Diversity and Morphological Phylogenetics of Marine Gammarid Amphipods in Thai Waters
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i Diversity and Morphological Phylogenetics of Marine Gammarid Amphipods in Thai Waters Koraon Wongkamhaeng A Thesis Submitted in Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Biology Prince of Songkla University 2013 Copyright of Prince of Songkla University i Thesis Title Diversity and Morphological Phylogenetics of Marine gammarid amphipods in Thai Waters Author MissKoraonWongkamhaeng Major Program Biology Major Advisor Examining Committee: ……………………………………… …………………….…Chairperson (Assoc. Prof. Dr.PornsilpPholpunthin) (DrAnuwatNateewathana) …………………….…Committee (DrJaruwatNabhitabhata) The Graduate School, Prince of Songkla University, has approved this thesis as fulfillment of the requirements for the Doctor of Philosophy Degree in Biology ………………………………………… (Assoc.Prof.Dr. TeerapolSrichaba) Dean of GraduateSchool iii This is to certify that the work here submitted is the result of the candidate’s own investigations. Due acknowledgement has been made of any assistance received. …………………………………………Signature (Assoc. Prof. Dr. Pornsilp Pholpunthin) Major Advisor …………………………………………Signature (Miss Koraon Wongkamhaeng) Candidate iv I hereby certify that this work has not been accepted in substance for any other degree, and is not being currently submitted in candidature for any degree. …………………………………………..Signature (Miss Koraon Wongkamhaeng) Candidate v ชื่อวิทยานพนธิ ์ ความหลากชนิดและวิวฒนาการชาตั พิ ันธุ์ทางสัณฐานวิทยาของแกมมาริด แอมฟิพอดในน่านน้ําไทย ผู้เขยนี นางสาวกรอร วงษ์กําแหง สาขาวิชา ชีววิทยา ปีการศึกษา 2555 บทคัดย่อ ศึกษาแกมมารดแอมฟิ ิพอดทะเลในน่านน้ําไทย พบแกมมาริดแอมฟิพอดทั้งหมด 33 ชนิด จาก 17 วงศ์ โดยแอมฟิพอด 8 ชนิดเป็นชนิดใหม่ของโลกได้แก่ Grandidierella halophilus, Grandidierella phetraensis, Cheiriphotis trifurcata, Ceradocus andamanensis, Maeropsis paphavasita, Parelasmopus siamensis, Rotomelita longipropoda และ Tethygeneia khanomensis แอมฟิพอด 8 ชนิดเป็นรายงานใหม่ในน่านน้ําไทยได้แก่ Amphilochus justi, Ampithoe africana, Ampithoe ramondi, Cymadusa pilipes, Paradexamine latifolia, Listriella longipalma, Waldeckia elephas และ Melita latiflagella และอีก 3 ชนดยิ ังไม่สามารถจําแนกชนิดได้ ได้แก่ Grandidierella sp.A, Chelicorophium sp.A และ Tiron sp.A ทําใหในป้ ัจจบุ ันมีรายงานของแอมฟิพอดในน่านน้ําไทย 63 ชนิด ทําการเปรียบเทียบวิวัฒนาการชาติพันธุ์ของการศึกษาก่อนหน้านี้กับการศึกษาครั้งนี้และ สร้างแผนภูมิแสดงวิวัฒนาการชาติพันธ์ของแอมฟิพอดในน่านน้ําไทย โดยใช้ลักษณะทั้งหมด 40 ลักษณะ พบว่ามีความสอดคล้องกับการจัดกลุ่มของแอมฟพอดในปิ ัจจุบันยกเว้นแอมฟิพอดวงศ์ Melitidae, Aoridae, Photidae และ Corophiidae แอมฟิพอดที่พบมีลักษณะทางนิเวศแตกต่างกัน แบ่งได้เป็นกลุ่มที่หากินตามหน้าดินและเหนือพื้นดิน (neritic) กลุ่มที่อยในดู่ นิ (nestler) กลุ่มที่สร้าง ท่อ (dominolous) และกลุ่มที่อาศัยอยู่กับสัตว์ชนิดอื่น (inquilinous) ทําการหาสายสัมพันธ์เชิง วิวัฒนาการของแอมฟิพอดในแต่ละสกุล vi Thesis Title Diversity and Morphological Phylogenetics of Marine gammarid amphipods in Thai Waters Author Miss Koraon Wongkamhaeng Major Program Biology Academic Year 2012 ABSTRACT The present study of marine gammaridean amphipod revealed 33 species in 17 families. Of these, 8 species were new to science namely Grandidierella halophilus, Grandidierella phetraensis, Cheiriphotis trifurcata, Ceradocus andamanensis, Maeropsis paphavasita, Parelasmopus siamensis, Rotomelita longipropoda and Tethygeneia khanomensis and 8 species were new record to Thai Waters namely Amphilochus justi, Ampithoe africana, Ampithoe ramondi, Cymadusa pilipes, Paradexamine latifolia, Listriella longipalma, Waldeckia elephas and Melita latiflagella and 3 species are unidentified namely Grandidierella sp.A, Chelicorophium sp.A and Tiron sp.A. Sixty three species are now known from Thai Waters. By considering to both previous phylogenetics and the results in this study, the phylogeny of the marine gammaridean Amphipoda in Thai Waters was created. Forty morphological characters were used in analyses. The phylogenetic tree shows the relationship between amphipod group which mostly follow the recent amphipod classification except the members Family Melitidae, Aoridae, Photidae and Corophiidae. The species occurred as a variety of ecological forms including neritic, nestler, domicolous and inquinous. Cladistic methods were used to examine phylogenetic relationships among the genera of marine gammaridean Amphipoda. vii ACKNOWLEDGEMENT The author is indebted to a number of individuals and organization for their support and encouragement towards the successful completion of this research work First of all, the author would like to express her sincere gratitude to the Examination committee Members, to Dr Anuwat Nateewathana, the Chairperson and Dr Jaruwat Nabhitabhata for their critical comments. The author is indebted and would like to express her deepest gratitude to her Advisors Assoc. Prof. Dr. Pornsilp Pholpunthin and Dr. Charle Oliver Coleman from Museum für Naturkunde, Germany and Dr Azman Abdul Rahim from Universiti Kebangsaan, Malaysia and for their valuable advices in conducting this research and their great contribution to the final draft of this thesis book. The supports provided by the following organizations during the field samplings and sorting of specimen are appreciated namely, Sichang Marine Science Research and Training Center and Phuket Marine Biological Research Center for laboratory facilities, amphipod specimen and reference books. The author would like to express her sincere thank to Mr. Pongrat Damrongrojwattana Mr Eknarin Rodcharoen Mr. Phuripong Meksuwan and all members of Plankton Research Unit and Cephalopods Researchl Unit, Department of Biology, Faculty of Science, Prince of Songkla University for their assistances during the field samplings and the laboratory works, their encouragements and suggestions. This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training grant BRT T253013. Thanks are due to the scholarship for an oversea research study from Graduate School, Prince of Songkla University for co-laboratory supported. Part of this work was supported by the Higher Education Research Promotion and National Research University Project of Thailand. Lastly, the author is indebted to her beloved parents and family for their understanding, supports and encouragements throughout this academic task. viii CONTENTS Page Abstract v Acknowledgements vii Contents viii List of Tables x List of Figures xi CHAPTER 1 Introduction 1 Literature Reviews 3 1.1 General Charateristics of gammarid amphipods 3 1.2 Systematics Resume of Order Amphipoda 3 1.3 Morphology of gammarid amphipod 5 1.4 Status of gammarid amphipods 14 1.5 Evolutionary Pattern of gammarid amphipod 14 CHAPTER 2 Materials and Methods 25 2.1 Sampling Stations 25 2.2 Sampling Methods 29 2.3 Specimen Collection 30 2.4 Phylogenetic Study 30 2.5 Phylogenetic analysis 30 CHAPTER 3 Result 39 3.1 Species list 39 3.2 Systematic account 44 3.3 Gammarid amphipod habitats 101 3.4 Gammarid amphipod ecology 102 3.5 Phylogenetic study 104 ix CONTENS (Continuted) Page CHAPTER 4 Discussion 108 4.1 Taxonomic study of gammarid amphipod in Thai Waters 108 4.2 Phylogenetic of gammarid amphipod in Thai-Water 111 4.3 Evolutionary Biology of the Amphipods in Thai Waters 114 CHAPTER 5 Conclusion 118 REFERENCE 119 BIOGRAPHY 130 x LIST OF TABLES Table Page 1 Comparatives of gammaridaen characters 21 2 Study sites in Andaman Sea 26 3 Study sites in easthern Gulf of Thailand 26 4 Study sites in western Gulf of Thailand 27 5 Study sites in lower western Gulf of Thailand 27 6 List of species found in this study 42 7 Gammarid amphipods species in Thai Water in different habitats 101 8 Gammarid amphipod ecological group 102 9 Gammarid Amphipod species in Thai Waters 108 xi LIST OF FIGURES Figure Page 1 General morphology of amphipods 3 2 Order Amphipoda 5 3 Amphipod head 6 4 Head structure of amphipods 7 5 Epistome 8 6 Lower lip 8 7 Mandible 9 8 The mandible modification pathways 10 9 First maxillae 10 10 Second maxillae 11 11 Thoracic appendages of amphipod 12 12 Gnathopod types 12 13 Pleon appendages 13 14 Pattern of evolution in the Gammaridea 16 15 World gammaroidean amphipods systematics 17 16 Systematic of 19 gammaridean superfamilies 18 17 The evolutionary tree of gammarid amphipod 19 18 The phylogenetic relationships among gammaridean families 19 19 Map of study sites 28 20 Head characters: free and coalesced 31 21 Head lateral cephalic lobe 32 22 Sternal spines 33 23 Mandibular molar 33 24 Maxilla 1 inner lobe 34 25 Maxilla 2 inner lobe 34 xii LIST OF FIGURES (continued) Figure Page 26 Labium outer lobes 35 27 Maxilliped inner plate 35 28 Amphilochus justi Azman, 2009 49 29 Ampithoe ramondi Audouin, 1826 52 30 Ampithoe africana K.H. Barnard, 1926 55 31 Cymadusa pilipes (Ledoyer, 1984) 57 32 Grandidierella sp.A, ♂ 65 33 Grandidierella sp.A, ♂ 66 34 Grandidierella sp.A, ♂ 67 35 Chelicorophium sp.♂ 70 36 Chelicorophium sp.♂ 71 37 Chelicorophium sp.♂ 72 38 Paradexamine latifolia Ren, 2006 ♀ 76 39 Listriella longipalma Othman&Morino, 2006 ♂ 80 40 Waldeckia elephas Hirayama & Kikuchi, 1980 ♂ 86 41 Melita latiflagella Ren&Andres, 2012 ♂ 90 42 Tiron sp. A, ♂ 97 43 Tiron sp. A, ♂ 98 44 Tiron sp. A, ♂ 99 45 Cladogram of the relationships of taxa within the amphipod in Thai-Waters 106 46 Characters used for defining clade 107 47 Cladogram of the relationships of the families within the amphipod 113 48 Cladogram of relationships of parvorders within the Senticaudata 114 49 Cladogram of the relationships of ecological groups within the 117 amphipod in Thai-Waters 1 CHAPTER 1 INTRODUCTION Gammarid