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Abstract Thermal performance curves (TPCs) are important tools for predicting the sensitivity of populations to climate change. However, the interactive ways that temperature affects multiple life‐history components lead to different fitness outcomes. These interactions are poorly understood for modular animals, especially over the lifespan of individual colonies, which limits our capacity to connect physiological and demographic responses.The goal of this study was to assess and compare the relationships between temperature and different life‐history components in a modular animal to reveal the mechanisms underlying TPCs for fitness.We reared replicated clones of the marine bryozoanBugula neritinaacross a thermal gradient (16 values) ranging from 23 to 32°C, which reflected the upper thermal range of seasonal variation in the field. TPCs were constructed for survival (measured as zooids states within a colony), growth rate, development to reproductive maturity and reproductive capacity, which were measured over much of the realized lifespan expected under field conditions (~30 days).The effect of temperature was more acute on zooid states rather than whole‐colony survival, and increased temperature increased the frequency of polypide regression. Most colonies reached reproductive maturity up to ~30°C, but growth rate and reproduction decreased at temperatures beyond ~25°C. The decline in reproductive capacity over temperatures above ~25°C was then due to the decline in the production of zooids capable of brooding embryos and zooids transitioning to regressed states up until about 30°C and transitioning to dead state beyond that.Higher temperatures are often considered to affect reproduction by interfering with gametogenesis and post‐zygotic pathways, but in modular animals, changes in growth rate and module states could indirectly cause temperature sensitivity of reproduction. Our study has implications for the role of temperature in driving the sampled population's dynamics by setting the number of generations that occur during the time window when temperatures are conducive to reproduction. Our results also have implications for the generality and predictability of temperature on population persistence across unitary and modular animals. Read the freePlain Language Summaryfor this article on the Journal blog. 

Abstract Dispersal can evolve as an adaptation to escape competition with conspecifics or kin. Locations with a low density of conspecifics, however, may also lead to reduced opportunities for mating, especially in sessile marine invertebrates with proximity-dependent mating success. Since there are few experimental investigations, we performed a series of field experiments using an experimentally tractable species (the bryozoan Bugula neritina) to test the hypothesis that the density, spatial arrangement, and genetic relatedness of neighbours differentially affect survival, growth, reproduction, paternity, and sperm dispersal. We manipulated the density and relatedness of neighbours and found that increased density reduced survival but not growth rate, and that there was no effect of relatedness on survival, growth, or fecundity, in contrast to previous studies. We also manipulated the distances to the nearest neighbour and used genetic markers to assign paternity within known mother–offspring groups to estimate how proximity affects mating success. Distance to the nearest neighbour did not affect the number of settlers produced, the paternity share, or the degree of multiple paternity. Overall, larger than expected sperm dispersal led to high multiple paternity, regardless of the distance to the nearest neighbour. Our results have important implications for understanding selection on dispersal distance: in this system, there are few disadvantages to the limited larval dispersal that does occur and limited advantages for larvae to disperse further than a few 10s of metres. 

Abstract While the presence of morphologically cryptic species is increasingly recognized, we still lack a useful understanding of what causes and maintains co‐occurring cryptic species and its consequences for the ecology, evolution, and conservation of communities. We sampled 724Pocilloporacorals from five habitat zones (the fringing reef, back reef, and fore reef at 5, 10, and 20 m) at four sites around the island of Moorea, French Polynesia. Using validated genetic markers, we identified six sympatric species ofPocillopora, most of which cannot be reliably identified based on morphology:P. meandrina(42.9%),P. tuahiniensis(25.1%),P. verrucosa(12.2%),P. acuta(10.4%),P. grandis(7.73%), andP.cf.effusa(2.76%). For 423 colonies (58% of the genetically identified hosts), we also usedpsbA^ncror ITS2 markers to identify symbiont species (Symbiodiniaceae). The relative abundance ofPocilloporaspecies differed across habitats within the reef. Sister taxaP. verrucosaandP. tuahiniensishad similar niche breadths and hosted the same specialist symbiont species (mostlyCladocopium pacificum) but the former was more common in the back reef and the latter more common deeper on the fore reef. In contrast, sister taxaP. meandrinaandP. grandishad the highest niche breadths and overlaps and tended to host the same specialist symbiont species (mostlyC. latusorum).Pocillopora acutahad the narrowest niche breadth and hosted the generalist, and more thermally tolerant,Durusdinium gynnii. Overall, there was a positive correlation between reef habitat niche breadth and symbiont niche breadth—Pocilloporaspecies with a broader habitat niche also had a broader symbiont niche. Our results show how fine‐scale variation within reefs plays an important role in the generation and coexistence of cryptic species. The results also have important implications for how niche differences affect community resilience, and for the success of coral restoration practices, in ways not previously appreciated. 

ABSTRACT Warming global temperatures have consequences for biological rates. Feeding rates reflect the intake of energy that fuels survival, growth and reproduction. However, temperature can also affect food abundance and quality, as well as feeding behavior, which all affect feeding rate, making it challenging to understand the pathways by which temperature affects the intake of energy. Therefore, we experimentally assessed how clearance rate varied across a thermal gradient in a filter-feeding colonial marine invertebrate (the bryozoan Bugula neritina). We also assessed how temperature affects phytoplankton as a food source, and zooid states within a colony that affect energy budgets and feeding behavior. Clearance rate increased linearly from 18°C to 32°C, a temperature range that the population experiences most of the year. However, temperature increased algal cell size, and decreased the proportion of feeding zooids, suggesting indirect effects of temperature on clearance rates. Temperature increased polypide regression, possibly as a stress response because satiation occurred quicker, or because phytoplankton quality declined. Temperature had a greater effect on clearance rate per feeding zooid than it did per total zooids. Together, these results suggest that the effect of temperature on clearance rate at the colony level is not just the outcome of individual zooids feeding more in direct response to temperature but also emerges from temperature increasing polypide regression and the remaining zooids increasing their feeding rates in response. Our study highlights some of the challenges for understanding why temperature affects feeding rates, especially for understudied, yet ecologically important, marine colonial organisms. 

Understanding population dynamics is a long-standing objective of ecology, but the need for progress in this area has become urgent. For coral reefs, achieving this objective is impeded by a lack of information on settlement versus post-settlement events in determining recruitment and population size. Declines in coral abundance are often inferred to be associated with reduced densities of recruits, which could arise from mechanisms occurring at larval settlement, or throughout post-settlement stages. This study uses annual measurements from 2008 to 2021 of coral cover, the density of coral settlers (S), the density of small corals (SC), and environmental conditions, to evaluate the roles of settlement versus post-settlement events in determining rates of coral recruitment and changes in coral cover at Moorea, French Polynesia. Coral cover, S, SC, and the SC:S ratio (a proxy for post-settlement success), and environmental conditions, were used in generalized additive models (GAMs) to show that: (a) coral cover was more strongly related to SC and SC:S than S, and (b) SC:S was highest when preceded by cool seawater, low concentrations of Chlorophyll a, and low flow speeds, and S showed evidence of declining with elevated temperature. Together, these results suggest that changes in coral cover in Moorea are more strongly influenced by post-settlement events than settlement. The key to understanding coral community resilience may lie in elucidating the factors attenuating the bottleneck between settlers and small corals. 

Dispersal emerges as an outcome of organismal traits and external forcings. However, it remains unclear how the emergent dispersal kernel evolves as a by-product of selection on the underlying traits. This question is particularly compelling in coastal marine systems, where dispersal is tied to development and reproduction and where directional currents bias larval dispersal downstream, causing selection for retention. We modeled the dynamics of a metapopulation along a finite coastline using an integral projection model and adaptive dynamics to understand how asymmetric coastal currents influence the evolution of larval (pelagic larval duration) and adult (spawning frequency) life history traits, which indirectly shape the evolution of marine dispersal kernels. Selection induced by alongshore currents favors the release of larvae over multiple time periods, allowing long pelagic larval durations and long-distance dispersal to be maintained in marine life cycles in situations where they were previously predicted to be selected against. Two evolutionarily stable strategies emerged: one with a long pelagic larval duration and many spawning events, resulting in a dispersal kernel with a larger mean and variance, and another with a short pelagic larval duration and few spawning events, resulting in a dispersal kernel with a smaller mean and variance. Our theory shows how coastal ocean flows are important agents of selection that can generate multiple, often co-occurring evolutionary outcomes for marine life history traits that affect dispersal. 

Pocillopora tuahiniensis sp. nov. is described based on mitochondrial and nuclear genomic data, algal symbiont genetic data, geographic isolation, and its distribution pattern within reefs that is distinct from other sympatric Pocillopora species (Johnston et al. 2022a, b). Mitochondrial and nuclear genomic data reveal that P. tuahiniensis sp. nov. is a unique species, sister to P. verrucosa, and in a clade different from that of P. meandrina (Johnston et al. 2022a). However, the gross in situ colony appearance of P. tuahiniensis sp. nov. cannot easily be differentiated from that of P. verrucosa or P. meandrina at Mo’orea. By sequencing the mtORF region, P. tuahiniensis sp. nov. can be easily distinguished from other Pocillopora species. Pocillopora tuahiniensis sp. nov. has so far been sampled in French Polynesia, Ducie Island, and Rapa Nui (Armstrong et al. 2023; Edmunds et al. 2016; Forsman et al. 2013; Gélin et al. 2017; Mayfield et al. 2015; Oury et al. 2021; Voolstra et al. 2023). On the fore reefs of Mo’orea, P. tuahiniensis sp. nov. is very abundant ≥10 m and is one of the most common Pocillopora species at these depths (Johnston et al. 2022b). It can also be found at a much lower abundance at shallow depths on the fore reef and back reef lagoon. The holotype is deposited at the Smithsonian Institution as USNM-SI 1522390 and the mtORF Genbank accession number is OP418359.  

For nearly 50 years, analyses of coral physiology have used small coral fragments (nubbins) to make inferences about larger colonies. However, scaling in corals shows that linear extrapolations from nubbins to whole colonies can be misleading, because polyps in nubbins are divorced of their morphologically complex and physiologically integrated corallum. We tested for the effects of integration among branches in determining size-dependent calcification of the coral Pocillopora spp. under elevated P CO 2 . Area-normalized net calcification was compared between branches (nubbins), aggregates of nubbins (complex morphologies without integration) and whole colonies (physiologically integrated) at 400 versus approximately 1000 µatm P CO 2 . Net calcification was unaffected by P CO 2 , but differed among colony types. Single nubbins grew faster than whole colonies, but when aggregated, nubbins changed calcification to match whole colonies even though they lacked integration among branches. Corallum morphology causes the phenotype of branching corals to differ from the summation of their branches. 

Abstract The severity of marine heatwaves (MHWs) that are increasingly impacting ocean ecosystems, including vulnerable coral reefs, has primarily been assessed using remotely sensed sea-surface temperatures (SSTs), without information relevant to heating across ecosystem depths. Here, using a rare combination of SST, high-resolution in-situ temperatures, and sea level anomalies observed over 15 years near Moorea, French Polynesia, we document subsurface MHWs that have been paradoxical in comparison to SST metrics and associated with unexpected coral bleaching across depths. Variations in the depth range and severity of MHWs was driven by mesoscale (10s to 100s of km) eddies that altered sea levels and thermocline depths and decreased (2007, 2017 and 2019) or increased (2012, 2015, 2016) internal-wave cooling. Pronounced eddy-induced reductions in internal waves during early 2019 contributed to a prolonged subsurface MHW and unexpectedly severe coral bleaching, with subsequent mortality offsetting almost a decade of coral recovery. Variability in mesoscale eddy fields, and thus thermocline depths, is expected to increase with climate change, which, along with strengthening and deepening stratification, could increase the occurrence of subsurface MHWs over ecosystems historically insulated from surface ocean heating by the cooling effects of internal waves. 

Abstract Dispersal has far‐reaching implications for individuals, populations, and communities, especially in sessile organisms. Escaping competition with conspecifics and with kin are theorized to be key factors leading to dispersal as an adaptation. However, manipulative approaches in systems in which adults are sessile but offspring have behaviors is required for a more complete understanding of how competition affects dispersal. Here, we integrate a series of experiments to study how dispersal affects the density and relatedness of neighbors, and how the density and relatedness of neighbors in turn affects fitness. In a marine bryozoan, we empirically estimated dispersal kernels and found that most larvae settled within ~1 m of the maternal colony, although some could potentially travel at least 10s of meters. Larvae neither actively preferred or avoided conspecifics or kin at settlement. We experimentally determined the effects of spreading sibling larvae by manipulating the density and relatedness of settlers and measuring components of fitness in the field. We found that settler density reduced maternal fitness when settler neighbors were siblings compared with when neighbors were unrelated or absent. Genetic markers also identified very few half sibs (and no full sibs) in adults from the natural population, and rarely close enough to directly interact. In this system, dispersal occurs over short distances (meters) yet, in contrast with expectations, there appears to be limited kinship between adult neighbors. Our results suggest that the limited dispersal increases early offspring mortality when siblings settle next to each other, rather than next to unrelated conspecifics, potentially reducing kinship in adult populations. High offspring production and multiple paternity could further dilute kinship at settlement and reduce selection for dispersal beyond the scale of 10s of meters.