The relative importance of local conditions and regional processes in structuring aquatic plant communities

1. The structure of biological communities reflects the influence of both local environmental conditions and processes such as dispersal that create patterns in species’ distribution across a region. 2. We extend explicit tests of the relative importance of local environmental conditions and regional spatial processes to aquatic plants, a group traditionally thought to be little limited by dispersal. We used partial canonical correspondence analysis and partial Mantel tests to analyse data from 98 lakes and ponds across Connecticut (northeastern United States). 3. We found that aquatic plant community structure reflects the influence of local conditions (pH, conductivity, water clarity, lake area, maximum depth) as well as regional processes. 4. Only 27% of variation in a presence/absence matrix was explained by environmental conditions and spatial processes such as dispersal. Of the total explained, 45% was related to environmental conditions and 40% to spatial processes. 5. Jaccard similarity declined with Euclidean distance between lakes, even after accounting for the increasing difference in environmental conditions, suggesting that dispersal limitation may influence community composition in the region. 6. The distribution of distances among lakes where species occurred was associated with dispersal‐related functional traits, providing additional evidence that dispersal ability varies among species in ways that affect community composition. 7. Although environmental and spatial variables explained a significant amount of variation in community structure, a substantial amount of stochasticity also affects these communities, probably associated with unpredictable colonisation and persistence of the plants.     

Patterns of parasite community dissimilarity: the significant role of land use and lack of distance‐decay in a bat–helminth system

Increasing community dissimilarity across geographic distance has been described for a wide variety of organisms and understanding its underlying causes is key to understanding mechanisms driving patterns of biodiversity. Both niche‐based and neutral processes may produce a distance decay relationship; however, disentangling their relative influence requires simultaneous examination of multiple potential drivers. Parasites represent a unique opportunity in which to study distance decay because community dissimilarity may depend on environmental requirements and dispersal capability of parasites as well also those of their hosts. We used big brown bats Eptesicus fuscus and their intestinal helminths to investigate: 1) independent contributions of geographic and environmental distances on dissimilarity of intestinal helminth component communities between populations of big brown bats; 2) which environmental variables best explained variation in community dissimilarity; and 3) whether similar patterns of decay with geographic or environmental distance were observed for within‐host population and within‐individual host parasite communities. We used compositional measures of community dissimilarity to examine how parasite communities may change with geographic distance and varying environmental conditions. Non‐spatial variables strongly influenced compositional parasite community dissimilarity over multiple community scales, and we observed little evidence for spatial processes such as distance decay. Environment surrounding roost sites better predicted helminth community dissimilarity than any other class of variables and landcover classes representing anthropogenic modification consistently explained variation in community structure. Our results indicate that human disturbance drives significant patterns of parasite community dissimilarity, most likely by changing the presence or abundance of intermediate hosts in an area.     

The relative importance of dispersal and local processes in structuring phytoplankton communities in a set of highly interconnected ponds

1. The recognition that both local and regional processes act together in shaping local communities makes determining their relative roles in natural communities central to understanding patterns in community structure. 2. We investigated the relative influence of these processes on the phytoplankton communities of a highly interconnected pond system. We sampled the phytoplankton communities of 28 ponds concurrently with 20 local environmental variables. 3. We found that phytoplankton community variation, in terms of both phytoplankton community composition (PCC) and diversity, was only significantly explained by local environmental variables. These were mainly associated with the contrasting clear‐water and turbid ecological states of the shallow ponds studied. Clear‐water conditions favoured only a few taxa, resulting in a significantly lower taxon diversity and richness under these conditions. 4. The failure to explain variation in PCC by a dispersal model based on the water flow between ponds points at very effective species sorting. This is attributed to the high population turn‐over rates and sensitivity to environmental conditions of phytoplankton communities. Some evidence was found, however, that dispersal influences local communities through mass effects between neighbouring ponds. 5. Overall, our results emphasize both the strong selection pressure that components of the food web exert on phytoplankton communities and the high potential of these communities to respond to such environmental change, thereby effectively opposing the homogenizing effects of continuous dispersal.     

Latitudinal gradient in the taxonomic composition of parasite communities

Although latitudinal gradients in diversity have been well studied, latitudinal variation in the taxonomic composition of communities has received less attention. Here, we use a large dataset including 950 surveys of helminth endoparasite communities in 650 species of vertebrate hosts to test for latitudinal changes in the relative contributions of trematodes, cestodes, nematodes and acanthocephalans to parasite assemblages. Although the species richness of helminth communities showed no consistent latitudinal variation, their taxonomic composition varied as a function of both host type and latitude. First, trematodes and acanthocephalans accounted for a higher proportion of species in helminth communities of fish, whereas nematodes achieved a higher proportion of the species in communities of bird and especially mammal hosts. Second, the proportion of trematodes in helminth communities of birds and mammals increased toward higher latitudes. Finally, the proportion of nematodes per community increased toward lower latitudes regardless of the type of host. We present tentative explanations for these patterns, and argue that new insights in parasite community ecology can be gained by searching for latitudinal gradients not only in parasite species richness, but also in the taxonomic composition of parasite assemblages.     

The predictability of helminth community structure in space: a comparison of fish populations from adjacent lakes

Patterns in helminth community structure can suggest that various processes are acting to shape parasite communities into organised, non-random assemblages of species. It is not clear, however, whether a pattern observed in one host population at one time would be observed again at another time, or at the same time in a different but comparable host population. Here, we test the repeatability of parasite community structure in space, and to a lesser extent time, with data on helminth parasites of two fish species, perch Perca fluviatilis and roach Rutilus rutilus, collected in different seasons from four adjacent lakes in Central Finland. Since populations of the same fish species harbour the same parasite species and were sampled in the same way, we would expect similar patterns in the structure of their helminth parasites if the same structuring processes are acting in all lakes. We found that no pairwise association between the most common helminth species were observed consistently between seasons within lakes, or among lakes during the same season. Similarly, nested subset patterns of species assembly were observed in some samples, but not consistently between seasons or among lakes. The lack of repeatability in space and between seasons shown by these analyses indicates that although helminth community structure often departs from randomness, it does not do so in a consistent and predictable manner. There may be some general, large-scale processes acting to structure helminth communities, but local or seasonal influences can often either mask their action, or play more important roles themselves.     

The structure of helminth communities

The existence of both interactive and isolationist communities of helminths, each of which may be produced by any one of several processes, necessitates a pluralistic view of helminth community structure. A scheme involving three hierarchical scales of communities of helminths is proposed. Interactions between species occur in infracommunities, and should be looked for there. Interactions are often clearer in smaller groups of species, either among core species or within guilds. Infracommunities are samples of helminth communities at two larger scales: the helminth community in the host population provides core species, largely specialists, and that in the host community provides the generalists and the satellite species. The richness of each of the two large-scale communities is affected by various ecological, historical and evolutionary factors. The different concepts which have been applied to parasite communities are based on these factors.     

Evaluating the role of regional and local processes in structuring a larval trematode metacommunity of Helisoma trivolvis

Metacommunity theory has advanced our understanding of how local and regional processes affect the structure of ecological communities. While parasites have largely been omitted from metacommunity research, parasite communities can provide the large sample sizes and discrete boundaries often required for evaluating metacommunity patterns. Here, we used assemblages of flatworm parasites that infect freshwater snails (Helisoma trivolvis) to evaluate three questions: 1) what factors affect individual host infections within ponds? 2) Is the parasite metacommunity structured among ponds? And 3) what is the relative role of local versus regional processes in determining metacommunity structure and species richness among ponds? We examined 10 821 snails from 96 sites in five park complexes in the San Francisco Bay area, California, and found 953 infections from six parasite groups. At the within‐pond level, infection status of host snails correlated positively with individual snail size and pond infection prevalence for all six parasite groups. Using an ordination method to test for metacommunity structure, we found that the parasite metacommunity was organized in a non‐random pattern with species responding individually along an environmental gradient. Based on a model selection approach involving local and regional predictors, parasite species richness and metacommunity structure correlated with both local abiotic (pH and total dissolved nitrogen) and biotic (non‐host mollusk density, and H. trivolvis biomass) factors, with little support for regional predictors. Overall, this trematode metacommunity most closely followed the predictions from the species sorting or mass effects metacommunity paradigm, in which community diversity is filtered by local site characteristics.     

Factors affecting the helminth community structure of adult collared peccaries in southern Texas

Four species of nematodes (Gongylonema pulchrum, Parabronema pecariae, Texicospirura turki, and Physocephalus sexalatus) and one species of cestode (Moniezia sp.) comprised the helminth fauna of adult collared peccaries (Tayassu tajacu) from the plains in southern Texas. The community structure of the helminth fauna of peccaries from this region was basically dissimilar to that from the more humid Gulf coastal prairies of southern Texas in composition (by the conspicuous absence of certain species) and relative abundance of shared species. The distributions of each of the three common species of helminths (G. pulchrum, T. turki, and P. sexalatus) were overdispersed. The effects of selected habitat variables operating across host subpopulations (delineated by condition and sex) and of the extrinsic variable of season on the dispersion patterns of the three common species of helminths were examined. The hypothesis that heterogeneity within the host population, rather than across the collective host population, is the main factor generating overdispersion in natural parasite populations was not confirmed for the three common species of helminths. Overdispersion in P. sexalatus resulted from seasonal changes across the collective host population, with the greatest abundances occurring during the cool season. Aggregated abundances of G. pulchrum resulted from variation generated across host sex subpopulations, while the dispersion patterns of T. turki appeared to be unaffected by the habitat variables examined in this study.     

A comparative hierarchical analysis of bacterioplankton and biofilm metacommunity structure in an interconnected pond system

It is unknown whether bacterioplankton and biofilm communities are structured by the same ecological processes, and whether they influence each other through continuous dispersal (known as mass effects). Using a hierarchical sampling approach we compared the relative importance of ecological processes structuring the dominant fraction (relative abundance ≥0.1%) of bacterioplankton and biofilm communities from three microhabitats (open water, Nuphar and Phragmites sites) at within‐ and among‐pond scale in a set of 14 interconnected shallow ponds. Our results demonstrate that while bacterioplankton and biofilm communities are highly distinct, a similar hierarchy of ecological processes is acting on them. For both community types, most variation in community composition was determined by pond identity and environmental variables, with no effect of space. The highest β‐diversity within each community type was observed among ponds, while microhabitat type (Nuphar, Phragmites, open water) significantly influenced biofilm communities but not bacterioplankton. Mass effects among bacterioplankton and biofilm communities were not detected, as suggested by the absence of within‐site covariation of biofilm and bacterioplankton communities. Both biofilm and plankton communities were thus highly structured by environmental factors (i.e., species sorting), with among‐lake variation being more important than within‐lake variation, whereas dispersal limitation and mass effects were not observed.     

The role of environmental and spatial processes in structuring native and non‐native fish communities across thousands of lakes

Quantifying the role of spatial patterns is an important goal in ecology to further understand patterns of community composition. We quantified the relative role of environmental conditions and regional spatial patterns that could be produced by environmental filtering and dispersal limitation on fish community composition for thousands of lakes. A database was assembled on fish community composition, lake morphology, water quality, climatic conditions, and hydrological connectivity for 9885 lakes in Ontario, Canada. We utilized a variation partitioning approach in conjunction with Moran's Eigenvector Maps (MEM) and Asymmetric Eigenvector Maps (AEM) to model spatial patterns that could be produced by human‐mediated and natural modes of dispersal. Across 9885 lakes and 100 fish species, environmental factors and spatial structure explained approximately 19% of the variation in fish community composition. Examining the proportional role of spatial structure and environmental conditions revealed that as much as 90% of the explained variation in native species assemblage composition is governed by environmental conditions. Conversely on average, 67% of the explained variation in non‐native assemblage composition can be related to human‐mediated dispersal. This study highlights the importance of including spatial structure and environmental conditions when explaining patterns of community composition to better discriminate between the ecological processes that underlie biogeographical patterns of communities composed of native and non‐native fish species.     

Community structure of helminth parasites of the tuna, Euthynnus affinis, from the Visakhapatnam coast, Bay of Bengal

An analysis is made of the community structure of the helminth parasites of the tuna Euthynnus affinis collected off the coast of Visakhapatnam, Bay of Bengal. The helminth fauna comprised 23 species, consisting of two monogeneans, 18 digeneans, one larval cestode, one nematode and one acanthocephalan. Didymozoids, represented by 14 species, were the dominant members of the parasite spectrum. The parasite assemblage is characterized by species richness as well as a high diversity. Most of the parasites found, the didymozoids in particular, were host specialists, exhibiting a high degree of host specificity. The fauna comprised three core species, four satellite species and many secondary species, with no potential for interaction among them but the degree of predictability is high in that each infracommunity comprised two or three core species superimposed by a few secondary species. Many host factors such as the varied diet, high vagility, long life span and endothermy appear to have contributed to the development of the species rich and diverse parasite communities in the tuna. There was evidence for a decrease in parasite density and an increase in diversity with increase in host size, indicating that host size has a profound influence on the component community structure.     

Speciation and diversification of parasite lineages: an analysis of congeneric parasite species in vertebrates

The evolutionary diversification of living organisms is a central research theme in evolutionary ecology, and yet it remains difficult to infer the action of evolutionary processes from patterns in the distribution of rates of diversification among related taxa. Using data from helminth parasite communities in 76 species of birds and 114 species of mammals, the influence of four factors that may either be associated with or modulate rates of parasite speciation were examined in a comparative analysis. Two measures of the relative number of congeneric parasite species per host species were used as indices of parasite diversification, and related to host body mass, host density, latitude, and whether the host is aquatic or terrestrial. The occurrence of congeneric parasites was not distributed randomly with respect to these factors. Aquatic bird species tended to harbour more congeneric parasites than terrestrial birds. Large-bodied mammal species, or those living at low latitudes, harboured more congeneric parasites than small-bodied mammals, or than those from higher latitudes. Host density had no apparent association with either measures of parasite diversification. These patterns, however, reflect only the present-day distribution of parasite diversification among host taxa, and not the evolutionary processes responsible for diversification, because the apparent effects of the factors investigated disappeared once corrections were made for host phylogeny. This indicates that features other than host body size, host density, latitude, and whether the habitat is terrestrial or aquatic, have been the key driving forces in the diversification of parasitic helminth lineages. This revised version was published online in July 2006 with corrections to the Cover Date.     

Macroinvertebrate community composition and diversity in ephemeral and perennial ponds on unregulated floodplain meadows in the UK

Ponds are common and abundant landscape features in temperate environments, particularly on floodplains where lateral connectivity with riverine systems persists. Despite their widespread occurrence and importance to regional diversity, research on the ecology and hydrology of temperate ephemeral and perennial floodplain ponds lags behind that of other shallow waterbodies. This study examines the aquatic macroinvertebrate diversity of 34 ponds (20 perennial and 14 ephemeral) on two unregulated riverine floodplain meadows in Leicestershire, UK. Perennial ponds supported nearly twice the diversity of ephemeral ponds. Despite frequent inundation of floodwater and connectivity with other floodplain waterbodies, ephemeral ponds supported distinct invertebrate communities when compared to perennial ponds. When the relative importance of physical, chemical, biological and spatial characteristics was examined, physical and chemical characteristics were found to account for more variation in community composition than biological or spatial variables. The results suggest that niche characteristics rather than neutral colonisation processes dominate the structure of invertebrate communities of floodplain ponds. The maintenance of pond networks with varying hydroperiod lengths and environmental characteristics should be encouraged as part of conservation management strategies to provide heterogeneous environmental conditions to support and enhance aquatic biodiversity at a landscape scale.     

Cross-Scale Variation in Biodiversity-Environment Links Illustrated by Coastal Sandflat Communities

Spatial variation in the composition of communities is the product of many biotic and environmental interactions. A neglected factor in the analysis of community distribution patterns is the multi-scale nature of the data, which has implications for understanding ecological processes and the development of conservation and environmental management practice. Drawing on recently established multivariate spatial analyses, we investigate whether including relationships between spatial structure and abiotic variables enable us to better discern patterns of species and communities across scales. Data comprised 1200 macrozoobenthic samples collected over an array of distances (30 cm to 1 km) in three New Zealand harbours, as well as commonly used abiotic variables, such as sediment characteristics and chlorophyll a concentrations, measured at the same scales. Moran’s eigenvector mapping was used to extract spatial scales at which communities were structured. Benthic communities, representing primarily bivalves, polychaetes and crustaceans, were spatially structured at four spatial scales, i.e. >100 m, 50–100 m, 50–15 m, and < 15 m. A broad selection of abiotic variables contributed to the large-scale variation, whereas a more limited set explained part of the fine-scale community structure. Across all scales, less than 30% of the variation in spatial structure was captured by our analysis. The large number of species (48) making up the 10 highest species scores based on redundancy analyses illustrate the variability of species-scale associations. Our results emphasise that abiotic variables and biodiversity are related at all scales investigated and stress the importance of assessing the relationship between environmental variables and the abundance and distribution of biological assemblages across a range of different scales.     

Deconstructing spatial patterns in species composition of ectoparasite communities: the relative contribution of host composition,environmental variables and geography

Aim We determined whether dissimilarity in species composition between parasite communities depends on geographic distance, environmental dissimilarity or host faunal dissimilarity, for different subsets of parasite species with different levels of host specificity. Location Communities of fleas parasitic on small mammals from 28 different regions of the Palaearctic. Method Dissimilarities in both parasite and host species composition were computed between each pair of regions using the Bray–Curtis index. Geographic distances between regions were also calculated, as were measures of environmental dissimilarity consisting of the pairwise Euclidean distances between regions derived from elevation, vegetation and climatic variables. The 136 flea species included in the dataset were divided into highly host‐specific species (using 1–2 host species per region, on average), moderately host‐specific species (2.2–4 hosts per region) and generalist species (>4 hosts per region). The relative influence of geographic distance, host faunal dissimilarity and environmental dissimilarity on dissimilarity of flea species composition among all regions was analysed for the entire set of flea species as well as for the three above subsets using multiple regressions on distance matrices. Results When including all flea species, dissimilarity in flea species composition was affected by all three independent variables, although the pure effect of dissimilarity in host species composition was the strongest. Results were different when the subsets of fleas differing in host specificity were treated separately. In particular, dissimilarity in species composition of highly host‐specific fleas increased solely with environmental dissimilarity, whereas dissimilarity for both moderately specific and non‐specific fleas increased with both geographic distance and dissimilarity in host species composition. Main conclusions Host specificity seems to dictate which of the three factors considered is most likely to affect the dissimilarity between flea communities. Counter‐intuitively, environmental dissimilarity played a key role in determining dissimilarity in species composition of highly host‐specific fleas, possibly because, although their presence in a region relies on the occurrence of particular host species, their abundance is itself mostly determined by climatic conditions. Our results show that deconstructing communities into subsets of species with different traits can make it easier to uncover the mechanisms shaping geographic patterns of diversity.     

Variation in parasite communities and health indices of juvenile Lepomis gibbosus across a gradient of watershed land-use and habitat quality

Parasites of fishes vary in community structure and species abundance in response to environmental conditions and pollutants. As a result, the use of parasites as bioindicators of habitat degradation has been proposed and successfully applied in recent years. Here, helminth parasites of juvenile pumpkinseed Lepomis gibbosus from three streams representing a gradient of watershed development and habitat quality were examined to assess variation in parasite communities. Health assessment indices were also generated for each host to quantify the influence of habitat on the observed health of individuals. A total of 22 parasite taxa were recovered from examined fish, comprising 11 digeneans, 3 cestodes, 2 acanthocephalans, 5 monogeneans and 1 crustacean. In the most disturbed stream, parasite species richness was lowest and total abundance was highest, while parasite abundance was lowest and diversity highest in the least disturbed stream. There was no significant difference in health indices among streams. Analysis of similarities (ANOSIM) and Bray–Curtis dissimilarity in species composition (SIMPER) identified Posthodiplostomum spp. and Actinocleidus sp. as the species driving parasite community dissimilarity. These taxa are relatively easy to identify to genus level and thus could be appropriate for use as indicators of environmental health, where increased abundance would suggest negative changes in habitat quality. However, larger scale study including more streams would be necessary to establish baseline community data before such implementation would be feasible.     

Variance partitioning of deconstructed periphyton communities: does the use of biological traits matter?

The use of species traits offers a promising approach to the understanding of the main processes underlying metacommunity patterns. We analyzed samples of periphytic algae in 30 environments of the Upper Paraná River floodplain in southeastern Brazil, to test the hypotheses that variation in species composition of algal groups with low dispersal abilities would be mainly explained by spatial variables; on the other hand, algal groups with higher dispersal abilities would be better explained by environmental variables. The variation in community structure was mainly correlated with environmental variables. This result is in line with a growing body of evidence indicating a predominant role of species-sorting processes. The more-refined prediction that the spatial variables would gradually become more important across a gradient of adherence or size was, however, not supported by our analyses. Also, the large unexplained variation suggested that these periphytic communities were assembled by idiosyncratic events, or that other variables that are often neglected in studies of aquatic metacommunities needed to be included.     

Spatial heterogeneity in the parasite communities of creek chub (Semotilus atromaculatus) in southeastern Nebraska

The intestinal helminth communities of creek chub (Semotilus atromaculatus) were studied in the streams of southeastern Nebraska to characterize spatial variation, to determine whether drainages act as regional species pools, and to examine the spatial patterning of individual parasite species within and among drainages. Creek chub were sampled in the summer of 2003 and the spring of 2004 at each of 12 sites distributed evenly among 3 drainages in the Big Nemaha River watershed. Four intestinal helminths were recovered: Allocreadium lobatum, Proteocephalus sp., Rhabdochona canadensis, and Paulisentis missouriensis. Host size had little or no effect on the composition of the parasite communities of creek chub, either among individual fish or among samples. In contrast, drainage and sample date explained 82% of the variation in mean infracommunity species richness among samples, and 62% of the variation in mean infracommunity abundance among samples. Drainage differences were determined by the distributions of P. missouriensis and R. canadensis; whereas, A. lobatum and Proteocephalus sp. were more uniformly distributed among drainages. Each drainage was characterized by a unique pattern of species diversity at infracommunity, component community, and drainage levels of organization.     

Evidence of stochasticity driving anuran metacommunity structure in the Pantanal wetlands

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Integrating environmental and spatial processes in ecological community dynamics

The processes controlling the abundances of species across multiple sites form the cornerstone of modern ecology. In these metacommunities, the relative importance of local environmental and regional spatial processes is currently hotly debated, especially in terms of the validity of neutral model. I collected 158 published data sets with information on community structure, environmental and spatial variables. I showed that approximately 50% of the variation in community composition is explained by both environmental and spatial variables. The majority of the data sets were structured by species-sorting dynamics (SS), followed by a combination of SS and mass-effect dynamics. While neutral processes were the only structuring process in 8% of the collected natural communities, disregarding neutral dispersal processes would result in missing important patterns in 37% of the studied communities. Moreover, metacommunity characteristics such as dispersal type, habitat type and spatial scale predicted part of the detected variation in metacommunity structure.