Biodiversity of marine zooplankton in Southeast Asia (Project-3: Plankton Group) Chapter 5
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Biodiversity of marine zooplankton in Southeast Asia (Project-3: Plankton Group) Chapter 5
Chapter 5 Biodiversity of marine zooplankton in Southeast Asia (Project-3: Plankton Group) Shuhei Nishida and Jun Nishikawa Atmosphere and Ocean Research Institute, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa 277-8564, Japan Introduction The ocean occupies more than 95% of the volume of biosphere on Earth. There is a wealth of diversity in ocean life, but we know only a small portion of it. Among others, zooplankton are distributed in any pelagic habitats in the sea, from coasts to offshore waters, and from the sea surface to the abyssal depths. Many of them are known to play important roles in marine ecosystems, including those in the food chain and matter transfer, but there are also many species whose distribution and ecology are mostly unknown. Southeast Asia is known as the center of marine biodiversity in the world, and this is referable to several unique settings of this region. First, the area has the Tethyan origin, which dates back to ca. 200 million years ago. It also has complex geologic history, including eustatic sea-level changes during the glacial- and inter-glacial periods, and frequent continental fusion and fission events through its geologic history. These resulted in the presence of many island chains and marginal seas, some of which have semi-enclosed deep basins, such as Sulu and Celebes Seas (e.g. Fleminger 1986). A large body of knowledge has accumulated on the high species diversity of marine fauna in this region. To pick up a few: there are more than 550 species of pelagic copepods known in this small region, accounting for one fourth of those known in the world; the MUSORSTOM Expedition, which aimed at re-discovering the primitive decapods “Neoglyhphaea”, resulted in records of >600 species of macrobenthos and demersal fishes as a biproduct, including discovery of more than 80 new species from the very narrow shelf in the northeastern Sulu Sea (e.g. Forest 1989); more recently, there was also the famous discovery of the coelacanth Latimeria menadoensis from near Manado, Sulawesi in 1998 (Pouyaud et al. 1999). All these indicate the ancient nature of the fauna, the extremely high species diversity in this area, and potential diversity of species still waiting for our investigation. However, the area has also been identified as a serious hotspot of biodiversity crisis owing to various human activities, such as: eutrophication, pollution by hazardous chemicals, destruction of habitats such as coral reefs, mangrove forests, and S. Nishida, M. D. Fortes and N. Miyazaki, eds. Coastal Marine Science in Southeast Asia —Synthesis Report of the Core University Program of the Japan Society for the Promotion of Science: Coastal Marine Science (2001–2010), pp. 59–71. © by TERRAPUB 2011. 60 S. N ISHIDA Table 1. AND J. NISHIKAWA List of members of the Plankton Group. Country Name Affiliation Indonesia Indonesia Japan Japan Japan Japan Japan Japan Japan Japan Japan Malaysia Malaysia Philippines Philippines Philippines Thailand Thailand Thailand Vietnam Vietnam Mulyadi Inneke FM Rumengan Susumu Ohtsuka Nozomu Iwasaki Tomohiko Kikuchi Shozo Sawamoto Hideo Sekiguchi Shuhei Nishida Jun Nishikawa Makoto Terazaki Tatsuki Toda BH Ross Othman Fatimah Md Yusoff Wilfredo L Campos Lourdes V Castillo* Ephrime B Metillo Khwanruan Srinui (Pinkaew) Ajcharaporn Piunsumboon Suree Satapoomin Nguyen Cho Nguyen Thi Thu Research Center for Biology, Indonesian Institute of Sciences Sam Raturangi University Hiroshima University Kochi University Yokohama National Uni versity Tokai University Mie University The University of Tokyo The University of Tokyo The University of Tokyo Soka University Universiti Kebangsaan Malaysia Universiti Putra Malaysia University of the Philippines Visayas University of the Philippines Los Bañ os Mindanao State University Burapha University Chulalongkorn University Phuket Marine Biological Center Institute of Oceanography, Nha Trang Institute of Marine Environment and Resources *Deceased seagrass beds, and overfishing. Under this circumstance, we have conducted researches into the biodiversity of zooplankton in Southeast Asia, as one of the field research projects of the Japan Society for the Promotion of Science (JSPS) on Coastal Marine Science during the years 2001–2010. We have also cooperated with the Census of Marine Zooplankton (CMarZ), a field project of the Census of Marine Life (CoML). This is being done with the multilateral cooperation of Japan and five countries in this region: Thailand, Malaysia, Indonesia, Philippines, and Vietnam. Research Planning The research on zooplankton in the present program was initially planned during the “Workshop on the Biodiversity Studies in the Coastal Waters of the East and Southeast Asia” held at Lankawi Island, October 2002, following preparatory communication among researchers in the six col- laborating countries. It was agreed that Plankton-Group comprises two core members from each Southeast Asian country, as a general rule, and Japanese collaborators specializing in major zooplankton taxa, resulting in the collaborators as listed in Table 1 (see Appendix-1 for details). The objectives of our research were: establishing past- and present status of zooplankton communities; elucidating mechanisms of generation/maintenance of biodiversity; elucidating functional role of biodiversity; and predicting the future of the marine ecosystems in this region. We have approached the last objective through: utilization of historical sample collections; training courses on methods of ecology and identification; fulfilling basic knowledge of biodiversity at species/ community levels; utilization of genetic tools for biodiversity analysis; and establishing databases. The sites for general and/or specific field researches are indicated in Fig. 1, including sites for general assessment of Biodiversity of marine zooplankton in Southeast Asia Fig. 1. 61 Research sites of the Plankton Group in the JSPS-CMS Program. zooplankton abundance and species composition, and those for taxonomic and faunal studies with a larger geographic coverage. There are also sites for trophicstructure studies of coral and seagrass communities, including zooplankton, and deep marginal basins such as the Sulu, Celebes, and South China Seas. As an essential strategic aspect, the core members were encouraged to seek and obtain research funds from their domestic sources for practical field research and analysis, since the support from the present project has been limited mainly to travel and meeting expenses. This appears to have been relatively well done, resulting in collaboration with various field projects, either domestic or bilateral-type, as referred to in the following sections. It was also essential to consider differences among collaborating countries in research expertise, needs of countries, and funding circumstances. This led to our basic strategy to put some flexibility in research planning in each country, in terms of, e.g., selection of research sites and seasons, sampling gears, and focal taxonomic groups, which appears to have been a good choice for realistic collaboration. Discovery of New Species Particular efforts have been paid to areas that we call “the hotspots”, where there have been few studies due to logistical and/ or technical difficulties. Hence, a comprehensive research has been conducted in major biodiversity hotspots such as embayed waters, coastal areas and marginal-seas of Southeast Asia, which have very complicated geography and geologic history. This resulted in the discovery of 62 S. N ISHIDA AND J. NISHIKAWA Table 2. List of new species of copepods, amphipods, and isopods described during the JSPSCMS Program by project members and collaborators. Sampling localities are also shown. “n. gen.” and “n. fam.” in parentheses indicate that the species also represent new genus and new family, respectively. Copepoda (holoplankton): 30 spp. (16 papers) Macandrewella stygiana Ohtsuka, Nishida & Nakaguchi, 2002; Okinawa Macandrewella omorii Ohtsuk a, Nishida & Nakaguchi, 2002; Okinawa Macandrewella serratipes Ohtsuka, Nishida & Nakaguchi, 2002; Okinawa Pontella bonei Mulyadi, 2003; Indonesia Pontella kleini Mulyadi, 2003; Indonesia Pontella vervoorti Mulyadi, 2003; Indonesia Neoscolecithrix japonica Ohtsuka, Boxshall & Fosshagen, 2003; Okinawa Scutogerulus boettgerschnacken Ohtsuka & Boxshall, 2004; Okinawa Pseudodiaptomus sulawesiensis Nishida & Rumengan, 2005; Sulawesi Tortanus vietnamicus Nishida & Cho, 2005; Vietnam Metacalanalis hakuhoae Ohtsuka, Nishida & Machida, 2005; Sulu Sea (n. gen.) Protoparamisophria biforaminis Ohtsuka, Nishida & Machida, 2005; Sulu Sea (n. gen.) Paraugaptiloides mirandipes Ohtsuka, Nishida & Machida, 2005; Sulu Sea Sarsarietellus suluensis Ohtsuka, Nishida & Machida, 2005; Sulu Sea Bradyetes pacificus Ohtsuka, Boxshall & Shimomura, 2005; Nansei Is. Lutamator paradiseus Ohtsuka, Boxshall & Shimomura, 2005; Nansei Is. Paracomantenna goi Ohtsuka, Boxshall & Shimomura, 2005; Nansei Is. Centropages maigo Ohtsu ka, Itoh & Mizushima, 2005; Japan Tortanus magnonyx Ohtsuka & Conway, 2005; Seychelles Acartia (Odontacartia) ohtsukai Ueda & Bucklin, 2006; Japan Pseudodiaptomus terazakii Walter, Ohtsuka & Castillo, 2006; Philippine Apocyclops ramkhamhaengi Chullasorn, Kangtia, Pinkaew & Ferrari, 2008; Thailand Kelleria indonesiana Mulyadi, 2009; Indonesia Kelleria javaensis Mulyadi, 2009; Indonesia Halicyclops ariakensis Ueda & Nagai, 2009; Japan Halicyclops continentalis Ueda & Nagai, 2009; Japan Halicyclops unc us Ueda & Nagai, 2009; Japan Three un-described species of Pontellopsis from Indonesia to be published in 2011 Copepoda (meroplankton/parasite): 16 spp. (10 papers) Hemicyclops tanakai Itoh & Nishida, 2002; Japan Hemicyclops javaensis Mulyadi, 2005; Indonesia Hemicyclops minutus Mulyadi, 2005; Indonesia Neomysidion rahotsu Ohtsuka, Boxshall & Harada, 2005; Japan (n. gen.) Umazuracola elongatus Ho, Ohtsuka & Nakadachi, 2006; Japan (n. fam.) Dactylopusioides malleus Shimono, Iwasaki & Kawai, 2007; Japan Maemonstrilla hyottoko Grygier & Ohtsuka, 2008; Okinawa (n. gen.) Maemonstrilla okame Grygier & Ohtsuka, 2008; Okinawa Maemonstrilla polka Grygier & Ohtsuka, 2008; Okinawa Maemonstrilla simplex Grygier & Ohtsuka, 2008; Okinawa Maemonstrilla spinicoxa Grygier & Ohtsuka, 2008; Okinawa Thysanote chalermwati Piasecki, Ohtsuka & Yoshizaki, 2008; Thailand Kensakia aiiroa Harris & Iwasaki, 2009; Malaysia Kensakia shimodensis Harris & Iwasaki, 2009; Japan Thalestris hokkaidoensis Takemori & Iwasaki, 2009; Japan Parenterognathus troglodytes Ohtsuka, Kitazawa & Boxshall, 2010; Japan (n. gen.) Amphipoda: 4 spp. (3 papers) Talorchestia morinoi Othman & Azman, 2007; Malaysia Listriella longipalma Othman & Morino, 2006; Malaysia Ceradocus mizani Lim, Azman & Othman, 2010; Malaysia Victoriopisa tinggiensis Lim, Azman & Othman, 2010; Malaysia Isopoda Metaphrixus setouchiensis Shimomura, Ohtsuka & Sakakihara, 2006; Japan Prodajus curviabdominalis Shimomura, Ohtsuka & Naito, 2005; Japan Biodiversity of marine zooplankton in Southeast Asia 63 Fig. 2. The copepod Tortanus vietnamicus, collected from a coral reef area in the middle of Vietnam during a nighttime sampling. It is thought that this species has escaped from conventional sampling, since they hide behind corals in the daytime (modified from Nishida and Cho 2005). many species new to science. This was more-or-less predicted in the earliest stage of the research, since taxonomic knowledge in this region had been largely based on the results from historical expeditions, and there were many types of habitats that had received little attention in previous researchers. Through cooperation from members and taxonomy experts who collaborated with the project through our taxonomic network, 29 planktonic copepods and 16 mero-planktonic or non-planktonic copepods, 4 amphipods, and 2 isopods have been described as new to science (Table 2, see Appendix-2 for references). In addition, 37 species of mysids have been described as new from Southeast Asia and Japanese waters by experts collaborating with the present project and CMarZ, both from new field sampling and examination of sample collections from previous research cruises. Accordingly, a total of ca. 90 new species have been described in 51 papers during the project. In addition, more than 50 undescribed species are still waiting for our analysis and description. Many of these species have been found from specific habitats that had been poorly investigated, such as estuaries, benthopelagic zones, coral reefs and marginal basins, and many of them are by no means “rare species”, sometimes comprising major components of zooplankton. As an example, Tortanus (Atortus) vietnamicus is a copepod discovered from a coral reef area in the middle of Vietnam (Fig. 2; Nishida and Cho 2005). They measure ca. 2 mm in total lengths, and are relatively large in size for copepods. They were collected by towing a small net from a pier at night. The copepods of this group, from the subgenus Atortus, are known to inhabit clear water and close to bottom substrates or structures, such as corals, in the daytime, and emerge up in the water column at night. So, it is very important to know the ecology and behavior of diverse 64 S. N ISHIDA AND groups of zooplankton for a full coverage of the fauna. A review of the geographic distribution of this group indicated presence of more than 10 species in the IndoPacific Region (Nishida and Cho 2005). They show a highly allopatric pattern of geographic distribution, suggesting speciation through isolation of populations that might have been enhanced by the eustatic sea level changes during the glacial-interglacial periods, which may have resulted in the repeated emergence of both land and deep-ocean barriers. Discovery of many more species is expected through finer geographic coverage of sampling. These copepods would also be a good model of allopatric speciation in coastal areas (see, e.g. Fleminger 1986). Coastal Habitats The coastal waters in Southeast Asia are characterized by the presence of highly diverse habitats such as coral reefs, mangrove forests, seagrass beds, and sandy beaches. To obtain basic information on the current status of zooplankton communities, their abundance and composition were studied in these representative habitats in the member countries (Fig. 1). These include the coastal waters in Vietnam (e.g. Thu 2005, Cho and Trinh 2006, 2008), the Straits of Malacca (e.g. Rezai et al. 2005, 2009, Yoshida et al. 2006), the Gulf of Thailand (e.g. Pinkaew 2003, Srinui 2007), the Philippine waters (e.g. copepods, fish larvae, and chaetognaths: Noblezada et al. 2004, Campos and Santillan 2005, Noblezada and Campos 2008), and the Indonesian waters (e.g. calanoid copepods: Mulyadi 2004, 2006). These data have been assessed for quality by expert members and incorporated into the database of the project (CMarZ-Asia Database, Fig. 4) for integration in the assessment of the current status of zooplankton biodiversity. Although the inventory of zooplankton species are still in preparation, the taxo- J. NISHIKAWA nomic literature on mysids, one of the most species-rich groups of zooplankton in Southeast Asia, have been fully catalogued (Sawamoto and Fukuoka 2005) with a total of 191 species reported from this region. As a matter of particular attention, an integrative, multidisciplinary research on coral reef ecosystems has been conducted in cooperation with the bilateral project between JSPS and VCC (Vice Counselor’s Committee), Malaysia. This was the first comprehensive research on the coral reef systems in Malaysia, encompassing islands and coasts in both eastern and western sides of Peninsular Malaysia, resulting in a number of papers, e.g., the stable-isotope study on food-web structure (e.g. Iwasaki et al. 2004), the community structure and health condition of coral reefs (Toda et al. 2007), abundance, composition and spatio-temporal variability of zooplankton (e.g. Nakajima et al. 2009a), and potential importance of coral mucus and dissolved organic matter (e.g. Nakajima et al. 2009b). Marginal Basins: Sulu, Celebes and South China Seas Comprising another set of habitats in Southeast Asia where research had been wanting, we investigated the Sulu and Celebes Seas, which have highly contrasting geographic features. Both basins are fairly deep with depths of more than 5000 m; the Sulu Sea is semi-enclosed with surrounding sills less than 420 m deep, mostly shallower than 200 m, hence the water exchange with outside is mostly limited to the epipelagic zone, while the Celebes Sea is less enclosed and with more typical open-oceanic conditions. The most striking oceanographic feature in the Sulu Sea is its highly homogenous water structure in meso- and bathypelagic zones, with high temperature of ca. 10°C from ca. 600 m through to the sea bottom of ca. 5000 m. Similar conditions of homogenous, high- Biodiversity of marine zooplankton in Southeast Asia temperature deep water have also been known in the Mediterranean and the Red Sea (e.g. Scotto di Carlo et al. 1984). We investigated these seas during the two cruises of the R/V Hakuho Maru, and compared the species diversity and community structure of meso-zooplankton between them, using a MOCNESS-1 as a main sampling device (Johnson et al. 2006, Nishikawa et al. 2007, Matsuura et al. 2010, Machida and Nishida 2010). Contrary to our expectation, there were no significant differences between the vertical patterns either in the total zooplankton abundance and biomass and in composition at higher taxonomic levels, such as copepods, chaetognaths, and cnidarians. Focusing on copepods, the most dominant group of zooplankton, the calanoids dominated in both seas, which is a general feature in pelagic communities, and again there was no significant difference in the order-level community structure between the seas. However, analyses at the families/species level revealed totally different features (Nishikawa et al. 2007, Nishikawa unpublished data). First, a total of 359 species were identified from the seas. This accounts for >15% of all known pelagic copepods, indicating fairly high species richness in this narrow area. Second, compared to the 314 species from the Celebes Sea, only 217 species were found from the Sulu Sea, accounting for 2/3 of the former. The vertical patterns of species richness clearly indicate that the reduction in the Sulu Sea is due to those in the mesopelagic zone. An analysis of the community structure at the species level also indicated marked differences in the mesopelagic zone between the two seas as compared with the epipelagic zone. It is also noted that the Sulu Sea is another source of species discovery. So far a total of 10 new or undescribed species have been found from the mesopelagic or benthopelagic zones only of the Sulu Sea (Ohtsuka et al. 2005, Nishikawa, unpublished data), suggesting 65 presence of species endemic to the Sulu Sea, inviting further research into the surrounding waters. In summary, the zooplankton diversity in these marginal seas are characterized by: (1) high species richness in the Celebes Sea; (2) reduced species richness in the Sulu Sea; and (3) different community structure in the deep water between the seas. We hypothesize that semi-enclosed marginal basins are other sites of speciation; the stable environmental gradients in the mesopelagic layer in the Celebes Sea may be an important factor for the observed high species richness; the semi-enclosed, warm, homogenous deep water in the Sulu Sea might have eliminated many species, allowed endemic species to evolve, and fewer species occupied broader niches as compared to the Celebes Sea. These views will give insights into our understanding of the mechanisms of speciation and species co-existence in the pelagic habitats. Jellyfish Fisheries and Jellyfish Fauna in Southeast Asia Jellyfish are one of the rare marine zooplankton that have been commercially exploited by humans for food. According to the FAO statistics, global jellyfish catch increased from 1970 to 2000, and it reached 3–5 × 105 Mt after 2000. At least 8 species of jellyfishes belonging to the order Rhizostomeae, class Scyphozoa, are known to be harvested in Southeast Asia. However, detailed information, such as target species, collection and processing methods, and the derived income of fishermen and the processing company, is not well known in most of the fishing grounds. Moreover, biological and ecological aspects of those large jellyfish are rarely studied in spite of their quantitative importance in the local coastal marine ecosystems. Since 2005, we have started the inves- 66 S. N ISHIDA AND J. NISHIKAWA THAILAND A B Fig. 3. A: Locations of the studied area for Jellyfish fisheries in Southeast Asia: Thanh Hoa in Vietnam, Bagan Datoh in Malaysia, and Kebumen in Indonesia, Ang Sila and Sri Racha in Thailand (modified from Nishikawa et al. 2008). B: Fishing boat full of jellyfish; inset, targeted species Rhopilema hispidum (modified from Nishikawa et al. 2007). tigations on the jellyfish fisheries (JF) at several fishery grounds in Southeast Asian countries, such as Vietnam, Indonesia, Malaysia, and Thailand, with the cooperation of local fishermen and fishery office staff (Fig. 3). Information was gathered from interview (with the owner of a jellyfish processing factory (JPF) and fishermen), sampling animals and other ecosystem constituents, and through reports of fishery statistics. Here we outline the results of our JF research, parts of which have been published in Nishikawa et al. (2008, 2009). In northern Vietnam, the harvesting season begins in April and ends in May. Two species, Rhopilema hispidum and R. esculentum are confirmed as commercially exploited, with the former species being caught in much higher abundance than the latter (Fig. 3B). Cyanea, Chrysaora, Sanderia, and Aequorea were also caught, but not used for processing. The number of Rhopilema jellyfish collected by fishermen is estimated at 800,000–1,200,000 individuals per fishery season, suggesting that the fishery can have an impact on jellyfish populations in the area. In Vietnam, 19 species belonging to 12 genera, 7 families and 3 orders of the class Scyphozoa were identified. This species number is more than two times higher than those previously known in Vietnam (9 species). In Central Java, Indonesia, main fishing season of jellyfish is from August to November. Main target species is Crambionella sp. that appears to be new to science. In Malaysia and unlike in most of the fishery grounds, the JF is carried out all year round at the Perak River estuary in Bagan Datoh, Malaysia. The fishermen set their fishing nets at the beginning of both low and high tides, usually once or twice a day. By utilizing the tidal current that transports jellyfish into the nets, they catch the jellyfish without towing them. The main harvested species is Acromitus hardenbergi, which is rarely collected in other areas and its biology is little known. Interestingly, Acromitus occurs abundantly only in this river, not in neighboring rivers, and it appears to prefer brackish water environments. To understand the factors controlling its mass occurrence only in the Perak River, research on its population dynamics, food habits, and trophic structure is now ongoing. Jellyfish fishery is also active in the eastern Gulf of Thailand. High season for the fishery corresponds to the SW 1.1 1.4 unknown Salt, Alum Salt, Alum, Calcium Hypochlorite Salt, Alum, (Soda) 1.2 Salt, Alum, Soda 1.4 Bagan Datoh Kukup Kebumen Malaysia Malaysia Indonesia Thailand Sri Racha, Ang Sila All year August−December June−August Than Hoa Vietnam All year Rhopilema hispidum (main), R. esculentum (rare) Acromitus hardenbergi (main), Rhopilema esculentum (rare), Lobonema smithii (?) (rare) Rhopilema hispidum Crambionella sp. Rhopilema hispidum (main), Lobonema or Lobonemoides (rare) April −May Salt, Alum Selling price to dealer* (USD kg−1) Chemicals used for processing Target species Fishing season Area Country To assess the present status of the impact of anthropogenic pollutants on ecosystems, we investigated the coastal waters and marginal seas of Southeast Asia and the equatorial Pacific in collaboration with a project supported by the JSPS Grant-in-aid for Scientific Research (2004-06). The levels of pollutants were analyzed for various ecosystem components, including pelagic- and benthic organisms, marine snow, and bottom sediments with special reference to the ecological properties of the organisms, such as their vertical distributions and trophic levels. Analysis of organotin compounds, organochlorines and polybrominated diphenyl ethers in the Sulu Sea revealed, for the first time, that these compounds are accumulated in the deep-sea animals in the area, while their levels are lower than in Comparison of jellyfish fisheries and processing methods in three sites in Southeast Anthropogenic Pollutants in Coastaland Deep-Sea Ecosystems Table 3. Asia. monsoon season, which probably transport or accumulate jellyfish to the eastern coastal area of the country. The gears used for catching jellyfish are dip-nets and hooks. The main harvested species is Rhopilema hispidum. Comparison of some aspects of the JF in five sites in Southeast Asia is shown in Table 3. It is interesting that the selling price of processed jellyfish is very similar between the fishery sites, 1.1–1.4 USD, although the price of commodities in general are different between the countries. We also investigated the symbionts on jellyfish (Ohtsuka et al. 2009). Scyphozoan jellyfishes harbored a wide variety of invertebrates and fishes. Juveniles of benthic organisms such as crabs and ophiuroids appear to be hitchhikers for dispersal, while juvenile fish utilize jellyfish as refugia against visual predators. Since edible jellyfish are associated with many kinds of symbionts, the JF possibly hinder recruitment of symbionts. *Most popular price of the most popular products, i.e. swimming bell of most popular species. Currency conversions are 1 USD = 3.47 RM, 17800 VND and 33.3 THB, respectively. 67 Biodiversity of marine zooplankton in Southeast Asia 68 S. N ISHIDA AND the other, more northern areas of the western Pacific (Ramu et al. 2006). An analysis of trace elements (TE) and stable isotope ratios of micronekton and fish in the Sulu Sea and the adjacent Celebes and Philippine Seas indicated high concentrations of Zn, Cu and Ag in non-migrant fish in deep-water, in contrast to high Rb levels in fish which migrate up to the epipelagic zone, probably resulting from differences in background levels of these TEs in each water environment or function of adaptation to deep-water by migrant and non-migrant species. Arsenic level in fish in the Sulu Sea was positively correlated with δ 15 N indicating biomagnification of the element (Asante et al. 2010). In an attempt to obtain basic information necessary to apply zooplankton as indicators of impact of anthropogenic pollutants, a series of experiments was conducted in collaboration with the Pollution Group. It was indicated that the copepod Apocyclops sp. from Sulawesi, Indonesia, is highly sensitive to acute toxicity of tributyltin, suggesting its potential usefulness in environmental assessment (Rumengan et al. 2009). Use of Genetic Markers in Biodiversity Research Genes, as well as morphology, provide important information in understanding the biodiversity of zooplankton. In addition to the analyses of the relationships between species and infra-specific genetic structure, genetic information is used in practical identification of species, such as those in immature and/or damaged specimens. In the present project, research was conducted on the genetic aspects of zooplankton biodiversity. We contributed barcode data of ca. 120 species of copepods and chaetognaths from the Asian Region. There have also been much advancements in the re-evaluation of mor- J. NISHIKAWA phological taxonomy of copepods and chaetognaths by using molecular markers (Machida and Nishida 2010, Miyamoto et al. 2010). Of particular importance in the ecosystem functioning in the Asian Region has been the identification of Calanus species by molecular markers (Nonomura et al. 2008), as summarized below. The copepods of the genus Calanus are among the most important animals both in abundance and biomass in zooplankton communities and play important roles in marine food chains and matter cycling. In the East Asian waters including the coastal waters of China and Japan, three species are distributed: C. sinicus, C. pacificus, and C. jashnovi. They are very similar in shape, and identification of younger stages of these species is almost impossible using morphology alone. For a better understanding of their life history, we applied molecular markers to identify these younger stages. By using three genetic regions as markers, we were able to distinguish the adults and immature copepodids of these species. As a result, it was revealed that the smaller and larger fifth copepodid stages (CVs) of Calanus that are abundant in the mesopelagic layer of Sagami Bay correspond to C. sinicus and C. jashnovi, respectively. The CVs of C. sinicus showed a bimodal distribution, and the deep population was very abundant, which is comparable to its epipelagic population. This and other life-history analyses with wider seasonal and geographic coverage indicated that the CV of C. sinicus is a diapausing stage and very widely distributed in the mesopelagic waters as south as the continental slope in the East China Sea during the seasons when the epipelagic water is too hot for their survival. CMarZ-Asia Database Finally we introduce CMarZ-Asia Database (http://www.cmarz-asia.org/db/ index.html), which is a database estab- Biodiversity of marine zooplankton in Southeast Asia 69 Fig. 4. An example of the species search using the CMarZ-Asia Database (http://www.cmarzasia.org/db/index.html), a comprehensive database on zooplankton information focusing on the Asian Region, containing a search system from species names and sea areas to distribution, sampling data, taxonomic illustrations, images, literature, and gene sequences. It also contains a search system of gene-sequence database (BLAST) to identify species from sequenced data of unidentified specimens. lished during the present project in collaboration with the Census of Marine Zooplankton (Fig. 4). This includes data sets of sample collection and species information, including taxonomic, distributional, and gene sequence (barcode) information, photographic images and taxonomic illustrations for species identification. The ecological data (distribution, abundance, biomass, diversity, etc.) collected/analyzed from different areas and/ or zooplankton-groups by the project members have been uploaded to this database with the metadata for the sample collection (dates, gears, depths, analysts, environmental data, etc.). Conclusion The present project has considerably increased our knowledge on the status of zooplankton biodiversity in East and Southeast Asia. However, much still remain to be addressed. The continuous discovery of many new species indicates high potential biodiversity of the region, necessitating continued research wherein communication and collaboration with taxonomic experts are essential, including education of experts in the collaborating countries themselves. On the other hand, the present research indicates that Southeast Asian water represents excellent habi- 70 S. N ISHIDA AND tats in elucidating the mechanisms generating and maintaining species diversity in the pelagic realm. This will invite further research on pelagic biodiversity through integrated morphological, genetic, and biogeographic studies. As for evaluation of the present status and future prediction of ecosystems, quantitative information on variability of zooplankton communities in space and time is still very limited. Combination of data rescue and mining, use of historical samples, and expanding database should be an inexpensive, promising strategy. J. NISHIKAWA Acknowledgements We thank the Japan Society for the Promotion of Science (JSPS) for supporting the Multilateral Core University Program: Coastal Marine Science. Thanks are also due to all members of the Plankton Group for their collaboration throughout the project. Special thanks are due to staff members and students of collaborating universities and institutes for their great help in the field and laboratory. On behalf of the Plankton-Group members, we dedicate this report to the memory of our good friend and colleague Lourdes V. 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