Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5507 No. 4: 16 Sept. 2024
Type: Article
Published: 2024-09-16
DOI: 10.11646/zootaxa.5507.4.1
Page range: 501-533
Abstract views: 85
PDF downloaded: 39

A Tale of Three Oceans—Taxonomy of the Holothuria (Thymiosycia) arenicola Semper, 1868 complex (Echinodermata: Holoturoidea: Holothuriidae)

Scientific Service of Heritage; Royal Belgian Institute of Natural Sciences; Vautierstraat 29; B-1000 Brussels; Belgium
Operational Direction Taxonomy & Phylogeny; Royal Belgian Institute of Natural Sciences; Vautierstraat 29; B-1000 Brussels; Belgium
Echinodermata sea cucumbers Holothuria conusalba Holothuria gracilis Holothuria kerriensis sp. nov. Holothuria milloti Holothuria rathbuni Holothuria strigosa Holothuria unicolor Holothuria zihuatanensis nomenclature

Abstract

The ubiquitous sea cucumber Holothuria (Thymiosycia) arenicola Semper, 1868, externally characterized by a double row of dark blotches of various sizes on its dorsal body wall and a cryptic behaviour, is generally assumed to have a wide tropical distribution, although it has not been reported from the Eastern Atlantic. Careful morphological examination, with emphasis on the ossicle assemblage, of type and non-type H. arenicola specimens sampled in the Indian, Pacific and Atlantic Ocean, its subjective synonyms and species with a similar colouration and habit, revealed that H. arenicola is often confused with other species. This paper formally separates the different species in the H. arenicola complex, one of them being a species new to science: Holothuria (Thymiosycia) kerriensis sp. nov. Additionally, we describe two other species that are often confused with H. arenicola: Holothuria (Lessonothuria) gracilis Semper, 1868 and H. (Thymiosycia) strigosa Selenka, 1867. The H. arenicola complex per se is keyed-out, with the ossicle assemblage of the musculature being recognised as an important, previously largely neglected, guide. This contribution highlights the importance of building and curating well-maintaned natural history collections to understand biodiversity through time and space.

 

References

  1. Abdel Razek, F.A., Abdel Rahman, S.H., Mona, M.H., El-Gamal, M.M. & Moussa, RM. (2007) An observation on the effect of environmental conditions on induced fission of the Mediterranean sand sea cucumber, Holothuria arenicola (Semper, 1868) in Egypt. SPC Beche de Mer Information Bulletin, 26, 33–34.
  2. Alvarado, J.J., Solís-Marin, F.A. & Ahearn, C. (2008) Equinodermos (Echinodermata) del Caribe Centroamericano. Revista de Biología Tropical, 56 (supplement 3), 37 –55.
  3. Aydin, M., Gurlek, M., Samyn, Y., Erguden, D. & Turan, C. (2019) First record of a Lessepsian migrant: the sea cucumber Holothuria (Theelothuria) hamata Pearson, 1913. Zootaxa, 4551 (1), 94–100. https://doi.org/10.11646/zootaxa.4551.1.7
  4. Bebber, D.P., Carine, M.A., Wood, J.R.I., Wortley, A.H., Harris, D.J., Prance, G.T., Davidse, G., Paige, J., Pennington, T.D., Robson, N.K.B. & Scotland, R.W. (2010) Herbaria are a major frontier for species discovery. Proceedings of the National Academy of Science, 107 (51), 22169–22171. https://doi.org/10.1073/pnas.1011841108
  5. Bedford, F.P. (1898) Report on the Holothurians collected by JS. Gardiner at Funafuti and Rotuma. Proceedings Zoological Society London, 1898, 834–848, pls. 52–53. https://doi.org/10.1111/j.1096-3642.1898.tb03186.x
  6. Bedford, F.P. (1899) Holothurians. In: Willey, A. (Ed.), Zoological results based on material from New Guinea, New Britain, Loyality Islands and elsewhere, collected during 1895–7. Part II. University Press, Cambridge, pp. 141–150, pl. xvii.
  7. Bell, F.J. (1887) Report on a collection of Echinodermata from the Andaman Islands. Proceedings of the Zoological Society of London, 1887, 139–145. https://doi.org/10.1111/j.1096-3642.1887.tb02953.x
  8. Bell, F.J. (1888) Report on a collection of Echinoderms made at Tuticorin, Madras. Proceedings of the Zoological Society of London, 1888, 383–389.
  9. Brandt, C. (1835) Echinodermata ordo Holothurina. In: Prodromus descriptionis animalium ab H. Mertensio in orbis terrarum circumnavigatione observatorum. l’Academie imperiale des sciences de St.-Petersbourg, St Petersburg, pp. 42–62.
  10. Briggs, J.C. & Bowen, B.W. (2012) Marine biogeographic provinces: a realignment with particular reference to fish distributions. Journal of Biogeography, 39, 12–30. https://doi.org/10.1111/j.1365-2699.2011.02613.x
  11. Briggs, J.C. & Bowen, B.W. (2013) Marine shelf habitat: biogeography and ecolution. Journal of Biogeography, 40, 1023–1035. https://doi.org/10.1111/jbi.12082
  12. Borrero-Pérez, G.H. & Vanegas-González, M.J. (2019) Holothuria (Mertensiothuria) viridiaurantia sp. nov. (Holothuriida, Holothuriidae), a new sea cucumber from the Eastern Pacific Ocean revealed by morphology and DNA barcoding. Zookeys, 892, 1–9. https://doi.org/10.3897/zookeys.893.36013
  13. Caso, M.E. (1955) Contribución al conocimiento de los holoturoideos de México. Algunas especies de holoturoideos litorales de la costa Atlántica Mexicana. Anales Instituto Biologia. Universidad Nacional Autonoma de México, 26, 501–525.
  14. Caso, M.E. (1965) Estudios sobre equinodermos de-Mexico. Contribucion al conocimiento de los holoturoideos de Zihuatanejo y de la isla de Ixtapa (primera parte). Anales Instituto Biologia. Universidad Nacional Autonoma de México, 36, 253–291.
  15. Caso, M.E. (1976) El estado actual del estudio de los equinodermos de México. Anales del Centro de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de México, 3 (1), 1–56.
  16. Cherbonnier, G. (1955) Résultats scientifiques des campagnes de la “Calypso”. Les Holothuries de la mer Rouge. Annales de l’Institut Océanographique de Monaco, New Series, 30, 129–183, pls. 22–49.
  17. Cherbonnier, G. (1963) Echinodermes des côtes du Cameroun récoltés par A. Crosnier en Décembre 1962-Janvier 1963. Bulletin du Muséum national d’Histoire naturelle, 2ième Série, 35 (2), 179–193.
  18. Cherbonnier, G. (1964) Note préliminaire sur les holothuries de l’Atlantic Sud. Bulletin du Muséum national d’Histoire naturelle, 2 ième série, 36, 4, 532–536.
  19. Cherbonnier, G. (1988) Echinodermes: Holothurides. Faune de Madagascar, 70, 1–292.
  20. Cherbonnier, G. & Féral, J.P. (1984) Les Holothuries de Nouvelle-Calédonie. Deuxième contribution (Première partie: Synallactidae et Holothuriidae). Bulletin du Muséum national d’Histoire naturelle Paris, 4ième Série, 6, section A (3), 659–700. https://doi.org/10.5962/p.285908
  21. Clark, A.M & Rowe, F.W.E. (1971) Monograph of shallow-water Indo-West Pacific echinoderms. Trusties of the British Museum of Natural History, London, 234 pp.
  22. Clark, H.L. (1901a) The Echinoderms of Porto Rico. Bulletin of the United States Fisheries Commission, 2, 231–263.
  23. Clark, H.L. (1901b) Bermudan echinoderms. Proceedings of the Boston Society of Natural History, 29, 339–345.
  24. Clark, H.L. (1919) The distribution of the littoral Echinoderms of the west Indies. Carnegie Institution of Washington Publication 281. Papers from the Department of Marine Biology of the Carnegie Institution of Washington, 13 (No. 3), 49–74
  25. Papers Department Marine Biology Carnegie Institution Washington. Publication 281, 13, 47–74, 3 pls.
  26. Clark, H.L. (1942) The Echinoderm Fauna of Bermuda. Bulletin of the Museum of Comparative Zoology, 89, 367–391.
  27. Conand, C., Michonneau, F., Paulay, G. & Bruggemann, H. (2010) Diversity of the Holothuroid Fauna (Echinodermata) at La Réunion (Western Indian Ocean). Western Indian Ocean Journal of Marine Science, 9, 145–151.
  28. Cutress, B.M. (1996) Changes in dermal ossicles during somatic growth in Caribbean littoral sea cucumbers (Echinodermata: Holothuroidea: Aspidochirotida. Bulletin of Marine Science, 58, 44–116.
  29. Daniel, A. & Halder, B.P. (1974) Holothuroidea of the Indian ocean with remarks on their distribution. Journal of the marine biological Association of India, 16, 412–436.
  30. Deichmann, E. (1926) Report on the Holothurians Collected by the Barbados-Antigua Expedition from the University of Iowa. Iowa Studies in Natural History, 11, 9–31 + pls. 1–3.
  31. Deichmann, E. (1930) The holothurians of the western part of the Atlantic Ocean. Bulletin Museum Comparative Zoology, Harvard, 71, 41–226, pls. 1 –24.
  32. Deichmann, E. (1958) The Holothuroidea collected by the VELERO III and IV during the years 1932 to 1954. Part II. Apsidochirota. Allan Hancock Pacific Expeditions, 11, 239–349, pls. 1–9.
  33. Deichmann, E. (1963) Shallow water holothurians known from the Caribbean waters. Studies Fauna Curaçao, Caribbean Islands, 63, 100–118.
  34. Domantay, J.S. (1934) Four additional species of littoral Holothurioidea of Port Galera Bay and adjacent waters. Natural Applied Science Bulletin, University Philippines, 4, 109–115.
  35. Domantay, J.S. (1936) The ecological distribution of the echinoderm fauna of Puerto Galera Marine Biological Station. Natural Applied Science Bulletin, University Philippines, 5, 385–403, pls. 1–7.
  36. Erwe, W., (1919) Holothurien aus dem Roten Meer. Mitteilungen aus dem Zoologischen Museum in Berlin, 9, 177–190.
  37. Féral, J.P. & Cherbonnier, G. (1986) Les holothuries. In: Guille, A., Laboutes, P. & Ménou, J.-L. (Eds.), Guide des étoiles de mer, oursins et autres échinodermes du lagon de Nouvelle-Calédonie. ORSTOM Editions (IRD), Nouméa, pp. 55–107.
  38. Fontaine, B., Perrard, A. & Bouchet Ph. (2012) 21 Years of shelf life between discovery and description of new species. Current Biology, 22 (22), R943–R944. https://doi.org/10.1016/j.cub.2012.10.029
  39. Gibbs, P.E., Clark, A.M. & Clark, C.M. (1976) Echinoderms from the Northern region of the Great Barrier Reef, Australia. Bulletin of the British Museum (Natural History) Zoology, 30 (4), 103–144, 1pl. https://doi.org/10.5962/bhl.part.2376
  40. Greeff, F.F.R. (1882) Echinoderme, beobachtet auf einer Reise nach der Guinea-Insel São-Tomé. Zoologische Anzeiger, 106, 114–120 + 135–139 + 156–159.
  41. Hasbun, C.R. & Lawrence, A.J. (2002) An annotated description of shallow water holothurians (Echinodermata: Holothuroidea) from Cayos Cochinos, Honduras. Revista Biologia Tropical, 50, 669–678.
  42. Heding, S.G. (1939) The Holothurians Collected During the Cruises of the M/S “Monsunen” in the tropical Pacific in 1934. Videnskabelige meddelelser fra den Dansk Naturhistoriske Forening, 102, 213–222.
  43. Hendler, G., Miller, J.E., Pawson, D.L. & Kier, P.M. (1995) Sea stars, sea urchins , and allies. Echinoderms of Florida and Caribbean. Smithsonian Institution Press, Washington, xi + 390 pp.
  44. Helfer, H. (1912) Uber einig von Dr Hartmeyer im Golf von Suez gesammelte Holothurien. Mitteilungen aus dem Zoologischen Museum in Berlin, 6, 327–324.
  45. Hérouard, E. (1893) Recherches sur les holothuries de la Mer Rouge. Archives de Zoologie expérimentale et générale, Troisième Série, 1, 125–138, pls. 7–8.
  46. Hérouard, E. (1929) Holothuries de la côte atlantique du Maroc et de Mauritanie. Bulletin de la Société des Sciences naturelles du Maroc, 9, 36–70, 1 pl.
  47. Hertlein, L.G. (1963) Contributions to the Biogeography of Cocos Island, including a Bibliography. Proceedings of the California Academy of Sciences, Series 4, 32, 8, 219–289.
  48. Hickman, C.P. (1998) A Field Guide to Sea Stars and other Echinoderms of Galapagos. Sugar Spring Press, Lexington, 83 pp.
  49. Honey-Escandon, M., Solis-Marin, F.A. & Laguarda-Figueras, A. (2008) Equinodermos (Echinodermata) del Pacifico Mexicano. Revista de Biologia Tropica, 56 (Supplement 3), 57–73.
  50. Koehler, R. & Vaney, C. (1905) Mission des Pêcheries de la Côte occidentale d’Afrique. II. Echinodermes. Actes Société Linnéenne, Bordeaux, 60, 58–66, pls. 4–6.
  51. Koehler, R. & Vaney, C. (1908) Holothuries recuiellies par l’Investigator dans l’Ocean Indien. II. Les Holothuries Littorales. Trustees Indian Museum, Calcutta, 54 pps., 3 pls.
  52. Kerr, A.M. & Kim, J. (2001). Phylogeny of Holothuroidea (Echinodermata) inferred from morphology. Zoological Journal of the Linnean Society, 133, 63–81. https://doi.org/10.1111/j.1096-3642.2001.tb00623.x
  53. Kerr, A.M., Janies, D.A., Clouse, R.M., Samyn, Y., Kuszak, J. & Kim, J. (2005) Molecular Phylogeny of Coral-Reef Sea Cuucmbers (Holothuriidae: Aspidochirotida) Based on 16S Mitochondrial Ribosomal DNA Sequence. Marine Biotechnology, 7, 53–60. https://doi.org/10.1007/s10126-004-0019-y
  54. Laguarda-Figueras, A., Solís-Marín, F.A., Durán-González, A., Hernández Pliego, P. & Del Valle-García, R. (2001) Holoturoideos (Echinodermata: Holothuroidea) del Caribe Mexicano: Puerto Morelos. Avicennia, 14, 7–46.
  55. Lampert, K. (1885) Die Seewalzen. Eine systematische Monographie. In: Semper, C., (Ed.), Reisen im Archipel der Philippinen. Teil 2. Wissenschaftliche Resultate. C.W. Kreidel, Wiesbaden, pp. 1–312, 1 pl.
  56. Lane, D.J.W., Marsh, L.M., VandenSpiegel, D. & Rowe, F.W.E. (2000) Echinoderm fauna of the South China Sea: an inventory and analysis of distribution patterns. Raffles Bulletin of Zoology, Supplement No. 8, 459–493.
  57. Levin, V.S. (1979) Aspidochirote holothurians of the upper sublittoral zone of Indo-West Pacific species composition and distribution. Biologya Morya, 5, 17–23.
  58. Levin, V.S. & Dao Tan Ho. (1989) Holothurians of the upper sublittoral zone of the coastal waters of Phukhanh Province (southern Vietnam). In: Zhirmunsky, AV. & Le T.P. (Eds.), Biology of the coastal waters of Vietnam. Benthic Marine invertebrates of southern Vietnam. Far East Science Center, Vladivostok, pp. 1–116.
  59. Liao, Y., (1980) The Aspidochirote holothurians of China.with the erection of a new genus. In: Jangoux, M. (Ed.), Echinoderms- present and past. Proceedings of the European Echinoderm Colloquium on echinoderms, Brussels. September 1979. Balkema Press, Rotterdam, pp. 115–120. https://doi.org/10.1201/9781003078913-21
  60. Liao, Y. (1984) The aspidochirote holothurians of China. Studia Marina Sinica, 23, 221–248.
  61. Liao, Y. (1997) Fauna Sinica. Phylum Echinodermata: Class Holothuroidea. A Major Project of the National Natural Science Foundation of China in the period of the Eighth Five-Year Plan (Supported by the State Science and Technology Commission of China, the National Natural Science Foundation of China and the Chinese Academy of Sciences). Science Press, Beijing, 334 pp.
  62. Liao, Y. (1998) The echinoderm fauna of Hainan Island. In: Morton, B. (Ed.) The Marine Biology of the South China Sea: Proceedings of the Third International Conference on the Marine Biology of the South China Sea. Hong Kong University Press, Hong Kong, pp. 75–82.
  63. Liao, Y. & Clark, A.M. (1995) The echinoderms of southern China. Science Press, Beijing, New York, iii + 614, 23 pls.
  64. Linnaeus, C. (1758). Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Vol. 1. 10th Revised Edition. Laurentius Salvius, Holmiae, 824 pp. https://doi.org/10.5962/bhl.title.542
  65. Lister, A.M. & Climate Change Research Group (2011) Natural History Collections as sources of long-term datasets. Trends in Ecology and Evolution, 26, 153–154. https://doi.org/10.1016/j.tree.2010.12.009
  66. Ludwig, H.L. (1875). Beiträge zur Kenntniss der Holothurien. Arbeiten aus dem zoologisch-zootomischen Institut Würzburg, 2, 77–118, pls. 6–7.
  67. Ludwig, H. (1880). Echinodermata. Zoologische Ergebnisse einer Reise in die Küstengebiet des Rothen Meeres von R. Kossmann, 1 (v), 1–7.
  68. Ludwig, H. (1883) Verzeichniss der Holothurien des Kieler Museums. Bericht der Oberhissischen Gesellschat für Natur-und Heilkunde, 22, 155–176.
  69. Ludwig, H. (1887) Die Von G. Chiercha auf der Fahrt de Kgl.- Ital. Korvette ‘Vettor Pisani’ gesammelten Holothurien. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographieder Tiere, 2, 1–36, pls. 1–2.
  70. Ludwig, H. (1888) Die van Dr. J Brock in Indischen Archipel gesammelten Holothurien. Zoologische Jahrbücher, Abteilung für Systematik, 3, 805–820, pl. 30. https://doi.org/10.5962/bhl.part.1932
  71. Ludwig, H. (1894) Reports on an exploration of the west coasts of Mexico, Central and South America, and off the Galapagos Islands in charge of Alexander Agassiz by the US Fish Commission Steamer Albatros. XII. The Holothurioidea. Memoirs of the Museum of Comparative Zoölogy at Harvard College Cambridge, 17, 1–183.
  72. Ludwig, H. (1889-92) Echinodermen: Die Seewalzen. In: Bronn, H.G. (Ed.), Broon’s Klassen und Ordnungen des Thierreichs. Band 2. Abteilung 3. Buch 1. CF. Winter’sche, Leipzig, pp. i–iv + 1–460, pls. 1–17.
  73. Ludwig, H. (1899) Echinodermen des Sanbargebietes. Wissenschatliche Ergebenisse der Reisen in Madagascar und Ostafrica in den jahren 1889-95. Abhandlungen der Senckenbergischen Naturforschen Gesellschaft, 21, 537–563.
  74. Maluf, L.Y. (1988) Composition and distribution of the central eastern Pacific echinoderm. Technical Report of the National History Museum of Los Angeles County, 2, 1–242.
  75. Martinez de Rodriguez, A. (1973) Contribution al estudio de los holoturoideos de Venezuela. Boletín Instituto Oceanográfico Universidad Oriente, Cumaná, 12, 41–50.
  76. Martinez de Rodriguez, A. & Mago Herminson, A. (1976) Contribucion al conocimientode los holoturoideos (Holothuroidea: Echinodermata) de la region oritental de Venezuela. Boletín Instituto Oceanográfico Universidad Oriente, Cumaná, 14, 187–197.
  77. Martinez, M.I., Solis-Marin, F.A. & Penchaszadeh, P.E. (2019) First report of Paelopatides (Synallactida, Synallactidae) from the SW Atlantic, with description of a new species from the deep-sea off Argentina. Zoologischer Anzeiger, 278, 21–27. https://doi.org/10.1016/j.jcz.2018.10.010
  78. Massin, C. (1993) The Holothurioidea (Echinodermata) collected during the Tyro Mauritania-II. Ecpedition 1988. Zoologische Mededelingen, 67, 397–429
  79. Massin, C. (1999) Reef-dwelling Holothuroidea (Echinodermata) of the Spermonde Archipelago (South-West Sulawesi, Indonesia). Zoologische Verhandelingen, 329, 1–144.
  80. Massin, C., Samyn, Y. & Thandar, A.S. (2004) The genus Labidodemas (Holothuroidae: Aspidochirotida) revisited with description of three new species and with re-positioning of Holothuria (Irenothuria) maccullochi Deichmann, 1958. Journal of Natural History, 38, 1811–1847.
  81. Miller, A.K., Kerr, A.M., Paulay, G., Reich, M., Wilson, N.G., Carvajal, J.I. & Rouse, G.W. (2017) Molecular phylogeny of extant Holothuroidea (Echinodermata). Molecular Phylogenetics and Evolution, 111, 110–13. https://doi.org/10.1016/j.ympev.2017.02.014
  82. Michonneau, F. (2015) Cryptic and not-so-cryptic species in the complex “Holothuria (Thymiosycia) impatiens (Forsskål, 1775) (Echinodermata: Holothuroidea: Holothuriidae). bioRxiv, 1–43. [published online] https://doi.org/10.1101/014225
  83. Mitsukuri, K. (1896) A List of Holothurians known to occur in Japan. Zoological Magazine, 8, 405–413.
  84. Mitsukuri, K. (1912) Studies on Actinopodous Holothurioidea. Journal College Science, Tokyo Imperial University, 29 (2), 1–284, 1–8 pls. https://doi.org/10.5962/bhl.title.37880
  85. Miller, J.E. & Pawson, D.L. (1984) Holothurians (Echinodermata: Holothuroidea). Memoirs of the Hourglass Cruises. Vol. VII. Part. I. Florida Department of Natural Resources, Marine Research Laboratory, St. Petersburg, Florida. 79 pps
  86. Mongiardino Koch, N., Tilic, E., Miller, A.S., Stiller, J. & Rouse, G.W. (2023) Confusion will be my epitaph: genome-scale discordance stifles phylogenetic resolution of Holothuroidea. Proceedings of the Royal Society B, 290 (2002), 1–11. https://doi.org/10.1098/rspb.2023.0988
  87. O’Loughlin, P.M., Paulay, G., VandenSpiegel, D. & Samyn, Y. (2007) New Holothuria species from Australia (Echinodermata: Holothuroidea: Holothuriidae), with comments on the origin of deep and cool holothuriids. Memoirs of Museum Victoria, 64, 35–52. https://doi.org/10.24199/j.mmv.2007.64.5
  88. Panning, A. (1929) Die Gattung Holothuria (1. Teil). Mitteilungen aus dem Zoologishen Staatinstitut und Zoologischen Museum in Hamburg, 44, 91–138.
  89. Panning, A. (1934) Die Gattung Holothuria (3. Teil). Mitteilungen aus dem Zoologishen Staatinstitut und Zoologischen Museum in Hamburg, 45, 65–84.
  90. Panning, A. (1935) Die Gattung Holothuria (4. Teil). Mitteilungen aus dem Zoologishen Staatinstitut und Zoologischen Museum in Hamburg, 45, 85–107.
  91. Panning, A. (1940) Holothurien von den Kanaren und von Dakar. Videnskabelige Meddelelser Dansk Naturhistorik Forening, København, 103, 523–546.
  92. Paulay, G. (2003) The Asteroidea, Echinoidea, and Holothuroidea (Echinodermata) of the Mariana Islands. Micronesica, 35–36, 563–583.
  93. Pawson, D.L. (1978) Echinoderm fauna of Ascension Island, South Atlantic Ocean. Smithsonian Contribution to the Marine Sciences No. 2. Smithsonian Institution Press, Washington, 31 pp. https://doi.org/10.5479/si.01960768.2.1
  94. Pawson, D.L. (1995) Echinoderms of the tropical island Pacific: status of their sytematics and notes on their ecology and biogeography. In: Maragos, J.E., Peterson, M.N.A., Eldredge, L.G., Bardach, J.E. & Takeuchi, H.F. (Eds.), Marine and Coastal Biodiversity in the Tropical Island Pacific Region. Vol. 1. Species Systematics and Information Management Priorities. Program on Environment East-West Centre, Honolulu, Hawaii, pp. 171–192.
  95. Pearson, J. (1913) Notes on the Holothurioidea of the Indian Ocean. I. The genus Holothuria. Spolia Zeylanica, 9 (34), 49–101. https://doi.org/10.5962/bhl.part.7317
  96. Pearson, J. (1914) Notes on the Holothurioidea of the Indian Ocean. II. The sub-genera Argiodia and Actinopyga. Spolia Zeylanica, 9 (35), 173–190. https://doi.org/10.5962/bhl.part.7318
  97. Price, A.R.G. (1982) Comparison between Echinoderm fauna’s of Arabian Gulf, SE Arabia, Red Sea and Gulfs of Aqaba and Suez. Fauna of Saudi Arabia, 4, 3–21.
  98. Price, A.R.G., Keeling, M.J. & O’Callagahan, C.J. (1999) Ocean-scale patterns of ‘biodiversity’ of Atlantic asteroids determined from taxonomic distinctness and other measures. Biological Journal of the Linnean Society, 66, 187–203. https://doi.org/10.1006/bijl.1998.0275
  99. Purcell, S.W., Lovatelli, A., Gonzalez-Wangüemert, M., Solis-Marin, F.A., Samyn, Y. & Conand, C. (2023) Commercially important sea cucumbers of the world. 2nd Edition. FAO Species Catalogue for Fishery Purposes. No. 6. Rev. 1. FAO, Rome, 245 pp.
  100. Rowe, F.W.E. (1969) A review of the family Holothuriidae (Holothuroidea: Aspidochirotida). Bulletin of the BritishMuseum (Natural History), Zoology, 18 (4), 119–170. https://doi.org/10.5962/bhl.part.18419
  101. Rowe, F.W.E. & Doty, J.E. (1977) The shallow water holothurians of Guam. Micronesica, 13, 217–250.
  102. Rowe, F.W.E. & Gates, J. (1995) Echinodermata. In: Wells, A. (Ed.), Zoological Catalogue of Australia. Vol. 33. CSIRO, Australia, Melbourne, pp. i–xiii + 1–510.
  103. Rowe, F.W.E. & Richmond, M.D. (1997) Echinodermata. In: Richmond, M.D. (Ed.), A guide to the seashores of eastern Africa and the western Indian Ocean Islands. The SEA Trust, Zanzibar, pp. 290–321.
  104. Samyn, Y. (2003) Shallow-water Holothuroidea (Echinodermata) from Kenya and Pemba Island, Tanzania. Studies in Afrotropical Zoology, 292, 1–158.
  105. Samyn, Y. & Tallon, I. (2005) Zoogeography of the shallow-water holothuroids of the western Indian Ocean. Journal of Biogeography, 32, 1523–1538. https://doi.org/10.1111/j.1365-2699.2005.01295.x
  106. Samyn, Y., VandenSpiegel, D. & Massin, C. (2006) Taxonomie des holothuries des Comores. Abc Taxa, 1, i–iii + 1–130.
  107. Samyn, Y. & VandenBerghe, E. (2000) Annotated checklist of the echinoderms from the Kiunga Marine National Reserve, Kenya. Part I: Echinoidea and Holothuroidea. Journal of East African Natural History, 89, 1–36. https://doi.org/10.2982/0012-8317(2000)89[1:ACOTEF]2.0.CO;2
  108. Samyn, Y. & Thandar, A.S. (2010) Two new species in the phyllophorid genus Massinium (Echinodermata: Holothuroidea) with redescription of Massinium magnum. Zootaxa, 2399 (1), 1–19. https://doi.org/10.11646/zootaxa.2399.1.1
  109. Sastry, D.R.K. (2005) Echinodermata of Andaman and Nicobar Islands, Bay of Bengal. An annotated list. Zoological Survey of India, Occasional paper 233, 1–207.
  110. Sastry, D.R.K. (2007) Echinodermata of India. An annotated list. Zoological Survey of India, Occasional paper, 271, 1–387
  111. Schmidt, W.J (1930) Die Skeletstücke der Stachhelhäuter als Biokristalle. Zoologische Jahrbücher Abteilung für Allgemeine Zoologie und Physiologie der Tiere, 47, 357–510.
  112. Schoppe, S. (2000) Echinoderm of the Philippines. a guide to common shallow water sea stars, brittle stars, sea urchins, sea cucumbers and feather stars. Times Editions, Singapore, 144 pp.
  113. Selenka, E. (1867) Beiträge zur Anatomie und Systematik der Holothurien. Zeitschrift für wissenschaftliche Zoologie, 17 (2), 291–374.
  114. Semper, C. (1868) Reisen im Archipel der Philippinen. Holothurien. 2. Wissenschaftliche Resultate. W. Engelmann, Leipzig, x + 288 pp., 40 pls. https://doi.org/10.5962/bhl.title.11687
  115. Sluiter, C.P. (1887) Die Evertebraten Aus Der Sammlung Des Königlichen Naturwissenschatlichen Vereins in Niederländisch Indien in Batavia. Natuurkundig Tijdschrift voor Nederlandsch-Indië, 47, 8ste Serie, 181–220, pls. 1–2.
  116. Sluiter, C.P. (1901) Die Holothurien der Siboga-Expedition. Siboga-Expeditie. Monograph 44, 1–142, pls. 1–11. https://doi.org/10.5962/bhl.title.85348
  117. Sluiter, C.P. (1910) Westindische Holothurien. Ergebnisse Zoologische Forschungereise nach Westindien von W. Kukenthal u. R. Hartmeyer. Zoologische Jahrbücher, Abteilung Systematik, Ökologie Geographie Tiere, Supplement 11 (Heft 2), 331–341.
  118. Suarez, A.V. & Tsutsui, N.D. (2004) The Value of Museum Collections for Research and Society. BioScience, 54 (1), 66–74. https://doi.org/10.1641/0006-3568(2004)054[0066:TVOMCF]2.0.CO;2
  119. Suárez, A.M. (1974) Lista de equinodermos cubanos recientes. Ciencias Investigaciones Marinas, 6, 1–72.
  120. Thandar, A.S, Zettler, M.L. & Arumugan, P. (2010) Additions to the sea cucumber fauna of Namibia and Angola, with descriptions of new taxa (Echinodermata: Holothuroidea). Zootaxa, 2655 (1), 1–24. https://doi.org/10.11646/zootaxa.2655.1.1
  121. Thandar, A.S. & Mjobo, S. (2014) On some sea cucumbers from Ghana (Echinodermata: Holothuroidea) with descriptions of a new genus and one new species. Zootaxa, 3900 (2), 243–254. https://doi.org/10.11646/zootaxa.3900.2.4
  122. Théel, H. (1886) Holothuroidea. Part 2. Report of the scientific Results of the Voyage of the “Challenger”, Zoologie, 39, 1–290, pls 1–16.
  123. Tikasingh ES.1963. The shallow water Holothurians of Curaçao, Aruba and Bonaire. Study Fauna Curaçoa, 14, 77–99.
  124. Tommasi, L.R. (1969) Lista dos Holothurioidea recentes do Brasil. Contribuições avulsas do Instituto Océanográfico, Universidade de São Paulo, Série: Oceanográfia biológica, 15, 1–29.
  125. Tortonese, E. (1977) Report on the echinoderms from the Gulf of Aqaba (Red Sea). Monitore zooligia italiano, New Series, Supplement IX, 273–290. https://doi.org/10.1080/03749444.1977.10736852
  126. Tortonese, E. (1980) Researches on the coast of Somalia. Littoral Echinodermata. Monitore Zoologica Italiano, NS Supplementi, 13, 99–139. https://doi.org/10.1080/00269786.1980.11758550
  127. Valle-Garcia del, R., Abreu Pérez, M., Rodriguez, R., Solis-Marin, F.A., Laguarda-Figueras, A. & Duran Gonzalez, A. (2008) Equinodermos (Echinodermata) del occidente del Archipiélago Sabana-Camagüey, Cuba. Revista Biologia Tropical, 56 (Supplement 3), 19–35.
  128. Verrill, A.E. (1902) Additions to the Fauna of the Bermudas from the Yale Expedition of 1901, with Notes on Other Species. Transactions of the Conneticut Academy of Arts and Sciences, 11, 15–62.
  129. Verrill, A.E. (1907) The Bermuda Islands. Part IV. Geology and Paleontology. Transactions of the Conneticut Academy of Arts and Sciences, 12, 143–146. https://doi.org/10.5962/bhl.part.28921