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Biodiversity and Biogeography of Polychaetes (Annelida): Globally and in Indonesia

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Biodiversity and Biogeography of Polychaetes (Annelida): Globally and in Indonesia Biodiversity and Biogeography of Polychaetes (Annelida): Globally and in Indonesia Joko Pamungkas A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy in Marine Science, University of Auckland October 2020 Chapte Abstract This thesis presents a review of the biodiversity of polychaete worms (Annelida) and their distribution across the globe. It also provides an evaluation of polychaete biodiversity studies in Indonesia and a description of a new polychaete species collected from Ambon Island, Province of Maluku, Indonesia. I reviewed polychaete data from the World Register of Marine Species (WoRMS), and found that 11,456 accepted polychaete species (1417 genera, 85 families) have been formally described by 835 first authors since the middle of the 18th century. A further 5200 more polychaete species are predicted to be discovered by the year 2100. The total number of polychaete species in the world by the end of the 21st century is thus anticipated to be about 16,700 species. While the number of both species and authors increased, the average number of polychaete species described per author decreased. This suggested increased difficulty in finding new polychaete species today as most conspicuous species may have been discovered. I analysed polychaete datasets from the Global Biodiversity Information Facility (GBIF), the Ocean Biogeographic Information System (OBIS), and my recently published checklist of Indonesian polychaete species, and identified 11 major biogeographic regions of polychaetes. They were: (1) North Atlantic & eastern and western parts of the Mediterranean, (2) Australia, (3) Indonesia, (4) New Zealand, (5) the Atlantic coasts of Spain and France, (6) Antarctica and the southern coast of Argentina, (7) Central Mediterranean Sea, (8) the western coast of the USA, (9) the eastern part of the Pacific Ocean, (10) Caribbean Sea and (11) Atlantic Ocean. Further, the latitudinal species richness gradient pattern of the animals was found to be asymmetrically bimodal, with similar peaks of richness in the northern (60oN) and southern (30oS) hemispheres, and a pronounced dip north of the Equator (15oN). The pattern is unlikely to be due to sampling bias, but rather a natural phenomenon most significantly correlated with sea temperature. I garnered all Indonesian polychaete species names from taxonomic and ecological literature, as well as from GBIF and OBIS, and found that since the middle of the 18th century, 713 polychaete species (55 families) have been identified from the Indonesian waters. Of these, 301 species (40 families) were described from the geographic region. Through time, most polychaete samples were collected during the Siboga Expedition at the turn of the 19th century, most of which were identified by European taxonomists and deposited in museums in the Netherlands, but now centralised at the Naturalis Biodiversity Center, Leiden. Marine benthic studies conducted by local scientists have yielded polychaete specimens, yet most were not identified to species level and not vouchered in a recognised institution. I discovered that the polychaete collections at the three largest polychaete repositories in Indonesia were mostly unidentified, unpublished, and not databased, suggesting that the taxonomic study of the polychaete fauna, at least locally, has been largely overlooked. I collected polychaete specimens from the Wallacea region and its surrounding waters, and discovered a new capitellid species formally named Capitella ambonensis Pamungkas, 2017. The species, which was discovered from a mangrove habitat on Ambon Island, differs from other Capitella species in the form of hooded hooks and the methylene blue staining pattern. Indeed, the polychaete material obtained from this fieldwork may contain more new species and may give a clue as to whether or not the Wallace’s Line is a real boundary for polychaete fauna. i Acknowledgement I would like to thank all the people who helped me during my doctoral study at the University of Auckland in New Zealand. First of all, I would like to express my gratitude and appreciation to my principal supervisor, Prof. Dr. Mark J. Costello, and to my co-supervisor, Dr. Christopher J. Glasby, without whom I would not have been able to undertake and complete my study. Constructive critical comments by Dr. Shane Lavery also improved the quality of this thesis. Further, I would also like to thank my fellow Ph.D students who supported me a lot during my studies. They were: Chhaya Chaudhary, Dinusha Jayathilake, Han-Yang Lin, Irawan Assad, Rakhshan Roohi, Tamlin Jefferson, Thomas Morris, Tri Arfianti and Qianshuo Zhao. This study would not have been possible without the technical assistance of my colleagues at the Museum Zoologicum Bogoriense, i.e., Alfiah, Nur R. Isnaningsih, Riena Prihandini and Ristiyanti M. Marwoto, and without the financial support from the New Zealand ASEAN Scholarship (NZAS) and Postgraduate Research Student Support (PReSS) funding by the University of Auckland. During my fieldwork in Indonesia, a number of people helped me a lot in collecting polychaete samples. I, therefore, would like to thank them: Achmad Husein Nyompa, Asriadin, Boby Otoluwa, E. Moniharapon, Habel Dimara, Hermes Sirlay, La Pay, Mochtar Djabar, Muhammad Bahri, Rikardo Huwae and Tendri Karey. Technical help by Natalia Abrego during my lab work at the University of Auckland is also highly appreciated. Lastly, special thanks to my beloved wife, Sri Murni Asih, who has accompanied me through this challenging process, and to my lovely children, i.e., Bintang Cahaya Langit, Annisa Zhafira and Felicity Queen (the latest one was “gifted” here), who have made my ‘learning journey’ in New Zealand more colourful. ii Table of Contents Abstract ............................................................................................................................. i Acknowledgement ............................................................................................................. ii Chapter 1: Thesis overview ............................................................................................... 1 1.1 General introduction ................................................................................................. 3 1.1.1 Polychaete worms: a brief review .................................................................... 3 1.1.2 The importance of species discovery ............................................................... 6 1.1.3 Global species distribution: why does it matter? .............................................. 7 1.1.4 Indonesian polychaetes: what is known? ......................................................... 8 1.2 Research questions .................................................................................................... 9 1.3 Thesis objectives and structure .................................................................................. 10 Chapter 2: Progress and perspectives in the discovery of polychaete worms (Annelida) of the world ................................................................................... 11 2.1 Introduction .............................................................................................................. 13 2.2 Materials and Methods .............................................................................................. 14 2.3 Results ...................................................................................................................... 16 2.3.1 Species richness .............................................................................................. 16 2.3.2 Species discovery and authors ......................................................................... 21 2.4 Discussion ................................................................................................................ 26 Chapter 3: Biogeography of polychaete worms (Annelida) of the world ........................ 29 3.1 Introduction .............................................................................................................. 31 3.2 Materials and Methods .............................................................................................. 32 3.2.1 Dataset collection and quality control .............................................................. 32 3.2.2 Polychaete biogeographic regions and indicator species .................................. 33 3.2.3 Analyses ......................................................................................................... 33 3.3 Results ...................................................................................................................... 34 3.3.1 Geographical distribution ................................................................................ 34 3.3.2 Latitudinal distribution .................................................................................... 40 3.4 Discussion ................................................................................................................ 44 3.4.1 Geographical distribution ................................................................................ 44 3.4.2 Latitudinal distribution .................................................................................... 46 Chapter 4: Status of polychaete (Annelida) taxonomy in Indonesia, including a checklist of Indonesian species .................................................... 49 4.1 Introduction .............................................................................................................
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