Regional differences in fishing behavior determine whether a marine reserve network enhances fishery yield.

A network of marine reserves can enhance yield in depleted fisheries by protecting populations, particularly large, old spawners that supply larvae for interspersed fishing grounds. The ability of marine reserves to enhance sustainable fisheries is much less evident. We report empirical evidence of a marine reserve network improving yield regionally for a sustainable spiny lobster fishery, apparently through the spillover of adult lobsters and behavioral adaptation by the fishing fleet. Results of a Before-After, Control-Impact analysis found catch, effort, and Catch-Per-Unit Effort increased after the establishment of marine reserves in the northern region of the fishery where fishers responded by fishing intensively at reserve borders, but declined in the southern region where they vacated once productive fishing grounds. The adaptation of the northern region of the fishery may have been aided by a history of collaboration between fishers, scientists, and managers, highlighting the value of collaborative research and education programs for preparing fisheries to operate productively within a seascape that includes a large marine reserve network.

A scaffolded and annotated reference genome of giant kelp (Macrocystis pyrifera).

Macrocystis pyrifera (giant kelp), is a brown macroalga of great ecological importance as a primary producer and structure-forming foundational species that provides habitat for hundreds of species. It has many commercial uses (e.g. source of alginate, fertilizer, cosmetics, feedstock). One of the limitations to exploiting giant kelp's economic potential and assisting in giant kelp conservation efforts is a lack of genomic tools like a high quality, contiguous reference genome with accurate gene annotations. Reference genomes attempt to capture the complete genomic sequence of an individual or species, and importantly provide a universal structure for comparison across a multitude of genetic experiments, both within and between species. We assembled the giant kelp genome of a haploid female gametophyte de novo using PacBio reads, then ordered contigs into chromosome level scaffolds using Hi-C. We found the giant kelp genome to be 537 MB, with a total of 35 scaffolds and 188 contigs. The assembly N50 is 13,669,674 with GC content of 50.37%. We assessed the genome completeness using BUSCO, and found giant kelp contained 94% of the BUSCO genes from the stramenopile clade. Annotation of the giant kelp genome revealed 25,919 genes. Additionally, we present genetic variation data based on 48 diploid giant kelp sporophytes from three different Southern California populations that confirms the population structure found in other studies of these populations. This work resulted in a high-quality giant kelp genome that greatly increases the genetic knowledge of this ecologically and economically vital species.

Seaweed biogeochemistry: Global assessment of C:N and C:P ratios and implications for ocean afforestation.

Algal carbon-to-nitrogen (C:N) and carbon-to-phosphorus (C:P) ratios are fundamental for understanding many oceanic biogeochemical processes, such as nutrient flux and climate regulation. We synthesized literature data (444 species, >400 locations) and collected original samples from Tasmania, Australia (51 species, 10 locations) to update the global ratios of seaweed carbon-to-nitrogen (C:N) and carbon-to-phosphorus (C:P). The updated global mean molar ratio for seaweed C:N is 20 (ranging from 6 to 123) and for C:P is 801 (ranging from 76 to 4102). The C:N and C:P ratios were significantly influenced by seawater inorganic nutrient concentrations and seasonality. Additionally, C:N ratios varied by phyla. Brown seaweeds (Ochrophyta, Phaeophyceae) had the highest mean C:N of 27.5 (range: 7.6-122.5), followed by green seaweeds (Chlorophyta) of 17.8 (6.2-54.3) and red seaweeds (Rhodophyta) of 14.8 (5.6-77.6). We used the updated C:N and C:P values to compare seaweed tissue stoichiometry with the most recently reported values for plankton community stoichiometry. Our results show that seaweeds have on average 2.8 and 4.0 times higher C:N and C:P than phytoplankton, indicating seaweeds can assimilate more carbon in their biomass for a given amount of nutrient resource. The stoichiometric comparison presented herein is central to the discourse on ocean afforestation (the deliberate replacement of phytoplankton with seaweeds to enhance the ocean biological carbon sink) by contributing to the understanding of the impact of nutrient reallocation from phytoplankton to seaweeds under large-scale seaweed cultivation.

Past climate-driven range shifts structuring intraspecific biodiversity levels of the giant kelp (Macrocystis pyrifera) at global scales.

The paradigm of past climate-driven range shifts structuring the distribution of marine intraspecific biodiversity lacks replication in biological models exposed to comparable limiting conditions in independent regions. This may lead to confounding effects unlinked to climate drivers. We aim to fill in this gap by asking whether the global distribution of intraspecific biodiversity of giant kelp (Macrocystis pyrifera) is explained by past climate changes occurring across the two hemispheres. We compared the species' population genetic diversity and structure inferred with microsatellite markers, with range shifts and long-term refugial regions predicted with species distribution modelling (SDM) from the last glacial maximum (LGM) to the present. The broad antitropical distribution of Macrocystis pyrifera is composed by six significantly differentiated genetic groups, for which current genetic diversity levels match the expectations of past climate changes. Range shifts from the LGM to the present structured low latitude refugial regions where genetic relics with higher and unique diversity were found (particularly in the Channel Islands of California and in Peru), while post-glacial expansions following ~ 40% range contraction explained extensive regions with homogenous reduced diversity. The estimated effect of past climate-driven range shifts was comparable between hemispheres, largely demonstrating that the distribution of intraspecific marine biodiversity can be structured by comparable evolutionary forces across the global ocean. Additionally, the differentiation and endemicity of regional genetic groups, confers high conservation value to these localized intraspecific biodiversity hotspots of giant kelp forests.

The Blob marine heatwave transforms California kelp forest ecosystems.

Ocean warming has both direct physiological and indirect ecological consequences for marine organisms. Sessile animals may be particularly vulnerable to anomalous warming given constraints in food acquisition and reproduction imposed by sessility. In temperate reef ecosystems, sessile suspension feeding invertebrates provide food for an array of mobile species and act as a critical trophic link between the plankton and the benthos. Using 14 years of seasonal benthic community data across five coastal reefs, we evaluated how communities of sessile invertebrates in southern California kelp forests responded to the "Blob", a period of anomalously high temperatures and low phytoplankton production. We show that this event had prolonged consequences for kelp forest ecosystems. Changes to community structure, including species invasions, have persisted six years post-Blob, suggesting that a climate-driven shift in California kelp forests is underway.

Detrital supply suppresses deforestation to maintain healthy kelp forest ecosystems.

Herbivores can reach extraordinary abundances in many ecosystems. When herbivore abundance is high, heavy grazing can severely defoliate primary producers and, in some cases, even drive ecosystem to undergo regime shifts from a high productivity state to a denuded, low productivity state. While the phenomenon of herbivore-driven regime shifts is well documented, we only partially understand the mechanisms underlying these events. Here, we combine herbivory experiments with 21 years of long-term monitoring data of kelp forest ecosystems to test the hypothesis that herbivores drive regime shifts when herbivory exceeds primary production. To test this hypothesis, we quantified how the foraging habits of an important group of marine herbivores-sea urchins-change with increases in sea urchin biomass and trigger regime shifts to a foundation species, giant kelp (Macrocystis pyrifera). Using experiments, we quantified how the grazing capacity of urchins increases as urchin biomass increases, then we combined these estimates of urchin grazing capacity with estimates of kelp production to predict when and where urchin grazing capacity exceeded kelp production. When grazing capacity exceeded kelp production, sea urchins caused a 50-fold reduction in giant kelp biomass. Our findings support the hypothesis that the balance between herbivory and production underlies herbivore-driven regime shifts in Southern California kelp forests and provides insight into when and where urchins are likely to force regime shifts in kelp forest ecosystems.

Disturbance structures canopy and understory productivity along an environmental gradient.

Disturbances often disproportionately impact different vegetation layers in forests and other vertically stratified ecosystems, shaping community structure and ecosystem function. However, disturbance-driven changes may be mediated by environmental conditions that affect habitat quality and species interactions. In a decade-long field experiment, we tested how kelp forest net primary productivity (NPP) responds to repeated canopy loss along a gradient in grazing and substrate suitability. We discovered that habitat quality can mediate the effects of intensified disturbance on canopy and understory NPP. Experimental annual and quarterly disturbances suppressed total macroalgal NPP, but effects were strongest in high-quality habitats that supported dense kelp canopies that were removed by disturbance. Understory macroalgae partly compensated for canopy NPP losses and this effect magnified with increasing habitat quality. Disturbance-driven increases in understory NPP were still rising after 5-10 years of disturbance, demonstrating the value of long-term experimentation for understanding ecosystem responses to changing disturbance regimes.

Improving the ability of a BACI design to detect impacts within a kelp-forest community.

Distinguishing between human impacts and natural variation in abundance remains difficult because most species exhibit complex patterns of variation in space and time. When ecological monitoring data are available, a before-after-control-impact (BACI) analysis can control natural spatial and temporal variation to better identify an impact and estimate its magnitude. However, populations with limited distributions and confounding spatial-temporal dynamics can violate core assumptions of BACI-type designs. In this study, we assessed how such properties affect the potential to identify impacts. Specifically, we quantified the conditions under which BACI analyses correctly (or incorrectly) identified simulated anthropogenic impacts in a spatially and temporally replicated data set of fish, macroalgal, and invertebrate species found on nearshore subtidal reefs in southern California, USA. We found BACI failed to assess very localized impacts, and had low power but high precision when assessing region-wide impacts. Power was highest for severe impacts of moderate spatial scale, and impacts were most easily detected in species with stable, widely distributed populations. Serial autocorrelation in the data greatly inflated false impact detection rates, and could be partly controlled for statistically, while spatial synchrony in dynamics had no consistent effect on power or false detection rates. Unfortunately, species that offer high power to detect real impacts were also more likely to detect impacts where none had occurred. However, considering power and false detection rates together can identify promising indicator species, and collectively analyzing data for similar species improved the net ability to assess impacts. These insights set expectations for the sizes and severities of impacts that BACI analyses can detect in real systems, point to the importance of serial autocorrelation (but not of spatial synchrony), and indicate how to choose the species, and groups of species, that can best identify impacts.

Variation in disturbance to a foundation species structures the dynamics of a benthic reef community.

Disturbance and foundation species can both have strong impacts on ecosystem structure and function, but studies of their interacting effects are hindered by the long life spans and slow growth of most foundation species. Here, we investigated the extent to which foundation species may mediate the impacts of disturbance on ecological communities, using the kelp forest ecosystem as a study system. Giant kelp (Macrocystis pyrifera) grows rapidly and experiences wave disturbance from winter storms. We developed and analyzed a model of the effects of variable storm regimes on giant kelp population dynamics and of the cascading effects on kelp-mediated competition between benthic community members in kelp forests. Simulations of severe storm regimes resulted in a greater abundance of understory macroalgae and a lower abundance of sessile invertebrates than did milder regimes. Both the cascading effects of periodic loss of giant kelp as well as the degree to which storms directly impacted the benthos (in the form of scouring) influenced the outcome of competition between benthic community members. The model's qualitative predictions were consistent with empirical data from a 20-yr time series of community dynamics, suggesting that interannual variability in disturbance that affects giant kelp abundance can have strong consequences for benthic community structure. Our findings point to the value of long-term studies in elucidating the interacting effects of disturbance and foundation species.

Evidence that spillover from Marine Protected Areas benefits the spiny lobster (Panulirus interruptus) fishery in southern California.

Marine Protected Areas (MPAs) are designed to enhance biodiversity and ecosystem services. Some MPAs are also established to benefit fisheries through increased egg and larval production, or the spillover of mobile juveniles and adults. Whether spillover influences fishery landings depend on the population status and movement patterns of target species both inside and outside of MPAs, as well as the status of the fishery and behavior of the fleet. We tested whether an increase in the lobster population inside two newly established MPAs influenced local catch, fishing effort, and catch-per-unit-effort (CPUE) within the sustainable California spiny lobster fishery. We found greater build-up of lobsters within MPAs relative to unprotected areas, and greater increases in fishing effort and total lobster catch, but not CPUE, in fishing zones containing MPAs vs. those without MPAs. Our results show that a 35% reduction in fishing area resulting from MPA designation was compensated for by a 225% increase in total catch after 6-years, thus indicating at a local scale that the trade-off of fishing ground for no-fishing zones benefitted the fishery.

Iron-Phosphorus Feedbacks Drive Multidecadal Oscillations in Baltic Sea Hypoxia.

Hypoxia has occurred intermittently in the Baltic Sea since the establishment of brackish-water conditions at ∼8,000 years B.P., principally as recurrent hypoxic events during the Holocene Thermal Maximum (HTM) and the Medieval Climate Anomaly (MCA). Sedimentary phosphorus release has been implicated as a key driver of these events, but previous paleoenvironmental reconstructions have lacked the sampling resolution to investigate feedbacks in past iron-phosphorus cycling on short timescales. Here we employ Laser Ablation (LA)-ICP-MS scanning of sediment cores to generate ultra-high resolution geochemical records of past hypoxic events. We show that in-phase multidecadal oscillations in hypoxia intensity and iron-phosphorus cycling occurred throughout these events. Using a box model, we demonstrate that such oscillations were likely driven by instabilities in the dynamics of iron-phosphorus cycling under preindustrial phosphorus loads, and modulated by external climate forcing. Oscillatory behavior could complicate the recovery from hypoxia during future trajectories of external loading reductions.

Foundation species promote community stability by increasing diversity in a giant kelp forest.

Foundation species structure communities, promote biodiversity, and stabilize ecosystem processes by creating locally stable environmental conditions. Despite their critical importance, the role of foundation species in stabilizing natural communities has seldom been quantified. In theory, the stability of a foundation species should promote community stability by enhancing species richness, altering the population fluctuations of individual species, or both. Here we tested the hypothesis that the stability of a marine foundation species, the giant kelp Macrocystis pyrifera, increased the stability of the aggregate biomass of a phylogenetically diverse assemblage of understory algae and sessile invertebrates that compete for space beneath the giant kelp canopy. To achieve this goal, we analyzed an 18-yr time series of the biomass of giant kelp and its associated benthic community collected from 32 plots distributed among nine shallow reefs in the Santa Barbara Channel, USA. We showed that the stability of understory algae and sessile invertebrates was positively and indirectly related to the stability of giant kelp, which primarily resulted from giant kelp's direct positive association with species richness. The stability of all community types was positively related to species richness via increased species stability and species asynchrony. The stabilizing effects of richness were three to four times stronger when algae and invertebrates were considered separately rather than in combination. Our finding that diversity-stability relationships were stronger in communities consisting of species with similar resource requirements suggests that competition for shared resources rather than differential responses to environmental conditions played a more important role in stabilizing the community. Increasing threats to structure-forming foundation species worldwide necessitates a detailed understanding of how they influence their associated community. This study is among the first to show that dampened temporal fluctuations in the biomass of a foundation species is an important determinant of the stability of the complex communities it supports.

Seascape genetics of the stalked kelp Pterygophora californica and comparative population genetics in the Santa Barbara Channel.

We conducted a population genetic analysis of the stalked kelp, Pterygophora californica, in the Santa Barbara Channel, California, USA. The results were compared with previous work on the genetic differentiation of giant kelp, Macrocystis pyrifera, in the same region. These two sympatric kelps not only share many life history and dispersal characteristics but also differ in that dislodged P. californica does not produce floating rafts with buoyant fertile sporophytes, commonly observed for M. pyrifera. We used a comparative population genetic approach with these two species to test the hypothesis that the ability to produce floating rafts increases the genetic connectivity among kelp patches in the Santa Barbara Channel. We quantified the association of habitat continuity and oceanographic distance with the genetic differentiation observed in stalked kelp, like previously conducted for giant kelp. We compared both overall (across all patches) and pairwise (between patches) genetic differentiation. We found that oceanographic transit time, habitat continuity, and geographic distance were all associated with genetic connectivity in P. californica, supporting similar previous findings for M. pyrifera. Controlling for differences in heterozygosity between kelp species using Jost's DEST , we showed that global differentiation and pairwise differentiation were similar among patches between the two kelp species, indicating that they have similar dispersal capabilities despite their differences in rafting ability. These results suggest that rafting sporophytes do not play a significant role in effective dispersal of M. pyrifera at ecologically relevant spatial and temporal scales.

Climate and fishing drive regime shifts in consumer-mediated nutrient cycling in kelp forests.

Globally, anthropogenic pressures are reducing the abundances of marine species and altering ecosystems through modification of trophic interactions. Yet, consumer declines also disrupt important bottom-up processes, like nutrient recycling, which are critical for ecosystem functioning. Consumer-mediated nutrient dynamics (CND) is now considered a major biogeochemical component of most ecosystems, but lacking long-term studies, it is difficult to predict how CND will respond to accelerating disturbances in the wake of global change. To aid such predictions, we coupled empirical ammonium excretion rates with an 18-year time series of the standing biomass of common benthic macroinvertebrates in southern California kelp forests. This time series of excretion rates encompassed an extended period of extreme ocean warming, disease outbreaks, and the abolishment of fishing at two of our study sites, allowing us to assess kelp forest CND across a wide range of environmental conditions. At their peak, reef invertebrates supplied an average of 18.3 ± 3.0 µmol NH4 +  m-2  hr-1 to kelp forests when sea stars were regionally abundant, but dropped to 3.5 ± 1.0 µmol NH4 +  m-2  hr-1 following their mass mortality due to disease during a prolonged period of extreme warming. However, a coincident increase in the abundance of the California spiny lobster, Palinurus interupptus (Randall, 1840), likely in response to both reduced fishing and a warmer ocean, compensated for much of the recycled ammonium lost to sea star mortality. Both lobsters and sea stars are widely recognized as key predators that can profoundly influence community structure in benthic marine systems. Our study is the first to demonstrate their importance in nutrient cycling, thus expanding their roles in the ecosystem. Climate change is increasing the frequency and severity of warming events, and rising human populations are intensifying fishing pressure in coastal ecosystems worldwide. Our study documents how these projected global changes can drive regime shifts in CND and fundamentally alter a critical ecosystem function.

Species insurance trumps spatial insurance in stabilizing biomass of a marine macroalgal metacommunity.

Because natural ecosystems are complex, it is difficult to predict how their variability scales across space and levels of organization. The species-insurance hypothesis predicts that asynchronous dynamics among species should reduce variability when biomass is aggregated either from local species populations to local multispecies communities, or from metapopulations to metacommunities. Similarly, the spatial-insurance hypothesis predicts that asynchronous spatial dynamics among either local populations or local communities should stabilize metapopulation biomass and metacommunity biomass, respectively. In combination, both species and spatial insurance reduce variation in metacommunity biomass over time, yet these insurances are rarely considered together in natural systems. We partitioned the extent that species insurance and spatial insurance reduced the annual variation in macroalgal biomass in a southern California kelp forest. We quantified variability and synchrony at two levels of organization (population and community) and two spatial scales (local plots and region) and quantified the strength of species and spatial insurance by comparing observed variability and synchrony in aggregate biomass to null models of independent species or spatial dynamics based on cyclic-shift permutation. Spatial insurance was weak, presumably because large-scale oceanographic processes in the study region led to high spatial synchrony at both population- and community-level biomass. Species insurance was stronger due to asynchronous dynamics among the metapopulations of a few common species. In particular, a regional decline in the dominant understory kelp species Pterygophora californica was compensated for by the rise of three subdominant species. These compensatory dynamics were associated with positive values of the Pacific Decadal Oscillation, indicating that differential species tolerances to warmer temperature and nutrient-poor conditions may underlie species insurance in this system. Our results illustrate how species insurance can stabilize aggregate community properties in natural ecosystems where environmental conditions vary over broad spatial scales.

Loss of foundation species: disturbance frequency outweighs severity in structuring kelp forest communities.

Disturbances often cause the disproportionate loss of foundation species but understanding how the frequency and severity of disturbance to such organisms influence biological communities remains unresolved. This gap in knowledge exists in part because of the rarity of ecologically meaningful studies capable of disentangling different elements of disturbance. Hence, we carried out a long-term (9 yr), large-scale (2,000 m2 plots), spatially replicated (4 sites) field experiment in which we manipulated disturbance to a globally distributed marine foundation species, the giant kelp Macrocystis pyrifera, and tracked community responses over time. To distinguish the effects of disturbance frequency and severity on the biodiversity and composition of temperate rocky reef communities, we simulated the repeated loss of giant kelp from destructive winter waves across a background of natural variation in disturbance. By following the response of over 200 taxa from the surrounding community, we discovered that the frequency of disturbance to giant kelp changed the biomass, diversity, and composition of community guilds in a manner commensurate with their dependence on the physical (i.e., benthic light and space), trophic (i.e., living and detrital biomass), and habitat (i.e., biogenic structure) resources mediated by this foundation species. Annual winter disturbance to giant kelp reduced living and detrital giant kelp biomass by 57% and 40%, respectively, enhanced bottom light by 22%, and halved the seafloor area covered by giant kelp holdfasts. Concomitantly, the biomass of understory algae and epilithic sessile invertebrates more than doubled, while the biomass of rock-boring clams, mobile invertebrates, and fishes decreased 30-61%. Frequent loss of giant kelp boosted understory algal richness by 82% and lowered sessile invertebrate richness by 13% but did not affect the biodiversity of mobile fauna. In contrast to changes driven by disturbance frequency, interannual variation in the severity of disturbance to giant kelp had weaker, less consistent effects, causing only modest changes in assemblages of sessile invertebrates, mobile invertebrate herbivores, and fishes. Our results broaden the foundation species concept by demonstrating that repeated disturbance to a dominant habitat-forming species can outweigh the influence of less frequent but severe disturbances for the surrounding community.

Improved estimates of net primary production, growth, and standing crop of Macrocystis pyrifera in Southern California.

The giant kelp Macrocystis pyrifera forms subtidal forests on shallow reefs in temperate regions of the world. It is one of the fastest-growing multicellular autotrophs on Earth and its high productivity supports diverse marine food webs. In 2008, we published a method for estimating biomass and net primary production (NPP) of giant kelp along with five years of data, to provide a more integrated measure of NPP than those yielded by previous methods. Our method combines monthly field measurements of standing crop and loss rates with a model of kelp biomass dynamics to estimate instantaneous mass-specific growth rates and NPP for each season of each year. We have since improved our approach to account for several previously unresolved sources of biomass loss. These improvements have led to a near doubling of our prior estimates of growth and NPP. At our site with the most persistent stand of giant kelp, NPP averages ~5.2 kg dry mass·m-2 ·yr-1 and results from the rapid growth (~3.5% per d) of a relatively small standing biomass (~0.4 kg dry mass/m2 on average) that turns over ~12 times annually. Here we provide revised estimates of seasonal biomass, growth, and NPP for the five years covered by our previous publication (2002-2006), along with more than a decade of additional data (2007-2017). We also present updated relationships for predicting giant kelp biomass and NPP from much more easily obtained measurements of frond density. These data can be used to understand the mechanisms that drive variation in giant kelp NPP at a wide range of temporal scales. No copyright or proprietary restrictions are associated with the use of this data set other than citation of this Data Paper.

Giant kelp, Macrocystis pyrifera, increases faunal diversity through physical engineering.

Foundation species define the ecosystems they live in, but ecologists have often characterized dominant plants as foundational without supporting evidence. Giant kelp has long been considered a marine foundation species due to its complex structure and high productivity; however, there is little quantitative evidence to evaluate this. Here, we apply structural equation modelling to a 15-year time series of reef community data to evaluate how giant kelp affects the reef community. Although species richness was positively associated with giant kelp biomass, most direct paths did not involve giant kelp. Instead, the foundational qualities of giant kelp were driven mostly by indirect effects attributed to its dominant physical structure and associated engineering influence on the ecosystem, rather than by its use as food by invertebrates and fishes. Giant kelp structure has indirect effects because it shades out understorey algae that compete with sessile invertebrates. When released from competition, sessile species in turn increase the diversity of mobile predators. Sea urchin grazing effects could have been misinterpreted as kelp effects, because sea urchins can overgraze giant kelp, understorey algae and sessile invertebrates alike. Our results confirm the high diversity and biomass associated with kelp forests, but highlight how species interactions and habitat attributes can be misconstrued as direct consequences of a foundation species like giant kelp.

Scale-specific drivers of kelp forest communities.

Identifying spatial scales of variation in natural communities and the processes driving them is critical for obtaining a predictive understanding of biodiversity. In this study, we focused on diverse communities inhabiting productive kelp forests on shallow subtidal rocky reefs in southern California, USA. We combined long-term community surveys from 86 sites with detailed environmental data to determine what structures assemblages of fishes, invertebrates and algae at multiple spatial scales. We identified the spatial scales of variation in species composition using a hierarchical analysis based on eigenfunctions, and assessed how sea surface temperature (SST), water column chlorophyll, giant kelp biomass, wave exposure and potential propagule delivery strength contributed to community variation at each scale. Spatial effects occurring at multiple scales explained 60% of the variation in fish assemblages and 52% of the variation in the assemblages of invertebrates and algae. Most variation occurred over broad spatial scales (> 200 km) consistent with spatial heterogeneity in SST and potential propagule delivery strength, while the latter also explained community variation at medium scales (65-200 km). Small scale (1-65 km) community variation was substantial but not linked to any of the measured drivers. Conclusions were consistent for both reef fishes and benthic invertebrates and algae, despite sharp differences in their adult mobility. Our results demonstrate the scale dependence of environmental drivers on kelp forest communities, showing that most species were strongly sorted along oceanographic conditions over various spatial scales. Such spatial effects must be integrated into models assessing the response of marine ecosystems to climate change.

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.