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Zoantharia (Cnidaria: Anthozoa: Hexacorallia) of the South China Sea and Gulf of Thailand: a species list based on past reports and new photographic records PDF

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Reimer et al.: Zoantharia in the South China Sea Taxonomy & Systematics RAFFLES BULLETIN OF ZOOLOGY 63: 334–356 Date of publication: 1 September 2015 http://zoobank.org/urn:lsid:zoobank.org:pub:7514BA25-FFD7-497D-8693-65262EEC127C Zoantharia (Cnidaria: Anthozoa: Hexacorallia) of the South China Sea and Gulf of Thailand: a species list based on past reports and new photographic records James Davis Reimer1, 2, 3, Hin Boo Wee3, Ang Put Jr.4 & Bert W. Hoeksema5 Abstract. This study is the first review of Zoantharia species in the South China Sea and Gulf of Thailand. In addition to past literature records, new field observations are added from previously unexamined countries and regions. In total 16 species are listed, 15 of which belong to suborder Brachycnemina, and only one to suborder Macrocnemina. Two species are undescribed. The lack of Macrocnemina species is not likely indicative of a low diversity of this suborder in the South China Sea and Gulf of Thailand, but instead of an absence of research below shallow subtidal depths. As the majority of the new records from this study were randomly compiled by researchers who are not experts of Zoantharia, specific surveys by experts are needed in these two marine regions. The present list should provide a solid basis for such future research. Key words. Southeast Asia, Indo-Pacific, new records, biodiversity, Zoanthus, Palythoa INTRODUCTION One data-deficient group in the South China Sea and Gulf of Thailand is the order Zoantharia (Cnidaria: Hexacorallia). The South China Sea is one of the most important marine Species in this order are widespread, and zoantharians are regions in terms of commercial shipping, and is currently found in most marine ecosystems from shallow tropical coral subject to contentious and overlapping territorial claims by reefs to the deep sea. Zoantharians are generally colonial, no less than eight countries or regions (People’s Republic of although solitary species also exist. Our understanding of China including Macau and Hong Kong, Republic of China species diversity and taxonomy of this group has traditionally (=Taiwan), the Philippines, Malaysia, Brunei, Indonesia, been limited by a combination of many factors, including Singapore, and Vietnam) due to the perceived value of its a lack of diagnostic characters (Ryland & Lancaster, resources, both mineral and biological. Directly to the west/ 2003; Reimer et al., 2004), high levels of intraspecific southwest of this is the shallow Gulf of Thailand, bordered morphological variation (Burnett et al., 1997; Reimer et by Malaysia, Thailand, and Cambodia. Despite their strategic al., 2004; Ong et al., 2013), and difficulty in performing location, biological knowledge of many marine taxa in these histological examinations due to sand incrustation in the two marine regions is less complete as compared to that body wall and coenenchyme (Reimer et al., 2010). of reef-dwelling scleractinians, which have received much attention recently (Hoeksema & Lane, 2014; Waheed & Recently, molecular techniques such as allozymes (Burnett et Hoeksema, 2014; Huang et al. 2015; Waheed et al., 2015). al., 1997) and DNA markers (Reimer et al., 2004; Sinniger For many taxa, even such basic data as species lists and et al., 2005) have helped to investigate the diversity of distributional information are absent. zoantharians, and have helped reorganise their taxonomy when needed (Sinniger et al., 2010; Fujii & Reimer, 2011, 2013). Many of these studies have focused on shallow- water zooxanthellate species of the Pacific, and combined with recent reports from various regions in the Pacific, the distribution of some species can now be speculated upon. 1Molecular Invertebrate Systematics and Ecology Laboratory, Faculty of Science, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan; Email: For example, many marine regions surrounding the South [email protected] (*corresponding author) China Sea and Gulf of Thailand, such as the Great Barrier 2Tropical Biosphere Research Center, University of the Ryukyus, 1 Senbaru, Nishihara, Reef (Australia) (Burnett et al., 1997), Singapore (Reimer Okinawa 903-0213, Japan & Todd, 2009), Palau (Reimer et al., 2014a), the central 3Institute of Oceanography and Environment (INOS), Universiti Malaysia Terengganu Coral Triangle (Reimer et al., 2014b), Taiwan (Reimer et (UMT), 21030 Kuala Terengganu, Terengganu, Malaysia al., 2011c, 2013), and Japan (Reimer et al., 2006c, etc.) have 4School of Life Sciences, Department of Biology, The Chinese University of Hong Kong, Shatin, N.T. Hong Kong SAR, China been generally well investigated for shallow-water species, 5Department of Marine Zoology, Naturalis Biodiversity Center, P.O. Box 9517, 2300 with species lists existing for each region. The aim of this RA Leiden, The Netherlands paper is to compile a list of the Zoantharia of the South China Sea. The production of a preliminary list will help © National University of Singapore ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print) link data and information between the various surrounding 334 RAFFLES BULLETIN OF ZOOLOGY 2015 regions, and also contribute to understanding of the fauna MATERIAL AND METHODS of the South China Sea. Previously published records in historical literature. We Aside from Pax & Müller (1957) in Vietnam and Reimer & collected data from both scientific reports and guidebooks. Todd (2009) in Singapore, no scientific reports have focused From reports, all data from Pax & Müller (1957) and Reimer on this group in the South China Sea and Gulf of Thailand. & Todd (2009) were included. Additionally, data for all Thus, in this study, we have compiled all previous reports of species from Kenting in southern Taiwan were included zoantharians in the South China Sea and Gulf of Thailand, from Reimer et al. (2011b, c, 2013). Data from Brunei combined them with our own field images, and provide the Darussalam from Reimer et al. (2012) were included, as first assessment of Zoantharia diversity in these regions. It is was a single record of Palythoa singaporensis from Pax & hoped that this study will provide a basis upon which future Müller (1956) from Singapore. Finally, we supplemented zoantharian research in this region can be built. these records with our own previous field images from work in the Gulf of Thailand, peninsular Malaysia, Layang-Layang Atoll, Borneo, and Hong Kong. Latitude and longitude for records and specimens were determined as accurately as Fig. 1. Map of the South China Sea and Gulf of Thailand showing locations of past literature records of Zoantharia (black dots) and newly reported photographic records in this study (pink dots). Note that within each location (Table 1) there may be more than one locality (details in Table 1 and text). 335 Reimer et al.: Zoantharia in the South China Sea possible, and data from original reports included (depth, Isaurus asymmetricus Haddon & Shackleton, 1891: described species identification). from the Torres Strait, Australia, included in this study as I. tuberculatus Gray, 1828 based on taxonomic revision as described in Muirhead & Ryland (1985); and Images from guidebooks were utilised, but added only to Palythoa singaporensis Pax & Müller, 1956: morphologically very the “remarks” section of each species, as such publications similar to Palythoa cf. toxica Walsh & Bowers, 1971, and has are not peer-reviewed literature, and furthermore usually been included as P. cf. toxica for this study. only include very approximate locations. As well, the three Zoantharia species listed below and Species identification and treatment. Species determinations mentioned in historical literature have been treated as follows: were made as in Reimer et al. (2014b) by consulting previous literature (listed with each species). However, some species Gemmaria philippinensis (Gray, 1867): described from the were only identified to “confers with” (cf.) or “affinity” (aff.) Philippines and based on information in von Heider (1899), levels. Aside from the data in Pax & Müller (1957), Reimer this species is unitary with “long” polyps, which fits well with & Todd (2009), and Reimer et al. (2012), very few records Sphenopus. However, researchers (e.g., von Heider, 1899) of zoantharians have been formally reported from the South have long speculated on its true identity. Since we have no China Sea and Gulf of Thailand regions. Therefore, we new information at our disposal, we have omitted this report from the present study; followed recent research (Burnett et al., 1994, 1995, 1997; Zoanthus robustus Carlgren, 1950: described from South Australia Reimer et al., 2006a, 2007, 2011a, 2012; Sinniger, 2006; and included in Pax & Müller (1957), included in this study Sinniger et al., 2010; Reimer & Todd, 2009; Reimer, 2010; as Z. robustus, as although this species has not been examined Fujii & Reimer, 2011) from neighboring regions and used phylogenetically, it is morphologically different from all other species names for which numerous references and accurate known Zoanthus species (for example, see figs. 1, 2 in Grant descriptions were available, unless specimens and/or images et al. (2010)); clearly did not match with previously published information. Palythoa stephensoni Carlgren, 1937: described from the Great Barrier Reef, and discussed in Pax & Müller (1957). In this Our treatment of the species listed in Pax & Müller (1957) study included as P. tuberculosa, based both on discussion in Burnett et al. (1994) and similar results for other species in also needs explanation. In this focused work, Pax & Müller Hibino et al. (2014). listed six Zoanthus spp., four Palythoa spp., and one Isaurus species, and six of these species were first described in this Unpublished photographic records. In this study, to publication. However, recent research (Burnett et al., 1997; supplement the scant data available from previous literature, Reimer et al., 2004) has shown that there are very high we included our own photographic records of Zoantharia from levels of synonymy in these two genera owing to repeated field work in the region (n = 94 new records). We include redescriptions of the same widely distributed species in a summary of the records from historical literature and the different localities. Thus, although we have included the images utilised in this study (Tables 1, 2), and photographic records from Vietnam listed in Pax & Müller (1957) in images of species with new records in the South China Sea this study, we have treated the species listed in their paper and Gulf of Thailand (Figs. 2, 4). For photographic images, as follows: species identifications were conducted as mentioned above. All details for each specimen or photographic record, Palythoa anthoplax Pax & Müller, 1957: included as Palythoa including latitude, longitude, and depth, are given in Table 1. cf. tuberculosa (Esper, 1805) based on phylogenetic results in Reimer et al. (2006c); Palythoa titanophila Pax & Müller, 1957: included as P. cf. Based on previous zoantharian studies (Reimer & Todd, tuberculosa based on external colony appearance and similar 2009), accuracy of identification of Zoanthus and Palythoa data in Hibino et al. (2014); species based on gross morphology from in situ photographs Zoanthus cavernarum Pax & Müller, 1957: included as Z. may approach ~90%: thus, the photographic image data in sansibaricus Carlgren, 1900 based on a description included in this study should not be considered as infallible. Only focused Pax & Müller (1957) compared with this species’ description; specimen collection and identification with a combination of Zoanthus cyanoides Pax & Müller, 1957: included as Z. sansibaricus both morphological and molecular examination should more based on a description included in Pax & Müller (1957) conclusively confirm the identities we have listed here from compared with this species’ description, as well as data in Burnett et al. (1997). Z. cyanoides is very similar to Z. fraseri the photographic records. (Pax & Müller, 1957), which has been stated to likely be Z. coppingeri (Burnett et al., 1997), which is in turn probably is In total, there are Zoantharia records from six regions (not Z. sansibaricus (Reimer, unpublished data); including Gray (1867)) in the South China Sea and Gulf of Zoanthus erythrochloros Pax & Müller, 1957: included as Z. Thailand in the literature (Fig. 1, black-marked regions). To sansibaricus based on phylogenetic results in Reimer et al. this, we have added photographic records from seven new (2004); regions (Fig. 1, pink-marked regions) and historical data Zoanthus gnophodes Pax & Müller, 1957: included as Z. sansibaricus from one additional region. Many of the records we report based on phylogenetic results in Reimer et al. (2004); on below are first records of species in the South China Sea Zoanthus vietnamensis Pax & Müller, 1957: included as Z. vietnamensis as this has been shown clearly to be a distinct and/or Gulf of Thailand, and also for various countries. Some species from other Pacific Zoanthus species (e.g., Burnett et regions include multiple localities (see Table 1 for details). al., 1997; Reimer et al., 2006a); 336 RAFFLES BULLETIN OF ZOOLOGY 2015 Notes Originally mentioned in von Heider (1899) as P. variabilis. Original description of P. anthoplax. Original description of Z. gnophodes. Original description of Z. gnophodes. Only record outside Australia. Original description of Z. cavernarum. Original description of P. titanophila. Original description of Z. erythrochloros. Date NA 1910 1910 1905 1910 1910 1908 arch 1953 ugust 1954 ugust 1954 December 1953 May 1954 and. 4 M 26 A 26 A 24 18 ail h T m) ulf of pth ( NA NA NA NA NA NA NA NA NA NA NA NA G e D e h t d n E E E E E E E E E E E hina Sea a Longitude NA 04°22'19" 04°22'19" 04°22'19" 04°22'19" 04°22'19" 07°04'55" 09°13'33" 09°15'43" 09°15'43" 09°14'32" 09°18'15" C 1 1 1 1 1 1 1 1 1 1 1 h ut from the So Latitude NA 09°39'56"N 09°39'56"N 09°39'56"N 09°39'56"N 9°39'56"N 10°18'09"N 12°10'59"N 12°11'52"N 12°11'52"N 12°17'18"N 12°35'58"N a) a (Cnidaria: Hexacoralli Locality Singapore Poulo Dama (Hon Cu Tron and islands), Vietnam Poulo Dama (Hon Cu Tron and islands), Vietnam Poulo Dama (Hon Cu Tron and islands), Vietnam Poulo Dama (Hon Cu Tron and islands), Vietnam Poulo Dama (Hon Cu Tron and islands), Vietnam Cape St. Jacques, Vietnam Southeast point of Hon Mieu Island, Nha Trang, Vietnam Hon Tre Island, Nha Trang, Vietnam Hon Tre Island, Nha Trang, Vietnam Hon Rua in Ca Dau Bay, Nha Trang, Vietnam Rocher Noir, Ca Dau Bay, Nha Trang, Vietnam ari h s of order Zoant Original identification Palythoa singaporensis Palythoa anthoplax Palythoa singaporensis Zoanthus gnophodes Zoanthus gnophodes Zoanthus robustus Palythoa singaporensis Zoanthus cavernarum Palythoa titanophila Isaurus asymmetricus Palythoa stephensoni Zoanthus erythrochloros d otographic recor Identification (this study) Palythoa cf. toxica Palythoa tuberculosa Palythoa cf. toxica Zoanthus sansibaricus Zoanthus sansibaricus Zoanthus robustus Palythoa cf. toxica Zoanthus sansibaricus Palythoa tuberculosa Isaurus tuberculatus Palythoa tuberculosa Zoanthus sansibaricus h w p en ge en en en en en en en en en en en en e ma m m m m m m m m m m m m s and n Specior im Speci Speci Speci Speci Speci Speci Speci Speci Speci Speci Speci Speci d or c cal re 1956 1957 1957 1957 1957 1957 1957 1957 1957 1957 1957 1957 ori er er er er er er er er er er er er Hist Müll Müll Müll Müll Müll Müll Müll Müll Müll Müll Müll Müll 1. y & & & & & & & & & & & & able Stud Pax Pax Pax Pax Pax Pax Pax Pax Pax Pax Pax Pax T 337 Reimer et al.: Zoantharia in the South China Sea of of es cription hodes. cription mensis. Not ginal desZ. gnop ginal desZ. vietna Ori Ori 6 6 6 6 6 6 6 6 6 6 6 6 54 54 –00 –00 –00 –00 –00 –00 –00 –00 –00 –00 –00 –00 er Date May 19 May 19 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 vembermber 2 eptemb2009 18 18 NoDece NoDece NoDece NoDece NoDece NoDece NoDece NoDece NoDece NoDece NoDece NoDece 4 S m) epth ( NA NA 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 1 to 3 9 to 12 NA D E E E E E E E E E E E E E E E ude 15" 15" 25" 25" 25" 25" 44" 44" 44" 44" 44" 32" 32" 19" 23" ngit °18' °18' °44' °44' °44' °44' °51' °51' °51' °51' °51' °51' °51' °45' °42' o 9 9 3 3 3 3 3 3 3 3 3 3 3 0 0 L 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 de 8"N 8"N 7"N 7"N 7"N 7"N 4"N 4"N 4"N 4"N 4"N 8"N 8"N 9"N 4"N u 5 5 3 3 3 3 1 1 1 1 1 1 1 4 4 Latit 2°35' 2°35' 1°09' 1°09' 1°09' 1°09' 1°13' 1°13' 1°13' 1°13' 1°13' 1°13' 1°13' 1°56' 1°59' 1 1 0 0 0 0 0 0 0 0 0 0 0 2 2 Locality Rocher Noir, Ca Dau Bay, Nha Trang, Vietnam Rocher Noir, Ca Dau Bay, Nha Trang, Vietnam Raffles Lighthouse, Singapore Raffles Lighthouse, Singapore Raffles Lighthouse, Singapore Raffles Lighthouse, Singapore usu Island, Singapore usu Island, Singapore usu Island, Singapore usu Island, Singapore usu Island, Singapore Lazarus Island, Singapore Lazarus Island, Singapore Ho-bi-hoo, Kenting, Taiwan Wanli Tung, Kenting, Taiwan K K K K K Original dentification Zoanthus gnophodes Zoanthus vietnamensis Palythoa tuberculosa Zoanthus vietnamensis Zoanthus sansibaricus Palythoa mutuki Palythoa tuberculosa Zoanthus vietnamensis Zoanthus sansibaricus Palythoa mutuki Palythoa sp. singapura Palythoa tuberculosa Zoanthus sansibaricus Acrozoanthus australiae Palythoa sp. tokashiki i Identification (this study) Zoanthus sansibaricus Zoanthus vietnamensis Palythoa tuberculosa Zoanthus vietnamensis Zoanthus sansibaricus Palythoa mutuki Palythoa tuberculosa Zoanthus vietnamensis Zoanthus sansibaricus Palythoa mutuki Palythoa cf. toxica Palythoa tuberculosa Zoanthus sansibaricus Acrozoanthus australiae Palythoa mizigama n e n n n n n n n n n n n n n n n eg e e e e e e e e e e e e e e e ma m m m m m m m m m m m m m m m pecir im peci peci peci peci peci peci peci peci peci peci peci peci peci peci peci So S S S S S S S S S S S S S S S 9 9 9 9 9 9 9 9 9 9 9 continued Müller 1957 Müller 1957 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 & Todd 200 et al. 2011b et al. 2011c able 1... Study Pax & Pax & Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer T 338 RAFFLES BULLETIN OF ZOOLOGY 2015 ate n. ns W W 3. Notes Only approximlocation know N=13 specime Observed by BHoeksema. Observed by BHoeksema; n= 1 er er 11 er er er er er er er 01 er 13 13 b b 0 b b b b b b b 2 b 0 0 Date 4 Septem2009 4 Septem2009 23 April 2 4 Septem2009 4 Septem2009 4 Septem2009 4 Septem2009 4 Septem2009 1 Septem2012 4 Septem2009 December 4 Septem2009 19 June 2 17 June 2 m) pth ( NA NA 13 2 to 5 1 1 1 to 2 2 3 to 5 5 to 8 3 to 5 9 NA NA e D E E E E E E E E E E E E E de 3" 5" 3" 3" 3" 3" 3" 3" 5" 5" 9" 5" 0" ngitu °42'2 NA °03'0 °42'2 °42'2 °42'2 °42'2 °42'2 °42'2 °46'0 °50'5 °45'1 °13'2 °07'2 o 0 5 0 0 0 0 0 0 0 0 0 4 4 L 2 1 2 2 2 2 2 2 2 2 2 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 de 4"N 4"N 4"N 4"N 4"N 4"N 4"N 4"N 6"N 9"N 9"N 6"N 4"N u 4 A 4 4 4 4 4 4 4 1 0 4 0 3 Latit 1°59' N 5°04' 1°59' 1°59' 1°59' 1°59' 1°59' 1°59' 1°57' 1°56' 1°56' 2°46' 2°47' 2 0 2 2 2 2 2 2 2 2 2 0 0 Locality Wanli Tung, Kenting, Taiwan Kenting, Taiwan Southwest of Pelong Rocks, northwestern Borneo, Brunei Darussalam Wanli Tung, Kenting, Taiwan Wanli Tung, Kenting, Taiwan Wanli Tung, Kenting, Taiwan Wanli Tung, Kenting, Taiwan Wanli Tung, Kenting, Taiwan Wanli Tung, Kenting, Taiwan Tiao Shi, Kenting, Taiwan Longkeng, Kenting, Taiwan Ho-bi-hoo, Kenting, Taiwan Batu Berlabuh, southeast Pulau Tioman, Pahang, Malaysia Tumok, west Pulau Tioman, Pahang, Malaysia Original identification Zoanthus kuroshio Isaurus tuberculatus Sphenopus marsupialis Palythoa tuberculosa Zoanthus vietnamensis Zoanthus sansibaricus Palythoa sp. sakurajimensis Palythoa mutuki Palythoa tuberculosa Palythoa tuberculosa Palythoa tuberculosa Palythoa tuberculosa Identification (this study) Zoanthus kuroshio Isaurus tuberculatus Sphenopus marsupialis Palythoa tuberculosa Zoanthus vietnamensis Zoanthus sansibaricus Palythoa sp. sakurajimensis Palythoa mutuki Palythoa tuberculosa Palythoa tuberculosa Palythoa tuberculosa Palythoa tuberculosa Zoanthus sansibaricus Zoanthus sansibaricus n e n n n n n n n n n n n meag me ge me me me me me me me me me me ge ge pecir im peci Ima peci peci peci peci peci peci peci peci peci peci Ima Ima So S S S S S S S S S S S c c 1 1 2 3 3 3 3 3 3 3 3 3 d 1 1 1 1 1 1 1 1 1 1 1 1 e 0 0 0 0 0 0 0 0 0 0 0 0 u 2 2 2 2 2 2 2 2 2 2 2 2 ntin al. al. al. al. al. al. al. al. al. al. al. al. co et et et et et et et et et et et et dy dy able 1... Study Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer Reimer This stu This stu T 339 Reimer et al.: Zoantharia in the South China Sea W W W W W W W W 2. W W 2. Notes Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema, n= Observed by BHoeksema. Observed by BHoeksema, n= 3 3 3 3 3 3 3 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 Date June 2 June 2 June 2 June 2 June 2 June 2 June 2 April 2 April 2 April 2 3 5 7 9 9 7 7 9 9 9 2 2 1 1 1 1 1 1 1 1 m) h ( A A A A A A A A A A pt N N N N N N N N N N e D E E E E E E E E E E ude 36" 23" 18" 25" 25" 20" 20" 05" 05" 06" ngit °07' °11' °06' °13' °13' °07' °07' °03' °03' °03' o 4 4 4 4 4 4 4 5 5 5 L 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 de 4"N 7"N 4"N 6"N 6"N 4"N 4"N 4"N 4"N 1"N u 0 3 1 0 0 3 3 4 4 4 Latit 2°48' 2°52' 2°54' 2°46' 2°46' 2°47' 2°47' 5°04' 5°04' 5°04' 0 0 0 0 0 0 0 0 0 0 Locality Tanjung Benut, west Pulau Tioman, Pahang, Malaysia Teluk Dalam, northeast Pulau Tioman, Pahang, Malaysia South side of Batu Malang, islet northwest of Pulau Tioman, Pahang, Malaysia Batu Berlabuh, southeast Pulau Tioman, Pahang, Malaysia Batu Berlabuh, southeast Pulau Tioman, Pahang, Malaysia Tumok, west Pulau Tioman, Pahang, Malaysia Tumok, west Pulau Tioman, Pahang, Malaysia Southwest of Pelong Rocks, northwestern Borneo, Brunei Darussalam Southwest of Pelong Rocks, northwestern Borneo, Brunei Darussalam South-southwest of Pelong Rocks, northwestern Borneo, Brunei Darussalam n nal atio gific Orinti e d i Identification (this study) Zoanthus sansibaricus Zoanthus sansibaricus Zoanthus sansibaricus Palythoa mutuki Palythoa heliodiscus Palythoa cf. toxica Palythoa tuberculosa Unidentified Palythoa sp. 1 Palythoa heliodiscus Palythoa mutuki n e pecimer imag Image Image Image Image Image Image Image Image Image Image So d e u n nti o y y y y y y y y y y c d d d d d d d d d d 1... y stu stu stu stu stu stu stu stu stu stu able Stud This This This This This This This This This This T 340 RAFFLES BULLETIN OF ZOOLOGY 2015 Notes Observed by BW Hoeksema, n=2. Large colonies cover seafloor. Observed by BW Hoeksema, n=5 Observed by BW Hoeksema. Observed by BW Hoeksema. Observed by BW Hoeksema, n=7. Observed by BW Hoeksema, n=2. Observed by BW Hoeksema. Observed by HB Wee. Observed by HB Wee, n=2. Observed by HB Wee. Observed by HB Wee, n=2. Date 30 June & 3 July 2013 30 June 2013 3 July 2013 1 July 2013 1 July 2013 1 July 2013 3 July 2013 18 April 2013 18 April 2013 3 July 2013 11 April 2013 m) pth ( NA NA NA NA NA NA NA 8 8 3 3 to 5 e D ongitude 3°01'41"E 3°01'41"E 3°01'41"E 3°02'10"E 3°02'10"E 3°02'22"E 2°59'51"E 3° 0'56"E 3° 0'56"E 3°01'44'E 3°03'25"E L 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 Latitude 05°45'42"N 05°45'42"N 05°45'42"N 05°45'37"N 05°45'37"N 05°46'01"N 05°43'56"N 5°47'20"N 5°47'20"N 5°44'23"N 5°37'18"N Locality Redang Kalong House Reef, east Pulau Redang, Terengganu, Malaysia Redang Kalong House Reef, east Pulau Redang, Terengganu, Malaysia Redang Kalong House Reef, east Pulau Redang, Terengganu, Malaysia Che Isa Islet off east Pulau Redang, Terengganu, Malaysia Che Isa Islet off east Pulau Redang, Terengganu, Malaysia Mak Cantek Islet off east Pulau Redang, Terengganu, Malaysia Trumbu Kili Island outh of Pulau Redang, Terengganu, Malaysia Teluk Dalam, Pulau Redang, Terengganu, Malaysia Teluk Dalam, Pulau Redang, Terengganu, Malaysia Ekor Tebu, Pulau Redang, Terengganu, Malaysia Pantai Pasir Cina, Pulau Bidong, Terengganu, Malaysia s n nal atio gific Orinti e d i Identification (this study) Zoanthus sansibaricus Palythoa mutuki Palythoa cf. toxica Palythoa tuberculosa Zoanthus sansibaricus Zoanthus sansibaricus Zoanthus sansibaricus Palythoa cf. toxica Zoanthus sansibaricus Palythoa tuberculosa Zoanthus sansibaricus n e meag ge ge ge ge ge ge ge ge ge ge ge pecir im Ima Ima Ima Ima Ima Ima Ima Ima Ima Ima Ima So d e u n nti o y y y y y y y y y y y c d d d d d d d d d d d 1... y stu stu stu stu stu stu stu stu stu stu stu able Stud This This This This This This This This This This This T 341 Reimer et al.: Zoantharia in the South China Sea e. e, e. e. e. e. We We We We We We W 4. W 7. W W W Notes served by HB served by HB n=3. served by HB served by HB served by HB served by HB Observed by BHoeksema, n= Observed by BHoeksema, n= Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. b b b b b b O O O O O O Date April 2013 July 2013 April 2013 April 2013 April 2013 April 2013 September 2012 September 2012 March 2013 March 2013 March 2013 11 5 7 19 19 20 25 24 24 27 28 m) pth ( 5 3 15 9 4 3 NA NA NA NA NA e D E E E E E E E E E E E ude 25" 32" 46" 33" 33" 01" 28" 24" 09" 02" 39" ngit °03' °02' °03' °00' °00' °00' °50' °01' °51' °51' °50' o 3 3 3 3 3 3 6 7 3 3 3 L 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Latitude 5°37'18"N 5°46'37"N 5°35'50"N 05°48'54"N 05°48'54"N 05°44'32"N 06°59'36"N 07°20'11"N 07°22'24"N 07°22'24"N 07°22'12"N Locality Pantai Pasir Cina, Pulau Bidong, Terengganu, Malaysia Pulau Paku, Pulau Redang, Terengganu, Malaysia Pulau Karah, Pulau Bidong, Terengganu, Malaysia Chagar Hutang, Pulau Redang, Terengganu, Malaysia Chagar Hutang, Pulau Redang, Terengganu, Malaysia Pasir Akar, Pulau Redang, Terengganu, Malaysia Tajau/Bak Bak, Kudat, Sabah, northernmost tip of Borneo, Malaysia Southeast Tanjung Siagut, Kudat, Sabah, northernmost tip of Borneo, Malaysia Coral Café, east Layang-Layang Atoll, Spratlys, Malaysia Wreck Point 1, east Layang-Layang Atoll, Spratlys, Malaysia Wreck Point 2, southeast Layang-Layang Atoll, Spratlys, Malaysia n nal atio gific Orinti e d i Identification (this study) Zoanthus vietnamensis Zoanthus sansibaricus Zoanthus sansibaricus Zoanthus sansibaricus Palythoa cf. toxica Palythoa cf. toxica Sphenopus marsupialis Sphenopus marsupialis Palythoa cf. toxica Palythoa cf. toxica Palythoa cf. toxica n e meag ge ge ge ge ge ge ge ge ge ge ge pecir im Ima Ima Ima Ima Ima Ima Ima Ima Ima Ima Ima So d e u n nti o y y y y y y y y y y y c d d d d d d d d d d d 1... y stu stu stu stu stu stu stu stu stu stu stu able Stud This This This This This This This This This This This T 342 RAFFLES BULLETIN OF ZOOLOGY 2015 W W W W W 4. W W W 4. W W Notes Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema, n= Observed by BHoeksema. Observed by BHoeksema. Observed by BHoeksema, n= Observed by BHoeksema. Observed by BHoeksema. 3 3 3 3 3 3 3 3 3 3 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 2 2 2 2 2 2 2 2 2 2 ate ch ch ch ch ch ch ch ch ch ch D ar ar ar ar ar ar ar ar ar ar M M M M M M M M M M 8 8 0 0 6 6 6 6 7 9 2 2 3 3 2 2 2 2 2 2 m) h ( A A A A A A A A A A pt N N N N N N N N N N e D E E E E E E E E E E ude 59" 59" 46" 46" 19" 19" 19" 55" 22" 32" ngit °49' °49' °49' °49' °47' °47' °47' °48' °47' °48' o 3 3 3 3 3 3 3 3 3 3 L 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 de 7"N 7"N 5"N 5"N 9"N 9"N 9"N 3"N 3"N 4"N u 0 0 5 5 1 1 1 5 3 4 Latit 7°23' 7°23' 7°21' 7°21' 7°22' 7°22' 7°22' 7°22' 7°22' 7°22' 0 0 0 0 0 0 0 0 0 0 Locality Navigator Lane, north Layang-Layang Atoll, Spratlys, Malaysia Navigator Lane, north Layang-Layang Atoll, Spratlys, Malaysia Runway, south Layang-Layang Atoll, Spratly Islands, Malaysia Runway, south Layang-Layang Atoll, Spratly Islands, Malaysia Shark Cave 2, west Layang-Layang Atoll, Spratlys, Malaysia Shark Cave 2, west Layang-Layang Atoll, Spratlys, Malaysia Shark Cave 2, west Layang-Layang Atoll, Spratlys, Malaysia Crack Reef, northwest Layang-Layang Atoll, Spratlys, Malaysia Wrasse Strip 2, northwest Layang-Layang Atoll, Spratlys, Malaysia Mid Reef, northwest Layang-Layang Atoll, Spratlys, Malaysia n nal atio gific Orinti e d i Identification (this study) Palythoa tuberculosa Palythoa heliodiscus Palythoa cf. toxica Acrozoanthus australiae Epizoanthus aff. illoricatus Palythoa cf. toxica Palythoa tuberculosa Palythoa tuberculosa Epizoanthus aff. illoricatus Palythoa heliodiscus n e pecimer imag Image Image Image Image Image Image Image Image Image Image So d e u n nti o y y y y y y y y y y c d d d d d d d d d d 1... y stu stu stu stu stu stu stu stu stu stu able Stud This This This This This This This This This This T 343

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