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The existence of sex‐specific differences in phenotypic traits is widely recognized. Yet they are often ignored in studies looking at the impact of global changes on marine organisms, particularly within the context of combined drivers that are known to elicit complex interactions. We tested sex‐specific physiological responses of the cosmopolitan and ecologically important marine copepod Acartia tonsa exposed to combined hypoxia and marine heatwave (MHW) conditions, both of which individually strongly affect marine ectotherms. Females and males were acutely exposed for 5 days to a combination of either control (18°C) or a high temperature mimicking a MHW (25°C), and normoxia (100% O2 sat.) or mild hypoxia (35% O2 sat.). Life‐history traits, as well as sex‐specific survival and physiological traits, were measured. Females had overall higher thermal tolerance levels and responded differently than males when exposed to the combined global change drivers investigated. Females also showed lower metabolic thermal sensitivity when compared to males. Additionally, the MHW exerted a dominant effect on the traits investigated, causing a lower survival and higher metabolic rate at 25°C. However, egg production rates appeared unaffected by hypoxia and MHW conditions. Our results showed that MHWs could strongly affect copepods' survival, that combined exposure to hypoxia and MHW exerted an interactive effect only on CTmax, and that sex‐specific vulnerability to these global change drivers could have major implications for population dynamics. Our results highlight the importance of considering the differences in the responses of females and males to rapid environmental changes to improve the implementation of climate‐smart conservation approaches. [ABSTRACT FROM AUTHOR]

Background: Annelids are one the most speciose and ecologically diverse groups of metazoans. Although a significant effort has been recently invested in sequencing genomes of a wide array of metazoans, many orders and families within the phylum Annelida are still represented by a single specimen of a single species. The genus of interstitial annelids Ophryotrocha (Dorvilleidae, Errantia, Annelida) is among these neglected groups, despite its extensive use as model organism in numerous studies on the evolution of life history, physiological and ecological traits. To compensate for the paucity of genomic information in this genus, we here obtained novel complete mitochondrial genomes of six Ophryotrocha species using next generation sequencing. In addition, we investigated the evolution of the reproductive mode in the Ophryotrocha genus using a phylogeny based on two mitochondrial markers (COXI and 16S rDNA) and one nuclear fragment (Histone H3). Results: Surprisingly, gene order was not conserved among the six Ophryotrocha species investigated, and varied greatly as compared to those found in other annelid species within the class Errantia. The mitogenome phylogeny for the six Ophryotrocha species displayed a separation of gonochoric and hermaphroditic species. However, this separation was not observed in the phylogeny based on the COX1, 16S rDNA, and H3 genes. Parsimony and Bayesian ancestral trait reconstruction indicated that gonochorism was the most parsimonious ancestral reproductive mode in Ophryotrocha spp. Conclusions: Our results highlight the remarkably high level of gene order variation among congeneric species, even in annelids. This encourages the need for additional mitogenome sequencing of annelid taxa in order to properly understand its mtDNA evolution, high biodiversity and phylogenetic relationships. [ABSTRACT FROM AUTHOR]

Ocean warming (OW) and acidification (OA) are key features of global change and are predicted to have negative consequences for marine species and ecosystems. At a smaller scale increasing oil and gas activities at northern high latitudes could lead to greater risk of petroleum pollution, potentially exacerbating the effects of such global stressors. However, knowledge of combined effects is limited. This study employed a scenario-based, collapsed design to investigate the impact of one local acute stressor (North Sea crude oil) and two chronic global drivers (pH for OA and temperature for OW), alone or in combination on aspects of the biology of larval stages of two key invertebrates: the northern shrimp (Pandalus borealis) and the green sea urchin (Strongylocentrotus droebachiensis). Both local and global drivers had negative effects on survival, development and growth of the larval stages. These effects were species- and stage-dependent. No statistical interactions were observed between local and global drivers and the combined effects of the two drivers were approximately equal to the sum of their separate effects. This study highlights the importance of adjusting regulation associated with oil spill prevention to maximize the resilience of marine organisms to predicted future global conditions. [ABSTRACT FROM AUTHOR]

Changes in environmental conditions can have a profound impact on organismal processes. For aquatic organisms, changes in dissolved oxygen concentration can have major repercussions, notably for physiological processes that have high long-term energetic costs and are more likely to rely on aerobic metabolic pathways. Here we propose a simple college or university-level low-cost laboratory activity that addresses the effect of differences in oxygen concentration on an important physiological property of planarian flatworms: their regeneration capacity. Students were asked to perform measurements and statistically compare the efficacy of the regeneration process between planarians kept in normoxic conditions and others kept in hypoxic conditions. The activity was evaluated by asking students to produce a short laboratory report with all of the sections that a scientific article generally comprises. This hands-on experiment has the benefit of going beyond how planarians are traditionally used in biodiversity classes (i.e. basic anatomy of flatworms) by combining behavioural, physiological and environmental observations while investigating the effects of an environmental variable on the regeneration response. We would further encourage professors to expand even more upon this framework by providing the students with an opportunity to actively engage in all steps inherent to the scientific process. [ABSTRACT FROM AUTHOR]

The O2 content of the global ocean has been declining progressively over the past decades, mainly because of human activities and global warming. Nevertheless, how long‐term deoxygenation affects macrobenthic communities, sediment biogeochemistry and their mutual feedback remains poorly understood. Here, we evaluate the response of the benthic assemblages and biogeochemical functioning to decreasing O2 concentrations along the persistent bottom‐water dissolved O2 gradient of the Estuary and Gulf of St. Lawrence (QC, Canada). We report several of non‐linear biodiversity and functional responses to decreasing O2 concentrations, and identify an O2 threshold that occurs at approximately at 63 μM. Below this threshold, macrobenthic community assemblages change, and bioturbation rates drastically decrease to near zero. Consequently, the sequence of electron acceptors used to metabolize the sedimentary organic matter is squeezed towards the sediment surface while reduced compounds accumulate closer (as much as 0.5–2.5 cm depending on the compound) to the sediment–water interface. Our results illustrate the capacity of bioturbating species to compensate for the biogeochemical consequences of hypoxia and can help to predict future changes in benthic ecosystems. [ABSTRACT FROM AUTHOR]

Linking variation in species' traits to large-scale environmental gradients can lend insight into the evolutionary processes that have shaped functional diversity and future responses to environmental change. Here, we ask how heat and cold tolerance vary as a function of latitude, elevation and climate extremes, using an extensive global dataset of ectotherm and endotherm thermal tolerance limits, while accounting for methodological variation in acclimation temperature, ramping rate and duration of exposure among studies. We show that previously reported relationships between thermal limits and latitude in ectotherms are robust to variation in methods. Heat tolerance of terrestrial ectotherms declined marginally towards higher latitudes and did not vary with elevation, whereas heat tolerance of freshwater and marine ectotherms declined more steeply with latitude. By contrast, cold tolerance limits declined steeply with latitude in marine, intertidal, freshwater and terrestrial ectotherms, and towards higher elevations on land. In all realms, both upper and lower thermal tolerance limits increased with extreme daily temperature, suggesting that different experienced climate extremes across realms explain the patterns, as predicted under the Climate Extremes Hypothesis. Statistically accounting for methodological variation in acclimation temperature, ramping rate and exposure duration improved model fits, and increased slopes with extreme ambient temperature. Our results suggest that fundamentally different patterns of thermal limits found among the earth's realms may be largely explained by differences in episodic thermal extremes among realms, updating global macrophysiological 'rules'. This article is part of the theme issue 'Physiological diversity, biodiversity patterns and global climate change: testing key hypotheses involving temperature and oxygen'. [ABSTRACT FROM AUTHOR]

Inter‐individual variation in phenotypic traits has long been considered as "noise" rather than meaningful phenotypic variation, with biological studies almost exclusively generating and reporting average responses for populations and species' average responses. Here, we compare the use of an individual approach in the investigation of extracellular acid–base regulation by the purple sea urchin Paracentrotus lividus challenged with elevated pCO2 and temperature conditions, with a more traditional approach which generates and formally compares mean values. We detected a high level of inter‐individual variation in acid–base regulation parameters both within and between treatments. Comparing individual and mean values for the first (apparent) dissociation constant of the coelomic fluid for individual sea urchins resulted in substantially different (calculated) acid–base parameters, and models with stronger statistical support. While the approach using means showed that coelomic pCO2 was influenced by seawater pCO2 and temperature combined, the individual approach indicated that it was in fact seawater temperature in isolation that had a significant effect on coelomic pCO2. On the other hand, coelomic [HCO3−] appeared to be primarily affected by seawater pCO2, and less by seawater temperature, irrespective of the approach adopted. As a consequence, we suggest that individual variation in physiological traits needs to be considered, and where appropriate taken into account, in global change biology studies. It could be argued that an approach reliant on mean values is a "procedural error." It produces an artefact, that is, a population's mean phenotype. While this may allow us to conduct relatively simple statistical analyses, it will not in all cases reflect, or take into account, the degree of (physiological) diversity present in natural populations. [ABSTRACT FROM AUTHOR]

Using a targeted metabolomic approach we investigated the effects of low seawater pH on energy metabolism in two late copepodite stages (CIV and CV) of the keystone Arctic copepod species Calanus glacialis. Exposure to decreasing seawater pH (from 8.0 to 7.0) caused increased ATP, ADP and NAD+ and decreased AMP concentrations in stage CIV, and increased ATP and phospho-L-arginine and decreased AMP concentrations in stage CV. Metabolic pathway enrichment analysis showed enrichment of the TCA cycle and a range of amino acid metabolic pathways in both stages. Concentrations of lactate, malate, fumarate and alpha-ketoglutarate (all involved in the TCA cycle) increased in stage CIV, whereas only alpha-ketoglutarate increased in stage CV. Based on the pattern of concentration changes in glucose, pyruvate, TCA cycle metabolites, and free amino acids, we hypothesise that ocean acidification will lead to a shift in energy production from carbohydrate metabolism in the glycolysis toward amino acid metabolism in the TCA cycle and oxidative phosphorylation in stage CIV. In stage CV, concentrations of most of the analysed free fatty acids increased, suggesting in particular that ocean acidification increases the metabolism of stored wax esters in this stage. Moreover, aminoacyl-tRNA biosynthesis was enriched in both stages indicating increased enzyme production to handle low pH stress. [ABSTRACT FROM AUTHOR]

Giant clams produce massive calcified shells with important biological (e.g., defensive) and ecological (e.g., habitat-forming) properties. Whereas elevated seawater temperature is known to alter giant clam shell structure, no study has examined the effects of a simultaneous increase in seawater temperature and partial pressure of carbon dioxide (pCO2) on shell mineralogical composition in these species. We investigated the effects of 60-days exposure to end-of-the-century projections for seawater temperature (+ 3 °C) and pCO2 (+ 500 µatm) on growth, mineralogy, and organic content of shells and scutes in juvenile Tridacna squamosa giant clams. Elevated temperature had no effect on growth rates or organic content, but did increase shell [24Mg]/[40Ca] as well as [40Ca] in newly-formed scutes. Elevated pCO2 increased shell growth and whole animal mass gain. In addition, we report the first evidence of an effect of elevated pCO2 on element/Ca ratios in giant clam shells, with significantly increased [137Ba]/[40Ca] in newly-formed shells. Simultaneous exposure to both drivers greatly increased inter-individual variation in mineral concentrations and resulted in reduced shell N-content which may signal the onset of physiological stress. Overall, our results indicate a greater influence of pCO2 on shell mineralogy in giant clams than previously recognized. [ABSTRACT FROM AUTHOR]

Phenotypic plasticity in parental care investment allows organisms to promptly respond to rapid environmental changes by potentially benefiting offspring survival and thus parental fitness. To date, a knowledge gap exists on whether plasticity in parental care behaviors can mediate responses to climate change in marine ectotherms. Here, we assessed the plasticity of parental care investment under elevated temperatures in a gonochoric marine annelid with biparental care, Ophryotrocha labronica, and investigated its role in maintaining the reproductive success of this species in a warming ocean. We measured the time individuals spent carrying out parental care activities across three phases of embryonic development, as well as the hatching success of the offspring as a proxy for reproductive success, at control (24℃) and elevated (27℃) temperature conditions. Under elevated temperature, we observed: (a) a significant decrease in total parental care activity, underpinned by a decreased in male and simultaneous parental care activity, in the late stage of embryonic development; and (b) a reduction in hatching success that was however not significantly related to changes in parental care activity levels. These findings, along with the observed unaltered somatic growth of parents and decreased brood size, suggest that potential cost‐benefit trade‐offs between offspring survival (i.e., immediate fitness) and parents' somatic condition (i.e., longer‐term fitness potential) may occur under ongoing ocean warming. Finally, our results suggest that plasticity in parental care behavior is a mechanism able to partially mitigate the negative effects of temperature‐dependent impacts. [ABSTRACT FROM AUTHOR]

In a recent mechanistic study, octopamine was shown to promote proton transport over the branchial epithelium in green crabs, Carcinus maenas. Here, we follow up on this finding by investigating the involvement of octopamine in an environmental and physiological context that challenges acid-base homeostasis, the response to short-term high p CO 2 exposure (400 Pa) in a brackish water environment. We show that hyperregulating green crabs experienced a respiratory acidosis as early as 6 h of exposure to hypercapnia, with a rise in hemolymph p CO 2 accompanied by a simultaneous drop of hemolymph pH. The slightly delayed increase in hemolymph HCO 3 − observed after 24 h helped to restore hemolymph pH to initial values by 48 h. Circulating levels of the biogenic amine octopamine were significantly higher in short-term high p CO 2 exposed crabs compared to control crabs after 48 h. Whole animal metabolic rates, intracellular levels of octopamine and cAMP, as well as branchial mitochondrial enzyme activities for complex I + III and citrate synthase were unchanged in posterior gill #7 after 48 h of hypercapnia. However, application of octopamine in gill respirometry experiments suppressed branchial metabolic rate in posterior gills of short-term high p CO 2 exposed animals. Furthermore, branchial enzyme activity of cytochrome C oxidase decreased in high p CO 2 exposed crabs after 48 h. Our results indicate that hyperregulating green crabs are capable of quickly counteracting a hypercapnia-induced respiratory acidosis. The role of octopamine in the acclimation of green crabs to short-term hypercapnia seems to entail the alteration of branchial metabolic pathways, possibly targeting mitochondrial cytochrome C in the gill. Our findings help advancing our current limited understanding of endocrine components in hypercapnia acclimation. Acid-base compensation upon short-term high p CO 2 exposure in hyperregulating green crabs started after 6 h and was accomplished by 48 h with the involvement of the biogenic amine octopamine, accumulation of hemolymph HCO 3 −, and regulation of mitochondrial complex IV (cytochrome C oxidase). [Display omitted] • Hyperregulating green crabs start compensating upon a short-term high p CO 2 exposure after 6 h. • Extracellular HCO 3 − to buffer hemolymph successfully by 48 h is retained by gills. • Octopamine plays a role in acid-base balance and compensation. • Exposure to short-term high p CO 2 results in decreased cytochrome C oxidase activity. • Octopamine might target branchial cytochrome C oxidase. [ABSTRACT FROM AUTHOR]

Tens of thousands of anthropogenic chemicals and wastes enter the marine environment each year as a consequence of the ever-increasing anthropogenic activities and demographic growth of the human population, which is majorly concentrated along coastal areas. Marine ecotoxicology has had a crucial role in helping shed light on the fate of chemicals in the environment, and improving our understanding of how they can affect natural ecosystems. However, chemical contamination is not occurring in isolation, but rather against a rapidly changing environmental horizon. Most environmental studies have been focusing on short-term within-generation responses of single life stages of single species to single stressors. As a consequence, one-dimensional ecotoxicology cannot enable us to appreciate the degree and magnitude of future impacts of chemicals on marine ecosystems. Current approaches that lack an evolutionary perspective within the context of ongoing and future local and global stressors will likely lead us to under or over estimations of the impacts that chemicals will exert on marine organisms. It is therefore urgent to define whether marine organisms can acclimate, i.e. adjust their phenotypes through transgenerational plasticity, or rapidly adapt, i.e. realign the population phenotypic performances to maximize fitness, to the new chemical environment within a selective horizon defined by global changes. To foster a significant advancement in this research area, we review briefly the history of ecotoxicology, synthesis our current understanding of the fate and impact of contaminants under global changes, and critically discuss the benefits and challenges of integrative approaches toward developing an evolutionary perspective in marine ecotoxicology: particularly through a multigenerational approach. The inclusion of multigenerational studies in Ecological Risk Assessment framework (ERA) would provide significant and more accurately information to help predict the risks of pollution in a rapidly changing ocean. Unlabelled Image • Chemical contamination is occurring against global changes. • The interaction of pollutants with global change drivers has been overlooked. • A new perspective to assess chemicals effects in the future oceanscape is need. • The integration of global change evolutionary biology and ecotoxicology is required. [ABSTRACT FROM AUTHOR]

Genome size, known also as the C-value, has been proposed as an important determinant of life-history variation in numerous animal taxa. We assessed the relationships between genome size and fitness-related life-history traits in six species of interstitial marine annelids of the genus Ophryotrocha. Life-history traits and genome size data obtained from 18 additional annelid species were included in our analyses to have a broader phylogenetic scope. Unexpectedly, genome sizes assessed here by flow cytometry in four Ophryotrocha species were three times larger than previously reported values obtained using Feulgen densitometry. This has implications for the hypothesis that harsh interstitial habitats select for small genomes in meiofaunal annelids. Within the genus Ophryotrocha, significant and positive relationships were found between genome size and nucleus size, and between genome size, age at first egg mass deposition, body size and lifespan. These relationships held up in the broader phylogenetic comparison. Our study provides evidence for the important role played by genome size in the evolution of life-history traits in annelids. [ABSTRACT FROM AUTHOR]

Trans-generational plasticity (TGP) represents a primary mechanism for guaranteeing species persistence under rapid global changes. To date, no study on TGP responses of marine organisms to global change scenarios in the ocean has been conducted on phylogenetically closely related species, and we thus lack a true appreciation for TGP inter-species variation. Consequently, we examined the tolerance and TGP of life-history and physiological traits in two annelid species within the genus Ophryotrocha: one rare (O. robusta) and one common (O. japonica). Both species were exposed over two generations to ocean acidification (OA) and warming (OW) in isolation and in combination (OAW). Warming scenarios led to a decrease in energy production together with an increase in energy requirements, which was lethal for O. robusta before viable offspring could be produced by the F1. Under OA conditions, O. robusta was able to reach the second generation, despite showing lower survival and reproductive performance when compared to control conditions. This was accompanied by a marked increase in fecundity and egg volume in F2 females, suggesting high capacity for TGP under OA. In contrast, O. japonica thrived under all scenarios across both generations, maintaining its fitness levels via adjusting its metabolomic profile. Overall, the two species investigated show a great deal of difference in their ability to tolerate and respond via TGP to future global changes. We emphasize the potential implications this can have for the determination of extinction risk, and consequently, the conservation of phylogenetically closely related species. [ABSTRACT FROM AUTHOR]

Little is known about the life-history trade-offs and limitations, and the physiological mechanisms that are associated with phenotypic adaptation to future ocean conditions. To address this knowledge gap, we investigated the within- and trans-generation life-history responses and aerobic capacity of a marine polychaete, Ophryotrocha labronica, to elevated temperature and elevated temperature combined with elevated salinity for its entire lifespan. In addition, transplants between treatments were carried out at both the egg mass and juvenile stage to identify the potential influence of developmental effects. Within-generation, life-history trade-offs caused by the timing of transplant were only detected under elevated temperature combined with elevated salinity conditions. Polychaetes transplanted at the egg mass stage grew slower and had lower activities of energy metabolism enzymes but reached a larger maximum body size and lived longer when compared with those transplanted as juveniles. Trans-generation exposure to both elevated temperature and elevated temperature and salinity conditions restored 20 and 21% of lifespan fecundity, respectively. Trans-generation exposure to elevated temperature conditions also resulted in a trade-off between juvenile growth rates and lifespan fecundity, with slower growers showing greater fecundity. Overall, our results suggest that future ocean conditions may select for slower growers. Furthermore, our results indicate that lifehistory trade-offs and limitations will be more prevalent with the shift of multiple global change drivers, and thus there will be greater constraints on adaptive potential. [ABSTRACT FROM AUTHOR]

Abstract The transition from the last pelagic larval stage to the first benthic juvenile stage in the complex life cycle of marine invertebrates, such as the American lobster Homarus americanus, a species of high economic importance, represents a delicate phase in these species development. Under future elevated p CO 2 conditions, ocean acidification and other elevated p CO 2 events can negatively affect crustaceans. This said their effects on the benthic settlement phase are virtually unknown. This study aimed to identify the effects of elevated seawater p CO 2 on stage V American lobsters exposed to seven p CO 2 levels. The survival, development time, metabolic and feeding rates, carapace composition, and energy metabolism enzyme function were investigated. Results suggested an increase in mortality, slower development and an increase in aerobic capacity with increasing p CO 2. Our study points to potential reduction in juvenile recruitment success as seawater p CO 2 increases, thus foreshadowing important socio-economic repercussions for the lobster fisheries and industry. Highlights • Responses were largely linear, traits being predictable across the p CO 2 gradient. • Oxygen consumption rates was not affected by elevated p CO 2 levels. • Exposure to elevated p CO 2 increased mortality and intermoult period. • Reduced aerobic capacity at high p CO 2 suggested energy metabolism reorganization. [ABSTRACT FROM AUTHOR]

Many aquatic animals grow into colonies of repeated, genetically identical, modules (zooids). Zooid interconnections enable colonies to behave as integrated functional units, while plastic responses to environmental changes may affect individual zooids. Plasticity includes the variable partitioning of resources to sexual reproduction, colony growth and maintenance. Maintenance often involves regeneration, which is also a routine part of the life history in some organisms, such as bryozoans. Here we investigate changes in regenerative capacity in the encrusting bryozoan Cryptosula pallasiana when cultured at different seawater p CO 2 levels. The proportion of active zooids showing polypide regeneration was highest at current oceanic pH (8.1), but decreased progressively as pH declined below that value, reaching a six-fold reduction at pH 7.0. The zone of budding of new zooids at the colony periphery declined in size below pH 7.7. Under elevated p CO 2 conditions, already experienced sporadically in coastal areas, skeletal corrosion was accompanied by the proportional reallocation of resources from polypide regeneration in old zooids to the budding of new zooids at the edge of the colony. Thus, future ocean acidification can affect colonial organisms by changing how they allocate resources, with potentially profound impacts on life-history patterns and ecological interactions. [ABSTRACT FROM AUTHOR]

Little is known of the capacity that marine metazoans have to evolve under rapid p CO2 changes. Consequently, we reared a marine polychaete, Ophryotrocha labronica, previously cultured for approximately 33 generations under a low/variable p H regime, under elevated and low p CO2 for six generations. The strain used was found to be tolerant to elevated p CO2 conditions. In generations F1 and F2 females' fecundity was significantly lower in the low p CO2 treatment. However, from generation F3 onwards there were no differences between p CO2 treatments, indicating that trans-generational effects enabled the restoration and maintenance of reproductive output. Whilst the initial fitness recovery was likely driven by trans-generational plasticity ( TGP), the results from reciprocal transplant assays, performed using F7 individuals, made it difficult to disentangle between whether TGP had persisted across multiple generations, or if evolutionary adaptation had occurred. Nonetheless, both are important mechanisms for persistence under climate change. Overall, our study highlights the importance of multi-generational experiments in more accurately determining marine metazoans' responses to changes in p CO2, and strengthens the case for exploring their use in conservation, by creating specific p CO2 tolerant strains of keystone ecosystem species. [ABSTRACT FROM AUTHOR]

An in situ reciprocal transplant experiment was carried around a volcanic CO 2 vent to evaluate the anti-predator responses of an anemone goby species exposed to ambient (∼380 μatm) and high (∼850 μatm) CO 2 sites. Overall, the anemone gobies displayed largely unaffected behaviors under high-CO 2 conditions suggesting an adaptive potential of Gobius incognitus to ocean acidification (OA) conditions. This is also supported by its 3-fold higher density recorded in the field under high CO 2. However, while fish exposed to ambient conditions showed an expected reduction in the swimming activity in the proximity of the predator between the pre- and post-exposure period, no such changes were detected in any of the other treatments where fish experienced acute and long-term high CO 2. This may suggest an OA effect on the goby antipredator strategy. Our findings contribute to the ongoing debate over the need for realistic predictions of the impacts of expected increased CO 2 concentration on fish, providing evidence from a natural high CO 2 system. [Display omitted] • This is the first assessment in the wild of behavioural responses of fish acutely and chronically exposed to elevated CO 2. • High density of anemone goby fish was recorded at high-CO 2 levels off a volcanic CO 2 vent in Vulcano island (Italy). • Acute and chronic exposure to elevated CO 2 did not affect most of the behaviours in adult G. incognitus. • Behavioural plasticity occurred under ocean acidification conditions suggesting potential local adaptation. [ABSTRACT FROM AUTHOR]

Aim In aquatic ecosystems, standing (lentic) and running (lotic) waters differ fundamentally in their stability and persistence, shaping the comparative population genetic structure, geographical range size and speciation rates of lentic versus lotic lineages. While the drivers of this pattern remain incompletely understood, the suite of traits making up the ability of a species to establish new populations is instrumental in determining such differences. Here we explore the degree to which the association between habitat type and geographical range size results from differences in dispersal ability or fundamental niche breadth in the members of the Enochrus bicolor complex, an aquatic beetle clade with species across the lentic-lotic divide. Location Western Mediterranean, with a special focus on North Africa, the Iberian Peninsula and Sicily. Methods DNA sequences for four loci were obtained from species of the E. bicolor complex and analysed using phylogenetic inference. Dispersal and establishment abilities were assessed in lentic-lotic species pairs of the complex, using flight wing morphometrics and thermal tolerance ranges as surrogates, respectively. Results There were clear differences in range size between the lotic and lentic taxa of the complex, which appears to have had a lotic origin with two transitions to standing waters. Only small differences were observed in temperature tolerance and acclimation ability between the two lotic-lentic sister species studied. By contrast, wing morphometrics revealed clear, consistent differences between lotic and lentic Enochrus species pairs, the latter having a higher dispersal capacity. Main conclusions We hypothesize that there have been two habitat shifts from lotic to lentic waters, which have allowed marked expansions in geographical range size in western Mediterranean species of the E. bicolor complex. Differences in dispersal rather than in establishment ability appear to underlie differences in geographical range extent, as transitions to lentic waters were associated with changes in wing morphology, but not in thermal tolerance range. In this lineage of water beetles, selection for dispersal in geologically short-lived lentic systems has driven the evolution of larger range sizes in lentic taxa compared with those of their lotic relatives. [ABSTRACT FROM AUTHOR]

Understanding how temperature determines the distribution of life is necessary to assess species' sensitivities to contemporary climate change. Here, we test the importance of temperature in limiting the geographic ranges of ectotherms by comparing the temperatures and areas that species occupy to the temperatures and areas species could potentially occupy on the basis of their physiological thermal tolerances. We find that marine species across all latitudes and terrestrial species from the tropics occupy temperatures that closely match their thermal tolerances. However, terrestrial species from temperate and polar latitudes are absent from warm, thermally tolerable areas that they could potentially occupy beyond their equatorward range limits, indicating that extreme temperature is often not the factor limiting their distributions at lower latitudes. This matches predictions from the hypothesis that adaptation to cold environments that facilitates survival in temperate and polar regions is associated with a performance trade-off that reduces species' abilities to contend in the tropics, possibly due to biotic exclusion. Our findings predict more direct responses to climate warming of marine ranges and cool range edges of terrestrial species., (© 2023. The Author(s).)

Crustaceans' endocrinology is a vastly understudied area of research. The major focus of the studies on this topic to date has been on the molting cycle (and in particular, the role of crustacean hyperglycemic hormone (CHH)), as well as the role of other hormones in facilitating physiological phenotypic adjustments to salinity changes. Additionally, while many recent studies have been conducted on the acclimation and adaptation capacity of crustaceans to a changing environment, only few have investigated internal hormonal balance especially with respect to an endocrine response to environmental challenges. Consequently, our study aimed to identify and characterize endocrine components of acid-base regulation in the European green crab, Carcinus maenas. We show that both the biogenic amine octopamine (OCT) and the CHH are regulatory components of branchial acid-base regulation. While OCT suppressed branchial proton excretion, CHH seemed to promote it. Both hormones were also capable of enhancing branchial ammonia excretion. Furthermore, mRNA abundance for branchial receptors (OCT-R), or G-protein receptor activated soluble guanylate cyclase (sGC1b), are affected by environmental change such as elevated pCO 2 (hypercapnia) and high environmental ammonia (HEA). Our findings support a role for both OCT and CHH in the general maintenance of steady-state acid-base maintenance in the gill, as well as regulating the acid-base response to environmental challenges that C. maenas encounters on a regular basis in the habitats it dwells in and more so in the future ocean., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Species with a wide distribution can experience significant regional variation in environmental conditions, to which they can acclimatize or adapt. Consequently, the geographic origin of an organism can influence its responses to environmental changes, and therefore its sensitivity to combined global change drivers. This study aimed at determining the physiological responses of the northern shrimp, Pandalus borealis, at different levels of biological organization and from four different geographic origins, exposed to elevated temperature and low pH to define its sensitivity to future ocean warming and acidification. Shrimp sampled within the northwest Atlantic were exposed for 30 days to combinations of three temperature (2, 6 or 10°C) and two pH levels (7.75 or 7.40). Survival, metabolic rates, whole-organism aerobic performance and cellular energetic capacity were assessed at the end of the exposure. Our results show that shrimp survival was negatively affected by temperature above 6°C and low pH, regardless of their origin. Additionally, shrimp from different origins show overall similar whole-organism performances: aerobic scope increasing with increasing temperature and decreasing with decreasing pH. Finally, the stability of aerobic metabolism appears to be related to cellular adjustments specific to shrimp origin. Our results show that the level of intraspecific variation differs among levels of biological organization: different cellular capacities lead to similar individual performances. Thus, the sensitivity of the northern shrimp to ocean warming and acidification is overall comparable among origins. Nonetheless, shrimp vulnerability to predicted global change scenarios for 2100 could differ among origins owing to different regional environmental conditions., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Ocean acidification (OA) affects marine organisms through various physiological and biological processes, yet our understanding of how these translate to large-scale population effects remains limited. Here, we integrated laboratory-based experimental results on the life history and physiological responses to OA of the American lobster, Homarus americanus, into a dynamic bioclimatic envelope model to project future climate change effects on species distribution, abundance, and fisheries catch potential. Ocean acidification effects on juvenile stages had the largest stage-specific impacts on the population, while cumulative effects across life stages significantly exerted the greatest impacts, albeit quite minimal. Reducing fishing pressure leads to overall increases in population abundance while setting minimum size limits also results in more higher-priced market-sized lobsters (> 1 lb), and could help mitigate the negative impacts of OA and concurrent stressors (warming, deoxygenation). However, the magnitude of increased effects of climate change overweighs any moderate population gains made by changes in fishing pressure and size limits, reinforcing that reducing greenhouse gas emissions is most pressing and that climate-adaptive fisheries management is necessary as a secondary role to ensure population resiliency. We suggest possible strategies to mitigate impacts by preserving important population demographics., (© 2021. The Author(s).)

Understanding how species' thermal limits have evolved across the tree of life is central to predicting species' responses to climate change. Here, using experimentally-derived estimates of thermal tolerance limits for over 2000 terrestrial and aquatic species, we show that most of the variation in thermal tolerance can be attributed to a combination of adaptation to current climatic extremes, and the existence of evolutionary 'attractors' that reflect either boundaries or optima in thermal tolerance limits. Our results also reveal deep-time climate legacies in ectotherms, whereby orders that originated in cold paleoclimates have presently lower cold tolerance limits than those with warm thermal ancestry. Conversely, heat tolerance appears unrelated to climate ancestry. Cold tolerance has evolved more quickly than heat tolerance in endotherms and ectotherms. If the past tempo of evolution for upper thermal limits continues, adaptive responses in thermal limits will have limited potential to rescue the large majority of species given the unprecedented rate of contemporary climate change.

Diving as a lifestyle has evolved on multiple occasions when air-breathing terrestrial animals invaded the aquatic realm, and diving performance shapes the ecology and behaviour of all air-breathing aquatic taxa, from small insects to great whales. Using the largest dataset yet assembled, we show that maximum dive duration increases predictably with body mass in both ectotherms and endotherms. Compared to endotherms, ectotherms can remain submerged for longer, but the mass scaling relationship for dive duration is much steeper in endotherms than in ectotherms. These differences in diving allometry can be fully explained by inherent differences between the two groups in their metabolic rate and how metabolism scales with body mass and temperature. Therefore, we suggest that similar constraints on oxygen storage and usage have shaped the evolutionary ecology of diving in all air-breathing animals, irrespective of their evolutionary history and metabolic mode. The steeper scaling relationship between body mass and dive duration in endotherms not only helps explain why the largest extant vertebrate divers are endothermic rather than ectothermic, but also fits well with the emerging consensus that large extinct tetrapod divers (e.g. plesiosaurs, ichthyosaurs and mosasaurs) were endothermic.

Regulation of extracellular acid-base balance, while maintaining energy metabolism, is recognised as an important aspect when defining an organism's sensitivity to environmental changes. This study investigated the haemolymph buffering capacity and energy metabolism (oxygen consumption, haemolymph [l-lactate] and [protein]) in early benthic juveniles (carapace length <40 mm) of the European lobster, Homarus gammarus , exposed to elevated temperature and P CO 2 At 13°C, H. gammarus juveniles were able to fully compensate for acid-base disturbances caused by the exposure to elevated seawater P CO 2  at levels associated with ocean acidification and carbon dioxide capture and storage (CCS) leakage scenarios, via haemolymph [HCO 3 - ] regulation. However, metabolic rate remained constant and food consumption decreased under elevated P CO 2 , indicating reduced energy availability. Juveniles at 17°C showed no ability to actively compensate haemolymph pH, resulting in decreased haemolymph pH particularly under CCS conditions. Early benthic juvenile lobsters at 17°C were not able to increase energy intake to offset increased energy demand and therefore appear to be unable to respond to acid-base disturbances due to increased P CO 2   at elevated temperature. Analysis of haemolymph metabolites suggests that, even under control conditions, juveniles were energetically limited. They exhibited high haemolymph [l-lactate], indicating recourse to anaerobic metabolism. Low haemolymph [protein] was linked to minimal non-bicarbonate buffering and reduced oxygen transport capacity. We discuss these results in the context of potential impacts of ongoing ocean change and CCS leakage scenarios on the development of juvenile H. gammarus and future lobster populations and stocks., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Global warming appears to favour smaller-bodied organisms, but whether larger species are also more vulnerable to thermal extremes, as suggested for past mass-extinction events, is still an open question. Here, we tested whether interspecific differences in thermal tolerance (heat and cold) of ectotherm organisms are linked to differences in their body mass and genome size (as a proxy for cell size). Since the vulnerability of larger, aquatic taxa to warming has been attributed to the oxygen limitation hypothesis, we also assessed how body mass and genome size modulate thermal tolerance in species with contrasting breathing modes, habitats and life stages. A database with the upper (CTmax) and lower (CTmin) critical thermal limits and their methodological aspects was assembled comprising more than 500 species of ectotherms. Our results demonstrate that thermal tolerance in ectotherms is dependent on body mass and genome size and these relationships became especially evident in prolonged experimental trials where energy efficiency gains importance. During long-term trials, CTmax was impaired in larger-bodied water-breathers, consistent with a role for oxygen limitation. Variation in CTmin was mostly explained by the combined effects of body mass and genome size and it was enhanced in larger-celled, air-breathing species during long-term trials, consistent with a role for depolarization of cell membranes. Our results also highlight the importance of accounting for phylogeny and exposure duration. Especially when considering long-term trials, the observed effects on thermal limits are more in line with the warming-induced reduction in body mass observed during long-term rearing experiments. This article is part of the theme issue 'Physiological diversity, biodiversity patterns and global climate change: testing key hypotheses involving temperature and oxygen'.

We are starting to understand the relationship between metabolic rate responses and species' ability to respond to exposure to high p CO 2 . However, most of our knowledge has come from investigations of single species. The examination of metabolic responses of closely related species with differing distributions around natural elevated CO 2 areas may be useful to inform our understanding of their adaptive significance. Furthermore, little is known about the physiological responses of marine invertebrate juveniles to high p CO 2 , despite the fact they are known to be sensitive to other stressors, often acting as bottlenecks for future species success. We conducted an in situ transplant experiment using juveniles of isopods found living inside and around a high p CO 2 vent (Ischia, Italy): the CO 2 'tolerant' Dynamene bifida and 'sensitive' Cymodoce truncata and Dynamene torelliae . This allowed us to test for any generality of the hypothesis that p CO 2 sensitive marine invertebrates may be those that experience trade-offs between energy metabolism and cellular homoeostasis under high p CO 2 conditions. Both sensitive species were able to maintain their energy metabolism under high p CO 2 conditions, but in C. truncata this may occur at the expense of [carbonic anhydrase], confirming our hypothesis. By comparison, the tolerant D. bifida appeared metabolically well adapted to high p CO 2 , being able to upregulate ATP production without recourse to anaerobiosis. These isopods are important keystone species; however, given they differ in their metabolic responses to future p CO 2 , shifts in the structure of the marine ecosystems they inhabit may be expected under future ocean acidification conditions., Competing Interests: All authors declare they have no conflict of interest. Ethical approval All applicable international, national, and/or institutional guidelines for the care and use of animals were followed.

An organism's physiological processes form the link between its life-history traits and the prevailing environmental conditions, especially in species with complex life cycles. Understanding how these processes respond to changing environmental conditions, thereby affecting organismal development, is critical if we are to predict the biological implications of current and future global climate change. However, much of our knowledge is derived from adults or single developmental stages. Consequently, we investigated the metabolic rate, organic content, carapace mineralization, growth, and survival across each larval stage of the European lobster Homarus gammarus, reared under current and predicted future ocean warming and acidification scenarios. Larvae exhibited stage-specific changes in the temperature sensitivity of their metabolic rate. Elevated Pco2 increased C∶N ratios and interacted with elevated temperature to affect carapace mineralization. These changes were linked to concomitant changes in survivorship and growth, from which it was concluded that bottlenecks were evident during H. gammarus larval development in stages I and IV, the transition phases between the embryonic and pelagic larval stages and between the larval and megalopa stages, respectively. We therefore suggest that natural changes in optimum temperature during ontogeny will be key to larvae survival in a future warmer ocean. The interactions of these natural changes with elevated temperature and Pco2 significantly alter physiological condition and body size of the last larval stage before the transition from a planktonic to a benthic life style. Thus, living and growing in warm, hypercapnic waters could compromise larval lobster growth, development, and recruitment.

Predicting species vulnerability to global warming requires a comprehensive, mechanistic understanding of sublethal and lethal thermal tolerances. To date, however, most studies investigating species physiological responses to increasing temperature have focused on the underlying physiological traits of either acute or chronic tolerance in isolation. Here we propose an integrative, synthetic approach including the investigation of multiple physiological traits (metabolic performance and thermal tolerance), and their plasticity, to provide more accurate and balanced predictions on species and assemblage vulnerability to both acute and chronic effects of global warming. We applied this approach to more accurately elucidate relative species vulnerability to warming within an assemblage of six caridean prawns occurring in the same geographic, hence macroclimatic, region, but living in different thermal habitats. Prawns were exposed to four incubation temperatures (10, 15, 20 and 25 °C) for 7 days, their metabolic rates and upper thermal limits were measured, and plasticity was calculated according to the concept of Reaction Norms, as well as Q10 for metabolism. Compared to species occupying narrower/more stable thermal niches, species inhabiting broader/more variable thermal environments (including the invasive Palaemon macrodactylus) are likely to be less vulnerable to extreme acute thermal events as a result of their higher upper thermal limits. Nevertheless, they may be at greater risk from chronic exposure to warming due to the greater metabolic costs they incur. Indeed, a trade-off between acute and chronic tolerance was apparent in the assemblage investigated. However, the invasive species P. macrodactylus represents an exception to this pattern, showing elevated thermal limits and plasticity of these limits, as well as a high metabolic control. In general, integrating multiple proxies for species physiological acute and chronic responses to increasing temperature helps providing more accurate predictions on species vulnerability to warming., (© 2014 John Wiley & Sons Ltd.)

Ocean acidification poses a global threat to biodiversity, yet species might have the capacity to adapt through evolutionary change. Here we summarize tools available to determine species' capacity for evolutionary adaptation to future ocean change and review the progress made to date with respect to ocean acidification. We focus on two key approaches: measuring standing genetic variation within populations and experimental evolution. We highlight benefits and challenges of each approach and recommend future research directions for understanding the modulating role of evolution in a changing ocean., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Metabolic rate determines the physiological and life-history performances of ectotherms. Thus, the extent to which such rates are sensitive and plastic to environmental perturbation is central to an organism's ability to function in a changing environment. Little is known of long-term metabolic plasticity and potential for metabolic adaptation in marine ectotherms exposed to elevated pCO2. Consequently, we carried out a series of in situ transplant experiments using a number of tolerant and sensitive polychaete species living around a natural CO2 vent system. Here, we show that a marine metazoan (i.e. Platynereis dumerilii) was able to adapt to chronic and elevated levels of pCO2. The vent population of P. dumerilii was physiologically and genetically different from nearby populations that experience low pCO2, as well as smaller in body size. By contrast, different populations of Amphiglena mediterranea showed marked physiological plasticity indicating that adaptation or acclimatization are both viable strategies for the successful colonization of elevated pCO2 environments. In addition, sensitive species showed either a reduced or increased metabolism when exposed acutely to elevated pCO2. Our findings may help explain, from a metabolic perspective, the occurrence of past mass extinction, as well as shed light on alternative pathways of resilience in species facing ongoing ocean acidification.

In the future, marine organisms will face the challenge of coping with multiple environmental changes associated with increased levels of atmospheric Pco(2), such as ocean warming and acidification. To predict how organisms may or may not meet these challenges, an in-depth understanding of the physiological and biochemical mechanisms underpinning organismal responses to climate change is needed. Here, we investigate the effects of elevated Pco(2) and temperature on the whole-organism and cellular physiology of the periwinkle Littorina littorea. Metabolic rates (measured as respiration rates), adenylate energy nucleotide concentrations and indexes, and end-product metabolite concentrations were measured. Compared with values for control conditions, snails decreased their respiration rate by 31% in response to elevated Pco(2) and by 15% in response to a combination of increased Pco(2) and temperature. Decreased respiration rates were associated with metabolic reduction and an increase in end-product metabolites in acidified treatments, indicating an increased reliance on anaerobic metabolism. There was also an interactive effect of elevated Pco(2) and temperature on total adenylate nucleotides, which was apparently compensated for by the maintenance of adenylate energy charge via AMP deaminase activity. Our findings suggest that marine intertidal organisms are likely to exhibit complex physiological responses to future environmental drivers, with likely negative effects on growth, population dynamics, and, ultimately, ecosystem processes.