Host-plant genotypic diversity mediates the distribution of an ecosystem engineer

Ecosystem engineers affect ecological communities by physically modifying the environment. Understanding the factors determining the distribution of engineers offers a powerful predictive tool for community ecology. In this study, we examine whether the goldenrod bunch gall midge (Rhopalomyia solidaginis) functions as an ecosystem engineer in an old-field ecosystem by altering the composition of arthropod species associated with a dominant host plant, Solidago altissima. We also examine the suite of factors that could affect the distribution and abundance of this ecosystem engineer. The presence of bunch galls increased species richness and altered the structure of associated arthropod communities. The best predictors of gall abundance were host-plant genotype and plot-level genotypic diversity. We found positive, nonadditive effects of genotypic diversity on gall abundance. Our results indicate that incorporating a genetic component in studies of ecosystem engineers can help predict their distribution and abundance, and ultimately their effects on biodiversity.     

Organization of Plethodon salamander communities: guild-based community assembly

A long-standing goal in evolutionary ecology is to determine whether the organization of communities is reflective of underlying deterministic processes. In this study, I examined patterns of species co-occurrence among eastern Plethodon salamanders and determined whether they were consistent with predictions from a guild model of competition-based community assembly. Using a database of 45 species and 4540 geographic sites, I found that patterns of co-occurrence were significantly nonrandom at both a regional and continental scale, and species of different size guilds were distributed more evenly in sites than was expected by chance. Sites with the highest species richness had consistent patterns of community composition, and with few exceptions, the same five species were present at all sites. Taken together, these results imply that larger Plethodon communities are assembled from simpler communities in a manner consistent with what is predicted through competitive mechanisms and suggest that stable species combinations are possible to achieve at various levels of species richness. These results also provide strong evidence consistent with the hypothesis that competitive-based community assembly is a general phenomenon in Plethodon and that interspecific competition is prevalent among the eastern species of this group.     

Zooplankton biodiversity and lake trophic state: explanations invoking resource abundance and distribution

While empirical studies linking biodiversity to local environmental gradients have emphasized the importance of lake trophic status (related to primary productivity), theoretical studies have implicated resource spatial heterogeneity and resource relative ratios as mechanisms behind these biodiversity patterns. To test the feasibility of these mechanisms in natural aquatic systems, the biodiversity of crustacean zooplankton communities along gradients of total phosphorus (TP) as well as the vertical heterogeneity and relative abundance of their phytoplankton resources were assessed in 18 lakes in Quebec, Canada. Zooplankton community richness was regressed against TP, the spatial distribution of phytoplankton spectral groups, and the relative biomass of spectral groups. Since species richness does not adequately capture ecological function and life history of different taxa, features which are important for mechanistic theories, relationships between zooplankton functional diversity (FD) and resource conditions were examined. Zooplankton species richness showed the previously established tendency to a unimodal relationship with TP, but functional diversity declined linearly over the same gradient. Changes in zooplankton functional diversity could be attributed to changes in both the spatial distribution and type of phytoplankton resource. In the studied lakes, spatial heterogeneity of phytoplankton groups declined with TP, even while biomass of all groups increased. Zooplankton functional diversity was positively related to increased heterogeneity in cyanobacteria spatial distribution. However, a smaller amount of variation in functional diversity was also positively related to the ratio of biomass in diatoms/chrysophytes to cyanobacteria. In all observed relationships, a greater variation of functional diversity than species richness measures was explained by measured factors, suggesting that functional measures of zooplankton communities will benefit ecological research attempting to identify mechanisms behind environmental gradients affecting diversity.     

A nearly neutral model of biodiversity

S. P. Hubbell's unified neutral theory of biodiversity has stimulated much new thinking about biodiversity. However, empirical support for the neutral theory is limited, and several observations are inconsistent with the predictions of the theory, including positive correlations between traits associated with competitive ability and species abundance and correlations between species diversity and ecosystem functioning. The neutral theory can be extended to explain these observations by allowing species to differ slightly in their competitive ability (fitness). Here, we show that even slight differences in fecundity can greatly reduce the time to extinction of competitors even when the community size is large and dispersal is spatially limited. In this case, species richness is dramatically reduced, and a markedly different species abundance distribution is predicted than under pure neutrality. In the nearly neutral model, species co-occur in the same community not because of, but in spite of, ecological differences. The more competitive species with higher fecundity tend to have higher abundance both in the metacommunity and in local communities. The nearly neutral perspective provides a theoretical framework that unites the sampling model of the neutral theory with theory of biodiversity affecting ecosystem function.     

Leaf quality, predators, and stochastic processes in the assembly of a diverse herbivore community

Ecological communities are structured by both deterministic, niche-based processes and stochastic processes such as dispersal. A pressing issue in ecology is to determine when and for which organisms each of these types of processes is important in community assembly. The roles of deterministic and stochastic processes have been studied for a variety of communities, but very few researchers have addressed their contribution to insect herbivore community structure. Insect herbivore niches are often described as largely shaped by the antagonistic pressures of predation and host plant defenses. However host plants are frequently discrete patches of habitat, and their spatial arrangement can affect herbivore dispersal patterns. We studied the roles of predation, host plant quality, and host spatial proximity for the assembly of a diverse insect herbivore community on Quercus alba (white oak) across two growing seasons. We examined abundances of feeding guilds to determine if ecologically similar species responded similarly to variation in niches. Most guilds responded similarly to leaf quality, preferring high-nitrogen, low-tannin host plants, particularly late in the growing season, while bird predation had little impact on herbivore abundance. The communities on the high-quality plants tended to be larger and, in some cases, have greater species richness. We analyzed community composition by correlating indices of community similarity with predator presence, leaf quality similarity, and host plant proximity. Birds did not affect community composition. Community similarity was significantly associated with distance between host plants and uncorrelated with leaf quality similarity. Thus although leaf quality significantly affected the total abundance of herbivores on a host plant, in some cases leading to increased species richness, dispersal limitation may weaken this relationship. The species composition of these communities may be driven by stochastic processes rather than variation in host plant characteristics or differential predation by insectivorous birds.     

A Two-Part Measure of Degree of Invasion for Cross-Community Comparisons

Abstract:  Invasibility is a critical feature of ecological communities, especially for management decisions. To date, invasibility has been measured in numerous ways. Although most researchers have used the richness (or number) of exotic species as a direct or indirect measure of community invasibility, others have used alternative measures such as the survival, density, or biomass of either a single or all exotic species. These different measures, even when obtained from the same communities, have produced inconsistent results and have made comparisons among communities difficult. Here, we propose a measure of the degree of invasion (DI) of a community as a surrogate for community invasibility. The measure is expressed as 2 independent components: exotic proportion of total species richness and exotic proportion of total species abundance (biomass or cover). By including richness and abundance, the measure reflects that the factors that control invasibility affect both of these components. Expressing exotic richness and abundance relative to the richness and abundance of all species in a community makes comparisons across communities of different sizes and resource availability possible and illustrates the importance of dominance of exotic species relative to natives, which is a primary management concern associated with exotic species.     

Building phenological models from presence/absence data for a butterfly fauna.

Species phenology is increasingly being used to explore the effects of climate change and other environmental stressors. Long-term monitoring data sets are essential for understanding both patterns manifest by individual species and more complex patterns evident at the community level. This study used records of 78 butterfly species observed on 626 days across 27 years at a site in northern California, USA, to build quadratic logistic regression models of the observation probability of each species for each day of the year. Daily species probabilities were summed to develop a potential aggregate species richness (PASR) model, indicating expected daily species richness. Daily positive and negative contributions to PASR were calculated, which can be used to target optimum sampling time frames. Residuals to PASR indicate a rate of decline of 0.12 species per year over the course of the study. When PASR was calculated for wet and dry years, wet years were found to delay group phenology by up to 17 days and reduce the maximum annual expected species from 32.36 to 30. Three tests to determine how well the PASR model reflected the butterfly fauna dynamics were all positive: We correlated probabilities developed with species presence/absence data to observed abundance by species, tested species' predicted phenological patterns against known biological characteristics, and compared the PASR curve to a spline-fitted curve calculated from the original species richness observations. Modeling individual species' flight windows was possible from presence/absence data, an approach that could be used on other similar records for butterfly communities with seasonal phenologies, and for common species with far fewer dates than used here. It also provided a method to assess sample frequency guidelines for other butterfly monitoring programs.     

Seaweed richness and herbivory increase rate of community recovery from disturbance

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.     

The effects of direct and indirect constraints on biological communities

Human activities are expected to result in a diversity of directional or stochastic constraints that affect species either directly or by indirectly impacting their resources. However, there is no theoretical framework to predict the complex and various effects of these constraints on ecological communities. We developed a dynamic model that mimics the use of different resource types by a community of competing species. We investigated the effects of different environmental constraints (affecting either directly the growth rate of species or having indirect effects on their resources) on several biodiversity indicators. Our results indicate that (i) in realistic community models (assuming uneven resource requirements among species) the effects of perturbations are strongly buffered compared to neutral models; (ii) the species richness of communities can be maximized for intermediate levels of direct constraints (unimodal response), even in the absence of trade-off between competitive ability and tolerance to constraints; (iii) no such unimodal response occurs with indirect constraints; (iv) an increase in the environmental (e.g., climatic) variance may have different effects on community biomass and species richness.     

Predation,recruitment and the dynamics of communities of coral-reef fishes

I investigated the ability of predators to influence the patterns of species richness and abundance of non-piscivorous fishes on small, artificial reefs replenished by natural recruitment. Periodic removal of predators effectively reduced the species richness and abundance of predators on removal reefs. The difference between the number of predators on control and removal reefs was greatest immediately following the removal of predators and attenuated between removals. During periods of recruitment, species richness and total abundance of recently-recruited, non-piscivorous fishes were generally greater on predator-removal reefs than on control reefs. Species richness and total abundance of resident non-piscivorous fishes were not affected by the removal of predators in the first year of the experiment. Both abundance and species richness of residents, however, were greater on the removal reefs during the second year of the experiment. The difference in the responses of the two age classes to the removal of predators suggests that predators may affect community patterns of older age classes through time-lagged effects on the survivorship of younger age classes. At the end of the experiment, species richness was positively related to abundance for recruits and residents. The effects of removing piscivorous fishes on the abundance of non-piscivorous fishes were similar for species considered separately. A greater number of species of recruit and resident fishes were more abundant on reefs from which predators had been removed. These data suggest that predators can play an important role in structuring communities of fishes on coral reefs.     

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.     

Small-scale grassland assembly patterns differ above and below the soil surface

The existence of deterministic assembly rules for plant communities remains an important and unresolved topic in ecology. Most studies examining community assembly have sampled aboveground species diversity and composition. However, plants also coexist belowground, and many coexistence theories invoke belowground competition as an explanation for aboveground patterns. We used next-generation sequencing that enables the identification of roots and rhizomes from mixed-species samples to measure coexisting species at small scales in temperate grasslands. We used comparable data from above (conventional methods) and below (molecular techniques) the soil surface (0.1 x 0.1 x 0.1 m volume). To detect evidence for nonrandom patterns in the direction of biotic or abiotic assembly processes, we used three assembly rules tests (richness variance, guild proportionality, and species co-occurrence indices) as well as pairwise association tests. We found support for biotic assembly rules aboveground, with lower variance in species richness than expected and more negative species associations. Belowground plant communities were structured more by abiotic processes, with greater variability in richness and guild proportionality than expected. Belowground assembly is largely driven by abiotic processes, with little evidence for competition-driven assembly, and this has implications for plant coexistence theories that are based on competition for soil resources.     

The comparative evidence relating to functional and neutral interpretations of biological communities

Neutral and functional theories provide rival interpretations of community patterns involving distribution, abundance, and diversity. One group of patterns describes the overall properties of species or sites, and derives principally from the frequency distribution of abundance among species. According to neutral theory, these patterns are determined by the number of individuals of novel type appearing each generation in the community, whereas functional theory relates them to the distribution of the extent of niches. A second group of patterns describes the spatial attributes of communities, and derives principally from the decay of similarity in species composition with distance. Neutral theory interprets these patterns as consequences of local dispersal alone, whereas the functional interpretation is that more distant sites are likely to be ecologically different. Neutral theory often provides good predictions of community patterns, yet is at variance with a wide range of experimental results involving the manipulation of environments or communities. One explanation for this discrepancy is that spatially explicit models where selection is generally weak, or where selection acts strongly on only a few species at each site, have essentially the same output as neutral models with respect to the distribution of abundance and the decay of similarity. Detecting a non-neutral signal in survey data requires careful spatial or phylogenetic analysis; we emphasize the potential utility of incorporating phylogenetic information in order to detect functional processes that lead to ecological variation among clades.     

Relative Effects of Disturbance on Red Imported Fire Ants and Native Ant Species in a Longleaf Pine Ecosystem

Abstract: The degree to which changes in community composition mediate the probability of colonization and spread of non‐native species is not well understood, especially in animal communities. High species richness may hinder the establishment of non‐native species. Distinguishing between this scenario and cases in which non‐native species become established in intact (lacking extensive anthropogenic soil disturbance) communities and subsequently diminish the abundance and richness of native species is challenging on the basis of observation alone. The red imported fire ant (Solenopsis invicta), an invasive species that occurs throughout much of the southeastern United States, is such an example. Rather than competitively displacing native species, fire ants may become established only in disturbed areas in which native species richness and abundance are already reduced. We used insecticide to reduce the abundance of native ants and fire ants in four experimental plots. We then observed the reassembly and reestablishment of the ants in these plots for 1 year after treatment. The abundance of fire ants in treated plots did not differ from abundance in control plots 1 year after treatment. Likewise, the abundance of native ants increased to levels comparable to those in control plots after 1 year. Our findings suggest that factors other than large reductions in ant abundance and species density (number of species per unit area) may affect the establishment of fire ants and that the response of native ants and fire ants to disturbance can be comparable.     

Modelling plant species richness using functional groups

Conservation biologists increasingly rely on spatial predictive models of biodiversity to support decision-making. Therefore, highly accurate and ecologically meaningful models are required at relatively broad spatial scales. While statistical techniques have been optimized to improve model accuracy, less focus has been given to the question: How does the autecology of a single species affect model quality? We compare a direct modelling approach versus a cumulative modelling approach for predicting plant species richness, where the latter gives more weight to the ecology of functional species groups. In the direct modelling approach, species richness is predicted by a single model calibrated for all species. In the cumulative modelling approach, the species were partitioned into functional groups, with each group calibrated separately and species richness of each group was cumulated to predict total species richness. We hypothesized that model accuracy depends on the ecology of individual species and that the cumulative modelling approach would predict species richness more accurately. The predictors explained plant species richness by ca. 25%. However, depending on the functional group the deviance explained varied from 3 to 67%. While both modelling approaches performed equally well, the models of the different functional groups highly varied in their quality and their spatial richness pattern. This variability helps to improve our understanding on how plant functional groups respond to ecological gradients.     

南亚热带森林3层次物种多度之广义泊松分布模型

生物群落中物种多度分布(species abundance distribution)呈典型的倒J形,即其中存在许多稀有种、少量常见种.物种多度分布模型研究有助于解决森林生态恢复中的物种配置等实际问题.本研究考察了一种过分散(over-dispersion,或称超分布,即方差大于均值)的离散型分布,即具有λ和α两个参数...     

Estimating species richness and accumulation by modeling species occurrence and detectability

A statistical model is developed for estimating species richness and accumulation by formulating these community-level attributes as functions of model-based estimators of species occurrence while accounting for imperfect detection of individual species. The model requires a sampling protocol wherein repeated observations are made at a collection of sample locations selected to be representative of the community. This temporal replication provides the data needed to resolve the ambiguity between species absence and nondetection when species are unobserved at sample locations. Estimates of species richness and accumulation are computed for two communities, an avian community and a butterfly community. Our model-based estimates suggest that detection failures in many bird species were attributed to low rates of occurrence, as opposed to simply low rates of detection. We estimate that the avian community contains a substantial number of uncommon species and that species richness greatly exceeds the number of species actually observed in the sample. In fact, predictions of species accumulation suggest that even doubling the number of sample locations would not have revealed all of the species in the community. In contrast, our analysis of the butterfly community suggests that many species are relatively common and that the estimated richness of species in the community is nearly equal to the number of species actually detected in the sample. Our predictions of species accumulation suggest that the number of sample locations actually used in the butterfly survey could have been cut in half and the asymptotic richness of species still would have been attained. Our approach of developing occurrence-based summaries of communities while allowing for imperfect detection of species is broadly applicable and should prove useful in the design and analysis of surveys of biodiversity.     

The role of time and species identities in spatial patterns of species richness and conservation

Many conservation actions are justified on the basis of managing biodiversity. Biodiversity, in terms of species richness, is largely the product of rare species. This is problematic because the intensity of sampling needed to characterize communities and patterns of rarity or to justify the use of surrogates has biased sampling in favor of space over time. However, environmental fluctuations interacting with community dynamics lead to temporal variations in where and when species occur, potentially affecting conservation planning by generating uncertainty about results of species distribution modeling (including range determinations), selection of surrogates for biodiversity, and the proportion of biodiversity composed of rare species. To have confidence in the evidence base for conservation actions, one must consider whether temporal replication is necessary to produce broad inferences. Using approximately 20 years of macrofaunal data from tidal flats in 2 harbors, we explored variation in the identity of rare, common, restricted range, and widespread species over time and space. Over time, rare taxa were more likely to increase in abundance or occurrence than to remain rare or disappear and to exhibit temporal patterns in their occurrence. Space–time congruency in ranges (i.e., spatially widespread taxa were also temporally widespread) was observed only where samples were collected across an environmental gradient. Fifteen percent of the taxa in both harbors changed over time from having spatially restricted ranges to having widespread ranges. Our findings suggest that rare species can provide stability against environmental change, because the majority of species were not random transients, but that selection of biodiversity surrogates requires temporal validation. Rarity needs to be considered both spatially and temporally, as species that occur randomly over time are likely to play a different role in ecosystem functioning than those exhibiting temporal structure (e.g., seasonality). Moreover, temporal structure offers the opportunity to place management and conservation activities within windows of maximum opportunity.     

Native fish diversity alters the effects of an invasive species on food webs

Aquatic communities have been altered by invasive species, with impacts on native biodiversity and ecosystem function. At the same time, native biodiversity may mitigate the effects of an invader. Common carp (Cyprinus carpio) is a ubiquitous, invasive fish species that strongly influences community and ecosystem processes. We used common carp to test whether the potential effects of an invasive species are altered across a range of species diversity in native communities. In mesocosms, treatments of zero, one, three, and six native fish species were used to represent the nested subset patterns observed in fish communities of lakes in Illinois, USA. The effect of the invader was tested across fish richness treatments by adding common carp to the native community and substituting native biomass with common carp. Native species and intraspecific effects reduced invader growth. The invader reduced native fish growth; however, the negative effect was minimized with increasing native richness. The zooplankton grazer community was modified by a top-down effect from the invader that increased the amount of phytoplankton. Neither the invader nor richness treatments influenced total phosphorus or community metabolism. Overall, the invader reduced resources for native species; and the effect scaled with how the invader was incorporated into the community. Higher native diversity mitigated the impact of the invader, confirming the need to consider biodiversity when predicting the impacts of invasive species.     

Linking richness, community variability, and invasion resistance with patch size

The influence of community dynamics on the success or failure of an invasion is of considerable interest. What has not been explored is the influence of patch size on the outcomes of invasions for communities with the same species pool. Here we use an empirically validated spatial model of a marine epibenthic community to examine the effects of patch size on community variability, species richness, invasion, and the relationships between these variables. We found that the qualitative form of the relationship between community variability and species richness is determined by the size of the model patch. In small patches, variability decreases with species richness, but beyond a critical patch size, variability increases with increasing richness. This occurs because in large patches large, long-lived colonies attain sufficient size to minimize mortality and dominate the community, leading to decreased species richness and community variability. This mechanism cannot operate on smaller patches where the size of colonies is limited by the patch size and mortality is high irrespective of species identity. Further, invasion resistance is strongly correlated with community variability. Thus, the relationship between species richness and invasion resistance is also determined by patch size. These patterns are generated largely by an inverse relationship between colony size and mortality, and they depend on the spatial nature and patch size of the community. Our results suggest that a continuum of possible relationships can exist between species richness, community variability, invasion resistance, and area. These relationships are emergent behaviors generated by the individual properties of the particular component species of a community.