Leaping frogs (Anura: Ranixalidae) of the Western Ghats of India: An integrated taxonomic review

Main Article Content

Neelesh Dahanukar
http://orcid.org/0000-0001-7162-9023
Nikhil Modak
http://orcid.org/0000-0003-1838-464X
Keerthi Krutha
http://orcid.org/0000-0003-3664-3924
P. O. Nameer
http://orcid.org/0000-0001-7110-6740
Anand D. Padhye
http://orcid.org/0000-0002-5971-4759
Sanjay Molur
http://orcid.org/0000-0001-8230-518X

Abstract

Leaping frogs of the family Ranixalidae are endemic to the Western Ghats of India and are currently placed in a single genus, Indirana.  Based on specimens collected from their entire range and a comprehensive study of type material defining all known species, we propose a revised taxonomy for the leaping frogs using an integrative approach including an analysis of the mitochondrial 16S rRNA and nuclear rhodopsin genes, as well as multivariate morphometrics. Both genetic and morphological analyses suggest that the genus Indirana is paraphyletic and a distinct monophyletic group, Walkerana gen. nov is described herein.  The new genus is separated from Indirana sensu stricto by an apomorphic character state of reduced webbing, with one phalange free on the first and second toe (vs. no free phalanges), two phalanges free on the third and fifth toe (vs. one free phalange), and three phalanges free on the fourth toe (vs. 2–2½ phalanges free).  This review includes (i) identification of lectotypes and redescription of three species of the genus Walkerana; (ii) identification of lectotypes for Indirana beddomii and I. semipalmata and their redescription; (iii) redescription of I. brachytarsus and I. gundia; and (iv) descriptions of four new species, namely, I. duboisi and I. tysoni from north of the Palghat gap, and I. yadera and I. sarojamma from south of the Palghat gap; and (iv) a key to the genera and species in the family Ranixalidae.

 

Article Details

Section
Monograph
Author Biographies

Neelesh Dahanukar, Indian Institute of Science Education and Research (IISER), G1 Block, Dr. Homi Bhabha Road, Pashan, Pune, Maharashtra 411008, India and Systematics, Ecology and Conservation Laboratory, Zoo Outreach Organization (ZOO), 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India

Neelesh Dahanukar works in ecology and evolution with an emphasis on mathematical and statistical analysis. He is also interested in taxonomy, distribution patterns and molecular phylogeny of fish and frogs. 

 

Nikhil Modak, Department of Biodiversity, MES Abasaheb Garware College, Karve Road, Pune, Maharashtra 411004, India

Nikhil Modak is a doctoral student interested in ecology, systematics and evolution of amphibians of the Western Ghats.  

 

Keerthi Krutha, Wildlife Information Liaison Development (WILD) Society, 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India

Keerthi Krutha works on ecology and conservation of Indian herpetofauna. 

 

P. O. Nameer, Centre for Wildlife Studies, College of Forestry, Kerala Agricultural University, Thrissur, Kerala 680656, India

P.O. Nameer, is a Professor and Head. His research interest include the taxonomy, biogeography and ecology of the vertebrates except fishes. 

 

Anand D. Padhye, Department of Zoology, MES Abasaheb Garware College, Karve Road, Pune, Maharashtra 411004, India and Institute of Natural History Education and Research (INHER), B1/602 Kumar Parisar, Kothrud, Pune, Maharashtra 411038, India

Anand D. Padhye is an Associate Professor working on systematics, ecology, diversity, distribution and evolution of amphibians. He is also secretary of Institute of Natural History Education and Research (INHER), Pune.

 

Sanjay Molur, Systematics, Ecology and Conservation Laboratory, Zoo Outreach Organization (ZOO), 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India and Wildlife Information Liaison Development (WILD) Society, 96 Kumudham Nagar, Vilankurichi Road, Coimbatore, Tamil Nadu 641035, India

Sanjay Molur works on species assessment and conservation.  His interests are in taxonomy, promoting scientific publications and in science communication.

 

References

Altschul, S.F., W. Gish, W. Miller, E.W. Myers & D.J. Lipman (1990). Basic local alignment search tool. Journal of Molecular Biology 215: 403–410; http://dx.doi.org/10.1016/S0022-2836(05)80360-2

Andrews, M.I., S. George & J. Joseph (2005). Amphibians in protected areas of Kerala. Zoos’ Print Journal 20(4): 1823–1831; http://dx.doi.org/10.11609/JoTT.ZPJ.1151b.1823-31

Annadale, N. (1909). Notes on Indian batrachia. Records of the Indian Museum 3: 282–286.

Biju, S.D. & S. Dutta (2004). Indirana leptodactyla. The IUCN Red List of Threatened Species 2004: e.T58312A11763407. Downloaded on 07 January 2016;

http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58312A11763407.en

Biju, S.D., S. Dutta, R. Inger & M.S. Ravichandran (2004). Indirana diplosticta. The IUCN Red List of Threatened Species 2004: e.T58309A11762447. Downloaded on 07 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58309A11762447.en

Biju, S.D. (2001). A synopsis to the frog fauna of the Western Ghats, India. Occasional Publications of the Indian Society for Conservation Biology 1: 1–24.

Biju, S.D. & S. Dutta (2004). Indirana leptodactyla. The IUCN Red List of Threatened Species 2004: e.T58312A11763407. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58312A11763407.en

Biju, S.D., S. Dutta & M.S. Ravichandran (2004a). Indirana beddomii. The IUCN Red List of Threatened Species 2004: e.T58307A11761967. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58307A11761967.en

Biju, S.D., S. Dutta & R. Inger (2004b). Indirana brachytarsus. The IUCN Red List of Threatened Species 2004: e.T58308A11762210. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58308A11762210.en

Biju, S.D., S. Dutta & R. Inger (2004c). Indirana gundia. The IUCN Red List of Threatened Species 2004: e.T58310A11762967. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58310A11762967.en

Biju, S.D., S. Dutta & R. Inger (2004d). Indirana longicrus. The IUCN Red List of Threatened Species 2004: e.T58313A11763661. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58313A11763661.en

Biju, S.D., S. Dutta & R. Inger (2004e). Indirana semipalmata. The IUCN Red List of Threatened Species 2004: e.T58315A11764064. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58315A11764064.en

Biju, S.D., S. Dutta & R. Inger (2004f). Indirana tenuilingua. The IUCN Red List of Threatened Species 2004: e.T58316A11764319. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58316A11764319.en

Biju, S.D., S. Dutta, A. Padhye & R. Inger (2004g). Indirana leithii. The IUCN Red List of Threatened Species 2004: e.T58311A11763222. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58311A11763222.en

Biju, S.D., S. Dutta, R. Inger & M.S. Ravichandran (2004h). Indirana diplosticta. The IUCN Red List of Threatened Species 2004: e.T58309A11762447. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58309A11762447.en

Biju, S.D., S.P. Vijayakumar & S. Dutta (2004i). Indirana phrynoderma. The IUCN Red List of Threatened Species 2004: e.T58314A11763836. Downloaded on 26 January 2016; http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.T58314A11763836.en

Bossuyt, F. & A. Dubois (2001). A review of the frog genus Philautus Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae). Zeylanica 6(1): 1–112.

Boulenger, G.A. (1882). Catalogue of the Batrachia Salientia S. Ecaudata in the Collection of the British Museum. 2nd Edition. Taylor and Francis, London. 495 pp., Plates I-XXX.

Boulenger, G.A. (1888). Descriptions of two new species of Rana. Annals and Magazine of Natural History 6(2): 506–508.

Boulenger, G.A. (1890). The fauna of British India including Ceylon and Burma. Reptilia and Batrachia. Taylor and Francis, 541pp.

Boulenger, G.A. (1918). On the Papuan Melanesian and North Australian species of the genus Rana. Annals and Magazine of Natural History 1(3): 236–242; http://dx.doi.org/10.1080/00222931808562307

Boulenger, G.A. (1920). A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum 20: 1–226.

Chanda, S.K. & K. Deuti (1997). Endemic amphibians of India. Records of the Zoological Survey of India 96: 63–79.

Chanda, S.K. (2002). Handbook of Indian Amphibians. Zoological Survey of India, Kolkata, 335 pp.

Dahanukar, N., K. Krutha, M.S. Paingankar, A.D. Padhye, N. Modak & S. Molur (2013). Endemic Asian chytrid strain infection in threatened and endemic anurans of the Northern Western Ghats, India. PLoS ONE 8(10): e77528; http://dx.doi.org/10.1371/journal.pone.0077528

Daniel, J.C. & A.G. Sekar (1989). Field guide to the amphibians of western India Part 4. Journal of the Bombay Natural History Society 86 (2): 194–202.

Dinesh, K.P., C. Radhakrishnan, K.V. Gururaja & G. Bhatta (2009). An annotated checklist of Amphibia of India with some insights into the pattern of species discoveries, distribution and endemism. Records of the Zoological Survey of India, Occasional Paper 302: 1–133.

Doornik, J.A. & H. Hansen (2008). An omnibus test for univariate and multivariate normality. Oxford Bulletin of Economics and Statistics 70: 927–939; http://dx.doi.org/10.1111/j.1468-0084.2008.00537.x

Dubois, A. (1986). Diagnose préliminaire d’un nouveau genre de Ranoidea (Amphibiens, Anoures) du sud de l’Inde. Alytes 4: 113–118.

Dubois, A. (1987a). Miscellanea taxinomica batrachologica (I). Alytes 5 (1–2): 7–95.

Dubois, A. (1987b). Miscellanea nomenclatorica batrachologica (XV). Alytes 5 (4): 175–176.

Dubois, A. (1992). Notes sur la classification des Ranidae (Amphibiens Anoures). Bulletin Mensuel de la Société Linnéenne de Lyon 61(10): 305–352.

Dubois, A. (1984). Note préliminaire sur le groupe de Rana limnocharis Gravenhorst, 1829 (Amphibiens, Anoures). Alytes 3: 143–159.

Dubois, A. (2003). True frogs (Ranidae), pp. 245–264.  In: Duellman, W.E. (ed.). Grzimek’s Animal Life Encyclopedia, Second Edition. Volume 6. Amphibians. Gale Group, Detroit.

Dutta, S.K. (1997). Amphibians of India and Sri Lanka. Odyssey Publishing House, Bhubaneshwar, 342pp+xxii.

Edgar, R.C. (2004). MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797; http://dx.doi.org/10.1093/nar/gkh340

Fischer, C.E.G. (1915). The habits of Rana semipalmata, Boulenger. Journal of the Bombay Natural History Society 24: 194.

Gopalan, S.V., A. Nair, K.S. Kumar, J. Merilä & S. George (2012). Morphology of Indirana semipalmata (Boulenger, 1882) (Amphibia; Anura) adults and tadpoles from the Western Ghats, India. Herpetology Notes 5: 263–273.

Günther, A. (1876). Third Report on collections of Indian reptiles obtained by the British Museum. Proceedings of the Zoological Society of London 1875: 567–577

Gururaja, K.V., S. Ali, G.R. Rao & T.V. Ramachandra (2008). Influence of land use changes in river basins on diversity and distribution of amphibians, pp. 94–112. In: Ramachandra, T.V. (eds.). Environment Education for Ecosystem Conservation. Capital Publishing Company, New Delhi.

Gururaja, K.V., S. Ali, G.R. Rao, V.D. Mukri & T.V. Ramachandra (2007). Biodiversity and ecological significance of Gundia river catchment. CES Technical Report 116: 1–26.

Hammer, Ø., D.A.T. Harper & P.D. Ryan (2001). Past: Paleontological Statistics Software Package for education and data analysis. Paleontología Electrónica 4: 1–9. http://palaeo-electronica.org/2001_1/past/issue1_01.html

Harris, R.J. (2001). A primer for multivariate statistics. Third Edition. Lawrence Erlbaum Associates Publishers, London.

Huberty, C.J. & S. Olejnik (2006). Applied MANOVA and Discriminant Analysis. 2nd Edition. Hohn Wiley and Sons, N.J., 488pp; http://dx.doi.org/10.1002/047178947X

Inger, R.F. & S.K. Dutta (1986). An overview of the amphibian fauna of India. Journal of the Bombay Natural History Society 83: 135–146.

Inger, R.F., H.B. Shaffer, M. Koshy & R. Bakde (1984). A report on a collection of amphibians and reptiles from the Ponmudi, Kerala, South India. Journal of the Bombay Natural History Society 81: 406–427.

Jesmina, A.S. & S. George (2015). New distribution records for the critically endangered frog Indirana gundia (Dubois, 1986) from Kerala part of Western Ghats, India. Biodiversity Data Journal 3: e5825; http://dx.doi.org/10.3897/BDJ.3.e5825

Johnsingh, A.J.T. (2001). The Kalakad–Mundanthurai Tiger Reserve: a global heritage of biological diversity. Current Science 80(3): 378–388.

Krishnamurthy, S.V. (2003). Amphibian assemblages in undisturbed and distrubed areas of Kudremukh National Park, central Western Ghats, India. Environmental Conservation 30(3): 274–282.

Krishnamurthy, S.V. & S. Katre (1993). Amphibian fauna of Sringeri Taluk (Chickamagalure District: Karnataka). Journal of Indian Institute of Science 73: 443–452.

Kumar, A., R. Chellam, B.C. Choudhury, D. Mudappa, K. Vasudevan, N.M. Ishwar & B.R. Noon (2002). Impact of rainforest fragmentation on small mammals and herpetofauna in the Western Ghats, South India. Final Technical Report, Wildlife Institute of India.

Kuramoto, M. & S.H. Joshy (2001). Scanning electron microscopic studies on spermatozoa of anurans from India and Sri Lanka. Amphibia-Reptilia 22: 303–308;

http://dx.doi.org/10.1163/156853801317050098

Laurent, R.F. (1986). Sous-classe des lissamphibiens (Lissamphibia). Systématique. pp. 594–797. In: Grassé, P. & M. Delsol (eds.). Traité de zoologie: anatomie, systématique, biologie, vol. XIV, Batraciens, fas. 1B.

Meier, R., G. Zhang & F. Ali (2008). The use of mean instead of smallest interspecific distances exaggerates the size of the “barcoding gap†and leads to misidentification. Systematic Biology 57: 809–813. http://dx.doi.org/10.1080/10635150802406343

Meyer, C.P. & G. Paulay (2005). DNA Barcoding: Error Rates Based on Comprehensive Sampling. PLoS Biology 3(12): e422; http://dx.doi.org/10.1371/journal.pbio.0030422

Minh, B.Q., M.A.T. Nguyen & A. von Haeseler (2013). Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30: 1188–1195;

http://dx.doi.org/10.1093/molbev/mst024

Modak, N., A. Padhye & N. Dahanukar (2014). Delimiting the distribution range of Indirana leithii (Boulenger, 1888) (Anura: Ranixalidae), an endemic threatened anuran of the Western Ghats, based on molecular and morphological analysis. Zootaxa 3796(1): 62–80; http://dx.doi.org/10.11646/zootaxa.3796.1.3

Modak, N., N. Dahanukar, N. Gosavi & A.D. Padhye (2015). Indirana salelkari, a new species of leaping frog (Anura: Ranixalidae) from Western Ghats of Goa, India. Journal of Threatened Taxa 7(9): 7493–7509; http://dx.doi.org/10.11609/JoTT.o4262.7493-509

Molur, S., K. Krutha, M.S. Paingankar & N. Dahanukar (2015). Asian strain of Batrachochytrium dendrobatidis is widespread in the Western Ghats, India. Diseases of Aquatic Organisms 112: 251–255; http://dx.doi.org/10.3354/dao02804

Myers, C.W. & W.E. Duellman (1982). A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Novitiates 2752: 1–32.

Nair, A., O. Daniel, S.V. Gopalan, S. George, K.S. Kumar, J. Merilä & A.G.F. Teacher (2011). Infectious disease screening of Indirana frogs from the Western Ghats biodiversity hotspot. Herpetological Review 42(4): 554–557.

Nair, A., S.V. Gopalan, S. George, K.S. Kumar, A.G.F. Teacher & J. Merilä (2012a). High cryptic diversity of endemic Indirana frogs in the Western Ghats biodiversity hotspot. Animal Conservation 15(5): 489–498; http://dx.doi.org/10.1111/j.1469-1795.2012.00539.x

Nair, A., S.V. Gopalan, S. George, K.S. Kumar, A.G.F. Teacher & J. Merilä (2012b). Endemic Indirana frogs of the Western Ghats biodiversity hotspot. Annales Zoologici Fennici 49: 257–286; http://dx.doi.org/10.5735/086.049.0501

Nei, M. & S. Kumar (2000). Molecular Evolution and Phylogenetics. Oxford University Press, New York, 333pp.

Nguyen, L.-T., H.A. Schmidt, A. von Haeseler & B.Q. Minh (2015). IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution 32: 268-274; http://dx.doi.org/10.1093/molbev/msu300

Padhye, A.D. & H.V. Ghate (2002). An overview of amphibian fauna of Maharashtra State. Zoos’ Print Journal 17(3): 735–740; http://dx.doi.org/10.11609/JoTT.ZPJ.17.3.735-40

Padhye, A.D. & H.V. Ghate (2012). Amphibia. Fauna of Maharashtra, Zoological Survey of India State Fauna Series 20 (1): 239–246.

Padhye, A.D., M. Mahabaleshwarkar & H.V. Ghate (2002). An overview of amphibian fauna of Pune District with special reference to their status in and around Pune City. Zoos’ Print Journal 17(4): 757–763; http://dx.doi.org/10.11609/JoTT.ZPJ.17.4.757-63

Padhye, A.D., N. Modak & N. Dahanukar (2014). Indirana chiravasi, a new species of leaping frog (Anura: Ranixalidae) from Western Ghats of India. Journal of Threatened Taxa 6(10): 6293–6312; http://dx.doi.org/10.11609/JoTT.o4068.6293-312

Parks, D.H., T. Mankowski, S. Zangooei, M.S. Porter, D.G. Armanini, D.J. Baird, M.G.I. Langille & R.G. Beiko (2013). GenGIS 2: Geospatial analysis of traditional and genetic biodiversity, with new gradient algorithms and an extensible plugin framework. PLoS One 8: 7, e69885; http://dx.doi.org/10.1371/journal.pone.0069885

Puillandre, N., A. Lambert, S. Brouillet & G. Achaz (2012). ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology 21: 1864–1877;

http://dx.doi.org/10.1111/j.1365-294X.2011.05239.x

Robin, V.V., A. Sinha & U. Ramakrishnan (2010). Ancient geographical gaps and paleo-climate shape the phylogeography of an endemic bird in the sky islands of southern India. PLoS ONE 5(10): e13321; http://dx.doi.org/10.1371/journal.pone.0013321

Schwarz, G. (1978). Estimating the dimension of a model. Annals of Statistics 6: 461–464.

Rambaut, A. (2009). FigTree, ver. 1.4.2. http://tree.bio.ed.ac.uk/software/figtree/ Accessed on 28 February 2015.

Rao, C.R.N. (1920). Some South Indian batrachians. Journal of the Bombay Natural History Society 27: 119–127.

Rao, C.R.N. (1937). On some new forms of Batrachia from S. India. Proceedings of the Indian Academy of Science (B) 6: 387–427; http://dx.doi.org/10.1007/BF03051434

Roelants, K., J. Jiang & F. Bossuyt (2004). Endemic ranid (Amphibia: Anura) genera in southern mountain ranges of the Indian subcontinent represent ancient frog lineages: evidence from molecular data. Molecular Phylogenetics and Evolution 31: 730–740; http://dx.doi.org/10.1016/j.ympev.2003.09.011

Satyamurti, T.S. (1967). The South Indian Amphibia in the Collection of the Madras Government Museum. Bulletin of the Madras Government Museum 7: 1–90.

Savage, J.M. & W.R. Heyer (1967). Variation and distribution in the treeâ€frog genus Phyllomedusa in Costa Rica, central America. Studies on Neotropical Fauna and Environment 5(2): 111–131; http://dx.doi.org/10.1080/01650526709360400

Srinivasulu, C. & I. Das (2008). The herpetofauna of Nallamala Hills, Eastern Ghats, India: An annotated checklist, with remarks on nomenclature, taxonomy, habitat use, adaptive types and biogeography. Asiatic Herpetological Research 11: 110–131.

Srinivasulu, C., M. Siliwal, A. Rajesh, B. Srinivasulu, P. Venkateshwarulu & V. Nagulu (2007). Diversity and distribution of amphibian fauna in Nagarjunsagar-Srisailam Tiger Reserve, Andhra Pradesh. Frog Leg 13: 3–6.

Tamura, K., G. Stecher, D. Peterson, A. Filipski & S. Kumar (2013). MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30: 2725–2729; http://dx.doi.org/10.1093/molbev/mst197

Thurston, E. (1888). Catalogue of the Batrachia, Salientia and Apoda (Frogs, Toads and Cecilians) of Southern India. Printed by the Superintendent, Government Press, Madras, 54pp.

van Bocxlaer, I., K. Roelants, S.D. Biju, J. Nagaraju & F. Bossuyt (2006). Late Cretaceous Vicariance in Gondwanan Amphibians. PLoS ONE 1(1): e74;

http://dx.doi.org/10.1371/journal.pone.0000074

van Bocxlaer, I., Biju, S.D., Willaert, B., Giri, V.B., Shouche, Y.S. & F. Bossuyt (2012). Mountain-associated clade endemism in an ancient frog family (Nyctibatrachidae) on the Indian subcontinent. Molecular Phylogenetics Evolution 62(3): 839-847; http://dx.doi.org/10.1016/j.ympev.2011.11.027

Vasudevan, K., A. Kumar & R. Chellam (2006). Species turnover: the case of stream amphibians of rainforests in the Western Ghats, southern India. Biodiversity and Conservation 15: 3515–3525; http://dx.doi.org/10.1007/s10531-004-3101-x

Vasudevan, K., A. Kumar, B.R. Noon & R. Chellam (2008). Density and Diversity of forest floor anurans in the rain forest of southern Western Ghats. Herpetologica 64: 207–215; http://dx.doi.org/10.1655/07-066.1

Vasudevan, K., A. Kumar & R. Chellam (2001). Structure and composition of rainforest floor amphibian communities in Kalakad–Mundanthurai Tiger Reserve. Current Science 80(3): 406–412.

Vences, M., M. Thomas, A. Van der Meijden, Y. Chiari & D.R. Vieites (2005). Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology 2: 5; http://dx.doi.org/10.1186/1742-9994-2-5

Xia, X. (2013). DAMBE5: A comprehensive software package for data analysis in molecular biology and evolution. Molecular Biology and Evolution 30: 1720–1728; http://dx.doi.org/10.1093/molbev/mst064