Do sulphuric acid and the brown alga <Emphasis Type="Italic">Desmarestia viridis</Emphasis> support community structure in Arctic kelp patches by altering grazing impact,distribution patterns,and behaviour of sea urchins?

Macrobenthic community structure and the distribution of the green sea urchin (Strongylocentrotus droebachiensis) were recorded inside and outside (=barrens) of kelp patches (Alaria esculenta) at Kongsfjordneset, Svalbard between August 2002 and October 2006. In manipulative field experiments, conducted at Kongsfjordneset, Svalbard in August 2002, the effect of the presence of the brown seaweed Desmarestia viridis on sea urchin distribution and kelp grazing was determined. Additionally, we studied the effect of sulphuric acid, which is produced and stored by D. viridis, on sea urchin movements in the laboratory at Ny-Ålesund, Svalbard, in May 2006. Sea urchin densities were two- to threefold lower in kelp patches than on barrens. The macrobenthic community inside kelp patches hosted 39% more species and was of different species composition than on barrens. Anchored pieces of the kelp A. esculenta were less consumed when surrounded by D. viridis than non-surrounded conspecifics. Changes in pH affected the behaviour of sea urchins. Exposing sea urchins to 500 μl seawater at pH 7.5 caused them to stop, while the exposure of as little as 25 μl at pH 1 caused sea urchins to move in the opposite direction. Acid-mediated escape responses in sea urchin behaviour suggest chemical protection by D. viridis as an additional mechanism to mechanical protection in the generation of kelp refuges. These results improve our understanding of how isolated kelp beds can persist over a wide range of environmental conditions, like wave-sheltered sites, and suggest that changes in community structure may be in part attributable to altered trophic interactions.     

Spatial variation and effects of habitat on temperate reef fish assemblages in northeastern New Zealand

Reef-associated fishes can respond to changes in habitat structure and the nature of their response can change with different spatial scales of observation. A structured hierarchical mensurative sampling design was used to sample temperate reef fish assemblages in northeastern New Zealand at several spatial scales over 2 years. The three spatial scales examined were tens of meters (transects), hundreds to thousands of meters (sites) and hundreds of kilometers (locations). We tested the hypothesis that fish assemblages differed between kelp forest habitat (relatively dense stands of the kelp, Ecklonia radiata (C. Agardh) J. Agardh, median depth=13.5 m) and barrens habitat (rocky reef dominated by turfing and encrusting red algae and the grazing urchin, Evechinus chloroticus (Valenciennes), median depth=6.7 m). Recently developed multivariate techniques were used to test for and quantify multivariate variation at different spatial scales. There were significant effects of habitat on the spatial distribution of fish assemblages, characterised by greater abundances or frequencies of Parika scaber, Chromis dispilus, Trachurus novaezelandiae, Nemadactylus douglasii, Bodianus unimaculatus, Odax pullus and Pseudolabrus miles in kelp forest habitat, and greater abundances or frequencies of Notolabrus celidotus, Notolabrus fucicola, Girella tricuspidata, Coris sandageri, Chironemus marmoratus, Parma alboscapularis, Scorpis violaceus and Kyphosus sydneyanus in barrens habitat. Some of the more common species, including Upeneichthys lineatus, Scorpis lineolatus and Cheilodactylus spectabilis showed no strong consistent effects of these two differing habitats on their distributions. There was, however, a significant Habitat×Locations interaction: effects of habitat did not occur at all locations. Variability was highest at the scale of individual transects and variability from site to site and from location to location was comparable. Spatial variation was large compared to inter-annual variation, which was minimal, and spatial patterns were consistent in the 2 years examined. Further experiments, including manipulations, are required to understand what mechanisms and processes might be driving these patterns. This study, coupled with results from previous studies, suggests that there may be a dynamic inter-play between effects of habitat on fish and effects of fish on biogenic habitat, such as kelp forests.     

The effects of an invasive habitat modifier on the biotic interactions between two native herbivorous species and benthic habitat in a subtidal rocky reef ecosystem

Range expanding species can have major impacts on marine ecosystems but experimental field based studies are often lacking. The urchin Centrostephanus rodgersii has recently undergone a southerly range expansion to the east coast of Tasmania, Australia. We manipulated densities of C. rodgersii and algal regrowth in urchin barrens habitat to test effects of the urchin on biotic interactions between two native herbivores, black-lip abalone (Haliotis rubra) and another urchin (Heliocidaris erythrogramma), and their benthic habitat. After 13 months, removals of only C. rodgersii resulted in overgrowth of barrens habitat by algae and sessile invertebrates. Densities of abalone increased (+92 %) only in patches from which C. rodgersii was removed and algal regrowth allowed. In contrast, densities of H. erythrogramma increased in all treatments (+45, +28, +25 %) in which C. rodgersii was removed, irrespective of the algal regrowth manipulations. These results suggest that C. rodgersii has a negative influence on the densities of abalone through competition for food and on densities of H. erythrogramma through competition for preferred habitat. Densities of abalone (+65 %) but not H. erythrogramma (+25 %), were lower in the patches from which C. rodgersii and canopy algae regrowth were removed relative to patches from which only C. rodgersii was removed (+92 and +28 %, respectively). These results suggest that C. rodgersii overgrazing of canopy-algae results in loss of structural complexity which could increase abalone susceptibility to predation, cause abalone to seek shelter in cryptic microhabitats and/or prevent their return to patches where canopy algae are absent. The ongoing spread of C. rodgersii and expansion of barrens habitat in eastern Tasmania will continue to negatively affect populations of these two native herbivores and their associated fisheries at a range of spatial scales. This example shows that habitat modifying species which become highly invasive can have disproportionate negative impacts on the structure and dynamics of the recipient community.     

Long Term Variability in the Structure of Kelp Communities in Northern Chile and the 1997–98 ENSO

This is the first study on the south eastern Pacific coast of South America which details long term, interannual variability in the structure of subtidal rocky-bottom kelp-dominated communities before, during, and after the El Niño Southern Oscillation (ENSO) event of 1997–1998 in northern Chile (23^°S). The temporal patterns of the main components of these ecosystems, which included Macrocystis integrifolia, Lessonia trabeculata, echinoids and asteroids, were evaluated seasonally between 1996 and 2004. M. integrifolia demonstrated high interannual variability in temporal patterns of abundance. The 1997–1998 ENSO did not significantly modify the temporal patterns of Macrocystis, although local extinction of M. integrifolia beds occurred during negative thermal anomalies in 1999–2000 (La Niña event), facilitating the establishment of urchin dominated “barren grounds”. The abundance of Lessonia trabeculata showed little temporal variability, and this species dominated the deeper regions of the kelp assemblage (8–13 m depth). The structure of the kelp communities in the study area is regulated by a trophic cascade which modulates alternation between kelp dominated areas and sea urchin barrens. In this context, frequent and intense upwelling of cold water high in nutrients favors the establishment and persistence of kelp assemblages. During ENSO, coastal upwellings can mitigate superficial warming of coastal water and increase the nutrient concentration in the water column. Superficial warming during the 1997–1998 ENSO induced spawning by different species of echinoderms, which resulted in major recruitment of these species during 1999. Top-down events, such as the decrease in densities of the asteroids after the 1997–1998 ENSO event, favored increases in densities of benthic grazers, which caused significant decreases in abundance of M. integrifolia. The re-establishment of the adult fraction of the carnivore (starfish) guild coincided with a decrease in the density of sea urchins and thus re-establishment of the kelp. In the temperate south eastern Pacific, oceanographic events, which act on different spatial-temporal scales, trigger trophic cascades that act at local levels, producing interannual variability in the structure of kelp communities. On the other hand, considering the high macroinvertebrate diversity associated with kelp assemblages, the transitions between kelp-dominated areas and sea urchin barrens do not appear to significantly affect the biodiversity of these assemblages of benthic invertebrates.     

Spatial and temporal variability among marine Bacteroidetes populations in the NW Mediterranean Sea

The abundance and structure of Bacteroidetes populations at diverse temporal and spatial scales were investigated in the Northwestern Mediterranean Sea. At a temporal scale, their relative abundance exhibited a marked seasonality, since it was higher in spring and decreased in winter. Similarly, Bacteroidetes community structure encompassed three main groups (winter, spring and summer-fall), which mimicked global bacterioplankton seasonality. At the spatial scale, relative abundances were similar in all surface samples along an inshore–offshore transect, but they decreased with depth. Analysis of the community structure identified four markedly different groups mostly related to different depths. Interestingly, seasonal changes in abundance and community structure were not synchronized. Furthermore, richness was higher when Bacteroidetes were less abundant. The variability of Bacteroidetes contributions to community structure in the temporal and spatial scales was correlated with different environmental factors: day length was the most important factor at the temporal scale, and salinity at the spatial scale. The community composition in terms of phylotypes changed significantly over time and along the depth gradients, but season or depth-specific phylogenetic clusters were not identified. Delineation of coherent Bacteroidetes sub-clusters should help to uncover higher resolution patterns within Bacteroidetes, and explore associations with environmental and biological variables.     

A COMPARISON OF DRAGON KELP,EUALARIA FISTULOSA, (PHAEOPHYCEAE) FECUNDITY IN URCHIN BARRENS AND NEARBY KELP BEDS THROUGHOUT THE ALEUTIAN ARCHIPELAGO1

The Aleutian Archipelago coastal ecosystem has undergone a dramatic change in community composition during the past two decades. Following the removal of ～99% of the sea otters, Enhydra lutris, from the ecosystem, changes to the benthic communities resulted in widespread losses to most of the region’s kelp beds and corresponding increases in the prevalence of urchin barrens. Within the urchin barrens, the few kelps that have remained are exposed to elevated light, nutrients and currents, all of which may enhance their physiological condition and thus result in greater fecundity. To explore this further, we examined patterns of sporophyte fecundity in the dominant canopy‐forming kelp, Eualaria fistulosa, in both urchin barrens and in nearby kelp beds at seven Aleutian Islands spanning a range of 800 km. We found that the average weight of E. fistulosa sporophyll bundles was significantly greater on sporophytes occurring in the urchin barrens than in the nearby kelp beds. Furthermore, the average number of zoospores released per cm² of sporophyll area was also significantly greater in individuals from the urchin barrens than the nearby kelp beds. When these two metrics were combined, our results suggest that individual E. fistulosa sporophytes occurring in the urchin barrens may produce as many as three times more zoospores than individual E. fistulosa sporophytes occurring in the nearby kelp beds, and thus they may contribute disproportionately to the following year’s sporophyte recruitment in both urchin barrens and the adjacent kelp beds.     

Predation risk influences feeding rates but competition structures space use for a common Pacific parrotfish

Not all hosts, communities or environments are equally hospitable for parasites. Direct and indirect interactions between parasites and their predators, competitors and the environment can influence variability in host exposure, susceptibility and subsequent infection, and these influences may vary across spatial scales. To determine the relative influences of abiotic, biotic and host characteristics on probability of infection across both local and estuary scales, we surveyed the oyster reef-dwelling mud crab Eurypanopeus depressus and its parasite Loxothylacus panopaei, an invasive castrating rhizocephalan, in a hierarchical design across >900 km of the southeastern USA. We quantified the density of hosts, predators of the parasite and host, the host’s oyster reef habitat, and environmental variables that might affect the parasite either directly or indirectly on oyster reefs within 10 estuaries throughout this biogeographic range. Our analyses revealed that both between and within estuary-scale variation and host characteristics influenced L. panopaei prevalence. Several additional biotic and abiotic factors were positive predictors of infection, including predator abundance and the depth of water inundation over reefs at high tide. We demonstrate that in addition to host characteristics, biotic and abiotic community-level variables both serve as large-scale indicators of parasite dynamics.     

Random movement pattern of the sea urchin Strongylocentrotus droebachiensis

We describe the fine-scale movement of the sea urchin Strongylocentrotus droebachiensis based on analyses of video recordings of undisturbed individuals in the two habitats which mainly differed in food availability, urchin barrens and grazing front. Urchin activity decreased as urchin density increased. Individuals alternated between moving and being stationary and their behaviour did not appear to be affected by either current velocity (within the range from 0 to 15 cm s^− 1) and temperature (2.3 to 6.0 °C). Movement of individuals at each location was compared to that predicted by a random walk model. Mean move length (linear distance between two stationary periods), turning angle and net squared displacement were calculated for each individual. The distribution of turning angles was uniform at each location and there was no evidence of a relationship between urchin density and either move length or urchin velocity. The random model predicted a higher dispersal rate at locations with low urchin densities, such as barrens habitats. However, the movement was sometimes greater or less than predicted by the model, suggesting the influence of local environmental factors. The deviation of individual paths from the model revealed that urchins can be stationary or adopt a local (displacement less than random), random or directional movement. The net daily distance displaced on the barrens, predicted by a random walk model, was similar to the observed movement recorded in our previous study of tagged urchins at one site, but less than that observed at a second site. We postulate that the random dispersal of urchins allows individuals on barrens to reach the kelp zone where food is more abundant although the time required to reach the kelp zone may be considerable (months to years). Urchins decrease their rate of dispersal once they reach the kelp zone so that they likely remain close to this abundant food sources for long periods.     

Global regime shift dynamics of catastrophic sea urchin overgrazing

A pronounced, widespread and persistent regime shift among marine ecosystems is observable on temperate rocky reefs as a result of sea urchin overgrazing. Here, we empirically define regime-shift dynamics for this grazing system which transitions between productive macroalgal beds and impoverished urchin barrens. Catastrophic in nature, urchin overgrazing in a well-studied Australian system demonstrates a discontinuous regime shift, which is of particular management concern as recovery of desirable macroalgal beds requires reducing grazers to well below the initial threshold of overgrazing. Generality of this regime-shift dynamic is explored across 13 rocky reef systems (spanning 11 different regions from both hemispheres) by compiling available survey data (totalling 10 901 quadrats surveyed in situ) plus experimental regime-shift responses (observed during a total of 57 in situ manipulations). The emergent and globally coherent pattern shows urchin grazing to cause a discontinuous ‘catastrophic’ regime shift, with hysteresis effect of approximately one order of magnitude in urchin biomass between critical thresholds of overgrazing and recovery. Different life-history traits appear to create asymmetry in the pace of overgrazing versus recovery. Once shifted, strong feedback mechanisms provide resilience for each alternative state thus defining the catastrophic nature of this regime shift. Importantly, human-derived stressors can act to erode resilience of desirable macroalgal beds while strengthening resilience of urchin barrens, thus exacerbating the risk, spatial extent and irreversibility of an unwanted regime shift for marine ecosystems.     

Changing Community States in the Gulf of Maine: Synergism Between Invaders, Overfishing and Climate Change

Human activities, including overfishing and species introductions, have had a dramatic impact on benthic communities in the Gulf of Maine within the past two decades. Prior to the 1970s, the climax community in the shallow subtidal was composed of Laminaria spp. kelp beds with an understory of arborescent red algae. In the 1980s, a population explosion of the green sea urchin, Strongylocentrotus droebachiensis, created an alternate community state, urchin barrens. Recently, a new community has been observed in former urchin barrens and kelp beds. This assemblage is principally composed of the introduced species: Codium fragile subsp. tomentosoides (green alga), Membranipora membranacea (bryozoan), Diplosoma listerianum (tunicate), Bonnemaisonia hamifera (red alga) and the opportunistic species Mytilus edulis (mussel) and Desmarestia aculeata (brown alga). In addition to changes in relative abundance, many of these species have greatly expanded their distribution and habitat selection. A model detailing mechanisms for the transition of the traditional kelp bed and urchin barren communities to others is presented and implications for this new community are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Distribution and biohistory of the endemic flora of the mid-Appalachian shale barrens

The mid-Appalachian shale barrens, first described by E. S. Steele in 1911, range within the eastern United States from south-central Pennsylvania to southwestern Virginia and adjacent West Virginia. These Paleozoic shaly outcrops are characterized by a steep southern exposure, typically an undercutting stream at the base, and a relatively sparse vegetation cover in contrast to the surrounding woodlands of the eastern deciduous forest. Lithologically, the barrens have thin, fissile, variously colored shale and siltstone fragments intermixed with sandstones and minor conglomerates. At least some of the barrens were colonizable sites since the late Tertiary. The sparsity of herbaceous vegetation is due chiefly to high insolation temperatures and low moisture conditions at the surface of the barrens, two physical parameters which eliminate most young seedlings. In addition to adapting to these severe constraints, the shale barren endemics require adequate root space, and, as obligate heliophytes, high sunlight intensity. Eighteen taxa are recognized as endemic to the shale barren region. Two (Phlox buckleyi, Trifolium virginicum) are paleoendemics, seven (Allium oxyphilum, Arabis serotina, Astragalus distortus var.distortus, Aster schistosus, Clematis coactilis, C. viticaulis, Solidago arguta var.harrisii) are neoschizoendemics with restricted ranges, six (Calystegia spithamaea ssp.purshiana, Clematis albicoma, Eriogonum allenii, Paronychia montana, Pseudotaenidia montana, Senecio antennariifolius) are holoschizoendemics, two (Antennaria virginica, Oenothera argillicola) are patroendemics which are ancestral to two or more species, and one (Helianthus laevigatus) is an apoendemic polyploid derivative. Possibly as many as six taxa (Antennaria virginica, Astragalus distortus var.distortus, Clematis albicoma, Eriogonum allenii, Oenothera argillicola, Senecio antennariifolius) are disjunct endemics with the vicariad in mid-central to western North America. The eighteen endemics are not equally distributed throughout the shale barren region, a fact which may reflect unequal evolutionary age, dissemination, and breeding structure. In general, speciation of the endemics appears to be largely an initial migration (from the southwestern United States) and geographic isolation of diploid populations or ecotypic differentiation of subsets of adjacent species within the Appalachian forests. Conservation of these endemics is strongly recommended.     

Communities in high definition: Spatial and environmental factors shape the micro-distribution of aquatic invertebrates

1. According to metacommunity theories, the structure of natural communities is the result of both environmental filtering and spatial processes, with their relative importance depending on factors including local habitat characteristics, functional features of organisms, and the spatial scale considered. However, few studies have explored environmental and spatial processes in riverine systems at local scales, explicitly incorporating spatial coordinates into multi-taxa distribution models. To address this gap, we conducted a small-scale study to discriminate between abiotic and biotic factors affecting the distribution of aquatic macroinvertebrates, applying metacommunity concepts.
2. We studied a mountain section in each of three perennial streams within the Po River Basin (northern Italy). We sampled macroinvertebrates both in summer and winter, using specific in situ 50-point random sampling grids. Environmental factors, including benthic organic matter (BOM), flow velocity, water depth, and substrate were recorded together with spatial coordinates for each sampling point. The relationships between community metrics (taxon richness, abundance, biomass, biomass–abundance ratio, and functional feeding groups) and explanatory variables (environmental and spatial) were assessed using generalised additive models. The influence of the explanatory variables on community structure was analysed with joint species distribution models.
3. Environmental variables—primarily BOM—were the main drivers affecting community metrics, whereas the effects of spatial variables varied among metrics, streams, and seasons. During summer, community structure was strongly affected by BOM and spatial position within the riverbed, the latter probably being a proxy for mass effects mediated by biotic and stochastic processes. In contrast, community structure was mainly shaped by hydraulic variables in winter.
4. Using macroinvertebrate communities as a model group, our results demonstrate that metacommunity concepts can explain small-scale variability in community structure. We found that both environmental filtering and biotic processes shape local communities, with the strength of these drivers depending on the season. These insights provide baseline knowledge that informs our understanding of ecological responses to environmental variability in contexts including restoration ecology, habitat suitability modelling, and biomonitoring.

     

Marine reserves demonstrate top-down control of community structure on temperate reefs

Replicated ecological studies in marine reserves and associated unprotected areas are valuable in examining top-down impacts on communities and the ecosystem-level effects of fishing. We carried out experimental studies in two temperate marine reserves to examine these top-down influences on shallow subtidal reef communities in northeastern New Zealand. Both reserves examined are known to support high densities of predators and tethering experiments showed that the chance of predation on the dominant sea urchin, Evechinus chloroticus, within both reserves was approximately 7 times higher relative to outside. Predation was most intense on the smallest size class (30-40 mm) of tethered urchins, the size at which urchins cease to exhibit cryptic behaviour. A high proportion of predation on large urchins could be attributed to the spiny lobster, Jasus edwardsii. Predation on the smaller classes was probably by both lobsters and predatory fish, predominantly the sparid Pagrus auratus. The density of adult Evechinus actively grazing the substratum in the urchin barrens habitat was found to be significantly lower at marine reserve sites (2.2ǂ.3 m^-2) relative to non-reserve sites (5.5ǂ.4 m^-2). There was no difference in the density of cryptic juveniles between reserve and non-reserve sites. Reserve populations were more bimodal, with urchins between 40 and 55 mm occurring at very low numbers. Experimental removal of Evechinus from the urchin barrens habitat over 12 months lead to a change from a crustose coralline algal habitat to a macroalgal dominated habitat. Such macroalgal habitats were found to be more extensive in both reserves, where urchin densities were lower, relative to the adjacent unprotected areas that were dominated by urchin barrens. The patterns observed provide evidence for a top-down role of predators in structuring shallow reef communities in northeastern New Zealand and demonstrate how marine reserves can reverse the indirect effects of fishing and re-establish community-level trophic cascades.     

Herbivory on coral reefs: community structure following mass mortalities of sea urchins

The community structure of Jamaican coral reefs has undergone drastic change since mass mortalities of the long-spined black sea urchin Diadema antillarum Philippi occurred in 1983. In the absence of Diadema, algal abundance has increased enormously, up to a mean of 95% cover or 4.6 kg wet weight · m ⁻². Coral cover, which was already low on some reefs following Hurricane Allen in 1980, has been further reduced by as much as 60% since 1983 by competition with algae. Densities of D. antillarum at 10 sites in 1986 ranged from 0 to 12% of pre-1983 levels. Other echinoids, which might potentially compensate for the lack of herbivory from D. antillarum, have not increased significantly in density. Numbers of herbivorous scarids and acanthurids also remain at relatively low levels, because of overfishing. In the absence of high densities of fish and sea urchins, it is likely that recent changes in community structure will continue, resulting in further replacement of corals by algae in shallow water. The impact of the urchin mass mortalities is qualitatively similar to previous experimental removals of this species. In both cases, removal of echinoids resulted in substantial increases in macroalgae. However, quantitatively, the responses of algal and coral communities to the natural die-off were significantly greater, probably due to wide differences in spatial and temporal scales of the respective perturbations.     

Kelp and sea urchin settlement mediated by biotic interactions with benthic coralline algal species

Species interactions can influence key ecological processes that support community assembly and composition. For example, coralline algae encompass extensive diversity and may play a major role in regime shifts from kelp forests to urchin-dominated barrens through their role in inducing invertebrate larval metamorphosis and influencing kelp spore settlement. In a series of laboratory experiments, we tested the hypothesis that different coralline communities facilitate the maintenance of either ecosystem state by either promoting or inhibiting early recruitment of kelps or urchins. Coralline algae significantly increased red urchin metamorphosis compared with a control, while they had varying effects on kelp settlement. Urchin metamorphosis and density of juvenile canopy kelps did not differ significantly across coralline species abundant in both kelp forests and urchin barrens, suggesting that recruitment of urchin and canopy kelps does not depend on specific corallines. Non-calcified fleshy red algal crusts promoted the highest mean urchin metamorphosis percentage and showed some of the lowest canopy kelp settlement. In contrast, settlement of one subcanopy kelp species was reduced on crustose corallines, but elevated on articulated corallines, suggesting that articulated corallines, typically absent in urchin barrens, may need to recover before this subcanopy kelp could return. Coralline species differed in surface bacterial microbiome composition; however, urchin metamorphosis was not significantly different when microbiomes were removed with antibiotics. Our results clarify the role played by coralline algal species in kelp forest community assembly and could have important implications for kelp forest recovery.     

The efficiency and effectiveness of different sea urchin removal methods for kelp forest restoration

Sea urchin overgrazing has caused widespread phase shifts from kelp forests to “urchin barrens” on many temperate reefs, reducing habitat complexity, productivity, and biodiversity. Sea urchin removal is increasingly used for kelp restoration; however, few studies have quantified the efficiency and effectiveness of different removal methods, resulting in limited understanding of their practicality. In this study, the efficiency (removal rate) and effectiveness (proportion removed) of four removal methods were evaluated in northeastern New Zealand. We compared culling or collecting sea urchins by either SCUBA or freediving in 128 small-scale plots (25 m²). We also evaluated the efficiency and effectiveness of culling in four large (1.6–2 ha) barren areas, scales relevant for restoration. On average, culling sea urchins was 1.9–4.4 times faster than collecting, and SCUBA was 1.5–3.3 times faster than freediving. Removal rates increased with sea urchin density, especially for culling on SCUBA, while freediving removal rates increased with experience. Effectiveness was lower in large-scale removals (86–93% of sea urchins ≥40 mm removed) compared to small-scale removals (98–99%), but sufficient for restoration objectives. Estimated time per area (using SCUBA culling) was similar across large-scale removals (49–57 hours/ha), despite an almost 2-fold variation in initial sea urchin densities (approximately 4–8 urchins/m²), suggesting area may better predict total removal time than simply number of sea urchins across low-density ranges. While sea urchin removal provides a rapid, feasible, and effective approach to restoring kelp in urchin barrens, restoration plans need to also address the causes of sea urchin overpopulation to ensure long-term benefits.     

Survival of kelp adjacent to areas grazed by sea urchins in New South Wales,Australia

Abstract The sea urchin Centrostephanus rodgersii grazes areas of reef free of large brown algae (the barrens habitat). Survival of the kelp Ecklonia radiata adjacent to patches of barrens habitat was compared to that in the centre of kelp forests (centre) and edges of forests not adjacent to patches of barrens habitat (ungrazed). Estimates of rates of instantaneous mortality for tagged plants, as described by the slope of a negative exponential model (± 95% CI) were: barrens, 0.078 ± 0.004; centre, 0.051 ± 0.004 and ungrazed edge, 0.065 ± 0.007. Survival of plants was greatest in the middle of forests and least on the margins of patches of barrens habitat. A significant proportion of mortality in the barrens and centre positions was caused by herbivorous fish. When these plants were excluded from analysis there were no significant differences in survival between the barrens and ungrazed positions. It is concluded that C. rodgersii has little impact on the abundance of Ecklonia outside sharply defined boundaries.     

Microbiome variation in corals with distinct depth distribution ranges across a shallow–mesophotic gradient (15–85 m)

Mesophotic coral ecosystems (MCEs) are generally poorly studied, and our knowledge of lower MCEs (below 60 m depth) is largely limited to visual surveys. Here, we provide a first detailed assessment of the prokaryotic community associated with scleractinian corals over a depth gradient to the lower mesophotic realm (15–85 m). Specimens of three Caribbean coral species exhibiting differences in their depth distribution ranges (Agaricia grahamae, Madracis pharensis and Stephanocoenia intersepta) were collected with a manned submersible on the island of Curaçao, and their prokaryotic communities assessed using 16S rRNA gene sequencing analysis. Corals with narrower depth distribution ranges (depth-specialists) were associated with a stable prokaryotic community, whereas corals with a broader niche range (depth-generalists) revealed a higher variability in their prokaryotic community. The observed depth effects match previously described patterns in Symbiodinium depth zonation. This highlights the contribution of structured microbial communities over depth to the coral’s ability to colonize a broader depth range.     

Canopy-Forming Seaweeds in Urchin-Dominated Systems in Eastern Canada: Structuring Forces or Simple Prey for Keystone Grazers?

Models of benthic community dynamics for the extensively studied, shallow rocky ecosystems in eastern Canada emphasize kelp-urchin interactions. These models may bias the perception of factors and processes that structure communities, for they largely overlook the possible contribution of other seaweeds to ecosystem resilience. We examined the persistence of the annual, acidic (H₂SO₄), brown seaweed Desmarestia viridis in urchin barrens at two sites in Newfoundland (Canada) throughout an entire growth season (February to October). We also compared changes in epifaunal assemblages in D. viridis and other conspicuous canopy-forming seaweeds, the non-acidic conspecific Desmarestia aculeata and kelp Agarum clathratum. We show that D. viridis can form large canopies within the 2-to-8 m depth range that represent a transient community state termed “Desmarestia bed”. The annual resurgence of Desmarestia beds and continuous occurrence of D. aculeata and A. clathratum, create biological structure for major recruitment pulses in invertebrate and fish assemblages (e.g. from quasi-absent gastropods to >150 000 recruits kg⁻¹ D. viridis). Many of these pulses phase with temperature-driven mass release of acid to the environment and die-off in D. viridis. We demonstrate experimentally that the chemical makeup of D. viridis and A. clathratum helps retard urchin grazing compared to D. aculeata and the highly consumed kelp Alaria esculenta. In light of our findings and related studies, we propose fundamental changes to the study of community shifts in shallow, rocky ecosystems in eastern Canada. In particular, we advocate the need to regard certain canopy-forming seaweeds as structuring forces interfering with top-down processes, rather than simple prey for keystone grazers. We also propose a novel, empirical model of ecological interactions for D. viridis. Overall, our study underscores the importance of studying organisms together with cross-scale environmental variability to better understand the factors and processes that shape marine communities.     

Contrasting spatial heterogeneity of sessile organisms within mussel (Perna perna L.) beds in relation to topographic variability

We examined the spatial heterogeneity in three sessile rocky shore organisms, the mussel Perna perna, the barnacle Octomeris angulosa (Sowerby) and the red alga Gelidium pristoides (Turn.) at a range of continuous local scales along horizontal transects within mid- and upper mussel beds of South African shores. We also examined the relationships between variability of organisms and topographic features (rock depressions, slope, aspect), and between mussel, barnacle and algal variability over the same scales. To estimate spatial heterogeneity, we analyzed scaling properties of semivariograms using a fractal approach. Relationships between different variables at the different scales were examined by cross-semivariograms. Spatial dependence of P. perna variability increased with spatial dependence of topographic variability, so that scaling regions of mussel and topographic distributions corresponded well. This relationship often improved with larger local scales (mussel cover increased with depressions, steeper slope and aspect towards waves), while at smaller spatial scales, variability in mussel cover was less well explained by variability in topography. The variability of the barnacle O. angulosa exhibited spatial dependence, even on topographically unstructured shores. In contrast, the distribution of the alga G. pristoides revealed high fractal dimensions, showing spatial independence on topographically unstructured shores. Algae also showed a very strong negative relationship with mussels at most local scales, and a negative relationship with barnacles in upper zones, especially at larger local scales. Barnacles may show clear spatial dependence because of hydrodynamics (at larger local scales) and the need to find a future mate in close proximity (at smaller local scales), while algae may show a strong negative relationship with mussels because of competition for space.