Diagnostic Methods for the Comprehensive Health Assessment of the Long-Spined Sea Urchin, Diadema antillarum
Using a dissecting microscope to examine D. antillarum. This animal was to be necropsied, so spines were cut prior to the exam.
view on EDIS


Diadema antillarum
Long spined Sea Urchin
Diagnostic methods

How to Cite

Francis-Floyd, Ruth. 2020. “Diagnostic Methods for the Comprehensive Health Assessment of the Long-Spined Sea Urchin, Diadema Antillarum”. EDIS 2020 (3). https://doi.org/10.32473/edis-vm244-2020.


This 55-page document contains a wealth of information on the long-spined sea urchin, which is an important animal in coral reef ecosystems. The publication discusses diagnostic methods for Diadema antillarum and provides reporting forms for laboratory and diagnostic work. Written by Ruth Francis-Floyd, Jan Landsberg, Roy Yanong, Yasu Kiryu, Shirley Baker, Deborah Pouder, William Sharp, Gabriel Delgado, Nicole Stacy, Tom Waltzek, Heather Walden, Roxanna Smolowitz, and Greg Beck, and published by the UF/IFAS Veterinary Medicine—Large Animal Clinical Sciences Department, May 2020.

view on EDIS


Applegate, J.R., Dombrowski, D.S., Christian, L.S., Bayer, M.P., Harms, C.A., Lewbart, G.A. 2016. Tricaine methanesulfonate (MS-222) sedationand anesthesia in the purple-spined sea urchin (Arbacia punctulata). J Zoo Wildl Med 47(4):1025-1033. https://doi.org/10.1638/2015-0288.1

Bak RPM, Carpay MJE, De Ruyter Van Steveninck ED (1984) Densities of the sea urchin Diadema antillarum before and after mass mortalities on the coral reefs on Curacao. Mar Ecology Prog Ser 17:105-108 https://doi.org/10.3354/meps017105

Bauer JC (1976) Growth, aggregation and maturation in the echinoid, Diadema antillarum. Bull Mar Sci 26 (2):273-277.

Bauer JC, Agerter C (1987) Isolation of bacteria pathogenic for the sea urchin Diadema antillarum (Echinodermata: Echinoidea). Bull Mar Sci 40:161-165

Bauer JC, Agerter CJ (1994) Isolation of potentially pathogenic bacterial flora from tropical sea urchins in selected West Atlantic and East Pacific sites. Bull Mar Sci 55:142-150

Bauer JC, Young CM (2000) Epidermal lesions and mortality caused by vibriosis in deep-sea Bahamian echinoids: a laboratory study. Dis Aquat Org 39:193-199 https://doi.org/10.3354/dao039193

Beck G, Miller R, Ebersol J (2009) Significance of immunological responses in the black-spined sea urchin, Diadema antillarum, to Caribbean-wide mass mortality. Proceedings of the 11th International Coral Reef Symposium (Ft. Lauderdale, FL): 221-225

Beck G, Miller R, Ebersol J (2014) Mass mortality and slow recovery of Diadema antillarum: could compromised immunity be a factor? Mar Biol 161:1001-1013. https://doi.org/10.1007/s00227-013-2382-6

Becker P, Gillan DC, Eeckhaut I (2007) Microbiological study of the body wall lesions of the echinoid Tripneustes gratilla. Dis Aquat Org 77:73-82 https://doi.org/10.3354/dao01821

Becker PT, Egea E, Eeckhaut I (2008) Characterization of the bacterial communities associated with the bald sea urchin disease of the echinoid Paracentrotus lividus. J Invertebr Pathol 98:136-147 https://doi.org/10.1016/j.jip.2007.12.002

Benítez-Villalobos F, Díaz Martínez JP, Martínez-García M (2009) Mass mortality of the sea urchin Diadema mexicanum in La Entrega at Bahias de Huatulco, Western Mexico. Coral Reefs 28:1017 https://doi.org/10.1007/s00338-009-0541-5

Biggar RB, Wenrich DH (1932) Studies on ciliates from Bermuda sea urchins. J Parasitol 18:252-257 https://doi.org/10.1186/1742-9994-6-10

Bodmer MDV, Rogers AD, Speight MR, Lubbock N, Exton DA (2015) Using an isolated population boom to explore barriers to recovery in the keystone Caribbean coral reef herbivore Diadema antillarum. Coral Reefs 34:1011-1021 https://doi.org/10.1007/s00338-015-1329-4

Boudouresque C-F, Nedelec H, Shepherd SA (1980) The decline of a population of the sea urchin Paracentrotus lividus in the Bay of Port-Cros (Var, France). Trav Sci Parc Nation Port-cros Fr 6:243-251

Buchwald RT, Feehan CJ, Scheibling RE, Simpson AGB (2015) Low temperature tolerance of a sea urchin pathogen: Implications for benthic community dynamics in a warming ocean. J Exp Mar Biol Ecol 469:1-9 https://doi.org/10.1016/j.jembe.2015.04.006

Carpenter RC (1988) Mass mortality of a Caribbean sea urchin: Immediate effects on community metabolism and other herbivores. Proc Natl Acad Sci USA 85:511-514 https://doi.org/10.1073/pnas.85.2.511

Carpenter RC (1990a) Mass mortality of Diadema antillarum I. Long-term effects on sea urchin population-dynamics and coral reef algal communities. Mar Biol 104:67-77 https://doi.org/10.1007/BF01313159

Carpenter RC (1990b) Mass mortality of Diadema antillarum II. Effects on population densities and grazing intensity of parrotfishes and surgeonfishes. Mar Biol 104:79-86 https://doi.org/10.1007/BF01313160

Cavey, MJ, K Märkel (1994) Echinoidea. In: Harrison FW, Chia F (eds) Microscopic Anatomy of Invertebrates, Vol. 14, Echinodermata. Wiley-Liss, Inc., New York, p345-400

Chiappone M, Rutten LM, Swanson DW, Miller SL (2008) Population status of the urchin Diadema antillarum in the Florida Keys 25 years after the Caribbean mass mortality. In: Proceedings of the Eleventh International Coral Reef Symposium, Ft. Lauderdale, Florida, 7-11 July 2008, p 706-710

Chiappone M, Swanson D, Miller S, Smith S (2002) Large-scale surveys on the Florida Reef Tract indicate poor recovery of the long-spined sea urchin Diadema antillarum. Coral Reefs 21:155-159

Clemente S, Lorenzo-Morales J, Mendoza JC, López C, Sangil C, Alves F, Kaufmann M, Hernández JC (2014) Sea urchin Diadema africanum mass mortality in the subtropical eastern Atlantic: role of waterborne bacteria in a warming ocean. Mar Ecol Prog Ser 506:1-14 https://doi.org/10.3354/meps10829

Dame, E A (2008) Assessing the effect of artificial habitat structure on translocation of the long-spined sea urchin, Diadema antillarum, in Curacao (Netherlands Antilles). Bull Mar Sci 82(2): 247-254

de Ruyter Van Steveninck ED, Bak RPM (1986) Changes in abundance of coral-reef bottom components related to mass mortality of the sea urchin Diadema antillarum. Mar Ecol Prog Ser 34:87-94 https://doi.org/10.3354/meps034087

de Ruyter van Steveninck ED, Breeman AM (1987) Deep water vegetations of Lobophora variegata (Phaeophyceae) in the coral reef of Curacao—population dynamics in relation to mass mortality of the sea urchin Diadema antillarum. Mar Ecol Prog Ser 36:81-90 https://doi.org/10.3354/meps036081

Dumont CP, Himmelman JH, Russell MP (2004) Sea urchin mass mortality associated with algal debris from ice scour. In: Heinzeller T, Nebelsick J (eds) Echinoderms: Munchen. Taylor & Francis, Munich, p 177-182 https://doi.org/10.1201/9780203970881.ch31

Dykova I, Lorenzo-Morales J, Kostka M, Valladares B, Peckova H (2011) Neoparamoeba branchiphila infections in moribund sea urchins Diadema aff. antillarum in Tenerife, Canary Islands, Spain. Dis Aquat Org 95:225-231 https://doi.org/10.3354/dao02361

Feehan CJ, Johnson-Mackinnon J, Scheibling RE, Lauzon-Guay JS, Simpson AG (2013) Validating the identity of Paramoeba invadens, the causative agent of recurrent mass mortality of sea urchins in Nova Scotia, Canada. Dis Aquat Org 103:209-227 https://doi.org/10.3354/dao02577

Feehan CJ, Scheibling RE (2014) Effects of sea urchin disease on coastal marine ecosystems. Mar Biol 161:1467-1485 https://doi.org/10.1007/s00227-014-2452-4

Feehan, C. J., Brown, M. S., Sharp, W. C., Lauzon‐Guay, J. S., & Adams, D. K. (2016). Fertilization limitation of Diadema antillarum on coral reefs in the Florida Keys. Ecology. 97(8):1897-1904. https://doi.org/10.1002/ecy.1461

Forcucci D (1994) Population density, recruitment and 1991 mortality event of Diadema antillarum in the Florida Keys. Bull Mar Sci 54:917-928

Francis-Floyd, R, Landsberg J, Yanong R, Baker S, Stacy N, Waltzek T, Kiryu Y, Hunt J, Sharp W, Smolowitz R, Beck G, Leber K, Vaughan D, Moe M. 2016. Final Report: Development of a Health Protocol for Hatchery-Reared Long-Spined Sea Urchin, Diadema antillarum. Florida Fish and Wildlife Conservation Commission Agreement No. 14044, SWG CFDA No. 15634. Tallahassee FL.

Furman B, Heck KL (2009) Differential impacts of echinoid grazers on coral recruitment. Bull Mar Sci 85:121-132

Garcia LS, Bruckner DA (1993) Examination of fecal specimens In: Diagnostic Medical Parasitology, 2nd Edition, American Society for Microbiology, Washington D.C., 764 pp.

Gardner TA, Cote IM, Gill JA, Grant A, Watkinson AR (2003) Long-term region-wide declines in Caribbean corals. Science 301:958-960 https://doi.org/10.1126/science.1086050

Gilles KW, Pearse JS (1986) Disease in sea urchins Strongylocentrotus purpuratus: experimental infection and bacterial virulence. Dis Aquat Org 1:105-114 https://doi.org/10.3354/dao001105

Girard D, Clemente S, Toledo-Guedes K, Brito A, Hernández JC (2012) A mass mortality of subtropical intertidal populations of the sea urchin Paracentrotus lividus: analysis of potential links with environmental conditions. Mar Ecol 33:377-385 https://doi.org/10.1111/j.1439-0485.2011.00491.x

Groliere CA, Puytorac Pd, Grain J (1980) Observations de quelques especes de cilies endocommensaux d'echinides du Golfe du Mexique et de la mer des Antilles. Protistol 16:233-239

Hagen NT (1987) Sea urchin outbreaks and nematode epizootics in Vestfjorden, northern Norway. Sarsia 72:213-229 https://doi.org/10.1080/00364827.1987.10419719

Hagen NT (1992) Macroparasitic epizootic disease: a potential mechanism for the termination of sea urchin outbreaks in Northern Norway? Mar Biol 114:469-478 https://doi.org/10.1007/BF00350039

Hagen NT (1996) Parasitic castration of the green echinoid Strongylocentrotus droebachiensis by the nematode endoparasite Echinomermella matsi: Reduced reproductive potential and reproductive death. Dis Aquat Org 24:215-226 https://doi.org/10.3354/dao024215

Harms, CA (2006) Echinoderms, In: Invertebrate Medicine, GA Lewbart (ED). Blackwell Publishing, Ames IA. Pp245-256. https://doi.org/10.1002/9780470344606.ch17

Hendler G (2013) Recent mass mortality of Strongylocentrotus purpuratus (Echinodermata: Echinoidea) at Malibu and a review of purple sea urchin kills elsewhere in California. Bull South Cal Acad Sci 112:19-37 https://doi.org/10.3160/0038-3872-112.1.19

Hughes TP, Keller BD, Jackson JBC, Boyle MJ (1985) Mass mortality of the echinoid Diadema antillarum Philippi in Jamaica. Bull Mar Sci 36:377-384

Hughes TP, Reed DC, Boyle M-J (1987) Herbivory on coral reefs: community structure following mass mortalities of sea urchins. J Exp Mar Biol Ecol 113:39-59 https://doi.org/10.1016/0022-0981(87)90081-5

Hunte W, Côté I, Tomascik T (1986) On the dynamics of the mass mortality of Diadema antillarum in Barbados. Coral Reefs 4:135-139 https://doi.org/10.1007/BF00427934

Jackson JBC (2008) Ecological extinction and evolution in the brave new ocean. Proc Natl Acad Sci USA 105:11458- 11465 https://doi.org/10.1073/pnas.0802812105

Jangoux M (1984) Diseases of echinoderms. Helgo Meeresunters 37:207-216 https://doi.org/10.1007/BF01989305

Jangoux M (1987a) Diseases of Echinodermata. I. Agents microorganisms and protistans Dis Aquat Org 2: 147-162

Jangoux M (1987b) Diseases of Echinodermata. II. Agents metazoans (Mesozoa to Bryozoa). Dis Aquat Org 2:205-234. https://doi.org/10.3354/dao002205

Jangoux M (1987c) Diseases of Echinodermata. III. Agents metazoans (Annelida to Pisces). Dis Aquat Org 3:59-83. https://doi.org/10.3354/dao003059

Jangoux M (1987d) Diseases of Echinodermata. IV. Structural abnormalities and general considerations on biotic diseases. Dis Aquat Org 3:221-229. https://doi.org/10.3354/dao003221

Jangoux M (1990) Diseases of Echinodermata. In: Kinne O (ed) Diseases of Marine Animals, Book 3. Biologische Anstalt Helgoland, Hamburg, Germany, pp. 439-567

Jellett JF, Wardlaw AC, Scheibling RE (1989) Experimental infection of the sea urchin (Strongylocentrotus droebachiensis) with a pathogenic amoeba (Paramoeba invadens) : quantitative changes in the coelomic fluid In vivo and cellular interaction in vitro. Dev Comp Immunol 13:428-429 https://doi.org/10.1016/0145-305X(89)90152-3

Jones GM, Scheibling RE (1985) Paramoeba sp. (Amoebida, Paramoebidae) as the possible causative agent of sea urchin mass mortality in Nova Scotia. J Parasitol 71:559-565 https://doi.org/10.2307/3281424

Jones GM, Schiebling RE, Hebda AJ, Miller RJ (1984) Amoebae in tissues of diseased echinoids (Strongylocentrosus droebachiensis) in Nova Scotia. In: Keegan BF, O'Connor BDS (eds). Proc Fifth Int. Echinoderm Conf. pp. 289-293

Jones IW, Rogers TE (1968) Studies on the endocommensal ciliate fauna of Caribbean sea urchins. Biol Bull 135:514-519 https://doi.org/10.2307/1539713

Jurgens LJ, Rogers-Bennett L, Raimondi PT, Schiebelhut LM, Dawson MN, Grosberg RK, Gaylord B (2015) Patterns of mass mortality among rocky shore invertebrates across 100 km of northeastern Pacific coastline. PloS One 10:e0126280 https://doi.org/10.1371/journal.pone.0126280

Kissling DL, Precht WF, Miller SL, Chiappone M (2014) Historical reconstruction of population density of the echinoid Diadema antillarum on Florida Keys shallow bank-barrier reefs. Bull Mar Sci 90:665-679 https://doi.org/10.5343/bms.2013.1022

Knowlton N (2001) Sea urchin recovery from mass mortality: new hope for Caribbean coral reefs? Proc Natl Acad Sci USA 98:4822-4824 https://doi.org/10.1073/pnas.091107198

Lessios HA (1988) Mass mortality of Diadema antillarum in the Caribbean: what have we learned? Annu Rev Ecol Syst 19:371-393 https://doi.org/10.1146/annurev.es.19.110188.002103

Lessios HA (1995) Diadema antillarum 10 years after mass mortality: still rare, despite help from a competitor. Proc Roy Soc Lond Series B: Biol Sci 259:331-337 https://doi.org/10.1098/rspb.1995.0049

Lessios HA (2005) Diadema antillarum populations in Panama twenty years following mass mortality. Coral Reefs 24:125-127 https://doi.org/10.1007/s00338-004-0443-5

Lessios, H. A. 2016. The great Diadema antillarum die-off: 30 years later. Annu Rev Mar Sci 8:267-283 https://doi.org/10.1146/annurev-marine-122414-033857

Lessios HA, Cubit JD, Robertson DR, Shulman MJ, Parker MR, Garrity SD, Levings SC (1984a) Mass mortality of Diadema antillarum on the Caribbean coast of Panama. Coral Reefs 3:173-182 https://doi.org/10.1007/BF00288252

Lessios H, Glynn P, Robertson D (1983) Mass mortalities of coral reef organisms. Science (New York, NY) 222:715 https://doi.org/10.1126/science.222.4625.715

Lessios HA, Robertson DR, Cubit JD (1984b) Spread of Diadema mass mortality through the Caribbean. Science 226:335-337 https://doi.org/10.1126/science.226.4672.335

Lester SE, Tobin ED, Behrens MD (2007) Disease dynamics and the potential role of thermal stress in the sea urchin, Strongylocentrotus purpuratus. Can J Fish Aquat Sci 64:314-323 https://doi.org/10.1139/f07-010

Levitan DR (1988) Algal-urchin biomass responses following mass mortality of Diadema antillarum Philippi at Saint John, U.S. Virgin Islands. J Exp Mar Biol Ecol 119:167-178 https://doi.org/10.1016/0022-0981(88)90231-6

Li T, Xu S (2000) Preliminary studies on the black mouth disease of sea urchin, Strongylocentrotus intermedius (Strongylocentrotidae Echinoidea). Mar Sci Qingdao 24:43-47

Liddell WD, Ohlhorst SL (1986) Changes in benthic community composition following the mass mortality of Diadema at Jamaica. J Exp Mar Biol Ecol 95:271-278 https://doi.org/10.1016/0022-0981(86)90259-5

Luna LG (1968) Manual of histologic staining methods of the Armed Forces Institutes of Pathology, 3rd edition. McGraw-Hill, New York

Maes P, Jangoux M (1984) The bald-sea-urchin disease: a biopathological approach. Helgol Meeresunters 37:217-224 https://doi.org/10.1007/BF01989306

Martony, M., Pouder, D., Yanong, R., Kiryu, Y., Landsberg, J.H., Isaza, R., Waltzek, T., Stacy, N., Giglio, R., Baker, S., Francis-Floyd, R. 2018. Establishing a diagnostic technique for coelomocentesis in the long-spined sea urchin Diadema antillarum. Journal of Aquatic Animal Health DOI:10.1002/aah.10043. https://doi.org/10.1002/aah.10043

Masuda Y, Tajima K, Ezura Y (2004) Resuscitation of Tenacibaculum sp., the causative bacterium of spotting disease of sea urchin Strongylocentroutus intermedius, from the viable but non-culturable state. Fish Sci 70:277-284 https://doi.org/10.1111/j.1444-2906.2003.00801.x

McClendon JF (1912) Echinochrome, a red substance in sea urchins. J of Biol Chem 11:435-441

McManus JW, Polsenberg JF (2004) Coral–algal phase shifts on coral reefs: Ecological and environmental aspects. Prog Oceanogr 60:263-279 https://doi.org/10.1016/j.pocean.2004.02.014

Miller R, Adams A, Ogden N, Ogden J, Ebersole J (2003) Diadema antillarum 17 years after mass mortality: Is recovery beginning on St. Croix? . Coral Reefs 22:181-187 https://doi.org/10.1007/s00338-003-0301-x

Miller RJ, Colodey AG (1983) Widespread mass mortalities of the green sea urchin in Nova Scotia, Canada. Mar Biol 73:263-267 https://doi.org/10.1007/BF00392252

Mumby PJ, Hedley JD, Zychaluk K, Harborne AR, Blackwell PG (2006) Revisiting the catastrophic die-off of the urchin Diadema antillarum on Caribbean coral reefs: Fresh insights on resilience from a simulation model. Ecol Model 196:131-148 https://doi.org/10.1016/j.ecolmodel.2005.11.035

Murillo-Castro MM, Cortés-Núñez J (1984) Alta mortalidad en la población del erizo de mar Diadema antillarum Philippi (Echinodermata: Echinoidea), en el Parque Nacional Cahuita, Limón, Costa Rica. Rev Biol Trop 32:167-169

Nagelkerken I, Smith GW, Snelders E, Karel M, James S (1999) Sea urchin Meoma ventricosa die-off in Curacao (Netherlands Antilles) associated with a pathogenic bacterium. Dis Aquat Org 38:71-74 https://doi.org/10.3354/dao038071

NOAA/FKNMS. 2015. Florida Keys National Marine Sanctuary 2011 Condition Report. http://sanctuaries.noaa.gov/science/condition/fknms/state.html. Accessed 24 March 2016

Noga EJ (2010) Fish Disease: Diagnosis and Treatment, 2nd Edition. Wiley Blackwell, Ames IA. Pp 13-48. https://doi.org/10.1002/9781118786758

Pearse JS, Costa DP, Yellin MB, Agegian CR (1977) Localized mass mortality of red sea urchin, Strongylocentrotus franciscanus, near Santa Cruz, California. Fish Bull 75:645-648

Phinney JT, Muller-Karger F, Dustan P, Sobel J (2001) Using remote sensing to reassess the mass mortality of Diadema antillarum 1983-1984. Conserv Biology 15:885-891 https://doi.org/10.1046/j.1523-1739.2001.015004885.x

Quintero-Hunter I, Grier H, Muscato M (1991) Enhancement of histological detail using metanil yellow as a counter-stain in periodic acid/Schiff’s hematoxylin staining of glycol methacrylate tissue sections. Biotech Histochem 66:169–172 https://doi.org/10.3109/10520299109109964

Roberts-Regan DL, Scheibling RE, Jellett JF (1988) Natural and experimentally induced lesions of the body wall of the sea urchin Strongylocentrotus. Dis Aquat Org 5:51-62 https://doi.org/10.3354/dao005051

Rogers, A. and K. Lorenzen. 2016. Does slow and variable recovery of Diadema antillarum on Caribbean fore-reefs reflect density-dependent habitat selection? Front Mar Sci Vol 3, Article 63 (DOI: 10.3389/FMARS.2016.00063). https://doi.org/10.3389/fmars.2016.00063

Scheibling RE, Feehan CM, Lauzon-Guay J-S (2010) Disease outbreaks associated with recent hurricanes cause mass mortality of sea urchins in Nova Scotia. Mar Ecol Prog Ser 408:109-116 https://doi.org/10.3354/meps08579

Scheibling RE, Hennigar AW (1997) Recurrent outbreaks of disease in sea urchins Strongylocentrotus droebachiensis in Nova Scotia: evidence for a link with large-scale meteorologic and oceanographic events. Mar Ecol Prog Ser 152:155-165 https://doi.org/10.3354/meps152155

Scheibling RE, Stephenson RL (1984) Mass mortality of Strongylocentrotus droebachiensis (Echinodermata: Echinoidea) off Nova Scotia, Canada. Mar Biol 78:153-164 https://doi.org/10.1007/BF00394695

Sharp, WC, Delgado GA, Hart JE, and Hunt, JH (2018). Comparing the behavior and morphology of wild-collected and hatchery-propagated long-spined sea urchins (Diadema antillarum): implications for coral reef ecosystem restoration. Bull Mar Sci 94(1):103-122. https://doi.org/10.5343/bms.2017.1068

Shimizu M (1994) Histopathological investigation of the spotted gonad disease in the sea urchin, Strongylocentrotus intermedius. J Invert Pathol 63:182-187 https://doi.org/10.1006/jipa.1994.1034

Sivertsen K (1996) Incidence, occurrence and distribution of the nematode Echinomermella matsi in its echinoid host, Strongylocentrotus droebachiensis, in northern Norway. Mar Biol:703 https://doi.org/10.1007/BF00351337

Skadsheim A, Christie H, Leinaas HP (1995) Population reductions of Strongylocentrotus droebachiensis (Echinodermata) in Norway and the distribution of its endoparasite Echinomermella matsi (Nematoda). Mar Ecol Prog Ser 119:199-209 https://doi.org/10.3354/meps119199

Sogabe Y, Tajima K, Tanaka R, Sawabe T, Ezura Y (2002) Development of 16S rRNA targeted PCR for the identification of Vibrio spp., the causative bacteria of the disease in cultured sea urchin Strongylocentrotus intermedius occurring at low water temperatures. Bull Jap Soc Sci Fish (Japan) 68:201-206 https://doi.org/10.2331/suisan.68.201

Stacy, N.I., Frankovich, T., Walden, H., Francis-Floyd, R. In Press. From macro- to microcosm: the art and beauty of plant material anddiatoms in sea urchin fecal cytology specimens. Vet Clin Pathol (In Press).

Stien A (1999) Effects of the parasitic nematode Echinomermella matsi on growth and survival of its host, the sea urchin Strongylocentrotus droebachiensis. Can J Zool 77:139-147 https://doi.org/10.1139/cjz-77-1-139 https://doi.org/10.1139/z98-159

Stien A, Leinaas HP, Halvorsen O, Christie H (1998) Population dynamics of the Echinomermella matsi (Nematoda) Strongylocentrotus droebachiensis (Echinoida) system: effects on host fecundity. Mar Ecol Prog Ser 163:193-201 https://doi.org/10.3354/meps163193

Tajima K, Cunha da Silva JRM, Lawrence JM (2007) Disease in sea urchins. In: Lawrence JM (ed) Edible Sea Urchins: Biology and Ecology, p 167-189 https://doi.org/10.1016/S0167-9309(07)80073-1

Tajima K, Hirano T, Nakano K, Ezura Y (1997a) Taxonomical study on the causative bacterium of spotting disease of sea urchin Strongylocentrotus intermedius. Fish Sci 63:897-900 https://doi.org/10.2331/fishsci.63.897

Tajima K, Hirano T, Shimizu M, Ezura Y (1997b) Isolation and pathogenicity of the causative bacterium of spotting disease of sea urchin Strongylocentrotus intermedius. Fish Sci 63:249-252 https://doi.org/10.2331/fishsci.63.249

Tajima K, Lawrence JM (2001) Disease in edible sea urchins. In: Lawrence JM (ed) Edible Sea Urchins: Biology and Ecology, Book 32. Elsevier, p 139-148 https://doi.org/10.1016/S0167-9309(01)80010-7

Tajima K, Shimizu M, Miura K, Ohsaki S, Nishihara Y, Ezura Y (1998) Seasonal fluctuations of Flexibacter sp. the causative bacterium of spotting disease of sea urchin Strongylocentrotus intermedius in the culturing facilities and coastal area. Fish Sci 64:6-9 https://doi.org/10.2331/fishsci.64.6

Takeuchi K, Tajima K, Iqbal MM, Sawabe T, Ezura Y (1999) Taxonomical and serological studies on the causative bacteria of the disease of sea urchin Strongylocentrotus intermedius occurring at low water temperatures. Fish Sci 65:264-268 https://doi.org/10.2331/fishsci.65.264

Taniuchi Y, Tajima K, Shimono I, Ezura Y (2002) Survival of Flexibacter sp. strain F-2, the causative bacterium of spotting disease of sea urchin Strongylocentrotus intermedius at low temperatures. Bull Jap Soc Sci Fish (Japan):46-51

Urdaneta-Morales S, De McLure MT (1966) The endocommensal ciliates of Venezuelan sea urchins. J Protozool 13:5-8 https://doi.org/10.1111/j.1550-7408.1966.tb01859.x

Uthicke S, Schaffelke B, Byrne M (2009) A boom-bust phylum? Ecological and evolutionary consequences of density variations in echinoderms. Ecol Monogr 79:3-24 https://doi.org/10.1890/07-2136.1

Violetta, H., Francis-Floyd, R., Yanong, R.P.E., Landsberg, J.H. 2017. Validation of health assessment criteria for Diadema antillarum. Proceedings of the 48th Conference of the International Association for Aquatic Animal Medicine, Cancun MX (May 22 2017).

Wang Y, Feng N, Li Q, Ding J, Zhan Y, Chang Y (2013a) Isolation and characterization of bacteria associated with a syndrome disease of sea urchin Strongylocentrotus intermedius in North China. Aquac Res 44:691-700 https://doi.org/10.1111/j.1365-2109.2011.03073.x

Wang YN, Chang YQ, Lawrence JM (2013b) Disease in sea urchins. In: Lawrence JM (ed) Sea Urchins: Biology and Ecology. Elsevier B. V., p 179-186 https://doi.org/10.1016/B978-0-12-396491-5.00012-5

Ward JR, Lafferty KD (2004) The elusive baseline of marine disease: are diseases in ocean ecosystems increasing? PLoS Biol 2:E120 https://doi.org/10.1371/journal.pbio.0020120

Weil E, Torres JL, Ashton M (2005) Population characteristics of the sea urchin Diadema antillarum in La Parguera, Puerto Rico, 17 years after the mass mortality event. Rev Biol Trop 53 Suppl 3:219-231

Williams EH, Bunkley-Williams L, Bruckner RJ, Bruckner AW, Ortiz-Corps EAR, Bowden-Kerby WA, Colon-Jones DE (1996) Recurring mass mortalities of the white-spined sea urchin, Tripneustes ventricosus, (Echinodermata: Echinoidea) in Puerto Rico. Caribb J Sci 32:111

Williams LB, Williams Jr EH, Bunkley Jr AG (1986) Isolated mortalities of the sea urchins Astropyga magnifica and Eucidaris tribuloides in Puerto Rico. Bull Mar Sci 38:391-393

Williams S, Yoshioka P, García Sais J (2010) Recruitment pattern of Diadema antillarum in La Parguera, Puerto Rico. Coral Reefs 29:809 https://doi.org/10.1007/s00338-010-0633-2

Ziegler A, Faber C, Bartolomaeus T (2009) Comparative morphology of the axial complex and interdependence of internal organ systems in sea urchins (Echinodermata: Echinoidea). Frontiers in Zoology 6:10 https://doi.org/10.1186/1742-9994-6-10

Copyright (c) 2020 UF/IFAS