Kelp-forest dynamics controlled by substrate complexity.

The factors that determine why ecosystems exhibit abrupt shifts in state are of paramount importance for management, conservation, and restoration efforts. Kelp forests are emblematic of such abruptly shifting ecosystems, transitioning from kelp-dominated to urchin-dominated states around the world with increasing frequency, yet the underlying processes and mechanisms that control their dynamics remain unclear. Here, we analyze four decades of data from biannual monitoring around San Nicolas Island, CA, to show that substrate complexity controls both the number of possible (alternative) states and the velocity with which shifts between states occur. The superposition of community dynamics with reconstructions of system stability landscapes reveals that shifts between alternative states at low-complexity sites reflect abrupt, high-velocity events initiated by pulse perturbations that rapidly propel species across dynamically unstable state-space. In contrast, high-complexity sites exhibit a single state of resilient kelp-urchin coexistence. Our analyses suggest that substrate complexity influences both top-down and bottom-up regulatory processes in kelp forests, highlight its influence on kelp-forest stability at both large (island-wide) and small (<10 m) spatial scales, and could be valuable for holistic kelp-forest management.

Projecting kelp (Ecklonia radiata) gametophyte thermal adaptation and persistence under climate change.

BACKGROUND AND AIMS: Kelp forests underpin temperate marine ecosystems but are declining due to ocean warming, causing loss of associated ecosystem services. Projections suggest significant future decline but often only consider the persistence of adult sporophytes. Kelps have a biphasic life cycle, and the haploid gametophyte can be more thermally tolerant than the sporophyte. Therefore, projections may be altered when considering the thermal tolerance of gametophytes. METHODS: We undertook thermal tolerance experiments to quantify the effect of temperature on gametophyte survival, relative growth rate (RGR) and sex ratio for three genetically distinct populations of Ecklonia radiata gametophytes from comparatively high, mid- and low latitudes (43°, 33° and 30°S). We then used these data to project the likely consequences of climate-induced thermal change on gametophyte persistence and performance across its eastern Australian range, using generalized additive and linear models. KEY RESULTS: All populations were adapted to local temperatures and their thermal maximum was 2-3 °C above current maximum in situ temperatures. The lowest latitude population was most thermally tolerant (~70 % survival up to 27 °C), while survival and RGR decreased beyond 25.5 and 20.5 °C for the mid- and low-latitude populations, respectively. Sex ratios were skewed towards females with increased temperature in the low- and high-latitude populations. Spatially explicit model projections under future ocean warming (2050-centred) revealed a minimal decline in survival (0-30 %) across populations, relative to present-day predictions. RGRs were also projected to decline minimally (0-2 % d-1). CONCLUSIONS: Our results contrast with projections for the sporophyte stage of E. radiata, which suggest a 257-km range contraction concurrent with loss of the low-latitude population by 2100. Thermal adaptation in E. radiata gametophytes suggests this life stage is likely resilient to future ocean warming and is unlikely to be a bottleneck for the future persistence of kelp.

Kelp forest food webs as hot spots for the accumulation of microplastic and polybrominated diphenyl ether pollutants.

Kelp forests (KFs) are one of the most significant marine ecosystems in the planet. They serve as a refuge for a wide variety of marine species of ecological and economic importance. Additionally, they aid with carbon sequestration, safeguard the coastline, and maintain water quality. Microplastic (MP) and polybrominated diphenyl ethers (PBDEs) concentrations were analyzed across trophic levels in KFs around Todos Santos Bay. Spatial variation patterns were compared at three sites in 2021 and temporal change at Todos Santos Island (TSI) in 2021 and 2022. We analyzed these MPs and PBDEs in water, primary producers (Macrocystis pyrifera), grazers (Strongylocentrotus purpuratus), predators (Semicossyphus pulcher), and kelp detritus. MPs were identified in all samples (11 synthetic and 1 semisynthetic polymer) and confirmed using Fourier-transform infrared microspectroscopy-attenuated total reflectance (µ-FTIR-ATR). The most abundant type of MP is polyester fibers. Statistically significant variations in MP concentration were found only in kelps, with the greatest average concentrations in medium-depth kelps from TSI in 2022 (0.73 ± 0.58 MP g-1 ww) and in the kelp detritus from TSI in 2021 (0.96 ± 0.64 MP g-1 ww). Similarly, PBDEs were found in all samples, with the largest concentration found in sea urchins from Punta San Miguel (0.93 ± 0.24 ng g-1 ww). The similarity of the polymers can indicate a trophic transfer of MPs. This study shows the extensive presence of MP and PBDE subtropical trophic web of a KF, but correlating these compounds in environmental samples is highly complex, influenced by numerous factors that could affect their presence and behavior. However, this suggests that there is a potential risk to the systems and the services that KFs offer.

Examining the reproductive success of bull kelp (Nereocystis luetkeana, Phaeophyceae, Laminariales) in climate change conditions.

Climate change is affecting marine ecosystems in many ways, including raising temperatures and leading to ocean acidification. From 2014 to 2016, an extensive marine heat wave extended along the west coast of North America and had devastating effects on numerous species, including bull kelp (Nereocystis luetkeana). Bull kelp is an important foundation species in coastal ecosystems and can be affected by marine heat waves and ocean acidification; however, the impacts have not been investigated on sensitive early life stages. To determine the effects of changing temperatures and carbonate levels on Northern California's bull kelp populations, we collected sporophylls from mature bull kelp individuals in Point Arena, CA. At the Bodega Marine Laboratory, we released spores from field-collected bull kelp, and cultured microscopic gametophytes in a common garden experiment with a fully factorial design crossing modern conditions (11.63 ± 0.54°C and pH 7.93 ± 0.26) with observed extreme climate conditions (15.56 ± 0.83°C and 7.64 ± 0.32 pH). Our results indicated that both increased temperature and decreased pH influenced growth and egg production of bull kelp microscopic stages. Increased temperature resulted in decreased gametophyte survival and offspring production. In contrast, decreased pH had less of an effect but resulted in increased gametophyte survival and offspring production. Additionally, increased temperature significantly impacted reproductive timing by causing female gametophytes to produce offspring earlier than under ambient temperature conditions. Our findings can inform better predictions of the impacts of climate change on coastal ecosystems and provide key insights into environmental dynamics regulating the bull kelp lifecycle.

Urchin grazing of kelp gametophytes in warming oceans.

Sea urchins can cause extensive damage to kelp forests, and their overgrazing can create extensive barren areas, leading to a loss of biodiversity. Barrens may persist when the recruitment of kelp, which occurs through the microscopic haploid gametophyte stage, is suppressed. However, the ecology of kelp gametophytes is poorly understood, and here we investigate if grazing by juvenile urchins on kelp gametophytes can suppress kelp recruitment and if this is exacerbated by climate change. We compared grazing of Ecklonia radiata gametophytes by two species of juvenile urchins, the tropical Tripneustes gratilla and the temperate Centrostephanus rodgersii, at winter (19°C), summer (23°C), and ocean warming (26°C) temperatures for the low-latitude range edge of E. radiata, which is vulnerable to ocean warming. We examined the rate of recovery of gametophytes following grazing and determined whether they survived and formed sporophytes after ingestion by sea urchins. Both T. gratilla and C. rodgersii grazed E. radiata gametophytes, reducing their abundance compared to no grazing controls. Surprisingly, temperature did not influence grazing rates, but gametophytes did not recover from grazing in the ocean warming (26°C) treatment. Gametophytes survived ingestion by both species of sea urchin and formed sporophytes after ingestion by T. gratilla, but not C. rodgersii. These results suggest complex grazer-gametophyte interactions, in which both negative (reduced abundance and poor recovery with warming) and positive (facilitated recruitment) effects are possible. Small grazers may play a more important role in kelp ecosystem function than previously thought and should be considered in our understanding of alternate stable states.

Parasites in kelp-forest food webs increase food-chain length, complexity, and specialization, but reduce connectance.

We explored whether parasites are important in kelp forests by examining their effects on a high-quality, high-resolution kelp-forest food web. After controlling for generic effects of network size, parasites affected kelp-forest food web structure in some ways consistent with other systems. Parasites increased the trophic span of the web, increasing top predator vulnerability and the longest chain length. Unique links associated with parasites, such as concomitant predation (consumption of parasites along with their hosts by predators) increased the frequency of network motifs involving mutual consumption and decreased niche contiguity of free-living species. However, parasites also affected kelp-forest food web structure in ways not seen in other systems. Kelp-forest parasites are richer and more specialized than other systems. As a result, parasites reduced diet generality and decreased connectance in the kelp forest. Although mutual consumption motifs increased in frequency, this motif type was still a small fraction of all possible motifs, so their increase in frequency was not enough to compensate for the decrease in connectance caused by adding many specialist parasite species.

Coupled changes in pH, temperature, and dissolved oxygen impact the physiology and ecology of herbivorous kelp forest grazers.

Understanding species' responses to upwelling may be especially important in light of ongoing environmental change. Upwelling frequency and intensity are expected to increase in the future, while ocean acidification and deoxygenation are expected to decrease the pH and dissolved oxygen (DO) of upwelled waters. However, the acute effects of a single upwelling event and the integrated effects of multiple upwelling events on marine organisms are poorly understood. Here, we use in situ measurements of pH, temperature, and DO to characterize the covariance of environmental conditions within upwelling-dominated kelp forest ecosystems. We then test the effects of acute (0-3 days) and chronic (1-3 months) upwelling on the performance of two species of kelp forest grazers, the echinoderm, Mesocentrotus franciscanus, and the gastropod, Promartynia pulligo. We exposed organisms to static conditions in a regression design to determine the shape of the relationship between upwelling and performance and provide insights into the potential effects in a variable environment. We found that respiration, grazing, growth, and net calcification decline linearly with increasing upwelling intensity for M. francicanus over both acute and chronic timescales. Promartynia pulligo exhibited decreased respiration, grazing, and net calcification with increased upwelling intensity after chronic exposure, but we did not detect an effect over acute timescales or on growth after chronic exposure. Given the highly correlated nature of pH, temperature, and DO in the California Current, our results suggest the relationship between upwelling intensity and growth in the 3-month trial could potentially be used to estimate growth integrated over long-term dynamic oceanographic conditions for M. franciscanus. Together, these results indicate current exposure to upwelling may reduce species performance and predicted future increases in upwelling frequency and intensity could affect ecosystem function by modifying the ecological roles of key species.

DNA reconciles morphology and colouration in the drunk blenny genus Scartichthys (Teleostei: Blenniidae) and provides insights into their evolutionary history.

The blenniids of the genus Scartichthys are one of the most common fishes of Central and South American Pacific coastal reefs. This being said, Scartichthys spp. remain difficult to identify in the field, and identification is particularly challenging across the c. 6000 km where three of the four currently accepted species are known to occur in sympatry. A reason for this is that the main taxonomic characters from traditional taxonomy are indeed elusive. In addition, at the same time, species can display multiple colour patterns in the field, depending on their ontogenetic stage, habitat association and reproductive behaviour. Overall, molecular characterization is warranted to help address these issues. In this study, the authors have used a novel approach to revise the genus by combining colouration, morphological and molecular data of representative specimens of the four currently valid species and seven described colour patterns. From this, the authors show that only three of the four species should be considered as valid; Scartichthys gigas (Steindachner, 1876), Scartichthys variolatus (Valenciennes, 1836) and Scartichthys viridis (Valenciennes, 1836), whereas Scartichthys crapulatus (Williams, 1990) should be synonymized with S. viridis. In the same way, the analyses in this study show that one of the colour patterns attributed so far only to S. gigas is characteristic of the juvenile stages of S. viridis. The time-calibrated phylogeny of this study shows that this genus is relatively young and that the estimated time of divergence between S. gigas and S. viridis is c. 1.71 Ma. In comparison, the Desventuradas and Juan Fernandez Islands endemic S. variolatus diverged c. 1.95 Ma. The results of this study help to clarify the taxonomy of Scartichthys.

Grazer behaviour can regulate large-scale patterning of community states.

Ecosystem patterning can arise from environmental heterogeneity, biological feedbacks that produce multiple persistent ecological states, or their interaction. One source of feedbacks is density-dependent changes in behaviour that regulate species interactions. By fitting state-space models to large-scale (~500 km) surveys on temperate rocky reefs, we find that behavioural feedbacks best explain why kelp and urchin barrens form either reef-wide patches or local mosaics. Best-supported models in California include feedbacks where starvation intensifies grazing across entire reefs create reef-scale, alternatively stable kelp- and urchin-dominated states (32% of reefs). Best-fitting models in New Zealand include the feedback of urchins avoiding dense kelp stands that can increase abrasion and predation risk, which drives a transition from shallower urchin-dominated to deeper kelp-dominated zones, with patchiness at 3-8 m depths with intermediate wave stress. Connecting locally studied processes with region-wide data, we highlight how behaviour can explain community patterning and why some systems exhibit community-wide alternative stable states.

Simplification, not "tropicalization", of temperate marine ecosystems under ocean warming and acidification.

Ocean warming is altering the biogeographical distribution of marine organisms. In the tropics, rising sea surface temperatures are restructuring coral reef communities with sensitive species being lost. At the biogeographical divide between temperate and tropical communities, warming is causing macroalgal forest loss and the spread of tropical corals, fishes and other species, termed "tropicalization". A lack of field research into the combined effects of warming and ocean acidification means there is a gap in our ability to understand and plan for changes in coastal ecosystems. Here, we focus on the tropicalization trajectory of temperate marine ecosystems becoming coral-dominated systems. We conducted field surveys and in situ transplants at natural analogues for present and future conditions under (i) ocean warming and (ii) both ocean warming and acidification at a transition zone between kelp and coral-dominated ecosystems. We show that increased herbivory by warm-water fishes exacerbates kelp forest loss and that ocean acidification negates any benefits of warming for range extending tropical corals growth and physiology at temperate latitudes. Our data show that, as the combined effects of ocean acidification and warming ratchet up, marine coastal ecosystems lose kelp forests but do not gain scleractinian corals. Ocean acidification plus warming leads to overall habitat loss and a shift to simple turf-dominated ecosystems, rather than the complex coral-dominated tropicalized systems often seen with warming alone. Simplification of marine habitats by increased CO2 levels cascades through the ecosystem and could have severe consequences for the provision of goods and services.

Homogenization and miniaturization of habitat structure in temperate marine forests.

Humans are rapidly transforming the structural configuration of the planet's ecosystems, but these changes and their ecological consequences remain poorly quantified in underwater habitats. Here, we show that the loss of forest-forming seaweeds and the rise of ground-covering 'turfs' across four continents consistently resulted in the miniaturization of underwater habitat structure, with seascapes converging towards flattened habitats with smaller habitable spaces. Globally, turf seascapes occupied a smaller architectural trait space and were structurally more similar across regions than marine forests, evidencing habitat homogenization. Surprisingly, such habitat convergence occurred despite turf seascapes consisting of vastly different species richness and with different taxa providing habitat architecture, as well as across disparate drivers of marine forest decline. Turf seascapes contained high sediment loads, with the miniaturization of habitat across 100s of km in mid-Western Australia resulting in reefs retaining an additional ~242 million tons of sediment (four orders of magnitude more than the sediments delivered fluvially annually). Together, this work demonstrates that the replacement of marine forests by turfs is a generalizable phenomenon that has profound consequences for the ecology of temperate reefs.

Impacts of ocean warming on kelp forest ecosystems.

Kelp forests represent some of the most diverse and productive habitats on Earth, and provide a range of ecosystem goods and services on which human populations depend. As the distribution and ecophysiology of kelp species is strongly influenced by temperature, recent warming trends in many regions have been linked with concurrent changes in kelp populations, communities and ecosystems. Over the past decade, the number of reports of ocean warming impacts on kelp forests has risen sharply. Here, I synthesise recent studies to highlight general patterns and trends. While kelp responses to climate change vary greatly between ocean basins, regions and species, there is compelling evidence to show that ocean warming poses an unequivocal threat to the persistence and integrity of kelp forest ecosystems in coming decades.

Intra-Annual Variability in Responses of a Canopy Forming Kelp to Cumulative Low Tide Heat Stress: Implications for Populations at the Trailing Range Edge.

Anthropogenic climate change is driving the redistribution of species at a global scale. For marine species, populations at trailing edges often live very close to their upper thermal limits and, as such, poleward range contractions are one of the most pervasive effects of ongoing and predicted warming. However, the mechanics of processes driving such contractions are poorly understood. Here, we examined the response of the habitat forming kelp, Laminaria digitata, to realistic terrestrial heatwave simulations akin to those experienced by intertidal populations persisting at the trailing range edge in the northeast Atlantic (SW England). We conducted experiments in both spring and autumn to determine temporal variability in the effects of heatwaves. In spring, heatwave scenarios caused minimal stress to L. digitata but in autumn all scenarios tested resulted in tissue being nonviable by the end of each assay. The effects of heatwave scenarios were only apparent after consecutive exposures, indicating erosion of resilience over time. Monthly field surveys corroborated experimental evidence as the prevalence of bleaching (an indication of physiological stress and tissue damage) in natural populations was greatest in autumn and early winter. Overall, our data showed that L. digitata populations in SW England persist close to their upper physiological limits for emersion stress in autumn. As the intensity of extreme warming events is likely to increase with anthropogenic climate change, thermal conditions experienced during periods of emersion will soon exceed physiological thresholds and will likely induce widespread mortality and consequent changes at the population level.

Nearshore seascape connectivity enhances seagrass meadow nursery function.

Diverse habitats composing coastal seascapes occur in close proximity, connected by the flux of materials and fauna across habitat boundaries. Understanding how seascape connectivity alters important ecosystem functions for fish, however, is not well established. For a seagrass-dominant seascape, we predicted that configuration and composition of adjacent habitats would alter habitat access for fauna and trophic subsidies, enhancing nursery function for juvenile fish. In an extensive Zostera marina seagrass meadow, we established sites adjacent to (1) highly complex and productive kelp forests (Nereocystis luetkeana), (2) unvegetated sand habitats, and (3) in the seagrass meadow interior. Using SCUBA, we conducted underwater observations of young-of the-year (YOY) rockfish (Sebastes spp.) recruitment across sites. Using generalized linear mixed effects models, we assessed the role of seascape adjacency relative to seagrass provisions (habitat complexity and prey) on YOY recruitment. YOY rockfish collections were used to trace sources of allochthonous vs. autochthonous primary production in the seagrass food web, via a δ13 C and δ15 N isotopic mixing model, and prey consumption using stomach contents. Overall, seagrass nursery function was strongly influenced by adjacent habitats and associated subsidies. Allochthonous N. luetkeana was the greatest source of energy assimilated by YOY rockfish within seagrass sites. In seagrass sites adjacent to N. luetkeana kelp forests, YOYs consumed higher quality prey, which corresponded with better body condition relative to sites adjacent to sand. Moreover, kelp forest adjacency enhanced YOY rockfish recruitment within the seagrass meadow, suggesting that habitat complexity is a key seascape feature influencing the nursery function of nearshore habitats. In general, to promote seascape connectivity, the conservation and restoration of nursery habitats should prioritize the inclusion of habitat mosaics of high structural complexity and productivity. We illustrate and emphasize the importance of using a seascape-level approach that considers linkages among habitats for the management of important nearshore ecosystem functions.

Sudden collapse of a mesopredator reveals its complementary role in mediating rocky reef regime shifts.

While changes in the abundance of keystone predators can have cascading effects resulting in regime shifts, the role of mesopredators in these processes remains underexplored. We conducted annual surveys of rocky reef communities that varied in the recovery of a keystone predator (sea otter, Enhydra lutris) and the mass mortality of a mesopredator (sunflower sea star, Pycnopodia helianthoides) due to an infectious wasting disease. By fitting a population model to empirical data, we show that sea otters had the greatest impact on the mortality of large sea urchins, but that Pycnopodia decline corresponded to a 311% increase in medium urchins and a 30% decline in kelp densities. Our results reveal that predator complementarity in size-selective prey consumption strengthens top-down control on urchins, affecting the resilience of alternative reef states by reinforcing the resilience of kelp forests and eroding the resilience of urchin barrens. We reveal previously underappreciated species interactions within a 'classic' trophic cascade and regime shift, highlighting the critical role of middle-level predators in mediating rocky reef state transitions.

Projected climate changes threaten ancient refugia of kelp forests in the North Atlantic.

Intraspecific genetic variability is critical for species adaptation and evolution and yet it is generally overlooked in projections of the biological consequences of climate change. We ask whether ongoing climate changes can cause the loss of important gene pools from North Atlantic relict kelp forests that persisted over glacial-interglacial cycles. We use ecological niche modelling to predict genetic diversity hotspots for eight species of large brown algae with different thermal tolerances (Arctic to warm temperate), estimated as regions of persistence throughout the Last Glacial Maximum (20,000 YBP), the warmer Mid-Holocene (6,000 YBP), and the present. Changes in the genetic diversity within ancient refugia were projected for the future (year 2100) under two contrasting climate change scenarios (RCP2.6 and RCP8.5). Models predicted distributions that matched empirical distributions in cross-validation, and identified distinct refugia at the low latitude ranges, which largely coincide among species with similar ecological niches. Transferred models into the future projected polewards expansions and substantial range losses in lower latitudes, where richer gene pools are expected (in Nova Scotia and Iberia for cold affinity species and Gibraltar, Alboran, and Morocco for warm-temperate species). These effects were projected for both scenarios but were intensified under the extreme RCP8.5 scenario, with the complete borealization (circum-Arctic colonization) of kelp forests, the redistribution of the biogeographical transitional zones of the North Atlantic, and the erosion of global gene pools across all species. As the geographic distribution of genetic variability is unknown for most marine species, our results represent a baseline for identification of locations potentially rich in unique phylogeographic lineages that are also climatic relics in threat of disappearing.

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.

From the predictable to the unexpected: kelp forest and benthic invertebrate community dynamics following decades of sea otter expansion.

The recovery of predators has the potential to restore ecosystems and fundamentally alter the services they provide. One iconic example of this is keystone predation by sea otters in the Northeast Pacific. Here, we combine spatial time series of sea otter abundance, canopy kelp area, and benthic invertebrate abundance from Washington State, USA, to examine the shifting consequences of sea otter reintroduction for kelp and kelp forest communities. We leverage the spatial variation in sea otter recovery to understand connections between sea otters and the kelp forest community. Sea otter increases created a pronounced decline in sea otter prey-particularly kelp-grazing sea urchins-and led to an expansion of canopy kelps from the late 1980s until roughly 2000. However, while sea otter and kelp population growth rates were positively correlated prior to 2002, this association disappeared over the last two decades. This disconnect occurred despite surveys showing that sea otter prey have continued to decline. Kelp area trends are decoupled from both sea otter and benthic invertebrate abundance at current densities. Variability in kelp abundance has declined in the most recent 15 years, as it has the synchrony in kelp abundance among sites. Together, these findings suggest that initial nearshore community responses to sea otter population expansion follow predictably from trophic cascade theory, but now, other factors may be as or more important in influencing community dynamics. Thus, the utility of sea otter predation in ecosystem restoration must be considered within the context of complex and shifting environmental conditions.

Climate-driven disparities among ecological interactions threaten kelp forest persistence.

The combination of ocean warming and acidification brings an uncertain future to kelp forests that occupy the warmest parts of their range. These forests are not only subject to the direct negative effects of ocean climate change, but also to a combination of unknown indirect effects associated with changing ecological landscapes. Here, we used mesocosm experiments to test the direct effects of ocean warming and acidification on kelp biomass and photosynthetic health, as well as climate-driven disparities in indirect effects involving key consumers (urchins and rock lobsters) and competitors (algal turf). Elevated water temperature directly reduced kelp biomass, while their turf-forming competitors expanded in response to ocean acidification and declining kelp canopy. Elevated temperatures also increased growth of urchins and, concurrently, the rate at which they thinned kelp canopy. Rock lobsters, which are renowned for keeping urchin populations in check, indirectly intensified negative pressures on kelp by reducing their consumption of urchins in response to elevated temperature. Overall, these results suggest that kelp forests situated towards the low-latitude margins of their distribution will need to adapt to ocean warming in order to persist in the future. What is less certain is how such adaptation in kelps can occur in the face of intensifying consumptive (via ocean warming) and competitive (via ocean acidification) pressures that affect key ecological interactions associated with their persistence. If such indirect effects counter adaptation to changing climate, they may erode the stability of kelp forests and increase the probability of regime shifts from complex habitat-forming species to more simple habitats dominated by algal turfs.

A phylogeographic investigation of the kelp genus Laminaria (Laminariales, Phaeophyceae), with emphasis on the South Atlantic Ocean.

The genus Laminaria has a wide distribution range compared with other kelp genera because it is found in both the North and the South Atlantic, on both sides of the North Pacific, as well as in the Mediterranean. Hypotheses behind this biogeographical pattern have been discussed by several authors but have not yet been fully evaluated with time-calibrated phylogenies. Based on the analysis of four molecular markers (ITS2, rbcL, atp8 and trnWI), our goal was to reassess the Laminaria species diversity in South Africa, assess its relationship with the other species distributed in the South Atlantic and reconstruct the historical biogeography of the genus. Our results confirm the occurrence of a single species, L. pallida, in southern Africa, and its sister relationship with the North Atlantic L. ochroleuca. Both species belonged to a clade containing the other South Atlantic species: L. abyssalis from Brazil, and the Mediterranean L. rodriguezii. Our time-calibrated phylogenies suggest that Laminaria originated in the northern Pacific around 25 mya, followed by at least two migration events through the Bering Strait after its opening (~5.32 mya). Today, the first is represented by L. solidungula in the Arctic, while the second gave rise to the rest of the Atlantic species. The colonization of the North Atlantic was followed by a gradual colonization southward along the west coast of Europe, into the Mediterranean (~2.07 mya) and two recent, but disconnected, migrations (~1.34 and 0.87 mya) across the equator, giving rise to L. abyssalis in Brazil and L. pallida in southern Africa, respectively.