Nursery-reared coral outplanting success in an upwelling-influenced area in Costa Rica

Introduction: Environmental and intrinsic factors such as seawater temperature, salinity, nutrient concentration, upwelling, species, and life history can influence coral outplant survival and growth, and in consequence, the effectiveness of restoration. Thus, it is key to understand how these factors can shape coral outplant performance to ensure the long-term success of a restoration program. Objective: To establish the survival and growth rate of outplanted coral nursery-reared colonies of branching Pocillopora spp. and massive corals Pavona gigantea, Pavona clavus, and Porites lobata in Bahía Culebra, North Pacific of Costa Rica, and to determine whether the site of origin of the coral fragment and the presence of seasonal upwelling affected the growth of Pocillopora outplants. Methods: From September 2020 to September 2021, we monitored the survival, health, and growth of 30 Pocillopora spp. colonies from six donor sites, and 31 fragments of massive species (P. gigantea [n = 18], P. clavus [n = 8], P. lobata [n = 5]) that were outplanted to a degraded reef. We recorded in situ seawater temperature, salinity, and nutrient concentration. Results: By the end of the year, 100 % of the Pocillopora spp. outplants survived. Survival was 71.4 % for P. clavus, 47.5 % for P. gigantea, and 20 % for P. lobata. Coral tissue loss and predation marks were observed at least once in 71 % of massive outplants. Pocillopora spp. colonies increased their initial area in 333.1 %, at a rate of 9.98 ± 1.69 cm yr-1. The site of origin of the outplant influenced growth rate, but not the presence of seasonal upwelling. Massive species fragments grew at a rate of 1.35 ± 0.24 cm yr-1 for P. clavus, 1.48 ± 0.21 cm yr-1 for P. gigantea, and 0.61 cm yr-1 for P. lobata, with no differences among them. Conclusions: Environmental conditions at site level allow for high survival and growth of Pocillopora spp. outplants, previously considered as sensitive to stressors, and point towards acclimation to seasonal upwelling. Although survival of massive species was lower, developing a multi-species approach is key to enhance restoration success.

Introducción: Factores ambientales e intrínsecos como la temperatura del agua, salinidad, concentración de nutrientes, afloramiento, especies e historia de vida pueden influir en la supervivencia y crecimiento de los trasplantes de coral y, en consecuencia, en la eficacia de la restauración. Por ello, entender cómo estos factores pueden moldear el desempeño de los trasplantes es clave para asegurar el éxito de un programa de restauración a largo plazo. Objetivo: Establecer la supervivencia y tasa de crecimiento de los trasplantes de colonias de coral previamente cultivadas en viveros del coral ramificado Pocillopora spp. y las especies de crecimiento masivo Pavona gigantea, Pavona clavus y Porites lobata en Bahía Culebra, Pacífico Norte de Costa Rica, y determinar si el sitio donante de origen del fragmento de coral y la presencia de afloramiento estacional afectó el crecimiento de los trasplantes de Pocillopora. Métodos: De setiembre 2020 a setiembre 2021, se monitoreó la supervivencia, salud y crecimiento de 30 colonias de Pocillopora spp. de seis sitios donantes diferentes y 31 fragmentos de especies masivas (P. gigantea [n = 18], P. clavus [n = 8], P. lobata [n = 5]) que se trasplantaron a un arrecife degradado. Se registró la temperatura del agua, la salinidad y concentración de nutrientes in situ. Resultados: Al final del año, el 100 % de los trasplantes de Pocillopora spp. sobrevivieron. La supervivencia fue de 71.4 % para P. clavus, 47.5 % para P. gigantea y 20 % para P. lobata. Se observaron pérdidas de tejido y marcas de depredación al menos una vez en un 71 % de los trasplantes masivos. Las colonias de Pocillopora spp. aumentaron su área inicial en un 333.1 %, a una tasa de 9.98 ± 1.69 cm año-1. El sitio de origen del trasplante tuvo efecto sobre la tasa de crecimiento, pero no la presencia de afloramiento estacional. Los fragmentos de especies masivas crecieron a tasas de 1.35 ± 0.24 cm año-1 (P. clavus), 1.48 ± cm año-1 (P. gigantea) y 0.61 cm año (P. lobata), sin diferencias entre ellas. Conclusiones: Las condiciones del sitio permiten una alta supervivencia y crecimiento de trasplantes de Pocillopora spp., que previamente habían sido considerados sensibles a factores estresantes, e indican aclimatación a las condiciones locales de afloramiento estacional. Pese a que la supervivencia de las especies masivas fue menor, es esencial desarrollar un enfoque multi-especie para aumentar el éxito de la restauración.

Pocillopora spp. growth analysis on restoration structures in an Eastern Tropical Pacific upwelling area.

Coral reefs in Culebra Bay (North Pacific of Costa Rica) are threatened by multiple anthropogenic disturbances including global warming, overfishing, eutrophication, and invasive species outbreaks. It is possible to assist their recovery by implementing ecological restoration techniques. This study used artificial hexagonal steel structures, called "spiders" to compare growth of Pocillopora spp. coral fragments of different sizes. Three initial fragment class sizes were used: 2, 5 and 8 cm, with each class size having 42 initial fragments. Changes in fragment length, width and area were measured monthly from January to December 2020. Results showed an overall survivorship of 70.21%, and no significant differences in survivorship and linear growth rate were detected between class sizes. The linear growth rates are 4.49 ± 1.19 cm year-1, 5.35 ± 1.48 cm year-1 and 3.25 ± 2.22 cm year-1 for the 2, 5 and 8 cm initial class sizes, respectively. Our results do not show significant differences in growth rates between the different initial fragment sizes. However, since small fragments (2 cm) present higher mortality during the first month, we recommend using larger fragments. In addition, coral fragments grew 48% more during the non-upwelling season, which may suggest that it might be more effective and safer to start the restoration efforts during this period.

Divergence in Heliconius flight behaviour is associated with local adaptation to different forest structures.

Microhabitat choice plays a major role in shaping local patterns of biodiversity. In butterflies, stratification in flight height has an important role in maintaining community diversity. Despite its presumed importance, the role of behavioural shifts in early stages of speciation in response to differences in habitat structure is yet to be established. Here, we investigated variation in flight height behaviour in two closely related Heliconius species, H. erato cyrbia and H. himera, which produce viable hybrids but are isolated across an environmental gradient, spanning lowland wet forest to high-altitude scrub forest. Speciation in this pair is associated with strong assortative mating, but ecological isolation and local adaptation are also considered essential for complete reproductive isolation. We quantified differences in flight height and forest structure across the environmental gradient and tested the importance of resource distribution in explaining flight behaviour. We then used common garden experiments to test whether differences in flight height reflect individual responses to resource distribution or genetically determined shifts in foraging behaviour. We found that the two species fly at different heights in the wild, and demonstrated that this can be explained by differences in the vertical distribution of plant resources. In both the wild and captivity, H. himera choose to fly lower and feed at lower positions, closely mirroring differences in resource availability in the wild. Given expectations that foraging efficiency contributes to survival and reproductive success, we suggest that foraging behaviour may reflect local adaptation to divergent forest structures. Our results highlight the potential role of habitat-dependent divergence in behaviour during the early stages of speciation.

La elección de microhábitat juega un papel determinante en los patrones locales de biodiversidad. En las mariposas, la estratificación (entendida como altura de vuelo) cumple un rol importante al promover la diversidad de las comunidades. A pesar de esta asumida importancia, todavía está por establecerse el rol de los cambios comportamentales en estadios tempranos de especiación en respuesta a diferencias en la estructura del hábitat. Aquí investigamos la variación en el comportamiento de estratificación en dos especies de mariposas Heliconius cercanamente emparentadas: H. erato cyribia y H. himera, las cuales producen híbridos viables, pero están aisladas a lo largo de un gradiente ambiental que va desde el bosque húmedo tropical de tierras bajas hasta el bosque de matorrales de altitud. La especiación en esta pareja de mariposas está asociada con un alto grado de emparejamiento selectivo, pero el aislamiento ecológico y la adaptación local también son considerados esenciales para un aislamiento reproductivo completo. Mostramos que las dos especies vuelan a alturas distintas, y demostramos que esto se explica por diferencias en la distribución vertical de los recursos florísticos. Posteriormente, usando experimentos controlados de jardín, exploramos si esta divergencia en altura de vuelo está determinada ambientalmente. Tanto en la naturaleza como en cautividad, H. himera escoge volar más bajo y alimentarse en lugares más bajos, lo que es consistente con las diferencias en la disponibilidad de recursos en la naturaleza. Dada la expectativa de que la eficiencia en el forrajeo contribuye al éxito reproductivo y de supervivencia, sugerimos que el comportamiento de forrajeo podría reflejar adaptación local a estructuras de bosque divergentes. Nuestros resultados resaltan el potencial rol de la divergencia dependiente del hábitat en el comportamiento en estadios tempranos de especiación.

Diversidad de los hábitats submareales de la Península de Santa Elena e Islas Murciélago, Pacífico Norte, Costa Rica

Introducción: Los hábitats costeros de todo el mundo están experimentando una presión cada vez mayor debido a la contaminación, el desarrollo costero, la pesca y el cambio climático. Identificar y registrar la biodiversidad costera es esencial para evaluar la salud, los cambios y el alcance de la pérdida de biodiversidad de los ecosistemas. Los hábitats costeros tropicales como los arrecifes de coral y los lechos de pastos marinos han sido el foco de investigación de los científicos durante las últimas décadas; sin embargo, se han descuidado otros ecosistemas, como los bosques de macroalgas, los fondos de sedimentos submareales (20-30 m de profundidad) y los arrecifes rocosos. Objetivo: Este trabajo reporta la biodiversidad marina asociada con arrecifes rocosos, arrecifes de coral, bosques de Sargazo y fondos submareales sedimentarios (20-30 m de profundidad), en un área tropical de afloramiento estacional (Península de Santa Elena, Bahía de Santa Elena e Islas Murciélago, Costa Rica). Métodos: Durante la Expedición Santa Elena (21 de abril-2 de mayo de 2018), se visitaron un total de 28 sitios para registrar la biodiversidad en cuatro hábitats diferentes (arrecifes rocosos y de coral, bosques de sargazo y fondos de sedimentos submareales), utilizando SCUBA, en muestreos sistemáticos, evaluaciones visuales y en algunos fondos de sedimentos se utilizó una draga de arrastre por 20 minutos. Resultados: Se identificaron un total de 254 taxones, siendo los peces óseos el grupo más diverso con 91 especies, seguido de los gasterópodos (25 spp.), las algas rojas (21 spp.) y los antozoos (19 spp.). Se reportan cuatro nuevos registros para el Pacífico continental de Costa Rica, incluida la subclase de una anémona tubícola (Ceriantharia). Conclusiones: Los resultados muestran que la península de Santa Elena, la bahía de Santa Elena y las islas Murciélago albergan hábitats poco comunes y no estudiados, como bosques del alga parda Sargassum, fondos de sedimentos, arrecifes construidos por el coral masivo Pavona gigantea, que es muy inusual, y comunidades de arrecifes rocosos dominadas por colonias calcáreas de poliquetos (Salmacina tribranchiata).

Introduction: Worldwide, coastal habitats are experiencing increasing pressure from pollution, coastal development, fisheries, and climate change. Identifying and recording coastal biodiversity is essential to assess ecosystem health, changes and the extent of biodiversity loss. Coastal tropical habitats such as coral reefs and seagrass beds have been the research focus for scientists during the last decades; however, other ecosystems have been neglected, such as macroalgae forests, subtidal (20-30 m deep) sedimentary bottoms, and rocky reefs. Objective: Our study reports the marine biodiversity associated with rocky reefs, coral reefs, Sargassum forests and sedimentary subtidal bottoms (20-30 m deep), in a tropical seasonal upwelling area (Santa Elena Peninsula, Santa Elena Bay, and Murciélago Islands, Costa Rica). Methods: During the 'Santa Elena Expedition' (April 21-May 2 2018), a total of 28 sites were visited in order to record the biodiversity across four different habitats (rocky and coral reefs, Sargassum forests, subtidal sedimentary bottoms), using SCUBA and both systematic surveys and visual assessments; in some sedimentary bottoms a 20-minute dredge tows were done. Results: A total of 254 taxa were identified, being bony fishes the most diverse group (91 species), followed by gastropods (25 spp.), red algae (21 spp.) and anthozoans (19 spp.). We report four new records for the Pacific mainland of Costa Rica, including the subclass of tube-dwelling sea anemone (Ceriantharia). Conclusions: Our results show that Santa Elena Peninsula, Santa Elena Bay, and Murciélago Islands harbor uncommon and unstudied habitats, such as Sargassum forests, sedimentary bottoms, reefs constructed by the stony coral Pavona gigantea -which is very unusual- and rocky reef communities dominated by calcareous colonies of polychaetes (Salmacina tribranchiata).

Forest stratification shapes allometry and flight morphology of tropical butterflies.

Studies of altitudinal and latitudinal gradients have identified links between the evolution of insect flight morphology, landscape structure and microclimate. Although lowland tropical rainforests offer steeper shifts in conditions between the canopy and the understorey, this vertical gradient has received far less attention. Butterflies, because of their great phenotypic plasticity, are excellent models to study selection pressures that mould flight morphology. We examined data collected over 5 years on 64 Nymphalidae butterflies in the Ecuadorian Chocó rainforest. We used phylogenetic methods to control for similarity resulting from common ancestry, and explore the relationships between species stratification and flight morphology. We hypothesized that species should show morphological adaptations related to differing micro-environments, associated with canopy and understorey. We found that butterfly species living in each stratum presented significantly different allometric slopes. Furthermore, a preference for the canopy was significantly associated with low wing area to thoracic volume ratios and high wing aspect ratios, but not with the relative distance to the wing centroid, consistent with extended use of fast flapping flight for canopy butterflies and slow gliding for the understorey. Our results suggest that microclimate differences in vertical gradients are a key factor in generating morphological diversity in flying insects.

Ecosistemas coralinos del Área de Conservación Osa, Costa Rica: estructura y necesidades de conservación / Osa Conservation Area (Costa Rica) coral ecosystems: structure and conservation needs

Resumen Costa Rica es considerado uno de los 20 países del mundo con mayor biodiversidad, entre los que destacan sus ecosistemas arrecifales. Más precisamente, el Pacífico Sur, el Área de Conservación Osa, se ha considerado como una de las zonas más diversas en estos ecosistemas. Estos arrecifes han sido estudiados desde inicios de los años 80's, pero enfocándose solamente en la cobertura coralinos de estos arrecifes. Por lo tanto, la presente investigación busca realizar una evaluación actualizada y con una metodología estándar él estado de varios componentes de los sistemas arrecifales de esta región. Entre el 2013 y el 2014, se visitaron cinco localidades: Dominicalito, Parque Nacional Marino Ballena, Reserva Biológica Isla del Caño, Península de Osa y Golfo Dulce, donde se evaluaron 27 sitios. En ellos se realizó la evaluación, a dos profundidades, de: cobertura del fondo, rugosidad del arrecife, diversidad y densidad de macroinvertebrados, así como la diversidad y biomasa de peces arrecifales. En términos generales los arrecifes de ACOSA, son moderadamente complejos y en ellos predominan el tapete algal (62.7%), mientras que la cobertura de coral vivo es moderada (16.5%). Se observó una diferencia entre la cobertura de tipos de sustratos por localidad, la cual puede estar atribuida a condiciones ambientales adversas (i.e. sedimentación, contaminación) para su desarrollo. Sin embargo, los arrecifes coralinos de Golfo Dulce e Isla del Caño muestran una notable recuperación al compararla con estudios previos. Se observaron 35 taxa de macroinvertebrados en los sitios estudiados, en su mayoría (58%) fueron poco abundantes u ocasionales. La mayor diversidad y densidad de macroinvertebrados se observó en Golfo Dulce y se determinaron diferencias significativas entre localidades. Este estudio brinda una línea base para muchos de estos organismos en varios de los sitios. Los equinoideos Diadema mexicanum y Eucidaris thouarsii fueron los organismos predominantes, y se reporta el cambute (Lobatus galeatus) en varios de los sitios de muestreo. De las 90 especies de peces, 10 fueron las de mayor ocurrencia y densidad. No hubo diferencias significativas en la diversidad y riqueza de especies de peces observadas entre localidades, pero si en la biomasa y en la composición de la comunidad íctica entre los sitios. Los sitios de la Isla del Caño fueron los más diversos de toda ACOSA, presentan biomasas importantes y de niveles tróficos altos. Mientras que Golfo Dulce difirió con respecto a las demás localidades. Según los resultados observados, los arrecifes de ACOSA presentan una alta biodiversidad. Las condiciones del de los ecosistemas indican que más esfuerzos de conservación, uso sostenible y manejo efectivo de loa recursos en tierra como en el mar deben ser implementados. El monitoreo ecológico va a proveer de la información sobre las tendencias y las relaciones entre el estado de los ecosistemas arrecifales y las condiciones ambientales.

Abstract Costa Rica is considered one of the 20 most biodiverse countries in the world, which includes it´s well known reefs ecosystems. Specifically, the South Pacific region, the Osa Conservation Area (ACOSA), holds one of the richest reefs of the country. Even though many of these reefs have been studied since the 1980´s, most of them had focused mainly on coral cover condition. Therefore, the present research aims to complete and update an evaluation of the main biological components of ACOSA´s reefs ecosystems, using a standard methodology. Between 2013 and 2014, five localities were visited: Dominicalito, Marino Ballena National Park, Caño Island Biological Reserve, Osa Peninsula and Golfo Dulce, on which 27 sites were evaluated. Transects at two depths in each site were done to estimate substrate cover, reef rugosity, macroinvertebrates diversity and density, and reef fishes composition and biomass. Results indicate that ACOSA´s reef are moderately complex and on most of them the substrate is dominated by turf (62.7%) and live coral cover is moderate (16.5%). Substrate cover was significantly different between localities, which can be attributed to adverse environmental conditions for reef development (e.g. sedimentation, pollution). However, reefs in Golfo Dulce and Caño Island BR showed a significant coral cover recovery since last studied. Of 35 taxa of macroinvertebrates identified, 58% had low abundance or were observed occasionally. Golfo Dulce presented the highest diversity and density of macroinvertebrates and significant differences were observed between localities. On this group, this study is providing an information baseline of most sites analyzed. The echinoids Diadema mexicanum and Eucidaris thouarsii were the most abundant and the conch (Lobatus galeatus) was reported for many of the sites. Of the 90 species of reef fishes documented, ten were common and with the highest densities. There were no significant differences in the diversity and richness of the reef fishes between localities, but fishes' biomass and composition of fish community were different between sites. Caño Island´s sites were the most diverse of ACOSA, and presented the highest biomass and trophic levels of the region. Golfo Dulce´s reef fish composition differs from the other localities. According to the results, ACOSA´s reef ecosystems have a high biodiversity. The ecosystems conditions observed indicate that more conservation efforts, sustainable use and effective management of land and marine resources should be implemented. Ecological monitoring will provide the information about trends and the relationship between reef ecosystems state and environmental conditions. Rev. Biol. Trop. 63 (Suppl. 1): 219-259. Epub 2015 April 01.

Reproductive traits of the symbiotic pea crab Austinotheresangelicus (Crustacea, Pinnotheridae) living in Saccostreapalmula (Bivalvia, Ostreidae), Pacific coast of Costa Rica.

Pea crabs of the family Pinnotheridae exhibit a symbiotic life style and live associated with a variety of different marine organisms, especially bivalves. Despite the fact that pea crabs can cause serious problems in bivalve aquaculture, the available information about the ecology of these crabs from Central America is extremely limited. Therefore, the present study aimed to describe different reproductive features of the pinnotherid crab Austinotheresangelicus associated with the oyster Saccostreapalmula in the Golfo de Nicoya, Pacific coast of Costa Rica. Monthly sampling was conducted from April to December 2012. Average carapace width (CW) of the 47 analyzed ovigerous females was 7.62 mm. The species produced on average 2677 ± 1754 recently -extruded embryos with an average volume of 0.020 ± 0.003 mm(3); embryo volume increased during embryogenesis by 21%, but did not vary significantly between developmental stages. Brood mass volume varied greatly (between 11.7 and 236.7 mm(3)), and increased significantly with female CW. Females invested on average 76.7% (minimum: 21.7%; maximum: 162.8%) of their body weight in brood production, which confirms a substantially higher energy allocation for embryo production in pinnotherid crabs compared to free-living decapods.