Disparate response of decapods to low pH: A meta-analysis of life history, physiology and behavior traits across life stages and environments.

We employed a meta-analysis to determine if the presumed resilience of decapods to ocean acidification extends to all biological aspects, environments, and life stages. Most response categories appeared unaffected by acidification. However, certain fitness-related traits (growth, survival, and, to some extent, calcification) were impacted. Acid-base balance and stress response scaled positively with reductions in pH, which maintains homeostasis, possibly at the cost of other processes. Juveniles were the only stage impacted by acidification, which is believed to reduce recruitment. We observed few differences in responses to acidification among decapods inhabiting contrasting environments. Our meta-analysis shows decapods as a group slightly to moderately sensitive to low pH, with impacts on some biological aspects rather than on all specific life stages or habitats. Although extreme pH scenarios may not occur in the open ocean, coastal and estuarine areas might experience lower pH levels in the near to medium future, posing potential challenges for decapods.

Size distribution and sex ratio between populations of the artisanal harvested land crab Cardisoma guanhumi (Decapoda: Gecarcinidae), with the estimation of relative growth and size at sexual maturity in Puerto Rico / Distribución de tallas y proporción de sexos entre poblaciones del cangrejo de tierra cosechado artesanalmente Cardisoma guanhumi (Decapoda: Gecarcinidae), con la estimación del crecimiento relativo y tamaño en la madurez sexual en Puerto Rico

Abstract Introduction: Knowledge of growth patterns, sex ratio, and sexual maturity are of importance to exploited populations. The land crab Cardisoma guanhumi is an artisanal and subsistence exploited species in Puerto Rico. However, the growth patterns and sexual maturity of the local populations are not known. Objectives: This study has a double objective: (1) to compare the size and sex structure between populations and (2) to model the relative growth of structures related to reproduction to estimate the average size of morphometric sexual maturity (MSM) for both males and females. Methods: A total of 2 849 specimens were captured from nine dispersed populations on the island between 2001 and 2020. Carapace width (CW) was measured as an estimator of the absolute size of all individuals, together with the propodus length (PL) in males and the abdomen width in females (AW). Differences in length structure between sexes and populations were tested by applying goodness-of-fit tests based on Kernel Density Estimators (KDE). The relative growth pattern was modeled adjusting a spline from which the maximum of its second derivative was calculated as an estimator of the MSM, and bootstrapping was used to generate confidence intervals. Results: Differences were found in size structures, between sexes, and between sites. Our estimates of morphometric sexual maturity resulted in a sexual maturity size for males between 57.9 and 79.0 mm CW, while in females, morphological maturity occurs between 43.8 to 51.5 mm CW. Conclusions: We found inter-population differences in body size that can be attributed to differences in the history of changes of land use and the exploitation biased towards larger individuals, though differences in recruitment should also be considered. Current regulations in Puerto Rico protect female crabs but not to larger male crabs. The regulation establishes that crabs smaller than 64 mm carapace width cannot be captured, leaving immature male crabs over 64 mm CW unprotected. We suggest considering different size limits depending on the sex of the crab. This will allow the full range of sizes where sexual maturity is reached to be protected, increasing the likelihood of the population's size to increase.

Resumen Introducción: El conocimiento de los patrones de crecimiento, la proporción de sexos y la madurez sexual son de suma importancia para las poblaciones explotadas. El cangrejo terrestre Cardisoma guanhumi es una especie explotada artesanalmente y para subsistencia en Puerto Rico. Sin embargo, se desconocen los patrones de crecimiento y la madurez sexual para la isla. Objetivos: Este estudio tiene un doble objetivo: (1) comparar el tamaño y la estructura sexual entre las poblaciones y (2) modelar el crecimiento relativo de estructuras relacionadas con la reproducción (propodo en machos y abdomen en hembras) para estimar el tamaño promedio de madurez sexual morfométrica (MSM). Métodos: Se capturaron un total de 2 849 especímenes de nueve poblaciones dispersas en la isla, entre 2001 y 2020. Se midió el ancho del caparazón (AC) como un estimador del tamaño absoluto de todos los individuos, junto con el largo de propodo en los machos y el ancho del abdomen en las hembras. Las diferencias en la estructura de tallas entre sexos y poblaciones se probaron aplicando la prueba de bondad de ajuste basado en estimaciones de densidad kernel (KDE). El patrón de crecimiento relativo se modeló ajustando un spline a partir del cual se calculó el máximo de su segunda derivada como estimador del MSM y se utilizó bootstrapping para generar intervalos de confianza. Resultados: Se encontraron diferencias en las estructuras de tallas, entre sexos y poblaciones. Nuestras estimaciones de madurez sexual morfométrica dieron como resultado un tamaño de madurez sexual para los machos entre 57.9 y 79.0 mm AC, mientras que, en las hembras, la madurez morfométrica se produce entre 43.8 y 51.5 mm AC. Conclusiones: La diferencia en el tamaño de la población se puede atribuir a las diferencias entre poblaciones en el cambio de uso de la tierra y la presión de explotación asociada con el sesgo hacia la captura de individuos más grandes o el reclutamiento. Las regulaciones actuales en Puerto Rico brindan protección a las hembras, pero no a los machos más grandes. El reglamento establece que no se pueden capturar cangrejos de menos de 64 mm de AC dejando desprotegidos los cangrejos machos inmaduros de más de 64 mm AC. Sugerimos considerar diferentes límites de tamaño según el sexo del cangrejo. Esto permitirá proteger un rango más amplio de tamaños donde se alcanza la madurez sexual, aumentando la probabilidad de un incremento en el tamaño de la población.

Impact on reproductive performance and body condition in a small limpet parasitized by a large castrator pea crab.

Parasitic castrators utilize the energy reserves that the host allocates on reproduction resulting in sterilization of the host. However, whether other processes (e.g., growth) are also impaired depends on the balance between the castrator's energy requirements and the available resources that the castrated host does not use in reproduction. We investigated whether the castrator pea crab Calytraeotheres garthi alters body condition, reproductive performance, and occurrence of copulation in its limpet host Bostrycapulus odites. For this purpose, we examined the anatomy of the ovary, the seminal receptacles, and the body condition of parasitized and unparasitized limpets. The histology of the female gonad of parasitized limpets revealed the ovarian tubules are smaller and characterized by a greater proportion of intertubular and intratubular free space compared with non-parasitized individuals. The body condition of female limpets from all sizes (during summer) and those larger than ~16 mm (during spring and autumn) were impacted by the pea crab. These results are in contrast to that previously reported in the comparatively larger limpet species Crepidula cachimilla, in which the same pea crab species does not alter or even increase the host body weight. We concluded this pea crab species could drastically impair its host's reproduction and body condition although deleterious effects are species-specific and likely depend on limpet body size. The histology of seminal receptacles revealed an uncommon disposition of spermatozoa (i.e., excessive debris and acrosomes detached from epithelium) in seminal receptacles of some parasitized limpets. However, this analysis failed to determine whether sperm derived from present matings or previous pea crab infection. Further studies are needed to address whether pea crabs interfere with the mating behavior of limpets and if the alterations in sperm disposition are a consequence of castration.

The ontogeny of the female reproductive system in the parasitic castrator pea crab Calyptraeotheres garthi: Implications for its mating system.

The knowledge of the mating system of pea crabs is still fragmentary as it remains dubious whether females copulate in the juvenile and free-living 'hard' or in the obligatory symbiotic stages (adult stage 'V' or intermediate stages II to IV). To discriminate between these two possibilities, we analysed the female seminal receptacles, vagina and opercula, and the sperm content in different stages of the pea crab Calyptraeotheres garthi. Our histology and scanning electron microscopy results revealed that in the hard stage the seminal receptacle is simple without secretory epithelia, and vagina and opercula are not controlled by musculature. In stages II to IV, the seminal receptacles, vagina, and opercula are under development and these structures reach maturity in stage V. These results suggest that females become receptive in stage V and not during predating stages. We found no spermatozoa in SR of 'hard' and stage II to IV females while these structures were loaded of sperm in most stage V, indicating that females start to mate in stage V. Our results support the notion that males of C. garthi roam among hosts in search for sedentary stage V females, as predicted by Baeza and Thiel's () model of mating systems for symbiotic crustaceans. Nevertheless, we failed to reveal whether females mate repeatedly: the accumulation of sperm in larger females might indicate occurrence of multiple copula or a high variability in male sperm transfer.

Functional morphology of the female reproductive system of a crab with highly extensible seminal receptacles and extreme sperm storage capacity.

We studied the functional morphology of the female reproductive system of the purple stone crab Danielethus crenulatus. The most remarkable feature is the relative storage capacity and extensibility of the seminal receptacles. These receptacles are a pair of simple sacs that lack internal structures dividing the internal lumen. Differences in seminal receptacle size and contents are accompanied by conspicuous changes in receptacle lining at a tissue level. Full seminal receptacles contain discrete sperm masses formed by hardened fluid and densely packed spermatophores. Different sperm masses are likely from different mates and their stratified disposition within the seminal receptacles is compatible with rival sperm displacement and last sperm precedence. Additionally, the anatomical structure of the vulva and vagina suggest active female control over copula. We discuss our results in the general context of sperm storage in brachyurans and the implications for the mating system of this species.

Hard-shell mating in Neohelice granulata: the role of ecdysone in female receptivity and mate attraction.

Most brachyuran females become receptive during the intermolt period, a condition considered "derived". However, as far as we know, studies testing the existence and function of pheromones in decapods are based on species which have mating linked to molting, a condition considered as "ancestral". For the first time, we studied some physiological and morphological processes involved in Neohelice granulata intermolt female crabs becoming receptive and potentially attracting males. We found that receptive females have mobile vulvae opercula due to a softening process of the cuticle hinge which showed lower calcium levels compared to the hinge of unreceptive females. Local softening of the hinge was stimulated by a low concentration of ecdysone during the intermolt period. A putative pheromone liberated by receptive females to attract males is presumed to be released through the mobile vulvae and not through the urine.

Predation risk modifies behaviour by shaping the response of identified brain neurons.

Interpopulation comparisons in species that show behavioural variations associated with particular ecological disparities offer good opportunities for assessing how environmental factors may foster specific functional adaptations in the brain. Yet, studies on the neural substrate that can account for interpopulation behavioural adaptations are scarce. Predation is one of the strongest driving forces for behavioural evolvability and, consequently, for shaping structural and functional brain adaptations. We analysed the escape response of crabs ITALIC! Neohelice granulatafrom two isolated populations exposed to different risks of avian predation. Individuals from the high-risk area proved to be more reactive to visual danger stimuli (VDS) than those from an area where predators are rare. Control experiments indicate that the response difference was specific for impending visual threats. Subsequently, we analysed the response to VDS of a group of giant brain neurons that are thought to play a main role in the visually guided escape response of the crab. Neurons from animals of the population with the stronger escape response were more responsive to VDS than neurons from animals of the less reactive population. Our results suggest a robust linkage between the pressure imposed by the predation risk, the response of identified neurons and the behavioural outcome.