Local Species Diversity Research Articles

Fish community composition and diversity at restored estuarine habitats in Tampa Bay, Florida, United States

Estuary restoration in Tampa Bay, Florida, United States, is an ongoing focus of natural resource managers because of pressure from an increasing coastal population, historic habitat loss, and restoration's importance to economic development, recreational activities, and fish habitat. A growing population can also limit future large‐scale restorations due to associations with cost and land availability. This limitation might be overcome by applying the habitat mosaic approach to restoration, which creates distinct habitat types at small spatial scales. This approach was applied to create three types of estuarine habitat, reconnected tidal creek, salt marsh, and tidal pond. The objectives of this study were to (1) initiate monitoring of a restored wetland mosaic and (2) determine how fish diversity and community structure vary among restored habitat types. Replicated sampling using a 3‐mm mesh seine was used to characterize the fish communities. Our results indicate that the habitat mosaic approach creates suitable habitat for a variety of fish species where 37% of fish species were captured in just one habitat type. In particular, the recreationally important Centropomus undecimalis (common snook) was more common in the mangrove‐lined creek and the non‐native Sarotherodon melanotheron (blackchin tilapia) was common in the tidal pond. Greater emphasis should be placed on applied restoration research to identify how habitat types within a larger restoration mosaic contribute to local species diversity and recreationally and commercially important fishes, while limiting non‐natives. This emphasis could reveal how restoration approaches can be modified to include habitat mosaics, maximizing their contribution to productive fish habitat.

Functional traits and environmental characteristics drive the degree of competitive intransitivity in European saltmarsh plant communities

Abstract Competitive intransitivity, the existence of loops in competitive hierarchies, is one mechanism that can promote the local coexistence of competitors and maintain high local species diversity, although its prevalence and importance remain largely unknown. A full understanding of local community assembly needs knowledge of how transitive and intransitive competitive interactions are linked to species functional traits and the strength of biotic and abiotic filters. We apply a recently developed statistical tool to quantitative data on central European inland saltmarsh plant communities to infer causal relationships between soil characteristics, species occurrences and functional traits, and we estimated coefficients of competition. We found a predominance of intransitive competitive hierarchies. The proportion of such hierarchies was positively correlated with local species richness and compositional variability. Average soil characteristics were not correlated with competitive intransitivity, whereas high soil pH and the high variability in local pH and soil salinity decreased the overall impact of competition on community composition. In pairwise comparisons of species, dissimilarity in morphology, resource demand and reproductive phenology was significantly negatively correlated with differences in competitive performance, while higher environmental dissimilarity was particularly linked to intermediate degrees of competitive superiority. Our results suggest that habitat filtering for similar traits might intensify competitive interactions, but might also give rise to intransitive competitive loops that subsequently promote species coexistence and permit species’ functional equivalence. Intransitive competition appears to increase local diversity and small scale‐species turnover. The observed local differences in competitive structures suggest frequent competitive plasticity and context‐dependent competitive interactions. Finally, our results support the view that local abundance distributions can be used to infer the strength and outcome of competitive interactions. Synthesis. Our results confirm that intransitive competitive interactions might be a strong force structuring local plant communities. Intransitivity needs to be considered when studying plant community assembly and species co‐existence.

Impact of stochastic migration on species diversity in meta-food webs consisting of several patches

The structure of space has an appreciable influence on the diversity and stability of ecosystems. So far, there are only few theoretical studies investigating the population dynamics of food webs consisting of many species that can migrate between several patches, and in most of these models migration is a continuous, deterministic process. However, when migration events are rare (for instance because the patches are far apart), migration is a stochastic process and should be modeled accordingly. We present computer simulations of a food web model of many species on a spatial network of several patches, combining deterministic local population dynamics with stochastic migration. We evaluate the influence of the migration rate and other model parameters on local and regional species diversity and on stability. We find that migration increases the number of surviving and coexisting populations by two effects. These are the rescue effect, which restores local populations that have gone extinct, and dynamical coexistence, which sustains local populations that could not persist in the absence of immigration. Both effects occur even when migration events are rare. Species diversity increases on local and regional scales with the frequency of migration events. Furthermore, we investigate the adiabatic limit in which population dynamics always reaches an equilibrium before the next migration event, and we investigate the possible long-term scenarios. While the final state often contains the same food web on all patches, we also find instances where two slightly different food webs coexist on different patches, even when initially each patch contained the same food web.

What shapes plant and animal diversity on urban golf courses?

Recent concern over increasing loss of biodiversity has prompted considerable interest in the role of urban green spaces as reservoirs of local biodiversity. This study assessed the diversity of three indicator taxa - plants, ants and birds - on golf courses spanning a wide range of environmental variation in terms of climate, elevation, course age, size and connectivity to native woodland. Species richness and community composition was further compared between contrasting on-course habitat types that reflect different management intensities. We identified a set of taxon-specific environmental correlates indicating an intricate interplay of landscape- and local-scale variables that affect local species diversity. Our results show that floristic diversity is positively associated with the amount of rainfall, whereas ant and bird diversity are related to local-scale factors, particularly the number of trees and the size of water features on a site. The amount of on-course native habitat was a strong predictor of plant and ant diversity and was also associated with the number of unique species at the site level; this reinforces the value of remnant habitat patches as local biodiversity reservoirs that represent mini hot-spots in an otherwise species-poor urban landscape. Community composition for all three taxa differed markedly between non-playing and playing areas, with boundary and remnant habitats generally having more diverse, species-rich communities. Our results suggest that local floral and faunal biodiversity on urban golf courses can be enhanced by creating woody non-playing areas and, especially, by preserving, restoring or expanding remnant habitats.

Regional forcing explains local species diversity and turnover on tropical islands

AbstractAimTo determine the role of regional forcing on plot‐level species diversity and composition, and to quantify the relative importance of biogeographical and climatic factors in explaining woody plant diversity and composition at the local‐, island‐ and archipelago‐scale.LocationForty‐one tropical islands of the Indo‐Pacific region from Madagascar to Hawai‘i Island.MethodsWe analysed the diversity and composition of tropical woody plant communities located across 113 plots, 41 islands and 19 archipelagos. We used generalized linear mixed‐effects models and generalized dissimilarity models to determine the role of regional forcing at the island and archipelago scale and to assess the relative importance of biogeographical (area and isolation of islands or archipelagos, geographical distance between plots) and climatic factors in explaining differences in local diversity and composition (species turnover). Analyses were conducted at different geographical scales (local, island and archipelago) and taxonomic levels (species, genus and family).ResultsVariation in local (plot‐level) diversity (as species density, the number of species per 100 woody plants) was primarily explained by island and archipelago identity. Maximum species density was positively correlated with the area of an island (or archipelago) and negatively correlated with the isolation of an archipelago. Local climatic variability was also a significant predictor of species density, but less important than regional forcing. Climate variables explained < 20% of the variation in species turnover across all plots. The importance of geographical distance between plots relative to climate in driving species turnover decreased from the species to family level, and from the regional to island level.Main conclusionsRegional forcing was the key driver of local diversity and composition on islands. Island area and archipelago isolation are likely driving local diversity through their effects on the pool of island species. Geographical distance between plots is the main factor explaining species turnover, while at higher taxonomic levels, climatic factors and niche conservatism are the main drivers.

Seed dispersal in both space and time is necessary for plant diversity maintenance in fragmented landscapes

Metacommunity theory emphasizes that seed dispersal not only limits but equally maintains plant diversity, though the latter receives little empirical attention. Discerning the temporal and spatial components of seed dispersal and understanding how their interaction shapes fragmented communities and maintains their diversity may be pivotal to further our ecological understanding of spatial and temporal seed dispersal and its implications for landscape‐scale conservation management. To investigate the relative importance of spatial and temporal seed dispersal and their roles in maintaining plant diversity, the herb layer and seed bank of grassland communities were inventoried in 77 sites across abandoned and intact rotational grazing networks in a 100 km2 fragmented grassland landscape in the Stockholm archipelago (Baltic Sea, Sweden). Besides analysing alpha‐ and beta‐diversity patterns, nestedness analyses connect deterministic community changes and diversity losses with dispersal‐related life‐history traits and habitat specialization to identify the mechanism driving community changes and maintaining local diversity. The loss of rotational grazing networks caused community diversity declines via non‐random extinctions of spatially and temporally poor dispersers, particularly among grassland specialists. Temporal seed dispersal halted further community disassembly, maintaining diversity in the abandoned grazing networks. Spatial dispersal within the intact grazing networks was found to be an overriding, homogenizing agent conserving diversity in both the herb layer and seed bank. This empirical evidence establishes how spatial and temporal seed dispersal interact to maintain diversity in fragmented landscapes. Poorly connected grasslands appear limited by spatial dispersal, yet are maintained by temporal seed dispersal. In fragmented landscapes where grazing networks are rarely present, temporal rather than spatial seed dispersal may be more important in maintaining species diversity, since effective spatial dispersal may be significantly diminished. The grazing network's efficacy at boosting spatial dispersal and upholding community diversity presents a powerful management tool to conserve local and regional species diversity.

Effects of pond management on biodiversity patterns and community structure of zooplankton in urban environments

As urban areas continue expanding, major cities become connected forming megacities. Urban encroachment into natural areas transforms the landscape into a built environment with heterogeneously distributed patches of novel habitat. Community structure within novel habitats is influenced by anthropogenic factors including fragmentation and species interactions. Alterations in complex biodiversity patterns may be used to assess how urban stressors impact community assemblages which, ultimately, may inform sustainable management decisions. To manage algal blooms, Aquashade® is applied directly to ponds. We investigated the effects of Aquashade®, nutrient loading and dispersal on local species diversity and compositional turnover of zooplankton communities from suburban ponds in Columbia, MD, USA using a mesocosm approach. We found that Aquashade® acted as an environmental filter by increasing local species diversity and decreasing compositional turnover. This ultimately could have an overall homogenizing effect on the regional species pool (or γ-diversity). The same pattern was observed in mesocosms that received simulated dispersal events of zooplankton. Nutrients, overall, increased autotrophic biomass and while Aquashade® had no effect on autotrophic biomass, the interaction of nutrients and Aquashade® similarly caused a homogenization of the zooplankton community. Additionally, there was an overall increase in cladoceran ephippia in mesocosms receiving Aquashade® compared to those not, suggesting there is a ‘trigger’ switching cladocerans from parthenogenic to sexual reproduction. Taken together, our results show the application of Aquashade®, nutrient loading and dispersal shift biodiversity patterns in urban zooplankton communities. We hypothesize these shifts originate at the resource level through alterations in the phytoplankton community either through composition or nutritive value in ponds receiving Aquashade®. Our study illustrates the importance of investigating multiple-scales of community assemblages when assessing impacts of anthropogenic stressors. Consideration of how to best manage green spaces may include planned un-managed areas to maintain regional biodiversity and heterogenous communities within the urban environment.

A multispecies approach reveals hot spots and cold spots of diversity and connectivity in invertebrate species with contrasting dispersal modes.

Genetic diversity is crucial for species' maintenance and persistence, yet is often overlooked in conservation studies. Species diversity is more often reported due to practical constraints, but it is unknown if these measures of diversity are correlated. In marine invertebrates, adults are often sessile or sedentary and populations exchange genes via dispersal of gametes and larvae. Species with a larval period are expected to have more connected populations than those without larval dispersal. We assessed the relationship between measures of species and genetic diversity, and between dispersal ability and connectivity. We compiled data on genetic patterns and life history traits in nine species across five phyla. Sampling sites spanned 600km in the northwest Mediterranean Sea and focused on a 50-km area near Marseilles, France. Comparative population genetic approaches yielded three main results. (i) Species without larvae showed higher levels of genetic structure than species with free-living larvae, but the role of larval type (lecithotrophic or planktotrophic) was negligible. (ii) A narrow area around Marseilles, subject to offshore advection, limited genetic connectivity in most species. (iii) We identified sites with significant positive contributions to overall genetic diversity across all species, corresponding with areas near low human population densities. In contrast, high levels of human activity corresponded with a negative contribution to overall genetic diversity. Genetic diversity within species was positively and significantly linearly related to local species diversity. Our study suggests that local contribution to overall genetic diversity should be taken into account for future conservation strategies.

Multiscale change in reef coral species diversity and composition in the Tropical Eastern Pacific

Both natural and anthropogenic factors are changing coral-reef structure and function worldwide. Long-term monitoring has revealed declines in the local composition and species diversity of reefs. Here we report changes in coral-reef community structure over 12 yr (2000–2012) at 17 sites and three spatial scales (reef, gulf and country) in the Tropical Eastern Pacific (Panama). We found a significant 4% annual decline in species population sizes at the country and gulf scales, with significant declines ranging from 3 to 32% at all but one reef. No significant temporal change in expected richness was found at the country scale or in the Gulf of Chiriqui, but a 7% annual decline in expected species richness was found in the Gulf of Panama. There was a 2% increase in community evenness in the Gulf of Chiriqui, but no change in the Gulf of Panama. Significant temporal turnover was found at the country and gulf scales and at 29% of the reefs, a finding mostly explained by changes in species abundance, and losses and gains of rare species. Temporal trends in alpha and beta diversity metrics were explained by water temperature maxima, anomalies and variation that occurred even in the absence of a strong El Nino warming event.

Effects of habitat manipulation by red grouper, Epinephelus morio, on faunal communities associated with excavations in Florida Bay

Organisms that either directly or indirectly modify the availability of resources can have profound effects on the structure and function of ecological communities. This holds true for organisms that enhance habitat complexity either as a function of their morphology or through mechanical activities, such as digging or burrowing. The net positive effects of engineers on local species diversity follows from activities that allow more species to live in modified habitats compared to unmodified ones. Here, we present the results of a multiyear observational study evaluating the effects of habitat modifications made by juvenile red grouper, Epinephelus morio (Valenciennes, 1828), on the fishes and motile macroinvertebrates associated with karst solution holes in Florida Bay, FL, USA. Because red grouper have multiple roles in these systems, acting as both habitat engineers and predators that consume a variety of benthic crustaceans and fishes, their influence on associated communities is complex. We found that the presence of red grouper resulted in deeper solution holes, increased abundance and richness of the associated fauna, especially among juvenile coral-reef fishes and motile invertebrates, and significant differences in the structure of faunal communities over time. These results suggest that red grouper shape unique communities, distinct from those of surrounding areas, and strongly influence the composition of communities associated with manipulated habitats.

Dimensions of biodiversity loss: Spatial mismatch in land-use impacts on species, functional and phylogenetic diversity of European bees.

AimAgricultural intensification and urbanization are important drivers of biodiversity change in Europe. Different aspects of bee community diversity vary in their sensitivity to these pressures, as well as independently influencing ecosystem service provision (pollination). To obtain a more comprehensive understanding of human impacts on bee diversity across Europe, we assess multiple, complementary indices of diversity.LocationOne Thousand four hundred and forty six sites across Europe.MethodsWe collated data on bee occurrence and abundance from the published literature and supplemented them with the PREDICTS database. Using Rao's Quadratic Entropy, we assessed how species, functional and phylogenetic diversity of 1,446 bee communities respond to land‐use characteristics including land‐use class, cropland intensity, human population density and distance to roads. We combined these models with statistically downscaled estimates of land use in 2005 to estimate and map—at a scale of approximately 1 km2—the losses in diversity relative to semi‐natural/natural baseline (the predicted diversity of an uninhabited grid square, consisting only of semi‐natural/natural vegetation).ResultsWe show that—relative to the predicted local diversity in uninhabited semi‐natural/natural habitat—half of all EU27 countries have lost over 10% of their average local species diversity and two‐thirds of countries have lost over 5% of their average local functional and phylogenetic diversity. All diversity measures were generally lower in pasture and higher‐intensity cropland than in semi‐natural/natural vegetation, but facets of diversity showed less consistent responses to human population density. These differences have led to marked spatial mismatches in losses: losses in phylogenetic diversity were in some areas almost 20 percentage points (pp.) more severe than losses in species diversity, but in other areas losses were almost 40 pp. less severe.Main conclusionsThese results highlight the importance of exploring multiple measures of diversity when prioritizing and evaluating conservation actions, as species‐diverse assemblages may be phylogenetically and functionally impoverished, potentially threatening pollination service provision.

Sapling growth rates reveal conspecific negative density dependence in a temperate forest.

Local tree species diversity is maintained in part by conspecific negative density dependence (CNDD). This pervasive mechanism occurs in a variety of forms and ecosystems, but research to date has been heavily skewed toward tree seedling survival in tropical forests. To evaluate CNDD more broadly, we investigated how sapling growth rates were affected by conspecific adult neighbors in a fully mapped 25.6 ha temperate deciduous forest. We examined growth rates as a function of the local adult tree neighborhood (via spatial autoregressive modeling) and compared the spatial positioning of faster‐growing and slower‐growing saplings with respect to adult conspecific and heterospecific trees (via bivariate point pattern analysis). In addition, to determine whether CNDD‐driven variation in growth rates leaves a corresponding spatial signal, we extended our point pattern analysis to a static, growth‐independent comparison of saplings and the next larger size class. We found that negative conspecific effects on sapling growth were most prevalent. Five of the nine species that were sufficiently abundant for analysis exhibited CNDD, while only one species showed evidence of a positive conspecific effect, and one or two species, depending on the analysis, displayed heterospecific effects. There was general agreement between the autoregressive models and the point pattern analyses based on sapling growth rates, but point pattern analyses based on single‐point‐in‐time size classes yielded results that differed markedly from the other two approaches. Our work adds to the growing body of evidence that CNDD is an important force in temperate forests, and demonstrates that this process extends to sapling growth rates. Further, our findings indicate that point pattern analyses based solely on size classes may fail to detect the process of interest (e.g., neighborhood‐driven variation in growth rates), in part due to the confounding of tree size and age.

The invasive herb Lupinus polyphyllus attracts bumblebees but reduces total arthropod abundance

Invasive plant species generally reduce the abundance and diversity of local plant species, which may translate into alterations at higher tropic levels, such as arthropods. Due to the diverse functional roles of arthropods in the ecosystems, it is critical to understand how arthropod communities are affected by plant invasions. Here, we investigated the impact of the invasive ornamental herb Lupinus polyphyllus (Lindl.) on arthropod communities during its main flowering period in southwestern Finland over two years. The total number of arthropods was about 46% smaller at the invaded sites than at the uninvaded sites in both study years, and this difference was mainly due to a lower abundance of beetles, Diptera, Lepidoptera, and ants. However, the number of bumblebees (particularly Bombus lucorum) was about twice as high at invaded sites compared with uninvaded sites, even though bumblebee richness did not differ between sites. There was no statistically significant difference between invaded and uninvaded sites in the abundances of the other arthropod groups considered (Hymenoptera (excluding bumblebees and ants), Hemiptera, and Arachnida). In addition, L. polyphyllus affected the relative abundance of four arthropod groups, with the order Lepidoptera being less common at invaded sites than at uninvaded sites, while the opposite was true for bumblebees, Hemiptera, and Arachnida. Overall, these results demonstrate that the negative impact of L. polyphyllus on biodiversity goes beyond its own trophic level, suggesting that this species has the potential to alter the abundance of different arthropod groups and, consequently, the structure of arthropod communities at a large scale.

Species Interactions Drive Fish Biodiversity Loss in a High-CO2 World

Accelerating climate change is eroding the functioning and stability of ecosystems by weakening the interactions among species that stabilize biological communities against change [1]. A key challenge to forecasting the future of ecosystems centers on how to extrapolate results from short-term, single-species studies to community-level responses that are mediated by key mechanisms such as competition, resource availability (bottom-up control), and predation (top-down control) [2]. We used CO2 vents as potential analogs of ocean acidification combined with insitu experiments to test current predictions of fish biodiversity loss and community change due to elevated CO2 [3] and to elucidate the potential mechanisms that drive such change. We show that high risk-taking behavior and competitive strength, combined with resource enrichment and collapse of predator populations, fostered already common species, enabling them to double their populations under acidified conditions. However, the release of these competitive dominants from predator control led to suppression of less common and subordinate competitors that did not benefit from resource enrichment and reduced predation. As a result, local biodiversity was lost and novel fish community compositions were created under elevated CO2. Our study identifies the species interactions most affected by ocean acidification, revealing potential sources of natural selection. We also reveal how diminished predator abundances can have cascading effects on local species diversity, mediated by complex species interactions. Reduced overfishing of predators could therefore act as a key action to stall diversity loss and ecosystem change in a high-CO2 world. VIDEO ABSTRACT.

Plant diversity in a water-meadow landscape: the role of irrigation ditches

Species-rich, extensively managed grasslands are among the most threatened ecosystems in Europe. The traditional technique of meadow irrigation, formerly practised in large parts of Europe, may be a viable management option that meets both ecological and economical objectives. While studies on plant diversity of the meadow matrix exist, the role of other landscape components for biodiversity conservation is unknown. We quantify the role of irrigation and drainage ditches compared with field paths, and analysed the local and landscape plant species composition and diversity in a water-meadow landscape. The meadows are located in the river Queich catchment (Upper Rhine valley, Germany). We recorded plant species cover on 18 meadows, adjacent ditches and field paths and analysed alpha, beta and gamma diversity within and among these landscape components. Irrigation and drainage ditches were more species rich than meadows and more heterogeneous in their species composition compared with meadows and paths. Ditches contributed one-third of the landscape’s plant species numbers, habitat for wetland species and refuges for high nature value grassland species. Fertilization of the adjacent meadow had no negative impact on the ditch diversity, while well-maintained ditches were more plant species rich than ditches filled with sediments. Traditional flood meadow irrigation should be taken into account when sustainable grassland management techniques are discussed that meet economical and ecological needs. A major contribution to landscape richness arises from the ditch structures as crucial part of the traditional irrigation technique.

Habitat heterogeneity hypothesis and edge effects in model metacommunities

Spatial heterogeneity is an inherent property of any living environment and is expected to favour biodiversity due to a broader niche space. Furthermore, edges between different habitats can provide additional possibilities for species coexistence. Using computer simulations, this study examines metacommunities consisting of several trophic levels in heterogeneous environments in order to explore the above hypotheses on a community level. We model heterogeneous landscapes by using two different sized resource pools and evaluate the combined effect of dispersal and heterogeneity on local and regional species diversity. This diversity is obtained by running population dynamics and evaluating the robustness (i.e., the fraction of surviving species). The main results for regional robustness are in agreement with the habitat heterogeneity hypothesis, as the largest robustness is found in heterogeneous systems with intermediate dispersal rates. This robustness is larger than in homogeneous systems with the same total amount of resources. We study the edge effect by arranging the two types of resources in two homogeneous blocks. Different edge responses in diversity are observed, depending on dispersal strength. Local robustness is highest for edge habitats that contain the smaller amount of resource in combination with intermediate dispersal. The results show that dispersal is relevant to correctly identify edge responses on community level.

Stochastic losses of fire-dependent endemic herbs revealed by a 65-year chronosequence of dispersal-limited woody plant encroachment.

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.

The difference between a rock and a biological hard place: epibionts in the rocky intertidal

In dynamic environments such as the rocky marine intertidal zone, epibiosis may promote the survival of some organisms. We compared the species composition and prevalence of the epibiotic community on different species of molluscan shells to those on neighboring rock at three sites and two tidal heights per site on San Juan Island, Washington (48°N, 123°W; summer 2014). Across all three sites and both tidal heights examined, the algal species dominating the bedrock did not match the dominant epibiotic species. Most epibiotic species were from a subset of the less common bedrock species. Within the lower tidal height, many potential host shells lacked epibionts, but the bedrock was heavily colonized. Our findings suggest that shells do not represent a microhabitat that is more beneficial to settlers than the surrounding bedrock. Epibiosis is unlikely to elevate local species diversity in this ecosystem because the few epibionts were generally from a subset of the species on the bedrock. Smooth-shelled limpets had a significantly greater prevalence of epibiosis in the lower than in the higher tidal height, but the prevalence of epibiosis did not differ significantly between tidal heights for carnivorous, spired-shell species. This variation among basibionts in pattern of prevalence across tidal heights suggests that the carnivorous snails are more likely than the limpets to range across intertidal heights.

Land-use intensification causes multitrophic homogenization of grassland communities.

Land-use intensification is a major driver of biodiversity loss. Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in β-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (α)-diversity and neglected biodiversity loss at larger spatial scales. Studies addressing β-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above- and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in α-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on β-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in β-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local α-diversity in aboveground groups, whereas the α-diversity increased in belowground groups. Correlations between the β-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.

Electric organ discharges of South African Marcusenius species (Teleostei: Mormyridae) and their effectiveness as indicators of local species diversity.

Recent morphological and genetic studies have revealed two new species of snoutfish in South Africa, Marcusenius caudisquamatus and M. krameri, which had been confused with M. pongolensis, the South African bulldog fish. All known mormyriform fish are nocturnal and emit electric organ discharges (EODs) for communication that are characteristic for their species. This paper examines whether or not the EODs of these three closely-related South African species can be differentiated from each other. An EOD pulse of a bulldog fish consists of a head-positive phase P, followed by a head-negative phase N of short duration. We measured and compared six variables of the EOD pulse waveform for South African samples for the three species from different locations using MANOVA, ANOVA and Discriminant Analysis, with M. devosi from Kenya as an outgroup. The EOD waveforms, normalized to the same P-phase amplitude, varied significantly from each other in four variables, most strongly in the amplitude of the N phase and the duration of the P phase. In two species, M. devosi and M. krameri, there was no evidence of difference between sexes, in contrast to M. pongolensis and M. caudisquamatus whose male pulses were of longer duration. M. devosi and M. krameri were statistically significantly independent of each other and of any other group studied. By contrast, the M. pongolensis specimens from different locations showed a high degree of variability amongst each other, including significant separation, and overlap with M. caudisquamatus.