New multidimensional functional diversity indices for a multifaceted framework in functional ecology

Functional diversity is increasingly identified as an important driver of ecosystem functioning. Various indices have been proposed to measure the functional diversity of a community, but there is still no consensus on which are most suitable. Indeed, none of the existing indices meets all the criteria required for general use. The main criteria are that they must be designed to deal with several traits, take into account abundances, and measure all the facets of functional diversity. Here we propose three indices to quantify each facet of functional diversity for a community with species distributed in a multidimensional functional space: functional richness (volume of the functional space occupied by the community), functional evenness (regularity of the distribution of abundance in this volume), and functional divergence (divergence in the distribution of abundance in this volume). Functional richness is estimated using the existing convex hull volume index. The new functional evenness index is based on the minimum spanning tree which links all the species in the multidimensional functional space. Then this new index quantifies the regularity with which species abundances are distributed along the spanning tree. Functional divergence is measured using a novel index which quantifies how species diverge in their distances (weighted by their abundance) from the center of gravity in the functional space. We show that none of the indices meets all the criteria required for a functional diversity index, but instead we show that the set of three complementary indices meets these criteria. Through simulations of artificial data sets, we demonstrate that functional divergence and functional evenness are independent of species richness and that the three functional diversity indices are independent of each other. Overall, our study suggests that decomposition of functional diversity into its three primary components provides a meaningful framework for its quantification and for the classification of existing functional diversity indices. This decomposition has the potential to shed light on the role of biodiversity on ecosystem functioning and on the influence of biotic and abiotic filters on the structure of species communities. Finally, we propose a general framework for applying these three functional diversity indices.     

Dominant species identity, not community evenness, regulates invasion in experimental grassland plant communities

While there has been extensive interest in understanding the relationship between diversity and invasibility of communities, most studies have only focused on one component of diversity: species richness. Although the number of species can affect community invasibility, other aspects of diversity, including species identity and community evenness, may be equally important. While several field studies have examined how invasibility varies with diversity by manipulating species identity or evenness, the results are often confounded by resource heterogeneity, site history, or disturbance. We designed a mesocosm experiment to examine explicitly the role of dominant species identity and evenness on the invasibility of grassland plant communities. We found that the identity of the dominant plant species, but not community evenness, significantly impacted invasibility. Using path analysis, we found that community composition (dominant species identity) reduced invasion by reducing early-season light availability and increasing late-season plant community biomass. Nitrogen availability was an important factor for the survival of invaders in the second year of the experiment. We also found significant direct effects of certain dominant species on invasion, although the mechanisms driving these effects remain unclear. The magnitude of dominant species effects on invasibility we observed are comparable to species richness effects observed in other studies, showing that species composition and dominant species can have strong effects on the invasibility of a community.     

旅游干扰下五台山不同植被景观区物种多样性特征

利用双向指示种分析方法(Two-way indicator species analysis,TWINSPAN)和6个物种多样性指数,研究了旅游干扰下五台山不同植被景观区物种多样性的特征.结果表明:1)TWINSPAN将所有样地划分为9类植被景观区,从Ⅰ区到Ⅸ区,随着旅游干扰程度的增加,植被景观大致由乔灌草区向灌草区、草本区和居民区方向变化.2)乔木层物种的丰富度和综合多样性随着旅游干扰的减小而趋于增加,但是其均匀度没有表现出明显的规律性.3)灌木层物种的丰富度和综合多样性也随着旅游干扰的减小而趋于增加,至于其均匀度,则呈现出在中度干扰下值最大,干扰很小的地方次之,在重度干扰下则最小.4)草本层物种的丰富度、均匀度和综合多样性指数均在旅游干扰适度的地方达到了最大值,在旅游干扰很小的地方,各种值则次之,在干扰严重的地方为最小.5)从整个植被层物种多样性的角度看,随着旅游干扰程度的减少,物种丰富度指数、均匀度指数和综合多样性指数均呈现趋于增加的趋势.表4参15     

Diversity and cover of a sessile animal assemblage does not predict its associated mobile fauna

Habitat-forming organisms often determine the structural properties and food resources available to a wide diversity of associated mobile species. Sessile invertebrate assemblages on marine hard substrates support an abundant fauna of mobile invertebrates whose associations with traits of their host assemblages are poorly known. To assess how changes to habitat-forming species are likely to affect their associated mobile fauna, the relationships between abundance, diversity and composition of mobile invertebrates and the diversity, cover and composition of the sessile assemblages they use as habitat were quantified in Sydney Harbour, Australia (33°50′S, 151°16′E). Similar compositions of sessile species were more likely to share a similar composition of mobile species, but univariate measures of the habitat (percent cover, species and functional diversity, prevalence of non-indigenous species) did not predict variation in associated mobile assemblages. These results demonstrate that in this habitat it is difficult to predict the diversity of marine assemblages based on common surrogate measures of biodiversity.     

Direct and indirect control of grassland community structure by litter, resources, and biomass

Multiple factors linked through complex networks of interaction including fertilization, aboveground biomass, and litter control the diversity of plant communities. The challenge of explaining plant diversity is to determine not only how each individual mechanism directly influences diversity, but how those mechanisms indirectly influence diversity through interactions with other mechanisms. This approach is well established in the study of plant species richness, but surprisingly little effort has been dedicated toward understanding the controls of community evenness, despite the recognition that this aspect of diversity can influence a variety of critical ecosystem functions. Similarly, studies of diversity have predominantly focused on the influence of shoot, rather than root, biomass, despite the fact that the majority of plant biomass is belowground in many natural communities. In this study, I examine the roles of belowground biomass, live aboveground biomass, litter, and light availability in controlling the species richness and evenness of a rough fescue grassland community using structural equation modeling. Litter was the primary mechanism structuring grassland diversity, with both richness and evenness declining with increasing litter cover. There were few relationships between shoot biomass, shading, and diversity, and more importantly, no relationship between root biomass and diversity. The lack of relationship between root biomass and species richness and evenness suggests that, even though root competition in grasslands is intense, belowground interactions may not play an important role in structuring community diversity or composition.     

Relationships between species richness, evenness, and abundance in a southwestern savanna

Species richness and evenness are components of biological diversity that may or may not be correlated with one another and with patterns of species abundance. We compared these attributes among flowering plants, grasshoppers, butterflies, lizards, summer birds, winter birds, and rodents across 48 plots in the grasslands and mesquite-oak savannas of southeastern Arizona. Species richness and evenness were uncorrelated or weakly negatively correlated for each taxonomic group, supporting the conclusion that richness alone is an incomplete measure of diversity. In each case, richness was positively correlated with one or more measures of abundance. By contrast, evenness usually was negatively correlated with the abundance variables, reflecting the fact that plots with high evenness generally were those where all species present were about equally uncommon. Therefore richness, but not evenness, usually was a positive predictor of places of conservation value, if these are defined as places where species of interest are especially abundant. Species diversity was more positively correlated with evenness than with richness among grasshoppers and flowering plants, in contrast to the other taxonomic groups, and the positive correlations between richness and abundance were comparatively weak for grasshoppers and plants as well. Both of these differences can be attributed to the fact that assemblages of plants and grasshoppers were numerically dominated by small subsets of common species (grasses and certain spur-throated grasshoppers) whose abundances differed greatly among plots in ways unrelated to species richness of the groups as a whole.     

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.     

Predators influence the tidal distribution of oysters (Crassostrea virginica)

Biotic and abiotic conditions can separately and synergistically influence the abundance and distribution of species and create vertical zonation patterns in marine systems. In Corpus Christi Bay, TX, USA, eastern oysters (Crassostrea virginica) are limited to intertidal habitats, while in adjacent estuaries, oysters not only grow subtidally, but thrive in these areas to the extent they are a viable commercial fishery. The purpose of this study was to assess how predators and abiotic conditions affect oyster mortality and growth at different tidal elevations. Anecdotal evidence suggests that abiotic conditions, primarily hypoxia and salinity, as well as oyster disease, limits oysters to intertidal areas. Yet, in Corpus Christi Bay, oysters are absent from subtidal areas where hypoxia is not known to occur. Infection by Perkinsus marinus (Dermo) is common in the study area, but previous work suggests that infection rates do not increase when oysters are transplanted subtidally. We investigated oyster tidal distributions by transplanting newly settled oysters into intertidal and subtidal areas. Predation on oysters was significantly greater in subtidal as compared to intertidal habitats. When protected from predators using cages, oyster survival significantly increased. Further, oysters in subtidal areas allocated significantly more resources to shell growth than did those in intertidal areas, and oysters are known to grow heavier shells in response to predators. Oyster settlement was not statistically different between inter and subtidal areas, and abiotic conditions measured during the study did not exceed known tolerance limits for oysters. Previous studies have shown that abiotic conditions influence oyster mortality and the success of restored oyster reefs. Our findings indicate that predators can also affect oyster distribution, and their effects should be evaluated when developing plans for oyster management and restoration.     

Parasitism as a determinant of community structure on intertidal flats

The burrowing and movement ability of the New Zealand cockle Austrovenus stutchburyi is reduced when infected by echinostome trematodes. Previous experimental evidence from a single site suggests that this parasite-induced behavioural change of a key bivalve can affect the structure of the surrounding benthic community. By using multiple regression analyses on data collected from 17 intertidal flats, we here show that cockle parasitism is associated with macrozoobenthic community structure on a larger spatial scale. Regressions were performed for animal abundance, biomass, species diversity and species richness separately, entering cockle parasitism (infection intensity), presence/absence of ghost shrimps (Callianassa filholi), cockle density, primary producer abundance and organic content, particle size, sorting coefficient and gravel content of the substrate as predictors. Next to ghost shrimps, cockle parasitism was the best predictor of animal abundance by affecting (mainly positively) 8 of the 49 most widespread species significantly. Cockle parasitism was also associated with the biomass of anthozoans (positively), nemerteans (negatively) and bivalves (positively), whereas overall animal biomass was positively related to the sorting coefficient of the substrate. Species diversity was positively associated with cockle parasitism and gravel content of the substrate. Species richness was significantly associated with cockle parasitism (positively), ghost shrimps (negatively) and abundance of primary producers (positively) in combination. The impact of cockle parasitism on benthic community structure is believed governed directly or indirectly by (1) reduced sediment disturbance, (2) increased surface structural complexity and (3) availability of larval trematodes as an additional food source.     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

人为干扰强度对村级景观结构特征的影响

以研究点与村落中心间的距离作为人为干扰强度的划分指标 ,把村落按重度人为干扰、中度人为干扰和轻度人为干扰 3个等级分为中心区、过渡区和基质区 ,分析不同人为干扰强度与村级景观特征的相互关系。结果表明 :随着人为干扰强度的增强 ,多样性指数、相对丰富度指数和均匀度指数均相应增大 ,在中度人为干扰强度区 ,上述 3种指数均最大 ,但优势度指数最小。随着干扰强度的增强 ,与人们生活密切相关的景观类型的分形维数减小 ,斑块的相似性增强     

Seed arrival and ecological filters interact to assemble high-diversity plant communities

Two prominent mechanisms proposed to structure biodiversity are niche-based ecological filtering and chance arrival of propagules from the species pool. Seed arrival is hypothesized to play a particularly strong role in high-diversity plant communities with large potential species pools and many rare species, but few studies have explored how seed arrival and local ecological filters interactively assemble species-rich communities in space and time. We experimentally manipulated seed arrival and multiple ecological filters in high-diversity, herbaceous-dominated groundcover communities in longleaf pine savannas, which contain the highest small-scale species richness in North America (up to > 40 species/m2). We tested three hypotheses: (1) local communities constitute relatively open-membership assemblages, in which increased seed arrival from the species pool strongly increases species richness; (2) ecological filters imposed by local fire intensity and soil moisture influence recruitment and richness of immigrating species; and (3) ecological filters increase similarity in the composition of immigrating species. In a two-year factorial field experiment, we manipulated local fire intensity by increasing pre-fire fuel loads, soil moisture using rain shelters and irrigation, and seed arrival by adding seeds from the local species pool. Seed arrival increased species richness regardless of fire intensity and soil moisture but interacted with both ecological filters to influence community assembly. High-intensity fire decreased richness of resident species, suggesting an important abiotic filter. In contrast, high-intensity fire increased recruitment and richness of immigrating species, presumably by decreasing effects of other ecological filters (competition and resource limitation) in postfire environments. Drought decreased recruitment and richness of immigrating species, whereas wet soil conditions increased recruitment but decreased or had little effect on richness. Moreover, some ecological filters (wet soil conditions and, to a lesser extent, high-intensity fire) increased similarity in the composition of immigrating species, illustrating conditions that influence deterministic community assembly in species-rich communities. Our experiment provides insights into how dispersal-assembly mechanisms may interact with niche-assembly mechanisms in space (spatial variation in disturbance) and time (temporal variation in resource availability) to structure high-diversity communities and can help guide conservation of threatened longleaf pine ecosystems in the face of habitat fragmentation and environmental change.     

Limited change in the diversity and structure of subtidal communities over four decades

A unique archive of photographs from 1969 to 1974 permitted a test of the hypothesis that the diversity and composition of contemporary epilithic communities on subtidal rock walls in the San Juan Islands, WA, USA, has changed over 40 years. Notably, the richness and diversity of sessile taxa was significantly higher in 2008–2011. Furthermore, the multivariate community structure of sessile and mobile taxa differed between the historic and modern eras. Historic communities were characterized by a high percent cover of bare rock and non-calcified algal crusts, consistent with the effects of grazing by chitons and urchins. The rate of sessile community turnover, an index less susceptible to spatial sampling artifacts than richness or diversity, was not significantly different between the two eras. Together with the naturally high spatial variability in these epilithic communities, and the limited replication of historic quadrats, we interpret cautiously the data as evidence of limited change despite a clear shift in temperature and predator (fish) guild composition. The lack of substantial change in rock wall communities may be due in part to their vertical topography, which limits physical disturbance and the preemption of space by weedy algae, two processes that are often associated with “phase-shifts” in other marine ecosystems.     

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.     

Spatial statistics for modeling of abundance and distribution of wildlife species in the Masai Mara ecosystem, Kenya

This study illustrates the use of modern statistical procedures for better wildlife management by addressing three key issues: determination of abundance, modeling of animal distributions and variability of diversity in space and time. Prior information in Markov Chain Monte Carlo (MCMC) methods is used to improve estimates of abundance. Measures of autocorrelation are included when modeling distributions of animal counts, and a diversity index to indicate species abundance and richness for large herbivores is developed. Data from the Masai Mara ecosystem in Kenya are used to develop and demonstrate these procedures. The new abundance estimates are up to 35% more accurate than those obtained by existing methods. Significant temporal changes in spatial patterns are found from a space-time analysis of elephant counts over a 20-year period, with strong interactions over 5 km and 6 months space and time separations, respectively. The new diversity index is sensitive to both high abundance and species richness and is also able to capture year to year variation. It indicates an overall marginal decrease in diversity for large herbivores in the Mara ecosystem. The space-time analyses and diversity index can easily be computed thereby providing tools for rapid decision making.     

The importance of agricultural lands for Himalayan birds in winter

The impacts of land‐use change on biodiversity in the Himalayas are poorly known, notwithstanding widespread deforestation and agricultural intensification in this highly biodiverse region. Although intact primary forests harbor many Himalayan birds during breeding, a large number of bird species use agricultural lands during winter. We assessed how Himalayan bird species richness, abundance, and composition during winter are affected by forest loss stemming from agriculture and grazing. Bird surveys along 12 elevational transects within primary forest, low‐intensity agriculture, mixed subsistence agriculture, and intensively grazed pastures in winter revealed that bird species richness and abundance were greatest in low‐intensity and mixed agriculture, intermediate in grazed pastures, and lowest in primary forest at both local and landscape scales; over twice as many species and individuals were recorded in low‐intensity agriculture than in primary forest. Bird communities in primary forests were distinct from those in all other land‐use classes, but only 4 species were unique to primary forests. Low‐, medium‐, and high‐intensity agriculture harbored 32 unique species. Of the species observed in primary forest, 80% had equal or greater abundance in low‐intensity agricultural lands, underscoring the value of these lands in retaining diverse community assemblages at high densities in winter. Among disturbed landscapes, bird species richness and abundance declined as land‐use intensity increased, especially in high‐intensity pastures. Our results suggest that agricultural landscapes are important for most Himalayan bird species in winter. But agricultural intensification—especially increased grazing—will likely result in biodiversity losses. Given that forest reserves alone may inadequately conserve Himalayan birds in winter, comprehensive conservation strategies in the region must go beyond protecting intact primary forests and ensure that low‐intensity agricultural lands are not extensively converted to high‐intensity pastures.     

Land‐Cover Change and Avian Diversity in the Conterminous United States

Abstract: Changes in land use and land cover have affected and will continue to affect biological diversity worldwide. Yet, understanding the spatially extensive effects of land‐cover change has been challenging because data that are consistent over space and time are lacking. We used the U.S. National Land Cover Dataset Land Cover Change Retrofit Product and North American Breeding Bird Survey data to examine land‐cover change and its associations with diversity of birds with principally terrestrial life cycles (landbirds) in the conterminous United States. We used mixed‐effects models and model selection to rank associations by ecoregion. Land cover in 3.22% of the area considered in our analyses changed from 1992 to 2001, and changes in species richness and abundance of birds were strongly associated with land‐cover changes. Changes in species richness and abundance were primarily associated with changes in nondominant types of land cover, yet in many ecoregions different types of land cover were associated with species richness than were associated with abundance. Conversion of natural land cover to anthropogenic land cover was more strongly associated with changes in bird species richness and abundance than persistence of natural land cover in nearly all ecoregions and different covariates were most strongly associated with species richness than with abundance in 11 of 17 ecoregions. Loss of grassland and shrubland affected bird species richness and abundance in forested ecoregions. Loss of wetland was associated with bird abundance in forested ecoregions. Our findings highlight the value of understanding changes in nondominant land cover types and their association with bird diversity in the United States.     

Biogenic disturbance determines invasion success in a subtidal soft-sediment system

Theoretically, disturbance and diversity can influence the success of invasive colonists if (1) resource limitation is a prime determinant of invasion success and (2) disturbance and diversity affect the availability of required resources. However, resource limitation is not of overriding importance in all systems, as exemplified by marine soft sediments, one of Earth's most widespread habitat types. Here, we tested the disturbance-invasion hypothesis in a marine soft-sediment system by altering rates of biogenic disturbance and tracking the natural colonization of plots by invasive species. Levels of sediment disturbance were controlled by manipulating densities of burrowing spatangoid urchins, the dominant biogenic sediment mixers in the system. Colonization success by two invasive species (a gobiid fish and a semelid bivalve) was greatest in plots with sediment disturbance rates < 500 cm(3) x m(-2) x d(-1), at the low end of the experimental disturbance gradient (0 to > 9000 cm(3) x m(-2) x d(-1)). Invasive colonization declined with increasing levels of sediment disturbance, counter to the disturbance-invasion hypothesis. Increased sediment disturbance by the urchins also reduced the richness and diversity of native macrofauna (particularly small, sedentary, surface feeders), though there was no evidence of increased availability of resources with increased disturbance that would have facilitated invasive colonization: sediment food resources (chlorophyll a and organic matter content) did not increase, and space and access to overlying water were not limited (low invertebrate abundance). Thus, our study revealed the importance of biogenic disturbance in promoting invasion resistance in a marine soft-sediment community, providing further evidence of the valuable role of bioturbation in soft-sediment systems (bioturbation also affects carbon processing, nutrient recycling, oxygen dynamics, benthic community structure, and so on.). Bioturbation rates are influenced by the presence and abundance of large burrowing species (like spatangoid urchins). Therefore, mass mortalities of large bioturbators could inflate invasion risk and alter other aspects of ecosystem performance in marine soft-sediment habitats.     

Increased primary production shifts the structure and composition of a terrestrial arthropod community

Numerous studies have examined relationships between primary production and biodiversity at higher trophic levels. However, altered production in plant communities is often tightly linked with concomitant shifts in diversity and composition, and most studies have not disentangled the direct effects of production on consumers. Furthermore, when studies do examine the effects of plant production on animals in terrestrial systems, they are primarily confined to a subset of taxonomic or functional groups instead of investigating the responses of the entire community. Using natural monocultures of the salt marsh cordgrass Spartina alterniflora, we were able to examine the impacts of increased plant production, independent of changes in plant composition and/or diversity, on the trophic structure, composition, and diversity of the entire arthropod community. If arthropod species richness increased with greater plant production, we predicted that it would be driven by: (1) an increase in the number of rare species, and/or (2) an increase in arthropod abundance. Our results largely supported our predictions: species richness of herbivores, detritivores, predators, and parasitoids increased monotonically with increasing levels of plant production, and the diversity of rare species also increased with plant production. However, rare species that accounted for this difference were predators, parasitoids, and detritivores, not herbivores. Herbivore species richness could be simply explained by the relationship between abundance and diversity. Using nonmetric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM), we also found significant changes in arthropod species composition with increasing levels of production. Our findings have important implications in the intertidal salt marsh, where human activities have increased nitrogen runoff into the marsh, and demonstrate that such nitrogen inputs cascade to affect community structure, diversity, and abundance in higher trophic levels.