Spatial heterogeneity influences native and nonnative plant species richness

Spatial heterogeneity may have differential effects on the distribution of native and nonnative plant species richness. We examined the effects of spatial heterogeneity on native and nonnative plant species richness distributions in the central part of Rocky Mountain National Park, Colorado, USA. Spatial heterogeneity around vegetation plots was characterized using landscape metrics, environmental/topographic variables (slope, aspect, elevation, and distance from stream or river), and soil variables (nitrogen, clay, and sand). The landscape metrics represented five components of landscape heterogeneity and were measured at four spatial extents (within varying radii of 120, 240, 480, and 960 m) using the FRAGSTATS landscape pattern analysis program. Akaike's Information Criterion adjusted for small sample size (AICc) was used to select the best models from a set of multiple linear regression models developed for native and nonnative plant species richness at four spatial extents and three levels of ecological hierarchy (i.e., landscape, land cover, and community). Both native and nonnative plant species richness were positively correlated with edge density, Simpson's diversity index and interspersion/juxtaposition index, and were negatively correlated with mean patch size. The amount of variation explained at four spatial extents and three hierarchical levels ranged from 30% to 70%. At the landscape level, the best models explained 43% of the variation in native plant species richness and 70% of the variation in nonnative plant species richness (240-m extent). In general, the amount of variation explained was always higher for nonnative plant species richness, and the inclusion of landscape metrics always significantly improved the models. The best models explained 66% of the variation in nonnative plant species richness for both the conifer land cover type and lodgepole pine community. The relative influence of the components of spatial heterogeneity differed for native and nonnative plant species richness and varied with the spatial extent of analysis and levels of ecological hierarchy. The study offers an approach to quantify spatial heterogeneity to improve models of plant biodiversity. The results demonstrate that ecologists must recognize the importance of spatial heterogeneity in managing native and nonnative plant species.     

Raising groundwater differentially affects mineralization and plant species abundance in dune slacks.

The experience with restoring high water levels (i.e., rewetting) within restoration ecology is limited, and information on changes in soil nutrient supply is scarce. A reduction in nutrient supply is needed to restore the desired oligotrophic vegetation. We determined the effects of restoration of high water levels on decomposition and net carbon (C), nitrogen (N), and phosphorus (P) mineralization rates in wet dune slacks and its consequences for the relative abundance of eutrophic vs. oligotrophic species in the vegetation. This was done by analyzing these variables for valleys that experienced a large groundwater rise vs. valleys that had a small groundwater rise but the same current water level. In addition, the influences of underlying factors (waterlogging, vegetation dieback, and soil dynamics prior to groundwater rise) were separated in a transplantation experiment. Short-term effects of large groundwater rise were a massive dieback of vegetation, increased thickness of the fermentation layer, increased microbial decomposition activity, increased C mineralization, and decreased net N mineralization. Net P mineralization was not affected. The relative abundance of oligotrophic vs. eutrophic species was greater at large groundwater rise. Changes in decomposition and mineralization by large groundwater rise were, however, not caused by the vegetation dieback, but due to previous soil conditions. Soils experiencing waterlogged conditions for 3-4 years or more prior to large groundwater rise had lower C and higher net N mineralization rates at waterlogged conditions than soils that had experienced aerobic conditions, presumably due to differences in labile soil C contents. Contrary to expectations induced by previously determined nutrient pulses and measured vegetation dieback, large groundwater rise resulted in lower soil nutrient supply rates and more oligotrophic vegetation. If these trends continue on the longer term, restoration of high water levels may be effective in restoration ecology to establish oligotrophic, wet vegetation in dune slacks.     

Grassland root communities: species distributions and how they are linked to aboveground abundance

There is little comprehensive information on the distribution of root systems among coexisting species, despite the expected importance of those distributions in determining the composition and diversity of plant communities. This gap in knowledge is particularly acute for grasslands, which possess large numbers of species with morphologically indistinguishable roots. In this study we adapted a molecular method, fluorescent fragment length polymorphism, to identify root fragments and determine species root distributions in two grasslands in Yellowstone National Park (YNP). Aboveground biomass was measured, and soil cores (2 cm in diameter) were collected to depths of 40 cm and 90 cm in an upland, dry grassland and a mesic, slope-bottom grassland, respectively, at peak foliar expansion. Cores were subdivided, and species that occurred in each 10-cm interval were identified. The results indicated that the average number of species in 10-cm intervals (31 cm3) throughout the sampled soil profile was 3.9 and 2.8 species at a dry grassland and a mesic grassland, respectively. By contrast, there was an average of 6.7 and 14.1 species per 0.5 m2, determined by the presence of shoot material, at dry and mesic sites, respectively. There was no relationship between soil depth and number of species per 10-cm interval in either grassland, despite the exponential decline of root biomass with soil depth at both sites. There also was no relationship between root frequency (i.e., the percentage of samples in which a species occurred) and soil depth for the vast majority of species at both sites. The preponderance of species were distributed throughout the soil profile at both sites. Assembly analyses indicated that species root occurrences were randomly assorted in all soil intervals at both sites, with the exception that Festuca idahoensis segregated from Artemisia tridentata and Pseudoroegnaria spicata in 10-20 cm soil at the dry grassland. Root frequency throughout the entire sampled soil profile was positively associated with shoot biomass among species. Together these results indicated the importance of large, well-proliferated root systems in establishing aboveground dominance. The findings suggest that spatial belowground segregation of species probably plays a minor role in fostering resource partitioning and species coexistence in these YNP grasslands.     

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.     

Pesticide treatments affect mountain pine beetle abundance and woodpecker foraging behavior.

In British Columbia, Canada, management efforts used to control mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks have included treatment of infested trees with an organic arsenic pesticide, monosodium methanearsonate (MSMA). Cumulative pesticide applications over a large geographic area have generated concerns about arsenic loading in the environment and potential toxicity to nontarget wildlife. We investigated woodpecker foraging patterns in infested stands with and without MSMA treatment using a combination of tree debarking indices, point count surveys, and radiotelemetry methods in addition to insect flight traps to measure mountain pine beetle emergence. Debarking indices indicated woodpecker foraging of MSMA-treated trees was significantly lower than nontreated trees in all sampling years. However, approximately 40% of MSMA trees had some evidence of foraging. Focal observations of foraging woodpeckers and point count surveys in MSMA treatment areas further confirmed that several species of woodpeckers regularly used MSMA stands during the breeding season. Radio-tagged Hairy (Picoides villosus) and Three-toed (Picoides dorsalis) Woodpeckers spent on average 13% and 23% (range 0-66%) of their time, respectively, in treated stands, despite the fact that these areas only comprised on average 1-2% of their core home range (1 km2). MSMA strongly reduced the emergence of several bark beetle (Coleoptera, Scolytidae) species including the mountain pine beetle, and there was a highly significant positive relationship between Dendroctonus beetle abundance and Three-toed Woodpecker abundance. This study identifies the potential negative impact that forest management practices using pesticides can have on woodpecker populations that depend on bark beetles and their host trees.     

Modelling the abundance of three key plant species in New Zealand hill-pasture using a decision tree approach

Decision tree models were developed to investigate and predict the relative abundance of three key pasture plants [ryegrass (Lolium perenne), browntop (Agrostis capillaris), and white clover (Trifolium repens)] with integration of a geographical information system (GIS) in a naturalised hill-pasture in the North Island, New Zealand, and were compared with regression models with respect to model fit and predictive accuracy. The results indicated that the decision tree models had a better model fit in terms of average squared error (ASE) and a higher percentage of adequately predicted cases in model validation than the corresponding regression models. These decision tree models clearly revealed the relative importance of environmental and management variables in influencing the abundance of these three species. Hill slope was the most significant environmental factor influencing the abundance of ryegrass while soil Olsen P and annual P fertilizer input were the most significant factors influencing the abundance of browntop, and white clover, respectively. Soil Olsen P of approximately 10 μg/g, or a slope of about 10.5° was critical points where the competition between ryegrass and browntop tended to come to an equilibrium. Integrating the decision tree models with a GIS in this study not only facilitated the model development and analyses, but also provided a useful decision support tool in pasture management such as in assisting precision fertilizer placement. The insights obtained from the decision tree models also have important implications for pasture management, for example, it is important to maintain a soil Olsen P higher than 10 μg/g in order to keep the dominance of ryegrass in the hill-pasture.     

Phylogeny, niches, and relative abundance in natural communities

Community structure refers to the number of species in a community and the pattern of distribution of individuals among those species. We use a novel way of representing community structure to show that abundance within closely related pairs of co-occurring tree species in a highly diverse Mexican forest is more equitable than is abundance within more distantly related pairs. This observation is at odds with the fundamental assumption of neutral models of community structure, i.e., that species are interchangeable. The observed patterns suggest niche apportionment, in which interaction is focused pairwise between congeners but falls away from the phylogenetic structure above the genus level. Thus niche processes may significantly affect community structure through regulating relative abundance in a substantial proportion of species, which in turn potentially enhances community stability. One such mechanism of stable coexistence has already been shown to be active in this forest.     

Energy use and animal abundance in litter and soil communities

Tools from metabolic scaling and food web theory were used to construct a general model of carbon flux by litter and soil invertebrates. The flux model was used to explore the energetic basis of invertebrate abundance and predicted that abundance should (1) scale linearly with net primary production; (2) be related to the body mass of animals as a power function, with an exponent between -0.65 and -0.85; (3) be related to the average body temperature of animals according to the Boltzmann factor, with an activation energy between 0.27 and 0.79 eV; and (4) decrease by a factor of 0.05 to 0.15 across trophic levels due to gross production efficiency of prey. Model predictions were generally supported by a global data set on invertebrate abundance that was amassed during the International Biological Programme, indicating that fundamental energetic principles explain a large degree of variation in invertebrate abundance across the globe.     

Environmental variability and allocation trade-offs maintain species diversity in a process-based model of succulent plant communities

Ecological theory suggests that environmental variability can promote coexistence, provided that species occupy differential niches. In this study, we focus on two questions: (1) Do allocation trade-offs provide a sufficient basis for niche differentiation in succulent plant communities? (2) What is the relative importance of different forms of environmental variability on species diversity and community composition? We approach these questions with a generic, individual-based simulation model. In our model, plants compete for water in a spatially explicit environment. Species differ in their size at maturity and in the allocation of carbon to roots, leaves and storage tissue. The model was fully specified with independent literature data. Model output was compared to characteristics of a species-rich community in the semi-arid Richtersveld (South Africa). The model reproduced the coexistence of plants with different sizes at maturity, the dominance of succulent shrubs, and the level of vegetation cover. We analyzed the effects of three forms of environmental variability: (a) temporal fluctuations in precipitation (rain and fog), (b) spatial heterogeneity of water supply due to run-on and run-off processes and (c) ‘rock pockets’ that limit root competition in space. The three types of variability had differential effects on diversity: diversity exhibited a strong hump-shaped response to temporal variation. Spatial variability increased diversity, with the strongest increase occurring at intermediate levels of temporal variability. Finally, rock pockets had the weakest effect, but contributed to diversity by providing refuges for small species, particularly at low temporal variability. The model thus shows that spatio-temporal variation of resource supply can maintain diversity over long time scales even in small systems, as is the case in the Richtersveld succulent communities. Trade-offs in allocation provide the basis for necessary niche differentiation. By describing resource competition between individual plants, our model provides a mechanistic basis for the link from species traits to community composition at given environmental conditions. It thereby contributes to an understanding of the forces shaping plant communities. Such an understanding is critical to reduce the threats environmental change poses to biodiversity and ecosystem services.     

Fire reduces morphospace occupation in plant communities

The two main assembly processes claimed to structure plant communities are habitat filtering and competitive interactions. The set of species growing in fire-prone communities has been filtered in such a way that species without fire-persistence traits have not successfully entered the community. Because plant traits are evolutionarily conserved and fire traits are correlated with other plant traits, communities under high fire frequency should not include all possible trait combinations, and thus the morphospace occupation by species in these communities should be lower than expected by chance (underoccupied). In contrast, communities under low fire frequency would lack the filtering factor, and thus their underoccupation of the morphospace is not expected. We test this prediction by comparing the morphospace occupation by species in communities located in the western Mediterranean Basin, five of them subject to high fire frequency (HiFi) and four to low fire frequency (LowFi). We first compile a set of morphological and functional traits for the species growing on the nine sites, then we compute the morphospace occupation of each site as a convex hull volume, and finally, to assert that our results are not a product of a random branching pattern of evolution, we simulate our traits under a null model of neutral evolution and compare the morphospace occupation of the simulated traits with the results from the empirical data. The results suggest that, as predicted, there is a clear differential morphospace occupation between communities under different fire regimes in such a way that the morphospace is underoccupied in HiFi communities only. The simulation of a neutral evolutionary model does not replicate the observed pattern of differential morphospace occupation, and thus it should be attributed to assembly processes. In conclusion, our results suggest that fire is a strong community assembling process, filtering the species that have fire-persistent traits and thus assembling phenotypically and phylogenetically clustered communities with vacant zones in the morphospace.     

Long-term dynamics of biotic and abiotic resistance to exotic species invasion in restored vernal pool plant communities

Invasion of native ecosystems by exotic species can seriously threaten native biodiversity, alter ecosystem function, and inhibit conservation. Moreover, restoration of native plant communities is often impeded by competition from exotic species. Exotic species invasion may be limited by unfavorable abiotic conditions and by competition with native species, but the relative importance of biotic and abiotic factors remains controversial and may vary during the invasion process. We used a long-term experiment involving restored vernal pool plant communities to characterize the temporal dynamics of exotic species invasion, and to evaluate the relative support for biotic and abiotic factors affecting invasion resistance. Experimental pools (n=256) were divided among controls and several seeding treatments. In most treatments, native vernal pool species were initially more abundant than exotic species, and pools that initially received more native seeds exhibited lower frequencies of exotic species over time. However, even densely seeded pools were eventually dominated by exotic species, following extreme climatic events that reduced both native and exotic plant densities across the study site. By the sixth year of the experiment, most pools supported more exotics than native vernal pool species, regardless of seeding treatment or pool depth. Although deeper pools were less invaded by exotic species, two exotics (Hordeum marinum and Lolium multiflorum) were able to colonize deeper pools as soon as the cover of native species was reduced by climatic extremes. Based on an information-theoretic analysis, the best model of invasion resistance included a nonlinear effect of seeding treatment and both linear and nonlinear effects of pool depth. Pool depth received more support as a predictor of invasion resistance, but seeding intensity was also strongly supported in multivariate models of invasion, and was the best predictor of resistance to invasion by H. marinum and L. multilorum. We conclude that extreme climatic events can facilitate exotic species invasions by both reducing abiotic constraints and weakening biotic resistance to invasion.     

Human impacts, plant invasion, and imperiled plant species in California.

Invasive species are one of the fastest growing conservation problems. These species homogenize the world's flora and fauna, threaten rare and endemic species, and impose large economic costs. Here, we examine the distribution of 834 of the more than 1000 exotic plant taxa that have become established in California, USA. Total species richness increases with net primary productivity; however, the exotic flora is richest in low-lying coastal sites that harbor large numbers of imperiled species, while native diversity is highest in areas with high mean elevation. Weedy and invasive exotics are more tightly linked to the distribution of imperiled species than the overall pool of exotic species. Structural equation modeling suggests that while human activities, such as urbanization and agriculture, facilitate the initial invasion by exotic plants, exotics spread ahead of the front of human development into areas with high numbers of threatened native plants. The range sizes of exotic taxa are an order of magnitude smaller than for comparable native taxa. The current small range size of exotic species implies that California has a significant "invasion debt" that will be paid as exotic plants expand their range and spread throughout the state.     

Interaction rules affect species coexistence in intransitive networks

Intransitive communities, those in which species' abilities cannot be ranked in a hierarchy, have been the focus of theoretical and empirical research, as intransitivity could help explain the maintenance of biodiversity. Here we show that models for intransitive competition embedding slightly different interaction rules can produce opposite patterns. In particular, we find that interactions in which an individual can be outcompeted by its neighbors, but cannot outcompete its neighbors, produce negative frequency dependence that, in turn, promotes coexistence. Whenever the interaction rule is modified toward symmetry (the individual and the neighbors can outcompete each other) the negative frequency dependence vanishes, producing different coexistence levels. Macroscopically, we find that asymmetric interactions yield highest biodiversity if species compete globally, while symmetric interactions favor highest biodiversity if competition takes place locally.     

Differences in gypsum plant communities associated with habitat fragmentation and livestock grazing.

The negative consequences of habitat fragmentation for plant communities have been documented in many regions of the world. In some fragmented habitats, livestock grazing has been proposed to be a dispersal mechanism reducing isolation between fragments. In others, grazing acts together with fragmentation in a way that increases habitat degradation. Iberian gypsum plant communities have been grazed and fragmented by agricultural practices for centuries. Although their conservation is considered a priority by the European Community, the effects of fragmentation on gypsum plant communities and the possible role of livestock grazing remain unknown. In addition, a substantial proportion of plant species growing in gypsum environments are gypsum specialists. They could be particularly affected by fragmentation, as was found for other habitat specialists (i.e., serpentine and calcareous specialists). In this study (1) we investigated the effect of fragmentation and grazing on gypsum plant community composition (species and life-forms), and (2) we tested to see if gypsum specialists were differently affected by fragmentation and grazing than habitat generalists. A vegetation survey was conducted in the largest gypsum outcrop of Europe (Middle Ebro Valley, northeast Spain). Fragmented and continuous sites in grazed and ungrazed areas were compared. Measurements related to species and composition of life-forms were contrasted first for the whole gypsum plant community and then specifically for the gypsum specialists. In the whole community, our results showed lower plant species diversity in fragmented sites, mainly due to the larger dominance of species more tolerant to fragmented habitat conditions. With livestock grazing, the plant species richness and the similarity in plant species composition between remnants was larger, suggesting that animals were acting as dispersal agents between fragments. As expected, gypsum specialists were less abundant in fragmented areas, and grazing led to the disappearance of the rare gypsum specialist Campanula fastigiata. According to our results, conservation strategies for gypsum plant communities in human-dominated landscapes should consider that fragmentation and grazing modify plant community composition affecting gypsum specialists in particular.     

Species and rotation frequency influence soil nitrogen in simplified tropical plant communities.

Among the many factors that potentially influence the rate at which nitrogen (N) becomes available to plants in terrestrial ecosystems are the identity and diversity of species composition, frequency of disturbance or stand turnover, and time. Replicated suites of investigator-designed communities afforded an opportunity to examine the effects of those factors on net N mineralization over a 12-year period. The communities consisted of large-stature perennial plants, comprising three tree species (Hyeronima alchorneoides, Cedrela odorata, and Cordia alliodora), a palm (Euterpe oleracea), and a large, perennial herb (Heliconia imbricata). Trees were grown in monoculture and in combination with the other two life-forms; tree monocultures were subjected to rotations of one or four years, or like the three-life-form systems, left uncut. The work was conducted on fertile soil in the humid lowlands of Costa Rica, a site with few abiotic constraints to plant growth. Rates of net N mineralization and nitrification were high, typically in the range of 0.2-0.8 microg x g(1) x d(-1), with net nitrification slightly higher than net mineralization, indicating preferential uptake of ammonium (NH4+) by plants and microbes. Net rates of N mineralization were about 30% lower in stands of one of the three tree species, Hyeronima, than in stands of the other two. Contrary to expectations, short-rotation management (one or four years) resulted in higher net rates of N mineralization than in uncut stands, whether the latter were composed of a single tree species or a combination of life-forms. Neither additional species richness nor replenishment of leached N augmented mineralization rates. The net rate at which N was supplied tended to be lowest in stands where demand for N was highest. Careful choice of species, coupled with low frequency of disturbance, can lead to maintenance of N within biomass and steady rates of within-system circulation, whereas pulses, whether caused by cutting and replanting or by the phenological traits of the species selected or combined, subject N supplies to leaching loss.     

A direct comparison of the consequences of plant genotypic and species diversity on communities and ecosystem function

Biodiversity loss is proceeding at an unprecedented rate, yet we lack a thorough understanding of the consequences of losing diversity at different scales. While species diversity is known to impact community and ecosystem processes, genotypic diversity is assumed to have relatively smaller effects. Nonetheless, a few recent studies suggest that genotypic diversity may have quantitatively similar ecological consequences compared to species diversity. Here we show that increasing either genotypic diversity of common evening primrose (Oenothera biennis) or species diversity of old-field plant species resulted in nearly equivalent increases (approximately 17%) in aboveground primary production. The predominant mechanism explaining this effect, niche complementarity, was similar for each type of diversity. Arthropod species richness also increased with both types of plant diversity, but the mechanisms leading to this effect differed: abundance-driven accumulation of arthropod species was important in plant genotypic polycultures, whereas resource specialization was important in plant species polycultures. Thus, similar increases in primary productivity differentially impacted higher trophic levels in response to each type of plant diversity. These results highlight important ecological similarities and differences between genotypic and species diversity and suggest that genotypic diversity may play a larger role in community and ecosystem processes than previously realized.     

昆嵛山盐肤木RhuschinensisMill．群落特征及其物种多样性

根据野外典型样方调查结果,以群落植物种类组成、群落外貌特征、群落垂直结构特征、群落多样性几项指标,分析山东半岛昆嵛山国家级自然保护区盐肤木Rhuschinensis群落特征。结果表明：盐肤木群落共出现种子植物66种,隶属于32科57属。在盐肤木群落属的区系组成中,群落植物主要由温带区系和热带区系成分组成,其中温带植物属占显著优势,其次为热带分布属,说明该群落具有明显的温带性质,并具有过渡特征。群落层次分明,有乔木层、灌木层、草本层3个层次,各层次的优势种依次为盐肤木和赤松Pinusdensiflora、麻栎Quercusacutissima、披针叶苔草Carexlanceolata。盐肤木群落的物种多样性由高到低依次为灌木层、乔木层、草本层,低均匀度导致物种丰富度最高的草本层具有最低的物种多样性。调查显示盐肤木幼苗幼树多,自我更新良好,群落处于稳定增长状态。通过对群落特征和物种多样性的研究,揭示了昆嵛山盐肤木群落的自然生态规律,为保护和开发利用盐肤木提供了科学依据。     

昆嵛山盐肤木Rhus chinensis Mill.群落特征及其物种多样性

根据野外典型样方调查结果,以群落植物种类组成、群落外貌特征、群落垂直结构特征、群落多样性几项指标,分析山东半岛昆嵛山国家级自然保护区盐肤木Rhus chinensis群落特征。结果表明:盐肤木群落共出现种子植物66种,隶属于32科57属。在盐肤木群落属的区系组成中,群落植物主要由温带区系和热带区系成分组成,其中温带植物属占显著优势,其次为热带分布属,说明该群落具有明显的温带性质,并具有过渡特征。群落层次分明,有乔木层、灌木层、草本层3个层次,各层次的优势种依次为盐肤木和赤松Pinus densiflora、麻栎Quercus acutissima、披针叶苔草Carex lanceolata。盐肤木群落的物种多样性由高到低依次为灌木层、乔木层、草本层,低均匀度导致物种丰富度最高的草本层具有最低的物种多样性。调查显示盐肤木幼苗幼树多,自我更新良好,群落处于稳定增长状态。通过对群落特征和物种多样性的研究,揭示了昆嵛山盐肤木群落的自然生态规律,为保护和开发利用盐肤木提供了科学依据。     

Invasive insect abundance varies across the biogeographic distribution of a native host plant.

Studies of biogeographic variation in species abundances are fundamental to understanding and predicting the impacts of invasive exotic species. We quantified the abundance of the introduced and now invasive biocontrol weevil, Rhinocyllus conicus, on a newly adopted native host plant, Cirsium canescens (Platte thistle), across the plant's distributional range. We used regression and structural equation analyses to examine variation in weevil abundance at 92-108 sites over three years in relation to variation in abiotic and biotic parameters hypothesized to be important in insect or plant dynamics and distribution. We found that R. conicus now occurs throughout the majority of the range of C. canescens, even in the center of the native plant's distribution where its coevolved, targeted weed host (Carduus nutans, musk thistle) is absent. In fact, weevil densities were greater in the center of the native plant's distribution in the Sand Hills formation than in peripheral sand patches closer to areas where the targeted 'exotic thistle is common. None of the macroclimatic variables examined were consistent predictors of the large-scale variation in weevil abundance on C. canescens. In addition to biogeographic position, the only other consistent predictor of weevil densities across sites was the number of flower heads per C. canescens plant. These results exclude the "spillover" hypothesis to explain nontarget feeding on this newly adopted native host species. Instead, the results are consistent with the alternative hypothesis that exotic weevil abundance on C. canescens is related to the local availability of native floral resources. Because C. canescens densities have declined with increases in R. conicus at sites central in the plant's distribution, these results suggest that isolated, peripheral populations of C. canescens are likely to be critical for persistence of Platte thistle. More generally, this study suggests that the persistence of a native species that is impacted by an exotic natural enemy may require preservation of populations in habitats outside the optimal portion of the native species' distribution.     

Linking abundance and diversity of sponge-associated microbial communities to metabolic differences in host sponges

Many sponge species contain large and diverse communities of microorganisms. Some of these microbes are suggested to be in a mutualistic interaction with their host sponges, but there is little evidence to support these hypotheses. Stable nitrogen isotope ratios of sponges in the Key Largo, Florida (USA) area grouped sponges into species with relatively low δ¹⁵N ratios and species with relatively high δ¹⁵N ratios. Using samples collected in June 2002 from Three Sisters Reef and Conch Reef in the Key Largo, Florida area, transmission electron microscopy (TEM) and denaturing gradient gel electrophoresis were performed on tissues of the sponges Ircinia felix and Aplysina cauliformis, which are in the low δ¹⁵N group, and on tissue of the sponge Niphates erecta, which is in the high δ¹⁵N group. Results showed that I. felix and A. cauliformis have large and diverse microbial communities, while N. erecta has a low biomass of one bacterial strain. As the low δ¹⁵N ratios indicated a microbial input of nitrogen, these results suggested that I. felix and A. cauliformis were receiving nitrogen from their associated microbial community, while N. erecta was obtaining nitrogen solely from external sources. Sequence analysis of the microbial communities showed a diversity of metabolic capabilities among the microbes of the low δ¹⁵N group, which are lacking in the high δ¹⁵N group, further supporting metabolic differences between the two groups. This research provides support for hypotheses of mutualisms between sponges and their associated microbial communities.