Description: Crustacean larvae have served as distinguished models in the field of Ecological Developmental Biology (“EcoDevo”) for many decades, a discipline that examines how developmental mechanisms and their resulting phenotype depend on the environmental context. A contemporary line of research in EcoDevo aims at gaining insights into the immediate tolerance of organisms and their evolutionary potential to adapt to the changing abiotic and biotic environmental conditions created by anthropogenic climate change. Thus, an EcoDevo perspective may be critical to understand and predict the future of organisms in a changing world. Many decapod crustaceans display a complex life cycle that includes pelagic larvae and, in many subgroups, benthic juvenile–adult stages so that a niche shift occurs during the transition from the larval to the juvenile phase. Already at hatching, the larvae possess a wealth of organ systems, many of which also characterise the adult animals, necessary for autonomously surviving and developing in the plankton and suited to respond adaptively to fluctuations of environmental drivers. They also display a rich behavioural repertoire that allows for responses to environmental key factors such as light, hydrostatic pressure, tidal currents, and temperature. Cells, tissues, and organs are at the basis of larval survival, and as the larvae develop, their organs continue to grow in size and complexity. To study organ development, researchers need a suite of state-of-the-art methods adapted to the usually very small size of the larvae. This review and the companion paper set out to provide an overview of methods to study organogenesis in decapod larvae. This first section focuses on larval rearing, preparation, and fixation, whereas the second describes methods to study cells, tissues, and organs.

Description: Understanding the response of biotic systems to multiple environmental drivers is one of the major concerns in ecology. The most common approach in multiple driver research includes the classification of interactive responses into categories (antagonistic, synergistic). However, there are situations where the use of classification schemes limits our understanding or cannot be applied. Here, we introduce and explore an approach that allows us to better appreciate variability in responses to multiple drivers. We then apply it to a case, comparing effects of heatwaves on performance of a cold-adapted species and a warm-adapted competitor. The heatwaves had a negative effect on the native (but not on the exotic) species and the approach highlighted that the exotic species was less responsive to multivariate environmental variation than the native species. Overall, we show how the proposed approach can enhance our understanding of variation in responses due to different driver intensities, species, genotypes, ontogeny, life-phases or among spatial scales at any level of biological organization.

Description: Current understanding of species capacities to respond to climate change is limited by the amount of information available about intraspecific variation in the responses. Therefore, we quantified between- and within- population variation in larval performance (survival, development, and growth to metamorphosis) of the shore crab Carcinus maenas in response to key environmental drivers (temperature, salinity) in 2 populations from regions with contrasting salinities (32-33 PSU: Helgoland, North Sea; 16-20 PSU: Kerteminde, Baltic Sea). We also accounted for the effect(s) of salinity experienced during embryogenesis, which differs between populations. We found contrasting patterns between populations and embryonic salinity conditions. In the Helgoland population, we observed a strong thermal mitigation of low salinity stress (TMLS) for all performance indicators, when embryos were kept in seawater. The negative effects of low salinity on survival were mitigated at increased temperatures; only at high temperatures were larvae exposed to low salinity able to sustain high growth rates and reduced developmental time, thereby metamorphosing with comparable levels of carbon and nitrogen to those reared in seawater. By contrast, larvae from the Kerteminde population showed a detrimental effect of low salinity, consistent with a maladaptive response and a weak TMLS. Low salinity experienced during embryogenesis pre-empted the development of TMLS in both populations, and reduced survival for the Kerteminde population, which is exposed to low salinity. Our study emphasises the importance of evaluating species responses to variation in temperature and salinity across populations; the existence of maladaptive responses and the importance of the maternal habitat should not be underestimated.

Description: Rhizocephalan cirripedes are a very unique group of parasites infecting decapod crustaceans, but apart from a few well- studied species, little is known on their ecology and impact on hosts. Here we report on the results of a 14-month study of infestations of the rhizocephalan Parasacculina leptodiae in the rocky shore crab Leptodius exaratus along the shores of Kuwait in the Persian Gulf (Arabian Gulf). Monthly samples along an intertidal gradient revealed a slightly higher prevalence of P. leptodiae in female (18%) compared to male crabs (11%) and marked differences in prevalence among the sampling sites. Crabs from more sheltered locations in Kuwait Bay showed lower prevalence of P. leptodiae compared to crabs from more exposed sites. Seasonal patterns were largely absent, but prevalence in female crabs showed some monthly variation depending on the site. Rhizocephalan prevalence was generally highest in both crab sexes at the lower shores. This possibly resulted from lower exposure of crabs to infective stages in the higher intertidal and movements of infected crabs to lower parts of the shore. Prevalence of ovigerous females significantly declined with increasing local parasite prevalence. This suggests that the well-known castrating effects of rhizocephalans on individual hosts can also affect local crab reproduction at the population level which has not been shown before. Our results indicate that the rhizocephalan P. leptodiae is a com- mon parasite of the rocky shore crab L. exaratus along the shores of Kuwait, with potential effects on the crab’s population dynamics which warrants further study.

Description: Predicting range expansion of invasive species is one of the key challenges in ecology. We modelled the phenological window for successful larval release and development (WLR) in order to predict poleward expansion of the invasive crab Hemigrapsus sanguineus along the Atlantic coast of North America and north Europe. WLR quantifies the number of opportunities (in days) when larval release leads to a successful completion of the larval phase; WLR depends on the effects of temperature on the duration of larval development and survival. Successful larval development is a necessary requirement for the establishment of self‐persistent local populations. WLR was computed from a mechanistic model, based on in situ temperature time series and a laboratory–calibrated curve predicting duration of larval development from temperature. As a validation step, we checked that model predictions of the time of larval settlement matched observations from the field for our local population (Helgoland, North Sea). We then applied our model to the North American shores because larvae from our European population showed, in the laboratory, similar responses to temperature to those of a North American population. WLR correctly predicted the northern distribution limit in North American shores, where the poleward expansion of H. sanguineus appear to have stalled (as of 2015). For north Europe, where H. sanguineus is a recent invader, WLR predicted ample room for poleward expansion towards NE England and S Norway. We also explored the importance of year‐to‐year variation in temperature for WLR and potential expansion: variations in WLR highlighted the role of heat waves as likely promoters of recruitment subsidising sink populations located at the distribution limits. Overall, phenological windows may be used as a part of a warning system enabling more targeted programs for monitoring.

Description: Understanding biological responses to environmental fluctuations (e.g. heatwaves) is a critical goal in ecology. Biological responses (e.g. survival) are usually measured with respect to different time reference frames, i.e. at specific chronological times (e.g. at specific dates) or biological times (e.g. at reproduction). Measuring responses on the biological frame is central to understand how environmental fluctuation modifies fitness and population persistence. We use a framework, based on partial differential equations (PDEs) to explore how responses to the time scale and magnitude of fluctuations in environmental variables (= drivers) depend on the choice of reference frame. The PDEs and simulations enabled us to identify different components, responsible for the phenological and eco-physiological effects of each driver on the response. The PDEs also highlight the conditions when the choice of reference frame affects the sensitivity of the response to a driver and the type of join effect of two drivers (additive or interactive) on the response. Experiments highlighted the importance of studying how environmental fluctuations affect biological time keeping mechanisms, to develop mechanistic models. Our main result, that the effect of the environmental fluctuations on the response depends on the scale used to measure time, applies to both field and laboratory conditions. In addition, our approach, applied to experimental conditions, can helps us quantify how biological time mediates the response of organisms to environmental fluctuations.

Description: Extensive marine benthos surveys have resulted in a solid understanding of the broad distribution pattern of seafloor biotopes in the southeastern North Sea (temperate northeast Atlantic region). However, due to the low spatial resolution of large-scale surveys, specific smaller-scale biotopes with scattered distribution have been insufficiently captured. Consequently, knowledge regarding the environmental characteristics and species inventories of some specific biotopes is still limited. We investigated the habitat characteristics and the macroinfauna (i.e., organisms in samples collected by a sediment grab and retained in a sieve with a mesh size of 1000 μm) of a spatially restricted, patchy coarse sediment (i.e., grain size fraction 〉500 μm accounting for ≥60% of the total sample mass) biotope in the German Bight over three consecutive years. Habitat and faunal characteristics were contrasted with four other benthic biotopes sampled at the same time to allow for a comparative evaluation. Our study revealed considerable fluctuations in grain size distribution among samples of the coarse sediment, potentially resulting from a frequent redistribution of sediments. A total number of 243 infauna taxa were identified at the 66 stations sampled over three consecutive years (16–33 stations per year) with a considerable proportion of endangered and rare species. The results highlight that previous studies have underestimated the species richness of the biotope. The focus on this previously poorly studied biotope type allowed us to detect species in the study region that were formerly unreported. The macro-infauna in the coarse sediments was characterized by comparatively high abundance and biomass, which may provide a rich food resource for organisms from higher trophic levels. Therefore, coarse sediments likely are an ecologically valuable seafloor biotope despite its limited coverage.