Metacommunity analysis of Mexican bats: environmentally mediated structure in an area of high geographic and environmental complexity

Aim We tested the hypothesis that distributions of Mexican bats are defined by shared responses to environmental gradients for the entire Mexican bat metacommunity and for each of four metaensembles (frugivores, nectarivores, gleaning insectivores, and aerial insectivores). Further, we identified the main environmental factors to which bats respond for multiple spatial extents. Location Mexico. Methods Using bat presence–absence data, as well as vegetation composition for each of 31 sites, we analysed metacommunity structure via a comprehensive, hierarchical approach that uses reciprocal averaging (RA) to detect latent environmental gradients corresponding to each metacommunity structure (e.g. Clementsian, Gleasonian, nested, random). Canonical correspondence analysis (CCA) was used to relate such gradients to variation in vegetation composition. Results For all bat species and for each ensemble, the primary gradient of ordination from RA, which is based on species data only, recovered an axis of humidity that matched that obtained for the first axis of the CCA ordination, which is based both on vegetation attributes and on species composition of sites. For the complete assemblage as well as for aerial and gleaning insectivores, analyses revealed Clementsian or quasi‐Clementsian structures with discrete compartments (distinctive groups of species along portions of an environmental gradient) coincident with the humidity gradient and with the Nearctic–Neotropical divide. Within‐compartment analysis further revealed Clementsian or quasi‐Clementsian structures corresponding to a gradient of elevational complexity that matched the second ordination axis in CCA. Frugivores had quasi‐nested structure, whereas nectarivores had Gleasonian structure. Main conclusions Our hierarchical approach to metacommunity analysis detected complex metacommunity structures associated with multiple environmental gradients at different spatial extents. More importantly, the resulting structures and their extent along environmental gradients are determined by ensemble‐specific characteristics and not by arbitrarily circumscribed study areas. This property renders compartment‐level analyses particularly useful for large‐scale ecological analyses in areas where more than one gradient may exist and species sorting may occur at multiple scales.     

Bat metacommunity structure on Caribbean islands and the role of endemics

Aim We evaluate characteristics of species ranges (i.e. coherence, species turnover and range boundary clumping) to determine the structure of bat metacommunities and metaensembles from Caribbean islands. We evaluate the effects of endemic species on that structure, and quantify associations between island characteristics and latent environmental gradients that structure these metacommunities and metaensembles. Location Sixty‐five Caribbean islands throughout the Bahamas, Greater Antilles and Lesser Antilles. Methods Metacommunity structure is an emergent property of a set of ecological communities at different sites defined by species distributions across geographic or environmental gradients. We analysed elements of metacommunity structure (coherence, range turnover and range boundary clumping) to determine the best‐fit pattern for metacommunities from all Caribbean islands, as well as from the Bahamas, the Greater Antilles and the Lesser Antilles separately. For each island group, analyses were conducted for all bats and for each of two broadly defined guilds (i.e. carnivores and herbivores). In addition, analyses were conducted for all species and for a subset in which endemic species were removed from the fauna. Spearman rank correlations identified island characteristics (area, elevation, latitude, longitude) that were associated significantly with island scores for ordination axes based on reciprocal averaging. Results Metacommunity structure for all bats and for carnivores was similar for each island group, with Clementsian distributions (i.e. discrete communities with groups of species replacing other groups of species along the gradient) for all islands, the Bahamas and the Lesser Antilles, but with nested distributions for the Greater Antilles. Herbivore distributions were random for the Bahamas, but were Clementsian for all other island groups. Removal of endemic species affected the best‐fit model of metacommunity structure in only 3 of 12 cases. In general, ordination scores for islands were correlated with longitude or latitude, but not with island area or elevation. Main conclusions Characteristics of bat species ranges and associated metacommunity structure were primarily dependent on the number and geographic arrangement of primary sources of colonization, and not on interspecific interactions, species‐specific levels of environmental tolerance, or the physical characteristics of islands. Endemic species did not greatly affect metacommunity structure in Caribbean bats.     

Plant metacommunity structure remains unchanged during biodiversity loss in English woodlands

The metacommunity concept provides important insights into large‐scale patterns and dynamics of distributions of interacting species. However, temporal change of metacommunity structure is little studied and has not been previously analysed in the context of biodiversity change. As metacommunity structure is determined by multiple species distributions, it is expected to change as a result of biodiversity loss. To examine this process, we analysed structural change of a southern English woodland metacommunity at two points in time, seven decades apart. During this interval, the metacommunity lost β‐diversity through taxonomic homogenization. We performed an ‘elements of metacommunity structure’ (EMS) analysis to examine metacommunity structure, based upon three structural elements: coherence (i.e. gaps in species range along a structuring gradient), spatial turnover (replacements), and species range boundary clumping. We predicted that metacommunity structure would decrease in spatial turnover and thus become more nested over time. We tested for change in individual structural elements with z‐scores and examined the role of spatial and environmental variables as potential structuring mechanisms through correlation with EMS ordination axes. Our results demonstrated that the metacommunity had a Clementsian structure that was maintained over time. Despite no change in broad structure, coherence and species range boundary clumping increased. Spatial turnover increased along the first structuring gradient but decreased on the second gradient. We hypothesise that this difference between gradients may reflect the presence of competing processes affecting spatial turnover. The mechanisms of biological structuring involved both environmental and spatial factors at the scale of the individual woodland. Therefore, our results suggest that broad metacommunity structure would not be a good landscape‐scale indicator for conservation status. Conversely, knowledge that metacommunity structure does not change over time could assist in long‐term conservation strategy because fundamental metacommunity structural processes are resistant to environmental change.     

A Complex Metacommunity Structure for Gastropods Along an Elevational Gradient

The metacommunity framework integrates species‐specific responses to environmental gradients to detect emergent patterns of mesoscale organization. Abiotic characteristics (temperature, precipitation) and associated vegetation types change with elevation in a predictable fashion, providing opportunities to decouple effects of environmental gradients per se from those of biogeographical or historical origin. Moreover, expected structure is different if a metacommunity along an elevational gradient is molded by idiosyncratic responses to abiotic variables (expectation=Gleasonian structure) than if such a metacommunity is molded by strong habitat preferences or specializations (expectation=Clementsian structure). We evaluated metacommunity structure for 13 species of gastropod from 15 sites along an elevational transect in the Luquillo Experimental Forest of Puerto Rico. Analyses were conducted separately for the primary axis and for the secondary axis of correspondence extracted via reciprocal averaging. The metacommunity exhibited quasi‐Clementsian structure along the primary axis, which represented a gradient of gastropod species specialization that was unassociated with elevation. The secondary axis represented environmental variation associated with elevation. Along this axis, the metacommunity exhibited Clementsian structure, with specialists characterizing each of three suites of sites that corresponded to three distinct forest types. These forest types are associated with low (tabonuco forest), mid‐ (palo colorado forest), or high (elfin forest) elevations. Thus, variation among sites in species composition reflected two independent processes: the first decoupled from elevational variation and its environmental correlates, and the second highly associated with environmental variation correlated with elevation. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Identifying the mechanisms that shape fungal community and metacommunity patterns in Yunnan,China

Fungi are key organisms in terrestrial ecosystems, functioning as decomposers, pathogens, and symbionts. Identifying the mechanisms that shape metacommunity patterns is likely to be critical for predicting how ecosystems will respond to global environmental change. Using fungal occurrence data and a hierarchical approach that combines three elements of metacommunity structure—coherence, turnover and boundary clumping—we identified the structures that best describe metacommunity patterns. We related these patterns to underlying environmental and spatial variables known to influence fungal distribution, and determined the relative importance of the environment and geographic distance in structuring fungal metacommunities. Fungal metacommunities had Clementsian and quasi-Clementsian structures, indicating that species distributions were compartmentalized along a dominant environmental gradient. This gradient was strongly associated with annual precipitation, precipitation seasonality and pH for the entire metacommunity. Variance partitioning revealed that the environment was relatively more important than geographic distance in explaining metacommunity patterns, indicating that niche-based processes are crucial in shaping species distributions among sites. However, the strength of the relationship between the latent gradient and environmental factors and the relative contributions of the environment and geographic distance to metacommunity structure varied across groups, suggesting that interactions among habitat, dispersal and life-history might be driving these differences.     

Vertebrate metacommunity structure along an extensive elevational gradient in the tropics: a comparison of bats,rodents and birds

Aim We evaluated the structure of metacommunities for each of three vertebrate orders (Chiroptera, Rodentia and Passeriformes) along an extensive elevational gradient. Using elevation as a proxy for variation in abiotic characteristics and the known elevational distributions of habitat types, we assessed the extent to which variation in those factors may structure each metacommunity based on taxon‐specific characteristics. Location Manu Biosphere Reserve in the Peruvian Andes. Methods Metacommunity structure is an emergent property of a set of species distributions across geographic or environmental gradients. We analysed elements of metacommunity structure (coherence, range turnover and range boundary clumping) to determine the best‐fit structure for each metacommunity along an elevational gradient comprising 13 250‐m elevational intervals and 58 species of rodent, 92 species of bat or 586 species of passerine. Results For each taxon, the environmental gradient along which the metacommunity was structured was highly correlated with elevation. Clementsian structure (i.e. groups of species replacing other such groups along the gradient) characterized rodents, with a group of species that was characteristic of rain forests and a group of species that was characteristic of higher elevation habitats (i.e. above 1500 m). Distributions of bats were strongly nested, with more montane communities comprising subsets of species at lower elevations. The structure of the passerine metacommunity was complex and most consistent with a quasi‐Clementsian structure. Main conclusions Each metacommunity exhibited a different structure along the same elevational gradient, and each structure can be accounted for by taxon‐specific responses to local environmental factors that vary predictably with elevation. The structures of rodent and bird metacommunities suggest species sorting associated with habitat specializations, whereas structure of the bat metacommunity is probably moulded by a combination of species‐specific tolerances to increasingly cold, low‐productivity environs of higher elevations and the diversity and abundance of food resources associated with particular habitat types.     

A comparative analysis of metacommunity types in the freshwater realm

Most metacommunity studies have taken a direct mechanistic approach, aiming to model the effects of local and regional processes on local communities within a metacommunity. An alternative approach is to focus on emergent patterns at the metacommunity level through applying the elements of metacommunity structure (EMS; Oikos, 97, 2002, 237) analysis. The EMS approach has very rarely been applied in the context of a comparative analysis of metacommunity types of main microbial, plant, and animal groups. Furthermore, to our knowledge, no study has associated metacommunity types with their potential ecological correlates in the freshwater realm. We assembled data for 45 freshwater metacommunities, incorporating biologically highly disparate organismal groups (i.e., bacteria, algae, macrophytes, invertebrates, and fish). We first examined ecological correlates (e.g., matrix properties, beta diversity, and average characteristics of a metacommunity, including body size, trophic group, ecosystem type, life form, and dispersal mode) of the three elements of metacommunity structure (i.e., coherence, turnover, and boundary clumping). Second, based on those three elements, we determined which metacommunity types prevailed in freshwater systems and which ecological correlates best discriminated among the observed metacommunity types. We found that the three elements of metacommunity structure were not strongly related to the ecological correlates, except that turnover was positively related to beta diversity. We observed six metacommunity types. The most common were Clementsian and quasi‐nested metacommunity types, whereas Random, quasi‐Clementsian, Gleasonian, and quasi‐Gleasonian types were less common. These six metacommunity types were best discriminated by beta diversity and the first axis of metacommunity ecological traits, ranging from metacommunities of producer organisms occurring in streams to those of large predatory organisms occurring in lakes. Our results showed that focusing on the emergent properties of multiple metacommunities provides information additional to that obtained in studies examining variation in local community structure within a metacommunity.     

Tropical metacommunities along elevational gradients: effects of forest type and other environmental factors

Elevational gradients provide a natural experiment for assessing the extent to which the structure of animal metacommunities is molded by biotic and abiotic characteristics that change gradually, or is molded by aspects of plant community composition and physiognomy that change in a more discrete fashion. We used a metacommunity framework to integrate species‐specific responses to environmental gradients as an approach to detect emergent patterns at the mesoscale in the Luquillo Mountains of Puerto Rico. Elements of metacommunity structure (coherence, species turnover and range boundary clumping) formed the basis for distinguishing among random, checkerboard, Gleasonian, Clementsian, evenly spaced and nested patterns. Paired elevational transects (300–1000 m a.s.l.) were sampled at 50 m intervals to decouple underlying environmental mechanisms: a mixed forest transect reflected changes in abiotic and biotic conditions, including forest type (i.e. tabonuco, palo colorado and elfin forests), whereas another transect reflected changes in environmental conditions but not forest type, as its constituent plots were located within palm forest. Based on distributional data (presence versus absence of species), the mixed forest transect exhibited Clementsian structure, whereas the palm forest transect exhibited quasi‐Gleasonian structure. In contrast, the distribution of modes in species abundance was random with respect to the latent environmental gradient in the mixed forest transect and clumped with respect to the latent environmental gradient in the palm forest transect. Such contrasts suggest that the environmental factors affecting abundance differed in form or type from those affecting distributional boundaries. Variation among elevational strata with respect to the first axis of correspondence from reciprocal averaging was highly correlated with elevation along each transect, even though axis scores were not correlated between mixed forest and palm forest transects. This suggests that the identity of the environmental characteristics, or the form of response by the fauna to those characteristics, differed between the two elevational transects. Despite the proximity of the transects, the patchy configuration of palm forest, and the pervasive distribution of the dominant palm species, the relative importance of abiotic variables and habitat in structuring gastropod metacommunities differed between transects, which is remarkable and attests to the sensitivity of metacommunity structure to environmental variation.     

小兴安岭阔叶红松林地表甲虫Metacommunity格局

Metacommunity理论框架为理解生物群落的时空格局及其构建机制提供了一个强有力的方法。然而,少有研究揭示土壤动物metacommunity的格局及其构建机制,小尺度空间的研究更是少见。于2015年分别在凉水和丰林小兴安岭典型阔叶红松林长期动态监测样地内,通过空间直观定位调查监测的方法获得地表甲虫群落,揭示小尺度空间(300 m)地表甲虫metacommunity的格局,并进一步分析地形、土壤因子与这种metacommunity格局的相关性。结果表明:镶嵌型(nestedness)和随机型(random)是小尺度空间地表甲虫metacommunity的常见格局。地表甲虫的metacommunity格局具有类群依赖性,步行虫科和葬甲科多形成镶嵌型格局,而隐翅虫科则仅形成随机型格局。地表甲虫metacommunity格局具有季节波动性,在相对温暖的季节易形成显著的镶嵌型格局,而在相对寒冷的季节多为随机型格局。凉水和丰林地表甲虫的metacommunity格局没有明显差异。地形和土壤因子对地表甲虫metacommunity格局有重要的影响,该影响过程相对复杂,依类群和时间而异。本实验表明镶嵌型和随机型是小尺度空间小兴安岭典型阔叶红松林地表甲虫的常见格局,地形、土壤因子对这些格局的形成有重要的影响。     

A comprehensive framework for the evaluation of metacommunity structure

The metacommunity framework is a powerful platform for evaluating patterns of species distribution in geographic or environmental space. Idealized patterns (checkerboard, Clementsian, evenly spaced, Gleasonian and nested distributions) give the framework shape. Each pattern represents an area in a multidimensional continuum of metacommunity structures; however, the current approach to analysis of spatial structure of metacommunities is incomplete. To address this, we describe additional non‐random structures and illustrate how they may be discerned via objective criteria. First, we distinguish three distinct forms of species loss in nested structures, which should improve identification of structuring mechanisms for nested patterns. Second, we define six quasi‐structures that are consistent with the conceptual underpinnings of Clementsian, Gleasonian, evenly spaced and nested distributions. Finally, we demonstrate how combinations of structures at smaller spatial extents may aggregate to form Clementsian structure at larger extents. These refinements should facilitate the identification of best‐fit patterns, associated structuring mechanisms, and informative scales of analysis and interpretation. This conceptual and analytical framework may be applied to network properties within communities (i.e. structure of interspecific interactions) and has broad application in ecology and biogeography.     

Metacommunities of spiders in grassland habitat fragments of an agricultural landscape

Arthropod communities in fragmented agricultural landscapes depend on local processes and the interactions between communities in the habitat islands. We aimed to study metacommunity structure of spiders, a group that is known for high dispersal power, local niche partitioning and for engaging in species interactions. While living in fragmented habitats could lead to nestedness, other ecological traits of spiders might equally lead to patterns dominated either by species interactions or habitat filtering. We asked, which community pattern will prevail in a typical agricultural landscape with isolated patches of semi-natural habitats. Such a situation was studied by sampling spiders in 28 grassland locations in a Hungarian agricultural landscape. We used the elements of metacommunity structure (EMS) framework to distinguish between alternative patterns that reveal community organization. The EMS analysis indicated coherent species ranges, high turnover and boundary clumping, suggesting Clementsian community organization. The greatest variation in species composition was explained by local habitat characteristics, indicating habitat filtering. The influence of dispersal could be detected by the significant effect of landscape composition, which was strongest at 500 m. We conclude that dispersal allows spiders to respond coherently to the environment, creating similar communities in similar habitats. Consistent habitat differences, such as species rich versus species poor vegetation, lead to recognisably different, recurrent communities. These characteristics make spiders a predictable and diverse source of natural enemies in agricultural landscapes. Sensitivity to habitat composition at medium distances warns us that landscape homogenization may alter these metacommunity processes.     

Bird dietary guild richness across latitudes,environments and biogeographic regions

Aim To integrate dietary knowledge and species distributions in order to examine the latitudinal, environmental, and biogeographical variation in the species richness of avian dietary guilds (herbivores, granivores, frugivores, nectarivores, aerial insectivores, terrestrial/arboreal insectivores, carnivores, scavengers, and omnivores). Location Global. Methods We used global breeding range maps and a comprehensive dietary database of all terrestrial bird species to calculate guild species richness for grid cells at 110 × 110 km resolution. We assessed congruence of guild species richness, quantified the steepness of latitudinal gradients and examined the covariation between species richness and climate, topography, habitat diversity and biogeographic history. We evaluated the potential of current environment and biogeographic history to explain global guild distribution and compare observed richness–environment relationships with those derived from random subsets of the global species pool. Results While most guilds (except herbivores and scavengers) showed strong congruence with overall bird richness, covariation in richness between guilds varied markedly. Guilds exhibited different peaks in species richness in geographical and multivariate environmental space, and observed richness–environment relationships mostly differed from random expectations. Latitudinal gradients in species richness were steepest for terrestrial/arboreal insectivores, intermediate for frugivores, granivores and carnivores, and shallower for all other guilds. Actual evapotranspiration emerged as the strongest climatic predictor for frugivores and insectivores, seasonality for nectarivores, and temperature for herbivores and scavengers (with opposite direction of temperature effect). Differences in species richness between biogeographic regions were strongest for frugivores and nectarivores and were evident for nectarivores, omnivores and scavengers when present‐day environment was statistically controlled for. Guild richness–environment relationships also varied between regions. Main conclusions Global associations of bird species richness with environmental and biogeographic variables show pronounced differences between guilds. Geographic patterns of bird diversity might thus result from several processes including evolutionary innovations in dietary preferences and environmental constraints on the distribution and diversification of food resources.     

Molecular and morphological evidence for conspecificity of two common Indo-Pacific species of Palythoa (Cnidaria: Anthozoa)

Most studies characterize metacommunities based on a single snapshot of the spatial structure, which may be inadequate for taxa with high migratory behavior (e.g., fish). Here, we applied elements of metacommunity structure to examine variations in the spatial distributions of stream fishes over time and to explore possible structuring mechanisms. Although the major environmental gradients influencing species distributions remained largely the same in time, the best-fit pattern of metacommunity structure varied according to sampling occasion and whether or not we included non-native species in the analyses. Quasi-Clementsian and Clementsian structures were the predominant best-fit structures, indicating the importance of species turnover among sites and the existence of more or less discrete community boundaries. The environmental gradient most correlated with metacommunity structure was defined by altitude, area of artificial ponds in the catchment, and dissolved oxygen content. Our results suggest that the best-fit metacommunity structure of the native species can change in time in this catchment due to seasonal changes in distribution patterns. However, the distribution of non-native species throughout the landscape homogenizes the temporal variability in metacommunity structure of native species. Further studies are necessary from other regions to examine best-fit metacommunity structures of stream fishes within relatively short environmental gradients.     

Effect of landscape context on fish metacommunity structuring in stream networks

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Bird metacommunity processes remain constant after 25 years of landscape changes

Metacommunity theory provides a framework to understand how ecological communities vary in space and time. However, few studies have investigated metacommunity processes in a context of long term changes. Environmental changes can impact species distribution and therefore the structure of metacommunities. Using two complementary methods, this study evaluated the temporal variability of bird metacommunity processes in an agricultural landscape after 25 years of changes in land-cover. Bird and landscape data were recorded in the same locations using a series of 256 point counts in 1982 and 2007. First, variance partitioning was applied to quantify the roles of environmental filtering (i.e. landscape composition variables) and spatial variables to organize bird metacommunities each year. Second, changes in the structure of the bird metacommunity were examined by quantifying three of its components: coherence, species turnover and species range boundary clumping. Our results demonstrate that landscape variables explained slightly more bird metacommunity patterns than spatial variation of unknown origin each year. The bird metacommunity had a Clementsian structure (i.e. grouped distribution of species along environmental gradients) which was correlated with a landscape gradient ranging from open farmland to wooded sites. This structure was similar each year. To conclude, the study shows that environmental filtering with specializations to different habitats is a major process in determining bird metacommunities in landscapes. Moreover, our results suggest that metacommunity structure can remain constant over time despite demographic and environmental changes.     

Relative importance of host environment,transmission potential and host phylogeny to the structure of parasite metacommunities

Identification of mechanisms that shape parasite community and metacommunity structures have important implications to host health, disease transmission, and the understanding of community assembly in general. Using a long‐term dataset on parasites from desert rodents, we examined the relative contributions of host traits that represent important aspects of parasite environment, transmission probability between host species, and host phylogeny to the structure of a parasite metacommunity as well as for taxonomically restricted parasite metacommunities (coccidians, ectoparasites and helminths). This was done using a combination of metacommunity analysis and variance partitioning based on canonical correspondence analysis. Coccidian and ectoparasite metacommunities did not exhibit coherent structure. In contrast, helminths and the full parasite metacommunity had Clementsian and quasi‐Clementsian structure, respectively, indicating that parasite species distributions for these metacommunities were compartmentalized along a dominant gradient. Variance decomposition indicated that characteristics associated with the host environment consistently explained more variation than did host traits associated with transmission opportunities or host phylogeny, indicating that the host environment is primary in shaping parasite species distributions among host species. Moreover, the importance of different types of host traits in structuring parasite metacommunities was consistent among taxonomic groups (i.e. full metacommunity, coccidians, and helminths) despite manifest differences in emergent structures (i.e. Clementsian, quasi‐Clementsian, and random) that arose in response to variation in host environment.     

Density compensation in New World bat communities

Understanding the role of competitive interactions in shaping the structure of communities has been one the most unrelenting challenges to ecology. Traditionally, competitive interactions were assumed to be the most important agent of deterministic structure, with overdispersed morphological patterns based on body size and trophic status as their hallmark. However, models of community organization based solely on morphology have yielded only equivocal results for many taxa. Fortunately, morphological patterns may not be the only indications of competitively induced deterministic structure. Herein, we explore the degree to which the structure of five feeding guilds (aerial insectivores, frugivores, gleaning animalivores, molossid insectivores, and nectarivores) from 15 New World bat communities reflects density compensation. Nonrandom associations between abundance and morphological distance were detected in five communities, in three feeding guilds, and with respect to four competitive scenarios. Nonetheless, patterns consistent with the hypothesis of density compensation were neither pervasive nor consistent in New World bat communities. Competitively induced community structure may exist under only narrow temporal and environmental conditions, and may not be characteristic of organization in most situations.     

Finding a common ordination for several data sets by individual differences scaling

Individual differences scaling is a multidimensional scaling method for finding a common ordination for several data sets. An individual ordination for each data set can then be derived from the common ordination by adjusting the axis lengths so as to maximize the correlations between observed proximities and individual ordination distances. The importance of the various axes for each data set and the mutual similarities and goodness of fit for the individual data sets are described by weight plots. As an example, 46 soft-water lakes in eastern Finland are ordinated on two dimensions according to 3 chemical data sets (water in summer and autumn, sediment) and 4 biological sets (major phytoplankton groups, phytoplankton, surface sediment diatom and cladoceran assemblages). The method seems to be effective as a means of ordination for obtaining the common ordination for the data sets. The major taxonomic groups gave the ordination which differed most clearly from the ordinations of the other data sets. Phytoplankton was most poorly ordinated in all the analyses. The other data sets were fairly coherent. When only biological data sets were ordinated, the diatoms and cladocerans showed rather different patterns. It seems that the cladocerans are best correlated with water chemistry, both according to weights in the joint analysis, and according to correlation between the axes from the biological data sets and the chemical variables.Abbreviations CCA = Canonical correspondence analysis - IDS = Individual differences scaling - MDS = multidimensional scaling - PCA = Principal components analysis     

Evaluating the role of biotic interactions in structuring communities using a gradient analysis of multiple interacting guilds

The extent to which ecological communities are coherent entities as opposed to mere intersections of individual species distributions has long been one of the fundamental questions of ecology. Gradient analysis is one commonly used tool for addressing this question; however, all such studies have used organisms from a single taxon or guild. This risks missing important connections due to non‐competitive interactions, which should be more likely to occur between members of different guilds. Such organisms are unlikely to compete for resources and can have complementary niches that promote non‐competitive interactions. We examined the abundances of taxa in four interacting guilds along an elevation gradient in a forest in the southern Appalachian mountains. A causal discovery algorithm was used to investigate the relative frequencies of interguild and intraguild interactions. These were approximately equally common once taxonomic richness was taken into account. We used elements of metacommunity structure analysis to study the extent to which species distributions are non‐independent and tested the hypothesis that combinations of two or more interacting guilds exhibit more coherence than single guilds. In this analysis, all guilds other than collembola were classified as Clementsian or quasi‐Clementsian. (Collembola were classified as random.) When sets of multiple guilds were examined, Clementsian and quasi‐Clementsian structures predominated. We also compared boundary conjunction, measured as Morisita’s index (MI) for these sets of guilds to the weighted average of the guilds’ MI values. Only sets of directly interacting guilds had higher‐than‐baseline boundary conjunction values, and such boundary conjunction values are found in all but one set of directly interacting guilds. Our results highlight the importance of inter‐guild interactions in structuring patterns of cooccurrence. Trophic interactions and plant–fungus symbioses (mutualistic and/or pathogen–host) appear particularly important.