In situ decrease in rhodolith growth associated with Arctic climate change.

Rhodoliths built by crustose coralline algae (CCA) are ecosystem engineers of global importance. In the Arctic photic zone, their three-dimensional growth emulates the habitat complexity of coral reefs but with a far slower growth rate, growing at micrometers per year rather than millimeters. While climate change is known to exert various impacts on the CCA's calcite skeleton, including geochemical and structural alterations, field observations of net growth over decade-long timescales are lacking. Here, we use a temporally explicit model to show that rising ocean temperatures over nearly 100 years were associated with reduced rhodolith growth at different depths in the Arctic. Over the past 90 years, the median growth rate was 85 µm year-1 but each °C increase in summer seawater temperature decreased growth by a mean of 8.9 µm (95% confidence intervals = 1.32-16.60 µm °C-1, p < .05). The decrease was expressed for rhodolith occurrences in 11 and 27 m water depth but not at 46 m, also having the shortest time series (1991-2015). Although increasing temperatures can spur plant growth, we suggest anthropogenic climate change has either exceeded the population thermal optimum for these CCA, or synergistic effects of warming, ocean acidification, and/or increasing turbidity impair rhodolith growth. Rhodoliths built by calcitic CCA are important habitat providers worldwide, so decreased growth would lead to yet another facet of anthropogenic habitat loss.

Global warming generates predictable extinctions of warm- and cold-water marine benthic invertebrates via thermal habitat loss.

Anthropogenic global warming is redistributing marine life and may threaten tropical benthic invertebrates with several potential extinction mechanisms. The net impact of climate change on geographical extinction risk nevertheless remains uncertain. Evidence of widespread climate-driven extinctions and of potentially unidentified mechanisms exists in the fossil record. We quantify organism extinction risk across thermal habitats, estimated by paleoclimate reconstructions, over the past 300 million years. Extinction patterns at seven known events of rapid global warming (hyperthermals) differ significantly from typical patterns, resembling those driven by global geometry under simulated global warming. As isotherms move poleward with warming, the interaction between the geometry of the globe and the temperature-latitude relationship causes an uneven loss of thermal habitat and a bimodal latitudinal distribution of extinctions. Genera with thermal optima warmer than ~21°C show raised extinction odds, while extinction odds continually increase for genera with optima below ~11°C. Genera preferring intermediate temperatures generally have no additional extinction risk during hyperthermals, except under extreme conditions as the end-Permian mass extinction. Widespread present-day climate-driven range shifts indicate that occupancy loss is already underway. Given the most-likely projections of modern warming, our model, validated by seven past hyperthermal events, indicates that sustained warming has the potential to annihilate cold-water habitat and its endemic species completely within centuries.

Extensive spatial impacts of oyster reefs on an intertidal mudflat community via predator facilitation.

Habitat engineers make strong and far-reaching imprints on ecosystem processes. In intertidal mudflats, the dominant primary producer, microphytobenthos (MPB), often forms high biomass patches around oyster reefs. We evaluate multiple hypotheses linking MPB with oyster reefs, including oyster biodeposition, meiofaunal grazing, and abiotic factors, aiming to help predict effects of reef removal or proliferation. We quantify spatial patterns of an Atlantic mudflat community and its environment around two large Crassostrea reefs before experimentally sacrificing one reef via burning. MPB biomass was enriched surrounding living oyster reefs although infaunal biomass and individual sizes were low. Structural equation modelling best supported the hypothesis that crab predation intensity, which decayed with distance from the reefs, locally freed MPB from grazing. Our results suggest that Crassostrea reef expansion may enrich local MPB patches and redirect trophic energy flows away from mudflat infauna, with potential implications for the sustainability of local fisheries and bird conservation.

Increase in marine provinciality over the last 250 million years governed more by climate change than plate tectonics.

Amidst long-term fluctuations of the abiotic environment, the degree to which life organizes into distinct biogeographic provinces (provinciality) can reveal the fundamental drivers of global biodiversity. Our understanding of present-day biogeography implies that changes in the distribution of continents across climatic zones have predictable effects on habitat distribution, dispersal barriers and the evolution of provinciality. To assess marine provinciality through the Phanerozoic, here we (a) simulate provinces based on palaeogeographic reconstructions and global climate models and (b) contrast them with empirically derived provinces that we define using network analysis of fossil occurrences. Simulated and empirical patterns match reasonably well and consistently suggest a greater than 15% increase in provinciality since the Mesozoic era. Although both factors played a role, the simulations imply that the effect of the latitudinal temperature gradient has been twice as important in determining marine provinciality as continental configuration.

Morphological response accompanying size reduction of belemnites during an Early Jurassic hyperthermal event modulated by life history.

One of the most common responses of marine ectotherms to rapid warming is a reduction in body size, but the underlying reasons are unclear. Body size reductions have been documented alongside rapid warming events in the fossil record, such as across the Pliensbachian-Toarcian boundary (PToB) event (~ 183 Mya). As individuals grow, parallel changes in morphology can indicate details of their ecological response to environmental crises, such as changes in resource acquisition, which may anticipate future climate impacts. Here we show that the morphological growth of a marine predator belemnite species (extinct coleoid cephalopods) changed significantly over the PToB warming event. Increasing robustness at different ontogenetic stages likely results from indirect consequences of warming, like resource scarcity or hypercalcification, pointing toward varying ecological tolerances among species. The results of this study stress the importance of taking life history into account as well as phylogeny when studying impacts of environmental stressors on marine organisms.

Victims of ancient hyperthermal events herald the fates of marine clades and traits under global warming.

Organismic groups vary non-randomly in their vulnerability to extinction. However, it is unclear whether the same groups are consistently vulnerable, regardless of the dominant extinction drivers, or whether certain drivers have their own distinctive and predictable victims. Given the challenges presented by anthropogenic global warming, we focus on changes in extinction selectivity trends during ancient hyperthermal events: geologically rapid episodes of global warming. Focusing on the fossil record of the last 300 million years, we identify clades and traits of marine ectotherms that were more prone to extinction under the onset of six hyperthermal events than during other times. Hyperthermals enhanced the vulnerability of marine fauna that host photosymbionts, particularly zooxanthellate corals, the reef environments they provide, and genera with actively burrowing or swimming adult life-stages. The extinction risk of larger sized fauna also increased relative to non-hyperthermal times, while genera with a poorly buffered internal physiology did not become more vulnerable on average during hyperthermals. Hyperthermal-vulnerable clades include rhynchonelliform brachiopods and bony fish, whereas resistant clades include cartilaginous fish, and ostreid and venerid bivalves. These extinction responses in the geological past mirror modern responses of these groups to warming, including range-shift magnitudes, population losses, and experimental performance under climate-related stressors. Accordingly, extinction mechanisms distinctive to rapid global warming may be indicated, including sensitivity to warming-induced seawater deoxygenation. In anticipation of modern warming-driven marine extinctions, the trends illustrated in the fossil record offer an expedient preview.

The biogeographical imprint of mass extinctions.

Mass extinctions are defined by extinction rates significantly above background levels and have had substantial consequences for the evolution of life. Geographically selective extinctions, subsequent originations and species redistributions may have changed global biogeographical structure, but quantification of this change is lacking. In order to assess quantitatively the biogeographical impact of mass extinctions, we outline time-traceable bioregions for benthic marine species across the Phanerozoic using a compositional network. Mass extinction events are visually recognizable in the geographical depiction of bioregions. The end-Permian extinction stands out with a severe reduction of provinciality. Time series of biogeographical turnover represent a novel aspect of the analysis of mass extinctions, confirming concentration of changes in the geographical distribution of benthic marine life.

Living to the range limit: consumer isotopic variation increases with environmental stress.

BACKGROUND: Theoretically, each species' ecological niche is phylogenetically-determined and expressed spatially as the species' range. However, environmental stress gradients may directly or indirectly decrease individual performance, such that the precise process delimiting a species range may not be revealed simply by studying abundance patterns. In the intertidal habitat the vertical ranges of marine species may be constrained by their abilities to tolerate thermal and desiccation stress, which may act directly or indirectly, the latter by limiting the availability of preferred trophic resources. Therefore, we expected individuals at greater shore heights to show greater variation in diet alongside lower indices of physiological condition. METHODS: We sampled the grazing gastropod Echinolittorina peruviana from the desert coastline of northern Chile at three shore heights, across eighteen regionally-representative shores. Stable isotope values (δ13C and δ15N) were extracted from E. peruviana and its putative food resources to estimate Bayesian ellipse area, carbon and nitrogen ranges and diet. Individual physiological condition was tracked by muscle % C and % N. RESULTS: There was an increase in isotopic variation at high shore levels, where E. peruviana's preferred resource, tide-deposited particulate organic matter (POM), appeared to decrease in dietary contribution, and was expected to be less abundant. Both muscle % C and % N of individuals decreased with height on the shore. DISCUSSION: Individuals at higher stress levels appear to be less discriminating in diet, likely because of abiotic forcing, which decreases both consumer mobility and the availability of a preferred resource. Abiotic stress might be expected to increase trophic variation in other selective dietary generalist species. Where this coincides with a lower physiological condition may be a direct factor in setting their range limit.

Coastal Upwelling Drives Intertidal Assemblage Structure and Trophic Ecology.

Similar environmental driving forces can produce similarity among geographically distant ecosystems. Coastal oceanic upwelling, for example, has been associated with elevated biomass and abundance patterns of certain functional groups, e.g., corticated macroalgae. In the upwelling system of Northern Chile, we examined measures of intertidal macrobenthic composition, structure and trophic ecology across eighteen shores varying in their proximity to two coastal upwelling centres, in a hierarchical sampling design (spatial scales of >1 and >10 km). The influence of coastal upwelling on intertidal communities was confirmed by the stable isotope values (δ13C and δ15N) of consumers, including a dominant suspension feeder, grazers, and their putative resources of POM, epilithic biofilm, and macroalgae. We highlight the utility of muscle δ15N from the suspension feeding mussel, Perumytilus purpuratus, as a proxy for upwelling, supported by satellite data and previous studies. Where possible, we used corrections for broader-scale trends, spatial autocorrelation, ontogenetic dietary shifts and spatial baseline isotopic variation prior to analysis. Our results showed macroalgal assemblage composition, and benthic consumer assemblage structure, varied significantly with the intertidal influence of coastal upwelling, especially contrasting bays and coastal headlands. Coastal topography also separated differences in consumer resource use. This suggested that coastal upwelling, itself driven by coastline topography, influences intertidal communities by advecting nearshore phytoplankton populations offshore and cooling coastal water temperatures. We recommend the isotopic values of benthic organisms, specifically long-lived suspension feeders, as in situ alternatives to offshore measurements of upwelling influence.