supported (Frost et
al. 2006). At the same time, a new genus Feihyla (Frost et al. 2006) was established to accommodate a single taxon
Philautus palpebralis Smith, 1924, previously referred to Chirixalus (e.g., Bourret 1939; Inger et al. 1999; Bossuyt
& Dubois 2001; Wilkinson et al. 2002; Delorme et al. 2005), in order to resolve the polyphyly of Chirixalus and
Chiromantis, thereby recognising two genera—Chiromantis sensu lato and Feihyla. Subsequent to the genus-level
changes by Frost et al. (2006), four species were transferred to Feihyla based on molecular evidence: Philautus
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 3
hansenae Cochran, 1927 (Chiromantis hansenae sensu Frost et al. 2006) by Aowphol et al. (2013); Chiromantis
inexpectatus Matsui, Shimada, and Sudin, 2014 by Chan et al. (2018); Rhacophorus kajau Dring, 1983 by Li et
al. (2013, supplemental data) and subsequently confirmed by Hertwig et al. (2013); and Ixalus vittatus Boulenger,
1887 (Chiromantis vittatus sensu Frost et al. 2006) by Fei et al. (2010). Further, one new species Feihyla fuhua was
described by Fei et al. (2010). With this, the genus Feihyla presently comprises six species widely distributed across
South, Southeast, and East Asia.
Most recently, another study (Chen et al. 2020) has once again proposed the recognition of the genus Chirixa-
lus, based on phylogenetic and geographical considerations. By implication, this has prompted the transfer of 14
Asian members from the genus Chiromantis to Chirixalus (Frost 2020), though in the absence of either molecular
evidence for several of these taxa or any morphological discussion. This action has also opened up questions con-
cerning: (1) the long overlooked morphological and phylogenetic affinities of the six species presently included in
the genus Feihyla; (2) the generic allocation of species to the genus Chirixalus as presently understood (Chen et al.
2020; Frost 2020) or the former Chiromantis sensu lato; and (3) the monophyly of the genus Feihyla, particularly
due to the previously shown phylogenetic position of ‘Feihyla vittata’ as being closer to Chirixalus + Chiromantis
(e.g., Frost et al. 2006; Yu et al. 2009; Hertwig et al. 2012; Meegaskumbura et al. 2015; Poyarkov et al. 2015; Biju
et al. 2016).
Of the above, the former two questions concerning species-level identification are central to any meaningful
phylogenetic resolution of the ChirixalusChiromantisFeihyla taxonomic conundrum. Several species in the cur-
rently recognised Asian genera Chirixalus and Feihyla were originally described in various rhacophorid genera such
as Ixalus (Boulenger 1887; Boulenger 1897; Smith 1924), Philautus (Cochran 1927; Roonwal & Kripalani 1966
“1961”; Chanda & Ghosh 1989), or Rhacophorus (Dring 1983). Most species have also been frequently transferred
between the genera Chirixalus and Chiromantis owing to unstable genus-level taxonomic definition (e.g., Ahl 1931;
Bourret 1942; Liem 1970; Bossuyt & Dubois 2001; Frost et al. 2006; Wostl et al. 2017; Chen et al. 2020). Even
some recent novelties were originally assigned to Chiromantis (e.g., Grismer et al. 2007; Mathew & Sen 2009; Chan
et al. 2011; Matsui et al. 2014; Riyanto & Kurniati 2014), probably due to uncertain knowledge of inter- and intra-
generic phylogenetic relationships and an incomplete morphological understanding of these groups, particularly
Feihyla. Furthermore, Chirixalus cherrapunjiae (Roonwal and Kripalani, 1966 “1961”), C. shyamrupus (Chanda
and Ghosh, 1989), C. senapatiensis (Mathew and Sen, 2009), and C. vittiger (Boulenger, 1897), still remain poorly
studied morphologically or phylogenetically, either due to their cursory original descriptions or lack of subsequent
collections. Altogether, this has led to phenotypic synapomorphies of the genera Feihyla and Chirixalus being over-
looked, particularly in terms of morphology and reproductive behaviour, resulting in taxonomic uncertainty.
At the genus-level, the monophyly of Feihyla with its current constituents remains largely unresolved (e.g.,
Wilkinson et al. 2002; Frost et al. 2006; Li et al. 2008; Yu et al. 2009; Meegaskumbura et al. 2015; Poyarkov et
al. 2015; Biju et al. 2016). Though, some recent phylogenies utilising a greater number of loci, albeit with limited
taxonomic representation, also suggest ‘Feihyla’ to be a monophyletic group (e.g., Li et al. 2009, 2013; Hertwig et
al. 2013; Chan et al. 2018; Chen et al. 2020). Nonetheless, the generic placement of species, as well as their intra-
and intergeneric systematic relationships based on morphology, remains altogether unattempted, despite occasional
new-species descriptions both in Chirixalus and Feihyla in the recent years (Riyanto & Kurniati 2014; Chan et al.
2011; Wilkinson et al. 2003; Mathew & Sen 2009; Fei et al. 2010; Matsui et al. 2014).
In this backdrop, we study all known species of the Asian tree frog genus Feihyla and revisit the generic alloca-
tion of species in the closely related genus Chirixalus, based on morphology, molecular affinities, and reproductive
modes including egg-laying, nesting behaviour, and clutch morphology of species. Further, on the basis of revised
species classification, we investigate the phylogenetic relationships of the three genera—Chirixalus, Chiromantis,
and Feihyla—and conclude with the need to recognise a new genus (Rohanixalus gen. nov.) to accommodate eight
previously known species. We delimit the geographical range of the genus Feihyla and clarify the distributions of
the Rohanixalus gen. nov. species in India, including new records based on fresh collections. Notes on tadpole
morphology, structure of male advertisement calls, breeding ecology, and natural history are also included for the
proposed type species of the new genus.
BIJU ET AL.
4 · Zootaxa 4878 (1) © 2020 Magnolia Press
Material and methods
Field studies. Field surveys, sampling, and natural history observations were carried out in India (across Northeast
states and Andaman Islands), Indonesia (Java and Sumatra), and Thailand (Chon Buri Province). Occasional natu-
ral history observations were also made in China (Guangxi Zhuang Autonomous Region and Yunnan Province).
Opportunistic searches were generally carried out at night and largely during the breeding season (May–August in
Northeast India and the Andaman Islands, June–August and January–March in Indonesia, and June–July in China).
Live animals (adults and tadpoles), egg clutches, and behaviour were photographed in the wild. Sampled individuals
were euthanized in Tricaine methanesulfonate (MS-222) solution. Prior to fixation, a portion of the thigh muscle or
liver in adults and tail muscle in tadpoles was extracted and stored in absolute ethanol for molecular studies. Adult
specimens were fixed in 4% formalin and subsequently preserved in 70% ethanol. Eggs and tadpoles were preserved
in 3.5–4% neutral buffered formalin. Referred specimens are available at the Zoological Survey of India–Andaman
and Nicobar Regional Centre (ZSI–ANRC), Port Blair, India; Systematics Lab, University of Delhi (SDBDU),
India; Museum Zoologicum Bogoriense (MZB), Research Center for Biology, Indonesian Institute of Sciences,
Indonesia; and Chulalongkorn University Museum of Natural History (CUMZ), Thailand.
Molecular study. Genomic DNA was extracted from tissue samples of the new populations by using Qiagen
DNeasy blood and tissue kit (Qiagen, Valencia, CA, USA). The following gene fragments (as desired for phylo-
genetic inference or barcoding analyses) were PCR-amplified and cycle-sequenced on both strands using previ-
ously published primer sets: ~315 bp segment of exon 1 of the nuclear rhodopsin gene (RHO: Rhod1A, Rhod1D)
(Bossuyt & Milinkovitch 2000), ~555 bp segment of the nuclear recombinase activating gene 1 (RAG1: Rag1-C,
Rag 1-E) (Biju & Bossuyt 2003), ~750 bp segment of the mitochondrial 12SrRNA (partial) + tRNA
VAL (complete) +
16SrRNA (partial) (H3296, L2519) (Richards & Moore 1996), and ~540 bp segment of the mitochondrial 16SrRNA
(16Sar, 16Sbr) (Simon et al. 1994). Purified PCR products were sequenced on both strands using BigDye Termi-
nator v3.1 Cycle Sequencing Kit on ABI 3730 automated DNA sequencer (Applied Biosystems). DNA sequences
were checked and assembled in ChromasPro v1.34 (Technelysium Pty Ltd.) and deposited in the GenBank under
accession numbers MW054213–MW054235 and MW055937–MW055975 (Table 1).
In order to infer the phylogenetic relationships between various Feihyla and ‘Chirixalus members, a multi-
gene dataset was assembled with representatives from all known rhacophorid genera. Three mitochondrial genes
(1,063 bp of 12SrRNA + tRNA
VAL + 16SrRNA) and two nuclear genes (562 bp RAG1 and 312 bp of RHO) were
sampled for a total of 125 taxa (Table 1). The alignments were created using ClustalW in MEGA 6.0 (Tamura et al.
2013). The coding gene alignments were checked by comparison with amino acid sequences; alignments for the
non-coding segments were visually optimised and ambiguously aligned regions with low confidence for positional
homology were excluded from phylogenetic analyses. Maximum Likelihood and Bayesian analyses were performed
on a total data matrix of 1,937 bp, partitioned by genes. The appropriate model of sequence evolution, i.e. General
Time Reversible (GTR) model with a proportion of invariant sites (+I) and gamma-distributed rate variation among
sites (+G), was independently determined for each gene by implementing Akaike Information Criteria in ModelTest
3.4 (Posada & Crandall 1998). Bayesian analyses were executed for 10 million generations in MrBayes (Ronquist
& Huelsenbeck 2003), using uniform priors and four Metropolis-Coupled Markov Chain Monte Carlo (MCMCMC)
chains, with sampling frequency of trees set at every 1000th generation. Bayesian posterior probabilities (BPP)
for the clades were summarised after discarding the first 25 percent of trees as burn-in (Huelsenbeck et al. 2001).
Maximum Likelihood clade support (BS) was assessed through 10,000 thorough bootstrap replicates using RAxML
7.3.0 (Stamatakis et al. 2008) as implemented in raxmlGUI 1.1 (Silvestro & Michalak 2012). Maximum Likelihood
(ML) searches were also performed in PAUP* (Swofford 2002), to assess the topology on an unpartitioned dataset,
with all parameters estimated.
To further assess the potential diversity within Feihyla and the new genus described in the present study, avail-
able homologous 16S sequences (short fragment of ~540 bp) for species assigned to these genera in the phylogenet-
ic analyses were retrieved from the GenBank (Table 1). This fragment represents the highest number of sequences
currently available for these groups. Two datasets were assembled: one for 15 Feihyla samples and another for 135
samples of the new genus, including new collections and outgroup taxa. Maximum Likelihood (ML) searches were
performed in PAUP* (Swofford 2002) using GTR+I+G as the best-fit model of DNA evolution. Bayesian analyses
for 10 million generations and RAxML support for 10,000 rapid bootstrap replicates were assessed as discussed
previously. For comparison of intra- and interspecific divergences, uncorrected pairwise genetic distances for the
16S rRNA were computed in PAUP* using all sites (Swofford 2002).
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 5
TABLE 1. List of taxa and DNA sequences included in the study. (1) Indicates taxa used for the multi-gene phylogenetic analysis of family Rhacophoridae; (2) indicates taxa used
for the 16S barcoding analyses of Feihyla and Rohanixalus gen. nov. Institutional abbreviations: SCUM (Zoological Museum of the School of Life Science, Sichuan University,
China); NMNS (National Museum of Natural Science, Taiwan); KIZ (Kunming Institute of Zoology, China); KUHE (Kyoto University, Graduate School of Human and Environ-
mental Studies, Japan); ZFMK (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Germany); ROM (Royal Ontario Museum, Department of Herpetology, Toronto,
Canada); THNHM (Thailand Natural History Museum, Thailand); HNNU (Department of Biology, Hunan Normal University, Changsha, China); WHT (Wildlife Heritage Trust,
Colombo, Sri Lanka); MVZ (Museum of Vertebrate Zoology, University of California Berkeley, USA); ZMH (Zoologisches Museum Hamburg, Universität Hamburg, Germany);
NMBE (Naturhistorisches Museum Bern, Switzerland); ZMKU (Zoological Museum, Kasetsart University, Bangkok, Thailand); MNHN (Muséum National d’histoire Naturelle,
Paris, France); SN (Shunqing field number); ACD (Arvin C. Deismos field number); SDBDU (Systematics Lab, University of Delhi, India); CUMZ (Chulalongkorn University
Museum of Natural History, Thailand); MZB (Museum Zoologicum Bogoriense, Research Center for Biology, Indonesian Institute of Sciences, Indonesia).
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
Subfamily Buergeriinae
1Buergeria oxycephala China: Hainan SCUM 050267YJ EU215556 GQ285758 EU215524 (1)
Subfamily Rhacophorinae
2Beddomixalus bijui India: Kadalar, Idukki, Kerala SDBDU 2011.1006 KU169946 KU169970 KU170017, KU169995 (1)
3Chirixalus doriae Myanmar: Kyaiphaung to Win Bo village, Kyaitong
Township
CAS 235526 MW055950 MW055937 MW055963, MW054213 (1)
4Chirixalus doriae (1) China NMNS 3183 AF458127 (1)
5Chirixalus doriae (2) Laos: Phou Louey National Biodiversity Conserva-
tion Area, Vieng Tong, Huaphahn Prov.
FMNH 255213 DQ283836 DQ283135 (1)
6Chirixalus doriae’ (3) China: Hainan SN 030051 EU215554 GQ285779 EU215527 (1)
7Chirixalus doriae’ (4) China: Simao, Yunnan KIZ 060821034 EU924539 EU924511 EF564444, EF564516 (1)
8Chirixalus doriae’ (5) China: Menglun, Yunnan KIZ 060821123 EF564445, EF564517 (1)
9Chirixalus doriae’ (6) China: Jindong, Yunnan KIZ 060821257 EF564446, EF564518 (1)
10 Chirixalus doriae’ (7) China: Simao, Yunnan KIZ 005Rao GQ285682 (1)
11 Chirixalus doriae (8) Laos: Viengthong district, Huaphahn Prov. FMNH 255215 GQ204657 GQ204602 GQ204772, GQ204721 (1)
12 Chirixalus doriae’ (9) Thailand: Phu Luang KUHE 19301 AB813159 (1)
13 Chirixalus nongkhorensis (1) Thailand: Nong Khor, Si Racha district, Chonburi
Prov.
CUMZ-A-7735 MW055951 MW055938 MW055964, MW054214 (1)
14 Chirixalus nongkhorensis (2) Laos FMNH 255378 GQ204659 GQ204604 GQ204774, GQ204723 (1)
15 Chirixalus nongkhorensis (3) Viet Nam: Ta Kou Mountain Natural Reserve KIZ 1056 KC465788 (1)
16 Chirixalus nongkhorensis (4) Thailand: Song Khla Prov. KUHE 19498 AB813158 (1)
17 Chiromantis petersii Africa MVZ 234168 GQ204784, GQ204733 (1)
......continued on the next page
BIJU ET AL.
6 · Zootaxa 4878 (1) © 2020 Magnolia Press
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
18 Chiromantis rufescens (1) Equatorial Guinea CAS 207599 DQ347356 DQ347237 DQ347004, DQ347297 (1)
19 Chiromantis rufescens (2) N.A. MNHN D734 AY880627 AY880539, AY880494 (1)
20 Chiromantis rufescens (3) Cameroon ZFMK 87811 KF991262, KF991282 (1)
21 Chiromantis xerampelina (1) N.A. MNHN 2000.2434 AY880540, AY880495 (1)
22 Chiromantis xerampelina (2) Africa MVZ 234606 GQ204785, GQ204734 (1)
23 Feihyla inexpectata (1) Malaysia: Camel Trophy field station, Maliau Basin
Conservation Area, Sandakan Division, Sabah
BORNENSIS22421 AB813160 (1), (2)
24 Feihyla inexpectata (2) Malaysia: Nepenthes Camp/Trails, Maliau Basin
Conservation Area, Sandakan Division, Sabah
ZMHA 12647 MG909569 (2)
25 Feihyla kajau (1) Malaysia: Camp 1, Gunung Mulu N. P., Sarawak,
Borneo
NMBE 1056500 KC961240, JN377362 (1), (2)
26 Feihyla kajau (2) Malaysia: Bintulu Division, Sarawak, Borneo FMNH 269090 KC465789 (2)
27 Feihyla kajau (3) Malaysia: Lambir N.P., Borneo KUHE 53591 AB847122 (2)
28 Feihyla kajau (4) Malaysia: Summit Road, Kubah N. P., Sarawak,
Borneo
NMBE 1057090 KC961241, KC961088 (2)
29 Feihyla kajau (5) Malaysia: Kubah N. P., Sarawak, Borneo ZMHAH 518 – , KT382330 (2)
30 Feihyla palpebralis Vietnam: Lam Dong KIZ 712 GQ285792 GQ285772 GQ285681 (1), (2)
31 Feihyla fuhua (1) China: Pingbian, Yunnan KIZ 080177 EU924541 EU924513 EU924625, EU924620 (1), (2)
32 Feihyla fuhua (2) China: Mt. Dawei, Yunnan SCUM 0606132L EU215576 GQ285773 EU215546 (1), (2)
33 Feihyla fuhua China NMNS 3104 AF458130 (2)
34 Feihyla fuhua N.A. N.A. –, JQ517287 (2)
35 Feihyla fuhua N.A. N.A. –, JQ621936 (2)
36 Feihyla vittiger Indonesia: Telaga Warna, Sukabumi, Jabar (West
Java)
MZBAmph27238 – , MW054231 (1), (2)
37 Ghatixalus asterops India N.A. KT359632 KT359636 KT359620, KT359626 (1)
38 Ghatixalus magnus India N.A. KT359630 KT359634 KT359618, KT359624 (1)
39 Ghatixalus variabilis India: Mukkurthi NP, Nilgiris, Tamil Nadu SDBDU 2008.4409 KU169932 KU169956 KU170006, KU169981 (1)
40 Gracixalus carinensis Vietnam: Sa Pa, Lao Cai ROM 39660 GQ285788 GQ285762 GQ285670 (1)
41 Gracixalus gracilipes China: Pingbian, Yunnan KIZ 060821196 EU924537 EU924509 EF564451, EF564523 (1)
......continued on the next page
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 7
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
42 Gracixalus jinxiuensis China: Mt. Dayao, Guangxi KIZ 061210YP EU215557 GQ285763 EU215525 (1)
43 Kurixalus appendiculatus Malaysia: Bukit Sarang, Sarawak, Borneo FMNH 267904 JQ060911 JQ060949, JQ060938 (1)
44 Kurixalus banaensis Vietnam: Krong Pa, Gia Lai ROM 32986 GQ285781 GQ285752 GQ285667 (1)
45 Kurixalus bisacculus Thailand: Pua district, Nan Prov. THNHM 10052 JQ060897 GU227280, GU227335 (1)
46 Kurixalus eiffingeri Japan KUHE 24111/A120 AY880625 –, DQ468673 (1)
47 Kurixalus hainanusChina: Mt. Diaoluo, Hainan HNNU A1180 EU215578 GQ285749 EU215548 (1)
48 Kurixalus idiootocus China: Lianhuachi, Taiwan SCUM 061107L EU215577 GQ285751 EU215547 (1)
49 Kurixalus odontotarsus China: Mengyang, Jinghong SCUM 060688L EU215579 GQ285750 EU215549 (1)
50 Kurixalus verrucosus Myanmar: Nagmung, Kachin CAS 224381 JQ060901 GU227274, GU227329 (1)
51 Leptomantis bimaculatus Philippines: Municipality of Tampakan, Cotabato
Prov, Mindanao
ACD 5395 KF933203 KF933272, – (1)
52 Leptomantis gauni Malaysia FMNH 235047 GQ204650 GQ204596 GQ204765, GQ204714 (1)
53 Liuixalus hainanus China: Mt. Diaoluo, Hainan SCUM 060401L GQ285785 GQ285757 GQ285671 (1)
54 Liuixalus ocellatus China: Mt. Wuzhi, Hainan HN0806045 GQ285784 GQ285755 GQ285672 (1)
55 Liuixalus romeri China: Mt. Shiwan, Guangxi KIZ 061205YP EU215559 GQ285756 EU215528 (1)
56 Mercurana myristicapalustris India: Chathankod, Thiruvananthapuram, Kerala SDBDU 2011.849 KU169945 KU169969 KU170016, KU169994 (1)
57 Nasutixalus jerdonii India: Mawphlang, Meghalaya SDBDU 2009.1166 KU169950 KU169974 KU230455, KU169999 (1)
58 Nasutixalus medogensis (1) India: Sessa, Arunachal Pradesh SDBDU 2009.1295 KU169955 KU169979 KU230460, KU170004 (1)
59 Nasutixalus medogensis (2) China: Motuo, Xizang 6255Rao GQ285782 GQ285753 GQ285679 (1)
60 Nyctixalus pictus Malaysia: Sarawak, Borneo MVZ 239460 GQ204666 GQ204613 GQ204783, GQ204732 (1)
61 Nyctixalus spinosus Philippines: Mindanao ACD 1043 DQ283827 DQ283114 (1)
62 Philautus abditus Vietnam: Buon Luoi, An Khe ROM33145 GQ285794 GQ285775 GQ285673 (1)
63 Philautus aurantium Malaysia: Mendolong, Borneo FMNH 233226 GQ204642 GQ204587 GQ204756, GQ204705 (1)
64 Philautus aurifasciatus Indonesia: Java ZRC1.1.5267 GQ204640 GQ204584 AY141805, GQ204702 (1)
65 Philautus ingeri Malaysia: Sarawak, Borneo FMNH 239280 AY880629 GQ204588 GQ204757, GQ204706 (1)
66 Philautus mjobergi Malaysia: Sarawak, Borneo FMNH 252411 GQ204644 GQ204590 GQ204759, GQ204708 (1)
67 Pseudophilautus amboli India: Castle Rock, Uttara Kannada, Karnataka SDBDU 2011.829 KU169933 KU169957 KU170007, KU169982 (1)
68 Pseudophilautus cavirostris Sri Lanka WHT 3299 GQ204622 GQ204561 FJ788137, FJ788156 (1)
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BIJU ET AL.
8 · Zootaxa 4878 (1) © 2020 Magnolia Press
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
69 Pseudophilautus limbus Sri Lanka: Haycock [Hiniduma] WHT 2700 GQ204553 AY141779, GQ204668 (1)
70 Pseudophilautus microtympa-
num
Sri Lanka: Central Hills WHT 5065 AF249126 GQ204563 AF249030, GQ204678 (1)
71 Pseudophilautus poppiae Sri Lanka: Rakwana Hills WHT 2779 GQ204616 GQ204555 FJ788136, FJ788155 (1)
72 Pseudophilautus sarasinorum Sri Lanka WHT 2481 GQ204614 GQ204552 AY141761, GQ204667 (1)
73 Pseudophilautus schmarda Sri Lanka WHT 2715 GQ204615 GQ204554 AY880617, GQ204669 (1)
74 Pseudophilautus wynaadensis India: Kalpetta, Waynaad, Kerala SDBDU 2010.334 KU169934 KU169958 KU170008, KU169983 (1)
75 Polypedates cruciger Sri Lanka WHT 2640 GQ204632 GQ204570 GQ204746, GQ204687 (1)
76 Polypedates leucomystax Indonesia: Java ZRC 1.15269 GQ204636 GQ204574 –, GQ204693 (1)
77 Polypedates macrotis Malaysia FMNH 239119 GQ204638 GQ204577 GQ204748, GQ204695 (1)
78 Polypedates maculatus Sri Lanka WHKANT GQ204637 GQ204576 GQ204747, GQ204694 (1)
79 Polypedates megacephalus China: Mt. Daiyun, Fujian SCUM 050508C EU215582 GQ285771 EU215552 (1)
80 Polypedates mutus China: Xishuangbanna, Yunnan SCUM 37940C EU215581 GQ285770 EU215551 (1)
81 Polypedates pseudocruciger India: Chathankod, Thiruvananthapuram, Kerala SDBDU 2006.4770 KU169935 KU169959 KU170009, KU169984 (1)
82 Polypedates otilophus Malaysia FMNH 239147 GQ204639 GQ204578 GQ204749, GQ204696 (1)
83 Raorchestes charius India: Coorg, Kodagu, Karnataka SDBDU 2011.814 KU169936 KU169960 KU170010, KU169985 (1)
84 Raorchestes glandulosus India: Coorg, Kodagu, Karnataka SDBDU 2011.817 KU169938 KU169962 KU170012, KU169987 (1)
85 Raorchestes griet India: Munnar, Idukki, Kerala SDBDU 2011.801 KU169941 KU169965 –, KU169990 (1)
86 Raorchestes gryllus Vietnam: Pac Ban, Tuyen Quang ROM 30288 GQ285796 GQ285777 GQ285674 (1)
87 Raorchestes jayarami India: Munnar, Idukki, Kerala SDBDU 2011.807 KU169940 KU169964 KU170013, KU169989 (1)
88 Raorchestes longchuanensis China: Longchuan, Yunnan 5RAO GQ285795 GQ285776 GQ285675 (1)
89 Raorchestes menglaensis China: Lvchun, Yunnan 060821286Rao GQ285797 GQ285778 GQ285676 (1)
90 Raorchestes resplendens India: Anamudi, Idukki, Kerala SDBDU 2009.1962 KU169939 KU169963 –, KU169988 (1)
91 Raorchestes signatus India: Avalanche, Nilgiris, Tamil Nadu SDBDU 2010.276 KU169937 KU169961 KU170011, KU169986 (1)
92 Raorchestes tinniens India: Munnar, Kerala SDBDU 2010.274 KU169942 KU169966 –, KU169991 (1)
93 Rhacophorus annamensis Vietnam FMNH 253934 GQ204653 GQ204598 GQ204768, GQ204717 (1)
94 Rhacophorus calcaneus Laos FMNH 256465 GQ204655 GQ204600 GQ204770, GQ204719 (1)
95 Rhacophorus kio China: Xishuangbanna, Yunnan SCUM 37941C EU215562 GQ285766 EU215532 (1)
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TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 9
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
96 Rhacophorus malabaricus India: Kalpetta, Wayanad, Kerala SDBDU 2007.6019 KU169943 KU169967 KU170014, KU169992 (1)
97 Rhacophorus pardalis Malaysia FMNH 231366 GQ204647 GQ204593 GQ204762, GQ204711 (1)
98 Rhacophorus reinwardtii Indonesia: Java ZRC 1.1.5273 GQ204656 GQ204601 GQ204771, GQ204720 (1)
99 Rhacophorus rhodopus China: Mengyang, Jinghong KIZ 060821037 EU215561 EU924532 EU215531 (1)
100 Rohanixalus baladika (1) Indonesia: Tapanuli, North Sumatra MZB Amph 23920 – , MW054232 (2)
101 Rohanixalus baladika (2) Indonesia: Tapanuli, North Sumatra MZB Amph 23951 – , MW054233 (1), (2)
102 Rohanixalus baladika (3) Indonesia: Rejanglebong, Bengkulu MZB Amph 26114 – , MW054234 (2)
103 Rohanixalus baladika (4) Indonesia: Berastagi, Karo, North Sumatra MZB Amph 31928 – , MW054235 (2)
104 Rohanixalus hansenae (1) Thailand: Nong Khor, Si Racha district, Chonburi
Prov.
CUMZ-A-7726 MW055952 MW055939 MW055965, MW054215 (1), (2)
105 Rohanixalus hansenae (2) Viet Nam: Buon Luoi, Kannack, Ankhe District,
Gia-Lai Province
FMNH 254444 DQ283835 DQ283134 (1), (2)
106 Rohanixalus hansenae Thailand: Nam Nao district, Phetchabun Prov. 0911Y2 – , KR827727 (2)
107 Rohanixalus hansenae Thailand: Nam Nao district, Phetchabun Prov. 0911Y1 – , KR827728 (2)
108 Rohanixalus hansenae Thailand: Kui Buri NP, Prachuap Khiri Khan Prov. 0933Y – , KR827729 (2)
109 Rohanixalus hansenae Thailand: Khao Ang Rui Ni WS, Chachoengsao
Prov.
0998Y – , KR827730 (2)
110 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAM 00613 – , KJ546836 (2)
111 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00619 – , KC357638 (2)
112 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00874 – , KC357640 (2)
113 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00870 – , KC357650 (2)
114 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00873 – , KC357651 (2)
115 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00875 – , KC357652 (2)
116 Rohanixalus hansenae Thailand: Chanthaburi Prov. ZMKUAM 00636 – , KJ546820 (2)
117 Rohanixalus hansenae Thailand: Chanthaburi Prov. ZMKUAM 00632 – , KJ546821 (2)
118 Rohanixalus hansenae Thailand: Chanthaburi Prov. ZMKUAM 00635 – , KJ546822 (2)
119 Rohanixalus hansenae Thailand: Nakhon Ratchasima Prov. ZMKUAM 00729 – , KJ546833 (2)
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BIJU ET AL.
10 · Zootaxa 4878 (1) © 2020 Magnolia Press
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
120 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAM 00689 – , KJ546834 (2)
121 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAM 00614 – , KJ546835 (2)
122 Rohanixalus hansenae Thailand: Kang Hang Meaw district, Chantaburi
Prov.
ZMKUAm00633 – , KC357639 (2)
123 Rohanixalus hansenae Thailand: Kang Hang Meaw district, Chantaburi
Prov.
ZMKUAm00634 – , KC357641 (2)
124 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00707 – , KC357642 (2)
125 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00711 – , KC357643 (2)
126 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00710 – , KC357644 (2)
127 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00718 – , KC357645 (2)
128 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00763 – , KC357646 (2)
129 Rohanixalus hansenae Thailand: Kang Hang Meaw district, Chantaburi
Prov.
ZMKUAm00712 – , KC357647 (2)
130 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00876 – , KC357648 (2)
131 Rohanixalus hansenae Thailand: Sriracha district, Chonburi Prov. ZMKUAm00877 – , KC357649 (2)
132 Rohanixalus hansenae Thailand: Nam Tok Hongkaew Prov. KUHE 34136 AB813161 (2)
133 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAm00971 – , KC357631 (2)
134 Rohanixalus hansenae Thailand: Nakhon Ratchasima Prov. ZMKUAM 00687 – , KJ546818 (2)
135 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAm00872 – , KC357632 (2)
136 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAm00977 – , KC357633 (2)
137 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAm00975 – , KC357634 (2)
138 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAm00974 – , KC357635 (2)
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TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 11
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
139 Rohanixalus hansenae Thailand: Phu Ruea district, Loei Prov. ZMKUAm00968 – , KC357636 (2)
140 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAm00972 – , KC357637 (2)
141 Rohanixalus hansenae Thailand: Nakhon Ratchasima Prov. ZMKUAM 01112 – , KJ546808 (2)
142 Rohanixalus hansenae Thailand: Nakhon Ratchasima Prov. ZMKUAM 00746 – , KJ546819 (2)
143 Rohanixalus hansenae Thailand: Loei Prov. ZMKUAM 00690 – , KJ546828 (2)
144 Rohanixalus hansenae Thailand: Chonburi Prov. ZMKUAM 00871 – , KJ546829 (2)
145 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAM 00723 – , KJ546831 (2)
146 Rohanixalus hansenae Thailand: Wang Nam Khieo district, Nakhon Ratch-
asima Prov.
ZMKUAM 00722 – , KJ546832 (2)
147 Rohanixalus hansenae Thailand: Surat Thani Prov. ZMKUAM 00949 – , KJ546827 (2)
148 Rohanixalus hansenae Thailand: Ban Ta Khun district, Surat Thani Prov. ZMKUAm00781 – , KC357625 (2)
149 Rohanixalus hansenae Thailand: Ban Ta Khun district, Surat Thani Prov. ZMKUAm00784 – , KC357626 (2)
150 Rohanixalus hansenae Thailand: Ban Ta Khun district, Surat Thani Prov. ZMKUAm00783 – , KC357627 (2)
151 Rohanixalus hansenae Thailand: Ban Ta Khun district, Surat Thani Prov. ZMKUAm00782 – , KC357628 (2)
152 Rohanixalus hansenae Thailand: Ban Ta Khun district, Surat Thani Prov. ZMKUAm00786 – , KC357629 (2)
153 Rohanixalus hansenae Thailand: Ban Ta Khun district, Surat Thani Prov. ZMKUAm00785 – , KC357630 (2)
154 Rohanixalus hansenae Thailand: Surat Thani Prov. ZMKUAM 00950 – , KJ546807 (2)
156 Rohanixalus hansenae Thailand: Sam Ngao district and Mueang district,
Tak Province
ZMKUAM 01121 – , KJ546809 (2)
157 Rohanixalus hansenae Thailand: Sam Ngao district and Mueang district,
Tak Province
ZMKUAM 01131 – , KJ546810 (2)
158 Rohanixalus hansenae Thailand: Sam Ngao district and Mueang district,
Tak Province
ZMKUAM 01114 – , KJ546811 (2)
159 Rohanixalus hansenae Thailand: Sam Ngao district and Mueang district,
Tak Province
ZMKUAM 01113 – , KJ546812 (2)
160 Rohanixalus hansenae Thailand: Sam Ngao district and Mueang district,
Tak Province
ZMKUAM 01130 – , KJ546813 (2)
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BIJU ET AL.
12 · Zootaxa 4878 (1) © 2020 Magnolia Press
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
161 Rohanixalus hansenae Thailand: Surat Thani Prov. ZMKUAM 00947 – , KJ546830 (2)
162 Rohanixalus cf. hansenae 1 (1) N.A. MNHN 1997.5453 AY880626 AY880579, AY880493 (1), (2)
163 Rohanixalus cf. hansenae 1 (2) China: Wenshan, Yunnan KIZ060821148 EF564448, EF564520 (1), (2)
164 Rohanixalus cf. hansenae 1 (3) Thailand: Sangkhla Buri KUHE 19441 AB813162 (1), (2)
165 Rohanixalus cf. hansenae 1 (4) China: Hainan Province 0031H KU840691 KU840481, KU840543 (1), (2)
166 Rohanixalus cf. hansenae 1Thailand: Mae Hong Son Prov. ZMKUAM 00896 – , KJ546838 (2)
167 Rohanixalus cf. hansenae 1Thailand: Mueang district, Mae Hong Son Prov. ZMKUAM 00678 – , KJ546837 (2)
168 Rohanixalus cf. hansenae 1Thailand: Mae Hong Son Prov. ZMKUAM 00681 – , KJ546826 (2)
169 Rohanixalus cf. hansenae 1Thailand: Kanchanaburi Prov. ZMKUAM 00825 – , KJ546825 (2)
170 Rohanixalus cf. hansenae 1Thailand: Mueang district, Mae Hong Son Prov. ZMKUAM 00682 – , KJ546824 (2)
171 Rohanixalus cf. hansenae 1Thailand: Mae Hong Son Prov. ZMKUAM 00667 – , KJ546823 (2)
172 Rohanixalus cf. hansenae 1Thailand: Mae Sod district, Tak Province ZMKUAM 01115 – , KJ546817 (2)
173 Rohanixalus cf. hansenae 1Thailand: Mae Sod district, Tak Province ZMKUAM 01117 – , KJ546816 (2)
174 Rohanixalus cf. hansenae 1Thailand: Mae Sod district, Tak Province ZMKUAM 01116 – , KJ546815 (2)
175 Rohanixalus cf. hansenae 1Thailand: Mae Sod district, Tak Province ZMKUAM 01118 – , KJ546814 (2)
176 Rohanixalus cf. hansenae 1Thailand: Kanchanaburi Prov. ZMKUAm00832 – , KC357669 (2)
177 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00819 – , KC357668 (2)
178 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00985 – , KC357667 (2)
179 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00818 – , KC357665 (2)
180 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00803 – , KC357664 (2)
181 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00828 – , KC357663 (2)
182 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00800 – , KC357662 (2)
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TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 13
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
183 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00799 – , KC357661 (2)
184 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00805 – , KC357660 (2)
185 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00806 – , KC357659 (2)
186 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00802 – , KC357658 (2)
187 Rohanixalus cf. hansenae 1Thailand: Thong Pha Phum district, Kanchanaburi
Prov.
ZMKUAm00831 – , KC357657 (2)
188 Rohanixalus cf. hansenae 1Thailand: Mueang district, Mae Hong Son Prov. ZMKUAm00672 – , KC357656 (2)
189 Rohanixalus cf. hansenae 1Thailand: Mueang district, Mae Hong Son Prov. ZMKUAm00671 – , KC357655 (2)
190 Rohanixalus cf. hansenae 1Thailand: Mueang district, Mae Hong Son Prov. ZMKUAm00668 – , KC357654 (2)
191 Rohanixalus cf. hansenae 1Thailand: Mueang district, Mae Hong Son Prov. ZMKUAm00670 – , KC357653 (2)
192 Rohanixalus cf. hansenae 1Vietnam: Dat, Lang Son K895 – , KR827725 (2)
193 Rohanixalus cf. hansenae 1Thailand: Huay Yang NP, Prachuap Khiri Khan
Prov.
0089Y – , KR827736 (2)
194 Rohanixalus cf. hansenae 1Thailand: Thung Sung social forest, Krabi Prov. 0304Y – , KR827737 (2)
195 Rohanixalus cf. hansenae 1Laos: Nathen, Phongsaly 2005.0109 – , KR827726 (2)
196 Rohanixalus cf. hansenae 2Thailand: Phu Hin Rong Kla NP, Phitsanulok Prov. 0901Y – , KR827731 (2)
197 Rohanixalus cf. hansenae 2Thailand: Thung Salaeng Luang NP, Phetchabun
Prov.
0133Y – , KR827732 (2)
198 Rohanixalus cf. hansenae 2Thailand: Thung Salaeng Luang NP, Phetchabun
Prov.
0013Y – , KR827733 (2)
199 Rohanixalus cf. hansenae 2Thailand: Nam Pad district, Uttaradit Prov. 0046Y – , KR827734 (2)
200 Rohanixalus cf. hansenae 2Thailand: Thung Salaeng Luang NP, Phetchabun
Prov.
0179Y – , KR827735 (2)
201 Rohanixalus senapatiensis (1) India: Manipur SDBDU 41657 MW055953 MW055940 MW055966, MW054216 (1), (2)
202 Rohanixalus senapatiensis (2) India: Manipur SDBDU 41658 MW055954 MW055941 MW055967, MW054217 (1), (2)
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BIJU ET AL.
14 · Zootaxa 4878 (1) © 2020 Magnolia Press
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
203 Rohanixalus senapatiensis (3) India: Nagaland SDBDU 41659 MW055955 MW055942 MW055968, MW054218 (1), (2)
204 Rohanixalus senapatiensis (4) India: Manipur SDBDU 41660 MW055956 MW055943 MW055969, MW054219 (1), (2)
205 Rohanixalus senapatiensis (5) India: Manipur SDBDU 41661 MW055957 MW055944 MW055970, MW054220 (1), (2)
206 Rohanixalus shyamrupus (1) India: Manipur SDBDU 41662 MW055958 MW055945 MW055971, MW054221 (1), (2)
207 Rohanixalus shyamrupus (3) India: Arunachal Pradesh SDBDU 41663 MW055959 MW055946 MW055972, MW054222 (1), (2)
208 Rohanixalus shyamrupus (4) India: Arunachal Pradesh SDBDU 41664 MW055960 MW055947 MW055973, MW054223 (1), (2)
209 Rohanixalus shyamrupus (5) India: Nagaland SDBDU 41665 – , MW054224 (2)
210 Rohanixalus shyamrupus (6) India: Nagaland SDBDU 41666 – , MW054225 (2)
211 Rohanixalus shyamrupus (7) India: Nagaland SDBDU 41667 – , MW054226 (2)
212 Rohanixalus shyamrupus (8) India: Nagaland SDBDU 41668 – , MW054227 (2)
213 Rohanixalus shyamrupus (9) India: Nagaland SDBDU 41669 – , MW054228 (2)
214 Rohanixalus cf. shyamrupus
(1)
Myanmar: Putao District, Kachin State CAS HERP 221212 –, KC692874 (1), (2)
215 Rohanixalus cf. shyamrupus
(2)
Myanmar: Putao District, Kachin State CAS HERP 221213 –, KC692876 (2)
216 Rohanixalus cf. shyamrupus
(3)
Myanmar: Putao District, Kachin State CAS HERP 244065 –, KC692875 (2)
217 Rohanixalus vittatus (1) Myanmar: Myitkyina District, Kachin state CAS HERP 241274 – , KC692881 (1), (2)
218 Rohanixalus vittatus (2) India: Rangat, Middle Andaman, Andaman and
Nicobar Islands
SDBDU 2019.4032 MW055961 MW055948 MW055974, MW054229 (1), (2)
219 Rohanixalus vittatus Thailand: Sangkhla Buri district, Kanchanaburi
Prov.
ZMKUAm00981 – , KC357620 (2)
220 Rohanixalus vittatus Thailand: Sangkhla Buri district, Kanchanaburi
Prov.
ZMKUAm00749 – , KC357621 (2)
221 Rohanixalus vittatus Thailand: Sangkhla Buri district, Kanchanaburi
Prov.
ZMKUAm00980 – , KC357622 (2)
222 Rohanixalus vittatus Thailand: Sangkhla Buri district, Kanchanaburi
Prov.
ZMKUAm00978 – , KC357623 (2)
......continued on the next page
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 15
TABLE 1. (Continued)
S.No Taxa Collection Locality Voucher Number GenBank Accession Numbers Analysis
RHO RAG1 12SrRNA, tRNA
VAL, 16SrRNA
223 Rohanixalus vittatus Thailand: Sangkhla Buri district, Kanchanaburi
Prov.
ZMKUAm00979 – , KC357624 (2)
224 Rohanixalus vittatus Myanmar: Dewei District, Tanintharyi Division CAS HERP 245916 – , KC692877 (2)
225 Rohanixalus vittatus Myanmar: Dewei District, Tanintharyi Division CAS HERP 245724 – , KC692878 (2)
226 Rohanixalus vittatus Myanmar: Dewei District, Tanintharyi Division CAS HERP 245915 – , KC692879 (2)
227 Rohanixalus vittatus Myanmar: Dewei District, Tanintharyi Division CAS HERP 245922 – , KC692880 (2)
228 Rohanixalus vittatus Myanmar: Myitkyina District, Kachin State CAS HERP 241113 – , KC692882 (2)
229 Rohanixalus sp. 1 (1) China NMNS 3184 AF458131 (1), (2)
230 Rohanixalus sp. 1 (2) China: Simao, Yunnan KIZ 060821090 EU924540 EU924512 EF564447, EF564519 (1), (2)
231 Rohanixalus sp. 1 (3) China: Yunnan, China KIZ 0001 Rao GQ285793 GQ285774 GQ285684 (1), (2)
232 Rohanixalus sp. 2 (4) Myanmar: Naung Layan, Loi Mwe WS, Kyaitong
Township
CAS 235551 MW055962 MW055949 MW055975, MW054230 (1), (2)
233 Rohanixalus sp. 2 (5) Laos: Viengthong district, Huaphahn Prov. FMNH 255217 GQ204658 GQ204603 GQ204773, GQ204722 (1), (2)
234 Taruga eques Sri Lanka WHT 2741 GQ204633 GQ204571 AY141801, GQ204689 (1)
235 Taruga fastigo Sri Lanka WHT 2783 GQ204634 GQ204572 AY141802, GQ204690 (1)
236 Taruga longinasus Sri Lanka WHTKAN1 GQ204635 GQ204573 GQ204745, GQ204691 (1)
237 Theloderma albopunctatum China: Jinping, Yunnan KIZ 060821201 EU924562 EU924534 EF564449, EF564521 (1)
238 Theloderma asperum Malaysia ZRC 1.1.9321 GQ204661 GQ204606 GQ204776, GQ204725 (1)
239 Theloderma moloch India: Arunachal Pradesh SDBDU 2011.345 KU169944 KU169968 KU170015, KU169993 (1)
240 Zhangixalus dugritei China: Baoxing, Sichuan SCUM 051001L EU215571 GQ285768 EU215541 (1)
241 Zhangixalus dennysi China: Jinxiu, Guangxi KIZ 060821050 EU924548 EU924520 EF564467, EF564539 (1)
242 Zhangixalus nigropunctatus China: Weining, Guizhou SCUM 070657L EU215563 GQ285767 EU215533 (1)
243 Zhangixalus smaragdinus China: Simao, Yunnan KIZ 060821140 EU924552 EU924524 EF564476, EF564548 (1)
Outgroup
244 Limnonectes magnus Philippine Islands 965 DQ347373 DQ347252 DQ347022, DQ347314 (1)
BIJU ET AL.
16 · Zootaxa 4878 (1) © 2020 Magnolia Press
External morphology. The new collections from India were morphologically compared with all the avail-
able Indian types and other museum specimens. Species reported from regions outside India were studied through
examination of types, original descriptions, and/or topotypes. Only adult individuals were used for morphometric
studies. Maturity and sex of the specimens were determined by examination of gonads through a small lateral or
ventral incision. 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 digital Vernier calipers or a binocular micro-
scope with an ocular micrometer: 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), SL (snout length, from the tip of the snout to the anterior
orbital border), EL (eye length, horizontal distance between the bony orbital borders), EN (distance from the front
of the eye to the nostril), NS (distance from the nostril to the tip of the snout), IN (internarial distance), TYD (great-
est tympanum diameter), TYE (distance from the tympanum to the back of the eye), 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), TFOL (total foot length, from heel to tip
of fourth toe). All measurements provided in the taxonomy section are in millimetres. Webbing is described as in
Biju et al. (2014b).
Larval study. The identity of the tadpoles was genetically confirmed based on barcoding of a 16SrRNA gene
fragment. Morphologically, the tadpoles were staged using Gosner (1960). The external morphology was described
following Grillitsch et al. (1993) and McDiarmid & Altig (1999). The following measurements were taken to the
nearest 0.01 mm using a digital Vernier caliper or a binocular microscope with an ocular micrometer: total length
(tl), snout-vent length (svl), distance from tip of snout to insertion of upper tail fin (su), maximum height of body
(bh), maximum width of body (bw), maximum diameter of eye (ed), interpupular distance (pp), internarial distance
(nn), naro-pupular distance (np), rostro-narial distance (rn), distance from tip of snout to opening of spiracle (ss),
distance from vent to tip of tail (vt), maximum height of tail (ht), tail muscle height (tmh), tail muscle width (tmw);
and oral disc width (odw).
Bioacoustics. Male vocalizations were recorded on Marantz PMD620 solid-state digital recorder (44.1 kHz
sampling rate, 16-bit resolution) with the aid of Sennheiser ME 66 unidirectional microphone positioned at about
30–50 cm from the calling individual. Dry and wet bulb air temperatures at the recording site were noted. For acous-
tic analysis, five temporal properties (call duration, call rise time, call fall time, number of pulses per call, and pulse
rate) and one spectral property (dominant frequency) for a single call of each species were measured using Raven
Pro 1.4 (Charif et al. 2010). Acoustic terminology and property definitions follow Bee et al. (2013).
Abbreviations. Abbreviations for museums and frequently used terms are as follows: SDBDU (Systematics
Lab, University of Delhi, India); NHM (Natural History Museum, London, United Kingdom) formerly BMNH
(British Museum of Natural History, London, United Kingdom); ZSIC (Zoological Survey of India, Kolkata, India);
CUMZ (Chulalongkorn University Museum of Natural History, Thailand); CIB (Herpetological Museum, Chengdu
Institute of Biology, Chinese Academy of Sciences, Chengdu, China); MZB (Museum Zoologicum Bogoriense,
Research Center for Biology, Indonesian Institute of Sciences, Indonesia); BORNEENSIS (Institute for Tropical
Biology and Conservation, University of Malaysia, Sabah; also known as BORN); CAS (California Academy of
Sciences, USA); LSUHC (La Sierra University Herpetological Collection, USA); MSNG (Museo Civico di Storia
Naturale di Genova, Italy); USNM (National Museum of Natural History, Division of Amphibians and Reptiles,
USA); ZRC (Zoological Reference Collection, Department of Zoology, University of Singapore, Singapore);
ZSI-E (Zoological Survey of India, Eastern Regional Station, Shillong, India).
Results
Phylogenetic relationships. The multi-gene phylogenetic analyses aimed to investigate the relationship of various
species previously assigned to Chirixalus and Feihyla, with respect to all other known genera of the subfamily Rha-
cophorinae. The recovered genus-level relationships were largely congruent with previously published phylogenies
(e.g., Yu et al. 2009; Meegaskumbura et al. 2015; Biju et al. 2016; Li et al. 2009, 2013; Hertwig et al. 2013; Chan
et al. 2018; Chen et al. 2020) (Fig. 1). An exception, however, was the relationship of various members of ‘Feihyla
and ‘Chirixalus’. Several studies have shown the paraphyletic relation of Feihyla with respect to a monophyletic
Chirixalus + Chiromantis or Chiromantis sensu lato (e.g., Wilkinson et al. 2002; Frost et al. 2006; Li et al. 2008; Yu
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 17
et al. 2009; Meegaskumbura et al. 2015; Poyarkov et al. 2015; Biju et al. 2016). Our analyses suggest members of
the morphologically recognised Feihyla palpebralis group and Feihyla vittiger group (see morphological grouping
of species under the taxonomic results) to have a sister-group relationship, albeit with weak support. On the other
hand, two members previously assigned to Feihyla’ and four to Chirixalus’ form another distinct and well sup-
ported lineage (formally described as Rohanixalus gen. nov. under taxonomic results), that has a moderately sup-
ported sister relationship with Chirixalus + Chiromantis. This is also congruent with the phylogenetic position of F.
vittata shown previously (e.g., Frost et al. 2006; Yu et al. 2009; Hertwig et al. 2012; Meegaskumbura et al. 2015;
Poyarkov et al. 2015; Biju et al. 2016). Certain studies utilising a larger number of loci but from only a few selected
taxa (e.g., Li et al. 2009, 2013; Hertwig et al. 2013; Chan et al. 2018; Chen et al. 2020) have earlier suggested
Feihyla (as previously understood) to be a monophyletic group. However, our study showed a closer but an unre-
solved to weakly-supported relationship of the genus Feihyla with the clade containing Ghatixalus + (Polypedates
+ Taruga), rather than the genera Chirixalus, Chiromantis, and Rohanixalus gen. nov. Increased sampling from the
genus Feihyla could further resolve its exact phylogenetic affinities. Nonetheless, in the light of largely unresolved
monophyly of the genus Feihyla, recognition of a new genus for the clade containing several previously attributed
Feihyla and Chirixalus members provides a phylogenetic resolution to the ChirixalusChiromantisFeihyla com-
plex and enables taxonomic stability that has been long lacking at the species level.
Within the new genus, all the five known species are phylogenetically distinct. Rohanixalus baladika has a
basal relationship compared to the other four members (R. hansenae, R. senapatiensis, R. shyamrupus, and R. vit-
tatus) and their closely related unidentified lineages (R. cf. shyamrupus, Rohanixalus sp. 1, Rohanixalus sp. 2, R.
cf. hansenae 1, and R. cf. hansenae 2). Among R. hansenae, R. senapatiensis, R. shyamrupus, and R. vittatus, the
latter three are more closely related to each other and broadly constitute the R. vittatus species group. Within this
group, two unidentified sister lineages, Rohanixalus sp. 1 and Rohanixalus sp. 2, show a close relationship with
R. senapatiensis + R. shyamrupus, rather than R. vittatus. On the other hand, R. hansenae, along with unidentified
lineages contained therein, constitute the R. hansenae species group.
In the genus Feihyla, the F. palpebralis species group, which represents Feihyla sensu stricto (Frost et al. 2006),
comprises F. palpebralis from Vietnam and populations representing F. fuhua from China. On the other hand, the
F. vittiger species group includes three species, F. kajau, F. inexpectata, and F. vittiger; the relationships within and
between the two groups, however, are yet not fully resolved.
Altogether, our results provide evidence for genus-level recognitions on the basis of phylogeny. This also facili-
tates the generic placement of previously confused taxa, while reiterating the need for future studies to add molecu-
lar data from all the members known across these genera, in order to comprehensively establish the species-level
relationships proposed based on our morphological studies.
DNA barcoding. The genus Feihyla included five species-level clades corresponding to four known species
(F. fuhua, F. inexpectata, F. kajau, and F. palpebralis) and one previously assigned to Chirixalus (hereafter, Feihyla
vittiger comb. nov.). Two major morphologically-identified species groups were shown to be phylogenetically
distinct: the Feihyla palpebralis group, with F. palpebralis and F. fuhua; and the Feihyla vittiger group with F.
inexpectata, F. kajau, and F. vittiger (Fig. 2a; Table 2). The analyses for genus Rohanixalus gen. nov. recovered 10
clades corresponding to two species previously attributed to the genus Feihyla (hereafter, Rohanixalus hansenae
comb. nov. and R. vittatus comb. nov.), three species previously assigned to Chirixalus (hereafter, Rohanixalus
baladika comb. nov., R. senapatiensis comb. nov., and R. shyamrupus comb. nov.), and five potentially unidenti-
fied lineages (R. cf. shyamrupus, Rohanixalus sp. 1, Rohanixalus sp. 2, R. cf. hansenae 1, and R. cf. hansenae 2).
Genetic identities at the species-level were determined by the phylogenetic position of topotypic material, except
for R. vittatus (see remarks under the taxonomic account of R. vittatus). Among the known species, the phylogenetic
position of Rohanixalus baladika, R. senapatiensis, R. shyamrupus, and Feihyla vittiger is clarified for the first time
in the study (Fig. 2b).
Based on the observed trend in genetic divergences among known species of the genus Rohanixalus, the mini-
mum inter-specific distance observed between R. senapatiensis and R. shyamrupus was used as a threshold criterion
and initial evidence for delineating potentially distinct evolutionary lineages or putative candidate species within the
genus (Fig. 2b; Tables 3). High levels of genetic divergence were observed across populations of R. hansenae and
R. vittatus. Although R. hansenae sensu lato was recovered as a well-supported monophyletic unit, one or two addi-
tional divergent lineages were observed. Several GenBank sequences identified as ‘Chirixalus vittatus’, Chiroman-
tis vittatus or ‘Feihyla vittata’ also nested within the various Rohanixalus hansenae sub-clades. On the other hand,
BIJU ET AL.
18 · Zootaxa 4878 (1) © 2020 Magnolia Press
sequences representing R. vittatus nested closer to R. senapatiensis and R. shyamrupus. Of these, Rohanixalus sp. 1
(‘vittatus from Yunnan, China) and Rohanixalus sp. 2 (‘vittatus’ from Myanmar) were more closely related to the
latter two species, whereas R. vittatus (from Myanmar + Thailand + Andamans, India) formed a sister relationship
with the rest (R. shyamrupus, R. senapatiensis, R. cf. shyamrupus, Rohanixalus sp. 1, and Rohanixalus sp. 2). De-
tailed comparisons of inter- and intraspecific genetic distances among the various known and unidentified lineages
are provided in Table 3.
TABLE 2. Inter and intraspecific uncorrected p-distances (in percent) in the genus Feihyla for the mitochondrial 16S
rRNA gene sequences. Values indicate average distances followed by range in parentheses.
Taxa 1 2 3 4 5
1Feihyla inexpectata 0.4
2Feihyla kajau 10.9
(10.7–11.1)
0.3
(0–0.6)
3Feihyla palpebralis 13.9 12.1
(11.9–12.1)
4Feihyla fuhua 12.1
(12.0–12.3)
9.8
(9.6–10.2)
11.0
(10.9–11.1)
0.1
(0–0.2)
5Feihyla vittiger 9.7
(9.5–9.9)
12.7
(12.6–12.8)
13.7 13.1
(13.0–13.2)
Taxonomic treatment
Class Amphibia Linnaeus, 1758
Order Anura Fischer von Waldheim, 1813
Family Rhacophoridae Hoffman, 1932
Subfamily Rhacophorinae Hoffman, 1932
I. Systematic review of genus Feihyla
Genus Feihyla
Original description. Feihyla Frost, Grant, Faivovich, Bain, Haas, Haddad, de S, Channing, Wilkinson, Donnellan,
Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch, Green, and Wheeler 2006. The amphibian tree of
life. Bulletin of the American Museum of Natural History, 297: 1–370.
Type species. Philautus palpebralis Smith, 1924 [= Feihyla palpebralis (Smith, 1924)]
Common name. Jelly-nest Tree Frogs
Etymology. The gender of this generic name is feminine. According to the original description, the generic no-
men of Feihyla is derived from Fei (the last name of Chinese herpetologist, Prof. Fei Liang) + hyla (Greek: vocative
form of Hylas, a traditional generic root for treefrogs) to honour the former’s extensive contributions to Chinese
herpetology (Frost et al. 2006).
Diagnosis. Small to medium-sized tree frogs (male SVL 17–28 mm, female SVL 23–34 mm); body rather
slender in appearance; skin shagreened to sparsely granular; a white streak extends along the upper lip margins,
either from below the eye up to the shoulder (in Feihyla palpebralis group) or from snout tip to the groin (in Feihyla
vittiger group); vomerine teeth absent; first two fingers opposable to the others; fingers free of web or with rudimen-
tary webbing; tips of fingers enlarged into discs with circum-marginal grooves; moderate webbing between toes, up
to or just above the third subarticular tubercle on either side of toe IV; eggs laid in terrestrial jelly nests (Fig. 3).
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 19
FIGURE 1. Maximum Likelihood tree showing phylogenetic relationships among 124 representative taxa from all recognised
genera of the subfamily Rhacophorinae. Relationships are inferred based on 1,937 bp of mitochondrial (12SrRNA, tRNAVAL,
16SrRNA) and nuclear (RHO and RAG1) genes. Numbers above and below the branches indicate Bayesian Posterior Probabili-
ties and RAxML bootstrap support values, respectively.
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20 · Zootaxa 4878 (1) © 2020 Magnolia Press
FIGURE 2. A–B. DNA Barcoding based on mitochondrial 16S rRNA sequences: A. genus Feihyla; B. genus Rohanixalus gen.
nov.; C. distribution of the new genus Rohanixalus. Images represent the corresponding species and their broad geographical
ranges.
Comparison. Genus Feihyla differs from the morphologically related genera Chirixalus and Chiromantis by its
slender body (vs. slender to robust); absence of vomerine teeth (vs. present); eggs laid in jelly nest (vs. foam nest)
(Fig. 3).
Distribution. Genus Feihyla currently has a disjunct distribution in southern China (Guangxi, Guizhou, and
Yunnan provinces) and possibly adjoining Vietnamese regions (Lao Cai Province), northern Vietnam (Tam Dao) to
central Vietnam (Lam Dong Province), and the southeast Asian islands of Borneo and Java (Fig 4).
Diversity. Prior to this study, there were six nominal taxa in the genus Feihyla (Frost et al. 2006; Aowphol et
al. 2013; Chan et al. 2018 ; Li et al. 2013; Fei et al. 2010). Based on the revised classification herein, two species
are transferred from Feihyla to Rohanixalus, while another two Chirixalus species are allocated to Feihyla (see
taxonomic remarks under the respective species accounts). Hereafter, the six currently recognised Feihyla members
are: F. fuhua, F. inexpectata, F. kajau, F. palpebralis, F. samkosensis comb. nov., and F. vittiger comb. nov.
Morphological grouping of species. Based on phenotypic traits, the present study identifies the following two
major species groups in this genus: (1) Feihyla palpebralis group (F. fuhua and F. palpebralis), and (2) Feihyla vit-
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 21
tiger group (F. inexpectata, F. kajau, F. samkosensis comb. nov., and F. vittiger comb. nov.). Taxonomic accounts
of all the recognised Feihyla species provided below are arranged by species-group.
FIGURE 3. Nest structure and egg clutch morphology in the closely related Asian genera Chirixalus, Feihyla, and Rohanixalus.
A. Foam nest in genus Chirixalus: an amplected pair of C. simus on a freshly laid foam nest; B. bubble nest in genus Rohanixa-
lus: R. vittatus female with a freshly laid egg clutch; C. jelly-nest in genus Feihyla: eggs of Feihyla fuhua; D. schematic illustra-
tion of egg clutch morphology in genus Rohanixalus; E. schematic illustration of egg clutch morphology in Feihyla palpebralis
group (Feihyla sensu stricto); F. schematic illustration of egg clutch morphology in Feihyla vittiger group; G. schematic illustra-
tion of egg clutch morphology in genus Chirixalus.
Feihyla palpebralis group
This group can be distinguished from the Feihyla vittiger group by the following suite of morphological characters:
medium-sized adults (SVL 25–34 mm); snout nearly pointed in dorsal view; first two fingers opposed to the others;
dorsum light yellowish-brown to reddish-brown with contrasting concave bands forming a ‘)(’-shaped marking or
faint discontinuous lines; a white streak along the upper lip margins, extending from the snout tip or below the eye
up to the shoulder; a weakly-developed dermal fringe along the outer margins of the hind limbs; whitish eggs with
light pigmentation on poles (Fig. 5).
Species included. Feihyla fuhua Fei, Ye, and Jiang, 2010 and Feihyla palpebralis (Smith, 1924).
Feihyla fuhua Fei, Ye, and Jiang, 2010
White-cheeked Jelly-nest Frog
(Figs. 1–5; Tables 1–2)
Original name and description. Feihyla fuhua Fei, Ye, and Jiang, 2010. Fei L., Ye C.Y., and Jiang J.P. 2010. A
new species of Rhacophoridae from Yunnan, China (Amphibia, Anura). Acta Zootaxonomica Sinica, 35: 413–417
BIJU ET AL.
22 · Zootaxa 4878 (1) © 2020 Magnolia Press
(in Chinese). Type. Holotype, CIB 584168, by original description. Type locality. “Qianjinxiang, Pingbian County
(22.58° N, 103.41° E, alt. 1040 m), Yunnan Province”, China. Current status of specific name. Valid name as
Feihyla fuhua Fei, Ye, and Jiang, 2010.
Diagnosis. Medium adult size (male SVL 25–29 mm); snout nearly pointed in dorsal view; dorsum shagreened
to sparsely granular; lateral surfaces of the head, including tympanic region, darker than dorsal colouration; a promi-
nent white or off-white streak extending from below the eye to the shoulder or beyond up to the groin; dorsum light
greyish-yellow or yellowish-brown, with contrasting reddish-brown concave bands forming a ‘)(‘-shaped marking;
presence of a pale white or cream pre-and post-orbital stripe, and a thin continuous or discontinuous reddish-brown
band between the eyes; first two fingers opposed to the others; foot webbing moderate, up to the second subarticular
tubercle on either side of toe IV (Fig. 5).
Distribution. Feihyla fuhua is currently known only from western and south-eastern Yunnan, western Guangxi
Province, and extreme western Guizhou Province in southern China. This species could be present in Vietnamese
regions (Lao Cai Province) (Fig. 4). Previous reports of ‘F. palpebralis’ from China are likely to refer to this spe-
cies (see remarks under F. palpebralis).
FIGURE 4. Distribution of genetically confirmed records of all known Feihyla and Rohanixalus members (indicated by circles)
and their respective type localities (indicated by triangles). Numbers corresponding to each species are indicated alongside the
distribution spots.
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 23
FIGURE 5. Members of the Feihyla palpebralis group and their nest structure. A–E. Feihyla fuhua from Mt. Dawei, Yunnan
Province, China. A. Dorsolateral view; B. lateral view (arrow indicating a white streak); C. dorsal view; D. freshly laid eggs
in a jelly-nest (arrow indicating pigmented pole); E. developing embryos; F–G. Feihyla palpebralis from the type locality,
Langbian Plateau, Lam Dong Province, Vietnam: F. dorsolateral view; G. freshly laid eggs in a jelly-nest (arrow indicating
pigmented pole); H–I. Feihyla fuhua from Maguan County, Yunnan Province, China: H. dorsolateral view (arrow indicating a
white streak); I. freshly laid eggs in a jelly-nest (arrow indicating pigmented pole).
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24 · Zootaxa 4878 (1) © 2020 Magnolia Press
Feihyla palpebralis (Smith, 1924)
Vietnamese Jelly-nest Frog
(Figs. 1–5; Tables 1–2)
Original name and description. Philautus palpebralis Smith, 1924. Smith M. A. 1924. New tree-frogs from Indo-
China and the Malay Peninsula. Proceedings of the Zoological Society of London, 1924: 225–234. Type. Holotype,
author’s number 2589, female (by original designation), presently NHM 1924.1.31.2. Type locality. “Langbian
Peaks, alt. 2000 m., S. Annam”, Vietnam. Current status of specific name. Valid name as Feihyla palpebralis
(Smith, 1924) (Frost et al. 2006).
Diagnosis. Medium adult size (male SVL 25–28 mm, female SVL 30 mm); snout pointed in dorsal view; dor-
sum shagreened to granular; white or light yellow streak extending from below the eye up to the shoulder; dorsum
light yellowish-brown or reddish-brown, occasionally with concave bands forming a ‘)(‘-shaped marking or faint
lines; presence of a pale white or cream pre- and post-orbital stripe, and continuous or discontinuous dark band be-
tween the eyes; limbs with pale cross-bands; first two fingers partially opposed to the others; rudimentary webbing
between the outer two fingers; foot webbing moderate, just below the second subarticular tubercle on either side of
toe IV (Fig. 5).
Distribution. Feihyla palpebralis is currently known only from the Langbian Peaks area of present-day central
Vietnam (Lam Dong Province) and probably northern Vietnam (Tam Dao). Previous studies reported this species
from southern China (Yunnan Province, Guangxi Province, and Guizhou Province) (e.g., Fei 1999; Fei et al. 2009,
2010). However, after recent examination of Chinese Feihyla specimens, including the type series of F. fuhua, we
found none that could be morphologically assigned to F. palpebralis. The original description of F. fuhua also did
not delist the F. palpebralis’ record from China. Hence, ‘F. palpebralis’ from China is considered to be misidentifi-
cation of F. fuhua (Fig. 4). Further studies are required to understand the geographical ranges of these two species.
Feihyla vittiger group
This group can be distinguished from the Feihyla palpebralis group by the following suite of morphological char-
acters: small to medium-sized adults (SVL 17–27 mm); snout nearly truncate in lateral view; the first two fingers
opposable to the others; webbing absent between fingers, except for a rudiment of web between fingers III and
IV at the base; foot webbing moderate, extending up to the second subarticular tubercle on either side of toe IV. A
prominent and useful character for differentiating Feihyla vittiger group is the unique colouration of its members:
upper arm, loreal, canthal and tympanic regions, lateral surfaces of abdomen, and anterior and posterior parts of
thigh non-pigmented (flesh coloured); and a narrow white streak that starts from the snout tip and extends along
the lateral surfaces of the head from below the eye up to the groin, clearly separating the dorsal and lateral body
colouration (Fig. 6). Geographically, this group is currently restricted to the Southeast Asian islands of Borneo and
Java; no reports exist from mainland Asia.
Species included. Feihyla inexpectata (Matsui, Shimada, and Sudin, 2014), Feihyla kajau (Dring, 1983), Fei-
hyla samkosensis (Grismer, Thy, Chav, and Holden, 2007) comb. nov., and Feihyla vittiger (Boulenger, 1897)
comb. nov.
Feihyla inexpectata (Matsui, Shimada, and Sudin, 2014)
Bornean Jelly-nest Frog
(Figs. 1–2, 4, 6; Tables 1–2)
Original name and description. Chiromantis inexpectatus Matsui, Shimada, and Sudin, 2014. Matsui M., Shi-
mada T., and Sudin A. 2014. First record of the tree-frog genus Chiromantis from Borneo with the description of a
new species (Amphibia: Rhacophoridae). Zoological Science, 31: 45–51. Type. Holotype, BORNEENSIS 22421,
by original designation. Type locality. “Camel Trophy field station (4°54’ N, 116°53’ E; ca. 1050 m a.s.l.) of the
Maliau Basin Conservation Area, Sandakan Division, Sabah”, Malaysia. Current status of specific name. Valid
name as Feihyla inexpectata (Matsui, Shimada, and Sudin, 2014) (Chan et al. 2018).
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 25
FIGURE 6. Members of the Feihyla vittiger group and their nest structure. A. F. inexpectata in dorsolateral view; B. F. kajau in
dorsolateral view; C. eggs of F. kajau; D. early development in F. kajau at approximately stage 19; E. F. vittiger in dorsolateral
view; F. an egg clutch of F. vittiger with developing embryos (approximately stage 21); G–M. Feihyla vittiger in life: G–J.
dorsolateral view; K. dorsal view; L. posterior view of thighs; M. lateral view.
Taxonomic remarks. The species was originally described as Chiromantis inexpectatus and compared with
all the congeners, except Chiromantis vittiger (Boulenger, 1897) presumably because of its placement in the genus
Philautus at that time. Chan et al. (2018) transferred Chiromantis inexpectatus to Feihyla, phylogenetically based
on its sister relationship with Feihyla kajau. We find that F. inexpectata is indeed morphologically as well as biogeo-
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26 · Zootaxa 4878 (1) © 2020 Magnolia Press
graphically closer to F. vittiger than any other known member of the genus. The reproductive mode in F. inexpectata
is yet unknown, but based on its morphological and phylogenetic relationship, it is most likely a jelly-nest species.
Diagnosis. Small-sized adults (male SVL 22–23 mm), with a slender body; snout nearly truncate in dorsal view,
and truncate in lateral view; dorsal skin shagreened to sparsely granular; dorsal colouration light yellow to light or
dark brown with a reddish tinge; dorsum with scattered contrasting dark brown spots; upper arm, loreal, canthal
and tympanic regions, lateral surfaces of abdomen, and anterior and posterior parts of thigh non-pigmented (flesh
coloured); a narrow white streak starting from the snout tip and extending along the lateral surfaces up to the groin,
separates the dorsal and lateral body colouration; the first two fingers opposed to the others; foot webbing moderate,
up to or slightly beyond the second subarticular tubercle on either side of toe IV.
Genetic divergence. Phylogenetically, F. inexpectata is closely related to members of the Feihyla vittiger group
(Fig. 1). For the 16S mitochondrial gene, it differs by average uncorrected genetic distances of: 10.7–11.1% from F.
kajau and 9.5–9.9% from F. vittiger. For interspecific genetic distances with other congeners, see Table 2.
Distribution. Feihyla inexpectata is currently known only from its type locality in Sabah, on the island of Bor-
neo in eastern Malaysia.
Feihyla kajau (Dring, 1983)
White-eared Jelly-nest Frog
(Figs. 1–2, 4, 6; Tables 1–2)
Original name and description. Rhacophorus kajau Dring, 1983. Dring J. C. M. 1983. Some new frogs from Sar-
awak. Amphibia-Reptilia, 4: 103–115. Type. Holotype, BMNH 1978.1757, by original designation. Type locality.
“Gunung Mulu, Fourth Division, Sarawak”, Malaysia. Current status of specific name. Valid name as Feihyla
kajau (Dring, 1983) (Li et al. 2013).
Diagnosis. Small-sized adults (male SVL 18–20 mm) with a slender body; snout rounded to truncate in dorsal
view, and truncate in lateral view; dorsal skin shagreened to sparsely granular with scattered spinules; dorsal co-
louration uniformly light green with minute white or yellowish speckles; upper arm, loreal, canthal and tympanic
regions, lateral surfaces of abdomen, and anterior and posterior parts of thigh, non-pigmented (flesh coloured); pres-
ence of a narrow white streak starting from the snout tip and extending along the lateral surfaces up to the groin, that
separates the dorsal and lateral body colouration; the first two fingers opposed to the others; foot webbing moderate,
not beyond the second tubercle on either side of toe IV; narrow crenulate dermal ridges present along the outer edges
of hand, forearm, foot, tarsus, and transversely below the vent. In life, the skin over the belly is translucent making
the internal organs, including mature ova, visible (Dring 1983).
Genetic divergence. Phylogenetically, F. kajau is closely related to members of the Feihyla vittiger group (Fig.
1). For the 16S mitochondrial gene, it differs by average uncorrected genetic distances of: 10.7–11.1% from F. inex-
pectata and 12.6–12.8% from F. vittiger. For interspecific genetic distances with other congeners, see Table 2.
Distribution. Feihyla kajau is currently known only from the Borneo island: Brunei; Sabah to Sarawak of
Malaysia; and Kalimantan of Indonesia.
Feihyla samkosensis (Grismer, Thy, Chav, and Holden, 2007) comb. nov.
Samkos Jelly-nest Frog
Original name and description. Chiromantis samkosensis Grismer, Thy, Chav, and Holden, 2007. Grismer L.L.,
Thy N., Chav T., and Holden J. 2007. A new species of Chiromantis Peters 1854 (Anura: Rhacophoridae) from
Phnom Samkos in the northwestern Cardamom Mountains, Cambodia. Herpetologica, 63: 392–400. Type. Holo-
type, ZRC1.11896, by original designation. Type locality. “the eastern flank of Phnom Samkos, Pursat Province,
Cardamom Mountains, Cambodia, 12° 08.817’ N, 103° 08.067’ E”, “at 501 m”. Current status of specific name.
Valid name as Feihyla samkosensis (Grismer, Thy, Chav, and Holden, 2007) comb. nov.
Taxonomic remarks. This species is morphologically close to members of the Feihyla vittiger group due to its
green dorsal colouration, truncate snout, absence of vomerine teeth, presence of a thick white line along the upper
lip from below the midpoint of eye to the shoulder, absence of dorsolateral or dorsal stripes, presence of scattered
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 27
small dark spots on dorsum, toes more than one-third webbed, and ventral skin semi-transparent. Hence, we provi-
sionally propose Chirixalus samkosensis (Grismer, Thy, Chav, and Holden, 2007) to be transferred to genus Feihyla.
Further molecular evidence is required for confirmation of this generic placement.
Diagnosis. Small-sized adults (male SVL 22.9 mm, female SVL 23.6 mm); green dorsal colouration; truncate
snout; absence of vomerine teeth; presence of a thick white line along the upper lip from below the midpoint of eye
to the shoulder; absence of dorsolateral or dorsal stripes; presence of scattered small dark spots on dorsum; first two
fingers opposed to the others; presence of web between fingers III and IV; toes more than one-third webbed, and
ventral skin semi-transparent (Grismer et al. 2007).
Distribution. Feihyla samkosensis is currently known only from its type locality on the eastern flank of Phnom
Samkos, Pursat Province, Cardamom Mountains, Cambodia.
Feihyla vittiger (Boulenger, 1897) comb. nov.
Javan Jelly-nest Frog
(Figs. 1–2, 4, 6; Tables 1–2)
Original name and description. Ixalus vittiger Boulenger, 1897. Boulenger G.A. 1897. Descriptions of new Malay
frogs. Annals and Magazine of Natural History, 19: 106–108. Type. Holotype, BMNH 1947.2.7.58. Type locality.
“Pengalengan”, West Java, Indonesia. Current status of specific name. Valid name as Feihyla vittiger (Boulenger,
1897) comb. nov.
Taxonomic remarks. This species is morphologically close to Feihyla inexpectata and the same is also con-
firmed phylogenetically based on the new sequence data included in our study (Fig. 1). Furthermore, its reproduc-
tive mode (eggs deposited in jelly nests) (Fig. 6) is possibly also a phenotypic synapomorphy for the genus Feihyla
(Li et al. 2009; Hertwig et al. 2013; present study). Hence, we propose that Chiromantis vittiger (Boulenger, 1897)
be transferred to genus Feihyla.
Diagnosis. Small to medium-sized adults (male SVL 24–27 mm, female SVL 23–26 mm) with a slender body;
snout rounded to truncate in dorsal view, and truncate in lateral view; dorsal skin shagreened to sparsely granular,
with or without scattered spinules; dorsal colouration variable, ranging from light lemon yellow to light or dark
brown, with or without a reddish tinge; dorsum with or without contrasting dark spots and minute speckles, even
within the same population; upper arm, loreal, canthal and tympanic regions, lateral surfaces of abdomen, and an-
terior and posterior parts of thigh, non-pigmented (flesh coloured); a white streak starting from the snout tip and
extending along the lateral surfaces up to the groin, separates the dorsal and lateral body colouration; the first two
fingers opposed to the others; foot webbing moderate, up to the second tubercle on either side of toe IV.
Genetic divergence. Phylogenetically, F. vittiger is closely related to members of the Feihyla vittiger group
(Fig. 1). For the 16S mitochondrial gene, it differs by average uncorrected genetic distances of: 9.5–9.9% from F.
inexpectata and 12.6–12.8% from F. kajau. For interspecific genetic distances with other congeners, see Table 2.
Distribution. Feihyla vittiger is currently known only from the Indonesian island of Java: Bogor, Pengalengan,
and Mount Halimun-Salak National Park in West Java, and Banwumas in Central Java.
II. Description of a new genus
Genus Rohanixalus gen. nov.
Zoobank: urn:lsid:zoobank.org:act:C5E1F838-322E-4304-BB13-348FB40B10E5
Etymology. The genus is named after Rohan Pethiyagoda, in appreciation of his contributions to herpetological and
ichthyological studies in Asia. The generic epithet is derived from Rohan + genus name Ixalus Dumeìril and Bibron,
1841 (a traditional suffix in rhacophorid generic names). For nomenclatural purposes, the gender of this generic
name is masculine.
Common name. Rohan’s Tree Frogs
Type species. Ixalus vittatus Boulenger, 1887 (= Rohanixalus vittatus comb. nov.)
Phylogenetic definition. Rohanixalus gen. nov. consists of the most inclusive clade that contains Rohanixalus
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28 · Zootaxa 4878 (1) © 2020 Magnolia Press
vittatus but not the type species of any of the currently recognised rhacophorid genera (Beddomixalus bijui, Buer-
geria buergeri, Chirixalus doriae, Chiromantis xerampelina, Feihyla palpebralis, Nasutixalus jerdonii, Ghatixalus
variabilis, Gracixalus gracilipes, Kurixalus eiffingeri, Leptomantis bimaculatus, Liuixalus romeri, Mercurana my-
risticapalustris, Nyctixalus margaritifer, Philautus aurifasciatus, Polypedates leucomystax, Pseudophilautus tem-
poralis, Raorchestes glandulosus, Rhacophorus reinwardtii, Taruga fastigo, Theloderma leporosum, and Zhangixa-
lus dugritei) (Fig. 1).
Diagnosis. Rohanixalus gen. nov. can be distinguished from other rhacophorid genera by the combination
of following characters: small to medium-sized adults (SVL 18–29 mm); body rather slender in appearance; skin
shagreened to sparsely granular; tympanum indistinct or weakly developed; vomerine teeth absent; the entire dor-
sum, lateral surfaces, and dorsal surfaces of limbs covered with fine dark brown speckles, some speckles clumping
together to form dark irregular spots or blotches over the dorsum; a pair of prominent to faint, continuous or discon-
tinuous, contrasting light coloured dorsolateral stripes starting from the tip of the snout, extending over the upper
eyelid margins, and ending close to the vent on either side; groin and anterior and posterior parts of thigh without
any prominent markings or colouration; the first two fingers opposed to the others; webbing between fingers absent,
except a rudiment of web at the base between fingers III and IV; foot webbing moderate, not beyond the second
subarticular tubercle on either side of toe IV; eggs laid in terrestrial bubble nests; freshly laid eggs light green in
colour, unpigmented on poles (Figs. 3, 7); and egg-attendance as a possible behavioural synapomorphy.
FIGURE 7. Members of the genus Rohanixalus. A. Dorsolateral view of R. hansenae from the type locality, Nong Khor,
Thailand; B. dorsolateral view of R. marginis (holotype) from Perlis State Park chalets, Perlis, Peninsular Malaysia; C. dorsal
view of R. cf. nauli, from Batang Gadis National Park, North Sumatra, Indonesia with a jelly-nest; D. dorsolateral view of R.
cf. nauli, from Batang Gadis National Park, North Sumatra, Indonesia; E. dorsolateral view of R. cf. nauli, from Bukit Barisan
Forest Park, North Sumatra, Indonesia; F. dorsolateral view of R. punctatus (Paratype, JBS 9274) from Gwa, Rakhine State,
Myanmar (CC Public domain 3.0; AmphibiaWeb); G. dorsolateral view of R. vittatus from Rangat, Middle Andaman Islands,
India; H. dorsolateral view of R. senapatiensis from Manipur, India; I. dorsolateral view of R. shyamrupus from Namdapha,
Arunachal Pradesh.
Comparison. Rohanixalus gen. nov. differs from all other rhacophorid members by the presence of a pair of
contrasting light-coloured dorsolateral stripes (with variable degree of prominence, prominent to faint, continu-
ous or discontinuous) starting from the snout tip, extending over the upper eyelid margins, and ending close to the
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 29
vent on either side (vs. absent, except in Rhacophorus lateralis), presence of prominent and dense minute speckles
throughout the dorsal and lateral surfaces of the body (including dorsum, lateral surfaces, and dorsal surface of
limbs) (vs. absent), and freshly laid eggs light green (vs. creamy white, except in some members of the Feihyla
vittiger group). Further, Rohanixalus differs from Philautus, Pseudophilautus, and Raorchestes in having an exo-
trophic mode of development with free-swimming tadpoles (vs. endotrophic or direct developing larvae), and the
first two fingers opposed to the others (vs. not opposable); it differs from Beddomixalus, Chirixalus, Chiromantis,
Ghatixalus, Leptomantis, Polypedates, Rhacophorus, Taruga, and Zhangixalus by eggs laid in bubble nests (vs.
foam nests), and absence of vomerine teeth (vs. presence). Specifically, Rohanixalus also differs from the various
foam-nesting genera individually or collectively by: relatively smaller adult size, SVL < 30 mm (vs. larger, SVL
> 30 mm: SVL 37–67 mm in Beddomixalus; SVL 39–82 mm in Ghatixalus; SVL 30–80 mm in Leptomantis; SVL
37–85 mm in Polypedates; SVL 30–100 mm in Rhacophorus; SVL 32–71 mm in Taruga; SVL 30–120 mm in
Zhangixalus), and absence of calcar on the distal end of tibia (vs. present, except in some species of Polypedates);
differs from Chirixalus by absence of dorsal lines (vs. present); differs from Chiromantis by relatively smaller adult
size, SVL < 30 mm (vs. larger, SVL 35–90 mm), a rudiment of web on fingers III and IV (vs. more extensive), and
tympanum indistinct or weakly developed (vs. distinct); differs from Leptomantis by the first two fingers opposed
to the others (vs. not opposable), absence of calcar on the distal end of tibia (vs. present), and absence of webbing
between fingers, except a rudiment of web on fingers III and IV (vs. present). Further, Rohanixalus differs from
Gracixalus, Kurixalus, Nasutixalus, Nyctixalus, and Theloderma by the first two fingers opposed to the others (vs.
not opposed), breeding and oviposition on leaves (vs. phytotelm-breeding and oviposition on walls of tree holes,
water-filled cavities, crevices, or on ground), and eggs laid in bubble nests (vs. gel encapsulated eggs). Specifically,
Rohanixalus also differs from Gracixalus, Kurixalus, and Nasutixalus, by its body rather slender in appearance (vs.
robust); differs from Gracixalus by absence of dorsal markings (vs. presence of ‘X’-shaped, inverted ‘V’ or ‘Y’-
shaped marking) and absence of spines on upper eyelids (vs. present); differs from Kurixalus by absence of dermal
fringes on forearms and tarsus (vs. present), and absence of vomerine teeth (vs. present); differs from Nasutixalus by
relatively smaller adult size, < 30 mm (vs. larger, 37–48mm), absence of oophagous tadpoles (vs. present), and iris
without ‘X’ mark (vs. present); specifically also differs from Nyctixalus and Theloderma by absence of spines, as-
perities, tubercles, or warts on dorsal skin (vs. present), and tympanum indistinct or weakly developed (vs. distinct).
Further, Rohanixalus differs from Mercurana by relatively smaller adult size, SVL < 30 mm (vs. larger, SVL 33–41
mm), first two fingers opposed to the others (vs. non opposed), absence of vomerine teeth (vs. present), oviposi-
tion and early development on leaves (vs. ground, eggs mixed with mud in shallow ground pits), and eggs laid in
bubble nests (vs. gel encapsulated eggs); differs from Feihyla by eggs laid in bubble nests (vs. jelly nests), freshly
laid eggs light green and unpigmented (vs. creamy white with pigmentation on poles, except in some members of
Feihyla vittiger group), and absence of a prominent white streak along the upper lip margins from below the eye
up to shoulder (vs. present). Further, Rohanixalus differs from Liuixalus by eggs laid in bubble nests on leaves (vs.
aquatic eggs), absence of ‘X’ shaped mark on dorsum (vs. present), and toes moderately webbed (vs. rudimentary
webbing). Rohanixalus also differs from Buergeria by eggs laid in bubble nests on leaves (vs. aquatic eggs), body
slender in appearance (vs. rather robust), dorsal skin without tubercles, granules, or ridges (vs. present), and absence
of vomerine teeth (vs. present). Rohanixalus is thus distinguished from all 21 other currently recognised rhacopho-
rid genera by a combination of characters, chiefly with respect to external morphology, breeding behaviour, oviposi-
tion, and development mode.
Distribution. The genus Rohanixalus gen. nov. is restricted to South, Southeast, and East Asia, where it is
currently known from India: Northeastern Indian states (Arunachal Pradesh, Assam, Manipur, Nagaland, Mizoram,
and Tripura) and the Andaman Islands; Bangladesh: Sylhet Division; Myanmar; Thailand; Laos; Cambodia; China
(Yunnan, Guangxi, Hainan, and Tibet); Vietnam; Malaysia; and the Sumatran island of Indonesia (Fig. 4).
Diversity. Currently there are eight nominal taxa in the new genus: Rohanixalus baladika (Riyanto and Kur-
niati, 2014) comb. nov.; Rohanixalus hansenae (Cochran, 1927) comb. nov.; Rohanixalus marginis (Chan, Gris-
mer, Anuar, Quah, Grismer, Wood, Muin, and Ahmad, 2011) comb. nov.; Rohanixalus nauli (Riyanto and Kurniati,
2014) comb. nov.; Rohanixalus punctatus (Wilkinson, Win, Thin, Lwin, Shein, and Tun, 2003) comb. nov.; Roha-
nixalus senapatiensis (Mathew and Sen, 2009) comb. nov.; Rohanixalus shyamrupus (Chanda and Ghosh, 1989)
comb. nov.; Rohanixalus vittatus (Boulenger, 1887) comb. nov.
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FIGURE 8. A–I. Rohanixalus vittatus (male, SDBDU 2019.4048) in life, from Andaman Islands, India: A. dorsolateral view; B.
dorsal view; C. ventral view; D. posterior view of thighs; E. enlarged view of skin over the tympanic region showing rounded
speckles; F. ventral view of hand; G. ventral view of foot; H. schematic illustration of webbing on foot; I. lateral view; J–P. vari-
ations in body colour and skin texture in R. vittatus at Andaman Islands, India: J. dorsolateral view (male, SDBDU 2019.4033);
K. dorsolateral view showing light green coloured mature ova through translucent skin (female, SDBDU 2019.4051); L. dor-
solateral view showing light green coloured mature ova through translucent skin (female, SDBDU 2019.4034); M. dorsal view
(male, SDBDU 2019.4037); N. dorsal view (male, 2019.4037); O. ventral view (female, SDBDU 2019.4034); P. ventral view
(male, SDBDU 2019.4037); Q. sampling localities of R. vittatus at Middle and North Andamans in the present study.
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 31
Rohanixalus vittatus (Boulenger, 1887) comb. nov.
Striped Bubble-nest Frog
(Figs. 1–4, 7–13; Tables 1, 3, 4)
Original name and description. Ixalus vittatus Boulenger, 1887. Boulenger G. A. 1887. An account of the batra-
chians obtained in Burma by M.L. Fea of the Genoa Civic Museum. Annali del Museo Civico di Storia Naturale
di Genova, Serie 2, 5: 418–424. Type. Lectotype, MSNG 29397. Type locality. “Bhamò” (= Bhamo, Myanmar).
Current status of specific name. Valid name as Rohanixalus vittatus (Boulenger, 1887) comb. nov.
Taxonomic remarks. Rohanixalus vittatus (Boulenger, 1887), the type species of the genus, was originally
described from “Bhamò” (= Bhamo), based on the collection of Col. M. L. Fea. Although Boulenger (1887) did not
provide further specific details concerning the collection locality of this taxon, his understanding of “Bhamo, as
used throughout the work [e.g., “Bhamò...(Upper Irawaddy)”; also often clearly distinguishing it from” other locali-
ties in “Kakhien Hills” (= Kachin Hills)], is likely to refer to the city Bhamo (not Bhamo district) situated close to
the banks of Irrawaddy River, Kachin state in northeastern part of Myanmar. It is also noteworthy that in the same
issue as Boulenger’s work (1887), Thorell (1887) more elaborately discussed the voyages of Col. M. L. Fea based
on his spider collections, indicating that Fea’s collections from Bhamo were gathered in “Nord della Birmania,
luno l’Iravaddi superiore” (= northern Burma, along the Upper Iravaddi) and that “Fra Rangoon e Bham seguendo
l’Iravaddi, corrono circa 900 miglia inglesi” (= approximately 900 British miles run between Yangon and Bhamo,
following the Iravaddi).
Rohanixalus vittatus is reported to occur widely across South and Southeast Asia. There have, however, been
no records of the species from the type locality ever since its original description. Aowphol et al. (2013) reported
sequence data referring to this taxon from two districts within the Kachin state (Myitkyina and Putao, both north of
Bhamo), and Dewei (= Dawei), Tanintharyi division in southwestern Myanmar. An additional sample from Shan
state in eastern Myanmar is also included in the present study. Phylogenetically, all the available samples from
Myanmar nest in four distinct lineages, suggesting that R. vittatus is a complex of multiple species (‘Feihyla vittata
Group I and Group II as per Aowphol et al. 2013; four lineages in the genus Rohanixalus in the present study, Figs.
1, 2). However, to which of these lineages does the name R. vittatus apply, is questionable. The geographical data for
the available Myanmar collections (available at California Academy of Sciences: https://www.calacademy.org/sci-
entists/herpetology-collection) suggests an elevation-linked distribution pattern. The populations from Putao in the
northernmost region of Myanmar (‘Feihyla vittata’ Group I of Aowphol et al. 2013; Rohanixalus cf. shyamrupus in
the present study, Figs. 1, 2) are from elevations of ~ 400–650 m asl. These are probably related to R. shyamrupus
that was originally described from the neighbouring Arunachal Pradesh state in India. Whereas, the populations
from lowland regions of Myitkyina and Dawei of Myanmar form another distinct lineage (‘Feihyla vittata’ Group
II of Aowphol et al. 2013; Rohanixalus vittatus in the present study, Figs. 1, 2), which is likely to be widely distrib-
uted based on disjunct records in lowlands of Myanmar, Thailand (Aowphol et al. 2013), and the Andaman Islands
of India (present study). Another population from elevations of >1500 m asl in eastern Myanmar (present study)
potentially represents a third lineage (included in ‘Feihyla vittata’ Group I of Aowphol et al. 2013; Rohanixalus sp.
1 in the present study, Figs. 1, 2) that nests along with a previously reported Laos sample ~770 m asl (Meegaskum-
bura et al. 2002). This lineage shows a sister relationship with samples from Yunnan region of China that falls in a
moderately high elevation zone (included in ‘Feihyla vittata Group I of Aowphol et al. 2013; Rohanixalus sp. 2 in
the present study, Figs. 1, 2).
Thus, based on its original type locality, Bhamo, R. vittatus is considered to be a low elevation species, widely
distributed in lowland areas right from northern (Myitkyina district in Kachin state) to southwestern Myanmar
(Dawei of Tanintharyi division), adjoining western Thailand (Sangkhla Buri district in Kanchanaburi province),
and the geographically close Andaman Islands of India. On the other hand, Rohanixalus sp. 1 and Rohanixalus sp.
2 potentially represent unidentified or undescribed endemic taxa restricted to relatively higher elevations. This ob-
servation may also be considered significant in the light of remarks made by Thorell (1887), although for arachnids,
noting that the fauna of Upper Iravaddi has a perfectly tropical aspect as that of Lower Iravaddi and Asia from the
south in general; whereas the fauna of the Kachin mountains has a very different character. Hence, based on the cur-
rently available molecular, morphological, and geographical information, we consider the identity of R. vittatus to
be as shown phylogenetically in Figures 1 and 2. New topotypic collections from Bhamo can ascertain the present
conclusion.
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32 · Zootaxa 4878 (1) © 2020 Magnolia Press
Diagnosis. Small to medium-sized adults (male SVL 21–26 mm, female SVL 24–29 mm) with slender body;
snout sub-elliptical to nearly pointed in dorsal view; tympanum externally obscure; dorsum shagreened to sparsely
granular; the entire dorsum, lateral surfaces (including tympanic region), and dorsal surfaces of limbs packed with
fine dark brown speckles, some speckles clumping together to form irregular dark spots or streaks over the dorsum;
limbs with lesser dense spots, thigh and tibia with irregularly scattered spots; a pair of prominent contrasting light
coloured dorsolateral stripes starting from the tip of the snout, extending over the upper eyelid margins, and ending
close to the vent on either side; a rudiment of web present between fingers III and IV at the base, webbing absent
between fingers I, II, and III; foot webbing moderate, up to the second subarticular tubercle on either side of toe IV;
ventral surfaces brownish or greyish white; eggs light green in colour and laid in bubble nests.
Genetic divergence. For the mitochondrial 16S gene, Rohanixalus vittatus differs from other genetically known
congeners (Fig. 1) by: 9.9–11.8% from R. baladika; 9.6–14.0% from R. hansenae; 9.0–10.1% from R. senapatien-
sis; and 8.6–10.1% from R. shyamrupus. For detailed intra-generic comparison see Table 3.
Distribution. Prior to this study, Rohanixalus vittatus was believed to be one of the most widely distributed
species in South and Southeast Asia, and reported from India, Bangladesh, Myanmar, Thailand, Laos, Cambodia,
Vietnam, up to southern China. In India, it was previously reported from the northeast states of Mizoram and Na-
galand (e.g., Dutta 1997; Deuti & Dutta 2002; Ao et al. 2003), excluding an earlier record from Assam that was
clarified as belonging to Nagaland (Dutta 1997). However, based on sampling in the present study, the previously
reported populations of Feihyla from regions across Northeast India represent Rohanixalus senapatiensis or R.
shyamrupus (Fig. 2; Table 1). Hence, the presence of R. vittatus in mainland India, and possibly Bangladesh, is
currently doubted and will require further studies or additional collections. Several other populations previously
identified as ‘vittatus from regions across South and Southeast Asia, including several examined populations from
China (Yunnan Province, Guangxi Province, and Tibet), are likely to represent other Rohanixalus species. Based on
our finding, we restrict the current distribution of Rohanixalus vittatus to low elevations ranging from sea level up
to 200 m asl in Myanmar and Thailand, with a new record from the Andaman Islands (Middle and North Andamans)
of India (Fig. 8).
Natural history and reproductive behaviour. The reproductive behaviour of Rohanixalus vittatus, including
egg-laying, nesting ecology, and clutch morphology, were studied by GG, SDB, and SG under natural conditions
in June 2019. The observations reported below are based on roughly 200 individuals in a population from Rangat,
Middle Andamans, India.
Day 1. Location: fringe vegetation in a small-sized rice paddy; weather condition: rained for about two hours
between 16:30–18:30 h.
Over 100 individuals were observed on the leaves of shrubs and herbs, including banana plants, from 0.1 to 2 m
above the water level on the fringes of a small paddy field of about 15 x 20 m. Seven mating pairs were observed in
axillary amplexus on the leaves of a single plant, aggressively advertising males were also observed around females.
A single gravid female was observed further. The skin over the female’s belly was translucent, making the internal
organs and mature ova visible through the ventral, dorsal, and lateral skin (Fig. 8), as also observed in females of
R. hansenae (Poo & Bickford 2013). The female was perched on a shrub about 0.3 m above the water level on the
edge of a waterlogged paddy field. At least six aggressively calling males were observed within a radius of about
0.5 m from the female. The female approached a calling male, which then mounted the female in axillary amplexus.
The amplected pair moved to a nearby suitable oviposition site, positioned itself and remained still for up to two
minutes. Then the female secreted a drop of mucous, followed by two eggs. The complete process of egg-laying was
completed within 15 minutes, starting from the release of the first egg until the unmounting of the male from am-
plexus. The female continued to sit over the eggs and release gelatinous secretion, with which she glazed the entire
egg clutch multiple times using both the hind limbs. The female remained on the egg clutch for nearly three hours
without much movement. The male usually remained in close proximity to the female, without any direct contact,
during most of the process (Fig. 9).
The freshly laid egg clutch was ovoid in shape (L 19 mm x B 16 mm x H 10 mm) and comprised of a sticky
colourless jelly matrix containing bubbles. The clutch consisted of 120 light green eggs, unpigmented on poles.
Due to the gelatinous and bubbly nature of the nest matrix, it was not possible to distinguish between or retain the
individual jelly layer of each egg. Hence, each ovum was measured excluding the jelly layer, 2.0 ± 0.2 mm (n = 30)
(Fig. 10).
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FIGURE 9. Male advertisement calls and reproductive behaviour in Rohanixalus vittatus from Andaman Islands, India. A–D.
Male advertisement calls: A. oscillogram for 10 s call segment; B. oscillogram for 1 s call segment; C. oscillogram for 0.1 s
call segment; D. spectrogram for 0.1 s call segment; E–I. sequence of reproductive behaviour: E. male and female in axillary
amplexus; F. lateral view of the amplected pair at the time of oviposition, with female depositing light green eggs in jelly-nest;
G. dorsal view of the amplected pair at the time of oviposition, with female clasping the egg clutch with her hind feet; H. male
dismounts while the female remains motionless with her hind legs stretched over the egg clutch; I. male departs while the female
continues to sit over the eggs, produces a gelatinous secretion from cloaca, and glazes it over the entire egg clutch by extending
both her hind feet in a circular motion.
Day 2. The specific egg clutch was observed on the next day, during the daytime at 09:00 h (air temperature
25°C). A female was found about 30 cm away, but not on the egg clutch. At around 18:00 h, the same female visited
the egg clutch and positioned herself on top of the egg mass providing cover. At around 19:00 h, the female again
secreted a viscous fluid from the cloaca and glazed the substance over the egg clutch by extending her hindlimbs in
a circular movement (Fig. 11).
Days 3-4. The egg clutch was observed over the subsequent two days. The same female was found sitting on the
egg clutch, but only during the nighttime, and remained in that position without much movement for long durations
or throughout the night. However, the gelatinous secretion was noted to be released only until the third day. On day
4, the light green coloured ova turned light grey and the jelly matrix became translucent (Figs. 10, 12). Further study
of the same egg clutch was abandoned.
Based on additional observations from multiple egg clutches and maternal egg-attending events in the field,
the following behavioural patterns were noted: (1) territorial behaviour or competition in males: seven male-male
combat events involving pushing, kicking, and dislodging from females were witnessed within roughly four hours
of observation. The fight bouts between males lasted from ~5–20 s. Out of the seven combat events, five involved
multiple males (Fig. 12); (2) multiple males mating with a single female: immediately after mounting of a male with
female and initiation of the egg-laying process, two additional males were observed to mount on either side of the
amplected pair (Fig. 12); (3) egg-attendance possibly only during the late evening and nighttime, but individuals can
be occasionally observed in nearby vegetation during the day time; (4) attendance of a single egg-clutch by mul-
tiple females (Fig. 11); (5) maternal attendance of the developing egg-clutch at all stages until hatching (Fig. 11);
(6) assisted release of hatchlings and young tadpoles from the bubble nest by the female upon disturbance: when
developed embryos are disturbed in the presence of a female, she rapidly and repeatedly kicks the egg clutch with
her hind feet, consequently dismantling the smooth clutch surface, dislodging the hatchlings and young tadpoles
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from the jelly matrix, and propelling them to the water below; (7) laying of fresh eggs over existing clutches or in
their close proximity (Fig. 11).
FIGURE 10. Freshly-laid egg clutch and embryonic development observed in the wild in Rohanixalus vittatus at Andamans
Islands, India. A. The first bout of eggs released from the cloaca of female, about one minute after the initiation of oviposition; B.
freshly-laid egg clutch after the completion of oviposition and glazing of the viscous secretion by female; C. differentiation of the
tail and eyes (stage 21) on day three; D. further differentiation in stage 22 larvae on day four; E. embryos with dark pigmentation
and developed eyes and tail (stage 23), showing movement inside the jelly-nest on day five; F. stage 24 larvae with reduced egg
yolk, ready to hatch after six days; G. young hatchlings (stage 24) before dropping out from the jelly-nest on day six.
Vocalisation. The calls of a Rohanixalus vittatus male (SDBDU 2019.4032) were recorded at Rangat, Middle
Andamans, India on 13 June 2019, by GG, SG, and SDB. The ambient temperature at the time of recording was
30°C (dry bulb) and 28.5°C (wet bulb). The males were observed to produce a single type of call with pulsatile
temporal structure. Calls were not delivered in groups and had uniform intervals. A typical male advertisement call
has a duration of 14.2 ms; a short rise time of about 0.5 ms and fall time of 13.6 ms; and six pulses delivered at a
rate of 472.5 pulses/second. The spectrum is characterized by a single broad peak with mean dominant frequency
of 4.9 kHz (Fig. 9).
We compared the call of R. vittatus with that of R. shyamrupus (SDBDU 4508 recorded at Nagaland, India,
by a Systematic Lab team), a closely related member of the genus. While the overall call structure was observed to
be similar in both species, the call of R. vittatus differed from that of R. shyamrupus primarily by faster pulse rate,
472.5 pulses/second (vs. slower, 173.4 pulses/second) and higher overall dominant frequency, 4.9 kHz (vs. lower,
3.9 kHz).
Based on available literature, we also compared R. vittatus and R. hansenae calls reported from two phyloge-
netically distinct populations in Thailand (‘Feihyla hansenae Groups I and II in Aowphol et al. 2013). Although
the comparison was limited due to methodological differences, especially with respect to acoustic terminologies, the
overall call structure in the two species was similar. However, the call of R. vittatus differed by relatively shorter call
duration, 21.2 ms (vs. longer, 26.89 ms in ‘Feihyla hansenae group I and 99.94 ms in ‘F. hansenae group II), and
lower dominant frequency of 4.9 kHz (vs. higher, 5.1–5.2 kHz for ‘F. hansenae groups I and II).
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 35
FIGURE 11. Maternal egg-attending behaviour observed in Rohanixalus vittatus at Andaman Islands, India. A. A portion of
banana leaf with seven egg clutches at different developmental stages (freshly-laid to stage 24) being attended by females; B.
female attending a one-day egg clutch; C. female attending a developing egg clutch with stage 22 larvae; D. female attending
a developing egg clutch with stage 23 larvae; E–H. attendance of egg clutches at different developmental stages by multiple
females.
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FIGURE 12. Male-male combats observed in Rohanixalus vittatus at Andaman Islands, India. A–C. Males presumably fighting
over perching positions; D. multiple males mating with a single female.
FIGURE 13. Tadpole of Rohanixalus vittatus from Andaman Islands, India. A–D. Gosner stage 35 larva: A. Lateral view, in
life; B. lateral view, in preservation; C. dorsal view, in preservation; and D. ventral view, in preservation; E–G. Oral apparatus
indicating variation in the tooth rows, keratinized jaw sheaths, and marginal and submarginal papillae at three different Gosner
stages: E. Stage 25; F. Stage 30; G. Stage 35. AL: anterior labium; A-1 and A-2: first and second anterior tooth rows; PL: pos-
terior labium; P-1, P-2, and P-3: first, second, and third posterior tooth rows; UJ: upper jaw sheath; LJ: lower jaw sheath; MP:
marginal papillae; SP: submarginal papillae.
Our recording of R. vittatus from Andamans could also not be reliably compared with that of the R. vittatus
population from Thailand (‘Feihyla vittata’ group II in Aowphol et al. 2013). Although the two were similar in
overall structure, they differed in duration (21.2 ms in Andamans population vs. 40.25 ms in Thailand population)
and the overall dominant frequency (4.9 kHz in Andamans population vs. 3.9 kHz in Thailand population). Further
acoustic studies are required for a proper understanding of intra- and interspecific variations among the calls of spe-
cies in the genus Rohanixalus.
Tadpole morphology. The larval description for Rohanixalus vittatus is based on Gosner stage 35 tadpole
(SDBDU 2019.4052) collected from a freshly laid egg clutch from Rangat, Middle Andamans, India on 15 June
2019, by GG, SDB, and SG.
External morphology. Small-sized tadpole (tl 24.75, svl 8.10, su 6.31); body oval-shaped in dorsal view (bh
3.68, bw 4.25); eyes rounded, positioned dorsolaterally (pp 3.95), eye diameter (ed 1.63) 38% of the body width (bw
4.25); snout shape rounded in dorsal and lateral view (from eyes to the base of upper labium); umbraculum absent;
nasolacrimal duct absent; narial depressions located closer to the snout than the eyes (rn 1.09, np 1.95, nn 1.89);
narial depressions and surrounding pigmentation not clearly visible in dorsal view; spiracle present as a muscular
flap on the left side of the body, opens mid-laterally (ss 5.65); a medial vent tube present, tubular in structure, and
with the aperture opening dextrally compared to the plane of the ventral fin; tail (vt 16.65, ht 4.62) composed of
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 37
bilateral myotomic muscle masses divided by V-shaped septa, with unequal tail membranes on either side of the tail
musculature; dorsal fin originates anterior to the tail body junction, whereas the ventral fin originates near the vent
tube; margin of the lower fin not parallel to the margin of tail muscle; tail musculature strong, extends up to the end
of the tail tip with a narrow terminal part; tail-muscle height (tmh 2.22) 60% of the body height (bh 3.68) and tail-
muscle width (tmw 1.82) 43% of body width (bw 4.25) (Fig. 13).
Mouth (oral disc): Oral disc elliptical in shape, anteroventrally positioned, moderately small, oral disc width
(odw 1.91) 45% of body width and 24% of snout to vent length; mouth consisting of lower and upper labia, with the
lower being larger; lateral emarginations present on both sides, bordered by continuous small blunt marginal papil-
lae; upper labium with a single row of small and blunt marginal papillae, marginal papillae not continuous through-
out the margin and contain a medial gap in between; lower labium with a row of elongated and blunt marginal
papillae, and a second row of blunt submarginal papillae, a ventral gap present in both the rows of papillae; two
upper tooth ridges and three lower ridges are present; uniserial tooth rows present for each tooth ridge, composed
of pointed and uniformly sized labial teeth except for some inconsiderable occasions (like the lateral ends where
the teeth tend to become smaller in size); A2 have natural gaps between them while A1 shows gaps made by teeth
degeneration; P1 indicates a gap comparatively smaller to A2; labial tooth row formula: 2(2)/3(1); keratinized up-
per and lower jaw sheaths, with an inverted, smooth, U-shaped upper beak and a V-shaped lower beak, both mostly
black with pale bases and serrated margins.
Development of mouth parts: The number of tooth rows and presence of papillae differs as development of the
tadpole progresses; upper labium possesses only marginal papillae throughout its various developmental stages,
whereas the lower labium possesses rows of marginal and submarginal papillae that increase in number and size as
the larvae develop; the row of submarginal papillae is not continuous. Tooth rows are developed in the early tadpole
stages (Gosner 25); A1 have no medial gap till stage 30 but have a wide medial gap by stage 35, after which the teeth
start degenerating; A2 have a wide medial gap from Gosner 25 and start degenerating at Gosner 35; P1 indicates a
narrow gap starting from Gosner 25; the oral disc is well-developed at stage 30, with two anterior (A1–A2) and three
posterior tooth rows (P1-P3); initially anterior tooth rows degenerate; A1 and A2 degenerate at Gosner 35; P1–P3
remain intact even in Gosner 35.
Rohanixalus baladika (Riyanto and Kurniati, 2014) comb. nov.
Sumatran Bubble-nest Frog
(Figs. 1–2, 4, 7, 14–15; Tables 1, 3, 4)
Original name and description. Chiromantis baladika Riyanto and Kurniati, 2014. Riyanto A. and Kurniati H.
2014. Three new species of Chiromantis Peters 1854 (Anura: Rhacophoridae). Russian Journal of Herpetology, 21:
65–73. Type. Holotype, MZB Amph. 17.935, by original designation. Type locality. “West Sumatra Province (1°
26’ 15.3 S, 101° 31’ 47.7 E; elevation 273 m a.s.l.) at temporary pool in palm oil plantation”, Indonesia. Current
status of specific name. Valid name as Rohanixalus baladika (Riyanto and Kurniati, 2014) comb. nov.
Taxonomic remarks. This taxon was originally described from West Sumatra based on five specimens. We
examined the type specimens and find that this species is closely related to members of the genus Rohanixalus
rather than Chirixalus due to the following characters: prominent and dense minute speckles throughout the dorsum,
lateral surfaces, and dorsal surface of limbs (vs. absent); and a pair of light coloured dorsolateral stripes (prominent
to faint, continuous or discontinuous) beginning from the snout tip, extending over the upper eyelid margins, and
ending close to the vent on either side (vs. absent) (Figs. 14, 15). Furthermore, the type series is morphologically
very similar to new collections from West Sumatra, the phylogenetic position of which is confirmed in the new
genus (Figs. 1, 2). Hence, based on the new molecular and morphological evidence presented herein, we propose
to formally reallocate Chiromantis baladika Riyanto and Kurniati, 2014 (= Chirixalus baladika) to Rohanixalus
baladika (Riyanto and Kurniati, 2014) comb. nov.
Furthermore, Rohanixalus baladika is morphologically close to R. marginis and R. nauli due to similar adult
size, male SVL 21–22 mm (vs. SVL 21 mm in R. nauli and SVL 22.8 mm in R. marginis), as well as the presence
of weak to prominent dorsolateral stripes and faint to prominent scattered dorsal spots in all the three species. Many
of the diagnostic characters mentioned in the original publication (Riyanto & Kurniati 2014) for distinguishing R.
baladika from R. marginis are variable within the types series (especially the prominence and thickness of the dorso-
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38 · Zootaxa 4878 (1) © 2020 Magnolia Press
lateral stripes) (Fig. 14), except for the absence of rudimentary webbing between fingers III and IV (vs. present in R.
marginis, but absent in R. nauli). While the comparison with R. nauli is based on the absence of dorsolateral stripes
and dorsal spots (Riyanto & Kurniati 2014), our examination of the holotype of R. nauli suggests the presence of
faint dorsolateral stripes (Fig. 15). Since the description of R. nauli is based on a single specimen, this character
cannot be considered reliable without considering possible variation. In the present study, we also examined new
collections from North and West Sumatra and found dorsal colouration as well as the prominence of dorsolateral
stripes to be variable among individuals (Fig. 14). These two species, along with R. marginis, are also found in close
geographical proximity. Our new collections also show R. baladika and R. nauli to be genetically close with a shal-
low divergence of 0.2–0.6% (Fig. 2). Hence, R. baladika and R. nauli are likely to be synonyms, pending further
validation.
FIGURE 14. Variations in dorsal markings and dorsolateral lines in Rohanixalus baladika. A. MZB Amph. 17.938 (paratype);
B. MZB Amph. 17.939 (paratype); C. MZB Amph. 17.936 (paratype); D. MZB Amph. 18684; E. MZB Amph. 15409; F. MZB
Amph. 16661; G. MZB Amph. 15655.
Diagnosis. Small-sized adults (male SVL 21–23 mm) with slender body; snout rounded to subovoid in dorsal
view; the entire dorsum, lateral surfaces, and dorsal surface of limbs covered with fine dark brown speckles, some
speckles clumping together to form dark irregular blotches or reticulation on the dorsum; a pair of narrow, promi-
nent to faint, continuous or discontinuous, dorsolateral stripes starting from the snout tip, extending over the eyes,
and ending close to the vent; the prominence of dorsolateral stripes and scattered dorsal spots highly variable among
individuals (e.g, dorsolateral stripes faint in Paratype MZB Amph. 17.938 and MZB Amph. 16661, Fig. 14); lateral
surfaces of the head, including tympanic region, darker than dorsal colouration; foot webbing moderate, up to the
second subarticular tubercle on either side of toe IV; eggs light green in colour and laid in bubble nests.
Genetic divergence. For the mitochondrial 16S gene, Rohanixalus baladika differs from other genetically
known congeners (Fig. 1) by: 8.3–11.3% from R. hansenae; 8.7–9.6% from R. senapatiensis; 9.4–10.1% from R.
shyamrupus; and 9.9–11.8% from R. vittatus. For detailed intra-generic comparison see Table 3. Further, our new
collections from North Sumatra and West Sumatra are divergent by 0.2–0.6%.
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 39
FIGURE 15. Holotypes of Rohanixalus baladika and R. nauli. A–J. Rohanixalus baladika, MZB Amph. 17.935: A. dorsal
view; B. ventral view; C. lateral view of head; D. ventral view of hand; E. ventral view of foot; F. schematic illustration of web-
bing on foot; G–L. Rohanixalus nauli, MZB.Amph. 14316: G. dorsal view; H. ventral view; I. lateral view of head; J. ventral
view of hand; K. ventral view of foot; L. schematic illustration of webbing on foot.
Distribution. Rohanixalus baladika is currently known only from West Sumatra and North Sumatra Province
in Indonesia.
Rohanixalus hansenae (Cochran, 1927) comb. nov.
Hansen’s Bubble-nest Frog
(Figs. 1–2, 4, 7, 16; Tables 1, 3, 4)
Original name and description. Philautus hansenae Cochran, 1927. Cochran D.M. 1927. New reptiles and batra-
chians collected by Dr. Hugh M. Smith in Siam. Proceedings of the Biological Society of Washington, 40: 179–192.
Type. Holotype, USNM 70109, by original designation. Type locality. “Nong Khor, southeastern Siam”, Thailand.
Current status of specific name. Valid name as Rohanixalus hansenae (Cochran, 1927) comb. nov.
Taxonomic remarks. Based on the morphological and phylogenetic evidence presented in this study, this spe-
cies is a member of the genus Rohanixalus. The taxon was originally described from southeastern Thailand and
subsequently reported to have a wide distribution within Thailand. Stuart and Emmett (2006) doubted the validity
of the species, however, R. hansenae was not only shown to be phylogenetically, morphologically, and acoustically
distinct from R. vittatus (Aowphol et al. 2013), but also suggested to comprise of two distinct lineages likely to be
separated geographically (as Feihyla hansenae Group I from central, eastern, and peninsular Thailand including
the type locality; and ‘Feihyla hansenae Group II largely from montane forests in northwestern and western Thai-
land) (Aowphol et al. 2013; Yodthong et al. 2014).We further investigated the levels of genetic divergence among
various available populations of Rohanixalus hansenae, including new collections from the type locality. Our phy-
logenetic results concur with the previous studies and show unidentified lineages among the various R. hansenae
populations, indicating that it could indeed represent a species complex, requiring detailed morphological studies.
Diagnosis. Small-sized adults (male SVL 18–21 mm, female SVL 24–25 mm) with slender body; snout nearly
pointed in dorsal view; dorsal colouration light to dark brown with a reddish tinge; the entire dorsum, lateral sur-
faces, and dorsal surface of limbs covered with fine dark brown speckles, some speckles clumping together to form
dark irregular blotches on the dorsum; a pair of light coloured and contrasting continuous dorsolateral stripes start-
ing from the snout tip, extending over the eyes, and ending close to the vent; lateral surfaces of the head, including
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40 · Zootaxa 4878 (1) © 2020 Magnolia Press
tympanic region, darker than dorsal colouration; foot webbing moderate, just above the second subarticular tubercle
on either side of toe IV; eggs light green in colour and laid in bubble nests.
Genetic divergence. For the mitochondrial 16S gene, Rohanixalus hansenae differs from other genetically
known congeners (Fig. 1) by: 8.3–11.3% from R. baladika; 8.6–10.7% from R. senapatiensis; 8.2–10.2% from
R. shyamrupus; and 9.6–14.0% from R. vittatus. For detailed intra-generic comparison see Table 3. Further, the
sampled populations of Rohanixalus hansenae show high uncorrected intraspecific divergences and the following
distances are observed between R. hansenae (typical) and its major sub-lineages: 4.4–8.0% for R. cf. hansenae 1
and 4.1–6.7% for R. cf. hansenae 2 (Table 3).
Distribution. Rohanixalus hansenae is widely distributed across Thailand, with presumed distribution in the
adjoining regions of Cambodia and Myanmar (Yodthong et al. 2014) (Fig. 4).
FIGURE 16. Rohanixalus hansenae in life, from the type locality Nong Khor, Thailand. A. Dorsolateral view (male); B. dor-
solateral view (female); C. dorsal view; D. enlarged view of the tympanic region showing star-shaped speckles; E. ventral view
(female); F. ventral view (male); G. ventral view of hand; H. ventral view of foot; I. schematic illustration of webbing on foot.
Rohanixalus marginis (Chan, Grismer, Anuar, Quah, Grismer, Wood, Muin, and Ahmad, 2011) comb. nov.
Malaysian Bubble-nest Frog
(Figs. 1–2, 4, 7; Tables 1, 3)
Original name and description. Chiromantis marginis Chan, Grismer, Anuar, Quah, Grismer, Wood, Muin, and
Ahmad 2011. Chan K. O., Grismer L.L., Anuar S., Quah E., Grismer J. L., Wood P. L. Jr., Muin M.A., and Ahmad N.
2011. A new species of Chiromantis Peters 1854 (Anura: Rhacophoridae) from Perlis State Park in extreme north-
ern peninsular Malaysia with additional herpetofaunal records for the park. Russian Journal of Herpetology, 18:
253–259. Type. Holotype, LSUHC 9700, by original designation. Type locality. “from the dam behind Perlis State
Park chalets, Perlis”, Peninsular Malaysia. Current status of specific name. Valid name as Rohanixalus marginis
(Chan, Grismer, Anuar, Quah, Grismer, Wood, Muin, and Ahmad 2011) comb. nov.
Taxonomic remarks. This taxon was described based on a single specimen from Peninsular Malaysia. Al-
though the species was originally assigned to genus Chiromantis (= Chirixalus), it is morphologically closer to
members of the genus Rohanixalus, primarily due to the following characters: prominent and dense minute speck-
les throughout the dorsum, lateral surfaces, and dorsal surface of limbs (vs. absent); and presence of light coloured
dorsolateral stripes beginning from the snout tip, extending over the upper eyelid margins, and ending close to the
vent on either side (vs. faint to prominent dorsal lines or absent). Further, the close relationship of R. marginis
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 41
with R. hansenae, R. punctatus and R. vittatus was also originally suggested (Chan et al. 2011). However, the
comparison of R. marginis with R. punctatus is considered weak as the degree of prominence of dorsal spots and
dorsolateral stripes is variable among R. punctatus individuals. Rohanixalus marginis is also morphologically
close to R. baladika and R. nauli (see taxonomic remarks under R. baladika for detailed discussion). Hence, based
on available evidence, we formally reallocate Chiromantis marginis Chan, Grismer, Anuar, Quah, Grismer, Wood,
Muin, and Ahmad 2011 (= Chirixalus marginis) to the genus Rohanixalus, as Rohanixalus marginis (Chan, Gris-
mer, Anuar, Quah, Grismer, Wood, Muin, and Ahmad 2011) comb. nov. Further molecular studies can clarify the
relationship of R. marginis with other Rohanixalus members, especially R. punctatus, with which it could be closely
related based on morphological and geographical considerations (see taxonomic remarks under R. punctatus for
further discussion).
Diagnosis. Small-sized adults (male SVL 23 mm) with slender body; tympanum indistinct, vomerine teeth
absent; dorsal colouration greyish-white; the entire dorsum, lateral surfaces, and dorsal surface of limbs covered
with fine dark brown speckles, some speckles clumping together to form dark irregular blotches or reticulation on
the dorsum; a pair of narrow continuous dorsolateral stripes starting from the snout tip, extending over the eyes, and
ending close to the vent on either side; webbing absent between fingers I, II, and III, basal webbing present between
fingers III and IV; foot webbing moderate, up to the second subarticular tubercle on either side of toe IV.
Distribution. Rohanixalus marginis is currently known only from its type locality, Perlis State Park in Penin-
sular Malaysia.
Rohanixalus nauli (Riyanto and Kurniati, 2014) comb. nov.
Nauli Bubble-nest Frog
(Figs. 1–2, 4, 7, 15; Tables 1, 3, 4)
Original name and description. Chiromantis nauli Riyanto and Kurniati, 2014. Riyanto A. and Kurniati H. 2014.
Three new species of Chiromantis Peters 1854 (Anura: Rhacophoridae). Russian Journal of Herpetology, 21: 65–73.
Type. Holotype, MZB.Amph.14.916, by original designation. Type locality. “Teluk Nauli, Sibolga, North Sumatra
Province”, Indonesia. Current status of specific name. Valid name as Rohanixalus nauli (Riyanto and Kurniati,
2014) comb. nov.
Taxonomic remarks. This taxon was originally described from North Sumatra based on a single specimen. We
examined the type specimen and found that this species is more closely related to members of the genus Roha-
nixalus than to Chirixalus. Further, R. nauli is morphologically similar to R. baladika described from West Sumatra
(for detailed discussion see taxonomic remarks under R. baladika). Subsequent new collections from North and
West Sumatra show that R. baladika and R. nauli are likely to represent a single species, with a shallow divergence
of 0.2–0.6%. Phylogenetically, the populations from North and West Sumatra are nested in the genus Rohanixalus
(Fig. 1). The dorsal colour and markings are also highly variable among the type specimens of R. baladika as well as
new collections from regions across North and West Sumatra, especially the prominence of dorsal spots and dorso-
lateral stripes (Fig. 14) that were originally used to distinguish the two species (Riyanto & Kurniati 2014). We also
report a Rohanixalus population from North Sumatra that was observed guarding a nest (Fig. 7), which is suggested
as a possible behavioural synapomorphy for diagnosis of the genus Rohanixalus (e.g., Li et al. 2009; Hertwig et
al. 2013; present study). Hence based on available evidence, we propose to formally reallocate Chiromantis nauli
Riyanto and Kurniati, 2014 (= Chirixalus nauli) as Rohanixalus nauli (Riyanto and Kurniati, 2014) comb. nov.
However, R. nauli is likely to be a junior subjective synonym of R. baladika, pending further investigation.
Diagnosis. Small-sized adults (male SVL 20.9 mm), with slender body; snout subelliptical to nearly pointed in
dorsal view; dorsal skin shagreened to sparsely granular; the entire dorsum, lateral surfaces, and dorsal surface of
limbs covered with fine dark brown speckles (in the type), some speckles clumping together to form dark irregular
blotches or reticulation on the dorsum; a pair of obscure dorsolateral stripes present in the type (stated as absent in
the original description); foot webbing moderate, up to the second subarticular tubercle on either side of toe IV; eggs
light green in colour and laid in bubble nests. Populations from north Sumatra were observed to have variable dorsal
markings and degree of prominence of the dorsolateral stripes (Fig. 14).
Genetic divergence. Sequence data from North and West Sumatra, potentially representing Rohanixalus nauli
and R. baladika respectively, shows shallow divergence of 0.2–0.6%.
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42 · Zootaxa 4878 (1) © 2020 Magnolia Press
TABLE 3. Inter and intraspecific uncorrected p-distances (in percent) in the genus Rohanixalus gen. nov. for the mitochondrial 16S rRNA gene sequences. Values indicate average
distances followed by range in parentheses.
Taxa 1 2 3 4 5 6 7 8 9 10
1Rohanixalus baladika 0.4
(0.2–0.6)
2Rohanixalus hansenae 9.3
(8.3–11.3)
1.2
(0–3.2)
3Rohanixalus cf. hansenae 110.7
(9.6–13.4)
5.7
(4.4–8.0)
2.5
(0–5.3)
4Rohanixalus cf. hansenae 211.6
(11.1–12.0)
5.2
(4.1–6.7)
5.4
(4.5–6.3)
0.5
(0–1.0)
5Rohanixalus shyamrupus 9.8
(9.4–10.1)
8.9
(8.2–10.2)
8.4
(7.3–10.0)
8.5
(8.3–9.2)
0.1
(0–0.4)
6Rohanixalus senapatiensis 9.3
(8.7–9.6)
9.2
(8.6–10.7)
9.4
(7.9–10.5)
9.4
(9.1–9.8)
3.2
(3.0–3.4)
0.1
(0–0.2)
7Rohanixalus cf.
shyamrupus
8.5
(7.9–8.8)
9.4
(8.8–10.6)
9.4
(8.2–10.9)
9.3
(9.0–9.7)
2.8 2.9
(2.8–3.2)
0.3
(0–0.4)
8Rohanixalus vittatus 11.0
(9.9–11.8)
10.9
(9.6–14.0)
11.9
(11.0–14.0)
12.1
(11.6–13.5)
9.3
(8.6–10.1)
9.6
(9.0–10.1)
8.7
(7.8–9.4)
0.9
(0–2.1)
9Rohanixalus sp. 1 9.7
(9.4–9.9)
9.2
(8.4–11.3)
9.4
(8.7–11.6)
9.6
(9.4–9.9)
5.0
(5.0–5.2)
5.6
(5.5–5.9)
4.8
(4.7–4.9)
9.2
(9.0–9.6)
0.3
(0.3–0.4)
10 Rohanixalus sp. 2 9.4
(8.8–9.9)
8.4
(7.2–10.4)
9.0
(8.4–10.7)
8.9
(8.5–9.2)
4.5
(4.5–4.6)
5.2
(5.1–5.3)
4.3
(4.2–4.4)
8.7
(7.6–9.2)
2.3
(2.8–3.4)
1.2
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 43
Distribution. Rohanixalus nauli is currently known only from its type locality in North Sumatra Province,
Indonesia.
TABLE 4. Morphometric measurements for six species of the new genus Rohanixalus. Measurement abbreviations and
museum acronyms are provided in the Material and methods section. HT (holotype); TT (topotype); RS (referred speci-
men); M (male). All measurements are in millimetres (mm).
Taxa Museum No. SVL HL HW SL EL TYD TYE
Rohanixalus baladika (HT) MZB Amp 17935 (M) 21.0 6.5 6.8 3.2 3.1 1.0 0.8
Rohanixalus hansenae (TT) CUMZ-A-7728 (M) 20.8 6.6 5.9 3.0 2.6 0.9 0.4
Rohanixalus nauli (HT) MZB Amp 17936 (M) 20.9 6.5 6.5 3.4 3.0 1.0 0.6
Rohanixalus senapatiensis (TT) SDBDU 41656 (M) 20.8 7.2 6.1 3.3 2.6 1.0 0.3
Rohanixalus shyamrupus (TT) SDBDU 41770 (M) 22.0 7.1 6.4 3.4 2.6 0.6 0.2
Rohanixalus vittatus (RS) SDBDU 2019.4032 (M) 32.1 7.8 6.5 3.6 2.6 1.0 0.4
Continued.
Taxa TYE EN NS IN FAL HAL TL SHL FOL TFOL
Rohanixalus baladika (HT) 0.8 1.3 1.0 1.9 3.6 5.9 10.7 10.5 8.6 13.1
Rohanixalus hansenae (TT) 0.4 1.8 1.2 2.0 3.4 5.8 9.3 10.5 8.3 13.9
Rohanixalus nauli (HT) 0.6 1.6 1.3 2.1 3.7 5.6 10.4 10.1 9.0 13.2
Rohanixalus senapatiensis (TT) 0.3 1.2 1.1 2.0 3.7 6.8 9.7 10.9 9.2 14.3
Rohanixalus shyamrupus (TT) 0.2 1.5 1.3 2.1 4.0 6.7 11.0 10.2 9.0 13.9
Rohanixalus vittatus (RS) 0.4 1.6 1.3 2.0 4.2 5.8 10.1 10.5 8.9 14.0
Rohanixalus punctatus (Wilkinson, Win, Thin, Lwin, Shein, and Tun, 2003) comb. nov.
Spotted Bubble-nest Frog
(Figs. 1–2, 4, 7; Tables 1, 3)
Original name and description. Chirixalus punctatus Wilkinson, Win, Thin, Lwin, Shein, and Tun 2003. Wilkin-
son J.A., Win, H., Thin T., Lwin K.S., Shein A.K. and Tun H. 2003. A new species of Chirixalus (Anura: Rhaco-
phoridae) from western Myanmar (Burma). Proceedings of the California Academy of Sciences, 54: 17–26. Type.
Holotype, CAS 221555, by original designation. Type locality. “Rakhine Yoma Elephant Wildlife Sanctuary Head-
quarters (17° 36’ 48.8” N 94° 36’ 50.2” E), Gwa, Gwa Township, Rakhine State”, Myanmar. Current status of spe-
cific name. Valid name as Rohanixalus punctatus (Wilkinson, Win, Thin, Lwin, Shein, and Tun 2003) comb. nov.
Taxonomic remarks. Based on morphological affinity, this species is closely related to members of the genus
Rohanixalus. Wilkinson et al. (2003) diagnosed the species on the basis of differences in dorsal colour and mark-
ings with the closely related R. vittatus, but subsequently stated in the variations section that the extent of spotting
varies, being “sparser in some individuals, restricted primarily to the dorsal aspect of the head whereas in others to
the entire dorsum. The accompanying paratype photograph (Joseph B. Slowinski field voucher number JBS 9274,
deposited in CAS; Wilkinson et al. 2003) also shows close similarities to another species, R. marginis, that was
described more recently from Peninsular Malaysia. Based on available evidence, we formally reallocate Chirixalus
punctatus Wilkinson, Win, Thin, Lwin, Shein, and Tun 2003 to the genus Rohanixalus, as Rohanixalus punctatus
(Wilkinson, Win, Thin, Lwin, Shein, and Tun 2003) comb. nov. Further molecular evidence can clarify the system-
atic relationship of R. punctatus with other Rohanixalus members, especially R. marginis, which could be a close
congener based on morphological and geographical considerations.
Diagnosis. Small-sized adults (male, SVL 23 mm), with slender body; snout sub-elliptical to nearly pointed
in dorsal view; tympanum externally obscure; dorsal skin smooth; the entire dorsum, lateral surfaces, and dorsal
surface of limbs covered with fine dark brown speckles; dorsum and limbs with or without prominent spots; a pair
of continuous dorsolateral stripes starting from the snout tip, extending over the eyes, and ending close to the vent;
webbing absent between fingers I, II, and III, basal webbing present between fingers III and IV; foot webbing mod-
erate, up to the second subarticular tubercle on either side of toe IV.
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44 · Zootaxa 4878 (1) © 2020 Magnolia Press
Distribution. Rohanixalus punctatus is currently known only from western Myanmar, the place of its original
description.
Rohanixalus senapatiensis (Mathew and Sen, 2009) comb. nov.
Senapati’s Bubble-nest Frog
(Figs. 1–2, 4, 7, 17; Tables 1, 3, 4)
Original name and description. Chirixalus senapatiensis Mathew and Sen, 2009. Mathew R. and Sen N. 2009.
Studies on little known amphibians of Northeast India. Records of the Zoological Survey of India, Occasional Pa-
pers, 293: 1–64. Type. Holotype, VA/ERS/ZSII/804 (ZSIE 804), by original designation. Type locality. “Mabing
river bed, Kangpokpi, Senapati district”, Manipur, India. Current status of specific name. Valid name as Roha-
nixalus senapatiensis (Mathew and Sen, 2009) comb. nov.
FIGURE 17. Feihyla senapatiensis in preservation and life. A–F. Holotype, VA/ERS/ZSII/804 (ZSIE 804), in preservation: A.
dorsal view; B. ventral view; C. lateral view; D. ventral view of hand; E. ventral view of foot; F. schematic illustration of web-
bing on foot; G–I. dorsolateral view, in life.
Taxonomic remarks. In the original description, Mathew and Sen (2009) distinguished this species from the
closely related Rohanixalus vittatus and R. shyamrupus mainly by its “Toes almost fully webbed” (vs. “Toes 3/4th
webbed” in R. vittatus), and certain dorsal colour differences such as “dorsum uniform light brown” (vs. “speckled
all over with purple” in R. vittatus) that are found to be variable among various species of Rohanixalus. These char-
acters mentioned by the authors are also not found in the type of R. senapatiensis, especially fully webbed toes.
Our examination of the holotype of Chirixalus senapatiensis also shows that it represents a member of the
genus Rohanixalus, mainly due to the presence of characters such as dorsolateral stripes and dorsal spots. We have
further confirmed the identity and generic placement of Rohanixalus senapatiensis (Mathew and Sen, 2009) comb.
nov., both morphologically and phylogenetically, based on new topotypic and additional collections from Northeast
India (Figs. 1, 2; Table 1).
Diagnosis. Small-sized adults (male SVL 18–21 mm, female SVL 24–25 mm) with a slender body; snout
nearly pointed in dorsal view; dorsal colouration light to dark brown with a reddish tinge; the entire dorsum, lateral
surfaces, and dorsal surface of limbs covered with fine dark brown speckles, some speckles clumping together to
form dark and irregular blotches on the dorsum; a dark continuous dorsolateral stripe starts from the tip of the snout,
extends through the eye, and reaches close to the vent on either side; lateral surfaces of the head, including tym-
panic region, darker than dorsal colouration; foot webbing moderate, extending just above the second subarticular
tubercle on either side of toe IV; eggs light green in colour and laid in bubble nests.
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 45
Genetic divergence. For the mitochondrial 16S gene, Rohanixalus senapatiensis differs from other genetically
known congeners (Fig. 1) by average uncorrected genetic distance of: 8.7–9.6% from R. baladika, 8.6–10.7% from
R. hansenae, 3.0–3.4% from R. shyamrupus, and 9.0–10.1% from R. vittatus. Some previously sampled populations
from the geographically close Putao district in Kachin state of Myanmar (‘Feihyla vittata Group I of Aowphol et al.
2013; R. cf. shyamrupus in the present study, Fig. 2) are related to R. senapatiensis by 2.8–3.2% divergence. Further
studies are required to ascertain the identity of Putao populations, as well as two additional unidentified lineages
(Rohanixalus sp. 1 and Rohanixalus sp. 2). For detailed intra-generic comparisons see Table 3.
Distribution. Rohanixalus senapatiensis was originally described based on a single specimen from Kangpokpi
in Senapati district of Manipur state, India. Based on subsequent surveys in Northeast India, this species is known
from Tamenglong and Churachandpur districts of Manipur; Phek district of Nagaland; Mizoram; and Tripura. The
previous report of R. vittatus from Mizoram by Deuti & Dutta (2002) is likely to represent R. senapatiensis, based
on the available specimen (ZSI A 9209, from Ngengpuri Wildlife Sanctuary) at ZSI Kolkata.
Rohanixalus shyamrupus (Chanda and Ghosh, 1989) comb. nov.
Shyamrup’s Bubble-nest Frog
(Figs. 1–2, 4, 7, 18–19; Tables 1, 3, 4)
Original name and description. Philautus shyamrupus Chanda and Ghosh, 1989. Chanda S. K. and Ghosh A.
K. 1989. A new frog of the genus Philautus Gistel, from the proposed Namdapha Biosphere Reserve, Arunachal
Pradesh, northeast India. Journal of the Bombay Natural History Society, 86: 215–217. Type. Holotype, ZSIC
A 7944 (formerly ZSI-KZ 313), by original designation. Type locality. “Hornbill, Namdapha Tiger Reserve”,
Arunachal Pradesh, India. Current status of specific name. Valid name as Rohanixalus shyamrupus (Chanda and
Ghosh, 1989) comb. nov.
Taxonomic remarks. Chanda & Ghosh (1989) originally placed this species in the genus Philautus. The origi-
nal description, as well as Chanda & Sarkar (1997), did not compare this taxon with any known Philautus species
from Eastern Himalayas or neighbouring regions, other than with unrelated ranid species (Dubois 1999; Bossuyt
& Dubois 2001). Based on the description and available photograph (Chanda & Sarkar 1997), Bossuyt & Dubois
(2001) suggested that the species is not a member of Philautus due to characters such as longitudinal bands or lines
on the body or flanks, smooth chest and belly, extensive webbing, and lack of inner and outer metatarsal tubercles.
They provisionally transferred it to the genus Chirixalus, without stating any potentially close taxa. We examined
the holotype of Philautus shyamrupus and found that it represents a member of the genus Rohanixalus due to
characters such as the presence of dorsolateral stripes and dorsal spots. We have further confirmed the identity and
generic placement of Rohanixalus shyamrupus (Chanda and Ghosh, 1989) comb. nov., both morphologically and
phylogenetically, based on new topotypic and additional collections from Northeast India (Figs. 1, 2; Table 1).
Diagnosis. Small-sized adults (male SVL 18–21 mm, female SVL 24–25 mm) with a slender body; snout
nearly pointed in dorsal view; dorsal colouration light to dark brown with a reddish tinge; the entire dorsum, lateral
surfaces, and dorsal surfaces of limbs covered with fine dark brown speckles, some speckles clumping together to
form dark and irregular blotches on the dorsum; a dark and continuous dorsolateral stripe starts from the tip of the
snout, extends through the eyes, and reaches close to the vent on either side; lateral surfaces of the head, including
tympanic region, darker than dorsum; foot webbing moderate, just above the second subarticular tubercle on either
side of toe IV. Further, the eggs are light green in colour and laid in bubble nests.
Genetic divergence. For the mitochondrial 16S gene, Rohanixalus shyamrupus differs from other genetically
known congeners (Fig. 1) by average uncorrected genetic distance of: 9.4–10.1% from R. baladika, 8.2–10.2% from
R. hansenae, 3.0–3.4% from R. senapatiensis, and 8.6–10.1% from R. vittatus. Some previously sampled popula-
tions from the geographically close Putao district in Kachin state of Myanmar (‘Feihyla vittata’ Group I of Aowphol
et al. 2013; R. cf. shyamrupus in the present study, Fig. 1) are related to R. shyamrupus by 2.8% divergence. Further
studies are required to ascertain the identity of Putao populations, as well as two additional unidentified lineages
(Rohanixalus sp. 1 and Rohanixalus sp. 2). For detailed intra-generic comparisons see Table 3.
Distribution. Rohanixalus shyamrupus was originally described from Changlang district in the state of Arunach-
al Pradesh, India. In the present study, we genetically confirm the identity and phylogenetic relationship of the spe-
cies based on topotypes from Namdapha Tiger Reserve, Changlang district. Additionally, this species is known from
BIJU ET AL.
46 · Zootaxa 4878 (1) © 2020 Magnolia Press
West Kameng district in Arunachal Pradesh; and Kohima and Peren districts of Nagaland. Previous reports of R.
vittatus from Nagaland (Romer 1951 “1949”; Kiyasetuo & Khare 1986) are likely to refer to this species.
FIGURE 18. Feihyla shyamrupus in preservation and life. A–G. Holotype, ZSIC A 7944 (formerly ZSI-KZ 313): A. dorsal
view; B. ventral view; C. lateral view; D. lateral view of head; E. ventral view of hand; F. ventral view of foot; G. schematic
illustration of webbing on foot; H–O. Topotypes from Northeast India, in life: H. dorsolateral view; I. dorsal view; J–K. dor-
solateral view; L. enlarged view of skin over the tympanic region showing star-shaped speckles; M. ventral view; N. ventral
view of hand; O. ventral view of foot.
FIGURE 19. Male advertisement calls of Feihyla shyamrupus recorded at Nagaland, India. A. Oscillogram of 10 s call seg-
ment; B. oscillogram of 1 s call segment; C. oscillogram of 0.1 s call segment; D. spectrogram of 0.1 s call segment.
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 47
FIGURE 20. Types of Philautus cherrapunjiae (= Rhacophorus cherrapunjiae comb. nov.) in preservation. A. Dorsal view;
B. ventral view; C. dorsal view.
III. Taxonomic identity of Chirixalus cherrapunjiae (Roonwal and Kripalani, 1966 “1961”)
This taxon was originally described as Philautus cherrapunjiae Roonwal and Kripalani, 1966 “1961” based on a
“recently metamorphosed frog” and “several young frogs and tadpoles” from the “vicinity of the Circuit House near
Cherrapunji (the Khasi-Jaintia Hills District, Assam, eastern India)”, now Meghalaya State, India. However, the
allocation of this species to the genus Philautus was considered erroneous, as it was stated to have free-swimming
tadpoles (vs. direct development in Philautus) (Dubois 1999; Bossuyt & Dubois 2001). Further, Bossuyt & Dubois
(2001) provisionally transferred the species to genus Chirixalus since the original description considered it close to
Philautus vittatus (= Chirixalus vittatus). Wilkinson et al. (2003) doubted this placement due to the lack of oppos-
able fingers in Philautus cherrapunjiae (vs. present in Chirixalus members), and suggested that it may be a member
of genus Rhacophorus, without any explanation. Subsequently, the species was reported from Arunachal Pradesh
without a voucher (Sarkar & Ray 2006), and from Assam but only with the type photographs (metamorphosed frog)
(Mathew & Sen 2010).
In the present study, the ‘type series’ was examined and found to be in an extremely poor condition (Fig. 20).
Based on the overall morphology, especially the digit tips and webbing, the specimens are similar to Rhacophorus
bipunctatus, which is a fairly common species in Cherrapunji. Metamorphosed individuals similar to those in the
original description, with SVL 12–14 mm, pale green or “Dirty grass green” dorsum, limbs “Dorsally dirty white”,
“coarse granulations on abdomen”, “fingers half webbed”, and “Toes three-fourths webbed”, have also been ob-
served along with foam nests and adults of Rhacophorus bipunctatus at the type locality. Although the tadpoles stud-
ied by Roonwal and Kripalani (1966 “1961”) are not available at ZSIC, their description indicates that they belong
to the genus Rhacophorus. For example, the number of tooth rows on the upper labium in Philautus cherrapunjiae is
five, of which rows 2–5 are divided (Roonwal & Kripalani 1966 “1961”) as observed also in Rhacophorus tadpoles
(for R. kio, Grosjean & Inthara 2016), whereas the tadpoles of Rohanixalus vittatus have two tooth rows of which
row 2 is divided (present study).
Hence, we herein propose to consider Philautus cherrapunjiae Roonwal and Kripalani, 1966 “1961” syn. nov.
a junior subjective synonym of Rhacophorus bipunctatus Ahl, 1927.
Discussion
Our study indicates the confused generic identities of some of the Asian rhacophorid frogs previously allocated to
Chirixalus and Feihyla. The characterization and recognition of Rohanixalus gen. nov. helps clarify the systematics
of this group, which has a complex taxonomic history. Members of Chirixalus, Feihyla, and Rohanixalus are com-
mon on wayside vegetation surrounding human-dominated landscapes, from sea level up to about 2500 m elevation
across South, Southeast, and East Asia. Yet, despite local abundance, their taxonomy had for long been confused
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48 · Zootaxa 4878 (1) © 2020 Magnolia Press
due to the lack of detailed morphological—or rather integrative molecular and morphological—studies. Our results
revealed the following genus-level units within the focal group: (1) Chiromantis Peters, 1854: an exclusively Afri-
can radiation with four currently recognised species (Chen et al. 2020)—C. kelleri, C. petersii, C. rufescens, and C.
xerampelina; (2) Chirixalus Boulenger, 1893, the Asian sister group of Chiromantis (e.g., Frost et al. 2006; Li et al.
2008; Yu et al. 2009; Biju et al. 2010; Hertwig et al. 2012; Meegaskumbura et al. 2015; Poyarkov et al. 2015; Biju
et al. 2016; Chan et al. 2018; Chen et al. 2020; present study), with five members recognised herein—C. doriae,
C. dudhwaensis, C. nongkhorensis, C. simus, and C. trilaksonoi; (3) Feihyla Frost, Grant, Faivovich, Bain, Haas,
Haddad, de S, Channing, Wilkinson, Donnellan, Raxworthy, Campbell, Blotto, Moler, Drewes, Nussbaum, Lynch,
Green, and Wheeler, 2006, an Asian radiation with six revised members (see results) divided into two major species
groups (Feihyla palpebralis group currently restricted to elevations of 800–2300 m asl in China and Vietnam; and
Feihyla vittiger group currently found only in insular Indonesia and Malaysia, from sea level up to ~1300 m asl
elevation); and (4) Rohanixalus gen. nov., the most speciose among these genera, with eight newly allocated mem-
bers (see results), and a wide distribution from sea level up to elevations of ~1700 m asl across Northeast India, the
Andamans, Bangladesh, Myanmar, Thailand, Cambodia, Vietnam, Laos, Malaysia, Indonesia, and Southern China
(Figs. 2, 4; Table 1).
Although our study reassesses the generic allocation of several taxa based on integrative evidence from mor-
phology, reproductive biology, or phylogeny, the placement of Feihyla samkosensis and three Rohanixalus species
(R. marginis, R. nauli, and R. punctatus) requires additional genetic validation. At the same time, several recent
descriptions that are based either on limited sampling (hence overlooking variation) or which lack appropriate
comparisons with closely related taxa (e.g., Chanda & Ghosh 1989; Wilkinson et al. 2003; Mathew & Sen 2009;
Chan et al. 2011; Riyanto & Kurniati 2014), also require further detailed morphological investigation. For example,
R. nauli is potentially a junior subjective synonym of R. baladika, due to lack of reliable morphological differ-
ences and a shallow genetic divergence (0.2–0.6% in 16S) observed between populations from North Sumatra (R.
baladika) and West Sumatra (R. nauli) in our study (Figs. 2, 4). Similarly, R. marginis from Peninsular Malaysia
is geographically and morphologically close to R. punctatus from southwestern Myanmar, especially given the
high degree of variation in colour and markings in these species. Hence, the taxonomic status of R. marginis may
require further validation based on molecular evidence, which is currently lacking for both the taxa. Furthermore,
based on available DNA sequence data (Wilkinson et al. 2002; Yu et al. 2008; Li et al. 2009; Meegaskumbura et
al. 2010; Aowphol et al. 2013; Matsui et al. 2014; Yodthong et al. 2014; Grosjean et al. 2015; Goutte et al. 2016;
present study) five unidentified putative candidate species from regions spanning China, Laos, Myanmar, Thailand,
and Vietnam, currently exist in the new genus Rohanixalus (‘Feihyla hansenae Group II and ‘F. vittata’ Group I in
Aowphol et al. 2013; Rohanixalus cf. hansenae 1, R. cf. hansenae 2, Rohanixalus sp. 1, Rohanixalus sp. 2, and R.
cf. shyamrupus in the present study). Detailed morphological studies of the various putative species within the new
genus, possibly with additional integrative evidence, will be essential to clarify their identity and the taxonomic
status. Considering the presence of undescribed cryptic diversity among previously known populations of the new
genus, future field studies could also potentially lead to the discovery of new taxa from unexplored regions across
its range in Asia. Furthermore, molecular data for other Rohanixalus members will also be required to fully resolve
relationships within this genus.
Morphological differentiation among closely related species is challenging in the genus Rohanixalus, espe-
cially given the high intraspecific morphological variation in body colouration and markings among individuals and
populations of the same species (see taxonomic remarks under species accounts). However, unusually high genetic
diversity is observed among members of the genus (Fig. 2; Table 3); the same was also discussed previously based
on population-level studies of members of the Rohanixalus vittatus group (‘Feihyla vittata’ group in Aowphol et
al. 2013; Yodthong et al. 2014). Hence, additional integrative evidence, especially from bioacoustics and larval
morphology, would prove to be useful for species-level differentiation in the genus Rohanixalus (e.g., Aowphol et
al. 2013), as shown for other widespread anuran groups in Asia comprising morphologically cryptic species (e.g.,
Kuramoto et al. 2007; Grosjean & Dubois 2011; Wang et al. 2014; Rowley et al. 2015; Vassilieva et al. 2016; Garg
& Biju 2017; Garg et al. 2018, 2019; Lyu et al. 2020).
Further sampling should include the enigmatic Rohanixalus vittatus from its type locality, along with generation
of molecular data. This taxon was originally described from Bhamo in Myanmar, and subsequently reported from the
entire range of the genus (e.g., Dutta 1997; Fei 1999; Inger et al. 1999; Deuti & Dutta 2002; Ao et al. 2003; Nguyen
et al. 2005; Stuart 2005; Stuart & Emmett 2006; Bain et al. 2007; Mathew & Sen 2010; Hasan et al. 2010; Li et al.
TREE FROGS OF GENUS FEIHYLA, AND A NEW RELATED GENUS Zootaxa 4878 (1) © 2020 Magnolia Press · 49
2010; Fei et al. 2012). However, our study suggests that R. vittatus is currently restricted to Myanmar and Thailand,
with an additional record from the geographically close Andaman Islands of India. This finding also constitutes the
first report of a rhacophorid from the Andamans, which is indeed surprising given the many fairly recent surveys of
the amphibian fauna of these islands (e.g., Sarkar 1990; Harikrishnan et al. 2012; Harikrishnan & Vasudevan 2018;
Rangasamy et al. 2018). Our finding of R. vittatus from Andamans is both taxonomically and biogeographically
significant, since the occurrence of the family Rhacophoridae on these islands has not been reported previously. The
find reiterates that Andaman amphibians have closer affinities with the fauna of peninsular Myanmar and nearby
regions (Mani 1974; Das 1999). The discovery of a common species from wayside vegetation well within human
inhabitations also indicates that extensive exploration of herpetofauna of these islands is still lacking.
One of the major impediments to comprehensive studies on this group of frogs was also the lack of generic-
level phylogenetic resolution of the ChiromantisChirixalusFeihyla complex. This has led to inappropriate generic
allocation of species, despite known affinities in terms of morphology and reproductive biology. Our reassessment
of genus and species-level taxonomy has found the latter two overlooked aspects reliable for allocations among the
four genera eventually recognised. Further integrative evidence from bioacoustics, larval morphology, and internal
anatomy may reveal additional genus-level synapomorphies, especially in Feihyla and Rohanixalus. At the same
time, a further phylogenetic resolution of intergeneric relationships certainly remains wanting. This could not be
fully achieved in the present work due to the unavailability of genetic data for several known taxa.
The Old World tree frog family Rhacophoridae is known to display a remarkable variety of reproductive modes,
including egg clutch morphology and developmental modes, which have also been used to extend their phenotypes
for purposes genus-level diagnosis (e.g., Hertwig et al. 2013; Meegaskumbura et al. 2015; Biju et al. 2016) par-
ticularly due to the lack of sufficient adult morphological synapomorphies. Among the groups under question in the
present study, Rohanixalus members can be distinguished from those in Chirixalus, Chiromantis, and Feihyla by
their egg clutch morphology, i.e., eggs light greenish, laid in bubble nests in Rohanixalus (vs. creamy white eggs
laid in foam nests in Chirixalus and Chiromantis; and creamy white eggs laid in jelly nests in Feihyla, except mem-
bers of the Feihyla vittiger group that possess light green eggs). Based on available information, the egg clutch mor-
phology is documented in the following Asian members: jelly-nests in Feihyla: F. fuhua (present study, Fig. 4), F.
palpebralis (present study, Fig. 4), F. kajau (Das et al. 2016; Sorokin & Steigerwald 2018), and F. vittiger (Kusrini
et al. 2017; Farits A, personal communication; and present study, Fig. 16); bubble nests in Rohanixalus: R. hansenae
(Poo & Bickford 2013; Poo & Low 2018); R. cf. nauli (Fajar Kaprawi, personal communication, Fig. 7); R. vittatus
(Wan et al. 2012; present study, Figs. 9–11). In contrast, all the Chirixalus members lay eggs in foam nests (Taylor
1962; Biswas 2000; Fei et al. 2009; Banerjee 2014; present study, Fig. 3). Although egg clutch information is cur-
rently lacking for some species (Feihyla inexpectata, Rohanixalus marginis, and R. punctatus), deposition of eggs in
terrestrial jelly nests or bubble nests appears to be reliable phenotypic traits for distinguishing these two genera from
Chirixalus and Chiromantis (Li et al. 2009; Hertwig et al. 2013; present study). Furthermore, many species assigned
to the genus Rohanixalus and the Feihyla vittiger group also show the egg attending behaviour—Rohanixalus: R.
hansenae (Sheridan & Ocopck 2008; Poo & Bickford 2013; Poo & Low 2018), R. vittatus (Wan et al. 2012; present
study, Fig. 11); Feihyla: F. kajau (Sorokin & Steigerwald 2018) and F. vittiger (Kusrini et al. 2017).
Interestingly, the egg clutch morphology also provides evolutionary insights. The nests of Chirixalus and Chiro-
mantis are composed of foam, a characteristic that also suggests their closer affinities to other foam-nesting genera
such as Ghatixalus, Leptomantis, Polypedates, Rhacophorus, Taruga, and Zhangixalus (e.g., Frost et al. 2006; Li
et al. 2009; Yu et al. 2009; Biju et al. 2010; Meegaskumbura et al. 2015; Biju et al. 2016; Chan et al. 2018; Chen
et al. 2020; present study). On the other hand, Rohanixalus frogs lay eggs in a gelatinous mass containing bubbles
(more prominent when freshly laid), whereas Feihyla have jelly-nests with a complete absence of bubbles even
in freshly laid egg clutches. Previous studies on the reproductive modes in Rhacophoridae from a phylogenetic
perspective (e.g, Hertwig et al. 2012; Meegaskumbura et al. 2015) have suggested an evolutionary shift from free-
swimming exotrophic and aquatic larval forms to endotrophic (direct developing) and more terrestrial nidicolous
stages, through gel-nesting ancestors, in order to achieve independence from water bodies during development. The
nesting behaviour in Chirixalus, Chiromantis, Feihyla, and Rohanixalus that varies from absence or presence of
bubbles within the gel to foam nesting may therefore indicate intermediate stages between gel-nesting and foam-
nesting. Future phylogenetic investigation in this regard could shed light on the evolution of reproductive strategies
in rhacophorids.
BIJU ET AL.
50 · Zootaxa 4878 (1) © 2020 Magnolia Press
Acknowledgments
We thank the following for permission to use their photographs: Anton Sorokin (Feihyla kajau eggs in Figs. 18C,
18D), Chan Kin Onn (Rohanixalus marginis in Fig. 5B), Fajar Kaprawi (Rohanixalus cf. nauli in Figs. 1, 5C–E),
Farits Alhadi (Feihyla vittiger in Figs. 1, 18G–M), Haegel Alif (Feihyla vittiger eggs in Fig. 18F), John Sullivan
(Feihyla kajau in Fig. 18B), Lars Fehlandt (Feihyla inexpectata in Figs. 1, 18A and F. kajau in Fig. 1), and Nguy-
en Thanh Luan (Feihyla palpebralis in Figs. 1, 3H). For field support and sampling, we thank A.N.D Akalabya
Sarmah, Abhijit Das, Rachunliu G. Kamei, and Stephen Mahony; Robin Suyesh for help in the acoustic analysis;
David C. Blackburn and Jens Vindum (California Academy of Science, USA) for providing tissues; Kailash Chan-
dra, Gopinathan Maheswaran, and Kaushik Deuti for access to specimens (ZSI, Kolkata) and museum support to
SDB and SG; Barry Clarke and David Gower (NHM, London) for access to specimens and museum support to
SDB; the State Forest Departments in India (Manipur, Tripura, Nagaland, and Arunachal Pradesh) for study permits
to SDB; Department of Environment and Forests, Andaman and Nicobar Islands, India for study permission and
logistic support to SC; Indonesian Institute of Sciences (LIPI) and Ministry of Environment and Forestry, Indonesia
for fieldwork permit and funding through the Indonesian Fauna Barcoding Grant (DIPA Puslit Biologi LIPI 2018)
to AH; Nongkham Subdistrict Administrative Organization, Royal Irrigation Department, Institute of Animal for
Scientific Purposes Development, and Chulalongkorn University Museum of Natural History (Thailand) for study
permission and assistance to PT; two anonymous reviewers and the Editor (Miguel Vences) for their helpful com-
ments and suggestions. SC thanks the Director, ZSI, Kolkata for encouragement and support; GG and SC thank SR
Chandramouli for initial support. This study was partially supported by the following grants to SDB: University of
Delhi Research and Development Grants between 2007–2016; DU-DST Purse Grants Phase I and Phase II, Depart-
ment of Science and Technology, Ministry of Science and Technology, Government of India. SG receives fellowship
from the Council for Scientific and Industrial Research, Government of India, as Research Associate, CSIR No.
9/45(1381)/2015-EMR-I. PT was supported by Grants for Development of New Faculty Staff and Ratchadaphisek-
somphot Endowment Fund for field surveys in Thailand. JR was supported by National Natural Science Foundation
of China Grant 31722049 to Jiatang Li. CG and KB receive Junior Research Fellowships from the University Grant
Commission, Government of India, for their Ph.D. programmes.
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... The early documented amphibian surveys on the islands of Andaman and Nicobar during the 19th century (Fitzinger 1860;Stoliczka 1870) and 20th century (e.g., Annandale 1917;Cherchi 1954;Pillai 1977Pillai 1991Mansukhani and Sarkar 1980;Mehta and Rao 1987;Sarkar 1990;Das 1994Das 1996Das 1998Daniels and David 1996) largely recorded the occurrence of species found in the neighbouring biogeographical regions, with sporadic descriptions of new taxa. The past nearly two decades have seen progress in description of new species and genera, new family records, and taxonomic clarifications, particularly with the aid of molecular techniques (e.g., Chandramouli et al. 2016Chandramouli et al. 2020aHarikrishnan and Vasudevan 2018;Biju et al. 2020). The recent surveys on these islands suggest the occurrence of 20 species representing 15 genera and five families (e.g., Harikrishnan and Vasudevan 2018;Rangasamy et al. 2018;Biju et al. 2020;Chandramouli et al. 2020a). ...
... The past nearly two decades have seen progress in description of new species and genera, new family records, and taxonomic clarifications, particularly with the aid of molecular techniques (e.g., Chandramouli et al. 2016Chandramouli et al. 2020aHarikrishnan and Vasudevan 2018;Biju et al. 2020). The recent surveys on these islands suggest the occurrence of 20 species representing 15 genera and five families (e.g., Harikrishnan and Vasudevan 2018;Rangasamy et al. 2018;Biju et al. 2020;Chandramouli et al. 2020a). Of these, 12 species are found in the Andaman group of islands, with five species and one genus known to be endemic. ...
Article
Full-text available
Recent studies on frogs of the Microhyla heymonsi species complex have demonstrated that high genetic variation exists among its various known populations from regions across Asia. We assessed the taxonomic identity of the Nicobar population of Microhyla cf. heymonsi and compared it to the typical Microhyla heymonsi from Taiwan and the two recently described species in this complex from Vietnam. Our study demonstrates that the Nicobar population is both genetically and morphologically divergent and warrants recognition as a new species, which we formally describe here as Microhyla nakkavaram sp. nov. The new species is closely related to M. daklakensis, M. heymonsi, and M. ninhthuanensis, but diagnosable from all three species by a suite of morphological characters, such as the presence of two small tubercles at mid-dorsum along with ( )-shaped markings, length of finger I longer than half the length of finger II, presence of three distinct metacarpal tubercles on hand, rudimentary foot webbing, as well as its slender body shape, granular dorsal skin texture, and other colour characters and body markings. Statistical analyses based on multiple morphometric characters also clearly separate our new taxon from M. heymonsi, with which it was previously confused. Our phylogenetic analyses based on the mitochondrial 16S rRNA locus find Microhyla nakkavaram sp. nov. to be nested in the Microhyla achatina species group, where it is delimited as a distinct species. This lineage shows genetic distances of ≥ 3.5% from all the other known congeners. Currently, the known distribution of the new taxon is restricted to the southernmost group of Nicobar Islands-Great Nicobar, Kondul, and Little Nicobar-where it is found in abundance across a wide range of habitats during the monsoon season. Our study underlines the need to reassess the identity of all the known populations of M. 'heymonsi' from other regions in Southeast and East Asia. To facilitate future taxonomic work in the light of our and other recent findings, we also provide a detailed redescription and revised diagnosis for M. heymonsi based on morphological examination of its century-old type material originating from Taiwan.
... The early documented amphibian surveys in the islands of Andaman and Nicobar during the 19th century (Fitzinger 1860;Stoliczka 1870) and 20th century (e.g., Annandale 1917;Cherchi 1954;Pillai 1977Pillai 1991Mansukhani and Sarkar 1980;Mehta and Rao 1987;Sarkar 1990;Das 1994Das 1996Das 1998Daniels and David 1996) largely recorded the occurrence of species found in the neighbouring biogeographical regions, with sporadic descriptions of new taxa. The past nearly two decades have witnessed progress in description of new species and genera, new family records, and taxonomic clarifications, particularly with the aid of molecular techniques (e.g., Chandramouli et al. 2016Chandramouli et al. 2020aChandramouli et al. 2020bHarikrishnan and Vasudevan 2018;Biju et al. 2020). The recent surveys on these islands suggest the occurrence of 20 species representing 15 genera and five families (e.g., Harikrishnan and Vasudevan 2018;Rangasamy et al. 2018;Biju et al. 2020;Chandramouli et al. 2020a). ...
... The past nearly two decades have witnessed progress in description of new species and genera, new family records, and taxonomic clarifications, particularly with the aid of molecular techniques (e.g., Chandramouli et al. 2016Chandramouli et al. 2020aChandramouli et al. 2020bHarikrishnan and Vasudevan 2018;Biju et al. 2020). The recent surveys on these islands suggest the occurrence of 20 species representing 15 genera and five families (e.g., Harikrishnan and Vasudevan 2018;Rangasamy et al. 2018;Biju et al. 2020;Chandramouli et al. 2020a). Of these, 12 species are found in the Andaman group of islands, with five species and one genus known to be endemic. ...