Showing papers in "Global Ecology and Biogeography in 2007"

Landscape modification and habitat fragmentation: a synthesis

Abstract: Landscape modification and habitat fragmentation are key drivers of global species loss. Their effects may be understood by focusing on: (1) individual species and the processes threatening them, and (2) human-perceived landscape patterns and their correlation with species and assemblages. Individual species may decline as a result of interacting exogenous and endogenous threats, including habitat loss, habitat degradation, habitat isolation, changes in the biology, behaviour, and interactions of species, as well as additional, stochastic threats. Human-perceived landscape patterns that are frequently correlated with species assemblages include the amount and structure of native vegetation, the prevalence of anthropogenic edges, the degree of landscape connectivity, and the structure and heterogeneity of modified areas. Extinction cascades are particularly likely to occur in landscapes with low native vegetation cover, low landscape connectivity, degraded native vegetation and intensive land use in modified areas, especially if keystone species or entire functional groups of species are lost. This review (1) demonstrates that species-oriented and pattern-oriented approaches to understanding the ecology of modified landscapes are highly complementary, (2) clarifies the links between a wide range of interconnected themes, and (3) provides clear and consistent terminology. Tangible research and management priorities are outlined that are likely to benefit the conservation of native species in modified landscapes around the world.

The importance of biotic interactions for modelling species distributions under climate change

Abstract: Aim There is a debate as to whether biotic interactions exert a dominant role in governing species distributions at macroecological scales. The prevailing idea is that climate is the key limiting factor; thus models that use present-day climate‐species range relationships are expected to provide reasonable means to quantify the impacts of climate change on species distributions. However, there is little empirical evidence that biotic interactions would not constrain species distributions at macroecological scales. We examine this idea, for the first time, and provide tests for two null hypotheses: ( H 0 1) ‐ biotic interactions do not exert a significant role in explaining current distributions of a particular species of butterfly (clouded Apollo, Parnassius mnemosyne ) in Europe; and ( H 0 2) ‐ biotic interactions do not exert a significant role in predictions of altered species’ ranges under climate change.

Spatial autocorrelation and the selection of simultaneous autoregressive models

Abstract: Aim Spatial autocorrelation is a frequent phenomenon in ecological data and can affect estimates of model coefficients and inference from statistical models. Here, we test the performance of three different simultaneous autoregressive (SAR) model types (spatial error = SAR err , lagged = SAR lag and mixed = SAR mix ) and common ordinary least squares (OLS) regression when accounting for spatial autocorrelation in species distribution data using four artificial data sets with known (but different) spatial autocorrelation structures. Methods We evaluate the performance of SAR models by examining spatial patterns in model residuals (with correlograms and residual maps), by comparing model parameter estimates with true values, and by assessing their type I error control with calibration curves. We calculate a total of 3240 SAR models and illustrate how the best models [in terms of minimum residual spatial autocorrelation (minRSA), maximum model fit ( R 2 ), or Akaike information criterion (AIC)] can be identified using model selection procedures. Results Our study shows that the performance of SAR models depends on model specification (i.e. model type, neighbourhood distance, coding styles of spatial weights matrices) and on the kind of spatial autocorrelation present. SAR model parameter estimates might not be more precise than those from OLS regressions in all cases. SAR err models were the most reliable SAR models and performed well in all cases (independent of the kind of spatial autocorrelation induced and whether models were selected by minRSA, R 2 or AIC), whereas OLS, SAR lag and SAR mix models showed weak type I error control and/or unpredictable biases in parameter estimates. Main conclusions SAR err models are recommended for use when dealing with spatially autocorrelated species distribution data. SAR lag and SAR mix might not always give better estimates of model coefficients than OLS, and can thus generate bias. Other spatial modelling techniques should be assessed comprehensively to test their predictive performance and accuracy for biogeographical and macroecological research.

Caribbean mangroves adjust to rising sea level through biotic controls on change in soil elevation

Abstract: Aim The long-term stability of coastal ecosystems such as mangroves and salt marshes depends upon the maintenance of soil elevations within the intertidal habitat as sea level changes. We examined the rates and processes of peat formation by mangroves of the Caribbean Region to better understand biological controls on habitat stability. Location Mangrove-dominated islands on the Caribbean coasts of Belize, Honduras and Panama were selected as study sites. Methods Biological processes controlling mangrove peat formation were manipulated (in Belize) by the addition of nutrients (nitrogen or phosphorus) to Rhizophora mangle (red mangrove), and the effects on the dynamics of soil elevation were determined over a 3-year period using rod surface elevation tables (RSET) and marker horizons. Peat composition and geological accretion rates were determined at all sites using radiocarbon-dated cores. Results The addition of nutrients to mangroves caused significant changes in rates of mangrove root accumulation, which influenced both the rate and direction of change in elevation. Areas with low root input lost elevation and those with high rates gained elevation. These findings were consistent with peat analyses at multiple Caribbean sites showing that deposits (up to 10 m in depth) were composed primarily of mangrove root matter. Comparison of radiocarbon-dated cores at the study sites with a sea-level curve for the western Atlantic indicated a tight coupling between peat building in Caribbean mangroves and sea-level rise over the Holocene. Main conclusions Mangroves common to the Caribbean region have adjusted to changing sea level mainly through subsurface accumulation of refractory mangrove roots. Without root and other organic inputs, submergence of these tidal forests is inevitable due to peat decomposition, physical compaction and eustatic sea-level rise. These findings have relevance for predicting the effects of sea-level rise and biophysical processes on tropical mangrove ecosystems.

Effects of incorporating spatial autocorrelation into the analysis of species distribution data

Abstract: Aim Spatial autocorrelation (SAC) in data, i.e. the higher similarity of closer samples, is a common phenomenon in ecology. SAC is starting to be considered in the analysis of species distribution data, and over the last 10 years several studies have incorporated SAC into statistical models (here termed ‘spatial models’). Here, I address the question of whether incorporating SAC affects estimates of model coefficients and inference from statistical models. Methods I review ecological studies that compare spatial and non-spatial models. Results In all cases coefficient estimates for environmental correlates of species distributions were affected by SAC, leading to a mis-estimation of on average c. 25%. Model fit was also improved by incorporating SAC. Main conclusions These biased estimates and incorrect model specifications have implications for predicting species occurrences under changing environmental conditions. Spatial models are therefore required to estimate correctly the effects of environmental drivers on species present distributions, for a statistically unbiased identification of the drivers of distribution, and hence for more accurate forecasts of future distributions.

Biotic interactions improve prediction of boreal bird distributions at macro‐scales

Abstract: Aim: The role of biotic interactions in influencing species distributions at macro-scales remains poorly understood. Here we test whether predictions of distributions for four boreal owl species at two macro-scales (10 × 10 km and 40 × 40 km grid resolutions) are improved by incorporating interactions with woodpeckers into climate envelope models. Location: Finland, northern Europe. Methods: Distribution data for fourowl and six woodpecker species, along with data for six land cover and three climatic variables, were collated from 2861 10 × 10 km grid cells. Generalized additive models were calibrated using a 50% random sample of the species data from western Finland, and by repeating this procedure 20 times for each of the four owl species. Models were fitted using three sets of explanatory variables: (1) climate only; (2) climate and land cover; and (3) climate, land cover and two woodpecker interaction variables. Models were evaluated using three approaches: (1) examination of explained deviance; (2) four-fold cross-validation using the model calibration data; and (3) comparison of predicted and observed values for independent grid cells in eastern Finland. The model accuracy for approaches (2) and (3) was measured using the area under the curve of a receiver operating characteristic plot. Results: At 10-km resolution, inclusion of the distribution of woodpeckers as a predictor variable significantly improved the explanatory power, cross-validation statistics and the predictive accuracy of the models. Inclusion of land cover led to similar improvements at 10-km resolution, although these improvements were less apparent at 40-km resolution for both land cover and biotic interactions. Main conclusions: Predictions of species distributions at macro-scales may be significantly improved by incorporating biotic interactions and land cover variables into models. Our results are important for models used to predict the impacts of climate change, and emphasize the need for comprehensive evaluation of the reliability of species-climate impact models. © 2007 The AuthorsGlobal Ecology and Biogeography, Journal compilation © 2007 Blackwell Publishing Ltd.

Altered stream-flow regimes and invasive plant species : the Tamarix case

Abstract: Aim To test the hypothesis that anthropogenic alteration of stream-flow regimes is a key driver of compositional shifts from native to introduced riparian plant species. Location The arid south-western United States; 24 river reaches in the Gila and Lower Colorado drainage basins of Arizona. Methods We compared the abundance of three dominant woody riparian taxa (native Populus fremontii and Salix gooddingii , and introduced Tamarix ) between river reaches that varied in stream-flow permanence (perennial vs. intermittent), presence or absence of an upstream flow-regulating dam, and presence or absence of municipal effluent as a stream water source. Results Populus and Salix were the dominant pioneer trees along the reaches with perennial flow and a natural flood regime. In contrast, Tamarix had high abundance (patch area and basal area) along reaches with intermittent stream flows (caused by natural and cultural factors), as well as those with dam-regulated flows. Main conclusions Stream-flow regimes are strong determinants of riparian vegetation structure, and hydrological alterations can drive dominance shifts to introduced species that have an adaptive suite of traits. Deep alluvial groundwater on intermittent rivers favours the deep-rooted, stress-adapted Tamarix over the shallower-rooted and more competitive Populus and Salix . On flow-regulated rivers, shifts in flood timing favour the reproductively opportunistic Tamarix over Populus and Salix , both of which have narrow germination windows . The prevailing hydrological conditions thus favour a new dominant pioneer species in the riparian corridors of the American Southwest. These results reaffirm the importance of reinstating stream-flow regimes (inclusive of groundwater flows) for re-establishing the native pioneer trees as the dominant forest type.

Environmental data sets matter in ecological niche modelling: an example with Solenopsis invicta and Solenopsis richteri.

Abstract: Aim In response to a recent paper suggesting the failure of ecological niche models to predict between native and introduced distributional areas of fire ants (Solenopsis invicta), we sought to assess methodological causes of this failure. Location Ecological niche models were developed on the species’ native distributional area in South America, and projected globally. Methods We developed ecological niche models based on six different environmental data sets, and compared their respective abilities to anticipate the North American invasive distributional area of the species. Results We show that models based on the ‘bioclimatic variables’ of the WorldClim data set indeed fail to predict the full invasive potential of the species, but that models based on four other data sets could predict this potential correctly. Main conclusions The difference in predictive abilities appears to centre on the complexity of the environmental variables involved. These results emphasize important influences of environmental data sets on the generality and ability of ecological niche models to anticipate novel phenomena, and offer a simpler explanation for the lack of predictive ability among native and invaded distributional areas than that of niche shifts.

Does size matter for dispersal distance

Abstract: Aim The aim of this study is to answer the questions: (1) do small organisms disperse farther than large, or vice versa; and (2) does the observed pattern differ for passive and active dispersers? These questions are central to several themes in biogeography (including microbial biogeography), macroecology, metacommunity ecology and conservation biology. Location The meta-analysis was conducted using published data collected worldwide. Methods We collected and analysed 795 data values in the peer-reviewed literature for direct observations of both maximal dispersal distance and mass of the dispersing organisms (e.g. seeds, not trees). Analysed taxa ranged in size from bacteria to whales. We applied macroecology analyses based on null models (using Monte Carlo randomizations) to test patterns relative to specific hypotheses. Results Collected dispersal distance and mass data spanned 9 and 21 orders of magnitude, respectively. Active dispersers dispersed significantly farther ( P < 0.001) and were significantly greater in mass ( P < 0.001) than passive dispersers. Overall, size matters: larger active dispersers attained greater maximum observed dispersal distances than smaller active dispersers. In contrast, passive-disperser distances were random with respect to propagule mass, but not uniformly random, in part due to sparse data available for tiny propagules. Conclusions Size is important to maximal dispersal distance for active dispersers, but not for passive dispersers. Claims that microbes disperse widely cannot be tested by current data based on direct observations of dispersal: indirect approaches will need to be applied. Distance‐mass relationships should contribute to a resolution of neutral and niche-based metacommunity theories by helping scale expectations for dispersal limitation. Also, distance‐mass relationships should inform analyses of latitudinal species richness and conservation biology topics such as fragmentation, umbrella species and taxonomic homogenization.

Small fish, big fish, red fish, blue fish: size-biased extinction risk of the world's freshwater and marine fishes.

Abstract: Aim In light of the current biodiversity crisis, there is a need to identify and protect species at greatest risk of extinction. Ecological theory and global-scale analyses of bird and mammal faunas suggest that small-bodied species are less vulnerable to extinction, yet this hypothesis remains untested for the largest group of vertebrates, fish. Here, we compare body-size distributions of freshwater and marine fishes under different levels of global extinction risk (i.e. listed as vulnerable, endangered or critically endangered according to the IUCN Red List of Threatened Species ) from different major sources of threat (habitat loss/degradation, human harvesting, invasive species and pollution). Location Global, freshwater and marine. Methods We collated maximum body length data for 22,800 freshwater and marine fishes and compared body-size frequency distributions after controlling for phylogeny. Results We found that large-bodied marine fishes are under greater threat of global extinction, whereas both small- and large-bodied freshwater species are more likely to be at risk. Our results support the notion that commercial fishing activities disproportionately threaten large-bodied marine and freshwater species, whereas habitat degradation and loss threaten smaller-bodied marine fishes. Main conclusions Our study provides compelling evidence that global fish extinction risk does not universally scale with body size. Given the central role of body size for trophic position and the functioning of food webs, human activities may have strikingly different effects on community organization and food web structure in freshwater and marine systems.

Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains

Abstract: Aim This research aims to understand the factors that shape elevational diversity gradients and how those factors vary with spatial grain. Specifically, we test the predictions of the species‐productivity hypothesis, species‐temperature hypothesis, the metabolic theory of ecology and the mid-domain effects null model. We also examine how the effects of productivity and temperature on richness depend on spatial grain. Location Deciduous forests along an elevational gradient in Great Smoky Mountains National Park, USA. Methods We sampled 22 leaf litter ant assemblages at three spatial grains, from 1-m 2 quadrats to 50 × 50 m plots using Winkler samplers. Results Across spatial grains, warmer sites had more species than did cooler sites, and primary productivity did not predict ant species richness. We found some support for the predictions of the metabolic theory of ecology, but no support for the mid-domain effects null model. Thus, our data are best explained by some version of a species‐temperature hypothesis. Main conclusions Our results suggest that temperature indirectly affects ant species diversity across spatial grains, perhaps by limiting access to resources. Warmer sites support more species because they support more individuals, thereby reducing the probability of local extinction. Many of our results from this elevational gradient agree with studies at more global scales, suggesting that some mechanisms shaping ant diversity gradients are common across scales.

Ice age legacies in the geographical distribution of tree species richness in europe

Abstract: Aim This study uses a high-resolution simulation of the Last Glacial Maximum (LGM) climate to assess: (1) whether LGM climate still affects the geographical species richness patterns in the European tree flora and (2) the relative importance of modern and LGM climate as controls of tree species richness in Europe. Location The parts of Europe that were unglaciated during the LGM. Methods Atlas data on the distributions of 55 tree species were linked with data on modern and LGM climate and climatic heterogeneity in a geographical information system with a 60-km grid. Four measures of species richness were computed: total richness, and richness of the 18 most restricted species, 19 species of medium incidence (intermediate species) and 18 most widespread species. We used ordinary least-squares regression and spatial autoregressive modelling to test and estimate the richness‐climate relationships. Results LGM climate constituted the best single set of explanatory variables for richness of restricted species, while modern climate and climatic heterogeneity was best for total and widespread species richness and richness of intermediate species, respectively. The autoregressive model with all climatic predictors was supported for all richness measures using an information-theoretic approach, albeit only weakly so for total species richness. Among the strongest relationships were increases in total and intermediate richness with climatic heterogeneity and in restricted richness with LGM growing-degree-days. Partial regression showed that climatic heterogeneity accounted for the largest unique variation fraction for intermediate richness, while LGM climate was particularly important for restricted richness. Main conclusions LGM climate appears to still affect geographical patterns of tree species richness in Europe, albeit the relative importance of modern and LGM climate depends on range size. Notably, LGM climate is a strong richness control for species with a restricted range, which appear to still be associated with their glacial refugia.

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Abstract: Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape-level and species-level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β-diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.

Fire persistence traits of plants along a productivity and disturbance gradient in mediterranean shrublands of south‐east Australia

Abstract: Aim To understand changes in fire persistence traits of plants along a latitudinal gradient, considering the interactions between productivity, community (fuel) structure and fire regime. Location A gradient in the south of Australia (latitude 33‐37 ° S; longitude 140‐ 143 ° E), including: Little Desert National Park (VIC), Big Desert Wilderness Park (VIC), Murray-Sunset National Park (VIC), Danggali Conservation Park (SA) and Ta rawi Nature Reserve (NSW). Methods We selected four areas along a latitudinal gradient for which information on fire history and vegetation was available. Then, we tested to what extent the four selected areas have different climate and different fire regimes. Plant cover values of different life forms provided an indication of the plant community structure and flammability, and the proportion of species with different fire persistence traits (resprouting, seedbank persistence) informed us on the trait selection. Results Precipitation decreases and temperature increases from south to north. Thus the selected sites represent a gradient from high productivity (low aridity) in the south to low productivity (high aridity) in the north. Fire statistics suggest that fire frequency parallels productivity. There is a tendency for life form dominance and community structure to shift in such a way that fuel connectivity is reduced towards the north. Resprouting species increase and obligate seeders decrease along the fire‐productivity gradient. Main conclusions Changes in plant traits are difficult to understand without simultaneous consideration of both the disturbance and the productivity gradients. In our study area, fire regime and productivity interact in such a way that decreases in productivity imply changes in fuel structure that produce a reduction in fire frequency. Resprouting species are better represented at the high fire‐productivity part of the gradient, while obligate seeders are better represented at the opposite end of the gradient. The results also emphasize the importance of considering not only climate changes but also changes in fuel structure to predict future fire regimes.

The global relationship between forest productivity and biomass

Abstract: Aim We aim to determine the empirical relationship between above-ground forest productivity and biomass. There are theoretical reasons to assume a relationship between forest structure and function, as both may be influenced by similar ecological factors such as moisture supply. Also, dynamic global vegetation model simulations imply that any increase in forest productivity driven by climate change will result in increases in biomass and therefore carbon storage. However, few studies have explored the strength and form of the relationship between forest productivity and biomass, whether in space or time. Location Global scale. Methods We collated a large data set of above-ground biomass (AGB) and above-ground net primary productivity (ANPP) and tested the extent to which spatial variation in forest biomass across the Earth can be predicted from forest productivity. Results The global ANPP–AGB relationship differs fundamentally from the strongly positive, linear relationship reported in earlier analyses, which mostly lacked tropical sites. AGB begins to peak at c. 15–20 Mg ha−1 year−1 ANPP, plateaus at ANPP > 20–25 Mg ha−1 year−1, and may actually decline at higher levels of production. Main conclusions High turnover rates in high-productivity forests may limit AGB by promoting the dominance of species with a low wood density. Predicted increases in ANPP will not necessarily favour increases in forest carbon storage, especially if changes in productivity are accompanied by compositional shifts.

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Abstract: Aim Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host‐parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites. Methods We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores. Results We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites. Main conclusions Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.

A continental‐scale analysis of tree cover in African savannas

Abstract: Aim We present a continental-scale analysis that explores the processes controlling woody community structure in tropical savannas. We analyse how biotic and abiotic factors interact to promote and modify tree cover, examine alternative ecological hypotheses and quantify disturbance effects using satellite estimates of tree cover. Location African savannas. Methods Tree cover is represented as a resource-driven potential cover related to rainfall and soil characteristics perturbed by natural and human factors such as fire, cattle grazing, human population and cultivation. Within this framework our approach combines semi-empirical modelling and information theory to identify the best models. Results Woody community structure across African savannas is best represented by a sigmoidal response of tree cover to mean annual precipitation (MAP), with a dependency on soil texture, which is modified by the separate effects of fire, domestic livestock, human population density and cultivation intensity. This model explains c. 66% of the variance in tree cover and appears consistent across the savanna regions of Africa. Main conclusions The analysis provides a new understanding of the importance and interaction of environmental and disturbance factors that create the broad spatial patterns of tree cover observed in African savannas. Woody cover increases with rainfall, but is modified by disturbances. These ‘perturbation’ effects depend on MAP regimes: in arid savannas (MAP < 400 mm) they are generally small (< 1% decrease in cover), while in semi-arid and mesic savannas (400–1600 mm), perturbations result in an average 2% (400 mm) to 23% (1600 mm) decrease in cover; fire frequency and human population have more influence than cattle, and cultivation appears, on average, to lead to small increases in woody cover. Wet savannas (1600–2200 mm) are controlled by perturbations that inhibit canopy closure and reduce tree cover by, on average, 24–34%. Full understanding of the processes determining savanna structure requires consideration of resource limitation and disturbance dynamics.

Niche properties of Central European spiders : shading, moisture and the evolution of the habitat niche

Abstract: Aim Niche theory emphasizes the importance of environmental conditions for the distribution and abundance of species. Using a macroecological approach our study aimed at identifying the important environmental gradients for spiders. We generated numerical values of niche position and niche width. We also investigated relationships between these niche properties as well as the degree of phylogenetic conservatism in order to draw conclusions about the evolution of the habitat niche. Location Central Europe: lowlands of Austria, Belgium, Germany, Luxembourg, the Netherlands and Switzerland. Methods We analysed 244 published spider communities from 70 habitat types by correspondence analysis. The resulting community scores were used to test for correlations with habitat characteristics. Species scores were used to derive niche position (mean scores) and niche width (standard deviation of scores). To test for niche conservatism we estimated variance components across the taxonomic hierarchy. Results The first two axes of the correspondence analysis were correlated with shading and moisture, respectively. Niche width had a hump-shaped relationship to both environmental gradients. β-diversity was strikingly higher in open habitats than in forests. Habitat niche conservatism was lower than phylogenetic conservatism in body size. Main conclusions Environmental factors are important drivers for the β-diversity of spiders, especially across open habitats. This underlines the importance of preserving the whole range of moisture conditions in open habitats. Narrow niches of species occurring at the ends of both environmental gradients indicate that adaptations to extreme habitats lead to constraints in ecological flexibility. Nevertheless, the habitat niche of species seems to evolve much faster than morphological or physiological traits.

Can niche-based distribution models outperform spatial interpolation?

Abstract: Aim Distribution modelling relates sparse data on species occurrence or abundance to environmental information to predict the population of a species at any point in space. Recently, the importance of spatial autocorrelation in distributions has been recognized. Spatial autocorrelation can be categorized as exogenous (stemming from autocorrelation in the underlying variables) or endogenous (stemming from activities of the organism itself, such as dispersal). Typically, one asks whether spatial models explain additional variability (endogenous) in comparison to a fully specified habitat model. We turned this question around and asked: can habitat models explain additional variation when spatial structure is accounted for in a fully specified spatially explicit model? The aim was to find out to what degree habitat models may be inadvertently capturing spatial structure rather than true explanatory mechanisms. Location We used data from 190 species of the North American Breeding Bird Survey covering the conterminous United States and southern Canada. Methods We built 13 different models on 190 bird species using regression trees. Our habitat-based models used climate and landcover variables as independent variables. We also used random variables and simulated ranges to validate our results. The two spatially explicit models included only geographical coordinates or a contagion term as independent variables. As another angle on the question of mechanism vs. spatial structure we pitted a model using related bird species as predictors against a model using randomly selected bird species. Results The spatially explicit models outperformed the traditional habitat models and the random predictor species outperformed the related predictor species. In addition, environmental variables produced a substantial R2 in predicting artificial ranges. Main conclusions We conclude that many explanatory variables with suitable spatial structure can work well in species distribution models. The predictive power of environmental variables is not necessarily mechanistic, and spatial interpolation can outperform environmental explanatory variables.

Energy and interspecific body size patterns of amphibian faunas in Europe and North America: anurans follow Bergmann's rule, urodeles its converse

Abstract: Aim To describe broad-scale geographical patterns of body size for European and North American amphibian faunas and to explore possible processes underlying these patterns. Specifically, we propose a heat balance hypothesis, as both heat conservation and heat gain determine the heat balance of ectotherms, and test it along with five other hypotheses that have a possible influence on body size gradients: size dependence, migration ability, primary productivity, seasonality and water availability. Location Western Europe and North America north of Mexico. Methods We processed distribution maps for native amphibian species to estimate the mean body size in 110 × 110 km cells and calculated eight environmental predictors to explore the relationship between environmental gradients and the observed patterns. We used least squares regression modelling and model selection approaches based on information theory to evaluate the relative support for each hypothesis. Results We found consistent body size gradients and similar relationships to environmental variables within each amphibian group in Europe and North America. Annual potential evapotranspiration, a measure of environmental energy, was the strongest predictor of mean body size in both regions. However, the contrasting responses to ambient energy in each group resulted in opposite geographical patterns, i.e. anurans increased in size from high- to low-energy areas in both continents and urodeles showed the opposite pattern. Main conclusions Our results support the heat balance hypothesis, suggesting that the thermoregulatory abilities of anurans would allow them to reach larger sizes in colder climates by optimizing the trade-off between heating and cooling rates, whereas a lack of such strategies among urodele faunas would explain why these organisms tend to be smaller in cooler areas. These findings may also have implications for the role of climate warming on the global decline of amphibians.

The role of environment and mid-domain effect on moth species richness along a tropical elevational gradient

Abstract: Aim The biodiversity of geometrid moths (Lepidoptera) along a complete tropical elevational gradient was studied for the first time. The patterns are described, and the role of geometric constraints and environmental factors is explored. Location The study was carried out along the Barva Transect (10° N, 84° W), a complete elevational gradient ranging from 40 to 2730 m a.s.l. in Braulio Carrillo National Park, Costa Rica, and adjacent areas. Methods Moths were sampled manually in 2003 and 2004 at 12 rain forest sites using light ‘towers’, each with two 15 W ultraviolet fluorescent tubes. We used abundance-based rarefaction, statistical estimation of true richness (Chao 1), geographically interpolated observed richness and Fisher's alpha as measures of local diversity. Results A total of 13,765 specimens representing 739 species were analysed. All four measures showed a hump-shaped pattern with maxima between 500 and 2100 m elevation. The two subfamilies showed richness and diversity maxima at either lower (Ennominae) or higher (Larentiinae) elevation than Geometridae as a whole. Among the four environmental factors tested, relative humidity yielded the highest correlation over the transect with the rarefaction-based richness estimates as well as with estimated true species richness of Geometridae as a whole and of Larentiinae, while rainfall explained the greatest variation of Ennominae richness. The elevational pattern of moth richness was discordant with both temperature and with tree species richness. A combination of all environmental factors in a stepwise multiple regression produced high values of r2 in Geometridae. The potential effects of geometric constraints (mid-domain effect, MDE) were investigated by comparing them with observed, interpolated richness. Overall, models fitted very well for Geometridae as a whole and for Ennominae, but less well for Larentiinae. Small-ranged species showed stronger deviations from model predictions than large-ranged species, and differed strikingly between the two subfamilies, suggesting that environmental factors play a more pronounced role for small-ranged species. We hypothesize that small-ranged species (at least of the Ennominae) may tend to be host specialists, whereas large-ranged species tend to be polyphagous. Based on interpolated ranges, mean elevational range for these moths was larger with increasing elevation, in accordance with Rapoport's elevational rule, although sampling effects may have exaggerated this pattern. The underlying mechanism remains unknown because Rapoport's ‘rescue’ hypothesis could not explain the observed pattern. Conclusions The results clearly show that moth diversity shows a hump-shaped pattern. However, remarkable variation exists with regard to taxon and range size. Both environmental and geometric factors are likely to contribute to the observed patterns.

Around the world in 10 million years: biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae)

Abstract: Aim The species-rich family of true toads (Anura: Bufonidae) has been the focus of several earlier studies investigating the biogeography of geographically widespread taxa. Herein, we employ newly developed Bayesian divergence estimate methods to investigate the biogeographical history of this group. Resulting age estimates are used to test several key temporal hypotheses including that the origin of the bufonid clade pre-dates Gondwanan vicariance (~105 million years ago, Ma). Area cladograms are also invoked to investigate the geographical origin of the family. Location Worldwide, except the Australia‐New Guinea plate, Madagascar and the Antarctic. Methods A phylogenetic hypothesis of the relationships among true toads was derived from analysis of 2521 bp of DNA data including fragments from three mitochondrial ( 12S , tRNA val , 16S ) and two nuclear ( RAG-1 , CXCR-4 ) genes. Analysis of multiple, unlinked loci with a Bayesian method for estimating divergence times allowed us to address the timing and biogeographical history of Bufonidae. Resulting divergence estimates permitted the investigation of alternative vicariance/dispersal scenarios that have been proposed for true toads. Results Our area cladogram resulting from phylogenetic analysis of DNA data supports a South American origin for Bufonidae. Divergence estimates indicate that the family originated earlier than had been suggested previously (78‐99 Ma). The age of the enigmatic Caribbean clade was dated to the late Palaeocene‐early Eocene. A return of bufonids to the New World in the Eocene was followed by rapid diversification and secondary expansion into South America by the early Oligocene (Rupelian). Main conclusions The South American origin of Bufonidae in the Upper Cretaceous was followed by relatively rapid expansion and radiation around the globe, ending with a return to the Americas via a Eurasian/North American land bridge in the Eocene. Though the exact route of this dispersal (Beringia or North Atlantic) remains unclear, an argument is made for the less frequently invoked North Atlantic connection. The origin of the enigmatic Caribbean lineage was found to be consistent with colonization following the bolide impact at the K/T boundary. These findings provide the first, firm foundation for understanding true toad divergence times and their truly remarkable and global radiation.

Effects of landscape structure and land-use intensity on similarity of plant and animal communities

Abstract: Aim Species richness in itself is not always sufficient to evaluate land management strategies for nature conservation. The exchange of species between local communities may be affected by landscape structure and land-use intensity. Thus, species turnover, and its inverse, community similarity, may be useful measures of landscape integrity from a diversity perspective. Location A European transect from France to Estonia. Methods We measured the similarity of plant, bird, wild bee, true bug, carabid beetle, hoverfly and spider communities sampled along gradients in landscape composition (e.g. total availability of semi-natural habitat), landscape configuration (e.g. fragmentation) and land-use intensity (e.g. pesticide loads). Results Total availability of semi-natural habitats had little effect on community similarity, except for bird communities, which were more homogeneous in more natural landscapes. Bee communities, in contrast, were less similar in landscapes with higher percentages of semi-natural habitats. Increased landscape fragmentation decreased similarity of true bug communities, while plant communities showed a nonlinear, U-shaped response. More intense land use, specifically increased pesticide burden, led to a homogenization of bee, bug and spider communities within sites. In these cases, habitat fragmentation interacted with pesticide load. Hoverfly and carabid beetle community similarity was differentially affected by higher pesticide levels: for carabid beetles similarity decreased, while for hoverflies we observed a U-shaped relationship. Main conclusions Our study demonstrates the effects of landscape composition, configuration and land-use intensity on the similarity of communities. It indicates reduced exchange of species between communities in landscapes dominated by agricultural activities. Taxonomic groups differed in their responses to environmental drivers and using but one group as an indicator for 'biodiversity' as such would thus not be advisable.

Searching for phylogenetic pattern in biological invasions

Abstract: It has been suggested that alien species with close indigenous relatives in the introduced range may have reduced chances of successful establishment and invasion (Darwin's naturalization hypothesis). Studies trying to test this have in fact been addressing four different hypotheses, and the same data can support some while rejecting others. In this paper, we argue that the phylogenetic pattern will change depending on the spatial and phylogenetic scales considered. Expectations and observations from invasion biology and the study of natural communities are that at the spatial scale relevant to competitive interactions, closely related species will be spatially separated, whereas at the regional scale, species in the same genera or families will tend to co-occur more often than by chance. We also argue that patterns in the relatedness of indigenous and naturalized plants are dependent on the continental/island setting, spatial occupancy levels, and on the group of organisms under scrutiny. Understanding how these factors create a phylogenetic pattern in invasions will help us predict which groups are more likely to invade where, and should contribute to general ecological theory.

Habitat and climate heterogeneity maintain beta‐diversity of birds among landscapes within ecoregions

Abstract: Aim Habitat and climate heterogeneity may affect patterns of species diversity from the relatively local scale of communities to the broad biogeographical scale of continents. However, the effects of heterogeneity on species diversity have not been studied as widely at intermediate scales although differences among landscapes in local climate and habitat should maintain beta-diversity. Location Bailey ecoregions in the USA. Methods Using a geographically extensive dataset on bird distribution and abundance in 35 ecoregions, we tested for the effects of habitat and climate heterogeneity on beta-diversity at two discrete spatial scales: among sample points within landscapes, and among landscapes within ecoregions. Results Landscape-level beta-diversity typically accounted for 50% or more of gamma-diversity and was significantly and positively related to habitat heterogeneity (elevational range within an ecoregion) and climate heterogeneity (variation in potential evapotranspiration). Contrary to predictions, point-level beta-diversity was negatively related to habitat and climate heterogeneity, perhaps because heterogeneity constrains alpha-diversity at the landscape level. The geographical spatial separation of landscapes within an ecoregion did not significantly affect beta-diversity at either scale. Main conclusions Our results suggest that habitat selection and adaptation to local climate may be the primary processes structuring bird diversity among landscapes within ecoregions, and that dispersal limitation has a lesser role in influencing beta-diversity among landscapes.

What determines conformity to Bergmann's rule?

Abstract: Aim Bergmann’s rule, the tendency of body size within species in bird and mammal populations to be positively correlated with latitude, is among the best known biogeographical generalizations. The factors behind such clines, however, are not well understood. Here we use a large data base of 79 mammalian carnivore species to examine the factors affecting latitudinal size clines. Location Worldwide. Methods We measured the skulls and teeth of carnivores in natural history museums, and calculated the amount of variation in size explained by latitude, supplementing our measurements with published data. We examined the effects of a number of variables on the tendency to show latitudinal clines. Results We found that geographical range and latitudinal extent are strongly related to size clines. Minimum temperatures across the range, net primary productivity and habitat diversity also have some, albeit much less, influence. Main conclusions We suggest that species with large geographical ranges are likely to encounter significant heterogeneity in those factors that influence body size, and are thus likely to exhibit size clines. However, the key factors that determine body size may not always operate along a latitudinal (or other geographical) cline, but be spatially linked to patches in the species range. One such important factor is likely to be food availability, which we show is a strong predictor of size in the brown bear ( Ursus arctos ) but is not associated with a latitudinal cline. We argue that the spatial distribution of key resources within the species range constitutes a significant predictor of carnivore body size.

Geographical and environmental range expansion through polyploidy in wild potatoes (Solanum section Petota)

Abstract: Aim To assess evidence for geographical and environmental range expansion through polyploidy in wild potatoes ( Solanum sect. Petota ). There are diploids, triploids, tetraploids, pentaploids and hexaploids in this group. Location Wild potatoes occur from the south-western USA (Utah and Colorado), throughout the tropical highlands of Mexico, Central America and the Andes, to Argentina, Chile and Uruguay. Methods We compiled 5447 reports of ploidy determination, covering 185 of the 187 species, of which 702 determinations are presented here for the first time. We assessed the frequency of cytotypes within species, and analysed the geographical and climatic distribution of ploidy levels. Results Thirty-six per cent of the species are entirely or partly polyploid. Multiple cytotypes exist in 21 species, mostly as diploid and triploid, but many more may await discovery. We report the first chromosome count (2 n = 24) for Solanum hintonii . Diploids occupy a larger area than polyploids, but diploid and tetraploid species have similar range sizes, and the two species with by far the largest range sizes are tetraploids. The fraction of the plants that are polyploids is much higher from Mexico to Ecuador than farther south. Compared with diploids, triploids tend to occur in warmer and drier areas, whereas higher-level polyploids tend to occur in relatively cold areas. Diploids are absent from Costa Rica to southern Colombia, the wettest part of the group’s range. Main conclusions These results suggest that polyploidy played an important role in this group’s environmental differentiation and range expansion.

Palms tracking climate change

Abstract: Aim Many species are currently expanding their ranges in response to climate change, but the mechanisms underlying these range expansions are in many cases poorly understood. In this paper we explore potential climatic factors governing the recent establishment of new palm populations far to the north of any other viable palm population in the world. Location Southern Switzerland, Europe, Asia and the world. Methods We identified ecological threshold values for the target species, Trachycarpus fortunei, based on gridded climate data, altitude and distributional records from the native range and applied them to the introduced range using local field monitoring and measured meteorological data as well as a bioclimatic model. Results We identified a strong relationship between minimum winter temperatures, influenced by growing season length and the distribution of the palm in its native range. Recent climate change strongly coincides with the palm's recent spread into southern Switzerland, which is in concert with the expansion of the global range of palms across various continents. Main conclusions Our results strongly suggest that the expansion of palms into (semi-)natural forests is driven by changes in winter temperature and growing season length and not by delayed population expansion. This implies that this rapid expansion is likely to continue in the future under a warming climate. Palms in general, and T. fortunei in particular, are significant bioindicators across continents for present-day climate change and reflect a global signal towards warmer conditions.

Species invasions and the changing biogeography of Australian freshwater fishes

Abstract: Aim By dissolving natural physical barriers to movement, human-mediated species introductions have dramatically reshuffled the present-day biogeography of freshwater fishes. The present study investigates whether the antiquity of Australia's freshwater ichthyofauna has been altered by the widespread invasion of non-indigenous fish species. Location Australia. Methods Using fish presence-absence data for historical and present-day species pools, we quantified changes in faunal similarity among major Australian drainage divisions and among river basins of north-eastern Australia according to the Sorensen index, and related these changes to major factors of catchment disturbance that significantly alter river processes. Results Human-mediated fish introductions have increased faunal similarity among primary drainages by an average of 3.0% (from 17.1% to 20.1% similarity). Over three-quarters of the pairwise changes in drainage similarity were positive, indicating a strong tendency for taxonomic homogenization caused primarily by the widespread introduction of Carassius auratus, Gambusia holbrooki, Oncorhynchus mykiss and Poecilia reticulata. Faunal homogenization was highest in drainages subjected to the greatest degree of disturbance associated with human settlement, infrastructure and change in land use. Scenarios of future species invasions and extinctions indicate the continued homogenization of Australian drainages. In contrast, highly idiosyncratic introductions of species in river basins of north-eastern Australia have decreased fish faunal similarity by an average of 1.4%. Main conclusions We found that invasive species have significantly changed the present-day biogeography of fish by homogenizing Australian drainages and differentiating north-eastern river basins. Decreased faunal similarity at smaller spatial scales is a result of high historical similarity in this region and reflects the dynamic nature of the homogenization process whereby sporadic introductions of new species initially decrease faunal similarity across basins. Our study points to the importance of understanding the role of invasive species in defining patterns of present-day biogeography and preserving the antiquity of Australia's freshwater biodiversity.

Size evolution in island lizards

Abstract: Aim The island rule, small animal gigantism and large animal dwarfism on islands, is a topic of much recent debate. While size evolution of insular lizards has been widely studied, whether or not they follow the island rule has never been investigated. I examined whether lizards show patterns consistent with the island rule. Location Islands worldwide. Methods I used literature data on the sizes of island–mainland population pairs in 59 species of lizards, spanning the entire size range of the group, and tested whether small insular lizards are larger than their mainland conspecifics and large insular lizards are smaller. I examined the influence of island area, island isolation, and dietary preferences on lizard size evolution. Results Using mean snout–vent length as an index of body size, I found that small lizards on islands become smaller than their mainland conspecifics, while large ones become larger still, opposite to predictions of the island rule. This was especially strong in carnivorous lizards; omnivorous and herbivorous species showed a pattern consistent with the island rule but this result was not statistically significant. No trends consistent with the island rule were found when maximum snout–vent length was used. Island area had, at best, a weak effect on body size. Using maximum snout–vent length as an index of body size resulted in most lizard populations appearing to be dwarfed on islands, but no such pattern was revealed when mean snout–vent length was used as a size index. Main conclusions I suggest that lizard body size is mostly influenced by resource availability, with large size allowing some lizard populations to exploit resources that are unavailable on the mainland. Lizards do not follow the island rule. Maximum snout–vent length may be biased by sampling effort, which should be taken into account when one uses this size index.