DNA barcoding insights into echinoderms from Situbondo Coral Reefs, East Java, Indonesia
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Abstract. Aulia ED, Muzaki FK, Lee S-K, Khim JS, Park J. 2025. DNA barcoding insights into echinoderms from Situbondo Coral Reefs, East Java, Indonesia. Biodiversitas 26: 3434-3445. Echinoderms are a key component of marine ecosystems, contributing to species richness and serving ecological functions. Given their importance, observing their diversity and ecology is essential. Despite the favorable coral reef conditions in Situbondo on Java Island, a comprehensive record of echinoderms has never been done. This research addresses this knowledge gap by collecting and identifying the species richness with morphology and molecular identification using the mitochondrial gene Cytochrome Oxidase I (COI) and 16S ribosomal RNA. Timed sampling for 40 minutes was carried out at intertidal reefs from depths between 0-5 m and in shallow reefs from 5-20 m, with a total of five sites. A total of 45 echinoderm species were identified, encompassing 6 Asteroidea, 5 Crinoidea, 8 Echinoidea, 15 Holothuroidea, and 11 Ophiuroidea. Of the 71 echinoderm samples collected, COI sequences were successfully amplified in 66% of the samples, while 16S rRNA sequences had a success rate of 53%. Notably, the COI sequences of five species and the 16S sequences of eight species were reported to GenBank for the first time, contributing to the expansion of publicly available reference data for echinoderm molecular identification. Furthermore, the present study highlights phylogenetic uncertainties based on maximum likelihood analysis of the COI sequences, particularly within two Asteroidea species and one Ophiuroidea species from the Ophiolepis clade. In addition, the findings suggest a potential range extension for Diadema clarki (Ikeda, 1939), indicating previously undocumented distribution patterns. These findings provide essential baseline data and monitoring for future marine biodiversity research, especially considering the area's significant coastal water utilization for tourism and fishing activities.
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References
Alcudia-Catalma MN, Diaz MGQ, Garcia RN, Ocampo PP, Laurena AC, Tecson-Mendoza EM. 2020. DNA barcoding and diversity analysis of 19 economically important Philippine sea cucumbers (Holothuroidea). Philipp J Sci 149 (2): 335-346. DOI: 10.56899/149.02.09.
Ali AI, Suryanti, Sulardiono B. 2017. Kelimpahan dan pola sebaran Echinodermata di Pulau Karimunjawa Jepara. Prosiding Seminar Nasional Hasil-Hasil Penelitian Perikanan dan Kelautan ke-VI. Universitas Diponegoro, Semarang. [Indonesian]
Améziane N, Eléaume M, Roux M. 2024. Classification of Isocrinida (Echinodermata: Crinoidea) with the description of a new extant genus and species from the western Pacific. Zool J Linn Soc 200 (4): 994-1012. DOI: 10.1093/zoolinnean/zlad101.
Amin MHF, Syukriya AJ, Irawan B, Pratiwi AI, Muttaqin Z, Winarni D. 2020. Taxonomic redescription of Colochirus quadrangularis (Echinodermata: Holothuroidea) from Surabaya Coastal Waters (East Java, Indonesia) with notes on new distinctive haplogroup of COI gene. Ecol Environ Conserv 26 (4): 1617-1622.
Andrimida A. 2022. New records of nudibranchs and a sacoglossan (Gastropoda: Heterobranchia) from Sempu Strait, Indonesia. Indo Pac J Ocean Life 6 (1): 1-9. DOI: 10.13057/oceanlife/o060101.
Arbi UY, Wirawati I, Widyastuti E, Hadiyanto, Ibrahim PS, Vimono IB, Pitriana P, Cappenberg HAW, Nugroho DA, Sani SY. 2022. Fauna Bentos Ekosistem Pesisir Indonesia. Epigraf Komunikata Prima, Sukoharjo. [Indonesian]
Aulia ED, Muzaki FK, Saptarini D, Setiawan E, Setiamarga D, Lutvianti ID, Rosyidah SK, Muhammad NA. 2021. Diversity of sea cucumber from intertidal area of Pacitan and Bangkalan, East Java, Indonesia. Biodiversitas 22 (4): 2136-2141. DOI: 10.13057/biodiv/d220463.
Aulia ED, Park J, Lee S-K, Khim JS. 2024. Revealing macrozoobenthos diversity of Java coral reefs, Indonesia: A review on research trends and species assemblages. Front Mar Sci 11: 1387984. DOI: 10.3389/fmars.2024.1387984.
Aziz A. 1981. Fauna Echinodermata dari terumbu karang Pulau Pari, Pulau-Pulau Seribu. Oseanologi di Indonesia 14: 41-50. [Indonesian]
Bahri S, Patech LR, Zulhalifah, Septiani DA, Siswadi. 2021. Distribution and diversity of echinoderms in the coastal waters of south beach of Lombok Island. Jurnal Biologi Tropis 21 (1): 22-31. DOI: 10.29303/jbt.v21i1.2320. [Indonesian]
Bravo G, Kaminsky J, Bagur M, Alonso CP, Rodríguez M, Fraysse C, Lovrich G, Bigatti G. 2023. Roving diver survey as a rapid and cost-effective methodology to register species richness in Sub-Antarctic kelp forests. Diversity 15 (3): 354. DOI: 10.3390/d15030354.
Bribiesca?Contreras G, Verbruggen H, Hugall AF, O’Hara TD. 2019. Global biogeographic structuring of tropical shallow?water brittle stars. J Biogeogr 46 (7): 1287-1299. DOI: 10.1111/jbi.13620.
Casillas S, Barbadilla A. 2017. Molecular population genetics. Genetics 205 (3): 1003-1035. DOI: 10.1534/genetics.116.196493.
Ceríaco LMP, Marques MP. 2025. Fluid-preserved zoological specimens in Portuguese Natural History collections: a historical overview and implications to collection management and research. Natural History Collections and Museomics 2: 1-69. DOI: 10.3897/nhcm.2.142114.
Chacón-Monge JL, Abarca-Odio JI, González-Sánchez K. 2024. Evaluating the reliability of DNA barcoding for Central American Pacific shallow water echinoderms identification: A molecular taxonomy and database accuracy analysis. Rev Biol Trop 72 (S1): e58997. DOI: 10.15517/rev.biol.trop..v72iS1.58997.
Chow S, Konishi K, Mekuchi M et al. 2016. DNA barcoding and morphological analyses revealed validity of Diadema clarki Ikeda, 1939 (Echinodermata, Echinoidea, Diadematidae). Zookeys 585: 1-16. DOI: 10.3897/zookeys.585.8161.
Clark AM, Rowe FEW. 1971. Monograph of Shallow-Water Indo-West Pacific Echinoderms. British Museum (Natural History), London.
Colin PL, Arneson C. 1995. Tropical Pacific invertebrates: A field guide to the marine invertebrates occurring on tropical Pacific coral reefs, seagrass beds and mangroves. Coral Reef Press, Irvine, California.
Coppard SE, Jessop H, Lessios HA. 2021. Phylogeography, colouration, and cryptic speciation across the Indo-Pacific in the sea urchin genus Echinothrix. Sci Rep 11: 16568. DOI: 10.1038/s41598-021-95872-0.
Costello MJ, Tsai P, Wong PS, Cheung AKL, Basher Z, Chaudhary C. 2017. Marine biogeographic realms and species endemicity. Nat Commun 8 (1): 1057. DOI: 10.1038/s41467-017-01121-2.
Dailami M, Jeni, Lapadi I, Sabariah V, Saleh FIE, Paisey AS, Dwiranti F, Saleky D, Toha AHA. 2023. DNA barcoding of sea urchin species in the bird’s head seascape Papua-Indonesia. IOP Conf Ser: Earth Environ Sci 1191: 012002. DOI: 10.1088/1755-1315/1191/1/012002.
Ennas C, Pasquini V, Abyaba H, Addis P, Sarà G, Pusceddu A. 2023. Sea cucumbers bioturbation potential outcomes on marine benthic trophic status under different temperature regimes. Sci Rep 13 (1): 11558. DOI: 10.1038/s41598-023-38543-6.
Foo SH, Taylor KH, Messing CG, Rouse GW, Tay TS, Tan KS, Huang D. 2021. Assessing the taxonomy of Heterometra-like feather stars (Echinodermata: Crinoidea: Himerometroidea) based on morphology and molecular data. Syst Biodivers 19 (7): 632-647. DOI: 10.1080/14772000.2021.1902418.
Fujita T. 2016. Brittle stars of Ophiodermatidae and Ophiolepididae (Echinodermata: Ophiuroidea: Ophiurida: Ophiurina) collected from the Singapore Strait. Raffles Bull Zool 34: 619-626.
Goh S, Quek RZB, Foo SH, Tay TS, Messing CG, Huang D. 2023. Phylogeny and morphology of Himerometroidea (Echinodermata: Crinoidea) feather stars in Singapore. Raffles Bull Zool 71: 92-105. DOI: 10.26107/RBZ-2023-0008.
Goharimanesh M, Stöhr S, Mirshamsi O, Ghassemzadeh F, Adriaens D. 2021. Interactive identification key to all brittle star families (Echinodermata; Ophiuroidea) leads to revised morphological descriptions. Eur J Taxon 766: 1-63. DOI: 10.5852/ejt.2021.766.1483.
González MJV, Borrero-Pérez GH. 2020. First records and new information on the associations of echinoderms with other phyla in the rocky reefs of northern Chocó, Colombian Pacific. Zookeys 921: 1-22. DOI: 10.3897/zookeys.921.32802.
Guzzi A, Alvaro MC, Cecchetto M, Schiaparelli S. 2023. Echinoids and crinoids from Terra Nova Bay (Ross Sea) based on a reverse taxonomy approach. Diversity 15 (7): 875. DOI: 10.3390/d15070875.
Hebert PDN, Cywinska A, Ball SL, deWaard JR. 2003a. Biological identifications through DNA barcodes. Proc Biol Sci 270 (1512): 313-321. DOI: 10.1098/rspb.2002.2218.
Hebert PDN, Ratnasingham S, deWaard JR. 2003b. Barcoding animal life: Cytochrome c oxiase subunit 1 divergences among closely related species. Proc Biol Sci 270: S96-S99. DOI: 10.1098/rsbl.2003.0025.
Hidayah Z, Nuzula NI. 2019. Pemetaan sebaran terumbu karang studi kasus Selat Madura, Jawa Timur. Jurnal Kelautan Tropis 22 (2): 127-134. DOI: 10.14710/jkt.v22i2.5634. [Indonesian]
Hoareau TB, Boissin E, Paulay G, Bruggemann JH. 2013. The Southwestern Indian Ocean as a potential marine evolutionary hotspot: Perspectives from comparative phylogeography of reef brittle-stars. J Biogeogr 40 (11): 2167-2179. DOI: 10.1111/jbi.12155.
Hoareau TB, Boissin E. 2010. Design of phylum?specific hybrid primers for DNA barcoding: Addressing the need for efficient COI amplification in the Echinodermata. Mol Ecol Resour 10 (6): 960-967. DOI: 10.1111/j.1755-0998.2010.02848.x.
Hodin J, Heyland A, Mercier A, Pernet B, Cohen DL, Hamel J-F, Allen JD, McAlister JS, Byrne M, Cisternas P, George SP. 2019. Culturing echinoderm larvae through metamorphosis. Methods Cell Biol 150: 125-169. DOI: 10.1016/bs.mcb.2018.11.004.
Hoeksema BW. 2017. The hidden biodiversity of tropical coral reefs. Biodiversity 18 (1): 8-12. DOI: 10.1080/14888386.2017.1307787.
Hoggett AK. 1991. The genus Macrophiothrix (Ophiuroidea: Ophiotrichidae) in Australian waters. Invertebr Taxon 4 (5): 1077-1146. DOI: 10.1071/IT9901077.
Huda MAI, Sudarmadji, Fajariyah S. 2017. Echinoidea Diversity in Intertidal Zone Jeding Beach Baluran National Park. Berkala Saintek 5(2): 61-65. DOI: 10.19184/bst.v5i2.5531. [Indonesian]
Hutomo M, Moosa MK. 2005. Indonesian marine and coastal biodiversity: Present status. Indian J Mar Sci 34 (1): 88-97.
Kalidi NS, Muskananfola MR, Suryanti S. 2023. Diversity and abundance of sea cucumber (Holothuroidea) resources in the Waters of Duroa Island, Tual City, Maluku, Indonesia. Biodiversitas 24 (11): 6002-6009. DOI: 10.13057/biodiv/d241120.
Khusnah A, Retnaningdyah C, Kurniawan N. 2019. Studies community structure of coral reef at Pasir Putih beach in Situbondo East Java, Indonesia. J Indones Tourism Dev Stud 7 (1): 32-38. DOI: 10.21776/ub.jitode.2019.07.01.05.
Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. J Mol Evol 16: 111-120. DOI: 10.1007/BF01731581.
Kogo I, Fujita T, Kubodera T. 2019. Shallow-water Comatulids (Echinodermata: Crinoidea) from Ambon and Lombok Islands, Indonesia. Spec Divers 24: 229-246. DOI: 10.12782/specdiv.24.229.
Laakmann S, Boos K, Knebelsberger T, Raupach MJ, Neumann H. 2016. Species identification of echinoderms from the North Sea by combining morphology and molecular data. Helgol Mar Res 70: 18. DOI: 10.1186/s10152-016-0468-5.
Lane DJW, Vimono IB. 2020. Two new species of sea star (Asteroidea, Echinodermata) from mesopelagic depths in the Sunda Strait, Indonesia. Raffles Bull Zool 68: 662-669. DOI: 10.26107/RBZ-2020-0081.
Layton KKS, Corstorphine EA, Hebert PDN. 2016. Exploring Canadian echinoderm diversity through DNA barcodes. PLoS One 11 (11): e0166118. DOI: 10.1371/journal.pone.0166118.
Lewis LS, Smith JE, Eynaud Y. 2018. Comparative metabolic ecology of tropical herbivorous echinoids on a coral reef. PLoS One 13 (1): e0190470. DOI: 10.1371/journal.pone.0190470.
Lin Y-J, Rabaoui L, Basali AU, Lopez M, Lindo R, Krishnakumar PK, Qurban MA, Prihartato PK, Cortes DL, Qasem A, Al-Abdulkader K, Roa-Ureta RH. 2021. Long-term ecological changes in fishes and macro-invertebrates in the world’s warmest coral reefs. Sci Total Environ 750: 142254. DOI: 10.1016/j.scitotenv.2020.142254.
Lopes EM, Pérez?Portela R, Paiva PC, Ventura CRR. 2016. The molecular phylogeny of the sea star Echinaster (Asteroidea: Echinasteridae) provides insights for genus taxonomy. Invertebr Biol 135 (3): 235-244. DOI: 10.1111/ivb.12135.
Madduppa H, Cahyani NKD, Anggoro AW, Subhan B, Jefri E, Sani LMI, Arafat D, Akbar N, Bengen DG. 2021. eDNA metabarcoding illuminates species diversity and composition of three phyla (chordata, mollusca and echinodermata) across Indonesian coral reefs. Biodivers Conserv 30: 3087-3114. DOI: 10.1007/s10531-021-02237-0.
Madeira P, Stefanni S, Ávila SP. 2018. Non-destructive tissue sampling and the use of PCR-RFLPs in two edible sea cucumbers from the north-eastern Atlantic, Holothuria mammata Grube, 1840 and H. sanctori Delle Chiaje, 1823 (Echinodermata: Holothuroidea). Eur Zool J 85 (1): 88-93. DOI: 10.1080/24750263.2018.1438529.
Mah CL, Blake DB. 2012. Global diversity and phylogeny of the Asteroidea (Echinodermata). PLoS One 7 (4): e35644. DOI: 10.1371/journal.pone.0035644.
McClanahan TR, Muthiga NA. 2020. Echinometra. In: Lawrence JM (eds). Sea Urchin: Biology and Ecology. Fourth Edition 43. Elsevier Inc., Amsterdam, Netherlands. DOI: 10.1016/B978-0-12-819570-3.00028-7.
McGaughran A. 2015. Integrating a population genomics focus into biogeographic and macroecological research. Front Ecol Evol 3: 132. DOI: 10.3389/fevo.2015.00132.
Moore AM, Tassakka ACM, Ambo-Rappe R, Yasir I, Smith DJ, Jompa J. 2019. Unexpected discovery of Diadema clarki in the Coral Triangle. Mar Biodivers 49: 2381-2399. DOI: 10.1007/s12526-019-00978-4.
Muthahharah A, Adiwibowo S. 2015. The impact of Pasir Putih beach’s tourism, Situbondo, on job and business opportunities. Sodality: J Sosiologi Pedesaan 3: 101-106. DOI: 10.22500/sodality.v3i2.11335. [Indonesian]
Muzaki FK, Saptarini D, Sari NWP, Wicaksono AD, Pragusta R, Lubab M, Humami DW, Tawakkal RI, Buharianto B. 2023. Workshop capacity building: Pelatihan monitoring terumbu karang bagi komunitas pesisir. Sewagati 7 (3): 370-376. DOI: 10.12962/j26139960.v7i3.503. [Indonesian]
Nethupul H, Stöhr S, Zhang H. 2022. Order Euryalida (Echinodermata, Ophiuroidea), new species and new records from the South China Sea and the Northwest Pacific seamounts. ZooKeys 1090: 161-216. DOI: 10.3897/zookeys.1090.76292.
Nugroho ED, Ardiansyah R, Kurniawan N, Widodo, Rahayu DA. 2023. Species identification of echinoderms from Gili Ketapang Island by combining morphology and molecular data. AACL Bioflux 16 (1): 135-150.
O’Loughlin PM, Bribiesca-Contreras G. 2015. New asterinid seastars from northwest Australia, with a revised key to Aquilonastra species (Echinodermata: Asteroidea). Mem Mus Vic 73: 27-40. DOI: 10.24199/j.mmv.2015.73.04.
Patantis G, Dewi AS, Fawzya YN, Nursid M. 2019. Identification of Beche-de-mers from Indonesia by molecular approach. Biodiversitas 20 (2): 537-543. DOI: 10.13057/biodiv/d200233.
Pawson DL. 2007. Phylum Echinodermata*. Zootaxa 1668 (1): 749-764. DOI: 10.11646/zootaxa.1668.1.31.
Pérez-Lloréns JL, Mouritsen OG. 2024. Sea cucumber: A scavenger overexploited, traded and turned into food (even a gastronomic delicacy). Intl J Gastron Food Sci 37: 100996. DOI: 10.1016/j.ijgfs.2024.100996.
Petovi? S, Krpo-?etkovi? J. 2016. How depth and substratum type affect diversity and distribution patterns of echinoderms on the continental. Acta Zool Bulg 68 (1): 89-96.
Pineda-Enríquez T, Bribiesca-Contreras G, Solís-Marín FA, Laguarda-Figueras A, O’Hara T. 2018. New species of the genus Ophiolepis Müller & Troschel, 1840 (Echinodermata: Ophiuroidea: Ophiolepididae). J Mar Biol Assoc UK 98 (8): 2049-2065. DOI: 10.1017/S0025315417001503.
Prata J, Christoffersen ML. 2024. Current knowledge of Holothuriida (Holothuroidea: Echinodermata) from Brazil. Front Mar Sci 11: 1333253. DOI: 10.3389/fmars.2024.1333253.
Prescott J, Riwu J, Prasetyo AP, Stacey N. 2017. The money side of livelihoods: Economics of an unregulated small-scale Indonesian sea cucumber fishery in the Timor Sea. Mar Policy 82: 197-205. DOI: 10.1016/j.marpol.2017.03.033.
Purcell SW, Conand C, Uthicke S, Byrne M. 2016. Ecological roles of exploited sea cucumbers. Oceanogr Mar Biol Ann Rev 54: 367-386. DOI: 10.1201/9781315368597-8.
Rodríguez-Barreras R. 2024. Recent molecular techniques to strengthen ecological studies in echinoderms. Rev Biol Trop 72 (S1): e58880. DOI: 10.15517/rev.biol.trop..v72iS1.58880.
Rojas-Montiel B, Reyes-Bonilla H, Calderon-Aguilera LE, Ramírez-Ortiz G, López-Pérez A, Hernández L, Rivera-Melo FJF. 2020. Echinoderm functional diversity does not correlate with the protection level of marine protected areas in the Mexican Pacific. Biodivers Conserv 29: 1871-1896. DOI: 10.1007/s10531-020-01952-4.
Setiawan R, Ula FA, Sijabat SF. 2019. Species Inventory of Brittle Stars (Ophiuroidea) at Bilik Coastal, Baluran National Park, East Java. Jurnal Kelautan 12 (2): 192-200. DOI: 10.21107/jk.v12i2.5838. [Indonesian]
Setyastuti A, Purwati P. 2015. Species list of Indonesian trepang. SPC Beche-de-mer Inform Bull 35: 19-25.
Setyastuti A, Wirawati I. 2018. The Banda Sea: A hotspot of deep-sea echinoderms diversity in Indonesia. IOP Conf Ser: Earth Environ Sci 184: 012003. DOI: 10.1088/1755-1315/184/1/012003.
Setyastuti A, Zamani NP, Purwati P. 2014. Trepang from Karimunjawa, Situbondo, Spermonde and Ambon. Oseanologi dan Limnologi di Indonesia 40 (2): 133-142. [Indonesian]
Setyastuti ANA, Purbiantoro W, Hadiyanto H. 2018. Spatial distribution of echinoderms in littoral area of Ambon Island, Eastern Indonesia. Biodiversitas 19 (5): 1919-1925. DOI: 10.13057/biodiv/d190544.
Shilling MD, Krueger-Hadfield SA, McClintock JB. 2022. Genetic evidence supports species delimitation of Luidia in the Northern Gulf of Mexico. Biol Bull 243 (1): 28-37. DOI: 10.1086/720972.
Siburian RHS, Tapilatu JR, Tapilatu ME. 2023. Discovery of habitat preferences and community structure of echinoderms in Kri, Raja Ampat, Indonesia. Biodiversitas 24 (7): 3968-3976. DOI: 10.13057/biodiv/d240735.
Sobczyk R, Presler P, Czortek P, Serigstad B, Pabis K. 2023. Diversity, distribution patterns and indicatory potential of echinoderm communities of the tropical East Atlantic (Gulf of Guinea): Influence of multiple natural and anthropogenic factors along a 25-1000 m depth gradient. Ecol Indic 156: 111108. DOI: 10.1016/j.ecolind.2023.111108.
Somma A, Zahida F, Yuda P. 2017. Abundance and distribution of sea urchin (Echinoidea) in Pasir Putih reef, Situbondo, Indonesia. Biota 2 (3): 111-115. DOI: 10.24002/biota.v2i3.1887. [Indonesian]
Sonet G, Smitz N, Vangestel C, Samyn Y. 2022. DNA barcoding echinoderms from the East Coast of South Africa. The challenge to maintain DNA data connected with taxonomy. PLoS One 17 (10): e0270321. DOI: 10.1371/journal.pone.0270321.
Stöhr S, O’Hara TD, Thuy B. 2012. Global diversity of brittle stars (Echinodermata: Ophiuroidea). PLoS One 7 (3): e31940. DOI: 10.1371/journal.pone.0031940.
Stöhr S, Weber AA-T, Boissin E, Chenuil A. 2020. Resolving the Ophioderma longicauda (Echinodermata: Ophiuroidea) cryptic species complex: Five sisters, three of them new. Eur J Taxon 607: 1-2: DOI: 10.5852/ejt.2020.600.
Stöhr S. 2012. Ophiuroid (Echinodermata) systematics—where do we come from, where do we stand and where should we go? Zoosymposia 7: 147-162. DOI: 10.11646/zoosymposia.7.1.14.
Sulardiono B, Hartoko A, Aini AN, Wulandari D, Budiharjo A. 2022. Genetic diversity of commercial sea cucumbers Stichopus (Echinoderm: Stichopodidae) based on DNA Barcoding in Karimunjawa, Indonesia. Biodiversitas 23 (2): 922-927. DOI: 10.13057/biodiv/d230234.
Summers MM, Messing CG, Rouse GW. 2014. Phylogeny of Comatulidae (Echinodermata: Crinoidea: Comatulida): A new classification and an assessment of morphological characters for crinoid taxonomy. Mol Phylogenet Evol 80: 319-339. DOI: 10.1016/j.ympev.2014.06.030.
Supono, Lane DJW, Susetiono. 2014. Echinoderm fauna of the Lembeh Strait, North Sulawesi: Inventory and distibution review. Mar Res Indones 39 (2): 51-61. DOI: 10.14203/mri.v39i2.85.
Tamura K, Stecher G, Kumar S. 2021. MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Mol Biol Evol 38 (7): 3022-3027. DOI: 10.1093/molbev/msab120.
Thuy B, Stöhr S. 2016. A new morphological phylogeny of the Ophiuroidea (Echinodermata) accords with molecular evidence and renders microfossils accessible for cladistics. PLoS One 11 (5): e0156140. DOI: 10.1371/journal.pone.0156140.
VandenSpiegel D, Lane DJW, Stampanato S, Jangoux M. 1998. The asteroid fauna (Echinodermata) of Singapore, with a distribution table and an illustrated identification to the species. Raffles Bull Zool 46 (2): 431-470.
Vimono IB, Borsa P, Hocdé R, Pouyaud L. 2023. Phylogeography of long-spined sea urchin Diadema setosum across the Indo-Malay Archipelago. Zool Stud 62: e39. DOI: 10.6620/ZS.2023.62-39.
Vimono IB, Lane DJW. 2021. Asteroidea (Echinodermata) of the South Java Deep-Sea biodiversity cruise, Indonesia. Raffles Bull Zool 36: 386-425. DOI: 10.26107/RBZ-2021-0044.
Virgili R, Cerrano C, Ponti M, Lasut MT, Reimer JD. 2020. Crinoid diversity and their symbiotic communities at Bangka Island (North Sulawesi, Indonesia). Mar Biodivers 50: 90. DOI: 10.1007/s12526-020-01097-1.
Ward RD, Holmes BH, O’Hara TD. 2008. DNA barcoding discriminates echinoderm species. Mol Ecol Resour 8 (6): 1202-1211. DOI: 10.1111/j.1755-0998.2008.02332.x.
Wirawati I, Setyastuti A, Purwati P. 2019. Timun Laut dari Perairan Dangkal Indonesia. Media Sains Nasional, Jakarta. [Indonesian]
Wolfe K, Byrne M. 2022. Overview of the Great Barrier Reef sea cucumber fishery with focus on vulnerable and endangered species. Biol Conserv 266: 109451. DOI: 10.1016/j.biocon.2022.109451.
WoRMS. 2024. Echinodermata. [accessed 2024 Jun 20]. https://www.marinespecies.org/aphia.php?p=taxdetails&id=1806.
Yasuda N, Hamaguchi M, Sasaki M, Nagai S, Saba M, Nadaoka K. 2006. Complete mitochondrial genome sequences for Crown-of-thorns starfish Acanthaster planci and Acanthaster brevispinus. BMC Genomics 7: 17. DOI: 10.1186/1471-2164-7-17.
Zhang R, Zhou Y, Mao J, Wang C, Zhang D. 2025. Hidden diversity of Freyastera (Asteroidea, Brisingida, Freyellidae) at great depth: Description of new species and remarks on species boundaries. Zoosyst Evol 101 (2): 735-760. DOI: 10.3897/zse.101.144918.
Zheng W, Sun S, Sha Z, Xiao N. 2022. Three new species and two new records of Echinothuriidae (Echinodermata: Echinothurioida) from seamounts in the Northwest Pacific Ocean: Diversity, phylogeny and biogeography of deep-sea echinothuriids. Front Mar Sci 9: 1036914. DOI: 10.3389/fmars.2022.1036914.