Spatial patterns of association at local and regional scales in coastal sand dune communities

Questions: Are positive understorey‐dominant associations important in physically severe dune communities and does the strength of positive associations vary with disturbance at the local scale and with stress at the regional scale? Do associational patterns observed at the neighbourhood scale predict diversity at higher scales? Location: Coastal sand dunes, Aquitaine (France). Methods: Associational patterns with five dominant species were recorded along a local gradient of disturbance and a 240‐km long regional gradient. Density, richness, cover and variance ratio of understorey species were recorded in quadrats located in dominant and in open areas. Spatial pattern of dominant plant species was recorded using a distance‐based method. Results: Positive understorey‐dominant associations were most frequent at both regional and local scale, although negative associations with understorey species were observed for one of the five dominants. At the regional scale, there was a shift in the magnitude of spatial associations, with higher positive associations in the most stressful sites, whereas spatial associations where not affected by the local disturbance gradient. Positive associations were not related to the size of the dominants but rather influenced by the identity of the dominant species. Conclusions: Our study highlights the potential crucial role of facilitation together with the importance of turnover of the dominants in explaining large‐scale variation in diversity. However, because positive associations may also be attributed to environmental heterogeneity or co‐occurrence of microhabitat preferences of species, experiments are needed to fully assess the relative importance of facilitation versus other drivers of community diversity.     

Urbanization negatively impacts frog diversity at continental,regional, and local scales

Urban environments are novel ecosystems, with increased chemical, sound, and light pollution differentially impacting many animals. Understanding the impacts of urban environments on biodiversity is the first step to understanding how to best mitigate biodiversity losses in an increasingly urbanizing world. Analyses with broad geographic and taxonomic coverage can offer critical context for informing urban biodiversity conservation. But such studies are currently lacking, especially for under-studied, but likely highly impacted, taxa such as frogs. Our objective was to document frog diversity in relation to urban environments at continental, regional, and local scales. We used FrogID data, ⁠an opportunistic citizen science dataset generated by volunteers recording calling frogs using a smartphone and validated by experts ⁠throughout continental Australia, to calculate species richness, Shannon diversity, and phylogenetic diversity of frogs in urban and non-urban areas, as well as along a continuous urbanization gradient. The overall species richness of frogs was, on average, 57% less in urban than non-urban areas across six ecoregions. Further, we found significantly lower frog diversity in urban environments compared with non-urban environments across the country, with an average reduction of 59% species richness, 86% Shannon diversity, and 72% phylogenetic diversity. We also found evidence for a steady decrease in frog diversity along an urbanization gradient, with no obvious thresholds. Our results highlight the negative impacts of urbanization, ⁠at a continental scale, ⁠on frog diversity, and clearly highlight the necessity to consider frog diversity in future urban land development decisions.     

A novel approach to understanding bird communities using informed diversity estimates at local and regional scales in northern California and southern Oregon

Assessment and preservation of biodiversity has been a central theme of conservation biology since the discipline's inception. However, when diversity estimates are based purely on measures of presence–absence, or even abundance, they do not directly assess in what way focal habitats support the life history needs of individual species making up biological communities. Here, we move beyond naïve measures of occurrence and introduce the concept of “informed diversity” indices which scale estimates of avian species richness and community assemblage by two critical phases of their life cycle: breeding and molt. We tested the validity of the “informed diversity” concept using bird capture data from multiple locations in northern California and southern Oregon to examine patterns of species richness among breeding, molting, and naïve (based solely on occurrence) bird communities at the landscape and local scales using linear regression, community similarity indices, and a Detrended Correspondence Analysis (DCA). At the landscape scale, we found a striking pattern of increased species richness for breeding, molting, and naïve bird communities further inland and at higher elevations throughout the study area. At the local scale, we found that some sites with species‐rich naïve communities were in fact species‐poor when informed by breeding status, indicating that naïve richness may mask more biologically meaningful patterns of diversity. We suggest that land managers use informed diversity estimates instead of naïve measures of diversity to identify ecologically valuable wildlife habitat.     

Exploring macroinvertebrate species distributions at regional and local scales across a sandy beach geographic continuum

Exposed sandy beaches are highly dynamic ecosystems where macroinvertebrate species cope with extremely variable environmental conditions. The majority of the beach ecology studies present exposed beaches as physically dominated ecosystems where abiotic factors largely determine the structure and distribution of macrobenthic communities. However, beach species patterns at different scales can be modified by the interaction between different environmental variables, including biotic interactions. In this study, we examined the role of different environmental variables for describing the regional and local scale distributions of common macrobenthic species across 39 beaches along the North coast of Spain. The analyses were carried out using boosted regression trees, a relatively new technique from the field of machine learning. Our study showed that the macroinvertebrate community on exposed beaches is not structured by a single physical factor, but instead by a complex set of drivers including the biotic compound. Thus, at a regional scale the macrobenthic community, in terms of number of species and abundance, was mainly explained by surrogates of food availability, such as chlorophyll a. The results also revealed that the local scale is a feasible way to construct general predictive species-environmental models, since relationships derived from different beaches showed similar responses for most of the species. However, additional information on aspects of beach species distribution can be obtained with large scale models. This study showed that species-environmental models should be validated against changes in spatial extent, and also illustrates the utility of BRTs as a powerful analysis tool for ecology data insight.     

Spider diversity in cereal fields: comparing factors at local, landscape and regional scales

Aim Factors acting at various scales may affect biodiversity, demanding analyses at multiple spatial scales in order to understand how community richness is determined. Here, we adopted a hierarchical approach to test the contribution of region, landscape heterogeneity, local management (organic vs. conventional) and location within field (edge vs. centre) to the species richness and abundance of spiders in cereals. Location Three regions of western and central Germany: Leine Bergland, Soester Boerde, and Lahn‐Dill Bergland. Methods Forty‐two paired organic and conventional winter wheat fields were compared. Field pairs were located in areas ranging from structurally simple to structurally complex landscapes. In May and June 2003, spiders were sampled using pitfall traps. Linear mixed models were used to determine the relationship of spider diversity and abundance with regional spatial factors and landscape heterogeneity within a 500‐m radius, as well as with local management and within‐field location. Results Within‐field location of the traps and landscape heterogeneity were the best predictors of species richness: more species were found in field edges and in heterogeneous landscapes. Region and local management had no effect on species richness. Activity density was higher in field edges and differed among regions. Main conclusions The diversity of farmland spiders was influenced by differences at two of the spatial scales (edge vs. centre, simple vs. complex landscapes), but not at the two others (field management, region), emphasizing the importance of analyses at a variety of spatial scales for an adequate explanation of patterns in biodiversity. Our study suggests that promoting heterogeneity in land use at landscape scales is one of the keys to promoting spider diversity in agroecosystems.     

Native and alien plant species richness in relation to spatial heterogeneity on a regional scale in Germany

Aim The aim of our study was to reveal relationships between richness patterns of native vs. alien plant species and spatial heterogeneity across varying landscape patterns at a regional scale. Location The study was carried out in the administrative district of Dessau (Germany), covering around 4000 km². Methods Data on plant distribution of the German vascular flora available in grid cells covering 5′ longitude and 3′ latitude (c. 32 km²) were divided into three status groups: native plants, archaeophytes (pre 1500 AD aliens) and neophytes (post 1500 AD aliens). Land use and abiotic data layers were intersected with 125 grid cells comprising the selected area. Using novel landscape ecological methods, we calculated 38 indices of landscape composition and configuration for each grid cell. Principal components analysis (PCA) with a set of 29 selected, low correlated landscape indices was followed by multiple linear regression analysis. Results PCA reduced 29 indices to eight principal components (PCs) that explained 80% cumulative variance. Multiple linear regression analysis was highly significant and explained 41% to 60% variance in plant species distribution (adjusted R²) with three significant PCs (tested for spatial autocorrelation) expressing moderate to high disturbance levels and high spatial heterogeneity. Comparing the significance of the PCs for the species groups, native plant species richness is most strongly associated with riverine ecosystems, followed by urban ecosystems, and then small‐scale rural ecosystems. Archaeophyte and neophyte richness are most strongly associated with urban ecosystems, followed by small‐scale rural ecosystems and riverine ecosystems for archaeophytes, and riverine ecosystems and small‐scale rural ecosystems for neophytes. Main conclusions Our overall results suggest that species richness of native and alien plants increases with moderate levels of natural and/or anthropogenic disturbances, coupled with high levels of habitat and structural heterogeneity in urban, riverine, and small‐scale rural ecosystems. Despite differences in the order of relevance of PCs for the three plant groups, we conclude that at the regional scale species richness patterns of native plants as well as alien plants are promoted by similar factors.     

Trait-based filtering from the regional species pool into local grassland communities

Aims For plants to establish in a local community from a pool of possible colonizers from the region, it must pass through a series of filters. Which of the filters is most important in this process has been much debated. In this study, we explored how species are filtered from the regional species pool into local communities. The aim was to determine if differences in species abundance and functional traits could explain which species from the regional species pool establish at the local scale and if the filtering differed between grassland communities.Methods This study took place in a cultivated landscape in southeastern Sweden. We estimated plant species abundance in 12 ex-arable field sites and 8 adjacent seminatural grassland sites and in a 100-m radius around the center of each site. We used Monte Carlo simulations to examine if species abundance and functional traits (height, seed mass, clonal abilities, specific leaf area and dispersal method) controlled the filtering of species from the regional pool into local communities.Important findings On average, only 28% of species found in the regional pool established in the ex-arable field sites and 45% in the seminatural grassland sites, indicating that the size of the regional species pool was not limiting local richness. For both grassland types, species abundance in the regional pool was positively correlated with species occurrence at the local scale. We found evidence for both species interaction filtering and dispersal limitation influencing the local assembly. Both local and regional processes were thus influencing the filtering of species from the regional species pool into local communities. In addition, the age of the communities influenced species filtering, indicating that community assembly and the importance of different filters in that process change over succession.     

Extinction,turnover and species diversity in an experimentally fragmented California annual grassland

Summary Many natural populations are subdivided among partially isolated habitat patches, but the influence of habitat patchiness per se on species immigration, extinction, and the resulting patterns of species diversity, has received virtually no experimental study. In an experiment designed to test the effects of habitat subdivision on local community structure, we compare the diversity and annual turnover of flowering plant species in 3 treatments of the same total area, but subdivided to different degrees. We experimentally fragmented a California winter annual grassland into isolated plots, two of 32 m², eight of 8 m², and 32 of 2 m², each treatment representing a combined area of 64 m². Insularization of the experimental habitat fragments is provided by grazing sheep. The effects of plot area on species diversity, extinction, and turnover are consistent with the MacArthur-Wilson model. Species richness increases with the degree of habitat subdivision. Extinction, immigration, and turnover, however, are relatively independent of the degree of subdivision. These experimental results contrast with predictions that habitat subdivision necessarily results in greater rates of extinction accompanied by reduced species diversity.     

Carbon fluxes and species diversity in grazed and fenced typical steppe grassland of Inner Mongolia,China

Aim Grasslands are dominant vegetation of China, support outstanding biodiversity and sequester bulk amount of atmospheric CO₂. These grasslands are highly degraded and fragmented due to remarkable anthropogenic and grazing loads. Chinese Government has made great attempt to restore by grazing exclusion. The relations of carbon fluxes with species composition and diversity in the communities sensitive to grazing by large herbivores are needed to be analysed under the global climate change scenario. The objective of present study was to comprehend the effects of grazing and fencing on the ecosystem structure and function of the typical steppe grassland.     

Spatial and temporal variability in California annual grassland: results from a long-term study

Abstract. We present data from the first 11 years of a longterm study of the dynamics of an annual grassland on serpentine soil in Jasper Ridge Biological Preserve, Northern California. Annual rainfall amounts and distributions varied greatly over the period 1982-1993, as did the amount and distribution of gopher disturbance. Temporal variation in gopher disturbance showed no relationship with rainfall, but spatial variation in disturbance frequency was related to soil depth. The disturbance regime experienced by the grassland is complex, both spatially and temporally, and most of the area is disturbed at least once every 3-5 years. Plant species abundances showed a variety of responses to climate variation and disturbance. Abundances of individual species in any given year could not be linked directly to rainfall amount due to hysteresis effects and other interactions. The grassland composition changed markedly over the study. Exclusion of gophers suggested that changing abundances of several species were linked to gopher disturbance. In particular, perennial species' abundances increased greatly in the years following exclosure, but then subsequently declined. Data on plant densities on gopher mounds disturbed at different times of year and in different years indicate that the local species composition remains distinct for a number of years following disturbance. Disturbance history is hence a major factor controlling local community variation. Changing species importances, a complex disturbance regime and the importance of disturbance history make prediction and modelling of this system difficult. It is suggested that the same is probably true for many plant communities, and that long-term studies must be an essential part of ecological research programs. This study illustrates the practical problems inherent in maintaining long-term field experiments and in analyzing complex time series data which suffer from inadvertent deviations from the original experimental design.     

Effects of competition, predation, and dispersal on species richness at local and regional scales

This study explores the consequences of predator-mediated coexistence among competitors for patterns of incidence and diversity at local and regional scales. We develop a model that draws on elements of metapopulation models of competitors and food chains by allowing competitors to coexist locally in the presence of predators but not in their absence. The model predicts that predators promote regional coexistence by greatly expanding the range of conditions under which two competitors persist at equilibrium. Predators could have positive or negative effects on mean local diversity within the region depending on their dispersal rates, those of the prey, and their effects on prey extinction rates. The presence of predators increased the abundance of inferior competitors, thereby expanding the conditions for positive relationships between local and regional diversity. The model also predicted positive correlations between local diversity of predators and prey. These predictions were supported by patterns of phytoplankton, zooplankton, and fish species richness among lakes. The model may help to resolve the apparent contrast between linear patterns of local and regional richness and experimental evidence for strong invasion resistance and rapid dispersal in zooplankton.     

Biodiversity and ecosystem functioning at local and regional spatial scales

Local niche complementarity among species (the partitioning of species based upon niche differentiation) is predicted to affect local ecosystem functioning positively. However, recent theory predicts that greater local diversity may hinder local ecosystem functioning when diversity is enhanced through source–sink dynamics. We suggest community assembly as a way to incorporate both the local and regional processes that determine biodiversity and its consequent effects on ecosystem functioning. From this, we propose a hump-shaped relationship between diversity and ecosystem functioning at local scales, but a linear increase of functioning with diversity at regional scales due to regional complementarity.     

The relationship between productivity and multiple aspects of biodiversity in six grassland communities

Biodiversity is a multifaceted concept but most studies examining the association between the biodiversity of a community and its productivity focus only on species richness. Consequently, studies are needed to examine how other facets of biodiversity vary with productivity if we want to have a better understanding of the distribution of biodiversity across our planet. We evaluated how a number of biodiversity measures (species richness, evenness, dominance, rarity, Simpson’s diversity, and Shannon–Weiner diversity) varied across natural productivity gradients at 6 grassland sites in the continental US. Variation in productivity did not account for a substantial amount of variation in any measure of biodiversity at small spatial scales (≈1 m²) at most sites. When productivity accounted for substantial variation in biodiversity, different measures of biodiversity responded to productivity in different ways. For example, dominance changed in a U-shaped fashion along a productivity gradient whereas richness increased in an asymptotic fashion. Consequently, diversity indices, which account for both species richness and evenness, varied in a hump-shaped fashion along the productivity gradient. Our results highlight that an exclusive focus on the association between species richness and productivity provides an incomplete picture of how a community’s biodiversity is related to its functioning.     

Vegetation responses to landscape structure at multiple scales across a Northern Wisconsin,USA, pine barrens landscape

Increasing awareness of the importance of scale and landscape structure to landscape processes and concern about loss of biodiversity has resulted in efforts to understand patterns of biodiversity across multiple scales. We examined plant species distributions and their relationships to landscape structure at varying spatial scales across a pine barrens landscape in northern Wisconsin, U.S.A. We recorded plant species cover in 1×1 m plots every 5 m along a 3575 m transect, along with variables describing macro- and micro-landscape structure. A total of 139 understory plant species were recorded. The distributions of many species appeared to be strongly associated with landscape structural features, such as distinct management patches and roads. TWINSPAN and detrended correspondence analysis (DCA) identified three groups of species that overlapped extensively in the ordination, possibly reflecting the relatively homogeneous nature of disturbance in the pine barrens landscape. Distribution of understory plants did not reflect all of the patch types we identified along the transect; plot ordination and classification resulted in three to five plot groups that differed in niche breadth. Wavelet transforms showed varying relationships between landscape features and plant diversity indices (Shannon–Weiner, Simpson's Dominance) at different resolutions. Wavelet variances indicated that patterns of Shannon diversity were dominated by coarse resolutions ranging from 900–1500 m, which may have been related to topography. Patterns of Simpson's Dominance were dominated by 700 m resolution, possibly associated with canopy cover. However, a strong correspondence between overstory patch type and diversity was found for several patch types at ranges of scales that varied by patch type. Effects of linear features such as roads were apparent in the wavelet transforms at resolutions of about 5–1000 m, suggesting roads may have an important impact on plant diversity at landscape scales. At broad scales, landscape context appeared more important to diversity than individual patches, suggesting that changes in structure at fine resolutions could alter overall diversity characteristics of the landscape. Therefore, a hierarchical perspective is necessary to recognize potential large-scale change resulting from small-scale activities.     

Variation in plant responses to neighbors at local and regional scales

Differences in the responses of plant species to neighbors may determine their distribution among contrasting environments, but no studies have compared variation in competitive or facilitative abilities both within and among environments. We determined whether the responses of plants to interspecific interactions varied at large scales (between environments) and small scales (among sites within an environment) across a tree line. We separated the effects of above- and belowground interactions on seedlings of grasses and trees grown in prairie or forest using vegetation removals at several sites. Species interactions generally had no significant effect on transplant survival. Competition reduced seedling growth by about 33%-89% in both prairie and forest environments. Despite the strong suppression of growth by neighbors, environment and species effects contributed more to variation in transplant performance than did neighbor removals. Responses to neighbors varied among transplant species but generally did not vary significantly between environments or among sites. With vegetation removed, grasses grew significantly faster in prairie and trees grew faster in forest. Thus, in the absence of neighbors, species showed distinct preferences for the environment in which they are most abundant. In summary, the responses of grasses and woody species to neighbors did not vary significantly at either large (between environments) or small (among sites) scales. These results suggest that species responses to interspecific interactions do not vary strongly with environment or smaller-scale site effects.     

Different factors affect the local distribution,persistence and spread of alien tree species in floodplain forests

Floodplain forests are characterized by high and increasing levels of invasions by plant species, but the factors that drive their spread are insufficiently understood. Using data from 708 plots surveyed twice (1998, 2008) supplemented with further data (management, stand age, distance to dispersal corridors, type of ecosystems invaded) we analyzed the factors which shape the local distribution, growth, persistence and spread of three invasive alien tree species (Acer negundo, Ailanthus altissima, Robinia pseudoacacia) in the National Park Donau-Auen in Austria. Using Generalized Linear Models (GLMs), we found that the distribution of the study species per plot is contingent on stand age (R. pseudoacacia), type of floodplain forest (A. negundo, A. altissima) and distance to the next water body (A. negundo). For all study species, colonization of new plots between both surveys is driven by short distance spread from already established invasion foci. Moreover, recipient habitats (softwood vs. hardwood floodplain forests) modify invasion success in species-specific ways. The probability of occurrence and colonization of plots located in softwood floodplain forests is higher for A. negundo, whereas A. altissima more frequently colonizes hardwood floodplain forests. Persistence of R. pseudoacacia decreases with stand age, whereas its growth rate is significantly higher in plots located in zones where management is allowed than in those which are not managed. Persistence and growth of the other two study species were not related to any explanatory variables analyzed. The on-going spread of the study species in the National Park Donau-Auen suggests that their local distribution is in disequilibrium, i.e. not all suitable habitat patches have yet been colonized. This implies that increased management efforts are necessary to reverse the spread of alien tree species in the study area and to maintain the high conservation value of this iconic area which protects one of the largest floodplain forests in Europe.     

Plant species richness and spatial organization at different small scales in western Mediterranean landscapes

The relationship between plant species richness and the space organization of the community at different small scales was studied. The study was based on 51 sites distributed along a belt from Central Spain to Portugal. Each site was analyzed with a transect cutting across the boundary between two neighboring patches of shrubland and grassland. Local spatial organization of vegetation was analyzed at different levels of detail and each transect was divided into successively smaller portions. The first division coincides with a physiognomic perception of the site in two patches (shrubland and grassland). The average spatial niche width of the species was used to calculate the spatial organization of the vegetation of each division in each site. The correlation between species richness and spatial organization depended on the block size under consideration. A physiognomic criterion, sectorizing the sites into patches of shrubland and grassland, determines noteworthy floristic changes but does not enable us to express satisfactorily the variability in plant richness. In order to account for this variation, other factors must be taken into account which act at a more detailed small-scale and which determine the internal variability of these patches. In the case studied, the species richness of the sites increases along with an increase in the percentage of species whose occupation of the space is relatively restricted within the site. Many of these species are, however, frequent within the whole of the territory studied. The results highlight the importance of the level of local scale at which the factors influencing occupation of the space, and consequentially, plant richness, preferentially act. This circumstance ought to be taken into consideration in strategies for the conservation of biological diversity, and based on the delimitation of protected spaces with criteria frequently linked to the physiognomy of the vegetation.Nomenclature: Follows T.G. Tutin et al. 1964-1980. Flora Europaea. Cambridge University Press, Cambridge     

Rabbit grazing and species diversity in a dune area

Summary The relation between density of rabbit populations and plant species diversity is discussed. In the dunes of the Dutch Frisian Island of Schiermonnikoog, the rabbit grazing pressure has been quantified on the basis of traces of recent grazing activity. Moderate grazing turned out to bring about maximal species richness. Current changes in grazing pressure (either decrease or increase) can result in plant species diversity that is lower than in other places with equal but stabilized grazing pressure.Nomenclature follows Heukels-van Ooststroom, Flora van Nederland, 18e druk, 1975.The work has been done by Mrs Ada Zeevalking-van Y peren and Mr Hein Zeevalking as part of their study for a doctoral degree.     

Native and introduced fish species richness in French lakes: local and regional influences

Aim To analyse the importance of local and regional influences on the patterns of species richness in natural and man‐made lakes and to infer the impacts of human‐mediated introductions on these patterns. Location France. Methods Species occurrence data were gathered for 25 natural and 51 man‐made lakes. Analysis is based on regression models of local richness against their related regional richness and lake environmental variables. Results Local native richness was mostly controlled by the regional richness. Conversely, local total richness was mainly explained by local variables. These statements apply to both natural and man‐made lakes. Lacustrine systems displayed weak resistance to invaders. Main conclusions Species introductions have apparently contributed to saturate fish communities in these systems even if no clear negative effect on the survival of native species (i.e. species extinction) is detectable so far.