Limitations to coral recovery along an environmental stress gradient.

Positive feedbacks driving habitat-forming species recovery and population growth are often lost as ecosystems degrade. For such systems, identifying mechanisms that limit the re-establishment of critical positive feedbacks is key to facilitating recovery. Theory predicts the primary drivers limiting system recovery shift from biological to physical as abiotic stress increases, but recent work has demonstrated that this seldom happens. We combined field and laboratory experiments to identify variation in limitations to coral recovery along an environmental stress gradient at Ningaloo Reef and Exmouth Gulf in northwest Australia. Many reefs in the region are coral depauperate due to recent cyclones and thermal stress. In general, recovery trajectories are prolonged due to limited coral recruitment. Consistent with theory, clearer water reefs under low thermal stress appear limited by biological interactions: competition with turf algae caused high mortality of newly settled corals and upright macroalgal stands drove mortality in transplanted juvenile corals. Laboratory experiments showed a positive relationship between crustose coralline algae cover and coral settlement, but only in the absence of sedimentation. Contrary to expectation, coral recovery does not appear limited by the survival or growth of recruits on turbid reefs under higher thermal stress, but to exceptionally low larval supply. Laboratory experiments showed that larval survival and settlement are unaffected by seawater quality across the study region. Rather, connectivity models predicted that many of the more turbid reefs in the Gulf are predominantly self seeded, receiving limited supply under degraded reef states. Overall, we find that the influence of oceanography can overwhelm the influences of physical and biological interactions on recovery potential at locations where environmental stressors are high, whereas populations in relatively benign physical conditions are predominantly structured by local ecological drivers. Such context-dependent information can help guide expectations and assist managers in optimizing strategies for spatial conservation planning for system recovery.

Early life history of the Caribbean coral Orbicella faveolata (Scleractinia: Merulinidae)

Introduction: Rehabilitation of hermatypic coral species that have declined in the Caribbean in recent decades is a priority. Production of sexual recruits is considered the best restoration method to aid affected populations. Objective: To gain knowledge of early life stages of Orbicella faveolata and to enhance production of new sexual recruits. Methods: Gamete bundles from the coral species O. faveolata were collected over two years (2018 and 2019) from Los Corales del Rosario y de San Bernardo Natural National Park, Cartagena, Colombia. Assisted fertilization, larval rearing, settlement (onto crustose coralline algae, CCA) and post settlement survival in laboratory conditions were monitored. Results: Embryonic and larval development were documented over 55 hours after the first cleavage, when larvae were fully developed and started pre-settlement behavior. Settlement began 7 days after first cleavage and after 37 days polyps had acquired zooxanthellae. Larval settlement was higher on Lythophyllum congestum and Titanoderma prototypum than in response to Porolithon pachydermum, Neogoniolithon sp., Hydrolithon sp., and Lythophyllum sp. Larvae did not settle on dead coral or on the negative control (sterilized seawater). After the first week post settlement survival was 59 % amongst O. faveolata recruits. During the second week, survival dropped to 42 %, and was further reduced to 0 % at the end of the third week. Conclusions: O. faveolata larvae require cues from certain CCA species to settle, they do not settle in absence of CCA. Increased larvae availability is possible through assisted fertilization in the laboratory, however, due to the high mortality in early post-settlement phases, additional research needs to be conducted in order to scale up larvae production and improve understanding of the cues that enhance settlement and the factors which cause post-settlement mortality.

Introducción: La rehabilitación de las especies de corales hermatípicos del Caribe que han disminuido en las últimas décadas es una prioridad. La producción de reclutas sexuales se considera el mejor método de restauración para ayudar a las poblaciones afectadas. Objetivo: Obtener conocimiento de las primeras etapas de la vida de O. faveolata y mejorar la producción de nuevos reclutas sexuales. Métodos: Por dos años (2018 y 2019), seis días después de luna llena en septiembre, se recolectaron paquetes gaméticos en arrecifes del Parque Nacional Natural Los Corales del Rosario y de San Bernardo, Cartagena, Colombia. Se siguió la fertilización asistida, la cría de larvas, el asentamiento y la supervivencia posterior al asentamiento en algas coralinas costrosas (ACC) en condiciones de laboratorio. Resultados: El desarrollo de embriones y larvas se documenta a lo largo de 55 h después del primer clivaje, cuando la larva está desarrollada completamente y comenzó el comportamiento previo al asentamiento. El asentamiento comienza 7 días después del primer clivaje y 37 días después, la mayoría de los pólipos presentan zooxantelas. El asentamiento larval fue más alto en Lythophyllum congestum y Titanoderma prototypum que en respuesta a Porolithon pachydermum, Neogoniolithon sp., Hydrolithon sp., y Lythophyllum sp. No hubo asentamiento sobre coral muerto ni en el control negativo (agua de mar esterilizada). La supervivencia bajó de un 59 % en la primera semana después del asentamiento, a 42 % durante la segunda semana y 0 % para el final de la tercera semana. Conclusiones: Las larvas de O. faveolata requieren señales de ciertas especies de ACC para asentarse, ellas no se asientan en ausencia de ACC. La disponibilidad de larvas es posible mediante la fertilización asistida en laboratorio. Sin embargo, debido a la alta mortalidad en las primeras fases posteriores al asentamiento, queda mucho por hacer para aumentar la producción de larvas y mejorar nuestro conocimiento y comprensión de las señales que mejoran el asentamiento y las que previenen o inhiben la supervivencia del recluta.

Coralline algal metabolites induce settlement and mediate the inductive effect of epiphytic microbes on coral larvae.

Settlement of invertebrates is a key process affecting the structure of marine communities and underpins the ability of benthic ecosystems to recover from disturbance. While it is known that specific crustose coralline algae (CCA) are important for settlement of some coral species, the role of algal chemical compounds versus surface microbial biofilms has long been ambiguous. Using a model system - a CCA of a genus that has been shown to induce high levels of settlement of Acropora corals (Titanoderma cf. tessellatum) and an abundant coral species (Acropora millepora)- we show that chemical effects of CCA are stronger than those from CCA surface microbial biofilms as drivers of coral settlement. Biofilms contributed to some extent to larval settlement via synergistic effects, where microbial cues were dependent on the CCA primary metabolism (production of dissolved organic carbon). We propose that optimal coral settlement is caused by complex biochemical communications among CCA, their epiphytic microbial community and coral larvae.

Variaciones adaptativas en la talla de la megafauna bentónica de fondos blandos tropicales en función de parámetros bióticos y abióticos / Adaptive variation in size of tropical soft bottom benthic megafauna related to biotic and abiotic factors

Resumen: Entender y predecir variaciones en el tamaño de los invertebrados megabentónicos sigue siendo un desafío importante en macroecología marina. Este estudio se realizó para identificar tendencias de cambio en la talla de la megafauna bentónica de fondos sedimentarios tropicales y conocer la influencia de variables que pueden determinar el tamaño de estos organismos, evaluando hipótesis y paradigmas ecológicos de mar profundo originados en zonas subtropicales y templadas. El área de estudio abarcó toda la plataforma continental del Caribe colombiano. Los ejemplares se recolectaron en 1998, 2001 y 2005 con red demersal semi-globo, entre 10 y 500 m de profundidad. Se seleccionaron las especies mejor representadas: Eudolium crosseanum, Cosmioconcha nitens, Nuculana acuta (moluscos), Astropecten alligator, Brissopsis atlantica, B. elongata (equinodermos), Anasimus latus, Chasmocarcinus cylindricus y Achelous spinicarpus (crustáceos). Para detectar tendencias significativas de cambio de tamaño, e inferir la influencia de variables bióticas y ambientales, se emplearon modelos aditivos generalizados, donde la talla fue la variable respuesta y las variables predictivas fueron: profundidad, temperatura, densidad intraespecífica e interespecífica, riqueza, latitud y longitud. Se midieron 7 000 ejemplares en total. Seis especies presentaron un incremento de la talla con el aumento de la profundidad y la disminución de la temperatura. Dichas especies abarcaron ámbitos batimétricos que superaron los 10 °C de diferencia entre zonas someras y profundas. Geográficamente se presentó una marcada disminución de tallas en el área de influencia de la desembocadura del río Magdalena, debida posiblemente a los fuertes cambios físicosquímicos que este río genera, principalmente porque es la zona con menos productividad primaria planctónica de toda el área de estudio. Hacia el norte de la desembocadura del río Magdalena se presentó un notorio aumento de tamaños entre los (74° - 71°W y 11° - 13°N), que puede deberse a la surgencia que se presenta en el norte del Caribe colombiano. La relación entre la densidad de individuos y la talla no fue clara. Sin embargo cinco especies mostraron una relación inversa con la densidad intraespecífica, y tres con la interespecífica. Temperatura y profundidad fueron las variables que mejor explicaron los cambios de talla detectados. La mayoría de especies presentaron un aumento de dimensiones corporales con la disminución de la temperatura en el perfil batimétrico. La tendencia al aumento de tallas con la profundidad es contraria a lo indicado por los modelos teóricos de talla óptima (pero acorde con algunos estudios más recientes), que predicen un decrecimiento del tamaño de los organismos con el incremento de la profundidad, debido a la disminución de nutrientes hacia zonas profundas del océano. Es posible que el aumento de tamaño sea una estrategia adaptativa para la maximización de la energía, como ocurre con muchos organismos en las porciones más frías de su ámbito de distribución. Futuros estudios en el Caribe deberían examinar variaciones en la talla de la megafauna bentónica en zonas más profundas (más de 500 m), donde la temperatura varía menos, razón por la cual otros factores pueden ser más importantes para determinar el tamaño de estos organismos.

Abstract: Understanding and predicting adaptations in body size of megabenthic invertebrates remains a major challenge in marine macroecology. This study was conducted in order to investigate size variations of benthic megafauna in the tropics and to identify the effect of biotic and abiotic factors that may produce changes to these organisms, testing unresolved hypothesis and paradigms of deep sea ecology from subtropical and temperate areas. The study area covered the continental shelf of the Colombian Caribbean. The samples were collected during 1998, 2001 and 2005, using semi-globe demersal net for a water depth of 10 to 500 m. The most common species were selected for further study: Eudolium crosseanum, Cosmioconcha nitens, Nuculana acuta (mollusks), Astropecten alligator, Brissopsis atlantica, B. elongata (equinoderms), Anasimus latus, Chasmocarcinus cylindricus and Achelous spinicarpus (crustaceans). Generalized Additive Models were used to detect significant changes in size and to infer the effects of biotic and environmental factors on organisms' size. The dependent variable was size and the predicting model variables were depth, temperature, intraspecific density, interspecific density, richness, latitude, and longitude. A total of 7 000 individuals were measured. Six species showed an increase in body size towards deeper and colder sites. These species inhabit shallow and deep environments that exceed a variation in temperature of 10 °C. There was a remarkable size reduction in areas affected by the Magdalena River, possibly due to major physicochemical changes caused by the river. This region has the lowest planktonic primary productivity within the study area. An increase in sizes was observed north of the Magdalena River (74° - 71°W & 11° - 13°N), which may be attributable to the coastal upwelling occurring in this part of Colombia. The relationship between the density of benthic organisms and size was not clear. However, five species showed an inverse relation with intraspecific density and three with interspecific density. Temperature and depth were the variables that best explained the variations in size. Most of the studied species showed an increase in body size when temperature dropped along the bathymetric range. The trend of increasing size in deeper zones is contrary to the prediction of the optimal size theoretical model (but consistent with recent studies), which indicates a reduction in organisms' size in the deep sea, due to food limitation with increasing depth. It is possible that this increase in size is an adaptation to maximize energy, which is frequently observed in the coldest habitats of several species. Future studies in Caribbean should examine variations in size of benthic megafauna towards deeper zones (more than 500 m), were temperature is less variable and then other factors can play a more important role determining the size of these organisms. Rev. Biol. Trop. 65 (3): 1002-1021. Epub 2017 September 01.

Density-dependent coral recruitment displays divergent responses during distinct early life-history stages.

Population growth involves demographic bottlenecks that regulate recruitment success during various early life-history stages. The success of each early life-history stage can vary in response to population density, interacting with intrinsic (e.g. behavioural) and environmental (e.g. competition, predation) factors. Here, we used the common reef-building coral Acropora millepora to investigate how density-dependence influences larval survival and settlement in laboratory experiments that isolated intrinsic effects, and post-settlement survival in a field experiment that examined interactions with environmental factors. Larval survival was exceptionally high (greater than 80%) and density-independent from 2.5 to 12 days following spawning. By contrast, there was a weak positive effect of larval density on settlement, driven by gregarious behaviour at the highest density. When larval supply was saturated, settlement was three times higher in crevices compared with exposed microhabitats, but a negative relationship between settler density and post-settlement survival in crevices and density-independent survival on exposed surfaces resulted in similar recruit densities just one month following settlement. Moreover, a negative relationship was found between turf algae and settler survival in crevices, whereas gregarious settlement improved settler survival on exposed surfaces. Overall, our findings reveal divergent responses by coral larvae and newly settled recruits to density-dependent regulation, mediated by intrinsic and environmental interactions.