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A review of Caribbean Copepoda associated with reef-dwelling cnidarians, and sponges Korzhavina, Oksana A.; Hoeksema, Bert W.; Ivanenko, Viatcheslav N.

Published in: Contributions to Zoology

DOI: 10.1163/18759866-20191411

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Citation for published version (APA): Korzhavina, O. A., Hoeksema, B. W., & Ivanenko, V. N. (2019). A review of Caribbean Copepoda associated with reef-dwelling cnidarians, echinoderms and sponges. Contributions to Zoology, 88(3), 297- 349. https://doi.org/10.1163/18759866-20191411

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A review of Caribbean Copepoda associated with reef-dwelling cnidarians, echinoderms and sponges

Oksana A. Korzhavina Department of Invertebrate Zoology, Biological Faculty, Lomonosov Moscow State University, Moscow, Russia

Bert W. Hoeksema and Systematics Group, Naturalis Biodiversity Center, Leiden, The Netherlands Groningen Institute for Evolutionary Life Sciences, University of Groningen, Groningen, The Netherlands

Viatcheslav N. Ivanenko Department of Invertebrate Zoology, Biological Faculty, Lomonosov Moscow State University, Moscow, Russia Taxonomy and Systematics Group, Naturalis Biodiversity Center, Leiden, The Netherlands [email protected]

Abstract

This review of associated with reef-dwelling cnidarians, sponges and echinoderms of the Greater Caribbean is based on published records, systematically arranged by the classification of symbiotic and their hosts, sampling sites, coordinates, depth and date of sampling, litera- ture sources, and three recent surveys (Cuba, St. Eustatius in the Eastern Caribbean and Curaçao in the Southern Caribbean). This resulted in totals of 532 records of 115 species of symbiotic copepods (47 gen- era, 17 families, three orders) hosted by 80 species of invertebrates, representing scleractinians (47%), octocorals (9%), echinoderms (3%), and sponges (1%). Among ten Caribbean ecoregions, the Greater Antilles (with 64 species of symbiotic copepods) as well as the Southern and Eastern Caribbean (with 46 and 17 species of copepods, respectively) are the most studied and best represented, whereas only six species of copepods are known from Bermuda, one from Southwestern Caribbean and none from the Gulf of Mexico. The absence of poecilostomatoid copepods (Anchimolgidae, Rhynchomolgidae and Xarifidae) on Caribbean stony corals as noted by Stock (1988) is confirmed. The results indicate that the diversity and ecology of Caribbean symbiotic copepods are still poorly investigated.

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298 KORZHAVINA et al.

Keywords

Crustacea – Cnidaria – Echinodermata – Porifera – reefs, diversity – symbiosis – parasitism

Introduction abundant on sponges (Stock & Kleeton, 1964; Stock, 1967; Boxshall & Huys, 1994; Kim, 2010; Symbiotic copepods are a widespread, numer- Varela et al., 2005b, 2007a, b, 2008; Varela, ous and diverse group of crustaceans living in 2010a, b, 2012). Twelve families of copepods association (parasitism, commensalism, mu- have been reported on diverse Caribbean tualism) with other marine ­(Gotto, echinoderms (Edwards, 1891; Emson & Mlad- 1979, 1993; Humes, 1985a, 1994; Ho, 2001). At enov, 1987; Emson et al., 1985; Hendler & Kim, least one third of all known copepods are 2010; Humes & Hendler, 1972, 1999; Humes & symbionts of marine fish and invertebrates. Ho, 1970, 1971; Humes & Stock, 1973; Humes, ­Symbiotic copepods are the most diverse in 1969a, 1998, 2000; Kim, 2009, 2010; Stock & the tropics, and only a small number of their Gooding, 1986; Stock & Humes, 1995; Stock potential marine invertebrate hosts has been et al., 1962, 1963a, b; Stock, 1968; Varela et al., explored so far (1.14% according to Humes, 2003, 2005b, 2008; Varela, 2010a, 2011a). Most 1994). A high degree of endemism as well as of these studies on Caribbean invertebrate- a remarkable difference in taxonomic com- associated copepods are from the last centu- position of copepods living in a poorly stud- ry and only 19 of them have been published ied symbiosis with Caribbean stony corals is since 2000 (Humes, 2000; Varela et al., 2003, noticed in comparison with the Indo-Pacific 2005a, b, 2007a, b, 2008; Varela & Lalana, 2007; (Stock, 1988). The paucity of knowledge of Kim, 2009, 2010; Hendler & Kim, 2010; Varela, symbiotic copepods of the Greater Caribbe- 2010a, b, 2011a, b, 2012; Ivanenko et al., 2017; an region has repeatedly been noted (Stock, Shelyakin et al., 2018; Garcia-Hernandez et al., 1973, 1975a, 1987, 1988; Humes & Hendler, 1972; 2019). Herriott & Immermann, 1979; Grygier, 1980; Caribbean reef corals are under threat from Ivanenko et al., 2017). climate change and local impacts (Carpen- The first study of symbiotic copepods liv- ter et al., 2008; Hughes et al., 2017) and some ing on cnidarians, echinoderms and sponges symbiotic (including parasitic) copepods po- of the Greater Caribbean was conducted by tentially may have an impact on the state of Edwards (1891), who discovered Diogenidium corals and other invertebrate hosts (Stock, nasutum (fig. 1c), living on the 1975a; Butter, 1979; Herriott & Immermann, Holothuria scabra, in the Bahamas (species 1979; Burke & Maidens, 2004; Ivanenko et al., authorities in tables 1–2). Taxonomic studies 2017; Shelyakin et al., 2018). Despite a long his- of the Caribbean symbiotic copepods were tory of marine biodiversity research in the continued by a number of researchers, who Caribbean, our knowledge of microscopic described large numbers of new species, as symbiotic copepods does not satisfy the needs reviewed by Gotto (1993). A number of cope- for defining priorities in conservation and the pods representing two families (the ectosym- development of management plans (Miloslav- biotic Asterocheridae and the endoparasitic ich et al., 2010; Zeppilli at al., 2015, 2018). The Corallovexiidae) have been found living on goal of our review is to analyze all published and in stony corals (Hoeksema et al., 2017b). data on copepods living in symbiosis with the Very few are reports have been published Caribbean reef-dwelling anthozoans, echino- about the Asterocheridae living and usually derms and spongesDownloaded as important from Brill.com02/21/2020 structural 01:34:50PM­ via Universiteit of Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 299

FIGURE 1 Poecilostomatoid , dorsal view. a – Hemicyclops columnaris (Clausidiidae), b – Corallovexia similis (Corallovexiidae), c – Diogenidium nasutum (Lichomolgidae), d – Macrochiron echinicolum (Macrochironidae), e – Pseudanthessius deficiens (Pseudanthessiidae), f – Doridicola astrophyticus (Rhynchomolgidae), g – Eupolymniphilus occidentalis Kim, 2009 (Sabelliphilidae), h – Meomicola amplectans, i – Presynaptiphilus amphiopli (Synapticolidae). After Stock et al. (1963b), Humes & Ho (1969), Humes & Hendler (1972), Stock (1973, 1975a), Humes & StockDownloaded (1973), Kim from (2009) Brill.com02/21/2020. 01:34:50PM via Universiteit of Groningen

300 KORZHAVINA et al. and functional components of Caribbean coral Material reefs in order to identify the least-studied ecoregions and groups of hosts as well as to The data are combined in the originally identify directions for further research. designed Database on Caribbean copepod crustaceans associated with reef-dwelling cor- als, echinoderms and sponges. This database Characteristics of the Greater Caribbean includes five main tables: Hosts, Symbionts, Samples, Sites, and Publications linked with The Greater Caribbean (The Caribbean s.l.) in each other and two combined tables Litera- the present review consists of the Caribbean ture Records and Sample Records; each record Sea plus the Gulf of Mexico and Bermuda contains data on the taxonomy of the host and (Spalding et al., 2007; Hoeksema et al., 2017a). its symbiont, the references to unique records The Caribbean Sea (Caribbean s.s.) is a large in the World of Copepods (Walter & Boxshall, semi-enclosed sea of the western Atlantic 2019), the number of associates per host, the Ocean with clear and warm water (22–29°C) nature of the association, the name and coor- and low tidal amplitude (0.4 m) (Kinder et al., dinates of the collection site, the depth and 1985). The Caribbean is enclosed by the land the date of collection, as well as their refer- masses of Central and South America (Bra- ence (Korzhavina & Ivanenko, 2019). In addi- zil) from the west and south. It is separated tion, data on samples of symbiotic copepods by island arcs of the Great and Lesser Antilles and their hosts collected at St. Eustatius (in in the north and east (Bayer, 1961; Spald- 2015, with 104 samples), Curaçao (in 2017, with ing et al., 2004; Alvarado, 2011). The Carib- 77 samples) and Cuba (in 2019, with 56 sam- bean is a unique biogeographic region with a ples) preserved in 96% ethanol and including number of endemic species (Rivera-Monroy underwater photographs have been added to et al., 2004; Alvarado, 2011). It is recognized as the table Sample Records. a global marine biodiversity hot spot and an important biogeographic coral reef province (Spalding et al., 2001; Roberts et al., 2002; Milo- Results and discussion slavich et al., 2010; Alvarado, 2011). The Greater Caribbean includes ten marine ecoregions: The database includes 532 records from 154 lo- Northern Gulf of Mexico, Southern Gulf of calities and 54 references published since 1891 Mexico, Floridian, Western Caribbean, Great- (Edwards, 1891). There are 115 species of cope- er Antillean, Southwestern Caribbean, South- pods (47 genera, 17 families, 3 orders) found ern Caribbean, Eastern Caribbean, Bahamian, in symbiosis with 80 invertebrate host species and Bermudian (Burke & Maidens, 2004; representing 58 genera, 39 families, 22 orders Spalding et al., 2007; Hoeksema et al., 2017a). and 7 classes of corals, sponges and echino- The Bahamian and Bermudian are adjacent to derms (figs. 1–7, tables 1–7). Only one species the temperate northwestern Atlantic. The ma- of copepods, the poecilostomatoid cyclopoid rine ecoregions of the Southern and Eastern Hemicyclops columnaris (syn Hemicyclops Caribbean are affected by biota from adjacent geminatus) was found in the area under con- Brazilian waters (Alvarado, 2011). The Gulf of sideration (Bahamas, Barbados, Curaçao, and Mexico has colder and more isolated water, Jamaica, associated with a burrowing ghost which is relatively poor in species (Felder & shrimp, a sponge, an ophiuroid, hermit crabs) Camp, 2009). and on the Pacific coast of Panama (on a stony

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 301 - - But ; Reference Garcia-Hernandez et al., 2017 Humes & Smith, 1974 Kim, 2009 Kim, 2009 1975a Stock, 1975a Stock, 1975a Stock, 1975a Stock, 1979 ter, 1975a Stock, & Immer Herriott mann, 1979 1975a Stock, 1975a Stock, 1975a Stock,

Site Site abbreviation** PR Pan Bah Bah Cur Cur Cur Cur Cur VI Cur Cur Cur

Host abbreviation* Po Act Op Po Sc Sc Sc Sc Sc Sc Sc Sc Sc

Diploria Diploria

(Linnaeus, 1758) (Linnaeus, (Verrill, 1901)) (Verrill, (Linnaeus, 1767) (as 1767) (Linnaeus, 1767) (Linnaeus, (Dana, 1846) (as 1846) (Dana, (Linnaeus, 1758) (Linnaeus, (Le Sueur, 1817) (Le Sueur, Ehrenberg, 1834 Ehrenberg, 1834 Ehrenberg, Lütken, 1856) Lütken, Milne Edwards & Haime, Milne Edwards Müller & Troschel, 1842 1842 Troschel, Müller & (Pallas, 1766) (Pallas, Azevedo et al., 2017 Azevedo (Dana, 1846)) (Dana, Ophiocoma riisei Ophiocoma Host species: valid name Host species: valid (and as in original record) lutea Clathrina annulata Bartholomea wendtii Ophiocoma (as Porifera cylindrus Dendrogyra fastigiata Eusmilia cylindrus Dendrogyra meandrites Meandrina stokesii Dichocoenia 1848 strigosa Pseudodiploria strigosa labyrinthiformis Diploria cavernosa Montastraea brasiliana Montastraea cavernosa Montastraea Stock, 1975 Stock, Stock, 1975 Stock, 1975 Stock, Stock, 1975 Stock, Humes, 1984 Humes, 1984 Humes, Stock, 1975 Stock, 1975 Stock,

Humes & Smith, 1974

Stock, 1975 Stock, 1975 Stock, sp. Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded

TABLE 1 Copepod Copepoda Calanoida Pseudocyclopidae fosshageni Ridgewayia Cyclopoida Clausidiidae columnaris Hemicyclops columnaris Hemicyclops Corallovexiidae baki Corallonoxia baki Corallonoxia longicauda Corallonoxia longicauda Corallonoxia Corallonoxia brevibrachium Corallovexia brevibrachium Corallovexia dorsospinosa minor Corallovexia dorsospinosa Corallovexia

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302 KORZHAVINA et al. ------Herri Herri Herri Herri ; ; ; ; Reference 1975a Stock, 1975a Stock, 1975a Stock, 1975a Stock, & Immer Herriott mann, 1979 1975a Stock, 1975a Stock, & Immermann, ott 1979 1975a Stock, & Immermann, ott 1979 1975a Stock, & Immermann, ott 1979 1975a Stock, & Immermann, ott 1979 & Immer Herriott mann, 1979 & Immer Herriott mann, 1979 & Immer Herriott mann, 1979

) Site Site abbreviation** Cur Cur Cur Cur VI Cur VI Cur, VI Cur, VI Cur, VI Cur, VI VI VI cont.

Host abbreviation* Sc Sc Sc Sc Sc Sc Sc Sc Sc Sc Sc Sc Sc

Diploria Diploria Manicina

(Dana, 1846) (Dana, (as 1846) (Dana, 1846) (Dana, (Linnaeus, 1758) (Linnaeus, (Ellis & Solander, 1786) (as 1786) (Ellis & Solander, (Houttuyn, 1772) (as 1772) (Houttuyn, (Müller)) 1772) (Houttuyn, 1772) (Houttuyn, 1772) (Houttuyn, (Linnaeus, 1758) (as 1758) (Linnaeus, 1758) (Linnaeus, (Lamarck, 1816) (Lamarck, 1816) (Lamarck, Wells, 1936) Wells, (Ellis & Solander, 1786)) (Ellis & Solander, mayori f. (Dana, 1846)) (Dana, Host species: valid name Host species: valid (and as in original record) natans Colpophyllia natans Colpophyllia strigosa Pseudodiploria natans Colpophyllia Manicina areolata areolata natans Colpophyllia Manicina areolata clivosa Pseudodiploria clivosa Diploria strigosa Pseudodiploria strigosa natans Colpophyllia palmata Acropora strigosa Pseudodiploria palmata Acropora meandrites Meandrina Stock, 1975 Stock, 1975 Stock, 1975 Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1975 Stock, 1975 Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1975 Stock, 1975 Stock, sp. sp. Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges ( echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded

TABLE 1 Copepod kristenseni Corallovexia longibrachium Corallovexia longibrachium Corallovexia longibrachium Corallovexia mediobrachium Corallovexia mediobrachium Corallovexia mediobrachium Corallovexia mediobrachium Corallovexia mixtibrachium Corallovexia similis Corallovexia similis Corallovexia Corallovexia Corallovexia

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 303 - - - Herriott & Immer Herriott mann, 1979 & Immer Herriott mann, 1979 1975a Stock, & Immer Herriott mann, 1979 1975a Stock, et al., 2005b Varela 1973 Stock, 1973 Stock, 1973 Stock, 1973 Stock, 1973 Stock, 1973 Stock, 1979 Stock, 1978 Stock, 1980 Grygier, et al., Ivanenko 2017 1970 Humes & Ho, 1970 Humes & Ho, VI VI Cur VI Cur Cub PR PR PR PR Cur Cur Flo Flo Bah SE Bar Bar Sc Sc Sc Sc Sc Oc Oc Oc Oc Oc Oc Oc Oc Oc Oc Oc Hol Hol -

Pseudop Plexaura Plexaura Semper, 1868 Semper, 1868 Semper, Pseudopterogorgia (Semper)) Milne Edwards & Milne Edwards Esper, 1794) Esper, (Linnaeus, 1767) (Linnaeus, 1767) (Linnaeus, (Dana, 1846) (Dana, (Ellis & Solander, 1786)) (Ellis & Solander, (Pallas, 1766) (Pallas, (Pallas, 1766) (as 1766) (Pallas, (Esper, 1794) (as 1794) (Esper, 1794) (Esper, (Ellis & Solander, 1786) (as 1786) (Ellis & Solander, Linnaeus, 1758 Linnaeus, Lamouroux, 1816 Lamouroux, (Duchassaing & Michelotti, (Duchassaing & Michelotti, (Pallas, 1766)) (Pallas, homomalla Bayer, 1951 (as 1951 Bayer, sp. sp. f. sp. Brandtothuria arenicola Brandtothuria Montastraea cavernosa Montastraea lamarckiana Mycetophyllia Haime, 1849 annularis Orbicella annularis Montastraea strigosa Pseudodiploria cavernosa Montastraea nutans Plexaurella 1860) mammosa Eunicea homomalla Plexaura homomalla homomalla Plexaura asbestinum Briareum Antillogorgia 1919) Kükenthal, acerosa Antillogorgia acerosa terogorgia Placogorgia Thesea Callogorgia ventalina Gorgonia arenicola (Thymiosycia) Holothuria arenicola (Thymiosycia) Holothuria (as Stock, 1975 Stock, Stock, 1978 Stock, Humes & Ho, 1970 Humes & Ho,

Stock, 1973 Stock,

Stock, 1973 Stock,

Stock, 1973 Stock,

Grygier, 1980 Grygier, Stock, 1973 Stock,

Stock, 1979 Stock,

Stock, 1973 Stock, 1973 Stock,

Humes & Ho, 1970 Humes & Ho,

sp. sp. sp. sp. sp. sp. Corallovexia Corallovexia Corallovexia Corallovexia ventrospinosa Corallovexia Lamippidae Enalcyonium euniceae Enalcyonium nudum Enalcyonium ramosum Enalcyonium variicauda Enalcyonium Lamippina aequalis Lamippina aequalis bouligandi Linaresia Magnippe caputmedusae caligicola Sphaerippe Sphaerippe Lichomolgidae deichmannae Diogenella impar Diogenella

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304 KORZHAVINA et al. - - Humes Humes Hen Hen ; ; ; ; Reference 1970 Humes & Ho, 1968 Stock, 1968 Stock, 1970 & Ho, 1968 Stock, 1970 & Ho, 1968 Stock, Hendler & Kim, 2010 Hendler & Kim, 2010 Hendler & Kim, 2010 Hendler & Kim, 2010 1968 Stock, dler & Kim, 2010 1891 Edwards, 1968 Stock, dler & Kim, 2010 Hendler & Kim, 2010 Humes & Stock, 1973

) Site Site abbreviation** MI PR MI, PR Jam, Bah, Cur, MI, PR PR Bah, PR Bar Bah Jam MI, PR Jam, Cur, Bah PR Jam, Bah, Jam Cur cont.

Host abbreviation* Hol Hol Hol Hol Hol Hol Hol Hol Hol Hol Hol Hol Hol Ec - Lud Muelleria Muelleria Selenka, (Ludwig)) (Forsskål, (Forsskål, Semper, 1868 Semper, 1868 Semper, Ludwig, 1875 Ludwig, 1875 Ludwig, 1875 Ludwig, 1875 Jaeger, 1833 Jaeger, Selenka, 1867 (Forskål, 1775)) (Forskål, (Ludwig))

(Selenka, 1867) (Selenka, 1867) (as (Selenka, 1867) (Selenka, 1867) A. Agassiz, 1863 A. Agassiz, Halodeima surinamensis Holothuria impatiens Holothuria Selenka, 1867) Ludwigothuria mexicana Ludwigothuria Host species: valid name Host species: valid (and as in original record) arenicola (Thymiosycia) Holothuria impatiens (Thymiosycia) Holothuria (as 1775) surinamensis (Semperothuria) Holothuria (as wig, 1875 mexicana (Halodeima) Holothuria (as glaberrima (Selenkothuria) Holothuria 1867 mexicana (Halodeima) Holothuria arenicola (Thymiosycia) Holothuria agassizii Actinopyga agassizi grisea (Halodeima) (Halodeima) Holothuria scabra (Metriatyla) Holothuria badionotus agassizii Actinopyga viridis Echinometra Humes & Stock, 1973 Humes & Stock, Stock, 1968 Stock,

Edwards, 1891 Edwards, 1891 Edwards, 1891 Edwards, 1891 Edwards, Stock, 1968 Stock,

Stock, 1968 Stock, 1968 Stock, 1968 Stock,

Stock, 1968 Stock, 1968 Stock, 1968 Stock, Humes & Ho, 1971 Humes & Ho,

Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges ( echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded

TABLE 1 Copepod seticauda Diogenella seticauda Diogenella seticauda Diogenella spinicauda Diogenella deforme Diogenidium deforme Diogenidium deforme Diogenidium nasutum Diogenidium nasutum Diogenidium nasutum Diogenidium nasutum Diogenidium spinulosum Diogenidium tectum Diogenidium Macrochironidae echinicolum Macrochiron

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 305 ; Stock, 1975c Stock, ; Humes & Stock, Humes & Stock, 1973 Humes & Stock, 1973 Kim, 2009 et al., 1963b Stock Humes & Hendler, 1999 et al., 1963b Stock Humes & Hendler, 1999 & Humes, Stock 1995 et al., 1963b Stock 1975c Stock, 1975c Stock, 2011a Varela, 1975c Stock, 1973 Humes, et al., 2003 Varela 1973 Humes, Humes & Stock; 1973 Humes & Stock, 1973 Bah, Bar, Bon, Bah, Bar, MI, PR Jam, Cur, SM Jam Cur Bel MI, SM Bel, Cur, Cur MI Bah, Jam, Cur Cur Cub Cur Ber Cub Ber Jam Cur, Bar, PR Ec Oc Po Hol Op Op Ec Ec Oc Oc Oc Oc Oc Oc Oc Oc Oc - Ophioder Ludwig, 1875 (Say, 1825) (as 1825) (Say, (Houttuyn, 1772) (Houttuyn, Müller & Troschel, 1842) Troschel, Müller & (Lamarck, 1816) (Lamarck, (Pallas, 1766) (Pallas, Bieschowsky (Esper, 1794) (Esper, Müller & Troschel, 1842 (as 1842 Troschel, Müller & (Linnaeus, 1758) (Linnaeus, Linnaeus, 1758 Linnaeus, (Lamarck, 1816) (Lamarck, (Pallas, 1766) (Pallas, (Lamouroux, 1821) (Lamouroux, (Say, 1825)) (Say, sp. Verrill, 1864 Verrill, sp. Lytechinus variegatus Lytechinus reniformis Renilla Porifera mexicana (Halodeima) Holothuria Ophioderma brevispina ma brevispinum Ophioderma cinerea Ophioderma cinereum ventricosa Meoma rosaceus Clypeaster laxa Muricea flexuosa Eunicea Plexaura homomalla Plexaura ventalina Gorgonia Pseudoplexaura porosa Pseudoplexaura acerosa Antillogorgia elastica Antillogorgia - - - Stock & Humes, 1995 & Humes, Stock

Stock, Humes & Stock, Humes & Stock, 1973 Humes & Stock, Stock, 1975 Stock, Stock, Humes & Good Stock, Humes & Good Stock, Humes & Good Stock, Humes & Stock, 1973 Humes & Stock, 1973 Humes & Stock,

Kim, 2009

Humes, 1973 Humes, 1973 Humes, 1973 Humes,

Stock, 1975 Stock, 1975 Stock, 1975 Stock, Sars, 1916 Sars, Macrochiron echinicolum Macrochiron sargassi Macrochiron Pseudanthessiidae acutus Pseudanthessius deficiens Pseudanthessius ing, 1963 deficiens Pseudanthessius ing, 1963 deficiens Pseudanthessius ing, 1963 exilicornis Pseudanthessius pectinifer Pseudanthessius Gooding, 1964 Rhynchomolgidae aequiseta Acanthomolgus affinis Acanthomolgus affinis Acanthomolgus affinis Acanthomolgus bayeri Acanthomolgus bayeri Acanthomolgus bayeri Acanthomolgus bilobipes Acanthomolgus bilobipes Acanthomolgus

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306 KORZHAVINA et al. ; Ortiz ; ; ; Reference 1975c Stock, 1973 Humes, 1975c Stock, 1975c Stock, et al., 2003 Varela 1975c Stock, 1975c Stock, 1975c Stock, 1973 Humes, 1975c Stock, 1975c Stock, 1975c Stock, et al., 2008 Varela 1973 Humes, 1975b Stock, 1969a Humes, 1975b Stock, et al., 1998 Goenaga, & Humes 1978 Goenaga, & Humes 1978 et al., 2008 Varela

) Site Site abbreviation** Cur SE Cur, Ber, Cur Cub Cur Cur Cur Ber SM SE, SM Cur, Cub Ber PR, SM Barad, Bon, Bah, Bar, FK, Cur, Cub, MI, PR Jam, PR PR Cub cont.

Host abbreviation* Oc Oc Oc Oc Oc Oc Oc Oc Oc Oc Oc Oc Act Act Anti Anti Op - -

Ho (Ellis, (Ellis, Stoichac (Gmelin, 1791)) (Le Sueur, 1817)) (Le Sueur,

(Ellis, 1768) (as 1768) (Ellis, (Gmelin, 1791) (as (Gmelin, 1791) (Lamarck, 1816) (Lamarck, (Le Sueur, 1817) (as 1817) (Le Sueur, (Esper, 1791) (Esper, Brook, 1889 Brook, Stoichactis helianthus Stoichactis Milne Edwards & Haime, 1857 Milne Edwards Linnaeus, 1758 Linnaeus, 1758 Linnaeus, (Ellis & Solander, 1786) (Ellis & Solander, Bayer, 1961 Bayer, (Kükenthal, 1911) (Kükenthal,

Bayer, 1961 Bayer, Duchassaing & Michelotti, 1860 Duchassaing & Michelotti,

(Lamouroux, 1821) (Lamouroux, sp. (Ellis), sp. Host species: valid name Host species: valid (and as in original record) americana Antillogorgia americana Pseudopterogorgia ventalina Gorgonia laciniata Eunicea Muricea flexuosa Eunicea tourneforti Eunicea clavigera Eunicea atlantica Muricea dichotoma Plexaurella ventalina Gorgonia mariae Gorgonia calyculata Eunicea Corynactis denticulosa mostichanthus denticulosus helianthus Stichodactyla tis anemone 1768)) Stichopathes Stichopathes lutkeni muricatum Astrophyton Stock, 1975 Stock, Humes, 1973 Humes, Stock, 1975 Stock, 1975 Stock, Stock, 1975 Stock, 1975 Stock,

Humes, 1969 Humes, 1969 Humes, Humes, 1973 Humes, Humes, 1973 Humes,

Stock, 1975 Stock,

Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1975 Stock, Humes & Goenaga, 1978 Humes & Goenaga, 1978 Humes & Goenaga,

Humes & Stock, 1983 Humes & Stock, Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges ( echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded

TABLE 1 Copepod dionyx Acanthomolgus gorgoniae Acanthomolgus intermedius Acanthomolgus intermedius Acanthomolgus longidactylus Acanthomolgus longifurca Acanthomolgus mononyx Acanthomolgus muriceanus Acanthomolgus seticornis Acanthomolgus triangulipes Acanthomolgus triangulipes Acanthomolgus verrucipes Acanthomolgus Aspidomolgus stoichactinus Aspidomolgus stoichactinus imparipes Calonastes imparipes Calonastes Critomolgus

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 307 ; Humes & Stock, Humes & Stock, 1973 1982 Humes, 1975c Stock, Kim, 2009 1969b Humes, 1969b Humes, 1969b Humes, 1969b Humes, Humes 1969b et al., 2003 Varela et al., 1963b Stock Humes & Hendler, 1972 Hendler & Kim, 2010 Hendler & Kim, 2010 Ivanenko, unpublished Bar, Jam, MI, PR Jam, Bar, PR PR Jam Jam Bah, Cur, Jam Bar, Bah, Bon, Cur, Jam Jam, Bah, Bar, MI, PR Bah, Bon, Cub, Jam Cur, Jam Cur, FK Bel Bel SE Op Act Cor Po Hol Hol Hol Hol Hol Ec Op Op Op Op (Selenka, 1867) (Selenka, 1867) (Lamarck, 1816) (Lamarck, Ludwig, 1875 (Weinland, 1860) (Weinland, (Say, 1825) (Say, (Selenka, 1867) (Selenka, 1867) Lütken, 1856 Lütken, (Lamarck, 1816) (Lamarck, Duchassaing & Michelotti, 1860 Duchassaing & Michelotti, sp. Astrophyton muricatum Astrophyton gigantea Condylactis florida Ricordea Porifera agassizii Actinopyga Isostichopus badionotus agassizii Actinopyga Isostichopus badionotus mexicana Holothuria ventricosa Meoma Amphioplus angulata Ophiothrix oerstedii Ophiothrix Ophiuroida ) Doridicola Doridicola

) (as Kim, 2009

Humes & Stock, Humes & Stock,

Humes & Hendler, Humes & Hendler, Humes & Stock, 1973 Humes & Stock,

( Stock, 1975 Stock, Suárez-Morales in Suárez-Morales in Suárez-Morales Stock, Humes & Gooding, Stock, Humes, 1969) Humes, Humes, 1982 Humes, ) ( (Humes, 1969) (as 1969) (Humes, 1969) (Humes, 1969) (Humes, sp.A sp sp Humes, 1982 Humes,

) Doridicola astrophyticus Doridicola Critomolgus astrophyticus Critomolgus (as 1973 titillans Critomolgus titillans antillianus Paramolgus Sabelliphilidae occidentalis Eupolymniphilus Synapticolidae Caribulus Caribulus sculptus Caribulus sculptus Scambicornus sculptus Caribulus sculptus Caribulus amplectans Meomicola 1963 Synaptiphilidae amphiopli Presynaptiphilus 1972 Thaumatopsyllidae chawayi Caribeopsyllus 1998 & Castellanos, Suárez-Morales chawayi Caribeopsyllus 1998 & Castellanos, Suárez-Morales Caribeopsyllus

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308 KORZHAVINA et al. ; Reference Hendler & Kim, 2010 et al., 2005b Varela 2010 Varela, Kim, 2010 Kim, 2010 2000 Humes, 2010 Varela, 2000 Humes, 2010b Varela, et al., 2007a Varela 2010b Varela, et al., 2007 Varela 2012 Varela, 2012 Varela, 1987 Stock, 1989 1987, Stock, Kim, 2010 Kim, 2010 Kim, 2010 et al., 2008 Varela & Huys, Boxshall 1994

) Site Site abbreviation** Bel Cub Cub Bah Jam Bel, Cub Bel Cub Cub Cub Cub Cub Cub Cur Cur Jam Bah Bah Cub Bel cont.

Host abbreviation* Op Po Po Po Cr Cr Cr Po Po Po Po Po Po Sc Sc Po Po Po Po Po

(Pourtalès, 1869) (Pourtalès, (Lamarck, 1814) (Lamarck, 1814) (Lamarck, Alcolado, 1984 Alcolado, (Say, 1825) (Say, (Duchassaing & Michelotti, (Duchassaing & Michelotti, (Ellis & Solander, 1786) (Ellis & Solander, (Linnaeus, 1758) (Linnaeus, AH Clark, 1909 (Schmidt, 1870) (Lamarck, 1814) (Lamarck, 1814) (Lamarck, Duchassaing & Michelotti, 1864 Duchassaing & Michelotti, sp. sp. Host species: valid name Host species: valid (and as in original record) angulata Ophiothrix vaginalis Callyspongia Callyspongia Porifera Crinoidea rubiginosus Davidaster grandis Nemaster dilatata Agelas vaginalis Callyspongia wiedenmayeri Agelas walpersii Ptilocaulis 1864) digitalis Niphates digitalis Niphates Manicina areolata annularis Orbicella Porifera Porifera Porifera Agelas clathrodes Agelas

Varela, 2010 Varela, Kim, 2010 Humes, 2000 Humes, 2000 Humes, 2000 Humes, Kim, 2010

Stock, 1987 Stock, 1987 Stock,

Varela, 2010 Varela, Kim, 2010 Kim, 2010

Varela, Ortiz & Lalana, Varela, Varela, 2010 Varela, Varela, Ortiz & Lalana, Varela, Boxshall & Huys, 1994 & Huys, Boxshall 1994 & Huys, Boxshall

Varela, 2013 Varela, Varela, 2012 Varela,

sp.A sp. Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges ( echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded

TABLE 1 Copepod Caribeopsyllus Asterocheridae Asterocheres antillensis Asterocheres bahamensis Asterocheres crinoidicola Asterocheres crinoidicola Asterocheres crinoidicola Asterocheres cubensis Asterocheres espinosai Asterocheres 2007 fernandezmilerai Asterocheres garridoi Asterocheres 2007 kimi Asterocheres lalanai Asterocheres maxillatus Asterocheres maxillatus Asterocheres oricurvus Asterocheres peniculatus Asterocheres plumosus Asterocheres reginae Asterocheres reginae Asterocheres

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 309 Kim, ; ; Varela, 2011a Varela, ; Stock & Humes, & Humes, Stock 1995 Kim, 2010 Kim, 2010 Kim, 2010 1989 Stock, Kim, 2010 1989 Stock, 1975b Stock, et al., 2005 Varela 1975b Stock, 1982 Humes, 1987 Stock, 1975b Stock, 1975b Stock, 2010 1987 Stock, & Humes, Stock 1995 & Humes, Stock 1995 & Humes, Stock 1995 1998 Humes, 1998 Humes, 1987 Stock, 1989 Stock, Cur Bah Bah, PR Jam Cur Bar Cur PR Cub Jam, Bah, Cur, MI, PR Cur MI PR Jam, Cur Cur, Bah, Cub, PR, SM Jam, Cur PR Bar Bar Cur Cur Ec Po Ec Ec Sc Sc Sc Cor Sc Act Sc Act Act Sc Ec Ec Ec Op Op Sc Sc Lütken, Lütken, (Say, 1825) (Say, Milne Edwards & Milne Edwards (Ellis, 1768) (Ellis, (Linnaeus, 1767) (Linnaeus, (Lamarck, 1816) (Lamarck, (Linnaeus, 1758) (Linnaeus, (Ellis & Solander, 1786) (Ellis & Solander, (Linnaeus, 1758) (Linnaeus, 1758) (Linnaeus, (Lamarck, 1816) (Lamarck, (Weinland, 1860) (Weinland, (Le Sueur, 1817) (Le Sueur, (Lamarck, 1816) (Lamarck, (Müller)) (Ellis & Solander, 1786) (Ellis & Solander, Duchassaing & Michelotti, 1860 Duchassaing & Michelotti, (Esper, 1795) (Esper, sp. Lytechinus variegatus Lytechinus Porifera tribuloides Eucidaris Echinoidea annularis Orbicella Meandrina meandrites Meandrina florida Ricordea lamarckiana Mycetophyllia Haime, 1849 gigantea Condylactis natans Colpophyllia crucifer Phymanthus helianthus Stichodactyla fragum Favia lucunter Echinometra ventricosus Tripneustes lucunter Echinometra angulata (Ophiothrix) Ophiothrix suensoni (Acanthophiothrix) Ophiothrix 1856 clivosa Pseudodiploria cavernosa Montastraea - Asco Varela, Ortiz & Varela, Stock, 1987 Stock,

Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1975 Stock, 1975 Stock, Stock & Humes, 1995 & Humes, Stock 1995 & Humes, Stock Kim, 2010 Stock, 1987 Stock,

Stock, 1987 Stock, Kim, 2010 Kim, 2010 Kim, 2010 Stock, 1987 Stock,

(T. Scott, 1898) (as Scott, (T. Stock, 1989 Stock,

Stock, 1989 Stock, Humes, 1998 Humes, 1998 Humes,

Stock & Humes, 1995 & Humes, Stock

T. Scott, 1898) Scott, T. Asterocheres simulans Asterocheres simulans myzon trisetatus Asterocheres unioviger Asterocheres unioviger Asterocheres foliata Asteropontella capillatus Asteropontius iuxtus Asteropontius longipalpus Asteropontius mycetophylliae Asteropontius Lalana, 2005 parvipalpus Asteropontius proximus Asteropontius ungellatus Asteropontius ungellatus Asteropontius faviae Asteropontopsis longipalpus Chelacheres longipalpus Chelacheres optans Chelacheres lunulifer Collocheres lunulifer Collocheres diploriae Hermacheres montastreae Hermacheres

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310 KORZHAVINA et al. Varela et al., Varela ; Reference Kim, 2010 Kim, 2010 Kim, 2010 Kim, 2010 1989 Stock, 2012 Varela, & Gooding, Stock 1986 2005b 2011b Varela, & Lalana, Varela 2007 1998 Humes, 1975d Stock, et al., 2008 Varela 2012 Varela, Kim, 2010 Humes & Hendler, 1999 Humes & Hendler, 1999 Humes & Hendler, 1999

) Site Site abbreviation** Bah Jam Jam Bar Cur Cub Cur Cub, Cub Cub MI Cur Cub Cub PR Bel Bel, FK Bel cont.

Host abbreviation* Po Po Po Po Sc Po Ec Oc Oc Op Sc Po Po Po Op Op Op (Clark, 1900) (Clark, 1900)

(Linnaeus, 1767) (Linnaeus, (Linnaeus, 1758) (Linnaeus, (Lamarck, 1814) (Lamarck, 1814) (Lamarck, (Pallas, 1766) (Pallas, (Higgin, 1877) Philippi, 1845 (Say, 1825) (Say, Lütken, 1856 Lütken, Lamouroux, 1816 Lamouroux, Host species: valid name Host species: valid (and as in original record) Porifera Porifera Porifera Porifera meandrites Meandrina vaginalis Callyspongia Diadema antillarum mammosa Eunicea asbestinum Briareum flaccida Ophiomyxa cavernosa Montastraea birotulata Iotrochota vaginalis Callyspongia Porifera ophiactoides Ophiocomella pumila Ophiocoma ophiactoides Ophiocomella ) Asterocheres Asterocheres ) (as Stock, 1989 Stock,

Varela, 2012 Varela, Kim, 2010 Kim, 2010 Kim, 2010

Kim, 2010

(

Humes, 1998 Humes,

(Scott T. & Scott A., 1894) & Scott T. (Scott Kim, 2010 (Johnsson, 1998) Stock, 1975 Stock, Stock & Gooding, 1986 Stock ( Varela, 2011 Varela,

) Humes & Hendler, 1999 Humes & Hendler, 1999 Humes & Hendler, 1999 Humes & Hendler,

Varela & Lalana, 2007 Varela

Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges ( echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded Kim, 2010

TABLE 1 Copepod aculeata Hetairosynella angulata Hetairosynella bifurcata Hetairosynella fastigatus Kimcheres fastigatus coronatum Meandromyzon humesi Neoasterocheres alatus Onychocheres antillensis Orecturus ortizi Orecturus bellulus Ophiurocheres rostratum Peltomyzon elongatus Scottocheres paraboecki Setacheres edentatum Stenomyzon Cancerillidae latus Ophiopsyllus latus Ophiopsyllus latus Ophiopsyllus

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 311 ; ; - - Va ; Stock et al., 1963a Stock Humes & Hendler, 1999 Emson et al., 1985 Emson & Mlad 1987 enov, Humes & Hendler, 1972 1992b Stock, 1992b Stock, 1992b Stock, et al., 2008 rela et al., 1963a Stock et al., 1962 Stock et al., 1962 Stock et al., 1962 Stock et al., 1962 Stock et al., 1962 Stock et al., 1962 Stock et al., 1962 Stock Bel, Cur Jam Bel, Ber, FK PR Cur Cur, Aru, Cub, PR Jam Jam MI, PR Jam MI PR Bah, Jam Jam Op Op Op Po Po Po Ec Hol Hol Hol Hol Hol Hol Ec - as ( Lud ) Semper, 1868 Semper, 1868 Semper, Holothuria Holothuria mixed Ludwig, 1875 Ludwig, 1875 and (Clark, 1900) (Selenka, 1867) (Selenka, 1867) (Selenka, 1867) (Lamarck, 1816) (Lamarck, (Lamarck, 1816) (Lamarck, 1816) (Lamarck, Schmidt, 1862 Schmidt, 1862 Schmidt, 1862 sp. Brandtothuria arenicola) Brandtothuria as Ophiocoma echinata Ophiocoma ophiactoides Ophiocomella Amphioplus nucula Chondrilla nucula Chondrilla nucula Chondrilla ventricosa Meoma Isostichopus badionotus Isostichopus badionotus arenicola (Thymiosycia) Holothuria surinamensis (Semperothuria) Holothuria wig, 1875 arenicola (Thymiosycia) Holothuria ( surinamensis (Semperothuria) ( surinamensis) Semperothuria Isostichopus badionotus ventricosa Meoma - Par Humes & Hendler, 1972 Humes & Hendler, Stock, 1992 Stock,

Wilson C.B., 1944 Wilson C.B., (as 1944 Wilson C.B., Stock, Humes & Gooding, Stock, Humes & Gooding, Stock, Stock, Humes & Gooding, Stock, Wilson C.B., 1944) Wilson C.B., Stock, Humes & Gooding, 1962 Stock, Humes & Gooding, 1962 Stock, Humes & Gooding, 1962 Stock, Humes & Gooding, 1962 Stock, Humes & Gooding, 1962 Stock, Stock, Humes & Gooding, 1962 Stock, Stock, Humes & Gooding, 1962 Stock, Ophiopsyllus reductus Ophiopsyllus 1963 reductus Ophiopsyllus 1963 ligatus Parophiopsyllus Entomolepididae verrucosa Parmulodes emarginata Parmulella verrucosa Parmulodes mulodes verrucosus Micropontiidae glaber Micropontius 1963 Nanaspididae exigua Nanaspis media Nanaspis pollens Nanaspis pollens Nanaspis pollens Nanaspis pollens Nanaspis pollens Nanaspis

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312 KORZHAVINA et al. - Reference et al., 1962 Stock et al., 1962 Stock

) Site Site abbreviation** MI, PR MI cont.

Host abbreviation* Hol Hol Semper, 1868 Semper, (Selenka, 1867)

Host species: valid name Host species: valid (and as in original record) parvula (Platyperona) Holothuria arenicola (Thymiosycia) Holothuria Stock, Humes & Gooding, Stock, Humes & Gooding, Stock, Copepod crustaceans recorded as associated with Caribbean reef-dwelling anthozoans, echinoderms and sponges ( echinoderms and sponges anthozoans, with Caribbean reef-dwelling as associated Copepod crustaceans recorded

TABLE 1 Copepod truncata Nanaspis 1962 truncata Nanaspis 1962 * Host Abbreviations: Act – Actiniaria, Anti – Antipatharia, Cor – Corallimorpharia, Cr – Crinoidea, Ec – Echinoidea, Hol – Holothuroidea, Oc – Octocorallia, Op – Ophiuroi Oc – Octocorallia, Cr – Crinoidea, Ec Echinoidea, Hol Holothuroidea, Anti – Antipatharia, Cor Corallimorpharia, – Actiniaria, Act * Host Abbreviations: Sc – Scleractinia. – Porifera, dea, Po Flo – Florida, Cub – Cuba, Cur Curacao, Ber – Bermuda, Bon Bonaire, Bel – Belize, – Baradal, Barad Bar – Barbados, Aru – Aruba, Bah Bahamas, Abbreviations: ** Site Virgin Islands. VI – SM – St. Martin, SE – St. Eustatius, Rico, PR – Puerto – Panama, Pan Key, Island, LK – Loggerhead MI – Magueyes – Jamaica, Jam Downloaded from Brill.com02/21/2020 01:34:50PM via Universiteit of Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 313

FIGURE 2 Cyclopoida. Lamippidae (a–d): a – Enalcyonium nudum, dorsal view; b – Linaresia bouligandi, dorsal view; c – Magnippe caputmedusae, ventral view; d – Sphaerippe caligicola, lateral view. Thaumatopsyl- lidae (e): nauplius (N1), metanauplius (MN), copepodid stages 1–5 (CI–CV)Downloaded of from Caribeopsyllus Brill.com02/21/2020 amphio 01:34:50PM- diae, dorsal view. After Stock (1973, 1978, 1979), Grygier (1980), Dojiri et al. (2008). via Universiteit of Groningen

314 KORZHAVINA et al. Stock, Stock, ; Humes, Humes, Kim, ; ; Ortiz et al., ; Reference 1975b Stock, 1982 1982 Humes, 1975b Stock, 1975b Stock, 1969a Humes, 1975b 1998 1975b Stock, 2010 Humes & Smith, 1974 Humes & Goenaga, 1978 Humes & Goenaga, 1978

Site abbreviation** MI, PR Jam, Bah, Cur, PR PR, SM Barad, MI Bon, Cub, Bah, Bar, MI, PR FK, Jam, Cur, PR Jam, Pan PR PR

Copepod family abbreviation* Ast Rhy Rhy Ast Rhy Ast Psec Rhy Rhy

) )

Humes, Humes, Humes, Stock, Stock, Stock, 1975 Stock, 1975 Stock, Humes &

Humes, 1982 Humes,

Humes & Humes & Humes, 1982 Humes,

( Doridicola titillans Doridicola Copepod species name parvipalpus Asteropontius 1975 titillans Critomolgus (as Aspidomolgus stoichactinus 1969 ungellatus Asteropontius Aspidomolgus stoichactinus 1969 ungellatus Asteropontius fosshageni Ridgewayia Smith, 1974 imparipes Calonastes 1978 Goenaga, imparipes Calonastes 1978 Goenaga,

and as name (Ellis, 1768) (as (Ellis, 1768) (Ellis, (Le Sueur, 1817) (Le Sueur, (Le Sueur, 1817) (Le Sueur, (Ellis, 1768)) (Ellis, (Weinland, 1860) (Weinland, 1860) (Weinland, (Le Sueur, 1817) (Le Sueur, (Ellis, 1768) and (Ellis, Brook, 1889 Brook, sp. Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans Caribbean reef-dwelling

Homostichanthus denticulosus TABLE 2 Host (valid Invertebrate in the original record) indicated Cnidaria Anthozoa Hexacorallia Actiniaria gigantea Condylactis gigantea Condylactis Corynactis denticulosa (as 1817)) (Le Sueur, crucifer Phymanthus helianthus Stichodactyla anemone Stoichactis helianthus Stoichactis helianthus Stichodactyla Antipatharia annulata Bartholomea Stichopathes Stichopathes lutkeni

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 315 - - Herriott Herriott Herriott ; ; Stock, 1975b Stock, 1975b Stock, & Immer Herriott mann, 1979 1975a Stock, & Immermann, 1979 1987 Stock, 1975a Stock, 1975a Stock, & Immer Herriott mann, 1979 1975a Stock, & Immermann, 1979 1975a Stock, 1975a Stock, 1975a Stock, 1975a Stock, 1975a Stock, 1987 Stock, 1987 Stock, PR PR VI VI Cur, Cur Cur Cur VI VI Cur, Cur Cur Cur Cur Cur Cur Cur Ast Rhy Cor Cor Ast Cor Cor Cor Cor Cor Cor Cor Cor Cor Ast Ast

Stock, Stock, Stock, Stock, Stock, Stock, Stock, Stock, Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1987 Stock, Stock, 1987 Stock,

Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1987 Stock,

Stock, 1975 Stock, Stock, 1975 Stock, 1975 Stock, sp. sp. Corallovexia mediobrahium Corallovexia Asteropontius longipalpus Asteropontius antillianus Paramolgus Corallovexia similis Corallovexia proximus Asteropontius kristenseni Corallovexia longibrachium Corallovexia 1975 mediobrachium Corallovexia 1975 (as 1975) Stock, mixtibrachium Corallovexia 1975 baki Corallonoxia longicauda Corallonoxia Corallonoxia brevibrachium Corallovexia 1975 baki Corallonoxia faviae Asteropontopsis maxillatus Asterocheres

Wells, 1936) Wells, (Linnaeus, 1758) (Linnaeus, (Muller)) . mayori f Ehrenberg, 1834 Ehrenberg, 1834 Ehrenberg, Milne Edwards & Milne Edwards (Houttuyn, 1772) (Houttuyn, 1772) (Houttuyn, 1772) (Houttuyn, 1772) (Houttuyn, 1772) (Houttuyn, (Pallas, 1766) (Pallas, (Linnaeus, 1758) (Linnaeus, (Lamarck, 1816) (Lamarck, 1816) (Lamarck, Duchassaing & Duchassaing & (Esper, 1795) (Esper, Colpophyllia natans Colpophyllia Manicina areolata Corallimorpharia florida Ricordea 1860 Michelotti, florida Ricordea 1860 Michelotti, Scleractinia palmata Acropora palmata Acropora natans Colpophyllia (as natans Colpophyllia natans Colpophyllia natans Colpophyllia natans Colpophyllia cylindrus Dendrogyra cylindrus Dendrogyra stokesii Dichocoenia Haime, 1848 labyrinthiformis Diploria fastigiata Eusmilia fragum Favia Manicina areolata (as

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316 KORZHAVINA et al. - - - Stock, 1975a Stock, 1975a Stock, Kim, 2010 1989 Stock, 1979 Butter, & Immer Herriott mann, 1979 1989 Stock, 1975a Stock, 1975a Stock, & Immer Herriott mann, 1979 1975a Stock, 1989 Stock, 1975b Stock, et al., 2005 Varela & Immer Herriott mann, 1979 1989 Stock, 1989 Stock, Reference

Site abbreviation** Cur Cur Bar Cur Cur VI Cur Cur Cur VI Cur Cur Cur Cub VI Cur Cur )

cont. (

Copepod family abbreviation* Cor Cor Ast Ast Cor Cor Ast Cor Cor Cor Cor Ast Ast Ast Cor Ast Ast

Stock, Stock, Varela, Varela, Stock, Stock, Stock, 1989 Stock, Stock, Stock,

Stock, 1987 Stock, Stock, 1975 Stock,

Stock, 1975 Stock, Kim, 2010 Stock, 1975 Stock, Stock, 1987 Stock,

Stock, 1989 Stock, Stock, 1975 Stock,

Stock, 1989 Stock,

sp. sp. sp. Copepod species name Corallovexia mediobrachium Corallovexia 1975 longibrachium Corallovexia 1975 capillatus Asteropontius iuxtus Asteropontius longicauda Corallonoxia Corallovexia coronatum Meandromyzon dorsospinosa minor Corallovexia 1975 dorsospinosa Corallovexia Corallovexia ventrospinosa Corallovexia montastreae Hermacheres rostratum Peltomyzon mycetophylliae Asteropontius Ortiz & Lalana, 2005 Corallovexia maxillatus Asterocheres foliata Asteropontella

and as Milne Edwards Milne Edwards Milne Edwards (Verrill, 1901)) (Verrill, (Ellis & Solander, (Ellis & Solander, name (Linnaeus, 1767) (Linnaeus, 1767) (Linnaeus, 1767) (Linnaeus, 1767) (Linnaeus, 1767) (Linnaeus, 1767) (Linnaeus, (Linnaeus, 1758) (Linnaeus, 1758) (Linnaeus, 1758) (Linnaeus, 1758) (Linnaeus, (Ellis & Solander, 1786) (Ellis & Solander, 1786) (Ellis & Solander, (Linnaeus, 1758) (Linnaeus, 1758) (Linnaeus, Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans Caribbean reef-dwelling sp.

Montastraea brasiliana Montastraea annularis Montastraea TABLE 2 Host (valid Invertebrate in the original record) indicated Manicina areolata Manicina areolata Meandrina meandrites Meandrina meandrites Meandrina meandrites Meandrina meandrites Meandrina cavernosa Montastraea (as cavernosa Montastraea cavernosa Montastraea cavernosa Montastraea cavernosa Montastraea cavernosa Montastraea lamarckiana Mycetophyllia & Haime, 1849 lamarckiana Mycetophyllia & Haime, 1849 annularis Orbicella (as 1786)) annularis Orbicella Downloaded from Brill.com02/21/2020 01:34:50PM via Universiteit of Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 317 ; - - - - Herriott Herriott Herriott ; ; Stock, 1975a Stock, & Immer Herriott mann, 1979 1987 Stock, 1975a Stock, & Immermann, 1979 & Immer Herriott mann, 1979 1975a Stock, 1975a Stock, & Immermann, 1979 & Immer Herriott mann, 1979 & Immer Herriott mann, 1979 1987 Stock, 1973 Humes & Stock, 1975c Stock, 1973 Stock, 1975c Stock, 1973 Stock, 1973 Humes & Stock, Cur VI Cur VI Cur, VI Cur VI Cur, VI VI Cur Jam Cur, Bar, Cur Cur Cur PR Cor Cor Ast Cor Cor Cor Cor Cor Cor Ast Rhy Lam Rhy Lam Rhy

Stock, 1975 Stock, Stock, Stock, Stock, 1975 Stock, Stock, Stock, Humes & Humes &

Stock, 1987 Stock, 1987 Stock, Stock, 1975 Stock,

Stock, 1973 Stock, 1973 Stock,

Stock, 1975 Stock, sp. sp. Corallovexia mediobrachium Corallovexia diploriae Hermacheres mediobrachium Corallovexia 1975 brevibrachium Corallovexia 1975 longibrachium Corallovexia mediobrachium Corallovexia 1975 similis Corallovexia Corallovexia diploriae Hermacheres bilobipes Acanthomolgus 1973 Stock, Lamippina aequalis dionyx Acanthomolgus Lamippina aequalis bilobipes Acanthomolgus 1973 Stock,

sp.) -

Biel (Gmelin, (Pallas, 1766)) (Pallas, elastica (Ellis & Solander, (Ellis & Solander, (Gmelin, 1791) (Dana, 1846) (Dana, 1846) (Dana, 1846) (Dana, 1846) (Dana, 1846) (Dana, 1846) (Dana, (Ellis & Solander, (Ellis & Solander, (Ellis & Solander, (Ellis & Solander, (Pallas, 1766) (Pallas, 1766) (Pallas, var. Pseudopterogorgia (Dana, 1846)) (Dana, Antillogorgia elastica Antillogorgia (Ellis & Solander, 1786) (Ellis & Solander, sp. (as sp. Diploria clivosa Diploria Diploria strigosa Diploria acerosa Pseudopterogorgia americana Pseudopterogorgia Orbicella annularis Orbicella clivosa Pseudodiploria 1786) (as 1786)) clivosa Pseudodiploria 1786) clivosa Pseudodiploria 1786) strigosa Pseudodiploria (as strigosa Pseudodiploria strigosa Pseudodiploria strigosa Pseudodiploria strigosa Pseudodiploria strigosa Pseudodiploria Octocorallia Alcyonacea acerosa Antillogorgia (as acerosa Antillogorgia americana Antillogorgia (as 1791)) Antillogorgia acerosa Antillogorgia 1929 (as schowsky, 1929) Bielschowsky,

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318 KORZHAVINA et al. Stock, Stock, ; Stock, 1973 Stock, & Lalana, 2007 Varela 1980 Grygier, 1973 Humes, 1975c Stock, 1975c Stock, 1975c Stock, 1975c Stock, 1973 Stock, 2011b Varela, 1975c Stock, et al., 2008 Varela 1973 Humes, 1973 Humes, 1975c 1975c Stock, et al., 2017 Ivanenko et al., 2003 Varela Reference

Site abbreviation** PR Cub Bah Ber Cur Cur Cur Cur PR Cub Cur Cub Ber SE Cur, Ber, SE, SM Cur, SE Cub )

cont. (

Copepod family abbreviation* Lam Ast Lam Rhy Rhy Rhy Rhy Rhy Lam Ast Rhy Rhy Rhy Rhy Rhy Lam Rhy

Stock, Stock, Stock, Stock, Stock, Stock, Stock, Stock, Humes, Humes, Humes, Humes,

Stock, 1975 Stock,

Stock, 1975 Stock, Stock, 1973 Stock,

Stock, 1973 Stock, Humes, 1973 Humes,

Stock, 1975 Stock, Grygier, 1980 Grygier,

Varela, 2011 Varela, Varela & Lalana, 2007 Varela

sp. Copepod species name variicauda Enalcyonium ortizi Orecturus caligicola Sphaerippe verrucipes Acanthomolgus 1973 mononyx Acanthomolgus affinis Acanthomolgus longidactylus Acanthomolgus 1975 intermedius Acanthomolgus 1975 euniceae Enalcyonium antillensis Orecturus longifurca Acanthomolgus triangulipes Acanthomolgus 1975 bayeri Acanthomolgus gorgoniae Acanthomolgus 1973 triangulipes Acanthomolgus 1975 Sphaerippe intermedius Acanthomolgus 1975

and as name (Pallas, 1766) (Pallas, 1766) (Pallas, Lamouroux, 1821) Lamouroux, Milne Edwards & Milne Edwards Linnaeus, 1758 Linnaeus, 1758 Linnaeus, 1758 Linnaeus, 1758 Linnaeus, Lamouroux, 1816 Lamouroux, 1816 Lamouroux, (Ellis & Solander, 1786) (Ellis & Solander, Bayer, 1961 Bayer,

Bayer, 1961 Bayer, Duchassaing &

(Lamouroux, 1821) (Lamouroux, 1821) (Lamouroux, Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans Caribbean reef-dwelling sp. sp.

Plexaura flexuosa Plexaura TABLE 2 Host (valid Invertebrate in the original record) indicated asbestinum Briareum asbestinum Briareum Callogorgia calyculata Eunicea clavigera Eunicea flexuosa Eunicea (as flexuosa Eunicea laciniata Eunicea 1860 Michelotti, mammosa Eunicea mammosa Eunicea tourneforti Eunicea Haime, 1857 mariae Gorgonia ventalina Gorgonia ventalina Gorgonia ventalina Gorgonia ventalina Gorgonia Muricea

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 319 ; Stock, Stock, ; ; Varela, Humes, 1973 Humes, 1975c 1975c Stock, 1979 Stock, 2011a Varela, 1975c Stock, 1973 Stock, 1973 Stock, 1975c Stock, et al., 2005b Varela 1973 Humes, et al., 2003 Varela 1978 Stock, 1973 Humes & Stock, Kim, 2010 2000 Humes, 2010 2000 Humes, Kim, 2010 1995 & Humes, Stock 2011a Varela, Ber Cur Flo Cub Cur PR PR SM Cub Ber Cub Flo SM Jam Bel, Cub Bel Jam MI, Jam, Cur, Bah, Cub, SM, PR Rhy Rhy Lam Rhy Rhy Lam Lam Rhy Lam Rhy Rhy Lam Mac Ast Ast Ast Ast Ast

Stock, 1978 Stock, Stock, 1975 Stock, Stock, 1975 Stock, Humes, 2000 Humes, 2000 Humes, 2000 Humes,

Stock & Stock

Stock, 1973 Stock,

Kim, 2010 Humes, 1973 Humes, 1973 Humes,

Stock, 1975 Stock, 1975 Stock, Sars, 1916 Sars, Stock, 1973 Stock, Stock, 1979 Stock,

sp. Acanthomolgus muriceanus Acanthomolgus 1973 Humes, aequiseta Acanthomolgus bouligandi Linaresia affinis Acanthomolgus affinis Acanthomolgus nudum Enalcyonium ramosum Enalcyonium seticornis Acanthomolgus Enalcyonium bayeri Acanthomolgus bayeri Acanthomolgus Magnippe caputmedusae sargassi Macrochiron crinoidicola Asterocheres crinoidicola Asterocheres crinoidicola Asterocheres unioviger Asterocheres longipalpus Chelacheres 1995 Humes, - Plex

Esper, 1794) Esper, (Houttuyn, 1772) (Houttuyn, (Pourtales, 1869) (Pourtales, (Esper, 1791) (Esper, (Linnaeus, 1758) (Linnaeus, (Esper, 1794) (as (Esper, 1794) (Esper, 1794) (Esper, homomalla (Duchassaing & (Kukenthal, 1911) (Kukenthal, Clark, 1909 (Pallas, 1766) (Pallas, f. Wright & Studer, 1889 & Studer, Wright Verrill, 1864 Verrill, sp. sp. Muricea atlantica Muricea laxa Muricea Placogorgia Plexaura homomalla Plexaura homomalla aura homomalla Plexaura homomalla Plexaura dichotoma Plexaurella nutans Plexaurella 1860) Michelotti, porosa Pseudoplexaura Pseudoplexaura Thesea Pennatulacea reniformis Renilla Echinodermata Crinoidea Crinoidea Comatulida rubiginosus Davidaster grandis Nemaster Echinoidea Echinoidea Camarodonta lucunter Echinometra

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320 KORZHAVINA et al. Varela et al., Varela ; Stock & Humes, 1995 & Humes, Stock 1973 Humes & Stock, 1995 & Humes, Stock 1973 Humes & Stock, 1995 & Humes, Stock 1973 Humes & Stock, Kim, 2010 et al., 1963b Stock & Gooding, Stock 1986 2005b et al., 1963b Stock et al., 1963a Stock Reference

Site abbreviation** PR Cur Cur MI, PR Jam, Bon, Cur, Cur MI Jam, Cur, Bah, Bar, Bah, PR MI Bah, Jam, Cur Cub, Jam Cur, Jam )

cont. (

Copepod family abbreviation* Ast Mac Ast Mac Ast Mac Ast Pse Ast Sync Mic Stock, Humes Stock, Humes & Humes & Humes &

Stock & Humes, & Humes, Stock T. Scott, 1898) Scott, T.

Stock, Humes & Stock, Kim, 2010

(T. Scott, 1898) Scott, (T. Stock & Gooding, Stock

Stock, Humes & Stock, Stock & Humes, & Humes, Stock

Ascomyzon simulans Ascomyzon Copepod species name optans Chelacheres 1995 echinicolum Macrochiron 1973 Stock, simulans Asterocheres (as echinicolum Macrochiron 1973 Stock, longipalpus Chelacheres 1995 echinicolum Macrochiron 1973 Stock, unioviger Asterocheres pectinifer Pseudanthessius & Gooding, 1964 alatus Onychocheres 1986 amplectans Meomicola Gooding, 1963 glaber Micropontius Gooding, 1963

and as name (Lamarck, 1816) (Lamarck, 1816) (Lamarck, (Linnaeus, 1758) (Linnaeus, (Lamarck, 1816) (Lamarck, 1816) (Lamarck, (Lamarck, 1816) (Lamarck, Philippi, 1845 (Linnaeus, 1758) (Linnaeus, A. Agassiz, 1863 A. Agassiz, (Lamarck, 1816) (Lamarck, 1816) (Lamarck, Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans Caribbean reef-dwelling

TABLE 2 Host (valid Invertebrate in the original record) indicated lucunter Echinometra viridis Echinometra variegatus Lytechinus variegatus Lytechinus ventricosus Tripneustes ventricosus Tripneustes Cidaroida tribuloides Eucidaris Clypeasteroida rosaceus Clypeaster Diadematoida Diadema antillarum Spatangoida ventricosa Meoma ventricosa Meoma

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 321 Varela Varela ; Humes & Hendler ; ; Stock et al., 1962 Stock 1995 & Humes, Stock 1969b Humes, 1969b Humes, Hendler & Kim, 2010 Hendler & Kim, 2010 1969b Humes, Hendler & Kim, 2010 1969b Humes, 1969b Humes, et al., 2003 1968 Stock, 1970 Ho, Hendler & Kim, 2010 1968 Stock, & Kim, 2010 et al., 1963b Stock 1891 Edwards, et al., 1962 Stock 1968 Stock,

Jam Cur Bah, Jam Bah, Jam Bah Bah, Jam Jam Jam Jam Bah, Cur, Cur, Bah, Bon, Cub, Jam MI, PR Jam, Bah, Cur, Bah, PR MI, PR Jam, Cur, Cur Bah MI, PR PR Nan Pse Sync Sync Lic Lic Sync Lic Sync Sync Lic Lic Lic Pse Lic Nan Lic

Stock & Stock

Stock, Humes Stock, Edwards, 1891 Edwards, 1891 Edwards, 1891 Edwards, 1891 Edwards, Humes, 1969) Humes, Stock, 1968 Stock,

Stock, 1968 Stock, 1968 Stock,

Humes & Ho, 1971 Humes & Ho,

Stock, Humes & Stock, (Humes, 1969) (Humes, 1969) (Humes, 1969) (as (Humes, Stock, Humes & Stock, sp. sp. Nanaspis pollens Nanaspis Gooding, 1962 exilicornis Pseudanthessius 1995 Humes, Caribulus sculptus Caribulus nasutum Diogenidium tectum Diogenidium sculptus Caribulus nasutum Diogenidium Caribulus sculptus Caribulus sculptus Scambicornus spinicauda Diogenella deforme Diogenidium nasutum Diogenidium deficiens Pseudanthessius & Gooding, 1963 nasutum Diogenidium truncata Nanaspis Gooding, 1962 deforme Diogenidium

Ludwig, Ludwig, Ludwig, Ludwig, Ludwig, Ludwig, (Ludwig)) (Selenka, Jaeger, 1833 Jaeger, Selenka, 1867 Selenka, 1867 (Selenka, 1867) (as (Selenka, 1867) (Selenka, 1867) (Selenka, 1867) (Lamarck, 1816) (Lamarck, 1816) (Lamarck, Selenka, 1867) Ludwigothuria mexicana Ludwigothuria Meoma ventricosa Meoma ventricosa Meoma Holothuroidea agassizii Actinopyga agassizi Muelleria agassizii Actinopyga agassizii Actinopyga agassizii Actinopyga grisea (Halodeima) Holothuria grisea (Halodeima) Holothuria mexicana (Halodeima) Holothuria 1875 mexicana (Halodeima) Holothuria 1875 mexicana (Halodeima) Holothuria 1875 mexicana (Halodeima) Holothuria 1875 mexicana (Halodeima) Holothuria 1875 mexicana (Halodeima) Holothuria 1875 (as scabra (Metriatyla) Holothuria parvula (Platyperona) Holothuria parvula (Selenka)) 1867) (as Microthele glaberrima (Selenkothuria) Holothuria Selenka, 1867

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322 KORZHAVINA et al. Humes & Hendler ; ; Stock, 1968 Stock, 1970 Ho, et al., 1962 Stock 1970 Humes & Ho, 1970 Humes & Ho, Hendler & Kim, 2010 1970 Humes & Ho, et al., 1962 Stock et al., 1962 Stock 1968 Stock, 1969b Humes, 1969b Humes, 1968 Stock, & Kim, 2010 Reference

Site abbreviation** MI, PR MI Bar Bar Bar MI Jam MI PR Jam Bah, Bar, MI, PR Jam, Bah, Bar, PR Jam, )

cont. (

Copepod family abbreviation* Lic Nan Lic Lic Lic Lic Nan Nan Lic Sync Sync Lic

Stock, 1968 Stock, Humes &

Stock, 1968 Stock,

Stock, 1968 Stock, 1968 Stock, 1968 Stock,

Stock, Humes & Stock, (Humes, 1969) (Humes, Humes & Ho, 1970 Humes & Ho,

Stock, Humes & Stock, Humes & Stock, sp. Copepod species name seticauda Diogenella pollens Nanaspis Gooding, 1962 deichmannae Diogenella 1970 Ho, impar Diogenella deforme Diogenidium seticauda Diogenella pollens Nanaspis Gooding, 1962 truncata Nanaspis Gooding, 1962 seticauda Diogenella Caribulus sculptus Caribulus spinulosum Diogenidium

Semper, Semper, Semper, Semper, Semper, Semper, Semper, (Semper, (Semper, and as name (Selenka, 1867) (Selenka, 1867) (Selenka, 1867) Halodeima surinamensis Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans Caribbean reef-dwelling Brandtothuria arenicola Brandtothuria

TABLE 2 Host (valid Invertebrate in the original record) indicated surinamensis (Semperothuria) Holothuria Ludwig, 1875 surinamensis (Semperothuria) Holothuria Ludwig, 1875 (as (Ludwig)) arenicola (Thymiosycia) Holothuria 1868 arenicola (Thymiosycia) Holothuria 1868 (as 1868)) arenicola (Thymiosycia) Holothuria 1868 arenicola (Thymiosycia) Holothuria 1868 arenicola (Thymiosycia) Holothuria 1868 arenicola (Thymiosycia) Holothuria 1868 impatiens (Thymiosycia) Holothuria 1775) (Forsskal, Isostichopus badionotus Isostichopus badionotus Isostichopus badionotus

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 323 Stock et al., Stock Emson et al., ; ; Stock et al., 1962 Stock et al., 1962 Stock et al., 1962 Stock et al., 2008 Varela 1973 Humes & Stock, unpublished Ivanenko, Humes & Hendler, 1972 Humes & Hendler, 1972 Humes & Hendler, 1999 1963a Humes & Hendler, 1999 Kim, 2009 Humes & Hendler, 1999 Emson & Mladenov, 1987 1985 Jam MI, PR Bah, Jam Cub MI, PR Jam, Bar, SE FK FK Bel, Cur Bel, FK Bah Bel Bel, Jam Nan Nan Nan Rhy Rhy Tha Can Synp Can Can Cla Can Can )

Humes &

Humes &

( Humes, 1984 Humes,

Humes &

Stock, Humes & Stock, Humes & Stock, Humes & Hendler, Humes & Hendler, Humes & Hendler,

Stock, Humes & Stock, Doridicola Doridicola Stock, Humes & Stock, Stock, Humes & Stock, sp. A sp. Humes & Stock, 1973 Humes & Stock,

Humes & Stock, 1983 Humes & Stock, ) (as Nanaspis exigua Nanaspis Gooding, 1962 media Nanaspis Gooding, 1962 pollens Nanaspis Gooding, 1962 Critomolgus astrophyticus Critomolgus 1973 Stock, astrophyticus Caribeopsyllus ligatus Parophiopsyllus 1972 Hendler, amphiopli Presynaptiphilus 1972 Hendler, reductus Ophiopsyllus Gooding, 1963 latus Ophiopsyllus 1999 columnaris Hemicyclops latus Ophiopsyllus 1999 reductus Ophiopsyllus Gooding, 1963 (Clark, 1900) (Clark, 1900) (Selenka, 1867) (Selenka, 1867) (Selenka, 1867) (Lamarck, 1816) (Lamarck, 1816) (Lamarck, (Lamarck, 1816) (Lamarck, Lutken, 1856) Lutken, Muller & Troschel, 1842 Troschel, Muller & Lutken, 1856 Lutken, sp. sp. Ophiocoma riisei Ophiocoma Isostichopus badionotus Isostichopus badionotus Isostichopus badionotus Ophiuroidea Euryalida muricatum Astrophyton muricatum Astrophyton Ophiurida Ophiuroida Amphioplus Amphioplus echinata Ophiocoma pumila Ophiocoma wendtii Ophiocoma (as ophiactoides Ophiocomella ophiactoides Ophiocomella

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324 KORZHAVINA et al. ; Humes & Hendler, Humes & Hendler, 1999 et al., 1963b Stock Humes & Hendler, 1999 1998 Humes, 1998 Humes, Hendler & Kim, 2010 Hendler & Kim, 2010 1998 Humes, Hendler & Kim, 2010 Kim, 2010 Kim, 2010 Kim, 2010 Kim, 2010 Kim, 2010 Kim, 2009 Kim, 2009 Kim, 2010 Kim, 2010 Kim, 2010 Reference

Site abbreviation** Bel MI, SM Bel, Cur, MI Bar Bel Bel Bar Bel Bah Jam Bah Bah Bah Jam Bah Bah Jam Jam )

cont. Copepod family abbreviation* Pse Pse Ast Ast Tha Tha Ast Tha Ast Ast Ast Ast Ast Sab Cla Ast Ast Ast Kim, 2009

Kim, 2010 Stock, Humes Stock, Humes Stock, Kim, 2010

Humes, 1984 Humes,

Kim, 2010 Kim, 2010

Kim, 2010

Humes, 1998 Humes, Kim, 2010 Suarez-Morales, Suarez-Morales, Suarez-Morales, Kim, 2010 Kim, 2010

Humes, 1998 Humes, 1998 Humes,

sp. A sp. Suarez-Morales & Castellanos, 1998 & Castellanos, Suarez-Morales 1998 & Castellanos, Suarez-Morales Copepod species name deficiens Pseudanthessius & Gooding, 1963 deficiens Pseudanthessius & Gooding, 1963 bellulus Ophiurocheres lunulifer Collocheres Caribeopsyllus chawayi Caribeopsyllus In: lunulifer Collocheres chawayi Caribeopsyllus In: bahamensis Asterocheres oricurvus Asterocheres peniculatus Asterocheres plumosus Asterocheres trisetatus Asterocheres occidentalis Eupolymniphilus columnaris Hemicyclops aculeata Hetairosynella angulata Hetairosynella bifurcata Hetairosynella

(Say, 1825) (Say, 1825) (Say, 1825) (Say, Lutken, Lutken, Muller & and as (Say, 1825)) (Say, name (Say, 1825) (Say, Muller & Troschel, Troschel, Muller & (Say, 1825) (Say, Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans ( Caribbean reef-dwelling Ophioderma cinereum

Ophioderma brevispinum TABLE 2 Host (valid Invertebrate in the original record) indicated Ophioderma brevispina (as Ophioderma cinerea 1842 (as 1842) Troschel, flaccida Ophiomyxa suensoni (Acanthophiothrix) Ophiothrix 1856 Lutken, angulata (Ophiothrix) Ophiothrix angulata (Ophiothrix) Ophiothrix angulata (Ophiothrix) Ophiothrix oerstedii (Ophiothrix) Ophiothrix 1856 Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 325

Kim, 2010 Kim, 2009 Kim, 2010 Garcia-Hernandez et al., 2019 et al., 2008 Varela 1994 & Huys, Boxshall 2010b Varela, 2010b Varela, et al., 2007 Varela 2010 Varela, et al., 2005b Varela et al., 2007a Varela 2012 Varela, Bar Jam PR PR Cub Bel Cub Cub Cub Cub Cub Cub Cub Ast Pse Ast Ast Ast Ast Ast Ast Ast Ast Ast Ast ) ) ) (as Varela, Varela, Varela, 2012 Varela, Kim, 2009 Kim, 2010 (

Kim, 2010

Varela, 2010 Varela,

Varela, Ortiz & Varela, Kim, 2010 Varela, 2010 Varela, Varela, Ortiz & Varela, Boxshall & Huys, & Huys, Boxshall & Huys, Boxshall (

Boeck, 1859 Kimcheres fastigatus Kimcheres fastigatus Asterocheres acutus Pseudanthessius edentatum Stenomyzon Copepoda reginae Asterocheres 1994 reginae Asterocheres 1994 cubensis Asterocheres fernandezmilerai Asterocheres 2010 garridoi Asterocheres Lalana, 2007 antillensis Asterocheres Asterocheres espinosai Asterocheres Lalana, 2007 humesi Neoasterocheres

(Lamarck, 1814) (Lamarck, 1814) (Lamarck, 1814) (Lamarck, Alcolado, 1984 Alcolado, (Duchassaing & (Schmidt, 1870) Azevedo, Padua, Moraes, Moraes, Padua, Azevedo, Duchassaing & Michelotti, Duchassaing & Michelotti, sp. sp. Porifera Porifera Porifera Calcarea Calcinea Clathrinida lutea Clathrina Muricy & Klautau, 2017 Rossi, Demospongiae Heteroscleromorpha Agelasida Agelas clathrodes Agelas dilatata Agelas 1864 wiedenmayeri Agelas Axinellida walpersii Ptilocaulis 1864) Michelotti, Haplosclerida Callyspongia vaginalis Callyspongia vaginalis Callyspongia vaginalis Callyspongia

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326 KORZHAVINA et al.

- - Varela Varela ; Varela, 2012 Varela, 2012 Varela, 2012 Varela, et al., 2008 Varela 1992b Stock, 1992b Stock, et al., 2008 Reference

Site abbreviation** Cub Cub Cub Cub Cur PR Cur, Aru, Cub, )

cont. Copepod family abbreviation* Ast Ast Ast Ast Ent Ent ) Stock, 1992 Stock, (Scott T. & Scott & Scott T. (Scott (Johnsson, 1998) Wilson C.B., 1944 Wilson C.B., Varela, 2013 Varela, Varela, 2012 Varela,

Parmulodes verrucosus Parmulodes Copepod species name paraboecki Setacheres kimi Asterocheres lalanai Asterocheres elongatus Scottocheres A., 1894) emarginata Parmulella verrucosa Parmulodes (as

and as name (Lamarck, 1814) (Lamarck, (Higgin, 1877) Schmidt, 1862 Schmidt, 1862 (Lamarck, 1814) (Lamarck, 1814) (Lamarck, Caribbean reef-dwelling anthozoans, echinoderms and sponges as hosts of echinoderms and sponges anthozoans, copepod crustaceans ( Caribbean reef-dwelling

TABLE 2 Host (valid Invertebrate in the original record) indicated vaginalis Callyspongia digitalis Niphates digitalis Niphates Poecilosclerida birotulata Iotrochota Verongimorpha Chondrillida nucula Chondrilla nucula Chondrilla Lam – Lamippidae, Lic Li Ent – Entomolepididae, Can – Cancerillidae, Cla Clausidiidae, Cor Corallovexiidae, Ast – Asterocheridae, * Copepod family Abbreviation: Sab – Sabel – Rhynchomolgidae, Rhy - Pseudocyclopidae, Psec – Pseudanthessiidae, Pse – Nanaspididae, Nan Mic – Micropontiidae, chomolgidae, Mac – Macrochironidae, Thaumatopsyllidae. Tha – – Synaptiphilidae, Synp – Synapticolidae, liphilidae, Sync Flo – Florida, Cub – Cuba, Cur Curacao, Ber – Bermuda, Bon Bonaire, Bel – Belize, – Baradal, Barad Bar – Barbados, Aru – Aruba, Bah Bahamas, Abbreviation: ** Site Virgin Islands. VI – SM – St. Martin, SE – St. Eustatius, Rico, PR – Puerto – Panama, Pan Key, Island, LK – Loggerhead MI – Magueyes – Jamaica, Jam

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 327 coral) (fig. 1a) (Humes, 1984; Stock, 1992a; Kim, Southern Gulf of Mexico (table 6, figs. 4–7; 2009). All other species of symbiotic cope- Spalding et al., 2007; Hoeksema et al., 2017a). pods are found in the Caribbean only, which Eight records are known for Bermuda: four may be the result of high endemism of their species of poecilostomatoid cyclopoids rep- host species (Miloslavich et al., 2010; Soest resenting genus Acanthomolgus and the si- et al., 2012; Zea et al., 2014; Ivanenko, 2016). phonostomatoid Ophiopsyllus reductus were The copepods were collected by washing found associated with alcyonaceans and an of the hosts in seawater or tap water (Butter, ophiuroid, respectively (Stock et al., 1963a; 1979; Garcia-Hernandez et al., 2017), a solu- Humes, 1973). Only one record is reported tion of formalin (Varela & Lalana, 2007; Varela for the Southwestern Caribbean: the cala- et al., 2003, 2005a, b, 2007a, b, 2008; Varela, noid Ridgewayia fosshageni aggregating on 2010a, b, 2011a, b, 2012), a solution of ethanol the actinairian Bartholomea annulata. Only (Stock et al., 1962, 1963a, b, Humes, 1969a, 1998, Aspidomolgus stoichactinus living on the ac- 2000; Humes & Stock, 1973; Stock, 1975a, b, d; tiniarian Stichodactyla helianthus and the Herriott & Immermann, 1979; Stock & Humes, corallimorpharian Corynactis denticulosa, 1995) or a solution of magnesium-chloride recorded as Homostichanthus denticulosus, is (Stock & Humes, 1995), dissecting of the host’s found in five ecoregions (Humes, 1969a; Stock, tissues or galls (see Stock et al., 1962; Humes & 1975b). Five species of copepods are found in Hendler, 1972, 1999; Humes & Goenaga, 1978; four ecoregions, viz. Caribulus sculptus living Stock, 1978; Grygier, 1980; Emson et al., 1985; on holothurians, Chelacheres longipalpus and Emson & Mladenov, 1987; Hendler & Kim, Macrochiron echinicolum (fig. 1d) found on sea 2010; Ivanenko et al., 2017), by a suction device urchins, Ophiopsyllus reductus living on ophi- (Humes & Stock, 1973) or by dissolving soft tis- uroids. Seventeen and 87 species of copepods sues of host corals with bleach (Stock 1975a, are recorded found in only two and one ecore- 1989). Most of the host invertebrates were col- gions, respectively (table 6). lected by SCUBA living down to 41 m depth. The data show that the three most inten- The only exceptions are the three of eight spe- sively explored ecoregions are the Bahamian, cies of the Caribbean copepods of the family Greater Antilles and Southern Caribbean, Lamippidae tentatively included in the da- with studies centered at Curaçao (123 records, tabase found in galls or tissue of octocorals 36 species of hosts, 49 species of copepods), collected by a submersible away from reefs at Puerto Rico (105, 22, 27) and the Bahamas (57, depths of 55–330 m (fig. 2 a–d) (Stock, 1978, 11, 19). The mosaic data show poor knowledge 1979; Grygier, 1980). It is noteworthy that there of most ecoregions as well as and many host are no data on symbiotic copepods associated taxa. This current state of the knowledge lim- with reef-dwelling sponges, echinoderms and its analysis of the distribution of symbiotic co- corals living at mesophotic depths; studies on pepods in the whole Caribbean. Well planned reef communities and coral-associated fauna studies of different Caribbean regions and the from mesophotic depths seems to be an im- application of modern methods of integrative portant task that has just started (Bongaerts taxonomy are needed to carry out such analy- et al. 2010, 2015; van der Meij et al., 2015; Hoek- ses (DeBiasse et al., 2016; Jossart et al., 2017; sema et al., 2017c; García-Hernández et al., Ivanenko et al., 2018). 2018; Veglia et al., 2018). A comparison of taxonomic names from Symbiotic copepods are reported from literature sources with their current nomen- eight out of ten ecoregions of the Greater clature revealed name changes for 29 (of 80)

Caribbean, but none from the Northern and species and 12 Downloaded(of 58) fromgenera Brill.com02/21/2020 of the hosts01:34:50PM via Universiteit of Groningen

328 KORZHAVINA et al.

FIGURE 3 Siphonostomatoida, dorsal view. a – Asterocheres unioviger, b – Cyclocheres sensilis (Asterocheridae), c – Parophiopsyllus ligatus (Cancerillidae), d – Micropontius glaber (Micropontiidae), e – Nanaspis pol- lens (Nanaspididae). After Stock et al. (1962, 1963a), Humes & Hendler (1972), Kim (2010). Downloaded from Brill.com02/21/2020 01:34:50PM via Universiteit of Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 329

TABLE 3 Numbers of records and species of Caribbean symbiotic copepods found on reef-dwelling anthozoans, echinoderms and sponges attributed to the host family

Host family Number of copepod Number of copepod records species

Echinodermata Holothuriidae 75 12 35 5 Ophiocomidae 29 3 Ophiotrichidae 19 2 Toxopneustidae 18 3 Echinometridae 17 3 Gorgonocephalidae 13 1 Ophiodermatidae 10 1 Amphiuridae 9 2 Brissidae 9 4 Clypeasteridae 6 1 Diadematidae 4 1 Comatulidae 3 1 Cidaridae 2 1 Ophiomyxidae 1 1 Cnidaria Faviidae 47 12 Plexauridae 29 16 Stichodactylidae 22 2 Gorgoniidae 21 6 Meandrinidae 20 5 Actiniidae 16 3 Montastraeidae 13 6 Merulinidae 7 2 Acroporidae 5 2 Corallimorphidae 4 2 Briareidae 4 2 Aiptasiidae 2 1 Antipathidae 2 1 Ricordeidae 2 1 Renillidae 1 1 Phymanthidae 1 1 Primnoidae 1 1 Porifera Clathrinidae 42 1 Chondrillidae 12 2

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330 KORZHAVINA et al.

TABLE 3 Numbers of records and species of Caribbean symbiotic copepods found on reef-dwelling anthozoans, echinoderms and sponges attributed to the host family (cont.)

Host family Number of copepod Number of copepod records species

Callyspongiidae 5 4 Agelasidae 4 3 Niphatidae 2 2 Iotrochotidae 1 1 Axinellidae 1 1

TABLE 4 Number of records and species per family of symbiotic copepods living on reef-dwelling Caribbean hosts (anthozoans, echinoderms and sponges)

Copepod family Number of records Number of species

Asterocheridae 103 45 Rhynchomolgidae 78 19 Corallovexiidae 66 11 Synapticolidae 53 3 Lichomolgidae 43 8 Cancerillidae 34 3 Pseudanthessiidae 21 4 Nanaspididae 19 4 Macrochironidae 18 2 Thaumatopsyllidae 17 1 Lamippidae 16 8 Entomolepididae 12 2 Synaptiphilidae 3 1 Clausidiidae 2 1 Micropontiidae 2 1 Pseudocyclopidae 2 1 Sabelliphilidae 1 1

(WoRMS, 2019). The taxonomic names are 41 records for which the hosts are identified changed for nine (of 115) species and six (of 47) only to taxonomic categories such as phy- genera of the copepods. There are six records lum, order or subclass. There are 81 records of symbiotic copepods identified to genus of copepods found on sponges, but hosts of (Asterocheres, Corallovexia, Corallonoxia, 14 of them are defined only to phylum (Kim, Critomolgus, Enalcyonium, Sphaerippe). These 2009, 2010). The absence of precise identifi- taxa are included in; among 16 hosts (for 18 cations and the necessity of linking outdated records) that have no identification at species host names with valid ones show the need for level; of these one is assigned to a phylum, specimen collections of not only copepods Downloaded from Brill.com02/21/2020 01:34:50PM three to a class and 12 to a genus. There are but also of their invertebrate hosts,via Universiteit as well of as Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 331

TABLE 5 Families of reef-dwelling Caribbean symbiotic copepods and their host families (anthozoans, echino- derms and sponges)*

Copepod family Host family Host class or type

Asterocheridae Actiniidae Ant Agelasidae Dem Axinellidae Dem Briareidae Ant Callyspongiidae Dem Cidaridae Ech Comatulidae Cri Diadematidae Ech Echinometridae Ech Iotrochotidae Dem Meandrinidae Ant Merulinidae Ant Montastraeidae Ant Niphatidae Dem Ophiomyxidae Oph Ophiotrichidae Oph Phymanthidae Ant Plexauridae Ant Ricordeidae Ant Stichodactylidae Ant Toxopneustidae Ech Cancerillidae Amphiuridae Oph Ophiocomidae Oph Clausidiidae Ophiocomidae Oph Corallovexiidae Acroporidae Ant Meandrinidae Ant Merulinidae Ant Montastraeidae Ant Faviidae Ant Entomolepididae Chondrillidae Dem Lamippidae Briareidae Ant Gorgoniidae Ant Plexauridae Ant Primnoidae Ant Lichomolgidae Holothuriidae Hol Stichopodidae Hol Macrochironidae Echinometridae Ech Renillidae Ant Toxopneustidae Ech Micropontiidae Brissidae Ech

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332 KORZHAVINA et al.

TABLE 5 Families of reef-dwelling Caribbean symbiotic copepods and their host families (anthozoans, echino- derms and sponges)* (cont.)

Copepod family Host family Host class or type

Nanaspididae Brissidae Ech Holothuriidae Hol Stichopodidae Hol Pseudanthessiidae Brissidae Ech Clypeasteridae Ech Holothuriidae Hol Ophiodermatidae Oph Rhynchomolgidae Actiniidae Ant Antipathidae Ant Corallimorphidae Ant Gorgoniidae Ant Gorgonocephalidae Oph Plexauridae Ant Ricordeidae Ant Stichodactylidae Ant Sabelliphilidae Porifera Po Synapticolidae Brissidae Ech Holothuriidae Hol Ophiotrichidae Oph Stichopodidae Hol Synaptiphilidae Amphiuridae Oph Thaumatopsyllidae Ophiotrichidae Oph

* Abbreviations: Ant – Anthozoa, Cri – Crinoidea, Dem – Demospongiae Ech – Echinoidea, Hol – Holothuroidea, Hom – Homoscleromorpha, Oph – Ophiuroidea, Po – Porifera. morphological and molecular studies of this dae and Meandrinidae (updated classification material (Rocha et al., 2014). The DNA-barcod- in Hoeksema & Cairns, 2019), respectively; 36 ing of the hosts, photographing of hosts alive and 16 of these records pertain to the endo- underwater, and photographing of host skel- parasitic copepod symbionts belonging to the etons along with basic locality data are impor- family Corallovexiidae. Most records from tant for both identification and maintaining holothurians from the families Holothuri- information about each copepod host and the idae and Stichopodidae (75 and 35 records), establishment of base line information about from ophiuroids the families Ophiocomi- their distributions (Hoeksema et al., 2011). dae and Ophiotrichidae (29 and 19 records) In total, there are 253, 197 and 81 records of are most common, from sea cucumbers the copepods found associated with echinoderms, family Stichopodidae (35 records) is well cnidarians and sponges, respectively (figs. 1–3). represented, alcyonaceans the family Plex- Forty-seven and 20 records of copepods are auridae (29), and corallimorpharians (four linked to the scleractinian coral families Favii- records) also are represented (table 3). Many Downloaded from Brill.com02/21/2020 01:34:50PM via Universiteit of Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 333

FIGURE 4 Caribbean copepods found on sponges (x; x = number of records and number of copepod species, respectively).

FIGURE 5 Caribbean copepods found on octocorals (Anthozoa: Octocorallia) (x; x – number of records and num- ber of copepod species, respectively). Downloaded from Brill.com02/21/2020 01:34:50PM via Universiteit of Groningen

334 KORZHAVINA et al.

FIGURE 6 Caribbean copepods found on hexacorallians (Anthozoa: Hexacorallia) (x; x – number of records and number of copepod species, respectively).

FIGURE 7 Caribbean copepods found on echinoderms (x; x – number of records and number of copepod species, respectively). Downloaded from Brill.com02/21/2020 01:34:50PM via Universiteit of Groningen

COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 335

Western Western Caribbean × × × × × Southwestern Southwestern Caribbean Southern Caribbean × × × × × × × × × × ×

Greater Greater Antilles × × × × × × × × × × × × × × × Floridian ×

Eastern Eastern Caribbean × × Bermuda × × × × Bahamian × × Varela, 2010 Varela, Stock, 1975 Stock, Humes, 1973 Humes, Stock, 1975 Stock, Stock, 1975 Stock,

Humes, 1969 Humes, Humes, 1973 Humes, Humes, 1973 Humes,

Stock, 1975 Stock,

Stock, 1975 Stock, Kim, 2010 Stock, 1975 Stock, Humes & Stock, 1973 Humes & Stock, Humes, 2000 Humes, Stock, 1975 Stock,

Varela, 2010 Varela, Stock, 1975 Stock, Humes, 1973 Humes, Varela, Ortiz & Lalana, 2007 Varela,

Stock, 1975 Stock, Varela, 2010 Varela, Varela et al., 2007 Varela Varela, 2013 Varela, Varela, 2012 Varela,

sp. Symbiotic copepods recorded on reef-dwelling anthozoans, echinoderms and sponges of the Greater Caribbean. The Gulf of echinoderms and sponges Caribbean. anthozoans, of the Greater on reef-dwelling and copepods recorded has no records Symbiotic Mexico is not included

TABLE 6 Copepod species / Ecoregion aequiseta Acanthomolgus affinis Acanthomolgus bayeri Acanthomolgus bilobipes Acanthomolgus dionyx Acanthomolgus gorgoniae Acanthomolgus intermedius Acanthomolgus longidactylus Acanthomolgus longifurca Acanthomolgus mononyx Acanthomolgus muriceanus Acanthomolgus seticornis Acanthomolgus triangulipes Acanthomolgus verrucipes Acanthomolgus Aspidomolgus stoichactinus Asterocheres antillensis Asterocheres bahamensis Asterocheres crinoidicola Asterocheres cubensis Asterocheres espinosai Asterocheres fernandezmilerai Asterocheres garridoi Asterocheres kimi Asterocheres lalanai Asterocheres

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336 KORZHAVINA et al.

Western Western Caribbean × × × × Southwestern Southwestern Caribbean Southern Caribbean × × × × × × × × ×

Greater Greater Antilles × × × × × × × × × × × × Floridian

Eastern Eastern Caribbean × × × Bermuda Bahamian × × × × × × × ) Varela, Ortiz & Lalana, Varela, cont. Stock, 1975 Stock, Stock, 1975 Stock, Kim, 2010 Stock, 1975 Stock,

Stock & Humes, 1995 & Humes, Stock Kim, 2010 Stock, 1987 Stock, Stock, 1987 Stock,

Kim, 2010 Suárez-Morales, 1998 Suárez-Morales, Kim, 2010 Kim, 2010 Kim, 2010 Stock, 1987 Stock,

(T. Scott, 1898) Scott, (T. Stock, 1989 Stock, Humes & Goenaga, 1978 Humes & Goenaga,

Boxshall & Huys, 1994 & Huys, Boxshall 1994 & Huys, Boxshall Stock, 1989 Stock,

Stock & Humes, 1995 & Humes, Stock

(Humes, 1969) (Humes, sp. A sp. Symbiotic copepods recorded on reef-dwelling anthozoans, echinoderms and sponges of the Greater Caribbean. The Gulf of echinoderms and sponges Caribbean. anthozoans, of the Greater on reef-dwelling and copepods recorded has no records Symbiotic Mexico is not included (

TABLE 6 Copepod species / Ecoregion maxillatus Asterocheres reginae Asterocheres oricurvus Asterocheres peniculatus Asterocheres plumosus Asterocheres reginae Asterocheres simulans Asterocheres trisetatus Asterocheres unioviger Asterocheres foliata Asteropontella capillatus Asteropontius iuxtus Asteropontius longipalpus Asteropontius mycetophylliae Asteropontius 2005, 2005 parvipalpus Asteropontius proximus Asteropontius ungellatus Asteropontius faviae Asteropontopsis imparipes Calonastes Caribeopsyllus chawayi Caribeopsyllus sculptus Caribulus longipalpus Chelacheres optans Chelacheres

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 337 × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × × ) ) Stock, 1975 Stock, Kim, 2009

Stock, 1975 Stock, Stock, 1975 Stock, Stock, 1975 Stock, Humes & Stock, 1973 Humes & Stock, Stock, 1975 Stock, Stock, 1975 Stock, ( Stock, 1975 Stock, Stock, 1968 Stock, Humes & Ho, 1970 Humes & Ho,

Stock, 1973 Stock,

Stock, 1975 Stock,

Stock, 1973 Stock, Stock, 1975 Stock,

Stock, 1973 Stock, Edwards, 1891 Edwards, Stock, 1968 Stock,

Stock, 1968 Stock, Humes, 1982 Humes,

Stock, 1973 Stock, Stock, 1968 Stock, Humes, 1998 Humes,

Humes & Ho, 1971 Humes & Ho,

(

Stock, 1975 Stock, Humes & Ho, 1970 Humes & Ho, Stock, 1975 Stock,

sp. sp. Collocheres lunulifer Collocheres baki Corallonoxia longicauda Corallonoxia brevibrachium Corallovexia dorsospinosa minor Corallovexia dorsospinosa Corallovexia kristenseni Corallovexia longibrachium Corallovexia mediobrachium Corallovexia mixtibrachium Corallovexia similis Corallovexia ventrospinosa Corallovexia Critomolgus astrophyticus Critomolgus titillans Critomolgus deichmannae Diogenella impar Diogenella seticauda Diogenella spinicauda Diogenella deforme Diogenidium nasutum Diogenidium spinulosum Diogenidium tectum Diogenidium Enalcyonium euniceae Enalcyonium nudum Enalcyonium ramosum Enalcyonium variicauda Enalcyonium occidentalis Eupolymniphilus

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338 KORZHAVINA et al.

Western Western Caribbean × × × Southwestern Southwestern Caribbean Southern Caribbean × × × × × × × ×

Greater Greater Antilles × × × × × × × × × × × × Floridian × × ×

Eastern Eastern Caribbean × × Bermuda × × Bahamian × × × × ) ) ) Stock, 1989 Stock,

cont. Stock, 1987 Stock,

Humes & Stock, 1973 Humes & Stock, Stock, 1978 Stock,

Varela, 2012 Varela,

Humes, 1984 Humes, Kim, 2010 Kim, 2010

Kim, 2010

(

Stock, 1987 Stock, Stock, Humes & Gooding, Stock,

Kim, 2010 Stock, Humes & Gooding, Stock, Sars, 1916 Sars, Stock & Gooding, 1986 Stock Stock, 1979 Stock, (

Stock, 1973 Stock, Stock, Humes & Gooding, 1963 Stock,

Humes & Hendler, 1999 Humes & Hendler, Stock, Humes & Gooding, 1962 Stock,

Stock, Humes & Gooding, 1962 Stock, Stock, Humes & Gooding, 1962 Stock, Stock, Humes & Gooding, 1962 Stock, Symbiotic copepods recorded on reef-dwelling anthozoans, echinoderms and sponges of the Greater Caribbean. The Gulf of echinoderms and sponges Caribbean. anthozoans, of the Greater on reef-dwelling and copepods recorded has no records Symbiotic Mexico is not included (

TABLE 6 Copepod species / Ecoregion columnaris Hemicyclops diploriae Hermacheres montastreae Hermacheres aculeata Hetairosynella angulata Hetairosynella bifurcata Hetairosynella fastigatus Kimcheres Lamippina aequalis bouligandi Linaresia echinicolum Macrochiron sargassi Macrochiron Magnippe caputmedusae coronatum Meandromyzon amplectans Meomicola 1963 glaber Micropontius exigua Nanaspis media Nanaspis pollens Nanaspis truncata Nanaspis humesi Neoasterocheres alatus Onychocheres latus Ophiopsyllus reductus Ophiopsyllus 1963

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 339 × × × × × × × × × × × × × × × × × × × × × × × ×

Humes & Hendler, 1972 Humes & Hendler, Stock & Humes, 1995 & Humes, Stock

Stock, Humes & Stock, Humes & Stock, Stock, Humes & Gooding, Stock, Kim, 2009 Kim, 2010

Humes & Hendler, 1972 Humes & Hendler,

Humes, 1998 Humes, Stock, 1975 Stock, Stock, 1992 Stock, Humes & Smith, 1974

(Scott T. & Scott A., 1894) & Scott T. (Scott (Johnsson, 1998) Wilson C.B., 1944 Wilson C.B., Stock, 1975 Stock, Grygier, 1980 Grygier,

Varela, 2011 Varela, Varela & Lalana, 2007 Varela

sp. Ophiurocheres bellulus Ophiurocheres antillensis Orecturus ortizi Orecturus antillianus Paramolgus emarginata Parmulella verrucosa Parmulodes ligatus Parophiopsyllus rostratum Peltomyzon amphiopli Presynaptiphilus acutus Pseudanthessius deficiens Pseudanthessius 1963 exilicornis Pseudanthessius pectinifer Pseudanthessius Gooding, 1964 pectinifer Pseudanthessius Gooding, 1964 fosshageni Ridgewayia elongatus Scottocheres paraboecki Setacheres caligicola Sphaerippe Sphaerippe edentatum Stenomyzon

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340 KORZHAVINA et al.

TABLE 7 Numbers of reef-dwelling species of Caribbean anthozoans, sponges, and echinoderms recorded as host for copepods

Taxa Estimated number of Number of species Percent of invertebrate host species associated with total species copepods

Subclass Hexacorallia (1) 45 21 47% Subclass Octocorallia (2) 195 18 9% Phylum Echinodermata (3, 4) 955 31 3% Phylum Porifera (5) 760 9 1% Total number 1955 79 4%

1 – Ivanenko, 2016, 2 – Bayer, 1961, 3 – Alvarado, 2011, 4 – Pawson et al., 2009, 5 – Perez et al., 2017

(25 of 81) records of copepods found on un- families of stony corals only. Three families identified sponges. (Lamippidae, Pseudanthessiidae, and Synap- Cyclopoida, representing mainly poecilo- tiphilidae) are found associated with four host stomatoids, with 317 records for 59 species is families; Macrochironidae and Nanaspididae the most diverse order of copepods found in are found with three host families; Canceril- symbiosis with corals, sponges and echino- lidae and Lichomolgidae are found with two derms; Siphonostomatoida with 170 records host families; seven families (Clausidiidae, of 55 species is the next (figs. 1–3, table 4). The Entomolepididae, Micropontiidae, Pseudo- order Calanoida is represented by the only cyclopidae, Sabelliphilidae, Synaptiphilidae, known symbiotic calanoid copepod Ridgeway- and Thaumatopsyllidae) are restricted to only ia fosshageni found associated with an actini- one family of invertebrate hosts (table 5). The arian at the Atlantic coast of Panama (Humes remarkable absence on Caribbean host cor- & Smith, 1974). The absence in the literature of als for copepods of the families Anchimolgi- any records of symbiotic harpacticoids is con- dae (124 species in 32 genera) and Xarifidae tradicted by the results recently obtained from (96 species in 6 genera) so far only found on samples of undescribed harpacticoids repre- Indo-Pacific scleractinians (Stock, 1988), is senting the family Laophontidae (see Yeom confirmed by literature data and results of our et al., 2018). Siphonostomatoid copepods of recent sampling (Cheng et al., 2016; Hoekse- the diverse but poorly investigated family As- ma et al., 2017b; table 1). To explain this distri- terocheridae and poecilostomatoid cyclopoids bution a study of phylogenetic relationships representing endoparasitic Corallovexiidae of Corallovexiidae with other families of the have the greatest diversity of associations and order and the additional search for endopar- the highest number of host families (table 5). asitic copepods living in Indo-Pacific stony Asterocheridae are found living on inverte- corals is needed. This proposed study should brates belonging to 22 families of cnidarians, include methods that enable dissolution only of echinoderms and sponges; Rhynchomolgidae soft coral tissue while the chitinous exoskel- are recorded from eight families of anthozoans etons of microscopic crustaceans to remain and echinoderms; Corallovexiidae from five intact.

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COPEPODS ASSOCIATED WITH CARIBBEAN INVERTEBRATES 341

Fifty of 115 species of symbiotic copepods six. The sea cucumber Holothuria arenicola are mentioned in literature only once; 83 of hosts six species of copepods (table 2). Three 115 species of copepods are reported from only scleractinian and two holoturian species host one species or one genus of the host. Only 13 five copepod species. The sea urchin Meoma species are reported in symbiosis with repre- ventricosa and the sea cucumber Actinopyga sentatives of different families. Four species agassizii host four copepod species; three cor- of copepods, the poecilostomatoids Eupolym- als and one sponge host three species each; niphilus occidentalis (fig. 1g), Hemicyclops co- 18 host species have been recorded with two lumnaris, Pseudanthessius deficiens (fig. 1e) associated species of symbiotic copepods. Nu- and the siphonostomatoid Nanaspis pollens merous findings of different species of symbi- (fig, 3e) are found in symbiosis with represen- otic copepods on the same hosts as well as the tatives of different classes. Of these species presence on these hosts of other symbionts only Hemicyclops columnaris is found associ- (shrimps, polychaetes, decapods, amphipods, ated with invertebrates representing differ- fish etc.) shows little knowledge of species re- ent phyla: echinoderms, corals, sponges and lationships in the symbiotic complexes (Stella . The finding on sponge and on a et al., 2011; Hoeksema et al., 2012). compound ascidian of the copepod Eupolym- Copepod crustaceans living in symbiosis niphilus occidentalis (family Sabelliphilidae) with the Caribbean reef invertebrates express typically living on tubicolous polychaetes re- diverse body shapes (cyclopiform, spherical, quires additional confirmation (Kim, 2009). flattened etc.), remarkably different body siz- Only two species of copepods are reported liv- es ranging from 0.25 mm (such as Collocheres ing on different classes: Pseudanthessius defi- lunulifer) to 4.75 mm (such as Corallovexia ciens is found on three species of echinoderms ventrospinosa), and various types of feeding to holothurians and echinoids; Nanaspis pol- apparatuses as well as host utilizations. Anal- lens is found on holothurians and ophiurioids. ysis of literature and sorting of samples shows Thus, most species of symbiotic copepods are a different, and sometimes very large number found associated with invertebrate hosts be- of individual copepods living on a single host. longing to a single genus or only one family. The most numerous are the poorly studied These finding suggest the need to study the ef- asterocherid copepods living on and in di- fect of host specificity more extensively. Simi- verse sponges: in one sample on one sponge lar studies of host switching events may show thousands of individual copepods belonging that this phenomenon has occurred several to several species (Schirl, 1973; Ivanenko & times during in the evolution of symbiotic co- Smurov, 1997; Ivanenko, 1998; Klinger et al., pepods, as it has among decapod 2019; present study). The diversity, host speci- taxa (Fransen & Hoeksema, 2014; Brinkmann ficity and phylogenetic relationships of these & Fransen, 2016; García-Hernández et al., copepods with other siphonostomatoid cope- 2016; Horká et al., 2016; Hoeksema & Fransen, pods, especially species found in association 2017; Hoeksema et al., 2018). The very nature with Caribbean stony corals are among the of the specificity of copepods to the host or to most interesting unexplored topics in coral the group of hosts requires a thorough sam- reef ecology. pling program as well as the use of molecular The symbiotic copepods are characterized methods (Ivanenko et al., 2018). by different ways of attachment to their host The number of copepod species found (loosely associated with or aggregating on on a single host species varies from one to the host’s surface, tightly attached to the host

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342 KORZHAVINA et al. by claw-like appendages, inducing a gall-like species of octocorals, 3% of echinoderms and structure, or residing inside of intestine, body of less than 1% species of sponges (table 7). cavity, or the host tissues). They also show They are found on 4% of potential hosts marked variation in the number of embryos of the Caribbean invertebrates which cor- present in the egg-sacs (from one embryo, as responds with previous data on symbiotic in Peltomyzon rostratum, to 50 embryos, as in copepods (Humes, 1994). The literature and Caribulus sculptus) or numerous embryos laid samples analyses indicate a poor knowledge in copepod-induced galls. The symbiotic co- of the diversity and distribution of symbiotic pepods have different types of lecithotrophic copepods living on different hosts in various and planktotrophic naupliar stages, some of ecoregions, and a particularly poor knowl- which, like Thaumatopsyllidae, are parasitic edge of copepods living on corals and sponges in the stomach of its host brittle star (fig. 2e; (Boxshall & Huys, 1994; García-Hernández Hendler & Kim, 2010). et al., 2019; present study) with unknown but Most of the ecological observations of the potentially high impact on their host and reef Caribbean symbiotic copepods are short com- community (Ho, 2001; Berkenbusch & Row- ments added to the taxonomic descriptions den, 2003; Hatcher et al., 2012; Shelyakin et al., and describe behavioral features and/or loca- 2018; Zeppilli et al., 2015, 2018). tion on the host. Exceptions are the study of Ophiopsyllus reductus parasitizing on shallow- water ophiuroids (Emson & Mladenov, 1987; Acknowledgements Emson et al., 1985), the ecological observations of the endoparasitic copepods of the family The preparation of the database, sampling in Corallovexiidae living in stony corals (Butter, Cuba and processing of the paper were sup- 1979; Herriott & Immermann, 1979), the stud- ported by the Russian Foundation for Basic ies of the life cycle and ecology of copepods Research (grant 18-54-34007). The data on of the family Thaumatopsyllidae parasitizing samples from St. Eustatius (2015) and Cura- living in the ophiuroids at naupliar stages and çao (2017) were obtained with support of a having non-feeding adult and subadult cope- Temminck-Fellowships to VNI by Naturalis podid stages (Suarez-Morales & Castellanos, Biodiversity Center. BWH and VNI want to 1998; Suarez-Morales & Tovar, 2004; Hendler thank staffs of the Caribbean Netherlands Sci- & Kim, 2010; Ferrari et al., 2010; Ferrari & von ence Institute (CNSI) at St. Eustatius and the Vaupel Klein, 2019), and the experiments on Carmabi Research Station at Curaçao for hos- calanoid copepods that were discovered in a pitality and assistance. Sorting of samples and host-specific association with only one spe- identification of copepods were conducted cies of actinarian (Humes & Smith, 1974). Re- with partial support of the Russian Founda- cent field trips to sample Caribbean symbiotic tion for Basic Research (grant 18-04-01192). copepods lead to the discovery of new species Jaaziel Emmanuel Garcia-Hernandez (Uni- of highly transformed gall-inducing copepods versity of Puerto Rico at Mayagüez) provided of the genus Sphaerippe (Lamippidae) caus- identification of sponges collected during ing the Multiple Purple Spot Syndrome previ- the field trips; Frank Ferrari commented on a ously found in Gorgonia ventalina (Ivanenko draft of the manuscript; Maickel Armenteros et al., 2017; Tracy et al., 2018). (University of Havana) hosted VNI and OAK The Caribbean symbiotic copepods are during the field trip to Cuba. found on 47% species of scleractinians, 9%

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