Saturday, November 14, 2020

[Herpetology • 2020] Rohanixalus gen. nov. • New Insights on the Systematics and Reproductive Behaviour in Tree Frogs (Anura, Rhacophoridae) of the Genus Feihyla, with Description of A New Related Genus from Asia


Rohanixalus vittatus (Boulenger, 1887) comb. nov.
 from Andaman Islands, India.

in Biju, Garg, Gokulakrishnan, Chandrakasan, Thammachoti, et al., 2020.
photo: S.D. Biju  facebook.com/BijuSDas 

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 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.

Keywords: Chiromantis, Chirixalus, egg attendance, gel-nesting, generic reallocation, integrative taxonomy, larval morphology, molecular phylogeny, Rohanixalus gen. nov., species groups


Class Amphibia Linnaeus, 1758 
Order Anura Fischer von Waldheim, 1813 

Family Rhacophoridae Hoffman, 1932 
Subfamily Rhacophorinae Hoffman, 1932

Genus Feihyla 

Original description. Feihyla Frost, et al., 2006
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 nomen 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).

Feihyla palpebralis group
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
 
Feihyla palpebralis (Smith, 1924)
Vietnamese Jelly-nest Frog
Philautus palpebralis Smith, 1924.


Feihyla vittiger group
Species included. Feihyla inexpectata (Matsui, Shimada, and Sudin, 2014), Feihyla kajau (Dring, 1983), Feihyla 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
Chiromantis inexpectatus Matsui, Shimada, and Sudin, 2014

Feihyla kajau (Dring, 1983)
White-eared Jelly-nest Frog
 Rhacophorus kajau Dring, 1983. Dring J. C. M. 1983.

Feihyla samkosensis (Grismer, Thy, Chav, and Holden, 2007) comb. nov.
Samkos Jelly-nest Frog
Chiromantis samkosensis Grismer, Thy, Chav, and Holden, 2007.

Feihyla vittiger (Boulenger, 1897) comb. nov.
Javan Jelly-nest Frog
 Ixalus vittiger Boulenger, 1897.





Genus Rohanixalus gen. nov.

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.)

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 NP, 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.
photos: Farits Haegel Alif Fajar (Indonesia), Chan Kin Onn (Singapore).

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, Grismer, 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.; Rohanixalus senapatiensis (Mathew and Sen, 2009) comb. nov.; Rohanixalus shyamrupus (Chanda and Ghosh, 1989) comb. nov.; Rohanixalus vittatus (Boulenger, 1887) comb. nov.


Rohanixalus vittatus in life, from Andaman Islands, India.




ปาดจิ๋วพม่า Rohanixalus vittatus (Boulenger, 1887)
ปาดจิ๋วศรีราชา Rohanixalus hansenae (Cochran, 1927) 
 ปาดจิ๋วมลายู Rohanixalus marginis (Chan, Grismer, Anuar, et al., 2011)

S. D. Biju, Sonali Garg, G. Gokulakrishnan, Sivaperuman Chandrakasan, Panupong Thammachoti, Jinlong Ren, C. Gopika, Karan Bisht, Amir Hamidy and Yogesh Shouche. 2020. 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). Zootaxa. 4878(1); 1–55. DOI: 10.11646/zootaxa.4878.1.1