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Discovery and Evolutionary Affinities of Five New Species of Amphibians from Bangladesh

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Discovery and Evolutionary Affinities of Five New Species of Amphibians from Bangladesh Discovery and evolutionary affinities of five new species of amphibians from Bangladesh Mohammad Sajid Ali Howlader Ecological Genetics Research Unit Department of Biosciences Faculty of Biological and Environmental Sciences University of Helsinki Finland Academic Dissertation To be presented for public examination with the permission of the Faculty of Biological and Environmental Sciences of the University of Helsinki in Latokartanonkaari 7 - B- rakennus, Luentosali 3 (A108) on 13th May 2016 at 12 noon. Helsinki 2016 Supervised by: Prof. Juha Merilä Department of Biosciences University of Helsinki, Finland Dr. Abhilash Nair Department of Biosciences University of Helsinki, Finland Thesis advisory committee: Prof. Jyrki Muona Zoology Unit, Finnish Museum of Natural History University of Helsinki, Finland Dr. Péter Poczai Botany Unit, Finnish Museum of Natural History University of Helsinki, Finland Reviewed by: Dr. Risto Väinölä Zoology Unit, Finnish Museum of Natural History University of Helsinki, Finland Dr. Ilari Eerikki Sääksjärvi Zoological Museum, Department of Biology University of Turku, Finland Examined by: Associate Prof. Tommi Nyman Department of Biology University of Eastern Finland, Finland Custos: Prof. Veijo Kaitala Department of Biosciences University of Helsinki, Finland Author‘s address: Ecological Genetics Research Unit, Department of Biosciences, PO Box 65 (Biocenter 3, Viikinkaari 1), University of Helsinki, Finland E-mail: [email protected], [email protected] Cover design by: Howlader, MSA ISBN 978-951-51-2105-9 (paperback) ISBN 978-951-51-2106-6 (PDF) Available at http://ethesis.helsinki.fi Unigrafia, Helsinki 2016. Disclaimer: This publication (or thesis) is not available for purposes of zoological nomenclature in accordance with the International Code of Zoological Nomenclature (ICZN), article 8.3 (“If a work contains a statement to the effect that all or any of the names or nomenclatural acts in it are disclaimed for nomenclatural purposes, the disclaimed names or acts are not available. Such a work may be a published work….”). Contents ABSTRACT ............................................................................................................................................. 5 INTRODUCTION ................................................................................................................................... 6 Conservation begins by species discovery ................................................................................... 6 Trends in species discovery: Amphibia ......................................................................................... 7 Species and systematics under molecular evolutionary framework ....................................... 9 Historical perspective to amphibian systematics in South Asia ............................................. 10 Amphibian diversity and conservation challenges in Bangladesh ........................................ 11 Aims .................................................................................................................................................. 13 MATERIALS AND METHODS ............................................................................................................ 14 Taxa and specimens ...................................................................................................................... 14 Morphological measurements ..................................................................................................... 15 Sequence analyses and phylogenetic methods.......................................................................... 16 Divergence time estimation ......................................................................................................... 17 Bioacoustic analyses ..................................................................................................................... 17 Ethics Statement ............................................................................................................................ 17 RESULTS AND DISCUSSION .............................................................................................................. 18 Phylogenetic relationships of Fejervarya and Zakerana genera ........................................... 18 New Species ..................................................................................................................................... 20 Zakerana and Fejervarya ......................................................................................................... 20 Euphlyctis .................................................................................................................................... 21 Microhyla .................................................................................................................................... 22 Potential threats ............................................................................................................................ 23 CONCLUSIONS AND FUTURE DIRECTIONS .................................................................................... 25 ACKNOWLEDGEMENTS .................................................................................................................... 26 REFERENCES ....................................................................................................................................... 27 The thesis is based on the following articles, which are referred to in the text by their Roman numerals: I. Howlader MSA (2011) A new species of Fejervarya (Anura: Dicroglossidae) from Bangladesh. Zootaxa 2761: 41–50. II. Howlader MSA, Nair A, Gopalan SV, Merilä J (2016) Molecular phylogeny of the genus Fejervarya (Anura: Dicroglossidae) with a description of new species from Dhaka. Submitted, manuscript. III. Howlader MSA, Nair A, Merilä J (2016) A new species of frog (Anura: Dicroglossidae) discovered from the mega city of Dhaka. PLoS ONE 11: e0149597. IV. Howlader MSA, Nair A, Gopalan SV, Merilä J (2015) A new species of Euphlyctis (Anura: Dicroglossidae) from Barisal, Bangladesh. PLoS ONE 10: e0116666. V. Howlader MSA, Nair A, Gopalan SV, Merilä J (2015) A new species of Microhyla (Anura: Microhylidae) from Nilphamari, Bangladesh. PLoS ONE 10: e0119825. Authors’ contributions I II III IV V Conceived and designed the MSAH MSAH, JM, AN MSAH, JM, AN MSAH, JM, AN MSAH, JM, AN study Performed the analyses MSAH MSAH, AN MSAH, AN MSAH, AN, JM MSAH, AN, JM Contributed study material MSAH MSAH, SVG MSAH MSAH, SVG MSAH, SVG Conduct the field work MSAH MSAH MSAH MSAH MSAH Manuscript preparation MSAH MSAH, JM, AN MSAH, JM, AN MSAH, JM, AN MSAH, JM, AN, Obtained permission for MSAH MSAH MSAH MSAH MSAH fieldwork and collection Received research funding ------- MSAH, JM MSAH, JM MSAH, JM MSAH, JM MSAH: Mohammad Sajid Ali Howlader, AN: Abhilash Nair, SVG: Sujith V Gopalan, JM: Juha Merilä © Magnolia Press (Chapter I) © Authors (Chapters III, IV and V) ABSTRACT Amphibians are the most threatened class of vertebrates. About 48% of the known amphibian species are threatened by extinction, and many species still remain undescribed, especially from tropical and sub-tropical countries such as Bangladesh. In contrast to India and Sri Lanka, amphibian diversity in Bangladesh is poorly known, and little effort has been put towards documenting the species diversity and resolving evolutionary affinities among amphibian taxa in this country. Hence, the actual diversity of amphibians in Bangladesh remains unknown. The aim of this dissertation work was to improve our knowledge of amphibian diversity in Bangladesh by identifying and describing new amphibian species and investigating their evolutionary relationships with closely related taxa. I used morphological and molecular phylogenetic methods to identify and describe one new genus and five new species from different genera. In addition to using traditional morphological comparisons, I also utilized mitochondrial gene fragments to estimate phylogenetic affinities among the studied taxa, with Maximum-likelihood and Bayesian methods. The first two chapters of the thesis focus on the amphibian genera Fejervarya and Zakerana, the latter which was previously embedded within Fejervarya. These chapters also include descriptions of two new species – Fejervarya asmati sp. nov. (now Zakerana asmati), as well as Fejervarya burigangaensis sp. nov. 1, respectively. In the third chapter, a new species (Zakerana dhaka sp. nov.) is described from the urban core of Dhaka, the capital of Bangladesh and one of the most densely populated mega cities in the world. In the fourth chapter, I describe Euphlyctis kalasgramensis sp. nov., which was earlier recognized as E. cyanophlyctis, and show that it is genetically highly divergent from the E. cyanophlyctis described from southern India. In the last and fifth chapter, I describe Microhyla nilphamariensis sp. nov. as a new species. It is a member of a highly genetically heterogeneous group of frogs that have been recognized as M. ornata for the past 173 years. In general, the results of the studies included in this dissertation advance our understanding of amphibian diversity in Bangladesh and adjacent regions, and show that discovery and description of new amphibian species from this region is still fairly easy. Consequently, it seems likely that more thorough sampling and further investigations in this region can uncover additional new amphibian species to science. Such studies, together with the discoveries
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