Echinoderms - Springer Link

Part 36

Echinoderms Juan José Alvarado and Jorge Cortés

Nidorellia armata, a common species from Pacific Costa Rica (Photo: Jorge Cortés)

J.J. Alvarado Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, San Pedro, San José 11501-2060, Costa Rica. e-mail: [email protected] J. Cortés Centro de Investigación en Ciencias del Mar y Limnologia (CIMAR), and Escuela de Biología, Universidad de Costa Rica, 11501-2060 San José, Costa Rica e-mail: [email protected]

I.S. Wehrtmann, J. Cortés (eds.) Marine Biodiversity of Costa Rica, Central America, © Springer Science + Business Media B.V. 2009

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Abstract Two hundred and twenty nine species of echinoderms (Phylum Echinodermata), including feather stars, sea stars, brittle stars, sea urchins, and sea cucumbers, are reported for Costa Rica. The Class Crinoidea is represented by six species from five genera, four families and one order; the Class Holothuroidea by 49 species, 24 genera, 15 families, and 6 orders; and the Asteroidea by 50 species, in 38 genera, 19 families, and 6 orders. The most diverse classes are Ophiuroidea (69 species, 31 genera, 11 families, and 2 orders) and Echinoidea (55 species, 33 genera, 17 families, and 11 orders). The genus Holothuria is the largest with 20 species. Of the 229 species reported, 187 are from the Pacific (103 from the Pacific mainland and 124 from Isla del Coco), 44 are from the Caribbean, and there were two species in common to both coasts (Ophioderma appresum and Ophiactis savigny). Most of the taxonomic research on echinoderms was done in the first half of the twentieth century. Here, we report 50 new records from Costa Rica. Due to the lack of collections from some areas of the country and from deepwaters, it is expected that more species will be found in the future, possibly including undescribed species.

Introduction The echinoderms (feather stars, sea stars, brittle stars, sea urchins, and sea cucumbers) are exclusively marine animals. The Phylum Echinodermata (literally: spiny skinned animals) is divided into five living classes: Crinoidea, Asteroidea, Ophiuroidea, Echinoidea, and Holothuroidea. About 6,500 species of living echinoderms and over 13,000 fossil species in many extinct classes have been described (Hendler et al. 1995). The echinoderms have important ecological roles such as the control of algal growth in Caribbean coral reefs by the black sea urchin, Diadema antillarum (Carpenter 1997). An example from the Pacific is the coral-eating sea star, Acanthaster planci, which reaches population densities in the Pacific that can devastate entire coral reefs (Moran 1986). The echinoderms from Costa Rica have been studied in a sporadic way. The first work dates from the late nineteenth and early twentieth century by the expeditions of the US Fish Commission Steamer “Albatross” in charge of Alexander Agassiz (Ludwig 1894, 1905; Agassiz 1898, 1904; Lütken & Mortensen 1899; Clark 1917). In the 1930s and the 1940s, as a result of the expeditions to the eastern Pacific by the Allan Hancock Foundation and the New York Zoological Society, there has been a significant amount of research on echinoderms (Ziesenhenne 1937, 1940, 1942, 1955; Clark 1939, 1940, 1948; Deichmann 1941, 1958). Among the numerous species reported in these papers, 11 species of echinoderms have been described as new. Many of these new species had been collected at Isla del Coco (summary in Hertlein 1963). One of these species recorded from Isla del Coco is Stichopus horrens (Fig. 36.1).

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Fig. 36.1 Stichopus horrens, a common species around Isla del Coco, Pacific Costa Rica (Photo: Graham Edgar)

More recent publications are ecologically oriented. Lawrence (1967) studied the lipid reserves in the gut of three species of sea urchins (Echinometra lucunter, Tripneustes ventricosus, and Lytechinus variegatus) collected in Portete, Limon. Bakus (1974) experimented with the toxicity of holothurians to fishes at Isla del Coco, finding that six of seven species tested were toxic. The density, distribution, and massive death of the black sea urchin, D. antillarum, at Parque Nacional Cahuita, were studied in 1977, 1980, 1983, 1992, 1999, and 2003 (Valdez & Villalobos 1978; Cortés 1981, 1994; Murillo & Cortés 1984; Alvarado et al. 2003). Guzmán (1988) and Guzmán & Cortés (1992) provided information on the density of the corallivorous sea star, A. planci, at Isla del Caño and Isla del Coco, respectively. Guzmán (1988), moreover, presented densities of the sea urchin D. mexicanum from Isla del Caño and Guzmán & Cortés (1992) from Isla del Coco. Rojas (1990) and Rojas et al. (1998) analyzed seven heavy metals in the sea cucumber, Holothuria mexicana, from Cahuita, and found that the concentrations of lead, iron, and copper are higher than in other organisms. Moreover, they suggested that the holothurians are organisms, suitable for the detection of metal pollution, because they feed on bottom sediment. Cortés et al. (1992) published on the massive death of organisms due to the uplift of the Caribbean coast during the Limón 1991 earthquake. Among the most affected was the boring sea urchin, E. lucunter. During the evaluation of the effects of the earthquake they collected a great number of skeletons of the irregular sea urchins, Meoma ventricosa and Echinoneus cyclostomus. The great abundance of skeletons of these sea urchins was used to reconstruct their population structure (Soto et al. in preparation). Lessios et al. (1996) reported the

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Fig. 36.2 Echinothrix diadema, in Costa Rica known exclusively from Isla del Coco (Photo: Graham Edgar)

presence of two Indo-Pacific diadematid sea urchins (Echinothrix diadema, Fig. 36.2, and E. calamaris) at Isla del Coco and proposed that their arrival was enhanced by the North Equatorial Counter Current during the 1982–1983 El Niño. The same authors suggest that the echinoderm fauna of Coco, Revillagigedo, and Clipperton Island, is a mixture of both east- and west-Pacific faunas. Lessios et al. (1998) postulated that Isla del Coco is a stepping stone between both sides of the Pacific Ocean, and found evidence of genetic flow across the largest marine biogeographic barrier of the world (the Eastern Pacific Barrier). At the end of the twentieth century and beginnings of the twenty-first century several researchers have being centering their attention on the biogeographic relationships of sea urchin populations between both sides of the Pacific by molecular analysis (Palumbi 1996; Lessios et al. 1999, 2001, 2003; McCartney et al. 2000; Zigler & Lessios 2003, 2004; Palumbi & Lessios 2005). Alvarado & Cortés (2004) reviewed the state of knowledge on echinoderms of Costa Rica and Central America until 2003 pointed out the needs for research on the area. The diversity, abundance, and distribution of echinoderms from Parque Nacional Marino Ballena, on the Pacific coast of Costa Rica was studied by Alvarado & Fernández (2005), and they suggested that sedimentation is having a negative effect on echinoderm diversity. Meanwhile, Bolaños et al. (2005) compared the diversity and distribution of echinoderms in two environments (seagrass and coral reefs) at the reef lagoon of Parque Nacional Cahuita. More recently, Hendler (2005) described two new species of brittle star of the genus Ophiothrix collected from Cahuita: O. stri and O. cimar. Furthermore, there are nine publications on fossil echinoderms of Costa Rica (Gabb 1881; Jackson 1917; Durham 1961; Aguilar 1978, 1997, 1999; Fisher 1985; Aguilar & Cortés 2001; Alvarado et al. 2006). Alvarado et al. (2006) made a review on the research done on that area in Costa Rica. The fossil fauna is composed of one crinoid and 45 species of echinoids. Most of the specimens are from the

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Cretaceous and Miocene, found in 11 geological formations throughout the country (Alvarado et al. 2006). In this part we present lists of the echinoderms of Costa Rica. To compile these lists we used the information in the literature, collections in the National Museum of Natural History, Smithsonian Institution (USNM), Colección Nacional de Equinodermos, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de Mexico (UNAM), and Museo de Zoología, Escuela de Biología, Universidad de Costa Rica (UCR). New records to Costa Rica are indicated in the list and in some cases specimens have been deposited at the Museo de Zoología (these specimens are indicated by UCR and a collection number) or in the National Museum of Natural History. We follow the taxonomy used by Maluf (1988).

Discussion The information in this part indicates that six species of crinoids, 50 asteroids, 69 ophiuroids, 55 echinoids, and 49 holothurians live in Costa Rica, comprising a total of 229 species (Species Lists 36.1 and 36.2 are included on the CD-Rom, Table 36.1). Forty four species are from the Caribbean, 185 from the whole Pacific (103 from the Pacific mainland and 124 from Isla del Coco) (Table 36.1), and there are two species occurring on both coasts in Costa Rica (Ophioderma appresum and Ophiactis savigny) (Fig. 36.3). Fifty species reported herein are new records for Costa Rica (Species Lists 36.1 and 36.2). Eleven species have been described from material collected in Costa Rica: two asteroids Pauliella aenigma (Albatross Expedition of 1891, in charge of Alexander Agassiz), and Tamaria obstipa, both from Isla del Coco. Two ophiuroids, Amphiodia vicina, collected in Bahía Ballena, Golfo de Nicoya during the Eastern Pacific Expeditions of the New York Zoological Society lead by William Beebe, and Ophionereis dictyota, collected at Chatham Bay, Isla del Coco in 1938, during the Allan Hancock Pacific Expeditions on the Velero III. Three echinoids, Hesperocidaris panamensis, and Centrocidaris doederleini, from material collected in Isla del Coco during the Albatross Expedition of 1891 (Clark 1948), and Encope cocosi, collected at Wafer Bay, Isla del Coco, on a sandy bottom between 4 and 8 m deep. Finally, four holothurians: Psolus diomedeae, Isla del Coco, collected during the Albatross Expedition in 1891, Pentamera beebi, Bahía Ballena, and Labidodemas americanum, Isla Tortuga, Golfo de Nicoya, both collected during the Eastern Pacific Expeditions of the New York Zoological Society (Deichmann 1938); and Euthyonidium veleronis, from Playa Blanca, Guanacaste, collected by the Allan Hancock Pacific Expedition of 1935 on the Velero III. Three species are endemic: the irregular sea urchin, E. cocosi and the sea stars Astropecten benthophilus and Persephonaster armiger from Isla del Coco. The sites with the highest number of species are Isla del Coco (124 species), Golfo de Nicoya (26), Playas del Coco (12), and Santa Elena (11), which are also

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Table 36.1 Number of species, genera, families, and orders of echinoderms from Costa Rica Class

Crinoidea Asteroidea Ophiuroidea Echinoidea Holothuroidea Total

Orders Families Genera Species Caribbean Whole Pacific Pacific Mainland (PM) Isla del Coco (IC) % species in common between PM and IC

1 4 5 6 4 2 0 2 0.00

6 19 38 50 7 43 14 34 14.71

2 11 31 69 17 55 36 30 40.00

11 17 33 55 12 42 25 31 3.65

6 15 24 49 4 45 28 27 37.04

26 66 131 229 44 187 103 124 32.26

Fig. 36.3 Ophiactis savigny from the Parque Nacional Marino Ballena, Pacific Costa Rica (Photo: Ingo Wehrtmann)

the sites visited by the expeditions of the 1930s. The number of species from other sites is expected to increase when more extensive collections will be carried out. Panamá and Belize are the Central American countries from which more species of echinoderms have been reported at 276 and 110, respectively (Table 36.2). These are also the best studied countries in the region. Four hundred and twenty seven species of echinoderms have been reported for Central America, but this certainly is an underestimation of the true diversity of echinoderms of the region. This

Crinoidea Asteroidea Ophiuroidea Echinoidea Holothuroidea Total Species in common with Costa Rica References

5 11 55 20 19 110 37 (33%) [24, 33, 34, 36, 40, 41, 51]

Belize

2 2 11 11 7 33 27 (82%) [9, 13, 22, 28, 37, 49, 52, 54]

Guatemala 9 3 7 18 8 45 26 (57%) [9, 13, 21, 22, 23, 32, 36, 37, 41, 44, 49, 52, 54]

Honduras 0 7 13 13 12 45 36 (80%) [6, 9, 13, 14, 22, 23, 37, 49, 52, 54, 60]

El Salvador 3 9 12 8 6 38 26 (76%) [9, 13, 22, 23, 36, 42, 49, 52, 54]

Nicaragua 6 50 69 55 49 229

Costa Rica

13 47 71 89 56 276 119 (43%) [9, 10, 12–14, 21, 23, 25, 31, 34, 35, 37, 41–43, 45, 49, 52–54, 57, 63, 67]

Panamá

20 86 135 98 88 427

CA

Table 36.2 Number of species of echinoderms reported for each Central American country. CA = number of species for the entire Central American region

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number represents 6.5% of the total diversity of described species of echinoderms. There are few studies from the other countries in the region, and many habitats and sites have not yet been sampled. The number of species in common between Costa Rica and the other Central American countries is presented in Table 36.2. Compared to the countries with low species diversity reported so far, the percentage of species in common is high, probably due to the fact that in those countries only the most abundant and widespread species have been reported. The number of species between Costa Rica (229) and Panama (276) is not substantially different, but only 43% of the species occur in both countries. This indicates, either that the faunas of each country are localized and differ widely, or that more species will be found in both countries as more areas and environments are studied. In conclusion, more studies are needed on the biodiversity of the echinoderms in Central America.

Specialists Gordon Hendler: Curator of Echinoderms, Natural History Museum of Los Angeles County, Los Angeles, California, USA – Worldwide. [email protected] Francisco A. Solís-Marín: Colección Nacional de Equinodermos, Laboratorio de Sistemática y Ecología de Equinodermos, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, México-Caribbean and tropical eastern Pacific. [email protected] Sabine Stöhr: Curator of Echinoderms, Swedish Museum of Natural History, Department of Invertebrate Zoology, Box 50007 SE-10405, Stockholm, Sweden-Worldwide. [email protected]

Collections There are collections of Costa Rican echinoderms in the Museo de Zoología of the Universidad de Costa Rica, Los Angeles County Museum, the National Museum of Natural History at the Smithsonian Institution, Colección Nacional de Equinodermos from the Instituto de Ciencias del Mar y Limnología at the Universidad Nacional Autónoma de México, the Natural History Museum of Santa Barbara, and in the Museum of Comparative Zoology, Harvard University.

Recommendations More studies are needed in deepwaters, in the Caribbean intertidal zone and in some sections of the coasts, for example, Bahía Salinas on the North Pacific. More research is needed in the rest of the Central American countries.

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Acknowledgments We appreciate the help received over the years by R. Vargas and A. Villalobos, Museo de Zoología, Escuela de Biología, Universidad de Costa Rica; Francisco SolísMarín, Colección Nacional de Equinodermos, Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México; Cinthya Ahearn, National Museum of Natural History, Smithsonian Institution; and the support with the literature by K. Hale and H. Albertson, RSMAS Library, University of Miami. We appreciate that Graham Edgar allowed us to use his excellent photos for this part. Funding to study the marine biodiversity of Costa Rica was provided by the Vicerrectoría de Investigación of the Universidad de Costa Rica (Projects 808-92-237; 80896-601) and by CONICIT (Project 90-326-BID). Comments by anonymous reviewers improved the paper.

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