Multivariate analysis of the bentho-demersal ichthyofauna along soft bottoms of the Eastern Atlantic: comparison between unvegetated substrates,seagrass meadows and sandy bottoms beneath sea-cage fish farms

By means of multivariate techniques, we studied: (1) the differences in the structure of bentho-demersal, non-cryptic, fish assemblages associated with unvegetated sandy substrates, vegetated meadows constituted by the seagrass Cymodocea nodosa and the bottoms under the influence of sea-cage fish farms; as well as (2) the persistence of these patterns with regard to different scales of spatial variability, across three islands of the Canarian Archipelago (Central East Atlantic). Our sampling strategy (involving three islands, with five locations per island, and two sites within each location) detected significant changes in the composition and structure of the fish assemblages between the three habitats. Fish assemblages associated with the unvegetated and vegetated bottoms were similar among the surveyed islands. In contrast, we observed a significant inter-island variability in the fish populations associated with the sea-cage fish farms. The presence of the sea-cage fish farms increased the overall fish abundance (184.8±49.8 ind 100 m⁻²) as compared to both the vegetated (38.8±9.7 ind 100 m⁻²) and unvegetated habitats (1.1±0.4 ind 100 m⁻²). Differences within and between the habitats were found to be associated with the relative abundance of a few fish species. The most abundant species were Xyrichthys novacula in the unvegetated bottoms and Diplodus annularis, Spondyliosoma cantharus and Mullus surmuletus on the seagrass meadows. Finally, we recorded an increase in the abundance of Heteroconger longissimus, Trachinus draco and Pagellus acarne in the bottoms beneath the sea-cage fish farms. These species, in addition to a group of large benthic chondrichthyes, were responsible for the differences between islands in the composition and structure of the demersal ichthyofauna beneath the sea-cage fish farms.     

Analysis of the influence of substrate variables on coral reef fish communities

A simple field technique to obtain a gross estimate of the surface area of a quadrat on a coral reef is described. This measure, termed the substrate rugosity index, was determined, in conjunction with two other substrate variables (vertical relief and coral species richness), in a series of 4 quadrats (10 to 40 m depth) along 4 transects. The mean substrate rugosity and vertical relief of a quadrat were highly correlated. A correlation analysis was made of the substrate variables and several reef fish community parameters (species richness, number of fishes and diversity). Species richness was highly correlated with substrate rugosity. This relationship was tested in two experimental quadrats and the results were generally in accord with those predicted. Stratification of the fish communities by body size revealed that the correlation with substrate rugosity was scale-dependent. The fish community parameters were poorly correlated with percentage substrate cover by corals (ramose and glomerate) and by sand. A significant area effect was determined for two species of sand-dwelling goby.     

Protection from fishing alters the species composition of fish assemblages in a temperate-tropical transition zone

Closure of areas to fishing is expected to result in an increase in the abundance of targeted species; however, changes to populations of species not targeted by fishermen will depend upon their role in the ecosystem and their relationship with targeted species. The effects of protection on targeted and non-targeted reef fish species at the Houtman Abrolhos Islands, Western Australia were studied using baited remote underwater stereo–video cameras. Video images were collected from shallow (8–12 m) and deep (22–26 m) reef sites inside a Marine Protected Area (MPA) at each of three island groups and from three replicate fished locations at each of these groups that span a temperate-tropical transition area. The MPAs were established in 1994 and vary in size from 13.72 km² at the Pelsaert group in the south to 22.29 km² at the Easter group to 27.44 km² at the Wallabi group in the north. The relative abundances of 137 fish species from 42 families were recorded. Large differences in fish assemblage structure existed between MPA and fished locations, and also between shallow and deep regions. Targeted fish species Plectropomus leopardus, Lethrinus miniatus, Lethrinus nebulosus, Pagrus auratus and Glaucosoma hebraicum were more abundant inside MPAs than in areas open to fishing. Their abundance inside MPAs was between 1.13 and 8 times greater than their abundance at fished locations. For non-targeted fish species many were more abundant in areas open to fishing, e.g. Coris auricularis, Thalassoma lutescens, Thalassoma lunare, Dascyllus trimaculatus, however others were conversely more abundant inside MPAs, e.g. Gymnothorax spp, Kyphosus sydneyanus, Scarus microhinos, Chromis westaustralis, Chaetodon spp. This study demonstrates that the removal of abundant targeted species from an ecosystem by fishing can indirectly impact non-fished species and alter the trophic structure of fish assemblages. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Selection of Islands for Conservation in the Urban Archipelago of Helsinki, Finland

Abstract: The occurrence of vascular plants was surveyed on 207 islands (size range 0.01–390.2 ha, number of plant species 1–449) offshore from the city of Helsinki in the Baltic Sea to examine the conservation value of these islands. We calculated a rarity score for each species (1/number of islands occupied by the species) and a biodiversity score for each island (sum of the rarity scores of each species present on the island). Positive correlations between species number and biodiversity score (r _s = 0.97, p < 0.001) and between biodiversity score and island area (r _s = 0.87, p < 0.001) indicated that these parameters are heavily dependent on island size. With the goal of including at least one occurrence (island) of all plant species, an iterative selection algorithm chose a set of 41 islands whose average size (29.3 ha) was four times the average size of all existing islands (7.0 ha). Strong nestedness ( N < 54) explains the concentration of plant species diversity on large islands. An operational strategy for selection of sites for protection is to complement the set produced by a selection algorithm with target species not yet included (e.g., endangered species with several occurrences). Comprehensive mapping and analysis of a taxonomic group will help integrate conservation biology into land-use planning and increase the quality of the networks of protected areas.     

Importance of dispersal routes that minimize open‐ocean movement to the genetic structure of island populations

Islands present a unique scenario in conservation biology, offering refuge yet imposing limitations on insular populations. The Kimberley region of northwestern Australia has more than 2500 islands that have recently come into focus as substantial conservation resources. It is therefore of great interest for managers to understand the driving forces of genetic structure of species within these island archipelagos. We used the ubiquitous bar‐shouldered skink (Ctenotus inornatus) as a model species to represent the influence of landscape factors on genetic structure across the Kimberley islands. On 41 islands and 4 mainland locations in a remote area of Australia, we genotyped individuals across 18 nuclear (microsatellite) markers. Measures of genetic differentiation and diversity were used in two complementary analyses. We used circuit theory and Mantel tests to examine the influence of the landscape matrix on population connectivity and linear regression and model selection based on Akaike's information criterion to investigate landscape controls on genetic diversity. Genetic differentiation between islands was best predicted with circuit‐theory models that accounted for the large difference in resistance to dispersal between land and ocean. In contrast, straight‐line distances were unrelated to either resistance distances or genetic differentiation. Instead, connectivity was determined by island‐hopping routes that allow organisms to minimize the distance of difficult ocean passages. Island populations of C. inornatus retained varying degrees of genetic diversity (N_A = 1.83 – 7.39), but it was greatest on islands closer to the mainland, in terms of resistance‐distance units. In contrast, genetic diversity was unrelated to island size. Our results highlight the potential for islands to contribute to both theoretical and applied conservation, provide strong evidence of the driving forces of population structure within undisturbed landscapes, and identify the islands most valuable for conservation based on their contributions to gene flow and genetic diversity.     

Eradication of Invasive Mammals on Islands Inhabited by Humans and Domestic Animals

Abstract: Non‐native invasive mammal species have caused major ecological change on many islands. To conserve native species diversity, invasive mammals have been eradicated from several islands not inhabited by humans. We reviewed the challenges associated with campaigns to eradicate invasive mammals from islands inhabited by humans and domestic animals. On these islands, detailed analyses of the social, cultural, and economic costs and benefits of eradication are required to increase the probability of local communities supporting the eradication campaign. The ecological benefits of eradication (e.g., improvement of endemic species’ probability of survival) are difficult to trade‐off against social and economic costs due to the lack of a common currency. Local communities may oppose an eradication campaign because of perceived health hazards, inconvenience, financial burdens, religious beliefs, or other cultural reasons. Besides these social challenges, the presence of humans and domestic animals also complicates eradication and biosecurity procedures (measures taken to reduce the probability of unwanted organisms colonizing an island to near zero). For example, houses, garbage‐disposal areas, and livestock‐feeding areas can provide refuges for certain mammals and therefore can decrease the probability of a successful eradication. Transport of humans and goods to an island increases the probability of inadvertent reintroduction of invasive mammals, and the establishment of permanent quarantine measures is required to minimize the probability of unwanted recolonization after eradication. We recommend a close collaboration between island communities, managers, and social scientists from the inception of an eradication campaign to increase the probability of achieving and maintaining an island permanently free of invasive mammals.     

Flows, droughts, and aliens: factors affecting the fish assemblage in a Sierra Nevada, California, stream

The fishes of Martis Creek, in the Sierra Nevada of California (USA), were sampled at four sites annually over 30 years, 1979-2008. This long-term data set was used to examine (1) the persistence and stability of the Martis Creek fish assemblage in the face of environmental stochasticity; (2) whether native and alien fishes responded differently to a natural hydrologic regime (e.g., timing and magnitude of high and low flows); and (3) the importance of various hydrologic and physical habitat variables in explaining the abundances of native and alien fish species through time. Our results showed that fish assemblages were persistent at all sample sites, but individual species exhibited marked interannual variability in density, biomass, and relative abundance. The density and biomass of native fishes generally declined over the period of study, whereas most alien species showed no significant long-term trends. Only alien rainbow trout increased in both density and biomass at all sites over time. Redundancy analysis identified three hydrologic variables (annual 7-day minimum discharge, maximum winter discharge, and number of distinct winter floods) and two habitat variables (percentage of pool habitat and percentage of gravel substrate) that each explained a significant portion of the annual variation in fish assemblage structure. For alien taxa, their proportional contribution to the total fish assemblage was inversely related to mean annual streamflow, one-day maximum discharge in both winter and spring, and the frequency of springtime floods. Results of this study highlight the need for continuous annual monitoring of streams with highly variable flow regimes to evaluate shifts in fish community structure. Apparent successes or failures in stream management may appear differently depending on the time series of available data.     

Quantifying the global threat to native birds from predation by non-native birds on small islands

Although invasive non-native species can adversely affect biodiversity in many ways, predation of native species by non-native species on islands can be severely damaging. Results of numerous studies document non-native birds preying on birds on islands, but our understanding of the number and type of species affected has been limited by the lack of a global review of these impacts. I identified the non-native bird species that have been recorded preying on birds, the locations where this predation occurred, and the bird species affected. Because the impacts of non-native birds can be particularly severe on small islands, I then identified the islands <500 km² around the world that are occupied by predatory non-native birds. By taking into account their life-history traits and predation history, I also identified the near-threatened and threatened bird species on these islands that they may prey on. The results indicated that predation by non-native birds was primarily a concern for threatened bird conservation on small islands; almost all predation impacts (91%) on near-threatened and threatened birds were recorded on islands, and median island size was 106 km². I also found non-native bird predation was a poorly known and widespread potential threat to avian biodiversity; worldwide, 194 islands of <500 km² were occupied by predatory non-native birds, but information on their impacts was unavailable for most of these islands. On them, where the impacts of non-native species can be severe, non-native birds may be preying on approximately 6% of the world's near-threatened and threatened bird species. Four non-native bird species I identified have been successfully eradicated from islands. If they were eradicated from the small islands they occupy, 70% of the near-threatened and threatened bird species I identified would no longer be affected by nest predation by non-native birds on small islands.     

Cryptobenthic fishes of the “Ciclopi Islands” marine reserve (central Mediterranean Sea): assemblage composition,structure and relations with habitat features

The fish assemblage of the Ciclopi Islands marine reserve, which is located along the eastern coast of Sicily (Central Mediterranean), was unknown until our study provided insight into the cryptobenthic fish assemblage inhabiting this integral reserve zone. Visual census surveys were periodically conducted along the littoral rocky shores of Lachea, the largest island of the Ciclopi archipelago. The fish assemblage, which comprised 20 species belonging to Blenniidae, Gobiidae, Tripterygiidae, Scorpaenidae and Gobiesocidae, was dominated by Scorpaena maderensis and Trypterigion delaisi. Species richness (S) did not show significant differences between the northern, eastern, southern and western island coasts, whereas both species diversity (H) and evenness (J) were significantly higher on the western coast than on the northern coast. The effect of the island coast on fish density was not significant with regard to the whole assemblage or to single species, except for S. maderensis and T. delaisi. The pattern of variation of assemblage parameters was mainly positively correlated with microhabitat variables, especially substratum composition and type of cover. The best predictors of fish total density were bottom slope, rugosity and substratum composition. Variations in relative species density were specified by a different combination of macro- and microhabitat variables. The density pattern of S. maderensis showed a positive relationship with depth. In T. delaisi, the influence of slope was interrelated to that of exposure to waves. This species preferentially inhabited flat habitat, sheltered from the open sea and covered by small boulders, but also more-exposed rockwalls. The population densities of Tripterygion melanurus and Lipophrys trigloides were negatively affected by depth, thus reflecting their preference for the upper infralittoral. T. melanurus was associated with a more complex habitat (stones and boulders), whereas L. trigloides was mainly censused on a narrow strip of solid rock between the waterline and 1 m depth.Communicated by R. Cattaneo-Vietti, Genova     

Threats to Avifauna on Oceanic Islands

Abstract:  Results of the study by Blackburn et al. (2004 a ) of avifauna on oceanic islands suggest that distance from the mainland and time since European colonization have major influences on species extinctions and that island area is a significant but secondary contributing factor. After augmenting the data of the study on geographical properties for some of the islands they examined, we used a causal analysis approach with structural equation modeling to reexamine their conclusions. In our model geographical properties of islands, such as island area and isolation, were considered constraints on biological factors, such as the number of introduced mammalian predators and existing number of avifauna, that can directly or indirectly influence extinction. Of the variables we tested, island area had the greatest total influence on the threat of extinction due to its direct and indirect effects on the size of island avifauna. Larger islands had both a greater number of threatened bird species and more avifauna, increasing the number of species that could become threatened with extinction. Island isolation also had a significant, positive, and direct effect on threats to island avifauna because islands farther from the mainland had fewer current extant avifauna. Time since European colonization had a significant negative, but relatively weaker, influence on threats compared with the traditional biogeographic factors of island area and distance to the mainland. We also tested the hypothesis that the amount of threat is proportionally lower on islands that have had more extinctions (i.e., there is a "filter effect"). Because the proportion of bird extinctions potentially explained only 2.3% of the variation in the proportion of threatened species on islands, our results did not support this hypothesis. Causal modeling provided a powerful tool for examining threat of extinction patterns of known and hypothesized pathways of influence.     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20 %, 22 % and 50 %, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53 %, 38 % and 17 %. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characterized byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition. Nomenclature: Van der Meijden et al. (1992) for phanerogams; Corley et al. (1981) for mosses; Grolle (1983) for hepatics.     

Drivers of inter-year variability of plant production and decomposers across contrasting island ecosystems

Despite the likely importance of inter-year dynamics of plant production and consumer biota for driving community- and ecosystem-level processes, very few studies have explored how and why these dynamics vary across contrasting ecosystems. We utilized a well-characterized system of 30 lake islands in the boreal forest zone of northern Sweden across which soil fertility and productivity vary considerably, with larger islands being more fertile and productive than smaller ones. In this system we assessed the inter-year dynamics of several measures of plant production and the soil microbial community (primary consumers in the decomposer food web) for each of nine years, and soil microfaunal groups (secondary and tertiary consumers) for each of six of those years. We found that, for measures of plant production and each of the three consumer trophic levels, inter-year dynamics were strongly affected by island size. Further, many variables were strongly affected by island size (and thus bottom-up regulation by soil fertility and resources) in some years, but not in other years, most likely due to inter-year variation in climatic conditions. For each of the plant and microbial variables for which we had nine years of data, we also determined the inter-year coefficient of variation (CV), an inverse measure of stability. We found that CVs of some measures of plant productivity were greater on large islands, whereas those of other measures were greater on smaller islands; CVs of microbial variables were unresponsive to island size. We also found that the effects of island size on the temporal dynamics of some variables were related to inter-year variability of macroclimatic variables. As such, our results show that the inter-year dynamics of both plant productivity and decomposer biota across each of three trophic levels, as well as the inter-year stability of plant productivity, differ greatly across contrasting ecosystems, with potentially important but largely overlooked implications for community and ecosystem processes.     

Tropical coastal habitats as surrogates of fish community structure, grazing, and fisheries value

Habitat maps are frequently invoked as surrogates of biodiversity to aid the design of networks of marine reserves. Maps are used to maximize habitat heterogeneity in reserves because this is likely to maximize the number of species protected. However, the technique's efficacy is limited by intra-habitat variability in the species present and their abundances. Although communities are expected to vary among patches of the same habitat, this variability is poorly documented and rarely incorporated into reserve planning. To examine intra-habitat variability in coral-reef fishes, we generated a data set from eight tropical coastal habitats and six islands in the Bahamian archipelago using underwater visual censuses. Firstly, we provide further support for habitat heterogeneity as a surrogate of biodiversity as each predefined habitat type supported a distinct assemblage of fishes. Intra-habitat variability in fish community structure at scales of hundreds of kilometers (among islands) was significant in at least 75% of the habitats studied, depending on whether presence/absence, density, or biomass data were used. Intra-habitat variability was positively correlated with the mean number of species in that habitat when density and biomass data were used. Such relationships provide a proxy for the assessment of intra-habitat variability when detailed quantitative data are scarce. Intra-habitat variability was examined in more detail for one habitat (forereefs visually dominated by Montastraea corals). Variability in community structure among islands was driven by small, demersal families (e.g., territorial pomacentrid and labrid fishes). Finally, we examined the ecological and economic significance of intra-habitat variability in fish assemblages on Montastraea reefs by identifying how this variability affects the composition and abundances of fishes in different functional groups, the key ecosystem process of parrotfish grazing, and the ecosystem service of value of commercially important finfish. There were significant differences in a range of functional groups and grazing, but not fisheries value. Variability at the scale of tens of kilometers (among reefs around an island) was less than that among islands. Caribbean marine reserves should be replicated at scales of hundreds of kilometers, particularly for species-rich habitats, to capture important intra-habitat variability in community structure, function, and an ecosystem process.     

Understanding Trait‐Dependent Community Disassembly: Dung Beetles,Density Functions,and Forest Fragmentation

Abstract: Anthropogenic disturbances such as fragmentation are rapidly altering biodiversity, yet a lack of attention to species traits and abundance patterns has made the results of most studies difficult to generalize. We determined traits of extinction‐prone species and present a novel strategy for classifying species according to their population‐level response to a gradient of disturbance intensity. We examined the effects of forest fragmentation on dung beetle communities in an archipelago of 33 islands recently created by flooding in Venezuela. Species richness, density, and biomass all declined sharply with decreasing island area and increasing island isolation. Species richness was highly nested, indicating that local extinctions occurred nonrandomly. The most sensitive dung beetle species appeared to require at least 85 ha of forest, more than many large vertebrates. Extinction‐prone species were either large‐bodied, forest specialists, or uncommon. These explanatory variables were unrelated, suggesting at least 3 underlying causes of extirpation. Large species showed high wing loading (body mass/wing area) and a distinct flight strategy that may increase their area requirements. Although forest specificity made most species sensitive to fragmentation, a few persistent habitat generalists dispersed across the matrix. Density functions classified species into 4 response groups on the basis of their change in density with decreasing species richness. Sensitive and persistent species both declined with increasing fragmentation intensity, but persistent species occurred on more islands, which may be due to their higher baseline densities. Compensatory species increased in abundance following the initial loss of sensitive species, but rapidly declined with increasing fragmentation. Supertramp species (widespread habitat generalists) may be poor competitors but strong dispersers; their abundance peaked following the decline of the other 3 groups. Nevertheless, even the least sensitive species were extirpated or rare on the smallest and most isolated islands.     

Evidence of dispersal limitation in soil microorganisms: isolation reduces species richness on mycorrhizal tree islands

Dispersal limitation plays an important role in a number of equilibrium and nonequilibrium theories about community ecology. In this study we use the framework of island biogeography to look for evidence of dispersal limitation in ectomycorrhizal fungal assemblages on "tree islands," patches of host trees located in a non-host vegetation matrix. Because of the potentially strong effects of island area on species richness and immigration, we chose to control island size by sampling tree islands consisting of a single host individual. Richness on tree islands was high, with estimates ranging up to 42 species of ectomycorrhizal fungi associating with a single host individual. Species richness decreased significantly with increasing isolation of tree islands, with our regression predicting a 50% decrease in species richness when tree islands are located distances of approximately 1 km from large patches of contiguous forests. Despite the fact that fungal fruit bodies produce large numbers of spores with high potential for long-distance travel, these results suggest that dispersal limitation is significant in ectomycorrhizal assemblages. There were no discernible effects of isolation or environment on the species identity of tree island fungal colonists. In contrast to the highly predictable patterns of tree island colonization we observed in a previous study on early successional forests, we suggest that over longer time periods the community assembly process becomes more dominated by stochastic immigration and local extinction events.     

Evaluating day–night changes in shallow Mediterranean rocky reef fish assemblages by visual census

Ecological information on coastal fish distribution patterns and habitat use can be greatly improved by nocturnal samplings and observations. To this purpose, the structure of a Mediterranean fish assemblage inhabiting the shallow rocky littoral of Linosa Island (Sicily Strait, Italy) was examined by using visual census to detect possible diel variations in species composition and abundance. Day–night fish distribution patterns were investigated by multivariate and univariate analyses. Overall, 42 fish taxa belonging to 19 families were recorded: 35 during the day and 24 during the night. Seventeen species were common to both diurnal and nocturnal assemblages. Within the diurnal assemblage, Chromis chromis was the most represented species (37.2%), followed by Thalassoma pavo (23.2%) and Sparisoma cretense (10.8%). Within the nocturnal assemblage, the most abundant taxon was Atherina spp. (33.9%), followed by Apogon imberbis (26.4%) and Boops boops (11.5%). Our results indicated wide variation in the abundance and species composition during the day and during the night. Multi Dimensional Scaling plot showed a clear-cut separation between the two assemblages and analysis of similarities found significant differences as well. SIMPER analysis revealed that ten species individually contributed by more than 2.5% to the dissimilarity between diurnal and nocturnal assemblages, T. pavo, C. chromis and S. cretense being the first three species in order of decreasing percentage. ANOVA performed on species richness and fish abundance detected significant differences between diurnal and nocturnal assemblages, the latter showing far lower average values for both variables.     

Grazing as a measure against grass encroachment in Dutch dry dune grassland: effects on vegetation and soil

A grazing experiment was started in 1984 and 1989 respectively, in two parts of a dune grassland in the nature reserve ‘Zwanenwater’, North Holland; a third part with similar geology and topography was used as a control area and not grazed. An evaluation of the effects of grazing on vegetation patterns, species composition, vegetation structure and humus form was made with the help of vegetation maps from 1986 and 1992 as well as field surveys. Dense tall-grass communities dominated byAmmophila arenaria increased over the period 1986–1992 in the grazed areas, and especially in the non-grazed area (increase in area to 20%, 22% and 50%, respectively). Open communities decreased in the grazed areas, but are still prevalent, while in the ungrazed area they virtually disappeared, with the result that the present percentage areas are 53%, 38% and 17%. Field survey data were classified by TWINSPAN producing four vegetation types. These occur more or less equally in grazed and ungrazed areas, albeit with different percentage areas: (1) open vegetation dominated byCorynephorus canescens; (2) open vegetation characteized, byKoeleria macrantha; (3) heathland dominated byEmpetrum nigrum; and (4) tall-grass communities dominated byAmmophila arenaria. Within a vegetation type, species composition was only marginally affected by grazing regime. Within the open communities the number of species, vegetation height, vegetation cover and soil organic horizons were not affected by grazing. In the tall-grass communities the number of species was significantly larger and the height of the vegetation significantly lower in the area grazed since 1984. In the heathland community the number of species and cover of the moss layer were significantly higher in the 1984 area and ectorganic and endorganic horizons significantly thicker in the ungrazed area. It is suggested that these effects are the result of an increased availability of light, but possibly also of a decreased stock of organic matter and nutrients, due to a decreased input of litter and accelerated rates of decomposition.     

Short-term variation and long-term changes in the oceanographic environment and zooplankton community in the vicinity of a sub-Antarctic archipelago

 Mesozooplankton community structure in the vicinity of the Prince Edward Islands (PEIs) was investigated during six surveys conducted in late austral summer (April/May) from 1996 to 1999. Zooplankton samples were collected by oblique tows using a Bongo net fitted with 300-μm mesh. Surface temperature, average temperature and chlorophyll a were measured in conjunction with each net tow. The positions of the Sub-Antarctic Front (SAF) and the Antarctic Polar Front (APF), in relation to the islands, were determined by CTD and/or XBT transects to the west of the islands (upstream). Both fronts were characterized by a high degree of latitudinal variation. Changes in position of the fronts occurred rapidly, the SAF moving up to ∼120 km in a 2-week period. Consequently, the oceanographic environment in the vicinity of the PEIs was subject to a high degree of intra- and inter-survey variation. The positions of the SAF and APF appeared to have a significant impact on phytoplankton biomass in the vicinity of the PEIs, possibly through the alteration of local oceanographic flow dynamics. Water retention over the island shelf in 1996, associated with location of the SAF far to the north of the PEIs, corresponded to enhanced chlorophyll-a concentrations (∼1.54 mg m⁻³). Conversely, when the fronts were close to the islands, as in 1997 and 1999, higher current velocity limited water retention and chlorophyll-a concentrations in the inter-island region were relatively low (∼0.4 mg m⁻³). Cluster analyses showed that, in many instances, there was greater similarity among zooplankton communities from different surveys than among communities within surveys, indicating that short-term variability exceeded inter-annual variability. The population structure of the copepod Calanus simillimus indicated that there was inter-annual variation in the timing of the biological season. Differences in the population structure of species, and consequently their contribution to abundance and biomass, may therefore have been an important contributor to inter-annual variation in community structure. Evidence is provided of a long-term southward shift in the position of the SAF. It is postulated that this may affect the PEIs by increasing the proportion of allochthonous energy input, because the PEIs now lie in the path of the front, altering the tropho-dynamics of the island ecosystem. Lower mesozooplankton biomass associated with warmer sub-Antarctic water may have important negative consequences for higher trophic levels that depend on mesozooplankton for food. Received: 10 June 2000 / Accepted: 22 September 2000     

Genetic variation in the mangrove periwinkle Littorina angulifera

Twenty populatios of Littorina angulifera, inhabiting islands composed of the mangrove tree Rhizophora mangle, were assayed at an esterase locus to determine whether genetic differentiation was associated with distance between populations. It was predicted, on the basis of larval dispersal in this species, that genetic differentiation between populations on islands separated by long distances should be greater than those on islands located near each other. A chi0square test of homogeneity revealed significant differences in esterase gene frequencies among the 20 island populations. However, there was no association of distance between islands and genetic heterogeneity. In addition, a cline in gene frequency was found to be assiciated with latitude. Factors responsible for the observed pattern of heterogeneity at the esterase locus are discussed.     

Scale-dependent variation in coral community similarity across sites, islands, and island groups

Community similarity is the proportion of species richness in a region that is shared on average among communities within that region. The slope of local richness (alpha diversity) regressed on regional richness (gamma diversity) can serve as an index of community similarity across regions with different regional richness. We examined community similarity in corals at three spatial scales (among transects at a site, sites on an island, and islands within an island group) across a 10 000-km longitudinal diversity gradient in the west-central Pacific Ocean. When alpha diversity was regressed on gamma diversity, the slopes, and thus community similarity, increased with scale (0.085, 0.261, and 0.407, respectively) because a greater proportion of gamma diversity was subsumed within alpha diversity as scale increased. Using standard randomization methods, we also examined how community similarity differed between observed and randomized assemblages and how this difference was affected by spatial separation of species within habitat types and specialization of species to three habitat types (reef flats, crests, and slopes). If spatial separation within habitat types and/or habitat specialization (i.e., underdispersion) occurs, fewer species are shared among assemblages than the random expectation. When the locations of individual coral colonies were randomized within and among habitat types, community similarity was 46-47% higher than that for observed assemblages at all three scales. We predicted that spatial separation of coral species within habitat types should increase with scale due to dispersal/extinction dynamics in this insular system, but that specialization of species to different habitat types should not change because habitat differences do not change with scale. However, neither habitat specialization nor spatial separation within habitat types differed among scales. At the two larger scales, each accounted for 22-24% of the difference in community similarity between observed and randomized assemblages. At the smallest scale (transect-site), neither spatial separation within habitat types nor habitat specialization had significant effects on community similarity, probably due to the small size of transect samples. The results suggest that coral species can disperse among islands in an island group as easily as they can among sites on an island over time scales that are relevant to their establishment and persistence on reefs.