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Marine Biodiversity

, Volume 41, Issue 2, pp 261–285 | Cite as

Echinoderm diversity in the Caribbean Sea

  • Juan José AlvaradoEmail author
Open Access
Original Paper

Abstract

The Caribbean is considered a unique biogeographic province, being one of the top five hotspots in the world for marine and terrestrial biodiversity. The echinoderm research on the Caribbean began in the middle of the nineteenth century, and during the first half of the twentieth century most of the species were described. The number of ecological studies increased in the 1970s that continues until today, mostly focused on Diadema antillarum. Based on an extensive review of published records, the Caribbean echinoderm fauna is composed of 433 species, 237 genera, 80 families, 29 orders in five classes, with four endemic species. The richest class is Ophiuroidea with 148 species followed by the class Asteroidea with 116 species. Mexico and Colombia are the richest countries with 182 and 180 species respectively, while Costa Rica and Guatemala are the less diverse with fewer than 50 species. In general terms, the Caribbean is very homogeneous in species composition; however, Colombia and Mexico are the countries more dissimilar in composition with respect to the rest of countries. This semi-enclosed sea represents 6.5% of the total diversity of the phylum worldwide, and is the second most diverse area in tropical America, after Gulf of Mexico. Six species of echinoderms are extracted for commercial use, mostly in Panama, Nicaragua, Colombia, Venezuela and Cuba. However, extraction for the souvenir trade, without any regulation and control, is one of the major threats for some animals—especially the sea star Oreaster reticulatus. Research priorities for this group in the Caribbean include exploration of the deep sea, the Central American region and the Lesser Antilles. Moreover, because of its rich biodiversity, research and conservation efforts have to be directed towards its preservation, avoiding illegal extraction, enforcing controls, and improving coastal zone management.

Keywords

Echinoderms Diversity Richness Similarity Historic account Caribbean 

Introduction

The Caribbean Sea is a large sea, closed off to the west and to the south by the Americas, and partly screened to the north and to the east by the island chains of the Greater and Lesser Antilles (Spalding 2004). The region extends over about 2,754,000 km2 in which 36–40 politically independent countries and territories can be found (Miloslavich and Klein 2005). Two large currents sweep towards this region from the Atlantic, the North Equatorial Current and Guyana current. A portion of their flow runs between the islands and into the Caribbean Sea, forming a generally westward flow known as the Caribbean current. The rest of the current is deflected to the northwest, fusing with the Gulf Stream (Spalding 2004).

The Caribbean is considered a unique biogeographic region. It is among the top five hotspots in the world for marine and terrestrial biodiversity, and it has many endemic species (Rivera-Monroy et al. 2004). Its complex geological history, begining 130 million years ago (Ma) and involving emergence of the Isthmus of Panama in the Pliocene (around 3.0–2.8 Ma), had major effects on the marine biodiversity. The separation of the tropical American ocean into two different realms produced isolation and environmental change that resulted in increased evolutionary divergence and radiation of species in extensive coral reefs, mangroves, seagrass beds, deep-shelf ecosystems and partially isolated deep basins and trenches (Collins 1996).

The echinoderm research on the Caribbean began in the middle of the nineteenth century with dredging explorations in Florida and Cuba (Agassiz 1869; Lyman 1869). The first half of the twentieth century increased the number of species described with collections from Puerto Rico (Clark 1901), the Lesser Antilles (Engel 1933), Jamaica (Fointaine 1953a, b, c), and the publication of monographs on groups such as Holothuroidea (Deichmann 1926, 1930, 1940, 1963). During the second half of the twentieth century research increased the number of local records in Cuba (Suárez 1974), Belize (Devaney 1974; Kier 1975; Hotchkiss 1982; Macurda 1982; Hendler and Pawson 2000), Panama (Chesher 1972), Venezuela (Zoppi de Roa 1967; Rodríguez 1969, 1973; Martínez and Mago 1975; Martínez 1987, 1989), Jamaica (Meyer 1973), Honduras (Lessios 1998; Hasbún and Lawrence 2002). During this period information on groups such as Asteroidea (Downey 1973; Clark and Downey 1992) and Crinoidea (Meyer et al. 1978) was augmented. This increase in research is partially a consequence of the work in Panama conducted by the Smithsonian Tropical Research Institution and research in other parts of the Caribbean by the Smithsonian Institution in Washington DC.

During the 1970s, there was an increase in ecological studies that continues to our times, mostly focused on Diadema antillarum (Ogden et al. 1973; Sammarco 1980, 1982a, b; Lessios et al. 1984; Lessios 2005; Weil et al. 2005; Mumby et al. 2006; Steiner and Williams 2006; among others). However, one of the most important works was the compilation by Hendler et al. (1995), containing keys, pictures and a detailed description of 144 species. It also contained general comments on their habitat, distribution and biology. In addition to taxonomy, research in the region has focused on topics such as bioerosion (Scoffin et al. 1980; Bak et al. 1984; Griffin et al. 2003; Brown-Saracino et al. 2007), biodiversity (Price et al. 1999), reproduction (Lessios 1981a; Guzmán and Guevara 2002a, b; Guzmán et al. 2003; Montealegre-Quijano and Gómez-Gaspar 2005), aquaculture (Buitrago and Lodeiros-Seijo 2005; Gómez and Gómez 2005), evolution (Lessios 1981b; Lessios et al. 2001; Lessios et al. 2003; Zigler and Lessios 2004), and competition (Parker and Shulman 1986). The majority of this research was focused on echinods. The Caribbean region likely ranks among the top tropical regions for echinoderms research.

More recently, more research has being carried out, in Colombia (Benavides-Serrato and Borrero-Pérez 2000; Benavidez-Serrato et al. 2005; Borrero-Pérez et al. 2002a, b, 2008), Cuba (Abreu-Pérez et al. 2005; Del Valle-García et al. 2005, 2008), Mexico (Laguarda-Figueras et al. 2005), Hispaniola Island (Herrera-Moreno and Betancourt 2004) and Puerto Rico (Benavides-Serrato 2006). In the past decade scientist from the Instituto de Investigaciones Marinas y Costeras (INVEMAR) have publish most on echinoderm diversity in both shallow and deep water habitats.

The aim of this review is to present an updated list of the echinoderms in the Caribbean region based on information from the literature and museum collections. A list of species is presented indicating their presence in each country, the similarities in species composition, and the possible causes of those patterns of distribution.

Material and methods

I reviewed the echinoderm species composition of the Caribbean coast from the Yucatán Peninsula in Mexico (Quintana Roo State) to Venezuela, including the arc of the Caribbean islands from Cuba to Trinidad & Tobago (Fig. 1). The ecoregions covered by this review included the Eastern Caribbean, Greater Antilles, Southern Caribbean, Southwestern Caribbean and Western Caribbean of the Tropical Northwestern Atlantic Province (Spalding et al. 2007). Existing published echinoderm studies were reviewed (Zoppi de Roa 1967; Chesher 1968, 1970; Flores and Martínez 1970; Serafy 1970; Meyer 1973; Devaney 1974; Martínez and Mago 1975; Meyer and Macurda 1976; Meyer et al. 1978; Caycedo 1978, 1979; Alvarez-Larrauri 1981; Hotchkiss 1982; Macurda 1982; Miller and Pawson 1984; Martínez 1987; Hendler 1988, 1995; Clark and Downey 1992; Hendler et al. 1995; Schoppe 1996; Lessios 1998; Benavides-Serrato and Borrero-Pérez 2000; Hendler and Pawson 2000; Pawson et al. 2001; Borrero-Pérez et al. 2002a, b; Gómez-Gaspar 2002; Gonzalez et al. 2002; Hasbún and Lawrence 2002; Fonseca and Arrivillaga 2003; Herrera-Moreno and Betancourt 2004; Abreu-Pérez et al. 2005; Benavidez-Serrato et al. 2005; Del Valle-García et al. 2005; Laguarda-Figueras et al. 2005; Benavides-Serrato 2006; Alvarado et al. 2008; Borrero-Pérez et al. 2008; Francisco and Pauls 2008), and extracted information from the Geology Collection of the California Academy of Sciences, San Francisco (http://www.calacademy.org/research/izg/iz_coll_db/index.asp). The taxa were validated with the Integrated Taxonomic Information System (http://www.itis.gov/), and the Ophiuroidea (Stöhr and O’Hara 2007) and Asteroidea databases (Mah 2009). For the taxonomic list, the criteria used in Alvarado et al. (2008) were applied.
Fig. 1

a Caribbean Sea countries and ecoregions. b Radial diagram of the number of species of echinoderms (axes) per country on the Caribbean. Gray circle countries with under the average number of species. Ecoregions: W Western Caribbean (blue dots), SW Southwestern Caribbean (green dots), GA Greater Antilles (red dots), E Eastern Caribbean (yellow dots), S Southern Caribbean (orange dots). PRC Puerto Rico, RDO Dominican Republic, HAI Haiti, JAM Jamaica, CUB Cuba, MEX Mexico, BEL Belize, GUA Guatemala, HON Honduras, NIC Nicaragua, CRC Costa Rica, PAN Panama, COL Colombia, VEN Venezuela, TYT Trinidad & Tobago, LESAN Lesser Antillers

I used the year of publication of the description of each currently valid species to examine patterns in the rate of species descriptions over time, and to construct species accumulation curves for the entire described fauna. I followed the criteria from Zapata and Robertson (2007), to predict the total fauna size using the logistic model in the PAST computer program.

Composition similarities for each country were estimated by a presence/absence matrix per class and for the complete phylum. With those matrixes, I applied a Euclidean distance similarity matrix and I compared the diversity of echinoderms with a cluster tree and non-metric multidimensional scaling (nMDS). I estimated the average taxonomic distinctness (Δ+) and its variation (Λ+) (Clarke and Warwick 2001). This index evaluated the taxonomic distance between each pair of individuals, defined by a Linnaean classification tree, and could be used with a presence/absence matrix without taking into consideration the sampling methodology used. This index is considered as one of the most precise indicators in a strict biodiversity sense (Clarke and Warwick 2001). It was used on five taxonomic levels: species, genus, family, order and class, with their respective weights ω = 20 (species in the same genera), 40 (same family but different genus), 60 (same order but different family), 80 (same class but different order) and 100 (different class). The statistical analysis was performed with the PRIMER 6.0 software.

Results

The list of echinoderms of the Caribbean includes 433 species, 237 genera, 80 families, 29 orders and five classes (Table 1), with four endemic species. The richest class was Ophiuroidea with 148 species, followed by the class Asteroidea with 116 species, then Echinoidea and Holothuroidea with 76 and 63 species respectively, and finally the class Crinoidea with 30 species (Table 2). Mexico and Colombia are the richest countries with 182 and 180 species respectively, followed by Panama with 155 species and Cuba with 145 species (Table 3). Eight countries possess between 100 and 182 species, six between 50 and 99 species, and only two countries possess less than 50 species, Costa Rica and Guatemala with 44 and 23 species respectively. There is an average of 105 species per country (Table 3). The average number of species per kilometer (spp./km) of coastline is 0.13 ± 0.07 spp./km, Belize being the richest country with 0.26 spp./km, and Cuba, Haiti, and Venezuela the poorest with 0.04 spp./km. The average number of spp./km2 of Economic Exclusive Zone (EEZ) is 0.0022 spp./km2 of EEZ. Based on this metric, Guatemala is the richest country with 0.0140 spp./km2 of EEZ, followed by Honduras (0.0052 spp./km2), and Venezuela and Jamaica are the poorest with 0.0003 spp./km2 of EEZ (Table 3).
Table 1

Taxonomic classification of the echinoderm fauna from Caribbean countries. PRC Puerto Rico, RDO Dominican Republic, HAI Haiti, JAM Jamaica, CUB Cuba, MEX Mexico, BEL Belize, GUA Guatemala, HON Honduras, NIC Nicaragua, CRC Costa Rica, PAN Panama, PAN Colombia, VEN Venezuela, TYT Trinidad & Tobago, LESAN Lesser Antillers, NCA no country assigned; X indicates presence

 

CUB

MEX

PRC

RDO

HAI

JAM

BEL

GUA

HON

NIC

CRC

PAN

COL

VEN

TYT

LESAN

NCA

Class Crinoidea

                 

Order Comatulida

                 

Family Colobometridae

                 

Analcidometra armata (Pourtàles, 1869)

  

X

X

 

X

X

 

X

 

X

X

     

Oligometra caribbea A.H. Clark, 1908

           

X

     

Family Comasteridae

                 

Comactinia echinoptera (Müller, 1840)

X

X

X

X

X

X

 

X

X

X

X

X

     

Comactinia meridionalis (L. Agassiz, 1865)

X

X

X

X

X

   

X

X

 

X

X

    

Davidaster discoidea (Carpenter, 1888)

X

 

X

X

 

X

X

X

X

X

X

X

X

  

X

 

Davidaster insolitus A.H. Clark, 1917

           

X

     

Davidaster rubiginosus (Pourtàles, 1867)

X

X

X

X

X

X

X

 

X

 

X

X

X

X

   

Leptonemaster venustus A.H. Clark, 1909

X

 

X

X

X

X

  

X

  

X

 

X

 

X

 

Nemaster grandis A.H. Clark, 1911

   

X

 

X

  

X

X

 

X

 

X

   

Neocomatella alata (Pourtàles, 1878)

 

X

               

Neocomatella pulchella (Pourtàles, 1867)

X

  

X

     

X

 

X

X

X

 

X

 

Family Charitometridae

                 

Crinometra brevipinna (Pourtàles, 1867)

 

X

 

X

 

X

   

X

   

X

X

X

 

Family Antedonidae

                 

Antedon bifida (Pennant, 1777)

             

X

   

Coccometra hagenii (Pourtàles, 1867)

X

                

Coccometra nigrolineata A.H. Clark, 1908

 

X

X

  

X

           

Ctenantedon kinziei Meyer, 1972

     

X

X

    

X

   

X

 

Hypalometra defecta (Carpenter, 1888)

   

X

       

X

     

Polimetra prolixa (Sladen, 1881)

   

X

             

Trichometra cubensis (Pourtàles, 1867)

X

   

X

    

X

  

X

    

Zenometra columnaris (Carpenter, 1881)

    

X

            

Family Atelecrinidae

                 

Atelecrinus balanoides Carpenter, 1881

X

    

X

      

X

 

X

X

 

Family Thalassometridae

                 

Stylometra spinifera (Carpenter, 1881)

   

X

 

X

X

 

X

    

X

   

Family Tropiometridae

                 

Tropiometra carinata (Lamarck, 1816)

   

X

         

X

X

X

 

Order Isocrinida

                 

Family Isocrinidae

                 

Endoxocrinus parrae (Gervais, 1835)

X

X

  

X

X

  

X

   

X

    

Cenocrinus asteriscus (Linnaeus, 1775)

   

X

             

Neocrinus decorus (Wyville-Thomson, 1864)

   

X

 

X

  

X

    

X

 

X

 

Order Cyrtocrinida

                 

Family Holopodidae

                 

Holopus rangii d'Orbigny, 1837

X

  

X

 

X

         

X

 

Order Millericrinida

                 

Family Bourgueticrinidae

                 

Democrinus brevis A.H. Clark, 1909

           

X

 

X

   

Democrinus conifer (A.H. Clark, 1909)

    

X

X

  

X

X

 

X

   

X

 

Democrinus rawsoni (Pourtàles, 1874)

   

X

     

X

 

X

   

X

 

Class Asteroidea

                 

Order Paxillosida

                 

Family Luidiidae

                 

Luidia alternata alternata (Say, 1825)

X

X

X

X

    

X

  

X

X

X

   

Luidia barbadensis Perrier, 1881

 

X

X

X

     

X

  

X

X

   

Luidia clathrata (Say, 1825)

X

X

X

X

  

X

 

X

X

 

X

X

X

   

Luidia heterozona barimae John & Clark, 1954

            

X

X

   

Luidia ludwigi scotti Bell, 1917

 

X

          

X

X

   

Luidia sarsi elegans Perrier, 1875

            

X

    

Luidia senegalensis (Lamarck, 1816)

X

 

X

X

X

X

  

X

X

X

X

X

X

 

X

 

Family Astropectinidae

                 

Astropecten acutiradiatus Tortonese, 1956

             

X

   

Astropecten alligator Perrier, 1881

   

X

    

X

X

  

X

    

Astropecten americanus Verrill, 1880

         

X

  

X

    

Astropecten antillensis Lütken, 1860

X

           

X

X

   

Astropecten articulatus (Say, 1825)

X

 

X

X

 

X

X

X

X

X

X

X

X

X

 

X

 

Astropecten brasiliensis Müller & Troschel, 1842

             

X

   

Astropecten caribemexicanensis Caso, 1990

 

X

               

Astropecten cingulatus Sladen, 1883

 

X

       

X

X

X

     

Astropecten comptus Verrill, 1915

 

X

               

Astropecten duplicatus Gray, 1840

X

X

X

X

 

X

X

 

X

X

    

X

  

Astropecten marginatus Gray, 1840

  

X

X

      

X

 

X

X

   

Astropecten nitidus Verrill, 1915

 

X

 

X

    

X

X

  

X

    

Blakiaster conicus Perrier, 1881

   

X

    

X

        

Dipsacaster antillensis Halpern, 1968

            

X

    

Persephonaster echinulatus H.L. Clark, 1941

 

X

      

X

X

 

X

X

    

Persephonaster leptactis H.L. Clark, 1941

X

                

Persephonaster patagiatus H.L. Clark, 1941

    

X

X

   

X

 

X

 

X

   

Plutonaster agassizi agassizi (Verrill, 1880)

            

X

    

Plutonaster efflorescens (Perrier, 1884)

X

 

X

  

X

       

X

   

Psilaster cassiope Sladen, 1889

            

X

X

   

Tethyaster grandis (Verrill, 1899)

 

X

 

X

        

X

    

Tethyaster vestitus (Say, 1825)

 

X

         

X

X

X

   

Family Goniopectinidae

                 

Goniopecten demonstrans Perrier, 1881

     

X

X

    

X

X

X

   

Prionaster elegans Verrill, 1899

 

X

       

X

  

X

X

   

Family Porcellanasteridae

                 

Styracaster elongatus Koehler, 1907

                

X

Thoracaster cylindratus Sladen, 1883

 

X

X

          

X

   

Order Notomyotida

                 

Family Benthopectinidae

                 

Benthopecten simplex simplex (Perrier, 1884)

 

X

  

X

           

X

Cheiraster (Barbadosaster) echinulatus (Perrier, 1875)

 

X

 

X

  

X

  

X

  

X

X

   

Cheiraster (Cheiraster) planus Verrill, 1915

 

X

   

X

  

X

   

X

 

X

  

Cheiraster (Cheiraster) sepitus (Verrill, 1885)

X

           

X

X

   

Cheiraster (Christopheraster) blakei A.M. Clark, 1981

X

X

   

X

X

 

X

   

X

    

Cheiraster (Christopheraster) mirabilis (Perrier, 1881)

X

 

X

  

X

   

X

  

X

    

Pectinaster gracilis Verrill, 1915

        

X

        

Order Valvatida

                 

Family Poraniidae

                 

Marginaster pectinatus Perrier, 1881

X

X

          

X

    

Family Asterinidae

                 

Asterina folium (Lütken, 1859)

 

X

X

   

X

    

X

X

X

   

Asterina hartmeyeri Döderlein, 1910

X

X

X

  

X

        

X

X

 

Asterinopsis pilosa (Perrier, 1881)

               

X

 

Paxillasterina pompom A.M. Clark, 1983

           

X

     

Stegnaster wesseli (Perrier, 1875)

     

X

      

X

X

X

X

 

Leilaster radians (Perrier, 1881)

 

X

               

Family Goniasteridae

                 

Anthenoides piercei Perrier, 1881

X

X

 

X

  

X

  

X

  

X

X

X

X

 

Apollonaster yucatanensis Halpern, 1970

 

X

          

X

    

Astroceramus brachyactis H.L. Clark, 1941

X

           

X

X

   

Ceramaster grenadensis (Perrier, 1881)

   

X

X

   

X

X

       

Circeraster americanus (A.H. Clark, 1916)

 

X

       

X

    

X

  

Cladaster rudis Verrill, 1899

                

X

Diplasiaster productus (A.H. Clark, 1917)

             

X

   

Floriaster maya Downey, 1980

 

X

               

Goniaster tessellatus (Lamarck, 1816)

 

X

           

X

   

Litonotaster intermedius (Perrier, 1884)

    

X

            

Nymphaster arenatus (Perrier, 1881)

 

X

  

X

   

X

X

 

X

X

X

   

Paragonaster grandis H.L Clark, 1941

X

X

               

Peltaster placenta (Müller & Troschel, 1842)

X

   

X

            

Plinthaster dentatus (Perrier, 1884)

        

X

X

 

X

X

X

X

  

Pseudarchaster gracilis gracilis (Sladen, 1889)

 

X

 

X

X

  

X

X

X

  

X

    

Rosaster alexandri (Perrier, 1881)

X

           

X

    

Tosia parva Perrier, 1881

 

X

 

X

        

X

    

Family Oreasteridae

                 

Oreaster reticulatus (Linnaeus, 1758)

X

X

X

X

X

 

X

X

X

X

X

X

X

X

   

Family Asteropsidae

                 

Poraniella echinulata (Perrier, 1881)

X

 

X

   

X

   

X

X

   

X

 

Family Mithrodiidae

                 

Mithrodia clavigera (Lamarck, 1816)

X

X

       

X

       

Family Ophidiasteridae

                 

Copidaster lymani A.H. Clark, 1948

X

     

X

    

X

     

Linckia bouvieri Perrier, 1875

X

X

               

Linckia guildingi Gray, 1840

X

X

X

X

X

X

X

X

  

X

X

 

X

   

Linckia nodosa Perrier, 1875

 

X

               

Narcissia trigonaria Sladen, 1889

 

X

         

X

     

Ophidiaster bullisi (Downey, 1970)

               

X

 

Ophidiaster guildingi Gray, 1840

X

X

X

X

  

X

    

X

   

X

 

Tamaria floridae (Perrier, 1881)

      

X

          

Tamaria halperni Downey, 1971

 

X

  

X

       

X

    

Order Velatida

                 

Family Caymanostellidae

                 

Caymanostella spinimarginata Belyaev, 1978

    

X

X

      

X

    

Family Solasteridae

                 

Laetmaster spectabilis (Perrier, 1881)

X

                

Lophaster furcifer (Düben & Koren, 1846)

X

                

Solaster caribbaeus Verrill, 1915

               

X

 

Solaster notophrynus Downey, 1971

               

X

 

Family Pterasteridae

                 

Calyptraster coa Sladen, 1882

               

X

 

Calyptraster personatus (Perrier, 1894)

    

X

       

X

  

X

 

Hymenaster lamprus H.L. Clark, 1923

     

X

           

Hymenaster rex Perrier, 1885

        

X

        

Pteraster abyssorum (Verrill, 1895)

    

X

X

  

X

        

Pteraster acicula Downey, 1970

X

          

X

X

    

Pteraster militarioides (O.F. Müller, 1776)

X

 

X

      

X

  

X

  

X

 

Pteraster personatus Sladen, 1891

            

X

    

Pteraster rugosus H.L. Clark, 1941

X

 

X

            

X

 

Family Korethrasteridae

                 

Remaster palmatus (Perrier, 1881)

X

     

X

        

X

 

Order Spinulosida

                 

Family Echinasteridae

                 

Echinaster brasiliensis Müller & Troschel, 1840

X

       

X

  

X

     

Echinaster echinophorus (Lamarck, 1816)

X

 

X

X

 

X

   

X

   

X

   

Echinaster guyanensis A.M. Clark, 1992

   

X

X

X

 

X

X

X

 

X

X

X

   

Echinaster modestus Perrier, 1881

 

X

X

X

 

X

X

  

X

   

X

 

X

 

Echinaster sentus (Say, 1825)

X

X

X

 

X

X

X

 

X

        

Echinaster serpentarius Müller & Troschel, 1842

 

X

          

X

    

Echinaster spinulosus Verrill, 1869

      

X

 

X

X

 

X

     

Henricia antillarum (Perrier, 1881)

 

X

   

X

  

X

X

     

X

 

Henricia downeyae A.M. Clark, 1987

    

X

            

Henricia sexradiata (Perrier, 1881)

                

X

Order Forcipulatida

                 

Family Asteriidae

                 

Sclerasterias contorta (Perrier, 1881)

             

X

 

X

 

Sclerasterias tanneri (Verrill, 1895)

        

X

        

Stephanasterias albula (Stimpson, 1853)

            

X

    

Family Labidiasteridae

                 

Coronaster briareus (Verrill, 1882)

 

X

          

X

    

Family Pedicellasteridae

                 

Pedicellaster pourtalesi Perrier, 1881

         

X

       

Family Zoroasteridae

                 

Doraster constellatus Downey, 1970

        

X

  

X

X

    

Mammaster sigsbeei (Perrier, 1894)

        

X

   

X

  

X

 

Zoroaster fulgens Wyville-Thomson, 1873

    

X

      

X

X

    

Order Brisingida

                 

Family Brisingidae

                 

Brisinga costata Verrill, 1888

    

X

        

X

   

Brisinga cricophora Sladen, 1889

               

X

 

Midgardia xandaros Downey, 1972

        

X

        

Novodinia pandina (Salden, 1889)

X

                

Novodinia antillensis (A.H. Clark, 1934)

  

X

              

Stegnobrisinga splendens H.L. Clark, 1926

        

X

    

X

   

Family Freyellidae

                 

Colpaster scutigerula Sladen, 1889

        

X

        

Class Ophiuroidea

                 

Order Phrynophiurida

                 

Family Ophiomyxidae

                 

Ophiomyxa flaccida (Say, 1825)

X

X

X

 

X

 

X

X

X

  

X

X

X

X

X

 

Ophiomyxa brevicauda Verrill, 1899

X

                

Ophiomyxa stimpsonii (Lyman, 1875)

            

X

    

Ophiomyxa tumida Lyman, 1883

            

X

    

Ophiobyrsa serpens Lyman, 1883

      

X

          

Ophiosyzygus disacanthus H.L. Clark, 1911

            

X

    

Ophioblenna antillensis Lütken, 1859

  

X

   

X

    

X

     

Family Asteronychidae

                 

Asteronyx loveni Müller & Troschel, 1842

 

X

           

X

   

Family Asteroschematidae

                 

Asterochema intectum Lyman, 1878

 

X

               

Asterochema oligactes (Pallas, 1788)

            

X

    

Family Gorgonocephalidae

                 

Astrocaneum herrerai (Oersted & Lütken, 1856)

 

X

               

Astrocnida isidis (Duchassaing, 1850)

            

X

    

Astrophyton muricatum (Lamarck, 1816)

X

X

X

X

X

X

X

X

X

X

X

X

X

X

 

X

 

Asteroporpa annulata Oesrsted & Lütken, 1856

X

X

X

         

X

    

Schizostella bifurcata A.H. Clark, 1952

  

X

   

X

          

Order Ophiurida

                 

Family Ophiuridae

                 

Amphiophiura fasciculata (Lyman, 1883)

 

X

               

Amphiophiura metabula H.L. Clark, 1915

            

X

    

Amphiophiura oedignatha H.L. Clark, 1915

            

X

    

Amphiophiura oligopora (H.L. Clark, 1913)

 

X

               

Amphipholizona delicata H.L. Clark, 1915

            

X

    

Anthophiura ingolfi Fasmer, 1930

 

X

               

Astrophiura permira Sladen, 1879

 

X

               

Ophiambix devaneyi Paterson, 1985

            

X

    

Ophioceramis januarii Lütken, 1856

 

X

               

Ophiolepis elegans Lütken, 1859

X

X

X

  

X

  

X

  

X

X

X

X

X

 

Ophiolepis gemma Hendler &Turner, 1987

X

X

    

X

          

Ophiolepis impressa Lütken, 1859

X

X

X

X

  

X

 

X

 

X

X

X

X

 

X

 

Ophiolepis kieri Hendler, 1979

X

          

X

     

Ophiolepis paucispina (Say, 1825)

X

X

X

 

X

X

X

    

X

  

X

X

 

Ophiolepis pawsoni Hendler, 1988

      

X

          

Ophiomastus secundus Lyman, 1878

            

X

    

Ophiomusium acuferum Lyman, 1875

 

X

          

X

  

X

 

Ophiomusium eburneun Lyman, 1869

            

X

    

Ophiomusium lymani Wyville-Thomsom, 1873

 

X

               

Ophiomusium testudo Lyman, 1875

            

X

    

Ophiomusium validum Ljungman, 1872

            

X

    

Ophiosphalma planum (Lyman, 1878)

 

X

               

Ophiostriatus atlanticus (Mortensen, 1933)

            

X

    

Ophiozonella nivea Lyman, 1875

 

X

               

Ophiura acervata (Lyman, 1869)

            

X

X

   

Ophiura falcifera (Lyman, 1869)

            

X

    

Ophiura fallax Cherbonnier, 1959

 

X

               

Ophiura tenera (Lyman, 1883)

            

X

    

Family Ophioleucidae

                 

Ophiernus adspersus Lyman, 1875

 

X

               

Family Ophiocomidae

                 

Ophiocoma echinata (Lamarck, 1816)

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

 

Ophiocoma paucigranulata Devaney, 1974

 

X

    

X

   

X

      

Ophiocoma pumila Lütken, 1859

X

X

X

X

X

X

X

 

X

  

X

 

X

X

X

 

Ophiocoma wendti Müller &Troschel, 1842

X

X

X

X

X

X

X

 

X

 

X

X

 

X

X

X

 

Ophiocomella ophiactoides (H.L. Clark, 1901)

X

X

X

X

 

X

X

    

X

 

X

X

X

 

Ophiopsila fulva Lyman, 1878

            

X

    

Ophiopsila hartmeyeri Koehler, 1913

X

X

        

X

 

X

X

   

Ophiopsila riisei Lütken, 1859

X

X

X

 

X

 

X

    

X

 

X

   

Ophiopsila vittata H.L. Clark, 1918

 

X

    

X

    

X

     

Family Ophionereididae

                 

Ophiochiton ternispinus Lyman,1883

 

X

               

Ophionereis dolabriformis John & A.M. Clark, 1954

            

X

    

Ophionereis olivacea H.L. Clark, 1901

X

X

X

   

X

    

X

 

X

   

Ophionereis reticulata (Say, 1825)

X

X

X

X

X

X

X

 

X

 

X

X

 

X

X

X

 

Ophionereis squamulosa Koehler,1913

X

X

X

 

X

 

X

    

X

  

X

  

Ophionereis vittata Hendler, 1995

X

     

X

    

X

   

X

 

Family Ophiodermatidae

                 

Bathypectinura heros (Lyman, 1879)

 

X

      

X

X

 

X

X

    

Ophioderma anitae Hotchkiss, 1982

      

X

          

Ophioderma appressum (Say, 1825)

X

X

X

X

X

X

X

 

X

 

X

X

X

X

X

X

 

Ophioderma brevicaudum Lütken, 1856

X

X

X

X

X

 

X

    

X

 

X

 

X

 

Ophioderma brevispinum (Say, 1825)

X

X

X

X

X

X

X

   

X

X

 

X

 

X

 

Ophioderma cinereum Müller & Troschel, 1842

X

X

X

X

X

X

X

 

X

 

X

X

 

X

X

X

 

Ophioderma ensiferum Hendler & Miller, 1984

X

     

X

          

Ophioderma guttatum Lütken, 1859

X

X

   

X

X

       

X

  

Ophioderma panamense Lütken, 1859

           

X

     

Ophioderma phoenium H.L. Clark, 1918

X

X

X

X

  

X

    

X

  

X

  

Ophioderma rubicundum Lütken, 1856

X

X

X

X

 

X

X

    

X

X

X

X

X

 

Ophioderma squamosissimum Lütken, 1856

X

 

X

   

X

          

Ophiopaepale goesiana Ljungman, 1872

            

X

    

Ophiurochaeta littoralis (Koehler, 1913)

      

X

          

Family Hemieuryalidae

                 

Hemieuryale pustulata Von Martens, 1867

      

X

    

X

     

Sigsbeia conifera Koehler, 1914

X

     

X

    

X

     

Sigsbeia murrhina Lyman, 1878

      

X

          

Ophiochondrus convolutus Lyman, 1869

            

X

    

Family Ophiacanthidae

                 

Ophiolimna littoralis Kolbe, 1899

X

                

Ophiocamax fasciculata Lyman, 1853

   

X

    

X

X

 

X

X

    

Ophiocamax hystrix Lyman, 1878

            

X

    

Ophiacantha echinulata Lyman, 1878

            

X

    

Ophiacantha mesembria H.L. Clark, 1915

            

X

    

Ophiacantha ophiactoides H.L. Clark, 1901

   

X

             

Ophiacantha oligacantha H.L. Clarck, 1918

   

X

             

Ophiocanthella troscheli (Lyman, 1878)

 

X

               

Ophiomitra valida Lyman, 1869

            

X

    

Ophiotrema gracilis Koehler, 1914

            

X

    

Ophiomitrella laevipellis (Lyman, 1883)

            

X

    

Ophiomitrella glabra (H.L. Clark, 1901)

   

X

  

X

          

Ophiomyces frutectosus Lyman, 1869

            

X

    

Ophiopristis hirsuta (Lyman, 1875)

            

X

    

Ophiopristis sertata (Lyman, 1869)

 

X

          

X

    

Ophiotreta valenciennesi rufescens Koehler, 1896

            

X

    

Family Ophiactidae

                 

Hemipholis elongata (Say, 1825)

X

 

X

   

X

    

X

  

X

  

Histampica duplicata (Lyman,1875)

            

X

    

Ophiactis algicola H.L. Clark, 1933

 

X

X

  

X

X

    

X

  

X

  

Ophiactis dispar Verrill, 1899

           

X

     

Ophiactis lymani Ljungman, 1871

              

X

  

Ophiactis muelleri Lütken, 1856

           

X

 

X

   

Ophiactis quinqueradia Ljungman, 1871

X

X

X

X

  

X

    

X

     

Ophiactis rubropoda Singletary, 1973

 

X

               

Ophiactis savignyi (Müller & Troschel, 1842)

X

X

X

X

X

 

X

 

X

 

X

X

X

X

   

Family Ophiolepididae

                 

Ophiothyreus goesi Ljungman, 1872

         

X

  

X

    

Family Amphiuridae

                 

Amphilimna mirabilis (H.L. Clark, 1941)

            

X

    

Amphilimna olivacea (Lyman, 1869)

 

X

         

X

X

    

Amphiodia atra (Stimpson, 1852)

            

X

X

   

Amphiodia planispina (Von Martens, 1867)

X

 

X

        

X

X

    

Amphiodia pulchella (Lyman, 1869)

X

X

X

  

X

X

    

X

  

X

X

 

Amphiodia riisei (Lütken, 1869)

           

X

     

Amphiodia trychna H.L. Clark, 1918

X

X

X

X

  

X

    

X

 

X

X

  

Amphiodia violacea (Lütken, 1856)

      

X

          

Amphioplus abditus (Verrill, 1871)

           

X

     

Amphioplus coniortodes H.L. Clark, 1918

X

X

           

X

   

Amphioplus sepultus Hendler, 1995

X

                

Amphioplus thrombodes H.L Clark, 1918

X

                

Amphioplus tumidus (Lyman, 1878)

            

X

    

Amphipholis gracillima (Stimpson, 1852)

X

 

X

   

X

       

X

  

Amphipholis januarii Ljungman, 1867

X

 

X

   

X

    

X

  

X

  

Amphipholis pachybactra H.L. Clark, 1918

           

X

  

X

  

Amphipholis squamata (Delle Chiaje, 1828)

X

X

X

   

X

X

X

  

X

 

X

   

Amphiura fibulata Koehler, 1913

X

X

    

X

          

Amphiura flexuosa Ljungman, 1867

  

X

              

Amphiura incisa Lyman, 1883

           

X

     

Amphiura otteri Ljungman, 1872

            

X

    

Amphiura palmeri Lyman, 1882

 

X

X

          

X

   

Amphiura rathbuni Koehler, 1914

            

X

X

   

Amphiura stimpsoni Lütken, 1859

X

X

X

 

X

X

X

       

X

X

 

Ophiocnida scabriuscula (Lütken, 1849)

X

X

X

  

X

     

X

 

X

X

X

 

Ophionema intricata Lütken, 1869

  

X

           

X

  

Ophionephthys limicola Lütken, 1869

X

 

X

   

X

    

X

   

X

 

Ophiophragmus cubanus (A.H. Clark, 1917)

X

     

X

          

Ophiophragmus filograneus (Lyman, 1875)

      

X

          

Ophiophragmus pulcher H.L. Clark, 1918

X

X

X

   

X

    

X

     

Ophiophragmus riisei (Lütken, 1859)

  

X

 

X

X

     

X

X

    

Ophiophragmus septus (Lütken, 1859)

X

 

X

   

X

       

X

  

Ophiophragmus wurdemani Lyman, 1860

             

X

   

Ophiostigma isocanthum (Say, 1825)

X

X

X

   

X

   

X

X

X

X

X

X

 

Ophiostigma siva Hendler, 1995

  

X

  

X

X

    

X

     

Family Ophiochitonidae

                 

Ophioplax ljungmani Lyman, 1875

            

X

    

Family Ophiotrichidae

                 

Ophiothrix angulata (Say, 1825)

X

X

X

X

X

 

X

    

X

X

X

   

Ophiothrix brachyactis H.L Clark, 1915

X

X

X

          

X

X

  

Ophiothrix cimar Hendler, 2005

          

X

X

     

Ophiothrix lineata Lyman, 1860

X

X

    

X

   

X

X

     

Ophiothrix oerstedii Lütken, 1856

X

X

X

X

X

X

X

    

X

X

X

X

X

 

Ophiothrix pallida Ljungman, 1872

         

X

       

Ophiothrix stri Hendler, 2005

          

X

X

     

Ophiothrix suensoni Lütken, 1856

X

X

X

X

X

 

X

   

X

X

X

X

   

Ophiothrix synoecina Schoppe, 1996

            

X

    

Class Echinoidea

                 

Order Cidaroida

                 

Family Cidaridae

                 

Eucidaris tribuloides (Lamarck, 1816)

X

X

X

X

X

X

X

X

X

X

X

X

X

X

   

Cidaris abyssicola (A. Agassiz, 1869)

 

X

               

Cidaris rugosa (H.L. Clark, 1907)

 

X

    

X

          

Calocidaris micans (Mortensen, 1903)

 

X

             

X

 

Genocidaris maculata A. Agassiz, 1869

 

X

               

Stylocidaris affinis (Philippi, 1845)

X

X

       

X

 

X

X

    

Stylocidaris lineata Mortensen, 1910

   

X

    

X

X

  

X

    

Tretocidaris bartletti (A. Agassiz, 1880)

   

X

    

X

   

X

    

Porocidaris purpurata (Wyville-Thomson, 1875)

         

X

       

Order Echinothuroida

                 

Family Echinothuriidae

                 

Araeosoma belli (Mortensen, 1903)

 

X

 

X

     

X

  

X

    

Araeosoma fenestratum (Wyville-Thomson, 1872)

   

X

        

X

    

Calveriosoma hystrix Wyville-Thomson, 1872

 

X

               

Hygrosoma petersi (A. Agassiz, 1880)

 

X

  

X

            

Phormosoma placenta (Wyville-Thomson, 1872)

    

X

   

X

X

 

X

X

    

Order Diadematoida

                 

Family Diadematidae

                 

Diadema antillarum (Philippi, 1845)

X

X

X

X

X

 

X

X

X

X

X

X

X

X

   

Centrostephanus longispinus rubricingulus H.L. Clark, 1921

 

X

 

X

             

Astropyga magnifica A.H. Clark, 1934

X

X

X

X

 

X

     

X

X

X

 

X

 

Family Aspidodiadematidae

                 

Aspidodiadema jacobyi A. Agassiz, 1880

 

X

 

X

  

X

  

X

  

X

    

Plesiodiadema antillarum (A. Agassiz, 1880)

    

X

          

X

 

Family Pedinidae

                 

Caenopedina cubensis A. Agassiz, 1869

      

X

          

Order Salenioida

                 

Family Saleniidae

                 

Salenia goesiana Loven, 1874

   

X

             

Salenocidaris profundi (Duncan, 1877)

    

X

            

Salenocidaris varispina (A. Agassiz, 1877)

    

X

            

Order Arbacioida

                 

Family Arbaciidae

            

X

    

Arbacia punctulata (Lamarck, 1816)

X

X

 

X

  

X

    

X

 

X

X

  

Podocidaris sculpta A. Agassiz, 1869

X

           

X

    

Coelopleurus floridanus A. Agassiz, 1871

 

X

 

X

  

X

    

X

X

    

Order Temnopleuroida

                 

Family Toxopneustidae

                 

Lytechinus euerces H.L. Clark, 1912

 

X

 

X

X

 

X

          

Lytechinus callipeplus H.L. Clark, 1912

            

X

    

Lytechinus variegatus (Lamarck, 1816)

X

X

X

X

X

X

X

X

X

 

X

X

X

X

   

Lytechinus williamsi Chesher, 1968

X

X

 

X

X

 

X

    

X

X

    

Pseudoboletia maculata Troschel, 1869

             

X

 

X

 

Tripneustes ventricosus (Lamarck, 1816)

X

X

X

X

X

 

X

 

X

 

X

X

X

X

   

Family Temnopleuridae

                 

Trigonocidaris albida (A. Agassiz, 1869)

   

X

        

X

    

Order Echinoida

                 

Family Echinidae

                 

Echinus gracilis A.Agassiz, 1869

 

X

               

Family Echinometridae

                 

Echinometra lucunter (Linnaeus, 1758)

X

X

X

X

X

X

X

X

X

X

X

X

X

X

   

Echinometra viridis A. Agassiz, 1863

X

X

X

X

X

 

X

X

X

 

X

X

X

X

   

Order Holectypoida

                 

Family Echinoneidae

                 

Echinoneus cyclostomus Leske, 1778

X

X

 

X

X

 

X

 

X

 

X

X

X

    

Order Clypeasteroida

                 

Family Clypeasteridae

                 

Clypeaster chesheri Serafy, 1970

 

X

 

X

    

X

  

X

X

    

Clypeaster cyclopilus H.L.Clark, 1941

             

X

   

Clypeaster euclastus H.L. Clark, 1941

   

X

    

X

   

X

X

   

Clypeaster lamprus A.H. Clark, 1914

   

X

    

X

X

  

X

X

   

Clypeaster luetkeni Mortensen, 1948

    

X

       

X

    

Clypeaster prostratus Ravenel, 1848

        

X

   

X

X

   

Clypeaster ravenelii (A.Agassiz, 1869)

 

X

               

Clypeaster rosaceus (Linnaeus, 1758)

X

X

X

X

X

X

X

 

X

  

X

X

X

   

Clypeaster speciosus Verrill, 1870

          

X

      

Clypeaster subdepressus (Gray, 1825)

X

X

 

X

 

X

X

    

X

X

X

   

Family Fibulariidae

                 

Echinocyamus grandiporus Mortensen, 1907

 

X

          

X

    

Family Mellitidae

                 

Encope aberrans Martens, 1867

 

X

               

Encope emarginata (Leske, 1778)

      

X

X

   

X

X

X

   

Encope michelini L. Agassiz, 1841

X

X

           

X

   

Leodia sexiesperforata (Leske, 1778)

X

X

X

X

 

X

X

 

X

 

X

X

X

X

 

X

 

Mellita quinquiesperforata (Leske, 1778)

X

 

X

X

 

X

  

X

X

X

X

X

X

   

Order Spatangoida

                 

Family Asterostomatidae

                 

Heterobrissus hystrix (A. Agassiz, 1880)

        

X

X

  

X

    

Linopneustes longispinus (A. Agassiz, 1878)

   

X

    

X

X

  

X

    

Palaeobrissus hilgardi (A. Agassiz, 1883)

   

X

        

X

    

Palaeopneustes cristatus A. Agassiz, 1873

   

X

    

X

   

X

    

Palaeopneustes tholoformis Chesher, 1968

   

X

        

X

    

Family Brissidae

                 

Brissopsis atlantica Mortensen, 1907

        

X

X

X

X

X

  

X

 

Brissopsis elongata Mortensen, 1907

  

X

X

  

X

 

X

  

X

X

X

   

Brissopsis mediterranea Mortensen, 1913

    

X

       

X

    

Brissus unicolor (Leske, 1778)

X

X

 

X

X

 

X

 

X

 

X

X

X

X

   

Meoma ventricosa ventricosa (Lamarck, 1816)

X

X

X

X

X

X

X

X

X

X

X

X

X

X

 

X

 

Plagiobrissus grandis (Gmelin, 1788)

X

X

X

X

 

X

X

    

X

X

    

Plethotaenia spatangoides (A. Agassiz, 1883)

         

X

       

Family Loveniidae

                 

Homolampas fragilis (A. Agassiz, 1869)

            

X

    

Family Schizasteridae

                 

Agassizia excentrica A. Agassiz, 1869

   

X

  

X

  

X

 

X

X

    

Brisaster fragilis (Düben & Koren, 1846)

       

X

         

Hypselaster limicolus (A. Agassiz, 1878)

        

X

   

X

    

Moira atropos (Lamarck, 1816)

X

 

X

X

  

X

 

X

  

X

X

X

   

Paraster doederleini Chesher, 1972

   

X

  

X

 

X

X

 

X

     

Paraster floridensis (Kier & Grant, 1965)

   

X

  

X

 

X

  

X

     

Schizaster orbignyanus A.Agassiz, 1880

            

X

X

   

Order Cassidulida

                 

Family Echinolampadidae

                 

Conolampas sigsbei (A. Agassiz, 1878)

   

X

    

X

X

       

Echinolampas depressa Gray, 1851

 

X

               

Family Cassidulidae

                 

Cassidulus caribaearum Lamarck, 1881

 

X

 

X

  

X

    

X

     

Class Holothuroidea

                 

Order Dendrochirotida

                 

Family Phyllophoridae

                 

Neothyonidium parvum (Ludwig, 1881)

  

X

        

X

     

Stolus cognatus (Lampert, 1885)

X

X

         

X

 

X

   

Phyllophorus occidentalis (Ludwig, 1875)

X

X

X

        

X

 

X

X

X

 

Thyone deichmannae Madsen, 1941

           

X

  

X

  

Thyone inermis Heller, 1868

X

          

X

  

X

  

Thyone pseudofusus Deichmann, 1930

 

X

    

X

    

X

  

X

X

 

Thyone sabanillaensis Deichmann, 1930

            

X

    

Thyone tanyspeira Pawson, 1976

        

X

        

Family Cucumariidae

                 

Acaudina suspecta Cherbonnier & Féral, 1981

            

X

    

Cucumaria pulcherrima (Ayres, 1854)

            

X

    

Duasmodactyla seguroensis (Deichmann, 1930)

 

X

X

  

X

       

X

   

Euthyonacta solida (Deichmann, 1930)

 

X

               

Leptopentacta deichmannae Domantay, 1953

            

X

    

Ocnus pygmaeus (Théel, 1886)

 

X

X

          

X

X

  

Ocnus surinamensis (Semper, 1868)

X

 

X

X

 

X

     

X

 

X

 

X

 

Ocnus suspectus (Ludwig, 1874)

X

X

X

  

X

X

    

X

   

X

 

Pentacta pygmaea (Théel, 1886)

             

X

   

Thyone cognata (Lampert, 1885)

    

X

            

Thyonella gemmata (Pourtàles, 1851)

X

X

         

X

     

Thyonella pervicax (Théel, 1886)

 

X

         

X

     

Family Sclerodactylidae

                 

Euthyonidiella destichada (Deichmann, 1930)

X

 

X

   

X

    

X

 

X

   

Euthyonidiella trita (Sluiter, 1910)

 

X

X

  

X

       

X

   

Sclerodactyla briareus (Lesueur, 1824)

             

X

   

Pseudothyone belli (Ludwig, 1886)

 

X

X

   

X

    

X

  

X

  

Family Psolidae

                 

Lissothuria antillensis Pawson, 1967

      

X

          

Lissothuria braziliensis (Theél, 1886)

            

X

    

Order Elasipodida

                 

Family Psychropotidae

                 

Benthodytes sanguinolenta (Theél, 1882)

        

X

        

Benthodytes typica Théel, 1882

    

X

            

Order Aspidochirotida

                 

Family Stichopodidae

                 

Astichopus multifidus (Sluiter, 1910)

X

X

X

X

 

X

     

X

X

X

   

Eostichopus arnesoni Cutress & Miller, 1982

  

X

    

X

         

Isostichopus macroparentheses (H.L. Clark, 1922)

 

X

   

X

X

          

Isostichopus badionotus (Selenka, 1867)

X

X

X

X

X

X

X

 

X

  

X

 

X

X

X

 

Family Holothuriidae

                 

Actinopyga agassizi (Selenka, 1867)

X

X

X

X

X

X

X

X

X

 

X

      

Holothuria (Cystipus) cubana Ludwig, 1874

X

X

X

X

  

X

    

X

 

X

   

Holothuria (Cystipus) occidentalis Ludwig, 1874

   

X

    

X

        

Holothuria (Cystipus) pseudofossor Deichmann, 1930

X

X

X

  

X

X

     

X

X

   

Holothuria (Halodeima) floridana Pourtàles, 1851

X

X

X

  

X

X

 

X

  

X

   

X

 

Holothuria (Halodeima) grisea Selenka, 1867

X

X

X

 

X

X

X

    

X

X

X

 

X

 

Holothuria (Halodeima) mexicana Ludwig, 1875

X

X

X

X

X

X

X

X

X

X

X

X

 

X

X

X

 

Holothuria (Platyperona) parvula (Selenka, 1867)

  

X

X

 

X

      

X

X

X

X

 

Holothuria (Platyperona) rowei Pawson & Gust, 1981

      

X

          

Holothuria (Semperothuria) densipedes Clark, 1901

  

X

              

Holothuria (Semperothuria) surinamensis Ludwig, 1874

X

X

X

  

X

X

      

X

X

X

 

Holothuria (Selenkothuria) glaberrima Selenka, 1867

X

 

X

X

X

X

  

X

  

X

X

 

X

X

 

Holothuria (Theelothuria) princeps Selenka, 1867

 

X

X

X

         

X

   

Holothuria (Theelothuria) imperator Deichmann, 1930

 

X

               

Holothuria (Thymiosycia) arenicola Semper, 1868

X

X

X

X

 

X

X

X

X

  

X

X

  

X

 

Holothuria (Thymiosycia) impatiens (Forskål, 1775)

X

X

X

  

X

X

    

X

X

X

X

X

 

Holothuria (Thymiosycia) thomasi Pawson & Caycedo, 1980

X

X

X

X

X

 

X

 

X

  

X

   

X

 

Order Molpadiida

                 

Family Molpadiidae

                 

Molpadia cubana Deichmann, 1940

           

X

     

Molpadia parva (Théel, 1882)

           

X

X

    

Order Apodida

                 

Family Synaptidae

                 

Epitomapta roseola (Verrill, 1873)

 

X

   

X

X

    

X

     

Euapta lappa (Müller, 1850)

X

X

X

X

 

X

X

   

X

X

  

X

X

 

Protankyra ramiurna Heding, 1928

  

X

              

Leptosynapta imswe Pawson, 1976

      

X

          

Leptosynapta tenuis (Ayres, 1851)

 

X

               

Leptosynapta multigranula H.L. Clark, 1924

 

X

    

X

          

Leptosynapta nannoplax Pawson, 1976

      

X

          

Leptosynapta parvipatina H.L. Clark, 1924

           

X

  

X

  

Leptosynapta roseogradia Pawson, 1976

      

X

          

Eupatinapta acanthia (A.H. Clark, 1899)

           

X

     

Synaptula hydriformis (Lesueur, 1824)

X

X

X

  

X

X

 

X

X

X

X

   

X

 

Family Chiridotidae

                 

Chiridota rotifera (Pourtàles, 1851)

X

X

X

X

 

X

X

    

X

 

X

X

X

 
Table 2

Species, genera, families and orders of Caribbean echinoderms

Class

Order

Family

Genus

Species

Crinoidea

4

10

23

30

Asteroidea

7

23

68

116

Ophiuroidea

2

16

63

148

Echinoidea

11

21

53

76

Holothuroidea

5

10

30

63

Total

29

80

237

433

Table 3

Number of echinoderm species per class in each country. Coastline information taken from the The world fact book (https://www.cia.gov/library/publications/the-world-factbook/geos) and Economic Exclusive Zone (EEZ) information was taken from www.seaaroundus.org/eez/eez.aspx

Country

Crinoidea

Asteroidea

Ophuiroidea

Echinoidea

Holothuroidea

Total

Coastline (km)

No. spp./km of coastline

EEZ (km2)

ssp./km2 EEZ

Mexicoa

7

44

63

36

32

182

905

0.20

88,874

0.0020

Colombia

7

51

62

47

13

180

1,760

0.10

472,891

0.0004

Panama

15

25

56

29

30

155

1,295

0.12

143,442

0.0011

Cuba

11

35

55

21

23

145

3,755

0.04

350,751

0.0004

Belize

5

18

58

27

26

134

516

0.26

35,351

0.0038

Venezuela

10

36

34

24

20

124

2,800

0.04

470,666

0.0003

Puerto Rico

7

22

48

14

30

121

501

0.24

205,529

0.0006

Dominican Republic

17

22

23

41

14

117

1,280

0.09

255,898

0.0005

Honduras

11

30

14

29

11

95

644

0.15

18,151

0.0052

Jamaica

15

20

20

10

21

86

1,022

0.08

258,137

0.0003

Lesser Antilles

11

21

23

7

17

79

Haiti

8

19

19

19

8

73

1,771

0.04

126,760

0.0006

Nicaragua

9

29

6

19

2

65

503

0.13

87,930

0.0007

Trinidad & Tobago

3

7

29

1

15

55

362

0.15

74,199

0.0007

Costa Rica

4

7

16

13

4

44

212

0.21

25,090

0.0018

Guatemala

2

5

4

8

4

23

150

0.15

1,642

0.0140

No country assigned

0

4

0

0

0

4

aShore line and economic exclusive zone are from Quintana Roo State

Among the Class Crinoidea, the Dominican Republic (RDO) is the richest country with 17 species, followed by Panama and Jamaica with 15 each (Table 3), while Trinidad & Tobago and Guatemala are the poorest with three and two respectively. Davidaster discoidea and D. rubiginosa were reported from 12 countries, while eight species were reported only from one country. The genera Davidaster and Democrinus were the most diverse with three species each.

Colombia was the richest country for the class Asteroidea with 51 species followed by Mexico with 44 species. For this class, Astropecten is the most diverse genus with 12 species. Astropecten articulatus and Oreaster reticulatus are reported from 13 countries, and 38 species are reported for only one country. Laetmaster spectabilis is endemic to Cuba. Four species were reported for international waters: Styracaster elongatus, Benthopecten simplex simplex, Cladaster rudis and Henricia sexradiata.

With respect to the Class Ophiuroidea, Mexico, Colombia, Belize and Panama exhibited the highest species richness with 63, 62, 58 and 56 species respectively, while Nicaragua and Guatemala were the least diverse with six and four species each (Table 3). Ophioderma was the most diverse genus with 11 species. Ophiocoma echinata was the only species reported in the entire region (16 countries or territories), while Astrophyton muricatum was reported from 15 countries and Ophioderma appressum from 14. For this class 71 species were reported from only one country, and the species Ophiothrix synoecina is endemic to Colombia, and O. stri and O. cimar are endemic to Costa Rica and Panama.

With respect to Echinoidea, Colombia was the richest country with 47 species, followed by the Dominican Republic with 41 (Table 3). Clypeaster was the most diverse genus with ten species; Meoma ventricosa ventricosa was reported in 15 countries and Echinometra lucunter and Eucidaris tribuloides in 14. In this class, 18 species were reported from only one country.

Mexico was the richest country for the Holothuroidea with 32 species, followed by Panama and Puerto Rico with 30 species each, while Guatemala, Costa Rica, and Nicaragua were the least diverse (Table 3). Holothuria was the most diverse genus with 16 species; Holothuria (Halodeima) mexicana was reported for 15 countries and Isostichopus badionotus for 12. Within this class, 23 species were reported for only one country.

Initially, the publication of echinoderm species descriptions in the Caribbean Sea was slow (Fig. 2a, b), with an average of 0.47 species described per year between 1740 and 1840. Between 1840 and 1920, there was a substantial increase (3.83 species described per year), followed by a slower rate between 1920 and the present, with 0.91 species described per year. Taking the entire period of 1740 to 2010, there were on average 1.61 species described per year. According to the logistic model (Fig. 2b), the curve will be reaching an asymptote in the next 50 years, with seven probable undescribed species to be named.
Fig. 2

a Number of new species of echinoderms from the Caribbean described by decade. b Accumulation curves of species descriptions for the total (gray line) and with the fitted logistic model (black line)

Colombia and Mexico, the most diverse countries, are the most dissimilar in comparison relative to the rest of the region (Figs. 3, 4, gray areas). The difference is primarily due to the composition of Asteroidea (Figs. 3c, 4c), Ophiuroidea (Figs. 3d, 4d) and Echinoidea (Figs. 3e, 4e), and to the influences from other regions like the Gulf of Mexico, Florida and Brazil. In general terms, the Caribbean is very homogenous in species composition between countries (Figs. 3, 4).
Fig. 3

Cluster tree (Euclidean distance) based on species presence/absence Bray-Curtis resemblance matrix: a Echinodermata, b Crinoidea, c Asteroidea, d Ophiuroidea, e Echinoidea, f Holothuroidea; between the Caribbean countries. Gray areas most similar countries

Fig. 4

Non-metric multiple dimensional scaling (NMDS) based on species presence/absence Bray-Curtis resemblance matrix: a Echinodermata, b Crinoidea, c Asteroidea, d Ophiuroidea, e Echinoidea, f Holothuroidea; between the Caribbean countries. Gray areas most similar countries

The taxonomic distinctness index (∆+) indicated that most of the countries are inside the 95% confidence limit of composition; Panama, Cuba, Mexico and Colombia (Fig. 5) being closer to the average of distinctness. Guatemala possesses few species (23 spp.) but exhibits a high value of distinctness (∆+), because those species represent many different groups with few representatives in each one, making the taxonomic distance greater. In Guatemala, each class is represented more or less equally by two to eight species, with very low variation (Fig. 5b). On the other hand, Trinidad & Tobago, which also has few species (55 spp.), has a lower distinctness value (∆+), because the species are unequally represented. The 55 species are dominated by two groups, Ophiuroidea (29 spp.) and Holothuroidea (15 spp.), while the other classes possess fewer species, so the average taxonomic distance is lower (Fig. 5a), with a greater variation (Fig. 5b).
Fig. 5

Taxonomic distinctness index: a average (delta: ∆+), and b their variation (lambda: Λ+) of the Caribbean echinoderms. Continuous line 95% confidence limit, discontinuous line+ average value

Discussion

The Caribbean Sea is a marine province rich in echinoderm species. It has 433 species, representing 6.5% of the total diversity of the phylum, according to numbers presented by Pawson (2007). This region possesses almost 10% of the world’s Ophiuroidea, and between 4 and 6% of the other classes. The Caribbean is second in the tropical Americas with respect to species numbers, after the Gulf of Mexico (522 species; Pawson et al. 2009). The Caribbean ranks above other regions like Brazil (329 species; Hadel et al. 1999), the Pacific of Central America (287; Alvarado et al. 2010), Baja California and the Gulf of California (231 species; Solís-Marín et al. 2005; Honey-Escandón et al. 2008), and the Galapagos Archipelago (198 species; Maluf 1991).

The Caribbean, with an apparent well-known echinoderm fauna (Fig. 2b), is highly homogeneous in composition, due mostly to the current patterns and the semi-closed nature of the region (Spalding 2004). This sea has been recognized as one of the most important coral reef biogeographic provinces (Spalding et al. 2001), with many cryptic habitats ideal for the development of the Ophiuroidea, which is the most diverse in the region. Moreover, additional species will probably be described when the deep areas of the Caribbean are studied, altering the predictive species description curve in Fig. 2b.

Mexico, Belize, Panama, Colombia and Cuba exhibit the highest species diversity, due in part to the rich coastal habitats, which are the most diverse in the region (Spalding et al. 2001), and to intensity of research in these areas. Relative to species description and publications, Colombia has taken the lead, with a higher sampling effort along all of its Economic Exclusive Zone, with collections from shallow and deep waters (Borrero-Pérez et al. 2002a, b, 2008; Benavidez-Serrato et al. 2005). An important feature of the region is the presence of the Mesoamerican reef, in Mexico, Belize, Guatemala and Honduras. This area could be considered a diversity hot spot.

In terms of ecoregions, the Southwestern Caribbean is the richest (Fig. 6), due to its central location in the region, the current patterns, and the influence of species from the north and the south. The Western Caribbean and the Greater Antilles ecoregions are the most similar due to their proximity but also due to the influence of species from the Gulf of Mexico, Florida and Bahamas. On the other hand, the Southern and Eastern Caribbean ecoregions are influenced by the faunas from Guyana and Brazil. This can also help explain why Mexico and Colombia are located on opposite sides of the nMDS (Fig. 4), having faunal influences from two different regions. All the countries from one ecoregion are very similar in composition (Figs. 3, 4).
Fig. 6

Cluster tree (Euclidean distance) based on species presence/absence Bray-Curtis resemblance matrix of echinoderms by ecoregion. W Western Caribbean, SW Southwestern Caribbean, GA Greater Antilles, E Eastern Caribbean, S Southern Caribbean

The Central American countries like Guatemala, Honduras, Nicaragua and Costa Rica, and the Lesser Antilles are the least studied. It is important to notice that in the case of the Central American countries, the social movements of the 1960s and 1970s shifted the attention to issues other other than science. This combined with the lack of economic resources, research institutions, and reduced science investments have resulted in this lack of knowledge. However, according to Fig. 1b, countries with low species numbers are probably undersampled, especially those in close proximity to highly diverse countries. Trinidad & Tobago, the Lesser Antilles, Haiti, Jamaica, Guatemala, Honduras, Nicaragua and Costa Rica are likely undersampled, when compared with the number of species reported from neighboring countries (Fig. 1). However, it is necessary to look at these analyses with care, because in some cases it could reflect the reality of the species richness due to the small size of the coastal zone, the degraded state of some coasts from natural and anthropogenic disturbances, and the lack of complex coastal morphology and habitat diversity.

Along the Caribbean Sea, seven species of echinoderms (four holothuroids and three echinoids) are extracted for commercial use. Among them are the sea cucumbers Astichopus multifidus, Holothuria mexicana, Actinopyga agassizi, and Isostichopus badionotus, the last being the most intensely harvested (Toral-Granda 2008). Extractions occur primarily in Panama, Nicaragua, Colombia, Venezuela, Mexico and Cuba. The exploitation of sea cucumbers without the use of a calculated fisheries management approach is probably the biggest threat to their populations within the region. For many of the species under legal or illegal fishing pressures there is little or no scientific information available on the biology, ecology, population abundance and dynamics (Toral-Granda 2008). Moreover, the situation is exacerbated because the majority of sea cucumber exploitations along the Caribbean (Buitrago and Boada 1996; Rodríguez-Millet and Pauls 1998; de la Fuente-Betancourt et al. 2001; Guzmán and Guevara 2002b) started with the consent of the local authorities, but without any knowledge about the ecology of the species (Guzmán et al. 2003). In the case of Panama, this activity is now banned, due to overfishing in 1997, during which 750,000 sea cucumbers were extracted in 30 days (Guzmán and Guevara 2002b).

Along the Caribbean there are small sea urchin fisheries focused on Echinometra spp., Lytechinus variegatus and Tripnesutes ventricosus. Granada and Martinique sea urchin harvests are the largest (Williams 2002). In the Margarita and Coche islands, Venezuela, L. variegatus and E. lucunter are extracted and processed on a small scale for human consumption (Gómez 1999). However, there is the fear that due to their naturally low population numbers these resources could be reduced by overfishing (Gómez and Gómez 2005).

In the case of the souvenir trade, Lunn et al. (2008) analysed the situation in Mexico, finding that annually 87,600 Oreaster reticulatus are extracted, with no regulation, and sold at prices between 1.8 and 60 US dollars. Similarly, Sloan (1984) indicated that this seastar might be under threat throughout the Caribbean, where some populations have been devastated. Mexico, Jamaica, Trinidad and Venezuela, among others, allow Oreaster extraction without apparent restrictions, which could increase demand in other countries (Guzmán and Guevara 2002a). Lunn et al. (2008) also mentioned the extraction of other echinoderms like Astropecten, Mellita and Tripnesutes as souvenirs in Mexico. However, there are no complete analyses of the exploitation of echinoderms in other places, like Colombia, Venezuela or Panama, where Oreaster as well as other echinoderms are extracted.

The lack of control in echinoderm extraction is a major threat in the Caribbean region. However, the greatest threats to this group of animals come from anthropogenic disturbances like sedimentation and coastal pollution. Sedimentation reduces recruitment success for some key species, such as sea urchins (i.e., Diadema antillarum; Vázquez-Domínguez 2003). Moreover, the loss of architectural complexity of the majority of coral reefs on the area (Alvarez-Filip et al. 2009), reduces the availability of substrate for recruitment and development. Agricultural pollution can produce endocrine disruptions in many species, slowing the recovery of critical populations (Rawlins et al. 1998). Coastal development has become a serious threat for endemic and cryptic species of ophiuroids in Colombia, Panama and Costa Rica, where the rock cavities made by sea urchins are being seriously degraded (Hendler 2005; Monroy-López and Solano 2005).

The research priorities for this group of animals in the Caribbean should be focused on the discovery of new species from deep waters, increased research in Central America and the Lesser Antilles, and increased ecological research in groups other than the echinoids. Future studies should also focus on (1) the role of marine protected areas for the recovery of key species, (2) population changes in fishing grounds, and (3) the impact of diseases in the region. Moreover, due to its rich biodiversity, relative to other tropical American waters, research and conservation efforts must be directed toward the preservation and conservation of that diversity, avoiding illegal extraction, enforcing controls, and improving coastal management.

Notes

Acknowledgements

I am grateful to those who facilitated access to literature or access to museum collections. Specially, I am grateful to the Census of Marine Life-Caribbean, CONACYT (México), CONICIT-MICIT (Costa Rica). I also would like to thank P. Miloslavich, G. Díaz-Erales, M. Chávez, J. Cortés, B. Bezy and C. Fernández, whose contributions greatly enriched this paper. I appreciated the comments of the three anonymous reviewers. This publication is dedicated to Balú.

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Copyright information

© Senckenberg, Gesellschaft für Naturforschung and Springer 2010

Authors and Affiliations

  1. 1.Centro de Investigación en Ciencias del Mar y Limnología (CIMAR)Universidad de Costa Rica (UCR)San PedroCosta Rica
  2. 2.Posgrado en Ciencias Marinas y Costeras (CIMACO)Universidad de Baja California Sur (UABCS)La PazMéxico

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