Wave Exposure Gradient Research Articles

Carbon stocks in Norwegian eelgrass meadows across environmental gradients

Seagrass meadows are well-known for their capacity to capture and store blue carbon in sediments. However carbon stocks vary significantly between meadows, spanning more than three orders of magnitude on both local and global scales. Understanding the drivers of seagrass carbon stocks could help improve strategies for incorporating blue carbon into management plans. Here, we measured sediment carbon stocks in eelgrass (Zostera marina) meadows and unvegetated areas along the Norwegian coast, spanning wide gradients in temperature, wave exposure, water depth, salinity, and eelgrass biomass. Carbon stocks were generally higher in eelgrass meadows than in adjacent unvegetated areas, yet they displayed considerable variation (400 − 30 000 g C m−2 at 50 cm sediment depth) even among nearby sites. Overall, the highest carbon stocks were found in deeper, muddier, sheltered meadows near river mouths. These sites likely have the highest input and retention of carbon from different sources. Consequently, they should be prioritized as conservation targets for preserving coastal blue carbon stocks. Despite ever-increasing efforts to quantify seagrass blue carbon globally, high uncertainties still persist, partly due to differing methodologies, processes, and environmental context. Blue carbon stock estimates could be improved through the coordination of standardised mapping and sampling methods.

Long‐Term Stability of Marine Forests Facing Moderate Gradual Warming in a Remote Biodiversity Hotspot

ABSTRACTAimOcean warming and marine heatwaves are rapidly reconfiguring the composition of seaweed forests—the world's largest coastal vegetated biome. Seaweed forest responses to climate change in remote locations, which constitute the majority of the forest biome, remain however poorly quantified. Here, we examine the temporal stability of the seaweed forests across a global seaweed biodiversity hotspot where several species are predicted to undergo severe range contractions in this century.LocationWestern south coast of Australia.MethodsSeaweed forest canopies were censused at 18 shallow (< 10 m) sheltered reefs between 1997 and 2006 and again between 2021 and 2024 (six sites per location). We also surveyed 24 sites to examine whether temporal changes differed over gradients of wave exposure and depth.ResultsSeaweed forest canopies across all locations showed surprisingly little change in biomass, cover, stand density and species composition over two decades, with strong spatial structuring across depth and exposure gradients persisting over time. The average thermal affinity of forest canopies (i.e., the community temperature index, CTI) did not track warming, suggesting that factors other than temperature (e.g., wave exposure and depth) are more important drivers of forest stand structure and/or that key thermal thresholds have not yet been crossed. Forests in the location with the most pronounced warming exhibited increased thermal bias over time (total bias of 0.8°C–2.2°C), indicating they were dominated by species with cooler affinities than their local temperatures.Main ConclusionsThe greater thermal bias in forests at the warmer edge of southern Australia suggests these will be more susceptible to future warming‐related compositional changes than forests in cooler locations. The relative stability we found contrasts with a current context of rapidly changing seaweed forests nationally and globally, highlighting the need to deepen our ecological understanding of the region so that future changes to its unique biodiversity and ecosystem services can be predicted and mitigated.

Notes from the past show how local variability can stymie urchins and the rise of the reds in the Gulf of Maine

AbstractThe impacts of global change—from shifts in climate to overfishing to land use change—can depend heavily on local abiotic context. Building an understanding of how to downscale global change scenarios to local impacts is often difficult, however, and requires historical data across large gradients of variability. Such data are often not available—particularly in peer reviewed or gray literature. However, these data can sometimes be gleaned from casual records of natural history—field notebooks, data sheet marginalia, course notes, and more. Here, we provide an example of one such approach for the Gulf of Maine, as we seek to understand how environmental context can influence local outcomes of region‐wide shifts in subtidal community structure. We explore a decade of hand‐drawn algal cover maps around Appledore Island made by Dr. Art Borror while teaching at the Shoals Marine Lab. Appledore's steep wave exposure gradient—from exposed to the open ocean to fully protected—provides a living laboratory to test interactions between global change and local conditions. We then recreate Borror's methods two and a half decades later. We show that overfishing‐driven urchin outbreaks in the 1980s were slowed or stopped by wave exposure and benthic topography. Similarly, local variation appears to have curtailed current invasions by filamentous red algae. Last, some formerly dominant kelps have disappeared over the past 40 years—an observation verified by subtidal surveys. Global change is altering life in the seas around us. While underutilized, solid natural history observations stand as a key resource for us to begin to understand how global change will translate to the heterogeneous mosaic of life in a future Gulf of Maine and other ecosystems around the world.

Spatiotemporal variability in population demography and morphology of the habitat-forming macroalga Saccorhiza polyschides in the Western English Channel.

Large brown macroalgae serve as foundation organisms along temperate and polar coastlines, providing a range of ecosystem services. Saccorhiza polyschides is a warm-temperate kelp-like species found in the northeast Atlantic, which is suggested to have proliferated in recent decades across the southern UK, possibly in response to increasing temperatures, physical disturbance and reduced competition. However, little is known about S. polyschides with regard to ecological functioning and population dynamics across its geographical range. Here we examined the population demography of S. polyschides populations in southwest UK, located within the species' range centre, to address a regional knowledge gap and to provide a baseline against which to detect future changes. Intertidal surveys were conducted during spring low tides at three sites along a gradient of wave exposure in Plymouth Sound (Western English Channel) over a period of 15 months. Density, cover, age, biomass and morphology of S. polyschides were quantified. Additionally, less frequent sampling of shallow subtidal reefs was conducted to compare intertidal and subtidal populations. We recorded pronounced seasonality, with fairly consistent demographic patterns across sites and depths. By late summer, S. polyschides was a dominant habitat-former on both intertidal and subtidal reefs, with maximum standing stock exceeding 13 000 g wet weight m-2. Saccorhiza polyschides is a conspicuous and abundant member of rocky reef assemblages in the region, providing complex and abundant biogenic habitat for associated organisms and high rates of primary productivity. However, its short-lived pseudo-annual life strategy is in stark contrast to dominant long-lived perennial laminarian kelps. As such, any replacement or reconfiguration of habitat-forming macroalgae due to ocean warming will probably have implications for local biodiversity and community composition. More broadly, our study demonstrates the importance of high-resolution cross-habitat surveys to generate robust baselines of kelp population demography, against which the ecological impacts of climate change and other stressors can be reliably detected.

Interactive effects of multiple stressors with significant wave height exposure on farmed Atlantic salmon (Salmo salar) welfare along an inshore-offshore gradient

Fish farming is the fastest growing food production sector worldwide and now accounts for most human fish consumption. Expansion of finfish aquaculture to exposed offshore marine environments is appealing where additional sheltered areas are unavailable. While more energetic environments may reduce waste accumulation and parasite exposure, effects on fish health and wellbeing are largely speculative. The multiple stressors faced by fish on offshore farms may interact synergistically and increase their cumulative impact. We used 20 months of health and welfare data from eight Atlantic salmon (Salmo salar) sea pen farms in Scotland along a wave exposure gradient to assess the effects of, and interactions between, environmental variables and management treatments on fish mortality and parasite loads. While farms showed high variability in mortality rate and sea lice infections, multi-level Bayesian modelling indicated that wave exposure primarily modulated effects of other variables. Higher exposure farms showed steeper increases in mortality with time and with extreme temperatures. Similarly, sea lice infections tended to increase with time, with higher exposure farms seeing steeper increases at higher Amoebic Gill Disease (AGD) scores and at high temperatures. The effect of AGD was greater at slow water speeds. Treatments against parasites were more frequent at low exposure farms, leading to uncertainty in their impact on welfare across farms. The support for interactive effects of wave exposure with other variables rather than strong direct effects suggests an accumulation of chronic and acute stressors. Expansion of aquaculture to more energetic offshore environments may have negative impacts on fish health in some circumstances, requiring adaptation of practices. In particular, the stronger increase in mortality over time may have implications for cycle length in different environments, and the more dramatic impacts of the warmest temperatures at high exposure farms call for consideration of the change in water temperature both inshore and offshore.

Local and large-scale spatial variation in a marine predator-prey interaction in the southwestern Atlantic.

Predator-prey interactions are a key ecological process which can be modified by environmental conditions over a range of spatial scales. Through two complementary short-term experiments, we assessed how local and large-scale environmental conditions affect a subtropical intertidal predator-prey interaction. At a local scale, we evaluated the effects of the degree of exposure to wave action and prey density on consumption rate and interaction strength using a whelk-barnacle system. Consumption rate decreased with wave exposure at experimentally reduced prey density but did not change at ambient density. Such an interactive effect occurred due to shifts in the whelk's feeding behaviour, likely linked to encounter rate and stress amelioration underpinned by prey density. Per capita interaction strength of the whelk on the barnacle weakened along the wave exposure gradient, but to a greater degree at reduced compared to ambient prey density. This confirms that environmental harshness can decrease the importance of predators, but the magnitude of change may be modified by density-dependent effects. A large-scale experiment did not reveal spatial patterns in the whelk-barnacle interaction, nor relationships to chlorophyll-a concentration or the minor change in sea temperature across the study area. Patterns in the size of consumed barnacles along the chlorophyll-a gradient suggest changes in food choice related to prey quality and size. We conclude that disentangling the effects of wave exposure and prey density revealed important potential mechanisms driving species locally. Large-scale variation in the whelk-barnacle interaction appeared to be linked to species' traits shaped by the environmental context.

An allozyme polymorphism is associated with a large chromosomal inversion in the marine snail Littorina fabalis.

Understanding the genetic targets of natural selection is one of the most challenging goals of population genetics. Some of the earliest candidate genes were identified from associations between allozyme allele frequencies and environmental variation. One such example is the clinal polymorphism in the arginine kinase (Ak) gene in the marine snail Littorina fabalis. While other enzyme loci do not show differences in allozyme frequencies among populations, the Ak alleles are near differential fixation across repeated wave exposure gradients in Europe. Here, we use this case to illustrate how a new sequencing toolbox can be employed to characterize the genomic architecture associated with historical candidate genes. We found that the Ak alleles differ by nine nonsynonymous substitutions, which perfectly explain the different migration patterns of the allozymes during electrophoresis. Moreover, by exploring the genomic context of the Ak gene, we found that the three main Ak alleles are located on different arrangements of a putative chromosomal inversion that reaches near fixation at the opposing ends of two transects covering a wave exposure gradient. This shows Ak is part of a large (3/4 of the chromosome) genomic block of differentiation, in which Ak is unlikely to be the only target of divergent selection. Nevertheless, the nonsynonymous substitutions among Ak alleles and the complete association of one allele with one inversion arrangement suggest that the Ak gene is a strong candidate to contribute to the adaptive significance of the inversion.

Patch dynamics driven by wave exposure in subtidal temperate seaweeds are exacerbated by warming oceans

Over the past decades, ocean temperatures have been steadily increasing and are projected to continue to do so, stressing many temperate marine organisms. Changing temperatures do not affect ecosystems in isolation, but interact with many other factors in shaping ecological communities. We investigated the changes over 2 decades in subtidal temperate seaweed communities over a wave exposure gradient in Western Australia, a global warming hotspot. We found higher diversity in the seaweed community and a higher proportion of biomass of species with a warm affinity (expressed as the tropicalization index: TI) over time. There was no decline in biomass of the dominant habitat-forming kelp Ecklonia radiata on low wave exposure reefs, while it was patchier and comprised a lower proportion of the total seaweed biomass on the medium and high wave exposure reefs. Furthermore, the presence of E. radiata was disproportionally associated with low abundances of seaweeds with warm affinity. The increasing patchiness of E. radiata likely provided a competitive release for other seaweeds, and the increase in abundance of Scytothalia dorycarpa likely provided a compensatory effect which resulted in a lower than expected TI. We found no indication of an ameliorating effect by wave exposure, and conclude that the patch dynamics driven by wave exposure are more likely exacerbated by increasing ocean temperatures on subtidal temperate reefs. If this continues, the reduction in E. radiata and increase in warm affiliated seaweeds will result in a more diverse seaweed community, but one with a lower standing biomass.

Explaining Recruitment Stochasticity at a Species’ Range Margin

Advancing our understanding of how environmental variability affects the distribution of organisms is crucial for ecology and conservation. The exploration of changes in demographic patterns close to species distribution margins is important as populations here may provide a window into future population changes also elsewhere. However, the knowledge of factors causing recruitment variation is still inadequate in many systems and this deficiency is particularly evident close to species’ distribution borders. We studied the spatiotemporal variability in recruit-adult dynamics in a blue mussel,Mytilus trossulus, population to get insights into how environmental variables drive variation in recruitment and how this variability affects adult population growth. Thirty sites along a wave exposure gradient were monitored during four consecutive years. From each site, mussels were collected both from artificial recruitment units and from natural mussel beds. Our results showed high year-to-year variation in recruitment strength with high spatial variation. Mussel recruitment to artificial units and later recruitment to the benthos correlated highly. Juvenile abundances 1 year later paralleled prior recruitment strengths and caused synchronous but time-lagged changes in adult cohorts. Seawater salinity was the strongest predictor for recruitment variation, whereas sea temperature and wave exposure had low predictive power for this early life stage. For juveniles and for adults in the benthos, wave exposure explained the variation best, whereas temperature and especially salinity explained less. The results indicate that (a) the studied blue mussel population is strongly driven by variation in recruitment strength that (b) drives the size of the later cohorts, and the population is possibly even (c) recruitment limited in some years. Our study predicts a challenging future for this range population, resulting from a higher frequency of recruitment failure caused by a deteriorating sea climate. Knowledge about factors underlying variation in recruitment is thus essential for forecasting the future of this range population and for conserving its future state.

Spatial variation in the structure of overwintering, remnant Saccorhiza polyschides sporophytes and their associated assemblages

AbstractUnderstanding the structure and richness of natural communities is a fundamental goal of marine ecology, and foundation species such as large macroalgae have a disproportionate role in structuring biodiversity. However, high-resolution information on assemblages associated with macroalgae is lacking for many species and regions. Saccorhiza polyschides is a warm-temperate kelp with a relatively short lifespan (12–18 months), large thallus and bulbous holdfast offering habitat for diverse assemblages. In the UK, S. polyschides populations are thought to have proliferated recently. Here, we quantified the density and habitat structure provided by S. polyschides along a gradient of wave exposure within Plymouth Sound, and examined the composition and diversity of associated faunal assemblages. Density varied significantly between sites but not by wave exposure, while biometric measurements were generally highly variable. Senescing holdfasts from sporophytes offered valuable habitat, with high abundance and richness of associated assemblages, although these varied markedly between sporophytes and sites. Faunal abundance, taxon richness and diversity were significantly higher at fully exposed sites than at moderately exposed sites. Internal volume of holdfasts was positively correlated with faunal abundance and taxon richness. We recorded more than 27 distinct taxa and up to ~600 individuals within a single holdfast. Taxa included three fish species, including a novel observation of the pipefish Nerophis lumbriciformis. Further work is needed to examine seasonality in habitat structure and associated diversity patterns but our study demonstrates that even remnant holdfasts from decaying sporophytes represent a valuable microhabitat that may provide shelter, protection and food during winter.

Variation in Population Structure and Standing Stocks of Kelp Along Multiple Environmental Gradients and Implications for Ecosystem Services

There is an increasing focus on ecosystem services provided by macroalgae worldwide, many which depend on its structural and morphological characteristics. Here we investigated how characteristics of canopy kelp plants of Laminaria hyperborea varies along the Norwegian part of the NE Atlantic. Ten characteristics related to size, allometry, density, biomass and epiphytes were analyzed along wide gradients of depth, wave exposure and latitude. The analyses were performed on a compiled dataset of 630 scuba samples from different research and monitoring projects along the Norwegian coast (58-71°N). The largest kelps, highest biomass, and highest biomass of epiphytic algae was found in Mid-Norway (63-65°N). While most size-related kelp characteristics were reduced with water depth, they were enlarged with wave exposure. The developed statistical models can be used to calculate site-specific values (and their variation) of the different characteristics at any location based on latitude, depth and wave exposure. These can further be used to map kelp derived ecosystem services. We predicted region-specific estimates of total plant weight, kelp density and kelp biomass. Moreover, the models were applied to estimate living biomass (56 million tonnes fresh weight), carbon standing stock (2.6 million tonnes carbon or 9.5 million tonnes CO2), and carbon sequestration potential (0.46 million tonnes per year) for the Norwegian kelp forests, that were estimated to cover about 5 355 km2. The site- and region-specific estimates are relevant to kelp forest management and for blue carbon accountings.

Global-scale species distributions predict temperature-related changes in species composition of rocky shore communities in Britain.

Changes in rocky shore community composition as responses to climatic fluctuations and anthropogenic warming can be shown by changes in average species thermal affinities. In this study, we derived thermal affinities for European Atlantic rocky intertidal species by matching their known distributions to patterns in average annual sea surface temperature. Average thermal affinities (the Community Temperature Index, CTI) tracked patterns in sea surface temperature from Portugal to Norway, but CTI for communities of macroalgae and plant species changed less than those composed of animal species. This reduced response was in line with the expectation that communities with a smaller range of thermal affinities among species would change less in composition along thermal gradients and over time. Local-scale patterns in CTI over wave exposure gradients suggested that canopy macroalgae allow species with ranges centred in cooler than local temperatures ('cold-affinity') to persist in otherwise too-warm conditions. In annual surveys of rocky shores, communities of animal species in Shetland showed a shift in dominance towards warm-affinity species ('thermophilization') with local warming from 1980 to 2018 but the community of plant and macroalgal species did not. From 2002 to 2018, communities in southwest Britain showed the reverse trend in CTI: declining average thermal affinities over a period of modest temperature decline. Despite the cooling, trends in species abundance were in line with the general mechanism of direction and magnitude of long-term trends depending on the difference between species thermal affinities and local temperatures. Cold-affinity species increased during cooling and warm-affinity ones decreased. The consistency of responses across different communities and with general expectations based on species thermal characteristics suggests strong predictive accuracy of responses of community composition to anthropogenic warming.

Applying continuous functional traits to large brown macroalgae: variation across tidal emersion and wave exposure gradients

Ecologists use a functional trait-based approach to seek a general understanding of organism—environment interactions, but, among primary producers, the empirical basis rests on vascular plants. We hypothesised that with increasing intertidal elevation, traits of large brown macroalgae would reflect a resource acquisition vs. conservation (stress tolerance) trade-off at species and community levels. Across the elevation gradient at four UK sites of varying wave exposure, we: (1) screened species’ relevant morphological traits, using principal component analysis to reduce dimensionality; and (2) up-scaled species’ traits using community weighted trait means (CWMs). The first principal component (PC1) strongly related to specific thallus area and thallus dry matter content, representing an acquisition–conservation trade-off. Although species generally shifted to the conservative end of this axis as elevation increased, mid-shore Ascophyllum nodosum sat at the extreme conservative end. PC2 related to holdfast ratio, thickness and length, with A. nodosum scoring higher than other mid-shore species. CWMs of PC1 decreased with elevation at two sites indicating a shift from ‘fast’ to ‘slow’ ecosystem functioning, but this relationship was disrupted by A. nodosum at the sheltered site, and by the up-shore extent of Laminaria digitata at the most exposed site. The anomalous traits of A. nodosum reflect its unique competitive strategy (slow, persistent growth) in the relatively stressful mid-shore. Seaweed functional traits show promise in linking species’ identities to their strategies and ecosystem contributions. However, because resource conservation traits can be related to competitive as well as stress-tolerance strategies, predicting seaweed trait responses to environmental stress gradients is challenging.

Wave stress and biotic facilitation drive community composition in a marginal hard‐bottom ecosystem

AbstractEcological patterns are inherently scale‐dependent and driven by the interplay of abiotic gradients and biotic processes. Despite the fundamental importance of such gradients, there are many gaps in our understanding of how abiotic stress gradients interplay with biotic processes and how these collectively affect species distributions. Using a hierarchical design, we sampled two communities separated by depth along wave exposure and salinity gradients to elucidate how these two gradients affect species composition in habitats formed by the foundation species Mytilus trossulus and Fucus vesiculosus. Specifically, we looked at the impacts of regional salinity and temperature, local wave exposure, and site‐dependent facilitation effects on the associated community composition. Wave exposure was the best predictor for species assembly structure, which was also affected by Mytilus biomass and by salinity and water temperature. While the tested variables provided robust explanations for community structure and density, they did not provide conclusive explanations for variation in species richness or evenness. Mytilus biomass had a stronger effect on the associated community with increasing wave exposure at the deeper depth, but the patterns were less obvious at the shallower depth. The latter was also the case for Fucus. These findings comply partly with theoretical predictions suggesting stronger facilitation effects in physically harsh environments. Our results indicate that environmental drivers are the main structuring forces that affect species assembly structure, but also foundation species are important. Thus, predicting changes in species distributions and biodiversity requires the simultaneous consideration of environmental gradients, as well as the structure and composition of foundation species and the interplay between these factors. This work advances our understanding of the processes that modulate species distributions in a marginal marine area and broadens the knowledge of how biological and environmental factors interplay and have an influence on hard‐bottom community structure in brackish water seas.

The effects of embryonic experience with predation risk vary across a wave exposure gradient

AbstractGradients in environmental stress can alter the ecological effects of predation risk to create variable landscapes of fear that shape prey antipredator responses. Prey obtain information about their risk environment not only from their immediate experiences with predators, but also from their previous experiences, especially if they occur during particularly sensitive windows in ontogeny. Embryonic development is often a key window when an individual's experiences can have lasting effects on behavior and fitness. The propensity of embryonic experiences with predation risk to affect prey performance, however, may vary across gradients of abiotic stress. Using a rocky intertidal system, we explored whether a dominant abiotic stressor—wave exposure—modifies the influence of embryonic experience with predation risk (from the green crab, Carcinus maenas) on prey (the carnivorous snail, Nucella lapillus) traits both at emergence and as one‐year‐olds exposed to current predation risk. We found that snails from wave‐exposed, but not sheltered, populations emerged smaller from development and grew less in the absence of current risk as one‐year‐olds if they experienced risk as embryos. However, exposure to current predation risk reduced the growth and growth efficiency of one‐year‐old snails from both wave‐exposed and sheltered populations. Our results demonstrate that increased environmental stress can modify predator–prey interactions and enhance prey reliance on early life experiences with predation risk, but that direct exposure to risk later in life can strongly affect prey performance across environmental stress gradients.

Untangling the contribution of genetic and environmental effects to shell differentiation across an environmental cline in a marine snail

The rough periwinkle Littorina saxatilis is characterised for showing shell phenotypic variation associated to environmental clines and this is found in parallel across shores. In order to disentangle the additive genetic effects and the environmental factors involved in this variation, we analysed natural populations subjected to a gradient in wave exposure and crab predation in 3 different Rias (primary estuaries) from Galicia (NW Spain). In addition to previously described “Crab” and “Wave” ecotypes, we studied a new phenotype (“Crab+”) associated to the most sheltered habitats of the gradient. We obtained estimates of heritability and Qst (between population genetic differentiation) in embryos and Pst (between population phenotypic differentiation) in adults for shell morphology traits. Heritability estimates were close to one for shell size and relative shell aperture, suggesting a low impact of environmental factors in embryos. Estimates of Qst between ecotypes were significant and typically much higher than published Fst estimates from neutral markers, indicating a role for selection on shaping additive genetic variation. Estimates of Pst between ecotypes for adults were substantially larger than embryos´ Qst, suggesting also a role of phenotypic plasticity in ecotype differentiation. Our results suggest that Pstvs Fst comparisons should be taken with caution because they might not reflect real additive genetic effects, but Pstvs Qst comparisons could shed light into the role of phenotypic plasticity.

Seascapes as drivers of herbivore assemblages in coral reef ecosystems

AbstractNotionally herbivorous fishes maintains a critical ecosystem function on coral reefs by grazing algae and maintaining highly productive algal turf assemblages. Current paradigms implicate habitat complexity, predation, and primary productivity as major drivers of the distribution and abundance of herbivorous fish, yet little is known about the relative contribution of these factors. Here, we compare bottom‐up and top‐down drivers of notional herbivore assemblages across an environmental gradient of wave exposure in the Palau archipelago. We surveyed herbivore assemblages at reef slopes (6–9 m) across 18 sites, and quantified proxies of top‐down control (predator biomass, habitat complexity) and bottom‐up drivers (net primary production, nutrients) at each site. Despite substantial variability in herbivore biomass throughout the archipelago (6–65 g/m2), general additive models indicate that neither top‐down nor bottom‐up drivers significantly predicted biomass or density of herbivores among sites. In contrast to expectations, herbivore biomass was highest at sites with high predator biomass, low structural complexity, and low benthic productivity. Rather, the highest biomass of herbivores was associated with shallow, tidally emergent, productive reef flats located adjacent to steep vertical walls (“drop‐offs”). The emergent nature of this neighboring habitat precluded occupation by territorial fishes and multiple species of herbivores were observed to make foraging runs into this habitat once tidally inundated. We hypothesize that this habitat configuration provides an important cross‐habitat resource subsidy. Multivariate ordination and permutation of herbivore communities revealed strong evidence for biogeographic partitioning throughout the archipelago (western, southwestern, inner eastern, and outer eastern clusters), contributing to an emerging picture that the habitat heterogeneity of seascapes can overwhelm the effects of conventional top‐down and bottom‐up structuring of herbivory on coral reefs.

Influence of wave climate on architecture and landscape characteristics of Posidonia oceanica meadows

AbstractSeagrass meadow characteristics, including distribution, shape, size and within‐meadow architectural features, may be influenced by various physical factors, including hydrodynamic forces. However, such influences have hardly been assessed for meadows of the ecologically important and endemic Mediterranean seagrass Posidonia oceanica. The distribution of P. oceanica meadows at five sites in the Maltese Islands was mapped to a depth of c. 15 m using a combination of aerial photography and SCUBA diving surveys. Estimates of wind‐generated wave energy and energy attenuated by depth were computed using the hydrodynamic model WEMo (Wave Exposure Model). Metrics for P. oceanica landscape features were calculated using FRAGSTATS for replicate 2500 m2 subsamples taken from the seagrass habitat maps in order to explore the influence of wave dynamics at the landscape scale. Data on within‐meadow architectural attributes were collected from five sites and analysed for relationships with wave energy. The results indicate that landscape and architectural features of P. oceanica meadows located within the 6–11 m depth range are significantly influenced by wave climate. Posidonia oceanica meadows tend to be patchier and have low overall cover, more complex patch shapes and reduced within‐patch architectural complexity along a wave exposure gradient from low to high energy. The findings from the present study provide new insight into the influence of hydrodynamic factors on the natural dynamism of P. oceanica meadow landscape and architecture, which has implications for the conservation and management of the habitat.

Distribution of macroalgae in Kongsfjorden, Svalbard

Macroalgal distribution varies with prevailing environmental gradients in fjords, particularly in glacial fjords such as Kongsfjorden, Svalbard. This survey by scuba-diving in 1996 and 1998 was the first to determine species composition, dominance and distribution of macroalgae along the entire length axis of a fjord in the Arctic. Macroalgal distribution was determined quantitatively by frame sampling and with digital video along transects at five hard-bottom locations from outer to inner fjord and interpreted against environmental gradients of depth (0–30 m), substratum, wave exposure and distance from inner glacial front. Most species identified (>70%) belonged to the Arctic and Arctic-temperate biogeographic groups, often with limited distribution to inner or outer part of the fjord. Dominant species or species with wide geographic distribution were typically widely distributed along the fjord axis. Peaks in biomass varied from 5 to 10 m at the middle-to-outer fjord, whereas in the inner bay it peaked at <5 m. The biomass distribution of macroalgae was mainly related to depth and distance from the inner glacial bay, and varied with wave exposure and sediment cover. Climate-related changes, after our study, indicate that the macroalgal belt has become shallower at Hansneset in middle fjord, with increased biomass in the upper sublittoral zone because of higher temperature and less sea ice. Distribution and production of macroalgae and fauna in shallow regions of the Arctic are likely to increase in the future, particularly in areas subjected to ocean warming, such as Kongsfjorden and other fjords and coastal areas in Svalbard.

Dislodgement force and shell morphology vary according to wave exposure in a tropical gastropod (Cittarium pica)

ABSTRACTWave exposure has strong influences on population density, morphology and behaviour of intertidal species in temperate zones, but little is known about how intertidal organisms in tropical regions respond to gradients in wave exposure. We tested whether dislodgement force and shell shape of a tropical gastropod, Cittarium pica, differs among shores that vary in wave exposure. After adjusting for body size, we found that C. pica from exposed shores required greater dislodgement force to remove them from the shore, had slightly larger opercula (the closure to the shell aperture), and were slightly squatter in shape (reduced in shell height relative to shell width) than C. pica from sheltered shores. These morphological adjustments are consistent with those observed in temperate gastropods, which are argued to represent adaptive responses to the risk of mortality associated with dislodgement.