Sorokin, Anton

Steigerwald, Emma

English

Cadernos Espinosanos (E-Journal)

127Phyllomedusa - 17(1), June 2018Received 28 July 2017Accepted 08 March 2018Distributed June 2018Short CommuniCationEvidence for parental care in Feihyla kajau (Anura: Rhacophoridae)Anton Sorokin1 and Emma Steigerwald21 Department of Biology, East Carolina University, NC, USA. E-mail: antonsrkn@gmail.com.2 Department of Environmental Science, Policy, and Management, University of California Berkeley, CA, USA.Keywords: anurans, Borneo, brooding, convergence, gel nesting, maternal care.Palavras-chave: anuros, Borneo, convergência, cuidado maternal, nidificação em gel, ninhada.Phyllomedusa 17(1):127–130, 2018© 2018 Universidade de São Paulo - ESALQ ISSN 1519-1397 (print) / ISSN 2316-9079 (online)doi: http://dx.doi.org/10.11606/issn.2316-9079.v17i1p127-130Rhacophoridae is a large (419 species) family of Old World tree frogs that have a variety of reproductive modes (AmphibiaWeb 2018). They reproduce by aquatic breeding, direct devel-opment, or laying eggs in foam nests or terrestrial “gel nests” (Meegaskumbura et al. 2015). The basal state to direct development and foam nesting in rhacophorids may be gel-nesting, in which terrestrial eggs are sheathed in a thick jelly (Meegaskumbura et al. 2015). Feihyla kajau (Dring, 1983), a Bornean endemic (Inger et al. 2004), is the only representative of the exclusively Southeast Asian genus Feihyla found on the island (Hertwig et al. 2013, AmphibiaWeb 2018). Like other rhacophorids with arboreal gel nests, F. kajau is relatively small and lays small egg clutches (Meegaskumbura et al. 2015). To date, there is no available information on parental egg attendance in this species.Feihyla kajau is remarkably similar to Neotropical glass frogs (Centrolenidae) in both appearance and behavior, despite the phylo-genetic distance between them. The resemblance of these taxa may be an example of convergent evolution between distantly related species that occupy a similar niche space. Like centrolenids, F. kajau is a small, arboreal frog that deposits gel-encapsulated eggs on vegetation overhanging streams (Figure 1); after hatching, the tadpoles complete development in the water (Ruiz-Carranza and Lynch 1991, Das et al. 2016). Centrolenids exhibit post-zygotic parental care of eggs after they are deposited (Vockenhuber et al. 2009, Delia et al. 2013, 2017)—a behavior heretofore unknown in F. kajau. Here we describe apparent parental care in F. kajau.Observations were made on 29 August 2014 in Sabah, Borneo (04°57'56.1'' N, 117°48'09.1'' E, WGS 84, 163 m a.s.l.). At 21:24 h, an amplectant pair of F. kajau was observed on the underside of a leaf (Figure 2A). Returning to the site 53 min later, we observed that the female had remained with the eggs, clasping them with 128Phyllomedusa - 17(1), June 2018her back feet (Figure 2B), but the male frog was not present. Multiple egg clutches at different stages of development were observed at this same site on five separate visits; however, no other parental care was observed.Possibly, F. kajau only attends the eggs for a short time after oviposition, as do some centrolenids that only brood for the remainder of the night of oviposition (Delia et al. 2017). Among glass frogs with first-night only care, it is the females who attend the eggs (Delia et al. 2017). Our observation of a female F. kajau, then, remains consistent in its parallel to Centrolenid behavior.Behaviors such as egg attendance and brooding have been repeatedly linked with egg survival in a wide range of gel-nesting species (Vockenhuber et al. 2009, Poo and Bickford 2013, Delia et al. 2013, 2017). Post-fertilization parental care has been recorded in approximately one tenth of frog species worldwide (Lehtinen and Nussbaum 2003). Although parental care remains sparsely recorded in Southeast Asian frogs (Poo and Bickford 2013), it is known in species closely related to F. kajau, such as F. (Chiromantis) hansenae (Cochran, 1927) (Sheridan and Ocock 2008, Poo and Bickford 2013) and F. vittata (Boulenger, 1887) (Wan et al. 2012). In these three species, the parent positions the posterior portion of its body and hind legs over the egg clutch.Arthropods, including katydids, ants, and wasps, are significant predators on gel-nesting frogs, accounting for as much as 90% of egg Figure 1.  Eggs of Feihyla kajau at varying levels of development. (A) A younger clutch. (B) A more developed clutch.A BSorokin and Steigerwald129Phyllomedusa - 17(1), June 2018mortality (Vockenhuber et al. 2009, Poo and Bickford 2013, Delia et al. 2017). There are no records of arthropods depredating eggs of F. kajau, but signs of invertebrate predation have been observed in egg clutches of F. kajau proximal to the present observation (A. Sorokin pers. obs.). In the closely related F. hansenae, there is intense arthropod predation pressure, which is the single largest cause of egg mortality (Poo and Bickford 2013). However, even a brief period of egg attendance can increase egg survivorship. In centrolenids, just a few hours of brooding provides eggs with days of protection against arthropods (Delia et al. 2017). Furthermore, brooded eggs are more likely to avoid desiccation during a dry period of weather, another principal cause of mortality for developing eggs. Attending parents hydrate their eggs through fluid transfer by contact with a highly vascularized patch below their pelvis (Delia et al. 2013, 2017).We still have much to learn about frog parental care, especially the tradeoffs shaping the evolution of this complex behavior. Little is known about this behavior in Southeast Asian frogs, with this note being the only observation in F. kajau. Further investigation of breeding in this species, particularly continuous observations immediately following oviposition, must be conducted to confirm that the behavior here observed constitutes parental care.Acknowledgments.—We thank Alys Granados for the opportunity to assist in her fieldwork in Borneo, Chien Lee for bringing the significance of the observation to our attention, and two anonymous reviewers for their helpful suggestions and edits.  A BFigure 2. (A) A pair of Feihyla kajau in amplexus on the underside of a leaf in Sabah, Borneo. (B) The female remains, attending the eggs and clasping them with her hind feet.Evidence for parental care in Feihyla kajau130Phyllomedusa - 17(1), June 2018ReferencesAmphibiaWeb. 2018. Amphibia Web: Information on amphibian biology and conservation. Eletronic Database accessible at https://amphibiaweb.org. University of California, Berkeley, USA. Captured on 13 June 2018.Das, I., H. Hedeir, Y. M. Pui, S. T. Hertwig, and A. Haas. 2016. Larval external morphology and development in Feihyla kajau (Dring, 1983) (Amphibia: Anura: Rhaco-phoridae). Raffles Bulletin of Zoology 64: 319–328.Delia, J., L. Bravo-Valencia, and K. M. Warkentin. 2017. Patterns of parental care in Neotropical glassfrogs: fieldwork alters hypotheses of sex-role evolution. Journal of Evolutionary Biology 30: 898–914.Delia, J., A. Ramírez-Bautista, and K. Summers. 2013. Parents adjust care in response to weather conditions and egg dehydration in a Neotropical glassfrog. Behavioral Ecology and Sociobiology 67: 557–569.Hertwig, S. T., M. Schweizer, I. Das, and A. Haas. 2013. Diversification in a biodiversity hotspot. The evolution of Southeast Asian rhacophorid tree frogs on Borneo (Amphibia: Anura: Rhacophoridae). Molecular Phylo-genetics and Evolution 68: 567–581.Inger, R., Iskandar, D., Das, I., Stuebing, R., Lakim, M., Yambun, P., & Mumpuni. 2004. Feihyla kajau. The IUCN Red List of Threatened Species. Electronic Database accessible at http://www.iucnredlist.org/details/58999/0. Captured on 12 July 2017.Lehtinen, R. M. and R. A. Nussbaum. 2003. Parental care: a phylogenetic perspective. Pp. 342–386 in B. G. M. Jamieson (ed.), Reproductive Biology and Phylogeny of Anura. Enfield. Science Publishers, Inc.Meegaskumbura, M., G. Senevirathne, S. D. Biju, S. Garg, S. Meegaskumbura, R. Pethiyagoda, J. Hanken, and C. J. Schneider. 2015. Patterns of reproductive-mode evolution in Old World tree frogs (Anura, Rhacophoridae). Zoologica Scripta 44: 509–522.Poo, S. and D. P. Bickford. 2013. The adaptive significance of egg attendance in a South-East Asian tree frog. Ethology 119: 671–679.Ruiz-Carranza, P.M. and J.D. Lynch. 1991. Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificacion generica. Lozania 57: 1–32.Sheridan, J. A. and J. F. Ocock. 2008. Parental care in Chiromantis hansenae (Anura: Rhacophoridae). Copeia 2008: 733–736.Vockenhuber, E. A., W. Hödl, and A. Amézquita. 2009. Glassy fathers do matter: egg attendance enhances embryonic survivorship in the glass frog Hyalino-batrachium valerioi. Journal of Herpetology 43: 340–344.Wan, J. P., Y. H. Sung, B. P. L. Chan, and G. Lu. 2012. Chiromantis vittatus (Two-striped Pigmy Tree Frog). Parental Care. Herpetological Review 43: 630–631.Editor: Jaime BertoluciSorokin and Steigerwald

Evidence for parental care in Feihyla kajau (Anura: Rhacophoridae)

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Evidence for parental care in Feihyla kajau (Anura: Rhacophoridae)

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