Vertical Distribution of Rocky Intertidal Organisms Shifts With Sea-Level Variability on the Northeast Pacific Coast.

Disentangling the effects of cyclical variability in environmental forcing and long-term climate change on natural communities is a major challenge for ecologists, managers, and policy makers across ecosystems. Here we examined whether the vertical distribution of rocky intertidal taxa has shifted with sea-level variability occurring at multiple temporal scales and/or long-term anthropogenic sea-level rise (SLR). Because of the distinct zonation characteristic of intertidal communities, any shift in tidal dynamics or average sea level is expected to have large impacts on community structure and function. We found that across the Northeast Pacific Coast (NPC), sea level exhibits cyclical seasonal variability, tidal amplitude exhibits ecologically significant variability coherent with the 18.6-year periodicity of lunar declination, and long-term sea-level rise is occurring. Intertidal taxa largely do not exhibit significant vertical distribution shifts coherent with short-term (monthly to annual) sea-level variability but do exhibit taxa-specific vertical distribution shifts coherent with cyclical changes in lunar declination and long-term SLR at decadal timescales. Finally, our results show that responses to cyclical celestial mechanics and SLR vary among taxa, primarily according to their vertical distribution. Long-term SLR is occurring on ecologically relevant scales, but the confounding effects of cyclical celestial mechanics make interpreting shifts in zonation or community structure challenging. Such cyclical dynamics alternatingly amplify and dampen long-term SLR impacts and may modify the impacts of other global change related stressors, such as extreme heat waves and swell events, on intertidal organisms living at the edge of their physiological tolerances. As a result, intertidal communities will likely experience cyclical periods of environmental stress and concomitant nonlinear shifts in structure and function as long-term climate change continues. Our results demonstrate that consistent, large-scale monitoring of marine ecosystems is critical for understanding natural variability in communities and documenting long-term change.

Un desafío crucial para ecólogos, gestores y responsables políticos en todos los ecosistemas es entender los efectos de la variabilidad cíclica de los forzamientos ambientales y el cambio climático a largo plazo en las comunidades naturales. Aquí examinamos si la distribución vertical de los taxones del intermareal rocoso ha cambiado con la variabilidad del nivel del mar que ocurre en múltiples escalas temporales o con el aumento del nivel del mar antropogénico a largo plazo (ANM). Se espera que cualquier cambio en la dinámica de las mareas o el nivel medio del mar tenga grandes impactos en la estructura y función de la comunidad debido a la distintiva zonación característica de las comunidades intermareales. Encontramos que a lo largo de la costa del Pacífico Noreste, el nivel del mar exhibe variabilidad estacional cíclica, que la amplitud de las mareas exhibe una variación ecológicamente significativa coherente con la variación en la declinación lunar con una periodicidad de 18.6 años, y que el aumento del nivel del mar a largo plazo está ocurriendo. Los taxones del intermareal no exhiben cambios significativos en su distribución vertical asociado a la variabilidad del nivel del mar a corto plazo (mensual a anual); sin embargo, muestran cambios específicos en la distribución vertical, variando para cada taxón en respuesta a los cambios cíclicos en la declinación lunar y al ANM a largo plazo en escala de décadas. Finalmente, nuestros resultados demuestran que las respuestas a las mecánicas celestes cíclicas y al ANM varían según los taxones, principalmente de acuerdo con su distribución vertical. El ANM a largo plazo está ocurriendo en escalas ecológicamente relevantes, pero los efectos confusos de las mecánicas celestes cíclicas hacen que interpretar los cambios en la zonación o la estructura de la comunidad sea un desafío. Tales dinámicas cíclicas alternadas amplifican y atenúan los impactos del ANM a largo plazo y pueden modificar los impactos de otros factores estresantes relacionados con el cambio global, como las olas de calor extremas y los eventos de oleaje, en los organismos intermareales que viven al límite de su tolerancia fisiológica. Como resultado, mientras continúe el cambio climático a largo plazo, las comunidades del intermareal probablemente experimentarán periodos cíclicos de estrés ambiental y cambios no lineales concurrentes de su estructura y función. Nuestros resultados demuestran que el monitoreo consistente y a gran escala de los ecosistemas marinos es fundamental para entender la variabilidad natural en las comunidades y documentar cambios a largo plazo.

A marine protected area network does not confer community structure resilience to a marine heatwave across coastal ecosystems.

Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014-2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no-take state marine reserves, and 76 partial-take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no-take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat-wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem-wide consequences resulting from acute climate-driven perturbations, and the potential role of regulatory protection in mitigating community structure changes.

Latitudinal variation in long-term stability of North American rocky intertidal communities.

Although long-term ecological stability is often discussed as a community attribute, it is typically investigated at the species level (e.g. density, biomass), or as a univariate metric (e.g. species diversity). To provide a more comprehensive assessment of long-term community stability, we used a multivariate similarity approach that included all species and their relative abundances. We used data from 74 sites sampled annually from 2006 to 2017 to examine broad temporal and spatial patterns of change within rocky intertidal communities along the west coast of North America. We explored relationships between community change (inverse of stability) and the following potential drivers of change/stability: (a) marine heatwave events; (b) three attributes of biodiversity: richness, diversity and evenness and (c) presence of the mussel, Mytilus californianus, a dominant space holder and foundation species in this system. At a broad scale, we found an inverse relationship between community stability and elevated water temperatures. In addition, we found substantial differences in stability among regions, with lower stability in the south, which may provide a glimpse into the patterns expected with a changing climate. At the site level, community stability was linked to high species richness and, perhaps counterintuitively, to low evenness, which could be a consequence of the dominance of mussels in this system. Synthesis. Assessments of long-term stability at the whole-community level are rarely done but are key to a comprehensive understanding of the impacts of climate change. In communities structured around a spatially dominant species, long-term stability can be linked to the stability of this 'foundation species', as well as to traditional predictors, such as species richness.

Cryptic diversity, geographical endemism and allopolyploidy in NE Pacific seaweeds.

BACKGROUND: Molecular markers are revealing a much more diverse and evolutionarily complex picture of marine biodiversity than previously anticipated. Cryptic and/or endemic marine species are continually being found throughout the world oceans, predominantly in inconspicuous tropical groups but also in larger, canopy-forming taxa from well studied temperate regions. Interspecific hybridization has also been found to be prevalent in many marine groups, for instance within dense congeneric assemblages, with introgressive gene-flow being the most common outcome. Here, using a congeneric phylogeographic approach, we investigated two monotypic and geographically complementary sister genera of north-east Pacific intertidal seaweeds (Hesperophycus and Pelvetiopsis), for which preliminary molecular tests revealed unexpected conflicts consistent with unrecognized cryptic diversity and hybridization. RESULTS: The three recovered mtDNA clades did not match a priori species delimitations. H. californicus was congruent, whereas widespread P. limitata encompassed two additional narrow-endemic species from California - P. arborescens (here genetically confirmed) and P. hybrida sp. nov. The congruence between the genotypic clusters and the mtDNA clades was absolute. Fixed heterozygosity was apparent in a high proportion of loci in P. limitata and P. hybrida, with genetic analyses showing that the latter was composed of both H. californicus and P. arborescens genomes. All four inferred species could be distinguished based on their general morphology. CONCLUSIONS: This study confirmed additional diversity and reticulation within NE Pacific Hesperophycus/Pelvetiopsis, including the validity of the much endangered, modern climatic relict P. arborescens, and the identification of a new, stable allopolyploid species (P. hybrida) with clearly discernable ancestry (♀ H. californicus x ♂ P. arborescens), morphology, and geographical distribution. Allopolyploid speciation is otherwise completely unknown in brown seaweeds, and its unique occurrence within this genus (P. limitata possibly representing a second example) remains enigmatic. The taxonomic separation of Hesperophycus and Pelvetiopsis is not supported and the genera should be synonymized; we retain only the latter. The transitional coastline between Point Conception and Monterey Bay represented a diversity hotspot for the genus and the likely sites of extraordinary evolutionary events of allopolyploid speciation at sympatric range contact zones. This study pinpoints how much diversity (and evolutionary processes) potentially remains undiscovered even on a conspicuous seaweed genus from the well-studied Californian intertidal shores let alone in other, less studied marine groups and regions/depths.

Fluctuations in population fecundity drive variation in demographic connectivity and metapopulation dynamics.

Demographic connectivity is vital to sustaining metapopulations yet often changes dramatically through time due to variation in the production and dispersal of offspring. However, the relative importance of variation in fecundity and dispersal in determining the connectivity and dynamics of metapopulations is poorly understood due to the paucity of comprehensive spatio-temporal data on these processes for most species. We quantified connectivity in metapopulations of a marine foundation species (giant kelp Macrocystis pyrifera) across 11 years and approximately 900 km of coastline by estimating population fecundity with satellite imagery and propagule dispersal using a high-resolution ocean circulation model. By varying the temporal complexity of different connectivity measures and comparing their ability to explain observed extinction-colonization dynamics, we discovered that fluctuations in population fecundity, rather than fluctuations in dispersal, are the dominant driver of variation in connectivity and contribute substantially to metapopulation recovery and persistence. Thus, for species with high variability in reproductive output and modest variability in dispersal (most plants, many animals), connectivity measures ignoring fluctuations in fecundity may overestimate connectivity and likelihoods of persistence, limiting their value for understanding and conserving metapopulations. However, we demonstrate how connectivity measures can be simplified while retaining utility, validating a practical solution for data-limited systems.

Interacting environmental mosaics drive geographic variation in mussel performance and predation vulnerability.

Although theory suggests geographic variation in species' performance is determined by multiple niche parameters, little consideration has been given to the spatial structure of interacting stressors that may shape local and regional vulnerability to global change. Here, we use spatially explicit mosaics of carbonate chemistry, food availability and temperature spanning 1280 km of coastline to test whether persistent, overlapping environmental mosaics mediate the growth and predation vulnerability of a critical foundation species, the mussel Mytilus californianus. We find growth was highest and predation vulnerability was lowest in dynamic environments with frequent exposure to low pH seawater and consistent food. In contrast, growth was lowest and predation vulnerability highest when exposure to low pH seawater was decoupled from high food availability, or in exceptionally warm locations. These results illustrate how interactions among multiple drivers can cause unexpected, yet persistent geographic mosaics of species performance, interactions and vulnerability to environmental change.

Seascape drivers of Macrocystis pyrifera population genetic structure in the northeast Pacific.

At small spatial and temporal scales, genetic differentiation is largely controlled by constraints on gene flow, while genetic diversity across a species' distribution is shaped on longer temporal and spatial scales. We assess the hypothesis that oceanographic transport and other seascape features explain different scales of genetic structure of giant kelp, Macrocystis pyrifera. We followed a hierarchical approach to perform a microsatellite-based analysis of genetic differentiation in Macrocystis across its distribution in the northeast Pacific. We used seascape genetic approaches to identify large-scale biogeographic population clusters and investigate whether they could be explained by oceanographic transport and other environmental drivers. We then modelled population genetic differentiation within clusters as a function of oceanographic transport and other environmental factors. Five geographic clusters were identified: Alaska/Canada, central California, continental Santa Barbara, California Channel Islands and mainland southern California/Baja California peninsula. The strongest break occurred between central and southern California, with mainland Santa Barbara sites forming a transition zone between the two. Breaks between clusters corresponded approximately to previously identified biogeographic breaks, but were not solely explained by oceanographic transport. An isolation-by-environment (IBE) pattern was observed where the northern and southern Channel Islands clustered together, but not with closer mainland sites, despite the greater distance between them. The strongest environmental association with this IBE pattern was observed with light extinction coefficient, which extends suitable habitat to deeper areas. Within clusters, we found support for previous results showing that oceanographic connectivity plays an important role in the population genetic structure of Macrocystis in the Northern hemisphere.

Connectivity structures local population dynamics: a long-term empirical test in a large metapopulation system.

Ecological theory predicts that demographic connectivity structures the dynamics of local populations within metapopulation systems, but empirical support has been constrained by major limitations in data and methodology. We tested this prediction for giant kelp Macrocystis pyrifera, a key habitat-forming species in temperate coastal ecosystems worldwide, in southern California, USA. We combined a long-term (22 years), large-scale (~500 km coastline), high-resolution census of abundance with novel patch delineation methods and an innovative connectivity measure incorporating oceanographic transport and source fecundity. Connectivity strongly predicted local dynamics (well-connected patches had lower probabilities of extinction and higher probabilities of colonization, leading to greater likelihoods of occupancy) but this relationship was mediated by patch size. Moreover, the relationship between connectivity and local population dynamics varied over time, possibly due to temporal variation in oceanographic transport processes. Surprisingly, connectivity had a smaller influence on colonization relative to extinction, possibly because local ecological factors differ greatly between extinct and extant patches. Our results provide the first comprehensive evidence that southern California giant kelp populations function as a metapopulation system, challenging the view that populations of this important foundation species are governed exclusively by self-replenishment.

Patch definition in metapopulation analysis: a graph theory approach to solve the mega-patch problem.

The manner in which patches are delineated in spatially realistic metapopulation models will influence the size, connectivity, and extinction and recolonization dynamics of those patches. Most commonly used patch-definition methods focus on identifying discrete, contiguous patches of habitat from a single temporal observation of species occurrence or from a model of habitat suitability. However, these approaches are not suitable for many metapopulation systems where entire patches may not be fully colonized at a given time. For these metapopulation systems, a single large patch of habitat may actually support multiple, interacting subpopulations. The interactions among these subpopulations will be ignored if the patch is treated as a single unit, a situation we term the "mega-patch problem." Mega-patches are characterized by variable intra-patch synchrony, artificially low inter-patch connectivity, and low extinction rates. One way to detect this problem is by using time series data to calculate demographic synchrony within mega-patches. We present a framework for identifying subpopulations in mega-patches using a combination of spatial autocorrelation and graph theory analyses. We apply our approach to southern California giant kelp (Macrocystis pyrifera) forests using a new, long-term (27 years), satellite-based data set of giant kelp canopy biomass. We define metapopulation patches using our method as well as several other commonly used patch delineation methodologies and examine the colonization and extinction dynamics of the metapopulation under each approach. We find that the relationships between patch characteristics such as area and connectivity and the demographic processes of colonizations and extinctions vary among the different patch-definition methods. Our spatial-analysis/graph-theoretic framework produces results that match theoretical expectations better than the other methods. This approach can be used to identify subpopulations in metapopulations where the distributions of organisms do not always reflect the distribution of suitable habitat.

Looking into the black box: simulating the role of self-fertilization and mortality in the genetic structure of Macrocystis pyrifera.

Patterns of spatial genetic structure (SGS), typically estimated by genotyping adults, integrate migration over multiple generations and measure the effective gene flow of populations. SGS results can be compared with direct ecological studies of dispersal or mating system to gain additional insights. When mismatches occur, simulations can be used to illuminate the causes of these mismatches. Here, we report a SGS and simulation-based study of self-fertilization in Macrocystis pyrifera, the giant kelp. We found that SGS is weaker than expected in M. pyrifera and used computer simulations to identify selfing and early mortality rates for which the individual heterozygosity distribution fits that of the observed data. Only one (of three) population showed both elevated kinship in the smallest distance class and a significant negative slope between kinship and geographical distance. All simulations had poor fit to the observed data unless mortality due to inbreeding depression was imposed. This mortality could only be imposed for selfing, as these were the only simulations to show an excess of homozygous individuals relative to the observed data. Thus, the expected data consistently achieved nonsignificant differences from the observed data only under models of selfing with mortality, with best fits between 32% and 42% selfing. Inbreeding depression ranged from 0.70 to 0.73. The results suggest that density-dependent mortality of early life stages is a significant force in structuring Macrocystis populations, with few highly homozygous individuals surviving. The success of these results should help to validate simulation approaches even in data-poor systems, as a means to estimate otherwise difficult-to-measure life cycle parameters.

Isolation by oceanographic distance explains genetic structure for Macrocystis pyrifera in the Santa Barbara Channel.

Ocean currents are expected to be the predominant environmental factor influencing the dispersal of planktonic larvae or spores; yet, their characterization as predictors of marine connectivity has been hindered by a lack of understanding of how best to use oceanographic data. We used a high-resolution oceanographic model output and Lagrangian particle simulations to derive oceanographic distances (hereafter called transport times) between sites studied for Macrocystis pyrifera genetic differentiation. We build upon the classical isolation-by-distance regression model by asking how much additional variability in genetic differentiation is explained when adding transport time as predictor. We explored the extent to which gene flow is dependent upon seasonal changes in ocean circulation. Because oceanographic transport between two sites is inherently asymmetric, we also compare the explanatory power of models using the minimum or the mean transport times. Finally, we compare the direction of connectivity as estimated by the oceanographic model and genetic assignment tests. We show that the minimum transport time had higher explanatory power than the mean transport time, revealing the importance of considering asymmetry in ocean currents when modelling gene flow. Genetic assignment tests were much less effective in determining asymmetry in gene flow. Summer-derived transport times, in particular for the month of June, which had the strongest current speed, greatest asymmetry and highest spore production, resulted in the best-fit model explaining twice the variability in genetic differentiation relative to models that use geographic distance or habitat continuity. The best overall model also included habitat continuity and explained 65% of the variation in genetic differentiation among sites.

Estimating blue carbon sequestration under coastal management scenarios.

Restoring and protecting "blue carbon" ecosystems - mangrove forests, tidal marshes, and seagrass meadows - are actions considered for increasing global carbon sequestration. To improve understanding of which management actions produce the greatest gains in sequestration, we used a spatially explicit model to compare carbon sequestration and its economic value over a broad spatial scale (2500 km of coastline in southeastern Australia) for four management scenarios: (1) Managed Retreat, (2) Managed Retreat Plus Levee Removal, (3) Erosion of High Risk Areas, (4) Erosion of Moderate to High Risk Areas. We found that carbon sequestration from avoiding erosion-related emissions (abatement) would far exceed sequestration from coastal restoration. If erosion were limited only to the areas with highest erosion risk, sequestration in the non-eroded area exceeded emissions by 4.2 million Mg CO2 by 2100. However, losing blue carbon ecosystems in both moderate and high erosion risk areas would result in net emissions of 23.0 million Mg CO2 by 2100. The removal of levees combined with managed retreat was the strategy that sequestered the most carbon. Across all time points, removal of levees increased sequestration by only an additional 1 to 3% compared to managed retreat alone. Compared to the baseline erosion scenario, the managed retreat scenario increased sequestration by 7.40 million Mg CO2 by 2030, 8.69 million Mg CO2 by 2050, and 16.6 million Mg CO2 by 2100. Associated economic value followed the same patterns, with large potential value loss from erosion greater than potential gains from conserving or restoring ecosystems. This study quantifies the potential benefits of managed retreat and preventing erosion in existing blue carbon ecosystems to help meet climate change mitigation goals by reducing carbon emissions.

Habitat continuity and geographic distance predict population genetic differentiation in giant kelp.

Isolation by distance (IBD) models are widely used to predict levels of genetic connectivity as a function of Euclidean distance, and although recent studies have used GIS-landscape ecological approaches to improve the predictability of spatial genetic structure, few if any have addressed the effect of habitat continuity on gene flow. Landscape effects on genetic connectivity are even less understood in marine populations, where habitat mapping is particularly challenging. In this study, we model spatial genetic structure of a habitat-structuring species, the giant kelp Macrocystis pyrifera, using highly variable microsatellite markers. GIS mapping was used to characterize habitat continuity and distance between sampling sites along the mainland coast of the Santa Barbara Channel, and their roles as predictors of genetic differentiation were evaluated. Mean dispersal distance (sigma) and effective population size (Ne) were estimated by comparing our IBD slope with those from simulations incorporating habitat continuity and spore dispersal characteristics of the study area. We found an allelic richness of 7-50 alleles/locus, which to our knowledge is the highest reported for macroalgae. The best regression model relating genetic distance to habitat variables included both geographic distance and habitat continuity, which were respectively, positively and negatively related to genetic distance. Our results provide strong support for a dependence of gene flow on both distance and habitat continuity and elucidate the combination of Ne and a that explained genetic differentiation.

Biogeography of ocean acidification: Differential field performance of transplanted mussels to upwelling-driven variation in carbonate chemistry.

Ocean acidification (OA) represents a serious challenge to marine ecosystems. Laboratory studies addressing OA indicate broadly negative effects for marine organisms, particularly those relying on calcification processes. Growing evidence also suggests OA combined with other environmental stressors may be even more deleterious. Scaling these laboratory studies to ecological performance in the field, where environmental heterogeneity may mediate responses, is a critical next step toward understanding OA impacts on natural communities. We leveraged an upwelling-driven pH mosaic along the California Current System to deconstruct the relative influences of pH, ocean temperature, and food availability on seasonal growth, condition and shell thickness of the ecologically dominant intertidal mussel Mytilus californianus. In 2011 and 2012, ecological performance of adult mussels from local and commonly sourced populations was measured at 8 rocky intertidal sites between central Oregon and southern California. Sites coincided with a large-scale network of intertidal pH sensors, allowing comparisons among pH and other environmental stressors. Adult California mussel growth and size varied latitudinally among sites and inter-annually, and mean shell thickness index and shell weight growth were reduced with low pH. Surprisingly, shell length growth and the ratio of tissue to shell weight were enhanced, not diminished as expected, by low pH. In contrast, and as expected, shell weight growth and shell thickness were both diminished by low pH, consistent with the idea that OA exposure can compromise shell-dependent defenses against predators or wave forces. We also found that adult mussel shell weight growth and relative tissue mass were negatively associated with increased pH variability. Including local pH conditions with previously documented influences of ocean temperature, food availability, aerial exposure, and origin site enhanced the explanatory power of models describing observed performance differences. Responses of local mussel populations differed from those of a common source population suggesting mussel performance partially depended on genetic or persistent phenotypic differences. In light of prior research showing deleterious effects of low pH on larval mussels, our results suggest a life history transition leading to greater resilience in at least some performance metrics to ocean acidification by adult California mussels. Our data also demonstrate "hot" (more extreme) and "cold" (less extreme) spots in both mussel responses and environmental conditions, a pattern that may enable mitigation approaches in response to future changes in climate.

Evidence That Microorganisms at the Animal-Water Interface Drive Sea Star Wasting Disease.

Sea star wasting (SSW) disease describes a condition affecting asteroids that resulted in significant Northeastern Pacific population decline following a mass mortality event in 2013. The etiology of SSW is unresolved. We hypothesized that SSW is a sequela of microbial organic matter remineralization near respiratory surfaces, one consequence of which may be limited O2 availability at the animal-water interface. Microbial assemblages inhabiting tissues and at the asteroid-water interface bore signatures of copiotroph proliferation before SSW onset, followed by the appearance of putatively facultative and strictly anaerobic taxa at the time of lesion genesis and as animals died. SSW lesions were induced in Pisaster ochraceus by enrichment with a variety of organic matter (OM) sources. These results together illustrate that depleted O2 conditions at the animal-water interface may be established by heterotrophic microbial activity in response to organic matter loading. SSW was also induced by modestly (∼39%) depleted O2 conditions in aquaria, suggesting that small perturbations in dissolved O2 may exacerbate the condition. SSW susceptibility between species was significantly and positively correlated with surface rugosity, a key determinant of diffusive boundary layer thickness. Tissues of SSW-affected individuals collected in 2013-2014 bore δ15N signatures reflecting anaerobic processes, which suggests that this phenomenon may have affected asteroids during mass mortality at the time. The impacts of enhanced microbial activity and subsequent O2 diffusion limitation may be more pronounced under higher temperatures due to lower O2 solubility, in more rugose asteroid species due to restricted hydrodynamic flow, and in larger specimens due to their lower surface area to volume ratios which affects diffusive respiratory potential.

Increased evolutionary rates and conserved transcriptional response following allopolyploidization in brown algae.

Genome mergers between independently evolving lineages, via allopolyploidy, can potentially lead to instantaneous sympatric speciation. However, little is known about the consequences of allopolyploidy and the resultant "genome shock" on genome evolution and expression beyond the plant and fungal branches of the Tree of Life. The aim of this study was to compare substitution rates and gene expression patterns in two allopolyploid brown algae (Phaeophyceae and Heterokonta) and their progenitors in the genus Pelvetiopsis N. L. Gardner in the north-east Pacific, and to date their relationships. We used RNA-seq data, all potential orthologues, and putative single-copy loci for phylogenomic, divergence, and gene expression analyses. The multispecies coalescent placed the origin of allopolyploids in the late Pleistocene (0.35-0.05 Ma). Homoeologues displayed increased nonsynonymous divergence compared with parental orthologues, consistent with relaxed selective constraint following allopolyploidization, including for genes with no evidence of pseudogenization or neofunctionalization. Patterns of homoeologue-orthologue expression conservation and expression level dominance were largely shared with both natural plant and fungal allopolyploids. Our results provide further support for common cross-Kingdom patterns of allopolyploid genome evolution and transcriptional responses, here in the evolutionarily distinct marine heterokont brown algae.

Reduction and recovery of keystone predation pressure after disease-related mass mortality.

Disturbances such as disease can reshape communities through interruption of ecological interactions. Changes to population demographics alter how effectively a species performs its ecological role. While a population may recover in density, this may not translate to recovery of ecological function. In 2013, a sea star wasting syndrome outbreak caused mass mortality of the keystone predator Pisaster ochraceus on the North American Pacific coast. We analyzed sea star counts, biomass, size distributions, and recruitment from long-term intertidal monitoring sites from San Diego to Alaska to assess regional trends in sea star recovery following the outbreak. Recruitment, an indicator of population recovery, has been spatially patchy and varied within and among regions of the coast. Despite sea star counts approaching predisease numbers, sea star biomass, a measure of predation potential on the mussel Mytilus californianus, has remained low. This indicates that post-outbreak populations have not regained their full predation pressure. The regional variability in percent of recovering sites suggested differences in factors promoting sea star recovery between regions but did not show consistent patterns in postoutbreak recruitment on a coast-wide scale. These results shape predictions of where changes in community composition are likely to occur in years following the disease outbreak and provide insight into how populations of keystone species resume their ecological roles following mortality-inducing disturbances.

Large-scale impacts of sea star wasting disease (SSWD) on intertidal sea stars and implications for recovery.

Disease outbreaks can have substantial impacts on wild populations, but the often patchy or anecdotal evidence of these impacts impedes our ability to understand outbreak dynamics. Recently however, a severe disease outbreak occurred in a group of very well-studied organisms-sea stars along the west coast of North America. We analyzed nearly two decades of data from a coordinated monitoring effort at 88 sites ranging from southern British Columbia to San Diego, California along with 2 sites near Sitka, Alaska to better understand the effects of sea star wasting disease (SSWD) on the keystone intertidal predator, Pisaster ochraceus. Quantitative surveys revealed unprecedented declines of P. ochraceus in 2014 and 2015 across nearly the entire geographic range of the species. The intensity of the impact of SSWD was not uniform across the affected area, with proportionally greater population declines in the southern regions relative to the north. The degree of population decline was unrelated to pre-outbreak P. ochraceus density, although these factors have been linked in other well-documented disease events. While elevated seawater temperatures were not broadly linked to the initial emergence of SSWD, anomalously high seawater temperatures in 2014 and 2015 might have exacerbated the disease's impact. Both before and after the onset of the SSWD outbreak, we documented higher recruitment of P. ochraceus in the north than in the south, and while some juveniles are surviving (as evidenced by transition of recruitment pulses to larger size classes), post-SSWD survivorship is lower than during pre-SSWD periods. In hindsight, our data suggest that the SSWD event defied prediction based on two factors found to be important in other marine disease events, sea water temperature and population density, and illustrate the importance of surveillance of natural populations as one element of an integrated approach to marine disease ecology. Low levels of SSWD-symptomatic sea stars are still present throughout the impacted range, thus the outlook for population recovery is uncertain.

Patterns of Mass Mortality among Rocky Shore Invertebrates across 100 km of Northeastern Pacific Coastline.

Mass mortalities in natural populations, particularly those that leave few survivors over large spatial areas, may cause long-term ecological perturbations. Yet mass mortalities may remain undocumented or poorly described due to challenges in responding rapidly to unforeseen events, scarcity of baseline data, and difficulties in quantifying rare or patchily distributed species, especially in remote or marine systems. Better chronicling the geographic pattern and intensity of mass mortalities is especially critical in the face of global changes predicted to alter regional disturbance regimes. Here, we couple replicated post-mortality surveys with preceding long-term surveys and historical data to describe a rapid and severe mass mortality of rocky shore invertebrates along the north-central California coast of the northeastern Pacific Ocean. In late August 2011, formerly abundant intertidal populations of the purple sea urchin (Strongylocentrotus purpuratus, a well-known ecosystem engineer), and the predatory six-armed sea star (Leptasterias sp.) were functionally extirpated from ~100 km of coastline. Other invertebrates, including the gumboot chiton (Cryptochiton stelleri) the ochre sea star (Pisaster ochraceus), and subtidal populations of purple sea urchins also exhibited elevated mortality. The pattern and extent of mortality suggest the potential for long-term population, community, and ecosystem consequences, recovery from which may depend on the different dispersal abilities of the affected species.

Settlement behavior of Chthamalus anisopoma larvae largely determines the adult distribution.

In the northern Gulf of California the adult distribution of the intertidal barnacle species, Chthamalus anisopoma, on exposed shores is approximately between 0.0 and 2.0 m above mean low water (MLW). The species is typically absent in protected (from wave splash) areas. In this study, I examined a series of alternative hypotheses relating to the factors that could be responsible for limiting the distribution. Post-settlement factors appear to be unimportant because settlement was largely restricted to areas within the adult distribution. Two processes could account for the high correlation between settlement and adult distributions. First, hydrodynamic factors could restrict deposition of larvae to sites that coincidently were in areas in which individuals could survive to maturity. Second, larvae may choose to settle only on sites where they can survive to maturity. Of the two, the later was supported as settlement could be induced on surfaces outside the adult distribution using transplanted adult conspecifics as cues. Thus, competent larvae were present outside the adult distribution of Chthamalus zone but did not settle under normal conditions. Also, there was no evidence that pre-emption of space by other sessile species, by itself, restricted the distribution of Chthamalus. Settlement within the existing adult distribution may be an evolutionary response to increased mortality for individuals settling outside the adult distribution compared to those settling within it.