Is habitat amount important for biodiversity in rocky shore systems? A study of South African mussel assemblages

Habitat-forming species on rocky shores are often subject to high levels of exploitation, but the effects of subsequent habitat loss and fragmentation on associated species and the ecosystem as a whole are poorly understood. In this study, the effects of habitat amount on the fauna associated with mussel beds were investigated, testing for the existence of threshold effects at small landscape scales. Specifically, the relationships between mussel or algal habitat amount and: associated biodiversity, associated macrofaunal abundance and density of mussel recruits were studied at three sites (Kidd’s Beach, Kayser’s Beach and Kini Bay) on the southern and south-eastern coasts of South Africa. Samples, including mussel-associated macrofauna, of 10 × 10 cm were taken from areas with 100 % mussel cover (Perna perna or a combination of P. perna and Mytilus galloprovincialis) at each site. The amount of habitat provided by mussels and algae surrounding the sampled areas was thereafter determined at the 4.0 m² scale. A number of significant positive relationships were found between the amount of surrounding mussel habitat and the abundances of several taxa (Anthozoa, Malacostraca and Nemertea). Likewise, there were positive relationships between the amount of surrounding algal habitat and total animal abundance as well as abundance of mussel recruits at one site, Kini Bay. In contrast, abundance of mussel recruits showed a significant negative relationship with the amount of mussel habitat at Kayser’s Beach. Significant negative relationships were also detected between the amount of mussel habitat and species richness and total abundance at Kidd’s Beach, and between amount of mussel habitat and the abundance of many taxa (Bivalvia, Gastropoda, Maxillopoda, Ophiuroidea, Polychaeta and Pycnogonida) at all three sites. No threshold effects were found, nor were significant relationships consistent across the investigated sites. The results indicate that the surrounding landscape is important in shaping the structure of communities associated with these mussel beds, with significant effects of the amount of surrounding habitat per se. The strength and the direction of habitat effects vary, however, between shores and probably with the scale of observation as well as with the studied dependent variables (e.g. diversity, abundance, mussel recruitment, species identity), indicating the complexity of the processes structuring macrofaunal communities on these shores.     

Bioengineers and their associated fauna respond differently to the effects of biogeography and upwelling

Temperature and primary production (often linked to nutrient supply) are two of the few factors influencing species diversity and abundances across mesoscale gradients, while at smaller scales the habitat complexity offered by bioengineers is important. Previous studies have illustrated effects of upwelling and biogeography on intertidal bioengineers, but it is not known if these processes influence assemblages associated with those bioengineers in a similar way. We examined the habitat structure offered by two species of mussels and their associated fauna in five regions across 3000 km and three biogeographic provinces of the South African coast, replicating upwelling and non-upwelling areas within each region. Upwelling and region influenced the structure of mussel beds (the density and size of mussels). In contrast, upwelling did not influence mesoscale differences in composition, abundance, and numbers of species of crustaceans, mollusks, or polychaetes in mussel beds. Regardless of trophic level or mode of reproduction, mussel bed fauna were influenced only by region. Regional differences were strongly influenced by biogeography. The associated fauna was, however, also strongly correlated with the structure of the habitats created by mussels. Our results support the importance of upwelling to a critical ecosystem engineer, but show that these effects do not extend directly to the assemblages of associated fauna, which are more influenced by regional-scale effects and biogeography. We suggest that mesoscale patterns in the associated fauna of this bioengineered habitat are driven by the direct effects of biogeography, combined with the influence of biogeography and upwelling on mussel bed structure.     

Estuarine Vegetated Habitats as Corridors for Predator Movements

Abstract: The spatial proximity of one habitat to another can strongly influence population and community dynamics. We investigated whether the proximity of intertidal oyster reefs to vegetated estuarine habitats, salt marshes, and seagrass beds, affects the abundance and community structure of benthic macroinvertebrates on reefs and predator-prey interactions between mobile predators and bivalves living on reefs. Benthic macroinvertebrate abundance was highest on reefs spatially separated from salt marshes. Macroinvertebrate species richness was highest on reefs separated from both salt marshes and seagrass beds. Comparisons of predation on juvenile bivalves transplanted to reefs for 7–12 days indicated that survivorship of clams was greatest on reefs spatially separated from both salt marshes and seagrass beds, whereas reef proximity to vegetated habitats did not affect the survivorship of oysters. The foraging behavior of blue crabs may explain patterns of macroinvertebrate abundance and clam survivorship among reefs with different proximity to vegetated habitats. In experiments conducted in 30-m² field enclosures, blue crabs had higher predation rates on hard clams transplanted onto artificial reefs adjacent to salt marshes or seagrass beds than on reefs separated from both habitats by unvegetated sand bottom. Thus, vegetated habitats appeared to act as corridors by facilitating the access of blue crabs to oyster reefs and enhancing the intensity of blue crab predation. Such an understanding of the effects of landscape characteristics of estuarine habitats on their value as habitats for estuarine organisms can be used to predict the consequences of habitat fragmentation on ecosystem function and to improve strategies for habitat and species conservation and restoration.     

Distribution of a marginal population of <Emphasis Type="Italic">Mytilus edulis</Emphasis>: responses to biotic and abiotic processes at different spatial scales

Physical and biological processes interact to produce pattern in nature. Pattern is scale dependent as processes generating pattern are heterogeneous in time and space. We tested some causes of variation in abundance and distribution of three marginal populations of sublittoral blue mussels, Mytilus edulis, in the non-tidal northeastern Baltic Sea. We studied the role of substrate inclination, perennial algae and siltation along local wave exposure gradients on mussel distribution over a regional salinity gradient. We found marked differences on regional scales (p < 0.001) with lower densities and biomasses of mussels with declining salinity. Along local gradients, mussel densities increased with increasing exposure (p < 0.001) and declining slope and sedimentation (p < 0.01). Site specifically, densities of blue mussels and the perennial red algae, Furcellaria lumbricalis, were positively related, results supported by a colonisation experiment. Also, young post-recruits showed significant relations to adult biomass, wave exposure, algal biomass, bottom slope and sediment cover. Findings showed that the relative importance of the determinants affecting blue mussels at the edge of their range vary with scale and are affected by the density and size structure of mussel populations. The study provides an indication of the types of factors that may be invoked as causes of spatial variation in marginal blue mussel populations and reinforces the need to consider multiple aspects when distributional patterns are assessed.     

Habitat modification affects recruitment of abalone in central New Zealand

The habitat experienced during early life-history stages can determine the number and quality of individuals that recruit to adult populations. In a field experiment, biogenic habitat complexity was manipulated (presence or absence of foliose macroalgae) at two depths (2–3 m and 5–6 m) and the habitat-dependent effects on recruitment of the black foot abalone (Haliotis iris) were examined at three field sites along the south coast of Wellington, New Zealand (41°20′S, 174°47′E), between July and November 2005. Recruit density (<5 weeks post-settlement) was measured on cobbles covered with crustose coralline algae. Habitats of low complexity (barrens treatments) had consistently greater densities of recruits than habitats of high complexity (algae treatments). However, recruits in algae habitats were larger, and for deep habitats, there was greater survival in algae habitats compared with barrens habitats. While depth had no significant effect on early recruit (<2 weeks post-settlement) density, late recruit (<5 weeks post-settlement) density was greater in shallow habitats, and so it seems recruit survival was greater in shallow habitats. In this experiment, algal habitat complexity had strong effects on early recruit abundance, but habitat-dependent variations in recruit growth and survival may modify initial patterns of abundance and determine recruitment to adult abalone populations.     

Ontogenetic changes in small-scale movement by recruits of an exploited mussel: implications for the fate of larvae settling on algae

Many South African populations of the brown mussel Perna perna have been depleted through over-exploitation by subsistence harvesters. This is problematic because recovery after disturbance is very slow, partly because recruits are largely associated with adult mussels. However, unlike large recruits of 3.5–10 mm that exhibit spatial structure related to that of adults, a very high proportion of settlers and small recruits (0–3.5 mm) occur on foliose algae. We tested the hypothesis that recruits on algae move to adult mussel beds after a period of growth, with the null hypothesis that they die at a certain size. We conducted an indirect field study comparing the ratios of large to small recruits in 100% cover mussel patches at locations with high and low algal cover. A second laboratory experiment analysed whether the size of recruits on algae affects their active movement behaviour in response to nearby mussel patches. Large/small recruit ratios were slightly, but not significantly greater in high than low algal cover locations. Both small (2–2.5 mm) and medium (4.5–5.5 mm) recruits remained on algae and moved very short distances throughout the laboratory experiment, while larger recruits (9–10 mm) moved significantly further distances and more often into mussel patches. The results suggest that very large recruits are able to migrate actively to nearby mussel patches, indicating ontogenetic shifts in this behaviour. However, the absence of a significant difference in ratios between field locations with high and low algal cover suggests many large recruits are accidentally dislodged from the algae and presumably die. Thus settlement of P. perna onto algae is likely to be wasted, with consequences for sustainable management of the mussel resource.     

Habitat complexity impacts persistence and species interactions in an intertidal whelk

Although experiments have shown that habitat structure may influence the distribution of species and species interactions, these effects are still not commonly integrated into studies of community dynamics. Since habitat structure often varies within and among communities, this may limit our understanding of how various factors influence communities. Here, we examined how mussel bed complexity (the presence and thickness of mussel layers) influenced the persistence of whelks (Nucella emarginata) and interactions with a top predator (ochre sea stars, Pisaster ochraceus) and prey (mussels, Mytilus californianus). Results from a mark?Crecapture experiment indicate that whelk recapture rates are higher in more complex habitats, and laboratory experiments demonstrate that habitat complexity affects whelk feeding, growth, and nonconsumptive interactions with a keystone predator. Habitat complexity therefore has direct effects on species and also may lead to trade-offs among feeding, refuge, and other factors, potentially influencing the distribution of whelks and the effects of both whelks and sea stars on intertidal communities. These results demonstrate that habitat structure may play an important role in intertidal communities and other habitats and should be further considered in the experimental design of future studies of community dynamics.     

Context-dependent species identity effects within a functional group of filter-feeding bivalves

We asked whether species richness or species identity contributed more to ecosystem function in a trait-based functional group, burrowing, filter-feeding bivalves (freshwater mussels: Unionidae), and whether their importance changed with environmental context and species composition. We conducted a manipulative experiment in a small river examining the effects of mussel assemblages varying from one to eight species on benthic algal standing crop across two sets of environmental conditions: extremely low discharge and high water temperature (summer); and moderate discharge and water temperature (fall). We found strong species identity effects within this guild, with one species (Actinonaias ligamentina) influencing accrual of benthic algae more than other species, but only under summer conditions. We suspect that this effect is due to a combination of the greater biomass of this species and its higher metabolic and excretion rates at warm summer temperatures, resulting in increased nitrogen subsidies to benthic algae. We also found that Actinonaias influenced the condition of other mussel species, likely through higher consumption, interference, or both. This study demonstrates that species within trait-based functional groups do not necessarily have the same effects on ecosystem properties, particularly under different environmental conditions.     

Density-dependent biodiversity effects on physical habitat modification by freshwater bivalves

Several decades of research have shown that biodiversity affects ecosystem processes associated with resource capture and the production of biomass within trophic levels. Although there are good reasons to expect that biodiversity influences non-trophic ecosystem processes, such as the physical creation or modification of habitat, studies investigating the role of biodiversity on physical processes are scarce. Here we report the results of a study using artificial streams to test the influence of freshwater mussel biodiversity on gravel erosion during high flows while manipulating mussel abundance. Mussel species vary in traits that should influence their effects on erosion, such as size, shell morphology, and burrowing behavior. We found that mussel species richness was associated with an increase in erosion at both low and high densities. Planned contrasts showed that the erosion observed in species mixtures was purely additive at low density, indicating that erosion in a species polyculture could routinely be predicted by the performance of monocultures. However, at high density certain combinations of species showed nonadditive effects on erosion, suggesting that organism abundance can fundamentally alter biodiversity effects. Although this may have been a result of altered species interactions at high density, our study design cannot confirm this.     

Community structure in Florida Escarpment seep and Snake Pit (Mid-Atlantic Ridge) vent mussel beds

Comparisons between invertebrate communities hosted by similar foundation species under different environmental conditions permit identification of patterns of species distributions that might be characteristic of the different ecosystems. Similarities and differences in community structure between two major types of chemosynthetic ecosystems were assessed by analyzing samples of invertebrates associated with Bathymodiolus heckerae Gustafson et al. mussel beds at the Florida Escarpment seep (Gulf of Mexico, 26°01.8N; 84°54.9W; October 2000) and B. puteoserpentis von Cosel et al. mussel beds at the Snake Pit vent (Mid-Atlantic Ridge, 23°22.1N; 44°56.9W; July 2001). Macrofaunal species richness was nearly twice as high in the seep mussel bed compared to the vent mussel bed, and only a single morphospecies, the ophiuroid Ophioctenella acies Tyler et al., was shared between the sites. Similarities between the two faunas at higher taxonomic levels (genus and family) were evident for only a small percentage of the total number of taxa, suggesting that evolutionary histories of many of these seep and vent macrofaunal taxa are not shared. The taxonomic distinctiveness of the seep and vent mussel-bed macrofaunal communities supports the hypothesis that environmental and oceanographic barriers prevent most taxa from occupying both types of habitats. Macrofaunal community heterogeneity among samples was similar in seep and vent mussel beds, indicating that spatial scales of processes regulating community variability may be similar in the two types of ecosystems. Suspension feeders were not represented in the macrofauna of seep or vent mussel beds. Primary consumers (deposit feeders and grazers) contributed more to the total abundance of macrofauna of seep mussel beds than vent mussel beds; secondary consumers (polychaetes and shrimp) were more abundant in the vent mussel beds.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-004-1304-z.Communicated by J.P. Grassle, New Brunswick     

Bird‐community responses to habitat creation in a long‐term,large‐scale natural experiment

Ecosystem function and resilience are compromised when habitats become fragmented due to land‐use change. This has led to national and international conservation strategies aimed at restoring habitat extent and improving functional connectivity (i.e., maintaining dispersal processes). However, biodiversity responses to landscape‐scale habitat creation and the relative importance of spatial and temporal scales are poorly understood, and there is disagreement over which conservation strategies should be prioritized. We used 160 years of historic post‐agricultural woodland creation as a natural experiment to evaluate biodiversity responses to habitat creation in a landscape context. Birds were surveyed in 101 secondary, broadleaf woodlands aged 10–160 years with ≥80% canopy cover and in landscapes with 0‐17% broadleaf woodland cover within 3000 m. We used piecewise structural equation modeling to examine the direct and indirect relationships between bird abundance and diversity, ecological continuity, patch characteristics, and landscape structure and quantified the relative conservation value of local and landscape scales for bird communities. Ecological continuity indirectly affected overall bird abundance and species richness through its effects on stand structure, but had a weaker influence (effect size near 0) on the abundance and diversity of species most closely associated with woodland habitats. This was probably because woodlands were rapidly colonized by woodland generalists in ≤10 years (minimum patch age) but were on average too young (median 50 years) to be colonized by woodland specialists. Local patch characteristics were relatively more important than landscape characteristics for bird communities. Based on our results, biodiversity responses to habitat creation depended on local‐ and landscape‐scale factors that interacted across time and space. We suggest that there is a need for further studies that focus on habitat creation in a landscape context and that knowledge gained from studies of habitat fragmentation and loss should be used to inform habitat creation with caution because the outcomes are not necessarily reciprocal.     

Macrofaunal responses to structural complexity are mediated by environmental variability and surrounding habitats

Investigating the context that surrounds each habitat is crucial to understand local responses of assemblages of species to habitats. Here, I tested whether responses of benthic macroinvertebrates to the structural complexity of experimental habitats were mediated by the characteristics of their surrounding habitats (i.e. rockpools or emergent-rock surfaces). Each type of surrounding habitat provided particular biotic (e.g. algal growth) and abiotic (e.g. temperature, water movement) conditions that were expected to affect benthic assemblages. The results show that (1) composition of entire assemblages was affected by the matrix and type of habitat; (2) effects of the matrix on the number of species varied depending on the different types of habitats; (3) abundant species showed specific responses to type of habitat, independently of the matrix; and (4) relationships between numbers of species and two major environmental variables (i.e. micro-algal biomass and sediment) varied depending on the type of habitats and the surrounding matrix. Generally, these findings demonstrate that understanding the consequences of the spatial structure of these habitats is essential to advance our knowledge on patterns of abundance and distributions of functionally important species and ultimately the structure of intertidal assemblages.     

Environmental determinants of distribution and foraging behaviour of cormorants (Phalacrocorax spp.) in temperate estuarine habitats

The distribution and behaviour of cormorants in estuarine environments was examined on the central coast of New South Wales, Australia, with respect to habitat associations at different spatial scales. No consistent variation in abundance was found for four species of cormorants (great Phalacrocorax carbo, pied P. varius, little black P. sulcirostris, and little pied P. melanoleucos) with state of tide (high and low) and time of day (early, middle, and late) at five estuarine locations. Differences in abundance were found among locations that were not confounded by short-term temporal variation (i.e. time of day and tide). Differences in abundance were detected among habitats (e.g. bays, creeks, and headlands) separated by hundreds of metres to kilometres in different estuaries. Cormorants of all species were rare on the open coast and near the entrance of estuaries. Abundances of cormorants varied greatly within and among creeks, bays, and river channels. Presence of seagrass beds explained much of this variation and most of the cormorants swimming and feeding were found near seagrass. Mapping of seagrass beds and the positions of cormorants at scales of metres to hundreds of metres showed a close relationship between the presence of swimming and roosting beds and the presence of seagrass beds for P. melanoleucos and for P. carbo. We argue that cormorants make decisions to visit particular estuarine habitats, especially those with seagrass, where many types of prey (e.g. fish and crustaceans) are probably most abundant. These choices must be interpreted in the context of decisions that cormorants make on scales of hundreds to thousands of kilometres during periodic excursions to the interior of Australia. Further, environmental threats to seagrass beds could impinge on these mobile visitors to the same extent as on more permanent residents. Received: 14 February 2000 / Accepted: 17 July 2000     

Roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis>) as an important predator on blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) populations in a brackish water environment,the northern Baltic Sea

Suspension-feeding bivalves are organisms of major functional importance in several aquatic environments around the world. They are also important food items for many fish and benthivorous seabirds. It has commonly been thought that predation pressure on blue mussel (Mytilus edulis) populations is negligible in the Baltic Sea, owing to the scarcity of major invertebrate predators such as starfish and crabs. It has recently been shown, however, that the blue mussel is the main food item for roach (Rutilus rutilus) in the archipelago areas of the western Gulf of Finland, where this freshwater fish species has become increasingly abundant, mainly due to increased eutrophication. To quantify the influence of roach predation on blue mussel populations we measured the standing biomass and size structure of the local blue mussel population and used a bioenergetic model to estimate mussel consumption by individual roach during two consecutive summers, 1997 and 1998. The results of the model were combined with existing data on roach abundance, giving annual consumption estimates of 75–105 kg blue mussel dry weight ha^–1 in the study area, approximately two-thirds of these consumed mussels being >10 mm. This corresponds to approximately one-third of the standing population of mussels >10 mm in the area. Our results suggest that the predation effects of vertebrates on Baltic blue mussel populations are not insignificant, as previously believed. Predation by roach and other predators may have an important structuring effect on unstable blue mussel communities within the Gulf of Finland, where the species lives at the edge of its range.Communicated by M. Kühl, Helsingør     

Density-dependent facilitation cascades determine epifaunal community structure in temperate Australian mangroves

Co-occurring foundation species can determine biological community structure via facilitation cascades. We examined the density dependencies of facilitation cascades, including how the density of a basal foundation species influences the density of secondary foundation species, and how the density of secondary foundation species influences community structure. The system in which we assessed density dependencies was a temperate mangrove forest in which pneumatophores trap the fucoid alga Hormosira banksii and provide substrate for the oyster, Saccostrea glomerata. The alga and oyster in turn determine benthic community structure. In the field, algal biomass was positively correlated with pneumatophore density. Oysters, by contrast, were highly over-dispersed and correlated with the presence/absence of pneumatophores. Epifaunal abundance and species richness were positively correlated with algal and oyster abundance, but their effects were independent. The positive effect of pneumatophore density on epifauna was primarily an indirect effect of trapping more algae. Pneumatophores did not directly influence invertebrate communities. Experiments revealed that, at very low pneumatophore densities, algal retention was insufficient to facilitate epifauna above that found on pneumatophores alone. At higher densities, however, increasing the density of pneumatophores increased algal retention, and the density and diversity of associated invertebrates. Shading by the mangrove canopy reduced algal biomass but did not modify the density-dependent nature of the cascade. Our results extend facilitation theory by showing that the density of both basal and secondary foundation species can be critical in triggering facilitation cascades. Our study also reveals that, where foundation species co-occur, multiple, independent cascades may arise from a single basal facilitator. These findings enhance our understanding of the role of density-dependent facilitation cascades in community assembly.     

Prioritizing sites for conservation based on similarity to historical baselines and feasibility of protection

The concept of shifting baselines in conservation science implies advocacy for the use of historical knowledge to inform these baselines but does not address the feasibility of restoring sites to those baselines. In many regions, conservation feasibility varies among sites due to differences in resource availability, statutory power, and land‐owner participation. We used zooarchaeological records to identify a historical baseline of the freshwater mussel community's composition before Euro‐American influence at a river‐reach scale (i.e., a kilometer stretch of river that is abiotically similar) in the Leon River of central Texas (U.S.A.). We evaluated how the community reference position and the feasibility of conservation might enable identification of sites where conservation actions would preserve historically representative communities and be likely to succeed. We devised a conceptual model that incorporated community information and landscape factors to link the best conservation areas to potential cost and conservation benefits. Using fuzzy ordination, we identified modern mussel beds that were most like the historical baseline. We then quantified housing density and land use near each river reach identified to estimate feasibility of habitat restoration. Using our conceptual framework, we identified reaches of high conservation value (i.e., contain the best mussel beds) and where restoration actions would be most likely to succeed. Reaches above Lake Belton were most similar in species composition and relative abundance to zooarchaeological sites. A subset of these mussel beds occurred in locations where conservation actions appeared most feasible. Our results show how to use zooarchaeological data (biodiversity data often readily available) and estimates of conservation feasibility to inform conservation priorities at a local spatial scale.     

Ecological and Economic Services Provided by Birds on Jamaican Blue Mountain Coffee Farms

Abstract: Coffee farms can support significant biodiversity, yet intensification of farming practices is degrading agricultural habitats and compromising ecosystem services such as biological pest control. The coffee berry borer (Hypothenemus hampei) is the world's primary coffee pest. Researchers have demonstrated that birds reduce insect abundance on coffee farms but have not documented avian control of the berry borer or quantified avian benefits to crop yield or farm income. We conducted a bird‐exclosure experiment on coffee farms in the Blue Mountains, Jamaica, to measure avian pest control of berry borers, identify potential predator species, associate predator abundance and borer reductions with vegetation complexity, and quantify resulting increases in coffee yield. Coffee plants excluded from foraging birds had significantly higher borer infestation, more borer broods, and greater berry damage than control plants. We identified 17 potential predator species (73% were wintering Neotropical migrants), and 3 primary species composed 67% of migrant detections. Average relative bird abundance and diversity and relative resident predator abundance increased with greater shade‐tree cover. Although migrant predators overall did not respond to vegetation complexity variables, the 3 primary species increased with proximity to noncoffee habitat patches. Lower infestation on control plants was correlated with higher total bird abundance, but not with predator abundance or vegetation complexity. Infestation of fruit was 1–14% lower on control plants, resulting in a greater quantity of saleable fruits that had a market value of US$44–$105/ha in 2005/2006. Landscape heterogeneity in this region may allow mobile predators to provide pest control broadly, despite localized farming intensities. These results provide the first evidence that birds control coffee berry borers and thus increase coffee yield and farm income, a potentially important conservation incentive for producers.     

Biotic habitat complexity controls species diversity and nutrient effects on net biomass production

Canopy-forming plants and algae commonly contribute to spatial variation in habitat complexity for associated organisms and thereby create a biotic patchiness of communities. In this study, we tested for interaction effects between biotic habitat complexity and resource availability on net biomass production and species diversity of understory macroalgae by factorial field manipulations of light, nutrients, and algal canopy cover in a subtidal rocky-shore community. Presence of algal canopy cover and/or artificial shadings limited net biomass production and facilitated species diversity. Artificial shadings reduced light to levels similar to those under canopy cover, and net biomass production was significantly and positively correlated to light availability. Considering the comparable and dependent experimental effects from shadings and canopy cover, the results strongly suggest that canopy cover controlled net biomass production and species diversity by limiting light and thereby limiting resource availability for community production. Canopy cover also controlled experimental nutrient effects by preventing a significant increase in net biomass production from nutrient enrichment recorded in ambient light (no shading). Changes in species diversity were mediated by changes in species dominance patterns and species evenness, where canopy cover and shadings facilitated slow-growing crust-forming species and suppressed spatial dominance by Fucus vesiculosus, which was the main contributor to net production of algal biomass. The demonstrated impacts of biotic habitat complexity on biomass production and local diversity contribute significantly to understanding the importance of functionally important species and biodiversity for ecosystem processes. In particular, this study demonstrates how loss of a dominant species and decreased habitat complexity change the response of the remaining assembly to resource loading. This is of potential significance for marine conservation since resource loading often promotes low habitat complexity and canopy species are among the first groups lost in degraded aquatic systems.     

A fractal approach for detecting spatial hierarchy and structure on mussel beds

Within beds of blue mussel (Mytilus edulis L.), individuals are aggregated into small patches, which in turn are incorporated into bigger patches, revealing a complex hierarchy of spatial structure. The present study was done to find the different scales of variation in the distribution of mussel biomass, and to describe the spatial heterogeneity on these scales. The three approaches compared for this purpose were fractal analysis, spatial autocorrelation and hierarchical (or nested) analysis of variances (ANOVA). The complexity (i.e. patchiness) of mussel aggregations was described with fractal dimension, calculated with the semivariogram method. Three intertidal mussel beds were studied on the west coast of Sweden. The distribution of wet biomass was studied along transects up to 128 m. The average biomasses of blue mussels on the three mussel beds were 1825낚, 179ᆩ and 576ᇖ g per0.1 m², respectively, and the fractal dimensions of the mussel distribution were 1.726ǂ.010, 1.842ǂ.014 and 1.939ǂ.029 on transects 1-3, respectively. Distributions of mussels revealed multiscaling behaviour. The fractal dimension significantly changed twice on different scales on the first bed (thus showing three scaling regions), the second and third beds revealed two and three scaling regions, respectively. High fractal dimension was followed by significant spatial autocorrelation on smaller scales. The fractal analysis detects the multiple scaling regions of spatial variance even when the spatial structure may not be distinguished significantly by conventional statistical inference. The study shows that the fractal analysis, the spatial autocorrelation analysis and the hierarchical ANOVA give complementary information about the spatial variability in mussel populations.