Marine Biodiversity Records

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New records of marine decapods and stomatopods in Área de Conservación Guanacaste (ACG): four years of marine biodiversity inventorying

  • Rita Vargas-Castillo
  • Jorge CortésEmail author
Open Access
Marine Record


The marine area of Área de Conservación Guanacaste (ACG) contains a 43,000 ha formal marine protected area, a 732 ha special management zone in Bahía Santa Elena, and 150 km of wild protected coastline. In an effort to broaden the biodiversity knowledge of all marine taxa present in the area, an inventory was started in 2015 (BioMar-ACG Project). This initiative is being funded by the Guanacaste Dry Forest Conservation Fund (GDFCF) in collaboration with government staff at ACG, and is carried out by Centro de Investigación en Ciencias del Mar y Limnología (CIMAR, Center for Research in Marine Science and Limnology) and Museo de Zoología (Zoology Museum), both from Universidad de Costa Rica (UCR). After four years of the project, 2650 specimens of marine decapod crustaceans and stomatopods have been collected, belonging to 209 species, out of which 99 are new records for ACG, four may be new species and nine (Cyrtoplax panamensis, Glyptoxanthus labyrinthicus, Pachyches marcortezensis, Petrolisthes donadio, Pylopagurus holmesi, Synalpheus pinkfloydi, Typton granulosus, Zenopontonia soror, Neogonodactylus pumilus) are new records for Costa Rica. With this contribution the total number of decapods (257 spp.) and stomatopods (14 spp.) for ACG is 271 species, more than half the species reported for Costa Rica, and more than a quarter of all crustaceans reported for the eastern tropical Pacific. The high concentration of species in ACG may be attributed to the diversity of habitats, the seasonal upwelling and to the recent sampling efforts. In only four years, the BioMar-ACG has increased the number of species in these groups of crustaceans by 37% over the past 85 years of previous studies in the ACG.


Decapods Stomatopods Biodiversity Inventory ACG 


The first study of marine decapods of Costa Rica was completed by Faxon (1895), and on stomatopods by Schmitt (1940) and Reaka and Manning (1980). Compilations of the biodiversity of marine decapod crustaceans and stomatopods in Costa Rica began with a publication by Moran and Dittel (1993), with an annotated list of anomuran and brachyuran crabs from the Pacific and Caribbean coasts. Castro and Vargas (1996) published an annotated list of decapods and stomatopods from Golfo Dulce, while Vargas and Cortés (1997) a list of stomatopods of the country. Afterwards, two other compilations on crustaceans were published for Penaeoidea, Sergestoidea, Caridea, Astacidea, Thalassinidea and Palinura in the Caribbean (Vargas and Cortés 1999a) and in the Pacific (Vargas y Cortés 1999b). Vargas and Cortés (2006) published a compilation of the Infraorder Anomura. Some years later, compilations were published in the book “Marine Biodiversity of Costa Rica, Central America” (Wehrtmann and Cortés 2009), which examines all groups of marine organisms reported for Costa Rica. Chapters include all crustacean groups known to be present in this country on the Pacific and Caribbean, including the best-known taxa: stomatopods (35 spp.) (Vargas 2009) and decapods (549 spp.) (Vargas and Wehrtmann 2009).

Área de Conservación Guanacaste (ACG) includes several terrestrial National Parks and Reserves and a Marine Sector that is 43,000 ha and 150 km of protected coastline ( It is one of the best-studied conservation areas in Central America, but previous biological research has focused primarily on the terrestrial part. In order to expand the knowledge of marine biodiversity in this region, the project entitled “Marine Biodiversity of Área de Conservación Guanacaste” (BioMar-ACG) was launched in 2015. The project has been funded by the Guanacaste Dry Forest Conservation Fund (GDFCF), and carried out by the Centro de Investigación en Ciencias del Mar y Limnología (CIMAR) and Museo de Zoología, both from Universidad de Costa Rica (UCR) (Cortés & Joyce in prep). The project has also received necessary support from the government staff of ACG, local marine “parataxonomists” supported by GDFCF and private foundations.

Following the compilation on marine biodiversity in ACG published by Cortés (2017), this paper presents an update of the inventory of marine decapod crustaceans and stomatopods, with new records for ACG as well as new records for the country. This is the result of work from mid 2015 to early 2019 by the BioMar-ACG project.


Sixty sites were visited in ACG, from Punta Descartes to Islas Murciélago (Table 1), including rocky beaches (most common) and muddy beaches, shallow and deep reefs, shallow and deep rocky areas, estuaries, mangroves and areas deeper than 45 m. The location, geographic coordinates, depth and substrate type were registered.
Table 1

Collection sites at Área de Conservación Guanacaste


Collection sites

Type of environment


Bahía El Hachal

Intertidal zone


Bajo El Jardín

Reef and algae covered coralline rocks


Bajo El Machetazo

Rocky bottom and small loose rocks


Bajo El Tigre

Rocky bottom and small loose rocks


Bajo El Viejón

Rocky bottom and small loose rocks


Bajo El Jardín

Rocky bottom and small loose rocks


Bahía Santa Elena, Pintadero, Playa Cocos

Rocky bottom and small loose rocks


Bajo Las Chavelas

Rocky bottom and small loose rocks


Bajo Los Mogotes

Rocky bottom and small loose rocks


Bajo La Rajada

Rocky bottom and small loose rocks


Bajo La Salvadita

Rocky bottom and small loose rocks


Bajo Los Muñecos

Rocky bottom and small loose rocks


Bajo Pintadero

Rocky bottom and small loose rocks


Bajo Pochote

Rocky bottom and small loose rocks


Bajo Rojo

Rocky bottom and small loose rocks


Bahía Thomas

Floating oyster baskets, rocky beach, rocky beach with sponges, live and dead Pocillopora


Bahía Cuajiniquil, Playa Cuajiniquil

Sand and mud bottoms


Isla Cocinera

Rocky intertidal zone and reefs


Isla David

Rocky bottom and small loose rocks


Isla Golondrina

Pavona clavus reef, dead coral


Isla Pelada

Rocky intertidal zone, rocks covered with algae


Esquina Conchal, Playa del Maíz, Playa Cactus, Isla San José

Rocky intertidal zone and dead Pocillopora


Arrecife al NE de Isla San Pedrito, arrecife muerto San Pedrito, San Pedrito

Coral reef and dead coral


Playa Junquillal, Junquillal esquina norte; Islote Junquillal, Junquillal, Esquina del Tamarindo

Rocky intertidal zone, polychaete reef, and Pocillopora



Coral reef, dead coral and rocky-sandy bottom


Manglar en Bahía Cuajiniquil, Manglar 3 Bocas, Manglar 4 × 4 Cuajiniquil

Mangrove, mud, rubble with sponges and algae


Manglar Santa Elena

Mangrove, muddy flats


Muelle lado derecho, muelle lado izquierdo, Cástula, Tiza, Hielera

Rocky intertidal and subtidal zones, sandy bottom, rubble


Piedra Claudio

Rocky bottom and small loose rocks with barnacles


Playa Corona

Rocky intertidal zone


Puerta de Iglesia, Isla San José

Rocky bottom


Playa Potrero Grande, Estero de Potrero Grande

Rocky intertidal zone, sandy beach, estuary, mangrove


Playa Macaya

Rocky intertidal zone, subtidal live coral


Punta Pochote

Dead corals


Playa Respingue

Rocky intertidal zone


Piedra Seca

Rocky bottom and small loose rocks with barnacles


Arrecife 2MM

Coral reef

In the rocky beaches during low tide, rocks were lifted to collect the organisms that live beneath them and the substrate was dug up to collect the organisms that live buried in the sediments. In the muddy beach areas, organisms were also collected from the mud. To collect the organisms that live within coral reefs, an indirect method was used: dead coral rocks were brought to the surface and broken apart to collect the organisms that mutually live inside. A similar method was used areas of rocky ocean floor, where collected specimens were brought to the surface. In the mangroves, specimens were collected during walks in the forest and intertidal zone, and they were taken from the roots and ground. Using a shovel, the substrate was dug up and sifted to look for those organisms that live buried in the mud. In the external part of the mangrove, a similar process was followed, but also fallen leaves and tree trunks were examined for organisms. Deep dredging sampling (down to 50 m depth) was carried out with a Van Veen dredge with a 25 × 25 cm opening. Other organisms were manually collected during dives, and some samples of crustaceans were associated with octocorals.

Collected specimens were placed in plastic containers with a net and then submerged in a bucket filled with seawater. Afterwards, the organisms were photographed in an improvised lab, a code was assigned to each specimen, a preliminary identification was provided and a sample of tissue was taken for barcoding. Once this process was finished, the organisms were sent to the Zoology Museum at Universidad de Costa Rica, where a definite identification was provided and the samples were catalogued and stored. The names of the species used are as in WoRMS (

Results and discussion

By early 2019, 2650 specimens have been collected (Table 2), corresponding to 209 species, 99 of which are new records for ACG, four could be new undescribed species, and nine are also new records for Costa Rica: Cyrtoplax panamensis (Fig. 1a), Glyptoxanthus labyrinthicus (Fig. 1b), Neogonodactylus pumilus (Fig. 1c), Pachyches marcortezensis (Fig. 1d), Petrolisthes donadio (Fig. 1e), Pylopagurus holmesi (Fig. 1f), Synalpheus pinkfloydi (Fig. 1g), Typton granulosus (Fig. 1h) and Zenopontonia soror (Fig. 1i). Cortés (2017) reported 172 species from ACG, 162 decapods and 10 stomatopods. In this study 209 species have been collected so far, 200 decapods and 9 stomatopods. Of the decapod species reported by Cortés (2017), 57 species of decapods and 5 species of stomatopod have not been collected yet. These belong mainly to organisms commonly found at depths greater than 40 m, that were collected by dredging. Taking this into account, the total number of species found in the area is 271, which includes 257 decapods and 14 stomatopods. During the last four years since the BioMar-ACG project started, 99 new records have been added to the known species from ACG (Table 2) This represents an increase of 37% over the almost 85 years of previous studies in the region.
Table 2

New records of marine decapods and stomatopods from Área de Conservación Guanacaste after four years of the BioMar-ACG project. In bold type: new records for Costa Rica




Family Albuneidae

 1) Albunea lucasia de Saussure, 1853


Family Alpheidae

 2) Alpheopsis sp. nov. A

BEJ, BTh, ISJ, Cua, Mue, Jun, BLM, MSE

 3) Alpheopsis sp. nov. B

BEJ, BTh, ISJ, Cua, Mue, Jun, BLM, MSE

 4) Alpheus bellimanus Lockington, 1877


 5) Alpheus bouvieri A. Milne-Edwards, 1878

ISJ, Jun, PCo, Cua, Mue, PRe, Pgr, MSE

 6) Alpheus cf. latus Kim & Abele, 1988


 7) Alpheus colombiensis Wicksten, 1988


 8) Alpheus cristulifrons Rathbun, 1900

ICo, Jun, ISJ, BEV, BTh, Mue, BRo

 9) Alpheus lottini Guérin-Méneville, 1838 [in Guérin-Méneville, 1829–1838]

BTh, BEH, IDa, Jun, PMa

 10) Alpheus malleator Dana, 1852

BTh, Jun, Mue, Cua, BEV

 11) Alpheus saxidomus Holthuis, 1980


 12) Alpheus tenuis Kim & Abele, 1988

MCu, PGr,

 13) Alpheus websteri Kingsley, 1880

IGo, ISJ, BTh, Cua, BLM, 2MM, BEJ

 14) Automate dolichognatha de Man, 1888

Mue, PRe

 15) Salmoneus serratidigitus (Coutière, 1898)

BTh, Cua

 16) Synalpheus cf. lockingtoni Coutière, 1909


 17) Synalpheus mexicanus Coutière, 1909


 18) Synalpheus nobilii Coutière, 1909

BTh, Cua, Jun, Mue

19) Synalpheus pinkfloydi Anker, Hultgren & De Grave, 2017


 20) Synalpheus sanjosei Coutière, 1909

BEH, BTh, ISJ, Mue, PMa, PPo

Family Axianassidae

 21) Axianassa mineri Boone, 1931

Cua, Mue

Family Axiidae

 22) Axiopsis baronai Squires, 1977


 23) Neaxius vivesi (Bouvieri, 1895)

Mue, ISJ

Family Cryptochiridae

 24) Opecarcinus crescentus (Edmondson, 1925)

BEJ, ICo, Jun

Family Diogenidae

 25) Calcinus obscurus Stimpson, 1859

BTh, Cua, ICo, ISJ, Jun, PCo, PGr, PMa, PRe

 26) Clibanarius albidigitus Nobili, 1901

ICo, Jun, MCu, MSE, Mue,

 27) Clibanarius lineatus (H. Milne Edwards, 1848)


Family Domeciidae

 28) Domecia hispida Eydoux & Souleyet, 1842


Family Epialtidae

 29) Acanthonyx petiverii H. Milne Edwards, 1834

BTh, BEM, IPe,

 30) Herbstia tumida (Stimpson, 1871)


 31) Notolopas lamellatus Stimpson, 1871


 32) Pelia pacifica A. Milne-Edwards, 1875

BTh, Jun, PGr

 33) Tyche lamellifrons Bell, 1836


Family Grapsidae

 34) Goniopsis pulchra (Lockington, 1877)


Family Hippolytidae

 35) Thor algicola Wicksten, 1987

BTh, BRo, ISJ, IGo, Jun, Cua, PGr

 36) Thor sp. nov.


Family Hymenoceridae

 37) Hymenocera picta Dana, 1852

BTh, 2MM

Family Inachidae

 38) Coryrhynchus vestitus (Stimpson, 1871)

BTh, BEJ, BLM, Cua, Jun, Mue, PCl

Family Leucosiidae

 39) Persephona townsendi Bell, 1855


 40) Uhlias ellipticus Stimpson, 1871


Family Menippidae

 41) Menippe obtusa Stimpson, 1859


Family Ocypodidae

 42) Leptuca beebei (Crane, 1941)

Cua, MCu, MSE

 43) Leptuca musica (Rathbun, 1914)


 44) Uca princeps (Smith, 1870)


 45) Uca stylifera (H. Milne Edwards, 1852)


 46) Ucides occidentalis (Ortmann, 1897)

MCu, PGr

Family Oziidae

 47) Eupilumnus xantusii (Stimpson, 1860)

BTh, BEM, ISJ, Mue

 48) Ozius perlatus Stimpson, 1860

ICo, Jun

 Family Paguridae

 49) Pagurus nanodes Haig & Harvey, 1991

BTh, IDa, Jun, Mue

 50) Phimochirus roseus (Benedict, 1892)


51) Pylopagurus holmesi Schmitt, 1921


Family Palaemonidae

 52) Ascidonia pusilla (Holthius, 1951)


 53) Harpiliopsis depressa (Stimpson, 1860)

BEH, BTh, IDa, Jun, PMa

 54) Palaemon ritteri Holmes 1895

BTh, Cua, ICo, ISJ, Mue, PCo, PRe, PGr

 55) Palaemonella holmesi (Nobili, 1907)

BTh, BLC, Bro, BEV, ISJ, Mat, Mue

 56) Pontonia mexicana Guérin-Méneville, 1855 [in Guérin-Méneville, 1855–1856]


 57) Pseudocoutierea elegans Holthuis, 1951


 58) Pseudoveleronia laevifrons (Holthuis, 1951)


59) Typton granulosus Ayon-Parente, Hendrickx & Galvan-Villa, 2015

BTh, BEV, Mue, MCu, PGr

60) Zenopontonia soror (Nobili, 1904)

BEH, BTh, IPe, Mue

Family Panopeidae

 61) Acantholobulus mirafloresensis (Abele & Kim, 1989)

BTh, BLR, Cua, Mue, MCu, PGr

 62) Lophopanopeus maculatus Rathbun, 1898


 63) Panopeus purpureus Lockington, 1877

BTh, Cua, MCu, MSE, PGr

 64) Prionoplax ciliata Smith, 1870


Family Parthenopidae

 65) Heterocrypta colombiana Garth, 1940

Cua, Mue

 66) Piloslambrus triangulus (Stimpson, 1860)


Family Penaeidae

 67) Penaeus stylirostris Stimpson, 1871


 68) Penaeus vannamei Boone, 1931


Family Pilumnidae

 69) Pilumnus limosus Smith, 1869

BTh, BEV, MCu, Mue

 70) Pilumnus stimpsoni Miers, 1886

BLM, BRo, BLS, BEV, IGo, ISP, Mue, PSe

Family Pinnotheridae

 71) Austinixa sp. indet


 72) Pinnixa sp. indet


 73) Pinnotheres sp. indet


 74) Tumidotheres sp. indet


Family Plagusiidae

 75) Plagusia immaculata Lamarck, 1818

BTh, PRe, PGr

Family Porcellanidae

 76) Megalobrachium erosum (Glassell, 1936)

BTh, Cua

 77) Neopisosoma mexicanum (Streets, 1871)

Jun, PGr

 78) Pachycheles calculosus Haig, 1960


79) Pachycheles marcortezensis Glassell, 1936


 80) Pachycheles spinidactylus Haig, 1957

ISJ, Jun, 2MM

81) Petrolisthes donadio Hiller & Werding, 2007

BTh, BLM, IGo, ISJ, ISP, Jun, Mat, Mue, Pse, PPo

 82) Polyonyx confinis Haig, 1960

Jun, Mue

 83) Ulloaia perpusillia Glassell, 1938


Family Portunidae

 84) Achelous tuberculatus Stimpson, 1860


Family Processidae

 85) Processa peruviana Wicksten, 1983

Cua, ISJ, Mue

Family Pseudorhombilidae

86) Cyrtoplax panamensis Ziesenhenne in Garth, 1940


Familia Raninidae

 87) Raninoides benedicti Rathbun, 1935


Family Rhynchocinetidae

 88) Cinetorhynchus sp. nov.


Family Sesarmidae

 89) Sesarma sulcatum Smith, 1870


Family Trapeziidae


 90) Trapezia digitalis Latreille, 1828

BEH, BTh, IDa, Jun, Mat, PMa

Family Upogebiidae


 91) Upogebia thistlei Williams, 1986

BTh, BSE,Jun, Mue

Family Xanthidae


92) Glyptoxanthus labyrinthicus (Stimpson, 1860)


 93) Lipaesthesius leeanus Rathbun, 1898


 94) Lipkemedaeus spinulifer (Rathbun, 1898)


 95) Platypodiella rotundata (Stimpson, 1860)

BTh, BEM, BRo, Cua, ICo, IGo, ISJ, Jun, MCu, Mue, PGr


 Family Gonodactylidae

96) Neogonodactylus pumilus (Manning, 1970)


 97) Neogonodactylus stanschi (Schmitt, 1940)

BEV, Mat

Family Squillidae

 98) Cloridopsis dubia (H. Milne Edwards, 1837)


 99) Meiosquilla dawsoni Manning, 1970


Fig. 1

New records of crustaceans for Costa Rica, with their BioMar-ACG sample code: (a) Cyrtoplax panamensis,, 17-BMACGRV-02644-ACG001862; (b) Glyptoxanthus labyrinthicus, 16-BMACGRV-02104-ACG005676; (c) Neogonodactylus pumilus, 16-BMACGRV-02407-ACG006086; (d) Pachyches marcortezensis, 15-BMACGRV-00507-ACG006967; (e) Petrolisthes donadio, 16-BMACGRV-02177-ACG009006; (f) Pylopagurus holmesi, 16-BMACGRV-02414-ACG007167; (g) Synalpheus pinkfloydi, 16-BMACGRV-02245-ACG003908; (h) Typton granulosus, 16-BMACGRV-01941-ACG001783, and (i) Zenopontonia soror, 15-BMACGRV-00590-ACG009037

The number of species of decapods and stomatopods reported for ACG (271) is the highest number found in Costa Rica and among the highest reported in the eastern tropical Pacific ETP (Table 3). The ETP extends from the Gulf of California to southern Ecuador and include several oceanic islands (Robertson and Kramer 2009). ACG has more than half the species reported for Costa Rica and about a quarter of decapods and stomatopods crustaceans reported for the ETP (Boschi 2000; Cortés et al. 2017) (Table 3). The high diversity of decapods and stomatopods at ACG may be due to the diversity of habitats: beaches of different wave and sediment regimes, islands and continental shorelines, mangrove forests, seagrass beds, coral reefs (live and dead), intertidal and submerged rocky platforms, sandy and muddy bottoms and deep areas (Cortés 2017). The region is exposed to season upwelling, with temperatures ranging from 15° to 30 °C (Cortés et al. 2014). And finally, the sampling effort makes a difference in how many species are reported from an area, as demonstrated in this paper.
Table 3

Richness of stomatopods and decapods in the eastern tropical Pacific (ETP)










Hendrickx 2005a, 2005b; M.E. Hendrickx, personal communication, 2019

Eastern Pacific




Boschi 2000; Cortés et al. 2017; Salgado-Barragán & Hendrickx 2010

Costa Rica




Vargas 2009; Vargas & Wehrtmann 2009





Lemaitre & Álvarez-León 1992; López & Jaimes 2014

ACG, Costa Rica




This study

Oaxaca, México




Bastida-Zavala et al. 2013

Galápagos, Ecuador




Hickman & Zimmermn 2000

Clipperton, France




Poupin et al. 2009

Isla del Coco, Costa Rica




Vargas-Castillo & Wehrtmann 2008; Cortés 2012

Bahía Culebra, Costa Rica




Cortés et al. 2012

Golfo Dulce, Costa Rica




Morales-Ramírez 2011; Castro & Vargas 1996

aIncludes pelagic and deep and shallow water species (M.E. Hendrickx, personal communication, 2019)

bIncludes only shallow water species

Up to now 60 sites have been sampled, with many more still to explore, such as the coastal area of Península Santa Elena, south of Playa Naranjo and offshore, as well as some habitats such as sandy beaches, and deep rocky and muddy bottoms. Definitely more species of decapods and stomatopods will be found increasing the biodiversity of crustaceans in Área de Conservación de Guanacaste.


El sector marino del Area de Conservación Guanacaste (ACG) posee 43,000 ha y 150 km de costa, en su mayoría poco estudiada. En un esfuerzo por conocer mejor la biodiversidad de todos los taxones marinos presentes en el área, se inicio en el 2015 un inventario (Proyecto BioMar-ACG). Esta iniciativa está siendo sustentada por la Guanacaste Dry Forest Conservation Fund (GDFCF) y desarrollada por el Centro de Investigación en Ciencias del Mar y Limnología (CIMAR) y el Museo de Zoología, ambos de la Universidad de Costa Rica (UCR). Transcurridos cuatro años del proyecto, se han recolectado 2650 especímenes de crustáceos decápodos y estomatópodos marinos pertenecientes a 209 especies, de las cuales 99 son nuevas para el ACG, cuatro podrían ser nuevas especies para la ciencia y nueve (Cyrtoplax panamensis, Glyptoxanthus labyrinthicus, Pachyches marcortezensis, Petrolisthes donadio, Pylopagurus holmesi, Synalpheus pinkfloydi, Typton granulosus, Zenopontonia soror y Neogonodactylus pumilus) son ampliaciones de ámbito y nuevos informes para el país. Con esta contribución aumenta el número de decápodos a 257 spp. y de estomatópodos a 14 spp. para un total de 271 spp. para ACG. Esto es más de la mitad las especies conocidas de estos grupos para Costa Rica y más una cuarta parte de las especies conocidas para el Pacífico Tropical Oriental. En cuatro años el proyecto BioMar-ACG ha aumentado el número de especies de decápodos y estomatópodos del ACG en 37% más que lo que se conocía a partir de estudios en los últimos 85 años.


The model for marine biodiversity inventory developed with the BioMar-ACG project, the partnership between government, academia, private funding and local parataxonomists, accelerates the rate of species discovery and reporting. It also makes species information and project results available in an open access format. In only for years of the project the number of decapods and stomatopods of ACG increased by 37% over the past 85 years of previus studies, for a total of 271 species. This number represnets more than half the species reported for Costa Rica, and more than a quarter of all decapods and stomatopods reported for the eastern tropical Pacific.



The authors acknowledge and thank the following people and organizations for their efforts and significant support to the BioMar-ACG project: Daniel Janzen, Winnie Hallwachs, Frank Joyce, María Marta Chavarría, Roger Blanco, Eric Palola, Yelba Vega, Gilberth Ampie, Diving Center Cuajiniquil, the Santa Elena Lodge, the Guanacaste Dry Forest Conservation Fund, the Wege Foundation, the New England BioLabs Foundation, the Wallace Genetic Foundation and the Centre for Biodiversity Genomics, University of Guelph. We thank Francilena Carranza for preparing the photographs and Eric Palola and anonymous reviewers for the review of the manuscript. Michel E. Hendrickx was very helpful in the preparation of the reviewed draft of the paper.

Authors’ contributions

RV collected and identified the specimens, wrote a first draft of the manuscript. JC, conceived the main project, helped with logistics and obtaining the funds, wrote some sections of the paper, helped prepared the images and the final draft of the paper. All authors read and approved the final manuscript.


Funding source indicated in the Acknowledgement section.

Ethics approval and consent to participate

Not applicable.

Consent for publication

Not applicable.

Competing interests

The authors declare that they have no competing interests.


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Authors and Affiliations

  1. 1.Museo de ZoologíaUniversidad de Costa RicaSan PedroCosta Rica
  2. 2.Centro de Investigación en Ciencias del Mar y Limnología (CIMAR)Universidad de Costa RicaSan PedroCosta Rica
  3. 3.Escuela de BiologíaUniversidad de Costa RicaSan PedroCosta Rica

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