Species–area relationships in Mediterranean-climate plant communities

Aim To determine the best‐fit model of species–area relationships for Mediterranean‐type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean‐climate regions. Methods The number of species was recorded from 1, 100 and 1000 m² nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r² values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi‐log slopes at 1–100 m² and 100–1000 m². Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.     

Species richness, endemism, and abundance patterns: tests of two fractal models in a serpentine grassland

Although scaling relationships that characterize fractal species distributions offer an exciting potential for unification in biogeography, empirical support for fractal theory remains the subject of debate. We synthesize and test multiple predictions of two interrelated fractal models and a null model of random placement using Californian serpentine grassland data describing the spatial location of over 37 000 individually identified plants. The endemics–area relationship and species‐abundance distribution recently derived from a community‐level fractal property performed poorly because of an inaccurate assumption of homogeneity among species. In contrast, a species‐level fractal model that incorporates species‐level differences predicted abundances well, but systematically overestimated endemism and predicted a species–area relationship that violated the observed power law. These findings indicate that in order to make predictions based on the existence of a power‐law species–area relationship, ecologists need a unifying theory of how the community‐level fractal property arises in the presence of species‐level distributional differences.     

Taxa-area relationship and neutral dynamics influence the diversity of fungal communities on senesced tree leaves

This study utilized individual senesced sugar maple and beech leaves as natural sampling units within which to quantify saprotrophic fungal diversity. Quantifying communities in individual leaves allowed us to determine if fungi display a classic taxa–area relationship (species richness increasing with area). We found a significant taxa–area relationship for sugar maple leaves, but not beech leaves, consistent with Wright's species‐energy theory. This suggests that energy availability as affected plant biochemistry is a key factor regulating the scaling relationships of fungal diversity. We also compared taxa rank abundance distributions to models associated with niche or neutral theories of community assembly, and tested the influence of leaf type as an environmental niche factor controlling fungal community composition. Among rank abundance distribution models, the zero‐sum model derived from neutral theory showed the best fit to our data. Leaf type explained only 5% of the variability in community composition. Habitat (vernal pool, upland or riparian forest floor) and site of collection explained > 40%, but could be attributed to either niche or neutral processes. Hence, although niche dynamics may regulate fungal communities at the habitat scale, our evidence points towards neutral assembly of saprotrophic fungi on individual leaves, with energy availability constraining the taxa–area relationship.     

Shapes and functions of species–area curves: a review of possible models

Aim This paper reviews possible candidate models that may be used in theoretical modelling and empirical studies of species–area relationships (SARs). The SAR is an important and well‐proven tool in ecology. The power and the exponential functions are by far the models that are best known and most frequently applied to species–area data, but they might not be the most appropriate. Recent work indicates that the shape of species–area curves in arithmetic space is often not convex but sigmoid and also has an upper asymptote. Methods Characteristics of six convex and eight sigmoid models are discussed and interpretations of different parameters summarized. The convex models include the power, exponential, Monod, negative exponential, asymptotic regression and rational functions, and the sigmoid models include the logistic, Gompertz, extreme value, Morgan–Mercer–Flodin, Hill, Michaelis–Menten, Lomolino and Chapman–Richards functions plus the cumulative Weibull and beta‐P distributions. Conclusions There are two main types of species–area curves: sample curves that are inherently convex and isolate curves, which are sigmoid. Both types may have an upper asymptote. A few have attempted to fit convex asymptotic and/or sigmoid models to species–area data instead of the power or exponential models. Some of these or other models reviewed in this paper should be useful, especially if species–area models are to be based more on biological processes and patterns in nature than mere curve fitting. The negative exponential function is an example of a convex model and the cumulative Weibull distribution an example of a sigmoid model that should prove useful. A location parameter may be added to these two and some of the other models to simulate absolute minimum area requirements.     

亚高寒草甸植物群落的中性理论验证

 该文以物种组成较为复杂的青藏高原东部亚高寒草甸为背景,结合最新的群落中性理论,以解释亚高寒草甸草本植物群落的物种分布格局和生物多样性的维持机制。通过对阴坡、阳坡和滩地3个生境进行随机取样调查,用中性模型对所得多样性数据进行拟合,并分别应用置信区间检验、拟合优度检验和多样性指数检验3种方法对拟合效果进行检验。研究结果表明,在拟合优度检验中,3个生境中中性理论预测和实际物种多度分布之间没有显著差异(p>0.05);实际观测值基本全部落入模型预测分布的95%的置信区间之内（仅滩地草本植物群落的63个物种中的1个以及阴坡草本植物群落75个物种中的2个偏离95%的置信区间）;对群落多样性的预测也和实际观测没有显著差异,其中丰富度预测拟合得最好(0.49相似文献   

塔里木荒漠河岸林不同生境群落物种多度分布格局

为解释塔里木荒漠河岸林群落构建和物种多度分布格局形成的机理, 本文以塔里木荒漠河岸林2个不同生境(沙地、河漫滩) 4 ha固定监测样地为研究对象, 基于两样地物种调查数据, 采用统计模型(对数级数模型、对数正态模型、泊松对数正态分布模型、Weibull分布模型)、生态位模型(生态位优先占领模型、断棍模型)和中性理论模型(复合群落零和多项式模型、Volkov模型)拟合荒漠河岸林群落物种多度分布, 并用K-S检验与赤池信息准则(AIC)筛选最优拟合模型。结果表明: (1)随生境恶化(土壤水分降低), 植物物种多度分布曲线变化减小, 群落物种多样性、多度和群落盖度降低, 常见种数减少。(2)选用的3类模型均可拟合荒漠河岸林不同生境群落物种多度分布格局, 统计模型和中性理论模型拟合效果均优于生态位模型。复合群落零和多项式模型对远离河岸的干旱沙地生境拟合效果最好; 对数正态模型和泊松对数正态模型对洪水漫溢的河漫滩生境拟合效果最优; 中性理论模型与统计模型无显著差异。初步推断中性过程在荒漠河岸林群落构建中发挥着主导作用, 但模型拟合结果只能作为推断群落构建过程的必要非充分条件, 不能排除生态位过程的潜在作用。     

鄱阳湖湿地典型草洲植物群落种-面积关系

为了解鄱阳湖湿地草洲植物群落的结构,对其3种典型草洲植物群落种-面积关系进行了研究。采用巢式样方法调查植物物种数量,并用3种非饱和曲线拟合种-面积方程。结果表明,基于实测数据,3个草洲物种数随着取样面积的增加先快速增加后趋于平稳;群落最小取样面积均为30 m~2。幂函数模型对3个草洲群落的种-面积关系拟合效果均为最佳,3个草洲群落的RSE分别为0.35、0.35和0.56,AAD分别为0.23、0.17和0.35,AARD分别为0.06、0.02和0.07,而指数模型、Fisher模型的拟合效果一般。指数模型计算得到的不同比例因子下3个草洲最小取样面积与实际情况最为吻合。这为鄱阳湖区进行植物群落数据采集的样方设置提供了理论参考。     

Bird species numbers in an archipelago of reeds at Lake Velence, Hungary

1 Bird species numbers were studied on 109 reed islands at Lake Velence, Hungary, in the 1993 and 1994 breeding seasons. The aim was to describe and account for the abundance and distribution patterns of the bird species. 2 It was expected that an exponential model would fit the calculated species–area curves. However, for the 1993 data, both the power function (LogS ~ LogArea) and the exponential (S ~ LogArea) models did so, while the power function, exponential and linear (S ~ A) models fitted the curves for the 1994 data. 3 The results showed that the pattern was not random: a collection of small islands held more species than a few large islands with the same total area. 4 The relative species richness of small islands is a result of the preference of most common passerine bird species for the edges of reed islands. Most individuals were found in the first 5 m of the reedbed, and no edge avoidance was detected on a local spatial scale. Large, rarer species (e.g. Great White Egret), however, were found to be dependent on large reed islands. 5 Comparison of results with two other studies on bird communities of reed islands revealed that the type of landscape matrix (e.g. deep water, shallow water or agricultural lands) among reed patches significantly influences bird communities. Deep water was dominated by grebes and coot, shallow water by reed‐nesting passerines, and farmed areas by reed‐ and bush‐nesting passerines.     

Does size really matter? Species–area relationships in human settlements

Aim To describe species–area relationships in human settlements and compare them with those from a non‐urban habitat. Location West‐central Mexico. Methods We surveyed breeding birds in 13 human settlements and five shrubland patches. We estimated bird species richness using an abundance‐based coverage estimator with equal sample sizes to eliminate biases related to sampling effort differences. To assess species–area relationships, we performed log–log linear regressions between the size of the studied patches and their estimated bird richness. We also used a logarithmic approach to determine how the species–area relationship asymptoted and made use of the Michaelis–Menten model to identify the size at which the studied patches reached their maximum species richness. We also investigated (1) possible relationships among the estimated bird richness and other variables known to affect urban‐dwelling birds (built cover, plant species richness, tree cover or human population density) and (2) changes in bird community composition related to the size of the studied human settlements. Results Species–area relationships exhibited different patterns among the studied habitats. The log–log regression slope was steeper in human settlements, while the intercept was higher in shrublands. The maximum number of species was more than twofold higher in shrublands. Human settlement patch size was the only variable significantly related to bird richness. Our community composition results show that two main bird groups are related to human settlement size, and that as the size of human settlements increases, bird community similarity in relation to the largest city increases. Main conclusions Human settlements act as ecological islands, with pronounced species–area relationships. Our results suggest that an important threshold for bird species richness and community composition is reached in human settlements > 10.2 km². This threshold is unlikely to be generalizable among bio‐regions, and thus should be quantified and considered when studying, managing and/or planning urban systems.     

Salt spray differentially affects water status, necrosis, and growth in coastal sandplain heathland species

Sandplain heathlands are disturbance-dependent plant communities that occur infrequently in coastal areas of the northeastern United States. We hypothesize that salt spray plays a role in maintaining the composition of the heathland community by excluding salt-intolerant species close to the ocean. We examined the distributions of Solidago nemoralis, Myrica pensylvanica, Pinus rigida, and Quercus spp. in heathlands and conducted greenhouse studies to determine whether different levels of salt spray tolerance explain patterns found in the field. We found that common heathland forb and shrub species grow closer to the ocean than successional woody species. In greenhouse experiments, these species differ in their water status, necrosis, and growth responses to salt spray. The tree species P. rigida and Q. rubra are more susceptible to salt spray than the common heathland species M. pensylvanica. Our results suggest that salt spray may prevent tree species in heathlands from growing close to the ocean and therefore might be an important factor in maintaining the characteristic community composition of these dwarf shrublands in coastal habitats.     

Effects of sampling protocol on the shapes of species richness curves

Aim Scheiner (Journal of Biogeography, 2009, 36 , 2005–2008) criticized several issues regarding the typology and analysis of species richness curves that were brought forward by Dengler (Journal of Biogeography, 2009, 36 , 728–744). In order to test these two sets of views in greater detail, we used a simulation model of ecological communities to demonstrate the effects of different sampling schemes on the shapes of species richness curves and their extrapolation capability. Methods We simulated five random communities with 100 species on a 64 × 64 grid using random fields. Then we sampled species–area relationships (SARs, contiguous plots) as well as species–sampling relationships (SSRs, non‐contiguous plots) from these communities, both for the full extent and the central quarter of the grid. Finally, we fitted different functions (power, quadratic power, logarithmic, Michaelis–Menten, Lomolino) to the obtained data and assessed their goodness‐of‐fit (Akaike weights) and their extrapolation capability (deviation of the predicted value from the true value). Results We found that power functions gave the best fit for SARs, while for SSRs saturation functions performed better. Curves constructed from data of 32² grid cells gave reasonable extrapolations for 64² grid cells for SARs, irrespective of whether samples were gathered from the full extent or the centre only. By contrast, SSRs worked well for extrapolation only in the latter case. Main conclusions SARs and SSRs have fundamentally different curve shapes. Both sampling strategies can be used for extrapolation of species richness to a target area, but only SARs allow for extrapolation to a larger area than that sampled. These results confirm a fundamental difference between SARs and area‐based SSRs and thus support their typological differentiation.     

Diversity measurements in shrubland communities of Galicia (NW Spain)

Diversity was studied in 10 communities, including the understory of native oak woodland, planted woodlands (pine and eucalypt), and shrublands in the strict sense (heathlands, broom shrublands, gorse shrublands).In each community, species richness, diversity, dominance and evenness were analysed. Differences were observed among communities with regard to species composition, richness in annual herbs, perennial herbs and shrubs, dominant plant families (Ericaceae, Papilionaceae) and diversification of shrub species.The possible relations between environmental stress and/or human influences on differences in diversity are discussed.     

The Distribution of Ocular Chlamydia Prevalence across Tanzanian Communities Where Trachoma Is Declining

BackgroundMathematical models predict an exponential distribution of infection prevalence across communities where a disease is disappearing. Trachoma control programs offer an opportunity to test this hypothesis, as the World Health Organization has targeted trachoma for elimination as a public health concern by the year 2020. Local programs may benefit if a single survey could reveal whether infection was headed towards elimination. Using data from a previously-published 2009 survey, we test the hypothesis that Chlamydia trachomatis prevalence across 75 Tanzanian communities where trachoma had been documented to be disappearing is exponentially distributed.Methods/FindingsWe fit multiple continuous distributions to the Tanzanian data and found the exponential gave the best approximation. Model selection by Akaike Information Criteria (AIC_c) suggested the exponential distribution had the most parsimonious fit to the data. Those distributions which do not include the exponential as a special or limiting case had much lower likelihoods of fitting the observed data. 95% confidence intervals for shape parameter estimates of those distributions which do include the exponential as a special or limiting case were consistent with the exponential. Lastly, goodness-of-fit testing was unable to reject the hypothesis that the prevalence data came from an exponential distribution.ConclusionsModels correctly predict that infection prevalence across communities where a disease is disappearing is best described by an exponential distribution. In Tanzanian communities where local control efforts had reduced the clinical signs of trachoma by 80% over 10 years, an exponential distribution gave the best fit to prevalence data. An exponential distribution has a relatively heavy tail, thus occasional high-prevalence communities are to be expected even when infection is disappearing. A single cross-sectional survey may be able to reveal whether elimination efforts are on-track.     

Effects of grazing on plant species composition and spatial distribution in rangelands of the Sahel

The effects of grazing by livestock on plant species composition and spatial distribution have been studied at Sadoré, Niger. Herbaceous species were recorded in plots of increasing size from 1/64 to 1024 m² in ten fallow plots subjected to five different grazing treatments over the previous three years. Treatments consisted of three intensities of grazing, and of protection from grazing for either 3 or 14 years. For all treatments, the number of species fitted a normal distribution with the log (ln) of the area inventoried. However, the fit improved slightly when the model included two successive log-normal distributions respectively considered as species distribution within a patch and at the patch mosaic scale. Across treatments, the optimal sampling areas averaged 3.8 ± 1.1 m² for the within-patch and 725 ± 113 m² for the mosaic scale distributions. It is argued that similarity between treatments in the overall log-normal distribution resulted from compensations between the divergent trends that affected species distribution within and across patches depending on the grazing status. Long-term protection resulted in a regular spatial arrangement of highly contrasted, but internally homogeneous patches. Heavy grazing ensued the dominance of a few species in contagious patches but also left niches for scattered individuals of other species. Tests of the relative frequency of each plant species, together with the average area needed to record that species, were used to characterize specific response to grazing. A majority of species encountered in old Sahelian fallows were either fostered by grazing, indifferent or tolerant to grazing. However, more than a third of the species appeared sensitive to heavy grazing, and no relationships were found between species response to grazing and palatability.     

Floristic inventory and diversity assessment of a lowland African Montane Rainforest at Korup,Cameroon and implications for conservation

Twenty modified‐Whittaker plots were stratified at different sampling locations from February to May of 2008 in the central zone of Korup National Park, Cameroon. Our interest was to assess floristic diversity and investigate their relationship with environmental variables. Diversity profiles and rank abundance–curves were used for diversity analysis while canonical correspondence analysis and species–response curves were used to investigate the relationships between the response and explanatory variables. Of the 66 families identified, the Rubiaceae (999 species) were the most abundant. The Sterculiaceae (basal area = 10.482 m² ha^?1) were the dominant family, while the co‐dominant families included the Ebenaceae (basal area = 9.092 m² ha^?1) and the Euphorbiaceae (basal area = 8.168 m² ha^?1). Soil variables explained 54.3% of total variation in family distribution. Canonical axes were related to different environmental gradients: axis1 was related to increasing canopy cover (r = 0.6951); axis 2, increasing Magnesium (r = 0.8465) and effective cation exchange capacity (r = 0.5899); axis 3, increasing effective cation exchange capacity (r = 0.5536); while axis 4, increasing Phosphorus concentration (r = 0.5232). Our results demonstrate the advantage which diversity profiles have over single or combination of indices, and the importance of using a combination of methodologies in diversity analysis.     

Checkerboard score–area relationships reveal spatial scales of plant community structure

Identifying the spatial scale at which particular mechanisms influence plant community assembly is crucial to understanding the mechanisms structuring communities. It has long been recognized that many elements of community structure are sensitive to area; however the majority of studies examining patterns of community structure use a single relatively small sampling area. As different assembly mechanisms likely cause patterns at different scales we investigate how plant species co‐occurrence patterns change with sampling unit scale. We use the checkerboard score as an index of species segregation, and examine species C‐score1–sampling area patterns in two ways. First, we show via numerical simulation that the C‐score–area relationship is necessarily hump shaped with respect to sample plot area. Second we examine empirical C‐score–area relationships in arctic tundra, grassland, boreal forest and tropical forest communities. The minimum sampling scale where species co‐occurrence patterns were significantly different from the null model expectation was at 0.1 m² in the tundra, 0.2 m² in grassland, and 0.2 ha in both the boreal and tropical forests. Species were most segregated in their co‐occurrence (maximum C‐score) at 0.3 m² in the tundra (0.54 3 0.54 m quadrats), 1.5 m² in the grassland (1.2 3 1.2 m quadrats), 0.26 ha in the tropical forest (71 3 71 m quadrats), and a maximum was not reached at the largest sampling scale of 1.4 ha in the boreal forest. The most important finding is that the dominant scales of community structure in these systems are large relative to plant body size, and hence we infer that the dominant mechanisms structuring these communities must be at similarly large scales. This provides a method for identifying the spatial scales at which communities are maximally structured; ecologists can use this information to develop hypotheses and experiments to test scale‐specific mechanisms that structure communities.     

Microbial community analysis in incompletely or destructively sampled systems

Analyses of microbial community dynamics are often constrained by the destructive, indirect, and incomplete nature of most sampling techniques. These methodological constraints compel assumptions that are rarely verified about the relationships among separate communities. We evaluated the consequences for community analysis of the common assumption that separate microbial communities are described by the same species abundance distribution. Sample data were generated from simulated communities in which the species abundance distributions were the same or were different. Samples from communities that had the same number of species or were described by the same species abundance distribution sometimes had significantly different numbers of species. Samples from simulated communities that had different species number-species abundance distribution combinations sometimes contained indistinguishable numbers of species. When sampling from independent communities described by unknown distributions (e.g., microbial communities on plant surfaces), the simulations showed that standardization of sample size (number of individuals or colony-forming units) does not guarantee samples of equal proportions of the total species in a community. Sample sizes that are logistically feasible for many microbial systems will provide only limited information for differentiating species numbers or species abundance distributions among separate communities over time. For ecologists studying destructively or incompletely sampled communities this seriously influences both the sample designs that are reasonable and the questions that can be addressed in such systems. Send offprint requests to: L. Kinkel.     

Dynamics of core and occasional species in the marine plankton: tintinnid ciliates in the north-west Mediterranean Sea

Aim To assess short‐term variability in the community composition and community structure of tintinnid ciliates, herbivores of the microzooplankton. Location North‐west Mediterranean Sea. Methods We sampled on 18 dates over a 4‐week period in 2004 at an open‐water site. Species were classified as ‘core species’, found on every date, or ‘occasional species’, absent on one or more dates. Species abundance distributions of the entire community, and separately the core and occasional species, were compared with geometric, log‐series and log‐normal distributions. Core and occasional species were compared in terms of the shell or lorica oral diameter (LOD), analogous to gape size. Results We found 11 core and 49 occasional species. Diversity metrics were stable compared with shifts in abundances. Core species accounted for the majority of individuals in all samples. On each date, 9–22 occasional species, representing 10–15% of the population, were found. Species richness of the occasionals was positively related to population size. The identities of the occasional species found were unrelated to the time between sampling. The species abundance distribution of the occasional population was best fit by a log‐series distribution, while that of the core species was best fit by a log‐normal distribution. The species abundance distribution of the entire community was best fit by a log‐series distribution. Most of the occasional species had LODs distinct from that of a core species and occupied size classes left empty by the core population. However, the most abundant and frequent of the occasional species had a LOD similar to that of a core species. Main conclusions Among tintinnids, which are planktonic protists, occasional species have a species abundance distribution pattern distinct from that of core species. Occasional species appeared to be composed of two groups, one of relatively abundant species and similar to core species, and a second group of ephemeral species with morphologies distinct from core species. The existence of two categories of occasional or rare species may be common: (1) those similar to, and thus perhaps able to replace, dominant species in the absence of a change in the environment; and (2) those distinct from dominant species and requiring different conditions to prosper.     

The portability of foodweb dynamics: reassembling an Australian eucalypt–psyllid–bird association within California

Aims To evaluate the role of native predators (birds) within an Australian foodweb (lerp psyllids and eucalyptus trees) reassembled in California. Location Eucalyptus groves within Santa Cruz, California. Methods We compared bird diversity and abundance between a eucalyptus grove infested with lerp psyllids and a grove that was uninfested, using point counts. We documented shifts in the foraging behaviour of birds between the groves using structured behavioural observations. Additionally, we judged the effect of bird foraging on lerp psyllid abundance using exclosure experiments. Results We found a greater richness and abundance of Californian birds within a psyllid infested eucalyptus grove compared to a matched non‐infested grove, and that Californian birds modify their foraging behaviour within the infested grove in order to concentrate on ingesting psyllids. This suggests that Californian birds could provide indirect top‐down benefits to eucalyptus trees similar to those observed in Australia. However, using bird exclosure experiments, we found no evidence of top‐down control of lerp psyllids by Californian birds. Main conclusions We suggest that physiological and foraging differences between Californian and Australian pysllid‐eating birds account for the failure to observe top‐down control of psyllid populations in California. The increasing rate of non‐indigenous species invasions has produced local biotas that are almost entirely composed of non‐indigenous species. This example illustrates the complex nature of cosmopolitan native‐exotic food webs, and the ecological insights obtainable through their study.