Top-down versus bottom-up: Grazing and upwelling regime alter patterns of primary productivity in a warm-temperate system.

Community structure is driven by the interaction of physical processes and biological interactions that can vary across environmental gradients and the strength of top-down control is expected to vary along gradients of primary productivity. In coastal marine systems, upwelling drives regional resource availability through the bottom-up effect of nutrient subsidies. This alters rates of primary production and is expected to alter algae-herbivore interactions in rocky intertidal habitats. Despite the potential for upwelling to alter these interactions, the interaction of upwelling and grazing pressure is poorly understood, particularly for warm-temperate systems. Using in situ herbivore exclusion experiments replicated across multiple upwelling regimes, we investigated the effects of both grazing pressure, upwelling, and their interactions on the sessile invertebrate community and biomass of macroalgal communities in a warm-temperate system. The sessile invertebrate cover showed indirect effects of grazing, being consistently low where algal biomass was high at upwelling sites and at nonupwelling sites when grazers were excluded. The macroalgal cover was greater at upwelling sites when grazers were excluded and there was a strong effect of succession throughout the experimental period. Grazing effects were greater at upwelling sites, particularly during winter months. There was a nonsignificant trend toward greater grazing pressure on early than later successional stages. Our results show that the positive bottom-up effects of nutrient supply on algal production do not overwhelm top-down control in this warm-temperate system but do have knock-on consequences for invertebrates that compete with macroalgae for space. We speculate that global increases in air and sea surface temperatures in warm-temperate systems will promote top-down effects in upwelling regions by increasing herbivore metabolic and growth rates.

Seasonal and spatial variability in rates of primary production and detritus release by intertidal stands of Laminaria digitata and Saccharina latissima on wave-exposed shores in the northeast Atlantic.

Coastal habitats are increasingly recognized as fundamentally important components of global carbon cycles, but the rates of carbon flow associated with marine macrophytes are not well resolved for many species in many regions. We quantified density, rates of primary productivity, and detritus production of intertidal stands of two common intertidal kelp species-Laminaria digitata (oarweed) and Saccharina latissima (sugar kelp)-on four NE Atlantic rocky shores over 22 months. The density of L. digitata was greater at exposed compared to moderately exposed shores but remained consistently low for S. latissima throughout the survey period. Individual productivity and erosion rates of L. digitata did not differ between exposed and moderately exposed shores but differed across exposure levels throughout the year at moderately exposed sites only. Productivity and erosion of S. latissima remained low on moderately exposed shores and showed no clear seasonal pattern. Patterns of productivity and total detrital production (erosion and dislodgement) per m2 of both L. digitata and S. latissima followed closely that of densities per m2, peaking in May during both survey years. Temperature and light were key factors affecting the productivity rates of L. digitata and S. latissima. Erosion rates of L. digitata were affected by wave exposure, temperature, light, grazing, and epiphyte cover, but only temperature-affected erosion of S. latissima. Production of biomass and detritus was greater in L. digitata than in S. latissima and exceeded previous estimates for subtidal and warmer-water affinity kelp populations (e.g., Laminaria ochroleuca). These biogenic habitats are clearly important contributors to the coastal carbon cycle that have been overlooked previously and should be included in future ecosystem models. Further work is required to determine the areal extent of kelp stands in intertidal and shallow subtidal habitats, which is needed to scale up local production estimates to entire coastlines.

Marine heat waves differentially affect functioning of native (Ostrea edulis) and invasive (Crassostrea [Magallana] gigas) oysters in tidal pools.

The frequency and duration of short-term extreme climatic events, such as marine heat waves (MHWs), are increasing worldwide. The rapid onset of MHWs can lead to short-term stress responses in organisms that may have lethal or sub-lethal effects. In addition, increased temperature variability and extremes are predicted to favour and facilitate the spread of non-native species, altering rates of key ecosystem processes and functions. It is possible, however, that compensatory mechanisms, such as increased feeding rates, may enable the maintenance of metabolic functioning and prevent detrimental temperature effects. Using a mesocosm-based approach, we experimentally tested for the effects of MHWs in tidal pools on the mortality, individual length, width and biomass, and respiration rates for both a native oyster, Ostrea edulis, and invasive oyster, Magallana gigas, with or without food supply. No mortality was recorded for either O. edulis or M. gigas for the duration of the four week experiment. Increases in length were greater in O. edulis compared to M. gigas but were not affected by temperature or food supply. Increases in width, however, did not differ between species but were reduced overall in heat wave treatments regardless of food supply. O. edulis gained more biomass than M. gigas in ambient treatments regardless of food supply but, in heat wave treatments, only gained greater biomass than M. gigas at additional levels of food supply. Respiration rates did not reflect changes in temperature or food supply in either species but differed through time, with greater rates post-heat wave in all treatments. Thermal responses of O. edulis and M. gigas to MHWs thus appear to be context dependent and, if food supply is sufficient, O. edulis may be able to maintain its presence in the intertidal. The ability of M. gigas to remain unaffected by fluctuating environmental conditions, however, suggests future resilience of invasive populations to climatic extremes that may result in competitive exclusion and a further decline in native oyster populations. This information is critical for developing effective management plans to ensure the sustainability of natural oyster populations whilst maintaining key ecosystem functioning.

Ocean warming and species range shifts affect rates of ecosystem functioning by altering consumer-resource interactions.

Recent warming trends have driven widespread changes in the performance and distribution of species in many regions, with consequent shifts in assemblage structure and ecosystem functioning. However, as responses to warming vary across species and regions, novel communities are emerging, particularly where warm-affinity range-expanding species have rapidly colonized communities still dominated by cold-affinity species. Such community reconfiguration may alter core ecosystem processes, such as productivity or nutrient cycling, yet it remains unclear whether novel communities function similarly to those they have replaced, and how continued warming will alter functioning in the near future. Using simplified kelp forest communities as a model system, we compared rates of respiration, consumption and secondary productivity between current cold-affinity and future warm-affinity kelp assemblages under both present-day temperatures and near-future warming in a series of mesocosm experiments. Overall, respiration rates of gastropods and amphipods increased with warming but did not differ between cold and warm affinity kelp assemblages. Consumption rates of three consumers (urchin, gastropod and amphipod) differed between kelp assemblages but only amphipod consumption rates increased with warming. A diet derived from warm-affinity kelp assemblages led to a decrease in growth and biomass of urchins, whereas the response of other consumers was variable depending on temperature treatment. These results suggest that climate-driven changes in assemblage structure of primary producers will alter per capita rates of ecosystem functioning, and that specific responses may vary in complex and unpredictable ways, with some mediated by warming more than others. Understanding how differences in life history and functional traits of dominant species will affect ecological interactions and, in turn, important ecosystem processes is crucial to understanding the wider implications of climate-driven community reconfiguration.

Habitat modification by Ascophyllum canopy negatively impacts macrofaunal communities on soft-sediment shores.

Canopy-forming macroalgae are known to act as ecosystem engineers, altering the physical parameters of the local environment, and as a result, driving changes in local biodiversity. Although a large body of evidence exists regarding macroalgal canopies on intertidal rocky shores, little is known regarding attached perennial species in soft sediment environments. The aim of this study was to assess whether the presence of an Ascophyllum nodosum canopy altered physical parameters, leading to the formation of different environmental conditions in the areas around the canopy and whether this led to changes in the local community. Sediment cores were taken in canopy-present and canopy-absent treatments at four sites over four sampling periods covering winter (November and January) through to spring (March and May) to assess modification of seven physical parameters: particle size, sand/silt/clay content, chlorophyll a, organic carbon, pore water content and temperature, as well as for macrofaunal diversity. Results revealed significant differences between treatments for all variables with the exception of clay content. Areas below the canopy were dominated by a high abundance of opportunistic species indicating a more disturbed environment, with increased levels of organic enrichment, anoxia and scouring found to be the principal sources of physical disturbance. In conclusion, differences in abiotic parameters between canopy and non-canopy areas in soft-sediment environments were driven both directly and indirectly by the presence of the algal canopy. This facilitated an alternative community composition that enhanced biodiversity within algal-sediment shores.