A fjord-like tropical ecosystem, Pacific coast of Costa Rica: overview of research in Golfo Dulce / Un ecosistema tropical similar a un fiordo, costa del Pacífico de Costa Rica: panorama de la investigación en el Golfo Dulce

Abstract Introduction: Coastal ecosystems worldwide are under the influence of local, regional and global stressors, such as pollution, eutrophication and climate change. Golfo Dulce is a relatively pristine and accessible deep tropical ecosystem that provides opportunities for comparative and collaborative research. Objective: To summarize published reports on past research conducted in this ecosystem, identify topics for further study, and suggest new research issues. Methods: A search was made on the web for reports based on research conducted in Golfo Dulce and published in scientific journals. Reports focusing on environmental parameters and on the biota were included. Results: A total of 123 studies that include data from Golfo Dulce are cited. The four topics more frequently addressed were reports based on the results of the R/V Victor Hensen expedition (1993-1994) and follow-up work on microbiology, studies on water parameters, research on vertebrates, and zooplankton studies. The reports focusing on vertical profiles of oxygen and temperature are discussed in detail, followed by those on the biota. Conclusions: Golfo Dulce has low oxygen concentrations below 50 m and is frequently anoxic at the 200 m deep basin with occasional formation of H2S. However, the ecosystem contains a relatively high diversity of identified organisms, from bacteria to whales. Of particular relevance for future studies are multidisciplinary surveys aiming at obtaining data on primary productivity, the diversity and biomass of the main groups of planktonic, demersal and benthic organisms, and the frequency and magnitude of the influx of deep offshore waters over the sill into the basin. These data, as well as the information gathered in the past, are essential for updating the trophic model developed more than 25 years ago and in support of new predictive models on the functioning of the ecosystem.

Resumen Introducción: Los ecosistemas costeros alrededor del mundo están bajo la influencia de tensores locales, regionales y globales, como la contaminación, la eutroficación y el cambio climático. El Golfo Dulce es un profundo ecosistema tropical relativamente inalterado y accesible, que provee oportunidades para la investigación comparativa y colaborativa. Objetivo: Resumir los informes publicados sobre investigaciones pasadas realizadas en el ecosistema, identificar tópicos para estudios futuros y sugerir nuevas áreas de investigación. Métodos: Se hizo una búsqueda en la red de informes basados en investigaciones hechas en Golfo Dulce y publicadas en revistas científicas. Fueron incluidos aquellos informes sobre parámetros ambientales y la biota. Resultados: Un total de 123 estudios que incluyen datos sobre Golfo Dulce son citados. Los cuatro tópicos citados con más frecuencia fueron: Los resultados de la expedición del R/V Victor Hensen (1993-1994) y estudios de seguimiento sobre microbiología, informes sobre parámetros acuáticos, investigaciones sobre vertebrados y estudios sobre zooplancton. Los informes sobre perfiles de oxígeno y temperatura son presentados con mayor detalle, seguidos por aquellos sobre la biota. Conclusiones: Golfo Dulce tiene bajas concentraciones de oxígeno por debajo de 50 m y es usualmente anóxico a 200 m en el fondo, con formación ocasional de H2S. Sin embargo, el ecosistema contiene una diversidad de organismos identificados relativamente alta, desde bacterias hasta ballenas. De relevancia particular para futuros estudios es, entre otros, la conducción, de muestreos multidisciplinarios orientados a obtener datos sobre productividad primaria, la diversidad y biomasa de los principales grupos de organismos planctónicos, demersales y bénticos, así como la frecuencia y magnitud del flujo de agua oceánica hacia el interior. Estos datos, así como los obtenidos en el pasado, son esenciales para actualizar el modelo trófico desarrollado hace más de 25 años, o en apoyo de nuevos modelos predictivos de funcionamiento del ecosistema.

Peces del Golfo de Nicoya, Pacífico, Costa Rica: actualización de las listas de los buques científicos Skimmer y Victor Hensen

Introducción: Desde mediados del siglo XX el estuario del Golfo de Nicoya (10o N & 85o W) ha sido la fuente principal de numerosos recursos pesqueros, como los moluscos, crustáceos y peces. El estuario está actualmente bajo impactos crecientes locales, regionales y globales. Las expediciones a bordo de los buques científicos Skimmer (1979-1980) y Víctor Hensen (1993-1994) aportaron datos multidisciplinarios en apoyo de políticas de manejo. Sin embargo, actualmente se necesita información actualizada. Objetivos: Actualizar las listas publicadas de peces recolectados en ambas expediciones y sugerir nuevas áreas de investigación para una futura evaluación del estado del ecosistema. Métodos: En el buque Skimmer las capturas de peces cubrieron un ámbito de profundidad de 10 a 50 m, con una red tipo otter. En el buque V. Hensen las capturas se hicieron entre 10 y 240 m de profundidad, con redes tipo otter y beam. En ambos cruceros se hizo a bordo la identificación de las especies y la medición de sus longitudes totales. Para actualizar las listas de especies se procedió a la verificación de los nombres vigentes con base en literatura reciente y la página World Register of Marine Species (WORMS). Agregamos a la lista los nombres de las especies identificadas en años recientes para el estuario y catalogados en la Colección Ictiológica del Museo de Zoología, Universidad de Costa Rica. Resultados: El total de especies identificadas es de 268 para ambos buques. Una especie de Syacium está pendiente de identificación. Del total, 19 especies corresponden a peces cartilaginosos y 249 a peces óseos. Los Scianeidae (corvinas) y los lenguados estuvieron representados por 31 y 32 especies, respectivamente. El género Symphurus (lenguados) fue el más diverso, con 11 especies. Dieciocho especies fueron identificadas después de la publicación de las listas originales. Ocho especies fueron descritas con base en ejemplares recolectados por ambos buques: Gymnothorax phalarus, Notarius biffi, N. cookei, Ophicthus mecoptera, O. melope, Sphoeroides rosenblatti, Trinectes xanthurus y Urotrygon cimar. Las especies presentes en más del 50 % de las 20 estaciones del Skimmer, fueron: Prionotus horrens (16), Sphoeroides trichocephalus (14), Syacium ovale (13) y Synodus scituliceps (13). En las 12 estaciones del V. Hensen, fueron: Porichthys margaritatus (9), S. scituliceps (8), Bollmannia chlamydes (6), Symphurus callopterus (6), S. elongatus (6) y Synodus evermanni (6). Los cinco peces de mayor longitud capturados por ambos buques, fueron: la raya látigo Hypanus longus (117 cm), el congrio Cynoponticus coniceps (103 cm), el bagre Sciades dowii (87 cm), la raya Narcine brancroftii (80 cm) y la corvina reina Cynoscion albus (75 cm). La longitud total máxima promedio de 137 especies medidas en el Skimmer fue de 26 cm y de 29 cm para las 164 medidas en el V. Hensen. Conclusiones: En vista del incremento del número de usuarios de los recursos estuarinos, es urgente una evaluación multidisciplinaria del ecosistema en apoyo a la propuesta de nuevas políticas de manejo. Los resultados facilitarían actualizar el Modelo Trófico del Golfo de Nicoya, una herramienta de manejo disponible desde 1998 y basada principalmente en datos obtenidos durante los cruceros del Skimmer y V. Hensen. Tensores del ecosistema, como la extracción de recursos, el ingreso de contaminantes y su posible relación con la disrupción endocrina, así como la presencia de barreras naturales como la hipoxia en la boca del estuario, deben ser incluidos en esa futura evaluación.

Introduction: Since the middle of the XXth century the Gulf of Nicoya estuary (10o N & 85o W) has been the main source of numerous fishing resources, such as mollusks, crustaceans, and fish. The estuary is presently under the increasing impact of local, regional and global stressors. The expeditions on board the R.V. Skimmer (1979-1980) and R.V. Hensen (1993-1994) research vessels provided multidisciplinary data on the estuary in support of management policies. However, updated information is still needed. Objectives: To update the published lists of fish species from both expeditions and make suggestions for new research areas for future evaluations of the ecosystem. Methods: The fish collections by the Skimmer covered a depth range of 10 to 50 m using an otter trawl net, and those aboard the Victor Hensen were made between 10 and 228 m depth using otter and beam trawl nets. The identification of the fish species and the measurement of its lenghts were made on board of both vessels. The species names were updated based on recent literature and the web page World Register of Marine Species (WORMS). The list of species identified in recent years based on voucher specimens from the estuary and deposited in the Ichthyology Collection of the Museum of Zoology, University of Costa Rica, was also included. Results: The total number of identified species from both vessels is 268. A species of Syacium is yet undescribed. Of the total, 19 are cartilaginous fishes and 249 are bony fishes. The Sciaenidae and the flatfishes were represented by 31 and 32 species, respectivley. The flatfish genus Symphurus was the most diverse, with 11 species. Eighteen species were identified after the publication of the original lists. Eight species were described based on specimens collected during the expeditions: Gymnothorax phalarus, Notarius biffi, N. cookei, Ophicthus mecoptera, O. melope, Sphoeroides rosenblatti, Trinectes xanthurus and Urotrygon cimar. The species present in more than 50% of the 20 Skimmer stations, were: Prionotus horrens (16 stations), Sphoeroides trichocephalus (14), Syacium ovale (13) and Synodus scituliceps (13). In the 12 stations of the V. Hensen there were: Porichthys margaritatus (9), S. scituliceps (8), Bollmannia chlamydes (6), Symphurus callopterus (6), S. elongatus (6), and Synodus evermanni (6). The five longer specimens collected by both vessels, were: the longtail sting ray Hypanus longus (117 cm), the pike conger Cynoponticus coniceps (103 cm), the brown sea catfish Sciades dowii (87 cm), the electric ray Narcine brancroftii (80 cm), and the queen corvina Cynoscion albus (75 cm). The average total length for 137 species measured during the Skimmer cruises was 26 cm, and 29 cm for the 164 measured during the V. Hensen survey. Conclusions: In view of the increasing number of users of the estuarine resources, it is urgent a multidisciplinary evaluation of the ecosystem in support of new management policies. The expected results may facilitate updating the Gulf of Nicoya Trophic Model, a management tool available for the estuary since 1998 and based mainly on information gathered during both expeditions. Ecosystem stressors, such as the extraction of resources, the input of pollutants and its possible link to endocrine disruption, as well as the presence of natural barriers such as hypoxia at deeper waters at the mouth of the estuary must be taken into account in a future evaluation.

Crustáceos (Decapoda y Stomatopoda) del R.V. Skimmer y R.V. Victor Hensen en el Golfo de Nicoya, Pacífico, Costa Rica / Crustaceans (Decapoda and Stomatopoda) from the R.V. Skimmer and R.V. Victor Hensen in the Gulf of Nicoya, Pacific, Costa Rica

Resumen El acceso a datos actualizados, sobre los nombres y distribución de las especies de crustáceos presentes en un área en particular, es un primer paso para evaluar cambios debidos a factores locales, regionales o globales, como la sobrepesca, la contaminación y el cambio climático. Los datos en este estudio están basados en las expediciones de los buques de investigación Skimmer (20 estaciones, 1979-1980) y Victor Hensen (1993-1994, 12 estaciones) en el estuario del Golfo de Nicoya. Los crustáceos fueron recolectados mediante redes de arrastre tipo Otter (Skimmer, malla 3.5 cm) y por tipos Otter y Beam (V. Hensen, mallas de 2.5 cm y 1.9 cm) a profundidades entre 10 y 228 m. Datos en dos estudios posteriores aportaron información para una zona de entre-mareas en la región superior del Golfo y para estaciones en la boca del estuario, lo que incrementa el ámbito de profundidad de 0 a 350 m. Se revisaron las publicaciones originales y se hicieron 32 actualizaciones de los nombres de las especies con base en la literatura reciente y en la página de red World Register of Marine Species (WORMS). El total de especies para ambas expediciones fue de 131, del que 119 fueron decápodos y 12 fueron estomatópodos. Los datos en los otros dos estudios, así como la revisión de las colecciones en el Museo de Zoología de la Universidad de Costa Rica, incrementó en 43 el número de registros, para un total de 174 especies en los sedimentos del estuario. Para el Skimmer el número mínimo de especies en una estación fue de cuatro (tres estaciones) y el máximo de 27, con un promedio de 12.3 especies / estación. Para el V. Hensen el mínimo de especies en una estación fue de ocho, con un máximo de 27 y un promedio de 17 especies / estación. Las especies presentes en 50 % o más de las 20 estaciones de la expedición del Skimmer, fueron: Callinectes arcuatus, Rimapenaeus faoe, Penaeus brevirostris, Achelous asper y Hepatus kossmanni. Para la expedición del V. Hensen las especies presentes en más del 50 % de las 12 estaciones, fueron: A. asper, Sicyonia disdorsalis, S. picta y Persephona subovata. Durante el estudio del Skimmer, un total de 15 especies fueron encontradas en solo una estación, mientras que para el V. Hensen el número fue de 26. Se digitalizaron dos matrices de datos de presencia-ausencia de las especies en las estaciones y con base en los nombres actualizados de las especies recolectadas por el Skimmer (57 especies x 20 estaciones) y V. Hensen (82 especies x 12 estaciones) se les aplicó un Análisis No Métrico Dimensional de Escala (NMDS) para obtener una distribución de las estaciones en un espacio bi-dimensional. Los resultados revelaron unos grupos heterogéneos de estaciones. Algunos sub-grupos de tres o cuatro estaciones concuerdan con su proximidad geográfica. Las cuatro estaciones del V. Hensen en la boca del estuario a profundidades mayores a los 60 m fueron separadas más claramente de las otras, y podrían indicar una transición, desde aguas estuarinas hacia aguas profundas, en la composición de la fauna de crustáceos. Evaluaciones futuras de la diversidad de crustáceos del Golfo de Nicoya deben de considerar la amplia distribución espacial de algunas especies y la restringida de otras. La variabilidad temporal es también importante en el estuario, tal como lo evidenció la oscilación de la población de Pinnixulala valerii en un periodo de tres años.(AU)

Abstract The access to updated data on the names of the crustacean species and their distribution in a given area is a first step to evaluate changes due to local, regional and global factors such as overfishing, pollution, and climatic change. Data in this study are based on the expeditions of the RV Skimmer (20 stations, 1979-1980) and RV Victor Hensen (1993-1994, 12 stations) in the Gulf of Nicoya estuary. The crustaceans were collected by means of an Otter trawl (Skimmer, mesh 3.5 cm) and by Otter and Beam trawls (V. Hensen, mesh 2.5 cm and 1.0 cm, respectively) at depths from 10 m to 228 m. Data from two later studies were also included, one from an intertidal flat in the upper Gulf and the other from stations at the mouth of the estuary, which expand the depth range from 0 to 350 m. The list of species in the original publications were updated and 32 corrections were made based on recent literature and the web page, World Register of Marine Species (WORMS). The total number of species for both surveys was 131, of which 119 were decapods and 12 were stomatopods. Data from the other two studies and from the crustacean collection deposited at the University of Costa Rica Zoology Museum added 43 records for a total of 174 species collected in sediments from the estuary. For the Skimmer, the minimum number of species found in one station was four (three stations) and the maximum was 27, with an average of 12.3 species / station. For the V. Hensen, the minimum of species found in one station was eight, with a maximm of 27 and an average of 17 species / station. The species present in 50 % or more of the 20 stations of the Skimmer expedition, were: Callinectes arcuatus, Rimapenaeus faoe, Penaeus brevirostris, Achelous asper and Hepatus kossmanni. For the V. Hensen expedition, the species present in 50 % or more of the 12 stations, were: A. asper, Sicyonia disdorsalis, S. picta, and Persephona subovata. During the Skimmer survey a total of 15 species were found at only one station, while for the V. Hensen the number was 26. Two presence-absence matrices based on the updated names of the crustaceans collected by the Skimmer (57 species x 20 stations) and V. Hensen (82 species x 12 stations) were analyzed by Non Metric Dimensional Scaling (NMDS) to display the distribution of stations in a two-dimensional space. The results revealed heterogeneous groups of stations. Several sub-groups of two or three stations agreed with their geographical proximity. The four V. Hensen stations, located at the mouth of the estuary at depths greater than 60 m, were separated more clearly from the others and may indicate a transition, from estuarine to deep waters, in the composition of the crustacean fauna. Future evaluations of the crustacean diversity of the Gulf of Nicoya must take into account the wide spatial distribution of some species and the patchy distribution of others. Temporal variability is also important in the estuary as evidenced by the population oscilallations of Pinnixulala valerii over a three year period.(AU)

Medusas (Cnidaria: Hydrozoa) de una zona de afloramiento costero, Bahía Culebra, Pacífico, Costa Rica / Medusae (Cnidaria: Hydrozoa) from a coastal upwelling zone, Culebra Bay, Pacific, Costa Rica

The hydromedusae have an important role in marine trophic webs due to their predatory feeding habits. This is the first study of this group of gelatinous marine zooplankton in a coastal upwelling area of Central America. The composition and abundance variability of hydromedusae were studied during six months in 1999 at four stations in Culebra Bay, Gulf of Papagayo, Pacific coast of Costa Rica (10º37’ N-85º40’ W). A total of 53 species were identified, of which 26 are new records for Costa Rica, 21 are new records for Central America, and eight are new records for the Eastern Tropical Pacific. The more abundant species (more than 30% of the total abundance) were Liriope tetraphylla, Solmundella bitentaculata and Aglaura hemistoma. Six species occurred throughout the sampling period, 10 were present only during the dry season (December-April), and 17 were so during the rainy season (May-November). Significant differences of medusan abundances were found between seasons (dry vs. rainy). Maximum abundance (2.1±4.3ind./m³) was recorded when upwelled deeper water influenced the Bay, as indicated by local higher oxygen concentrations and lower water temperatures. The relatively high species richness of medusae found in Culebra Bay is probably related to factors like the pristine condition of the Bay, the arrival of oceanic species transported by the Equatorial Counter Current (ECC), the eastward shoaling of the Costa Rica Dome, and local currents. Illustrations of the 15 more important species are included to facilitate their identification and foster future work in the region.

Las hidromedusas tienen un papel importante en redes tróficas marinas debido a sus hábitos alimenticios depredadores. Este es el primer estudio de este grupo del zooplancton gelatinoso en un area de afloramiento costero en América Central. Durante seis meses de 1999, se estudió la abundancia de hidromedusas en cuatro estaciones en Bahía Culebra, Golfo de Papagayo, costa Pacífica de Costa Rica (10º 37’N-85º40’W). Se identificó un total de 53 especies de las que 26 son registros nuevos para Costa Rica, 21 son registros nuevos para América Central y 8 son nuevos registros para el Pacífico Oriental Tropical. Las especies más abundantes durante el estudio (con más de 30% de la abundancia total) fueron Liriope tetraphylla, Solmundella bitentaculata y Aglaura hemistoma. Seis especies se presentaron en todos los muestreos, 10 especies se presentaron únicamente durante la época seca y 17 se presentaron únicamente durante la época lluviosa. Se mostraron diferencias significativas entre las épocas lluviosa y seca. La máxima abundancia promedio de hidromedusas (2.1±4.3 ind./m³) fue encontrada durante las fechas que se espera el afloramiento costero, indicado por alta concentración de oxígeno y baja temperatura. La rica composición de especies encontrada en Bahía Culebra puede ser el resultado de varios factores, incluyendo la condición prístina de la bahía, el transporte de especies por la Contra Corriente Nor-Ecuatorial (NECC) y los aportes de origen terrestre. Se incluyen ilustraciones de las 15 especies más importantes para facilitar su identificación y promover estudios futuros en la región.

Crustaceans from a tropical estuarine sand-mud flat, Pacific, Costa Rica, (1984-1988) revisited

The availability of data sets for time periods of more than a year is scarce for tropical environments. Advances in hardware and software speed-up the re-analysis of old data sets and facilitates the description of population oscillations. Using recent taxonomic literature and software we have updated and re-analized the information on crustacean diversity and population fluctuations from a set of cores collected at a mud-sand flat in the mid upper Gulf of Nicoya estuary, Pacific coast of Costa Rica (1984-1988). A total of 112 morphological species of macroinvertebrates was found, of which 29 were crustaceans. Taxonomic problems, maily with the peracarids, prevented the identification of a group of species. The abundance patterns of the crab Pinnixa valerii, the ostracod Cyprideis pacifica, and the cumacean Coricuma nicoyensis were analized with the Generalized Additive Models of the free software R. The models evidenced a variety of population oscillations during the sampling period. These oscillations probably included perturbations induced by external factors, like the strong red tide events of 1985. In additon, early on 1984 the populations might have been at an altered state due to the inpact of El Niño 1982-83. Thus, the oscillations observed during the study period departed from the expected seasonality (dry vs rainy) pattern and are thus considered atypical for this tropical estuarine tidal-flat. Crustacean diversity and population peaks were within the range of examples found in worldwide literature. However, abundances of the cumacean C. nicoyensis, an endemic species, are the highest reported for a tropical estuary. Comparative data from tropical tidal flat crustaceans continues to be scarce. Crustaceans (total vs groups) had population changes in response to the deployment of predator exclusion cages during the dry and rainy seasons of 1985. Temporal and spatial patchiness characterized the abundances of P. valeri, C. pacifica and C. nicoyenis.

La disponibilidad de grupos de datos recolectados por periodos mayores a un año es escasa para ambientes marinos tropicales. Avances en hardware y software pueden acelerar el re-análisis de grupos antiguos de datos y facilitar la descripción de oscilaciones poblacionales ocultas. Mediante el uso de literatura taxonómica y programas de cómputo recientes actualizamos y re-analizamos la información sobre diversidad de crustáceos y oscilaciones poblacionales en un grupo muestras recolectadas con barreno en una planicie arenoso-fangosa en la región media superior del estuario del Golfo de Nicoya, costa Pacífica de Costa Rica (1984-1988). Se encontró un total de 112 morfo-especies de macro-invertebrados, de las que 29 fueron crustáceos. Problemas taxonómicos, especialmente con los peracáridos, impidieron la identificación completa de un grupo de especies. Las oscilaciones en la abundancia del cangrejo Pinnixa valerii, el ostrácodo Cyprideis pacifica, y el cumáceo Coricuma nicoyensis fueron analizadas con los Modelos Generales Aditivos (GAM) del programa de cómputo R de acceso libre. Los modelos evidenciaron una variedad de oscilaciones poblacionales durante el periodo de toma de muestras. Estas oscilaciones probablemente incluyen perturbaciones inducidas por factores externos, como los fuertes eventos de mareas rojas de 1985. Además, al inicio de 1984 las poblaciones podrían haber estado alteradas debido al impacto de El Niño 1982-83. Entonces, las oscilaciones observadas durante el estudio se alejan del patrón estacional (seco vs lluvioso) y son consideradas atípicas para esta planicie tropical estuarina arenoso-fangosa. La diversidad de crustáceos y las oscilaciones poblacionales fueron similares a las publicadas en la literatura mundial. Sin embargo, la abundancias del cumáceo Coricuma nicoyensis, una especie endémica, son las más altas informadas para un estuario tropical. Datos comparativos sobre planicies arenoso-fangosas en el trópico continúan siendo escasos. Los crustáceos (total vs grupos) tuvieron cambios poblacionales en respuesta a la colocación de jaulas con malla para excluir macro-depredadores durante las estaciones seca y lluviosa de 1985. Parches en el tiempo y el espacio caracterizaron a las abundancias de P. valeri, C. pacifica y C. nicoyenis.

[Medusae (Cnidaria: Hydrozoa) from a coastal upwelling zone, Culebra Bay, Pacific, Costa Rica]. / Medusas (Cnidaria: Hydrozoa) de una zona de afloramiento costero, Bahía Culebra, Pacífico, Costa Rica.

The hydromedusae have an important role in marine trophic webs due to their predatory feeding habits. This is the first study of this group of gelatinous marine zooplankton in a coastal upwelling area of Central America. The composition and abundance variability of hydromedusae were studied during six months in 1999 at four stations in Culebra Bay, Gulf of Papagayo, Pacific coast of Costa Rica (10 degrees 37' N-85 degrees 40' W). A total of 53 species were identified, of which 26 are new records for Costa Rica, 21 are new records for Central America, and eight are new records for the Eastern Tropical Pacific. The more abundant species (more than 30% of the total abundance) were Liriope tetraphylla, Solmundella bitentaculata and Aglaura hemistoma. Six species occurred throughout the sampling period, 10 were present only during the dry season (December-April), and 17 were so during the rainy season (May-November). Significant differences of medusan abundances were found between seasons (dry vs. rainy). Maximum abundance (2.1 +/- 4.3ind./m3) was recorded when upwelled deeper water influenced the Bay, as indicated by local higher oxygen concentrations and lower water temperatures. The relatively high species richness of medusae found in Culebra Bay is probably related to factors like the pristine condition of the Bay, the arrival of oceanic species transported by the Equatorial Counter Current (ECC), the eastward shoaling of the Costa Rica Dome, and local currents. Illustrations of the 15 more important species are included to facilitate their identification and foster future work in the region.

Crustaceans from a tropical estuarine sand-mud flat, Pacific, Costa Rica, (1984-1988) revisited.

The availability of data sets for time periods of more than a year is scarce for tropical environments. Advances in hardware and software speed-up the re-analysis of old data sets and facilitates the description of population oscillations. Using recent taxonomic literature and software we have updated and re-analized the information on crustacean diversity and population fluctuations from a set of cores collected at a mud-sand flat in the mid upper Gulf of Nicoya estuary, Pacific coast of Costa Rica (1984-1988). A total of 112 morphological species of macroinvertebrates was found, of which 29 were crustaceans. Taxonomic problems, maily with the peracarids, prevented the identification of a group of species. The abundance patterns of the crab Pinnixa valerii, the ostracod Cyprideis pacifica, and the cumacean Coricuma nicoyensis were analized with the Generalized Additive Models of the free software R. The models evidenced a variety of population oscillations during the sampling period. These oscillations probably included perturbations induced by external factors, like the strong red tide events of 1985. In additon, early on 1984 the populations might have been at an altered state due to the inpact of El Niño 1982-83. Thus, the oscillations observed during the study period departed from the expected seasonality (dry vs rainy) pattern and are thus considered atypical for this tropical estuarine tidal-flat. Crustacean diversity and population peaks were within the range of examples found in worldwide literature. However, abundances of the cumacean C. nicoyensis, an endemic species, are the highest reported for a tropical estuary. Comparative data from tropical tidal flat crustaceans continues to be scarce. Crustaceans (total vs groups) had population changes in response to the deployment of predator exclusion cages during the dry and rainy seasons of 1985. Temporal and spatial patchiness characterized the abundances of P. valeri, C. pacifica and C. nicoyenis.

Molluscan assemblage from a tropical intertidal estuarine sand-mud flat, Gulf of Nicoya, Pacific, Costa Rica (1984-1987).

The availability of data sets covering more than a year is scarce for tropical environments. Advances in hardware and software speed-up the re-analysis of old data sets and facilitates the identification of hidden data patterns. From February 1984 to April 1987 (49 sampling dates), core samples (17.7cm2, 15cm deep) were collected at low tide at a sand-mud flat in the mid upper Gulf of Nicoya estuary, Pacific, Costa Rica. Predator exclusion experiments (cages 0.5 x 0.5 x 0.2m, galvanized wire, mesh size 5mm), were conducted at the site in 1985 (dry and rainy seasons sets). Samples were preserved with 5% buffered formalin in sea water stained with Rose Bengal, and washed after 24 hours on a 500 micron mesh sieve. The 1 120 cores yielded a total of 112 morphological species of which the mollusks were represented by 23 species, and included the bivalves Tellina rubescens, Tagelus bourgeoisae, Dosinia dunkeri and Leukoma asperrima, and the gastropods, Natica unifasciata, Nassarius luteostomus, Costoanachis rugosa and Turbonilla sp. The 23 species are indicative of a relatively rich sedimentary molluscan fauna. T. bourgeoisae had a seasonal oscillation, with higher abundances during the rainy seasons. T. rubescens was not seasonal, but presented an oscillation with peaks at about 1.5 year intervals. Many empty shells of Cosmioconcha modesta, lower number of N. luteostomus and a few of T. rubescens were found with boreholes by the predatory snail N. unifasciata. T. rubescens was not significantly more abundant inside or outside cages. T. bourgeoisae showed a significant increase within the caged areas. The numerical fluctuations of the mollusks became more important during the rainy season of 1985. Red tide outbreaks in the Gulf of Nicoya in 1985 may have had an impact on the molluscan populations.

Molluscan assemblage from a tropical intertidal estuarine sand-mud flat, Gulf of Nicoya, Pacific, Costa Rica (1984-1987)

The availability of data sets covering more than a year is scarce for tropical environments. Advances in hardware and software speed-up the re-analysis of old data sets and facilitates the identification of hidden data patterns. From February 1984 to April 1987 (49 sampling dates), core samples (17.7cm², 15cm deep) were collected at low tide at a sand-mud flat in the mid upper Gulf of Nicoya estuary, Pacific, Costa Rica. Predator exclusion experiments (cages 0.5x0.5x0.2m, galvanized wire, mesh size 5mm), were conducted at the site in 1985 (dry and rainy seasons sets). Samples were preserved with 5% buffered formalin in sea water stained with Rose Bengal, and washed after 24 hours on a 500 micron mesh sieve. The 1 120 cores yielded a total of 112 morphological species of which the mollusks were represented by 23 species, and included the bivalves Tellina rubescens, Tagelus bourgeoisae, Dosinia dunkeri and Leukoma asperrima, and the gastropods, Natica unifasciata, Nassarius luteostomus, Costoanachis rugosa and Turbonilla sp. The 23 species are indicative of a relatively rich sedimentary molluscan fauna. T. bourgeoisae had a seasonal oscillation, with higher abundancesduring the rainy seasons. T. rubescens was not seasonal, but presented an oscillation with peaks at about 1.5 year intervals. Many empty shells of Cosmioconcha modesta, lower number of N. luteostomus and a few of T. rubescens were found with boreholes by the predatory snail N. unifasciata. T. rubescens was not significantly more abundant inside or outside cages. T. bourgeoisae showed a significant increase within the caged areas. The numerical fluctuations of the mollusks became more important during the rainy season of 1985. Red tide outbreaks in the Gulf of Nicoya in 1985 may have had an impact on the molluscan populations. Rev. Biol. Trop. 59 (3): 1135-1148. Epub 2011 September 01.

La disponibilidad de datos con una cobertura mayor al año es rara para ambientes tropicales. Avances en computación y programas de cómputo facilitan el re-análisis de antiguos datos y la identificación de patrones ocultos en ellos. desde febrero de 1984 hasta abril de 1987 (49 fechas de colecta), muestras de barreno (17.7cm², 15cm de profundidad) fueron recolectadas durante la marea baja en una planicie arenosa-fangosa en la región media del estuario del Golfo de Nicoya, Costa Rica. Experimentos de exclusión de depredadores (jaulas 0.5x0.5x0.2m, malla galvanizada, poro de 5mm), fueron realizados en el sitio en 1985 (estación seca vs estación lluviosa). Las muestras fueron preservadas con formalina al 5% en agua de mar teñida con Rosa de Bengala y lavadas después de 24 horas en un tamiz de 500 micras de poro de malla. Los 1 120 barrenos produjeron un total de 112 especie morfológicas, de las cuales los moluscos estuvieron representados por 23 especies que incluyen los bivalvos: Tellina rubescens, Tagelus bourgeoisae, Dosinia dunkeri y Leukoma asperrima, y los gastrópodos, Natica unifasciata, Nassarius luteostomus, Costoanachis rugosa y Turbonilla sp. Las 23 especies son indicadoras de una fauna de moluscos relativamente rica. T. bourgeoisae presentó una oscilación estacional, con mayor abundancia durante la estación de lluvias. T. rubescens no fue estacional, pero presentó una oscilación con máximos a intervalos de cerca de 1.5 años. Muchas conchas vacías de Cosmioconcha modesta, un poco menos de N. luteostomus y unas pocas de T. rubescens fueron encontradas con perforaciones hechas por el depredador N. unifasciata. T. rubescens no fue significativamente más abundande dentro o fuera de las jaulas. T. bourgeoisae mostró una falta de abundancia estacional, pero una densidad significativamente mayor dentro de las jaulas. Las fluctuaciones poblacionales de los moluscos fueron más importantes durante la estación de lluvias cuando se les compara con las fluctuaciones de los poliquetos, crustáceos y otros grupos. Mareas rojas en el Golfo de Nicoya durante 1985 podrían haber tenido un impacto en la dinámica de las poblaciones de moluscos.

Hidrocarburos de petróleo, disueltos y dispersos, en cuatro ambientes costeros de Costa Rica / Oil pollution status expressed as the fraction of dissolved and dispersed petroleum hydrocarbons

Entre los años 2000 a 2002 se recolectaron y se analizaron 130 muestras de agua de mar en las costas de Costa Rica, para determinarles la concentración de hidrocarburos de petróleo disueltos y dispersos (HPDD), referida a equivalentes de criseno en hexano, y expresada como µg·L-1. Se utilizó la técnica analítica de fluorescencia molecular. 42 muestras correspondieron al Caribe (Moín) y 81 al Pacífico (Bahía Culebra, Golfo de Nicoya, Golfo Dulce y en una ocasión se tomaron muestras a lo largo del Estero de Puntarenas). En la zona costera de Moín el promedio de la concentración de HPDD fue 0.10 µg·L-1, con una desviación estándar de ± 0.18 µg·L-1, en un ámbito desde concentraciones no detectables por el método, (nd), hasta 0.65 µg·L-1. En el Pacífico las concentraciones para todo el período del estudio, estuvieron entre nd y 0.37 µg·L-1. En la Bahía Culebra no se detectaron HPDD en ninguna de las muestras (n = 12); en el Golfo de Nicoya el promedio y la desviación estándar fueron 0.04 µg·L-1 ± 0.09 µg·L-1 (n = 51), en un ámbito desde nd hasta 0.33 µg·L-1. En el Golfo Dulce el promedio y la desviación estándar fueron 0.05 µg·L-1 ± 0.11 µg·L-1 (n = 18), en el ámbito desde nd a 0.37 µg·L-1. Durante las fechas en que se llevó a cabo el muestreo, no se encontró una situación de contaminación por petróleo en los ecosistemas costeros estudiados, puesto que las concentraciones de HPDD no rebasaron el límite de 10 µg·L-1 referido a equivalentes de criseno, considerado típico de áreas oceánicas contaminadas. En Bahía Culebra fue la primera vez que se hizo el estudio de la contaminación por petróleo y se comprobó que en las fechas de muestreo la zona estuvo libre de esta contaminación. El promedio y la desviación estándar en el Estero de Puntarenas fueron 1.21 µg·L-1 ± 2.10 µg·L-1 (n = 7), en un ámbito de 0.17 a 5.91 µg·L-1. Estos resultados evidenciaron una situación de contaminación moderada y la naturaleza estuarina de este cuerpo de agua costera, que distribuye irregularmente los HPDD descargados en ella

Four coastal ecosystems with contrasting characteristics were sampled in Costa Rica (2000 – 2002). Oil pollution status, expressed as the fraction of dissolved/dispersed petroleum hydrocarbons related to chrysene equivalents, was determined by the molecular fluorescence analytical technique. A total of 130 water samples were taken, from the Caribbean (Moín Bay), and from the Pacific (Bahía Culebra, Gulf of Nicoya and Dulce Gulf). On one occasion, seven samples along the Puntarenas estuary were also analysed. In Moín the mean and standard deviation were 0.10 µg·L-1 ± 0.18 µg·L-1, ranging from non detectable (nd) to 0.65 µg·L-1. For the Pacific ecosystems the total range was from nd to 0.37 µg·L-1. In Bahía Culebra no fluorescence signals were obtained. In the Gulf of Nicoya the mean and standard deviation were 0.04 µg·L-1 ± 0.09 µg·L-1, from nd to 0.33 µg·L-1. Values in Dulce Gulf were 0.05 µg·L-1 ± 0.11 µg·L-1, from nd to 0.37 µg·L-1. Along the Puntarenas estuary the range was 0.17 to 5.91 µg·L-1, with a mean of 1.21 µg·L-1 and a standard deviation of ± 2.10 µg·L-1. The four coastal ecosystems had concentrations below the 10 µg·L-1 limit for polluted oceanic areas. The Puntarenas estuary reflects the influence of antropogenic activities from and around the City of Puntarenas. These levels are considered low for inshore waters

Metales traza en sedimentos costeros de Costa Rica / Trace metals in coastal sediments from Costa Rica

Se determinó la concentración de hierro, plomo, cobre y zinc, en sedimentos de cuatro ecosistemas costeros de Costa Rica, por espectroscopia de absorción atómica, con llama o con horno de grafito. En el Pacífico se escogieron Bahía Culebra (Golfo de Papagayo), el Golfo de Nicoya y la bahía de Golfito (Golfo Dulce) y en el Caribe la bahía de Moín. Las muestras fueron recolectadas entre los años 2000 y 2002, y los valores promedio de las concentraciones de los elementos, en todos los ecosistemas, siguieron una misma tendencia: Fe > Zn > Cu > Pb. No se encontró ningún tipo de patrón temporal de distribución de los metales en ninguno de los ecosistemas. Las concentraciones medias de hierro y de cobre fueron mayores en la bahía de Golfito (5.8% y 87 µg / g, respectivamente) y menores en la bahía de Moín (3.4% y 52 µg / g. respectivamente). La concentración media de zinc fue mayor también en la bahía de Golfito (96 µg / g), pero menor en Bahía Culebra (66 µg / g). La concentración de plomo fue mayor en la bahía de Moín (6.4 µg / g) y menor en Bahía Culebra (3.0 µg / g). La concentración de plomo fue alta en el Caribe y en la bahía de Golfito; los valores de concentración promedio obtenidos para el hierro, el plomo, el cobre y el zinc, fueron máximos en la bahía de Golfito, comparados con los otros ecosistemas. Con base en los datos obtenidos, se concluyó que la Bahía Culebra fue la zona más limpia; la bahía de Golfito la más contaminada y en un lugar intermedio la bahía de Moín y el Golfo de Nicoya

Marine sediment samples from four coastal ecosystems in Costa Rica were taken between the years 2000-2002 and their iron, lead, copper and zinc concentrations were determined by the atomic absorption technique with flame or graphite furnace. In the Pacific coast, Culebra Bay (Papagayo Gulf), Gulf of Nicoya, and Golfito Bay (Dulce Gulf), were selected as representative sites, and Moín Bay, at the Caribbean coast. Mean metal concentrations for all ecosystems followed the same pattern: Fe > Zn > Cu > Pb. No temporal pattern was found for any metal. Iron and copper mean concentrations were higher in Golfito Bay (5.8% and 87 µg / g, respectively) and lower in Moín Bay (3.4% and 52 µg / g, respectively). Zinc mean concentration was also higher in Golfito Bay (96 µg / g), but lower in Culebra Bay (66 µg / g). Lead mean concentration was higher in Moín Bay (6.4 µg / g) and lower in Culebra Bay (3.0 µg / g). Lead highest concentrations occurred in the Caribbean and in Golfito Bay, and for the rest of the elements the maximum values were found in Golfito Bay. On the basis of data obtained in this work, Culebra Bay was considered a relatively unpolluted location; Golfito Bay was more contaminated, and Moín Bay and the Gulf of Nicoya showed an intermediate condition

[Oil pollution status expressed as the fraction of dissolved and dispersed petroleum hydrocarbons]. / Hidrocarburos de petróleo, disueltos y dispersos, en cuatro ambientes costeros de Costa Rica.

Four coastal ecosystems with contrasting characteristics were sampled in Costa Rica (2000-2002). Oil pollution status, expressed as the fraction of dissolved/dispersed petroleum hydrocarbons related to chrysene equivalents, was determined by the molecular fluorescence analytical technique. A total of 130 water samples were taken, from the Caribbean (Moín Bay), and from the Pacific (Bahía Culebra, Gulf of Nicoya and Dulce Gulf). On one occasion, seven samples along the Puntarenas estuary were also analysed. In Moín the mean and standard deviation were 0.10 microg x L(-1) +/- 0.18 micro x L(-1), ranging from non detectable (nd) to 0.65 microg x L(-1). For the Pacific ecosystems the total range was from nd to 0.37 microg x L(-1). In Bahia Culebra no fluorescence signals were obtained. In the Gulf of Nicoya the mean and standard deviation were 0.04 microg x L(-1) +/- 0.09 microg x L(-1), from nd to 0.33 microg x L(-1). Values in Dulce Gulf were 0.05 microg x L(-1) +/- 0.11 microg x L(-1), from nd to 0.37 microg x L(-1). Along the Puntarenas estuary the range was 0.17 to 5.91 microg x L(-1), with a mean of 1.21 microg x L(-1) and a standard deviation of +/- 2.10 microg x L(-1). The four coastal ecosystems had concentrations below the 10 microg x L(-1) limit for polluted oceanic areas. The Puntarenas estuary reflects the influence of antropogenic activities from and around the City of Puntarenas. These levels are considered low for inshore waters.

[Trace metals in coastal sediments from Costa Rica]. / Metales traza en sedimentos costeros de Costa Rica.

Marine sediment samples from four coastal ecosystems in Costa Rica were taken between the years 2000-2002 and their iron, lead, copper and zinc concentrations were determined by the atomic absorption technique with flame or graphite furnace. In the Pacific coast, Culebra Bay (Papagayo Gulf), Gulf of Nicoya, and Golfito Bay (Dulce Gulf), were selected as representative sites, and Moín Bay, at the Caribbean coast. Mean metal concentrations for all ecosystems followed the same pattern: Fe > Zn > Cu > Pb. No temporal pattern was found for any metal. Iron and copper mean concentrations were higher in Golfito Bay (5.8% and 87 microg/g, respectively) and lower in Moín Bay (3.4% and 52 microg/g, respectively). Zinc mean concentration was also higher in Golfito Bay (96 microg/g), but lower in Culebra Bay (66 microg/g). Lead mean concentration was higher in Moín Bay (6.4 microg/g) and lower in Culebra Bay (3.0 microg/g). Lead highest concentrations occurred in the Caribbean and in Golfito Bay, and for the rest of the elements the maximum values were found in Golfito Bay. On the basis of data obtained in this work, Culebra Bay was considered a relatively unpolluted location; Golfito Bay was more contaminated, and Moín Bay and the Gulf of Nicoya showed an intermediate condition.

Abundance of Petrolisthes armatus (Crustacea: Porcellanidae) on a tropical estuarine intertidal rocky beach, Gulf of Nicoya estuary, Costa Rica

Population of the porcellanid crab Petrolisthes armatus was studied on a rocky intertidal beach located at the Punta Morales peninsula in the mid upper Gulf of Nicoya estuary, Pacific coast of Costa Rica, from December 1997 to November 1998 (14 dates). Horizontal plankton tows (280 micron mesh net) were also made to verify the presence of P. armatus larvae. Crabs were collected every 3 m along three 18 m long transects, at two sites on the beach, by placing a bottomless bucket fringed with canvas to prevent the organisms from escaping under the rim. A total of 15,382 P. armatus were collected. Only 146 (0.95%) crabs had a carapace length longer than 10 mm, and 8995 (58.5%) were in the size range of 2 to 4.5 mm. The remaining crabs 6241 (40.5%) were in the size range of 4.6 to 10 mm. Male and female P. armatus were represented by 2777 and 3518 individuals respectively, with a sex ratio of 1:1.26. Ovigerous females were found at all dates and included 2937 individuals (83% of females). Plankton tows yielded only 73 larvae of P. armatus (Zoea 1), with a density of 1.2 larvae/m3. No statistically significant seasonal trends in the population of this species were detected. P. armatus appears to reproduce continuously the year around in Punta Morales, and some peaks of abundance were present during the dry and rainy seasons. These trends are similar to trends reported for other crustacean species in the Gulf of Nicoya.