*Dirección para correspondencia:

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Abstract

Isla del Coco (also known as Cocos Island) is an oceanic island in the Eastern Tropical Pacific; it is part of the largest national park of Costa Rica and a UNESCO World Heritage Site. The island has been visited since the 16^th Century due to its abundance of freshwater and wood. Marine biodiversity studies of the island started in the late 19^th Century, with an intense period of research in the 1930’s, and again from the mid 1990’s to the present. The information is scattered and, in some cases, in old publications that are difficult to access. Here I have compiled published records of the marine organisms of the island. At least 1688 species are recorded, with the gastropods (383 species), bony fishes (354 spp.) and crustaceans (at least 263 spp.) being the most species-rich groups; 45 species are endemic to Isla del Coco National Park (2.7% of the total). The number of species per kilometer of coastline and by square kilometer of seabed shallower than 200m deep are the highest recorded in the Eastern Tropical Pacific. Although the marine biodiversity of Isla del Coco is relatively well known, there are regions that need more exploration, for example, the south side, the pelagic environments, and deeper waters. Also, several groups of organisms, such as the flatworms, nematodes, nemerteans, and gelatinous zooplankton, have been observed around the Island but have been poorly studied or not at all.

Key words: Marine biodiversity, Costa Rica, Isla del Coco, Cocos Island, Eastern Pacific, endemic species.

Resumen

La Isla del Coco es una isla oceánica en el Pacífico Tropical Oriental; es parte del Parque Nacional más grande de Costa Rica y es un sitio de Patrimonio Mundial. La isla ha sido visitada desde el Siglo XVI por su abundancia de agua dulce y árboles. Estudios de biodiversidad marina de la isla empezaron a finales del Siglo XIX, con un intenso período de investigación en la década de 1930, y de nuevo desde mediados de la década de 1990 al presente. La información sobre organismos marinos se encuentra dispersa y en algunos casos en publicaciones antiguas. En el presente trabajo se recopilan todos los registros publicados de organismos marinos de la  isla. Al menos 1688 especies han sido registradas, con los gasterópodos (383 especies), peces óseos (354 spp.) y crustáceos (al menos 263 spp.) como los grupos con más especies; de esas, 45 son especies endémicas del Parque Nacional Isla del Coco (2.7% del total). El número de especies por kilómetro de costa y por kilómetro cuadrado de lecho marino de menos de 200m de profundidad son los más altos de cualquier sitio estudiado. Aunque se conoce relativamente bien la  biodiversidad marina de la Isla del Coco,  hay regiones, por ejemplo, el lado sur, los  ambientes pelágicos, y las zonas más  profundas que requieren de más exploración. También, varios grupos de organismos han sido observados en la isla pero muy poco estudiados o no del todo, por ejemplo los gusanos planos, nemátodos y el plancton gelatinosos.

Palabras clave: Biodiversidad marina, Costa Rica, Isla del Coco, Pacífico Oriental, especies endémicas.

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Isla del Coco (Cocos Island) is an oceanic island in the Eastern Tropical Pacific Ocean, about 500km from mainland Costa Rica (Cortés 2008), and biogeographically it is part of the  Ocean  Island  Province  (sensu  Robertson & Cramer 2009). It is a highly diverse area with the highest number of endemic species in Costa Rica (Cortés & Wehrtmann 2009, Cortés 2013a). It was declared a National Park in 1978, a UNESCO World Heritage Site in 1997, and a Ramsar site in 1998. It is important for Costa Rica, due to of its biological and historical richness and also because the Territorial Sea surrounding the island makes the marine area of Costa Rica more than 10 times its terrestrial area (Cortés & Wehrtmann 2009, Cortés 2013b).

Marine  biodiversity  studies  at  Isla  del Coco started in the late 19^th Century with the expedition of the US Fisheries Commission Steamer Albatross (see a history of marine research at what is now Parque Nacional Isla del Coco (Isla del Coco National Park) in Cortés 2008). Since then, many studies have been done but there is still more to be discovered. Hertlein (1963) compiled what was known about marine species of the Island. In this paper I update the list of marine organisms of Isla del Coco National Park, compare it with the species from the mainland, and identify areas for future research.

Materials and Methods

Published   records   of   marine   species of  Parque  Nacional  Isla  del  Coco  (PNIC) were searched and species lists compiled. An important source was the book “Marine Biodiversity of Costa Rica, Central America” edited by Wehrtmann and Cortés (2009). Here I included overlooked and newly published records. The number of species from PNIC is compared to the numbers from the rest of the Pacific of Costa Rica. Including the mainland, the Coco Volcanic Cordillera, the pelagic areas, and the Costa Rica Thermal Dome.

The perimeter of Isla del Coco is 28.8km, and it is surrounded by a platform that drops off at around 180-200m. The perimeter at 100m is 54.4km and at 200m is 71.4km. The area less than 50m deep around the island is 91.5km², less than 100m is 133km², and less than 200m deep  is  318km²  (O.G.  Lizano,  pers.  comm. 2011).  The  Linear  Biodiversity  Index  (LBI) and the Area Biodiversity Index (ABI) were calculated as in Wehrtmann et al. (2009). To calculate the LBI, the number of species was divided by the perimeter of the island. The ABI was calculated by dividing the number of species by the area down to the 200m isobaths; the depth within which almost all the species had been recorded.

Marine Biodiversity of Isla del Coco National Park

At least 1688 species of marine organisms (Appendix 1) have been reported from Isla del Coco National Park (PNIC). The most speciesrich groups are the gastropods (383 species), bony fishes (354 spp.) and crustaceans (at least 263 spp.) (Table 1). Close to 4,700 species of marine organisms have been reported for Costa Rica (Table 1); of these species, 747 or ~16% have been only reported from PNIC but not from other areas of Costa Rica. The percentages by taxonomic group ranged from 0 to 100%. All brown algae, echiuran, marine insects, and reptiles from PNIC are also found in the rest of Costa Rica. While all reported branchiopods, ostracods, phoronids, and crinoids, represented by one or two species, known from Costa Rica are reported from PNIC. Some groups have a disproportionate percentage of known species at PNIC, scyphozoans (40%), echinoderms as a phylum (42.6%), and its classes, asteroids (66.7%) echinoids (36.4%) and holothurians (40.4%), brachiopods (75.0%), and within the chordates, cephalochordates (50%) appendicularians (70%) and thaliacians (75%).

Species of several groups have been reported from other areas of Costa Rica than PNIC (Table 2). Some of these taxa we know are absent from PNIC, for example the seagrasses and mangroves, and some of the species associated to this ecosystems. Other groups have been observed, photographed or collected but there are no published accounts of them. Within these groups we have nematodes, nemerteans, ascideans, and parasites of fishes and turtles. Free-living flat worms have been observed along the mainland coast of Costa Rica as well as at PNIC, but there are no publications. Of other taxa we do not know if they are present or not, for example, marine fungi, cumaceans and kinorhynchans (Table 2). Plus there must be other marine groups that have been reported from the Eastern Tropical Pacific, for example, loriciferans (Heiner & Neuhaus 2007) that might be present at PNIC.

Forty five species or 2.7% of the species known  from  PNIC  are  endemic  (Tables  3, 4), and this represents 47.4% of all endemic marine species of Costa Rica (95 spp.). The number of endemic species is relatively low, but that is common in marine environments. The list of endemic marine species is presented in Table 3, as well as the reference to the publication where the species was described. Between 1893 and 1971, 16 species were described, while 29 were described from 1981 to 2011. Most endemic species are fishes (33.3% of all endemics from PNIC) and most were described in the last 30 years (11 of the 15 species). Crustacea is the next group with most endemism, 28.9%, followed by the mollusks, 15.5%, all very well studied groups (Table 4). Within a particular group, the brachiopods have the highest percentage of endemism, 16.7% followed by the sponges and polyplacophorans with 12.5% (Table 4).

Biodiversity indices used to compare species diversity at Isla del Coco with that at the Costa Rican coast revealed significantly higher values at PNIC than at the coast. For example, the Linear Biodiversity Index for Isla del Coco is 58.6, which is significantly higher than the highest  value  (3.8)  found  by  Wehrtmann  et al. (2009) for the Costa Rican Pacific coast. The same happens when comparing the Area Biodiversity Index of continental shelves. The value for the 200m isobaths of PNIC is 5.3, compared to the highest value of 0.3 reported by Wehrtmann et al. (2009). The LBI and ABI values for the Costa Rican coastline and continental platform, respectively, were the highest when compared to other countries in the region (Wehrtmann  et  al.  2009).  By  all  measures Isla del Coco is a very rich area in the eastern tropical Pacific.

Status of the endemic species

Forty five species are endemic to PNIC (Table 2). While some are abundant, such as the calcified hydrozoan Stylaster cocosensis, described in 1991 by Stephen D. Cairns, others have not been seen since they were described, for example, the sand dollar, Encope cocosi. This species had not been found alive since it was described by H.L. Clark in 1948. However,  in  January  2007  a  specimen  recently dead was dredged from deep water. In 1986 using the research submersible Johnson-SeaLink (Harbor Branch Oceanographic Institution,  Fort  Pierce,  Florida,  USA)  diving  to several hundred meters, collected specimens that resulted in new species and some were endemic (Cairns 1991a, b). We don not know the status of some of those endemics because no submersible with the depth capacity of the Johnson-Sea-Link has been back to the island. There  is  now  another  submersible  operating more regularly at the island, the DeepSee (Undersea Hunter Group, Puntarenas, Costa Rica), with a depth capability of 450m (Cortés & Blum 2008). We have been able to observed several  of  the  deepwater  endemics  collected in 1986, some are relatively abundant. Eleven endemic species have been described in the last decade so it’s possible that eventually they will be found in other areas. Some species of fishes which were initially classified as endemic to one of the oceanic islands of the eastern Tropical Pacific are now reported from one or more of the other oceanic islands. For example, Stegastes arcifrons which have been found in the three oceanic islands, Galápagos, Malpelo and Isla del Coco or Serranus tico and Halichoeres discolor, found in Isla del Coco and Malpelo. Starr et al. (2012) indicated in their study of deepwater fishes of Isla del Coco National Park and Las Gemelas Seamount that probably deep areas in the eastern tropical Pacific will have similar species, but more studies and collections are needed.

Discussion

Isla del Coco National Park has a rich marine biodiversity with some groups having been studied for many years and numerous scientists. For example, fishes and mollusks, especially gastropods, are relatively well known while other groups such as cyanobacteria, gelatinous zooplankton, nematodes and flatworms  have  been  poorly  studied  or  never at all even though we know they are on the island. As a result of recent expeditions (2006-2012) many new records of species have been reported (Dean et al. 2010a, 2012, Sibaja-Cordero et al. 2012), including a phylum, Phoronida (Dean et al. 2010b), and new species are being discovered, even of conspicuous groups such as octocorals (Breedy & Cortés 2011, Breedy et al. 2012). Reports of new records and descriptions of new species are being prepared at the present time.

Hertlein (1963) did a compilation of published marine species of Isla del Coco, and included a biogeographic analysis of the flora and fauna of the island, plus an annotated bibliography. He reported 334 species (Table 5), with the gastropods (62 species) as the most species-rich group, followed by bony fishes (59) and crustaceans (56). The number of species and of different taxonomic groups has increased  significantly  but  the  same  pattern of the most species-rich groups is maintained. Wehrtmann et al. (2009) reported 1,142 marine species for Isla del Coco National Park, with the most species-rich groups, in the same order, being the same as above. Here, 546 more species were added to the list of marine species of Isla del Coco National Park, and more will be added in the near future as other groups, depths and areas of the island are being studied.

Hickman (2009), in his study of the marine invertebrate  biota  of  the  Galápagos  Islands, found that while some groups of species are depauperated others displayed high diversity when compared to mainland Ecuador. Similar patters were observed at Isla del Coco National Park. These patterns can be attributed to several possible factors likely acting in concert, both for source populations from elsewhere as well as established populations at PNIC: variation in the dispersal potential to and from PNIC, the probability of recruitment at PNIC, and the potential for survival and continued recruitment based on local environmental conditions. Species with long-lived larvae will have a chance of dispersing more than others if they find the type of environmental conditions necessary to survive and reproduce. For example, the absence of seagrasses and the low number of species of bivalves may be due to the lack of soft sediments where they can live.

Areas for future research

The least studied area of PNIC is the south side due to the normally rough sea conditions on that side (Lizano 2008). From a few observations, several species and environments in the south are different from the north in species density and composition, probably due to the currents that flow there (Cortés & Blum 2008). More sampling should be done on that side in the future in the shallow and deepwaters of the south for better understanding the biodiversity of PNIC, and the effect of currents on that biodiversity.

There are several groups of organisms which have been observed and in some cases collected but for which there are no publications. Examples include cyanobacteria, sponges, flatworms, and nematodes (Table 2). For a few groups, especially the best known, there are some publications on their biogeographic relationships. Several species of stomatopods (Manning 1972), most reef building corals (Cortés 1986, 2011, Glynn & Ault 2000), some mollusks (Montoya & Kaiser 1988), sea urchins (Lessios et al. 1998), and about one third of the shore-fishes (Robertson et al. 2004) are related western Pacific species. More molecular work is needed to discover cryptic species (e.g. Knowlton 2000, Boulay et al. in prep.), and the genetic connectivity (e.g. Lessios & Robertson 2006) of PNIC populations with other areas.

Polidoro et al. (2012) indicated the importance of species-specific information regarding population trends and extinction risks for developing conservation strategies. To do this we must first know what is there, which this paper intends to fulfill. Then we need to know what is the status of the populations, how they are changing over time, and what is affecting them. Unfortunately for most groups this information is unknown.

Acknowledgements

I thank the following scientists for their help with their group of specialty and/or review of  the  manuscript:  Fabián Acuña,  Juan  José Alvarado, Peter Auster, Gilbert Barrantes, Odalisca Breedy, Richard Brusca, William Bussing, Allan Carillo, Allen Collins, Harlan Dean, Ana Dittel, Cindy Fernández, Cristian Pacheco, Christian Emig, José Leal, Laurence Madin,  Ross  Robertson,  Eva  Salas,  Astrid Sánchez, Jeffrey Sibaja-Cordero, Rick Starr, Robert  van  Syos,  Benjamin Victor  and  Rita Vargas. Omar Lizano for providing the data on perimeter and areas of the marine sections of Isla del Coco National Park (PNIC). Research at PNIC has been funded by the Vicerrectoría de Investigación and CIMAR of the Universidad de Costa Rica, Conservation International (CI), Fonds Français pour l’Environnement Mondial  (FFEM)  and  the  Consejo  Nacional de Rectores de las Universidades Públicas de Costa Rica (CONARE). The preparation of this publication was advanced significantly during my stay as visiting Professor at Newcastle University, Newcastle, United Kingdom. Support has been received from Área de Conservación Marina Isla del Coco (ACMIC) and the Undersea Hunter Group.

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*Correspondencia a:
Jorge Cortés. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Ciudad de la Investigación, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica; jorge.cortes@ucr.ac.cr. Escuela de Biología, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica

1. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Ciudad de la Investigación, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica; jorge.cortes@ucr.ac.cr
2. Escuela de Biología, Universidad de Costa Rica, San Pedro, 11501-2060 San José, Costa Rica
Note: Number references as used in the Tables.

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