Showing papers in "Global Ecology and Biogeography in 2009"

A global study of relationships between leaf traits, climate and soil measures of nutrient fertility

Abstract: Aim This first global quantification of the relationship between leaf traits and soil nutrient fertility reflects the trade-off between growth and nutrient conservation. The power of soils versus climate in predicting leaf trait values is assessed in bivariate and multivariate analyses and is compared with the distribution of growth forms (as a discrete classification of vegetation) across gradients of soil fertility and climate. Location All continents except for Antarctica. Methods Data on specific leaf area (SLA), leaf N concentration (LNC), leaf P concentration (LPC) and leaf N:P were collected for 474 species distributed across 99 sites (809 records), together with abiotic information from each study site. Individual and combined effects of soils and climate on leaf traits were quantified using maximum likelihood methods. Differences in occurrence of growth form across soil fertility and climate were determined by one-way ANOVA. Results There was a consistent increase in SLA, LNC and LPC with increasing soil fertility. SLA was related to proxies of N supply, LNC to both soil total N and P and LPC was only related to proxies of P supply. Soil nutrient measures explained more variance in leaf traits among sites than climate in bivariate analysis. Multivariate analysis showed that climate interacted with soil nutrients for SLA and area-based LNC. Mass-based LNC and LPC were determined mostly by soil fertility, but soil P was highly correlated to precipitation. Relationships of leaf traits to soil nutrients were stronger than those of growth form versus soil nutrients. In contrast, climate determined distribution of growth form more strongly than it did leaf traits. Main conclusions We provide the first global quantification of the trade-off between traits associated with growth and resource conservation ‘strategies’ in relation to soil fertility. Precipitation but not temperature affected this trade-off. Continuous leaf traits might be better predictors of plant responses to nutrient supply than growth form, but growth forms reflect important aspects of plant species distribution with climate.

Predicting the past distribution of species climatic niches.

Abstract: Predicting past distributions of species climatic niches, hindcasting, by using climate envelope models (CEMs) is emerging as an exciting research area. CEMs are used to examine veiled evolutionary questions about extinctions, locations of past refugia and migration pathways, or to propose hypotheses concerning the past population structure of species in phylogeographical studies. CEMs are sensitive to theoretical assumptions, to model classes and to projections in non-analogous climates, among other issues. Studies hindcasting the climatic niches of species often make reference to these limitations. However, to obtain strong scientific inferences, we must not only be aware of these potential limitations but we must also overcome them. Here, I review the literature on hindcasting CEMs. I discuss the theoretical assumptions behind niche modelling, i.e. the stability of climatic niches through time and the equilibrium of species with climate. I also summarize a set of ‘recommended practices’ to improve hindcasting. The studies reviewed: (1) rarely test the theoretical assumptions behind niche modelling such as the stability of species climatic niches through time and the equilibrium of species with climate; (2) they only use one model class (72% of the studies) and one palaeoclimatic reconstruction (62.5%) to calibrate their models; (3) they do not check for the occurrence of non-analogous climates (97%); and (4) they do not use independent data to validate the models (72%). Ignoring the theoretical assumptions behind niche modelling and using inadequate methods for hindcasting CEMs may well entail a cascade of errors and naive ecological and evolutionary inferences. We should also push integrative research lines linking macroecology, physiology, population biology, palaeontology, evolutionary biology and CEMs for a better understanding of niche dynamics across space and time.

Global analysis of bird elevational diversity

Abstract: Aim Elevational gradients distributed across the globe are a powerful test system for understanding biodiversity. Here I use a comprehensive set of bird elevational gradients to test the main drivers of diversity, including sampling, area, mid-domain effect, temperature, temperature and water availability, and hypotheses of evolutionary history. Location Seventy-eight elevational gradients of bird diversity from mountains in both hemispheres spanning 24.5 ° S to 48.2 ° N, including gradients from various climates, biogeographical regions and habitat types. Methods Data on bird elevational diversity were taken from the literature. Of the 150 datasets found or compiled, only those with a high, unbiased sampling effort were used in analyses. Datasets sampled all birds, all breeding birds or all forest birds; a few studies detailed seasonal, elevational shifts. Eighteen predictions of diversity theory were tested, including three sets of interactions. Results Birds display four distinct diversity patterns in nearly equal frequency on mountains: decreasing diversity, low-elevation plateaus, low-elevation plateaus with mid-peaks, and unimodal mid-elevational peaks. Bird elevational diversity strongly supports current climate as the main driver of diversity, particularly combined trends in temperature and water availability. Bird diversity on humid mountains is either decreasing or shows a low-elevation plateau in diversity, while on dry mountains it is unimodal or a broad, low-elevation plateau usually with a mid-elevation maximum. The predictions of sampling, area and mid-domain effect were not consistently supported globally. The only evolutionary hypothesis with preliminary support was niche conservatism. Main conclusions Both water and temperature variables are needed to comprehensively predict elevational diversity patterns for birds. This result is consistent for breeding and forest birds, for both hemispheres, and for local- or regional-scale montane gradients. More analyses are needed to discern whether the mechanism underlying these relationships is ecological, based on direct physiological limitations or indirect food resource limitations, or historical, based on phylogenetic niche conservation or other evolutionary trends related to climate. The species‐area and middomain effects are not supported as primary drivers of elevational diversity in birds.

Global pattern of NPP to GPP ratio derived from MODIS data: effects of ecosystem type, geographical location and climate

Abstract: To examine the global pattern of the net primary production (NPP)/gross primary production (GPP) ratio of the Earth's land area along geographical and climatic gradients. The global planetary ecosystem. The 4-year average annual NPP/GPP ratio of the Earth's land area was calculated using 2000-03 Moderate Resolution Imaging Spectroradiometer (MODIS) data. The global pattern of the NPP/GPP ratio was investigated by comparing it among each typical terrestrial ecosystem and plotting it along a geographical and climatic gradient, including latitude, altitude, temperature and precipitation. The global terrestrial ecosystem had an average NPP/GPP ratio value of 0.52 with minor variation from 2000 to 2003. However, the NPP/GPP ratio showed considerable spatial variation associated with ecosystem type, geographical location and climate. Densely vegetated ecosystems had a lower NPP/GPP ratio than sparsely vegetated ecosystems. Forest ecosystems had a lower NPP/GPP ratio than shrub and herbaceous ecosystems. Geographically, the NPP/GPP ratio increased with altitude. In the Southern Hemisphere, the NPP/GPP ratio decreased along latitude from 30 degrees to 10 degrees and it exhibited high fluctuation in the Northern Hemisphere. Climatically, the NPP/GPP ratio exhibited a decreasing trend along enhanced precipitation when it was less than 2300 mm year(-1) and a static trend when the annual precipitation was over 2300 mm. The NPP/GPP ratio showed a decreasing trend along temperature when it was between -20 degrees C and 10 degrees C, and showed an increasing trend along rising temperature when it was between -10 degrees C and 20 degrees C. Within each ecosystem, the NPP/GPP ratio revealed a similar trend to the global trend along temperature and precipitation. The NPP/GPP ratio exhibited a pattern depending on the main climatic characteristics such as temperature and precipitation and geographical factors such as latitude and altitude. The findings of this research challenge the widely held assumption that the NPP/GPP ratio is consistent regardless of ecosystem type.

Global-scale patterns of nutrient resorption associated with latitude, temperature and precipitation

Abstract: Aim Nutrient resorption from senescing leaves is an important mechanism of nutrient conservation in plants, but the patterns of nutrient resorption at the global scale are unknown. Because soil nutrients vary along climatic gradients, we hypothesize that nutrient resorption changes with latitude, temperature and precipitation. Location Global. Methods We conducted a meta-analysis on a global data set collected from published literature on nitrogen (N) and phosphorus (P) resorption of woody plants. Results For all data pooled, both N resorption efficiency (NRE) and P resorption efficiency (PRE) were significantly related to latitude, mean annual temperature (MAT) and mean annual precipitation (MAP): NRE increased with latitude but decreased with MAT and MAP. In contrast, PRE decreased with latitude but increased with MAT and MAP. When functional groups (shrub versus tree, coniferous versus broadleaf and evergreen versus deciduous) were examined individually, the patterns of NRE and PRE in relation to latitude, MAT and MAP were generally similar. Main conclusions The relationships between N and P resorption and latitude, MAT and MAP indicate the existence of geographical patterns of plant nutrient conservation strategies in relation to temperature and precipitation at the global scale, particularly for PRE, which can be an indicator for P limitation in the tropics and selective pressure shaping the evolution of plant traits. Our results suggest that, although the magnitude of plant nutrient resorption might be regulated by local factors such as substrate, spatial patterns are also controlled by temperature or precipitation.

Ensemble models predict Important Bird Areas in southern Africa will become less effective for conserving endemic birds under climate change.

Abstract: DST-NRF Centre of Excellence for Invasion Biology, the University of Pretoria, EU FP6 MACIS species targeted project (Minimisation of and Adaptation to Climate change: Impacts on biodiversity, contract No.: 044399) and EU FP6 ECOCHANGE integrated project (Challenges in assessing and forecasting biodiversity and ecosystem changes in Europe).

Global trends in senesced-leaf nitrogen and phosphorus

Abstract: Aim Senesced-leaf litter plays an important role in the functioning of terrestrial ecosystems. While green-leaf nutrients have been reported to be affected by climatic factors at the global scale, the global patterns of senesced-leaf nutrients are not well understood. Location Global. Methods Here, bringing together a global dataset of senesced-leaf N and P spanning 1253 observations and 638 plant species at 365 sites and of associated mean climatic indices, we describe the world-wide trends in senesced-leaf N and P and their stoichiometric ratios. Results Concentration of senesced-leaf N was highest in tropical forests, intermediate in boreal, temperate, and mediterranean forests and grasslands, and lowest in tundra, whereas P concentration was highest in grasslands, lowest in tropical forests and intermediate in other ecosystems. Tropical forests had the highest N : P and C : P ratios in senesced leaves. When all data were pooled, N concentration significantly increased, but senesced-leaf P concentration decreased with increasing mean annual temperature (MAT) and mean annual precipitation (MAP). The N : P and C : P ratios also increased with MAT and MAP, but C : N ratios decreased. Plant functional type (PFT), i.e. life-form (grass, herb, shrub or tree), phylogeny (angiosperm versus gymnosperm) and leaf habit (deciduous versus evergreen), affected senesced-leaf N, P, N : P, C : N and C : P with a ranking of senesced-leaf N from high to low: forbs ≈ shrubs ≈ trees > grasses, while the ranking of P was forbs ≈ shrubs ≈ trees < grasses. The climatic trends of senesced-leaf N and P and their stoichiometric ratios were similar between PFTs. Main conclusions Globally, senesced-leaf N and P concentrations differed among ecosystem types, from tropical forest to tundra. Differences were significantly related to global climate variables such as MAT and MAP and also related to plant functional types. These results at the global scale suggest that nutrient feedback to soil through leaf senescence depends on both the climatic conditions and the plant composition of an ecosystem.

Importance of abiotic stress as a range‐limit determinant for European plants: insights from species responses to climatic gradients

Abstract: Aim We examined whether species occurrences are primarily limited by physiological tolerance in the abiotically more stressful end of climatic gradients (the asymmetric abiotic stress limitation (AASL) hypothesis) and the geographical predictions of this hypothesis: abiotic stress mainly determines upper-latitudinal and upper-altitudinal species range limits, and the importance of abiotic stress for these range limits increases the further northwards and upwards a species occurs. Location Europe and the Swiss Alps. Methods The AASL hypothesis predicts that species have skewed responses to climatic gradients, with a steep decline towards the more stressful conditions. Based on presence-absence data we examined the shape of plant species responses (measured as probability of occurrence) along three climatic gradients across latitudes in Europe (1577 species) and altitudes in the Swiss Alps (284 species) using Huisman-Olff-Fresco, generalized linear and generalized additive models. Results We found that almost half of the species from Europe and one-third from the Swiss Alps showed responses consistent with the predictions of the AASL hypothesis. Cold temperatures and a short growing season seemed to determine the upper-latitudinal and upper-altitudinal range limits of up to one-third of the species, while drought provided an important constraint at lower-latitudinal range limits for up to one-fifth of the species. We found a biome-dependent influence of abiotic stress and no clear support for abiotic stress as a stronger upper range-limit determinant for species with higher latitudinal and altitudinal distributions. However, the overall influence of climate as a range-limit determinant increased with latitude. Main conclusions Our results support the AASL hypothesis for almost half of the studied species, and suggest that temperature-related stress controls the upper-latitudinal and upper-altitudinal range limits of a large proportion of these species, while other factors including drought stress may be important at the lower range limits.

Niche shift versus niche conservatism? Climatic characteristics of the native and invasive ranges of the Mediterranean house gecko (Hemidactylus turcicus)

Abstract: Aim The use of ecological niche models (ENMs) to predict potential distributions of species is steadily increasing. A necessary assumption is that climatic niches are conservative, but recent findings of niche shifts during biological invasion indicate that this assumption is not always valid. Selection of predictor variables may be one reason for the observed shifts. In this paper we assess differences in climatic niches in the native and invaded ranges of the Mediterranean house gecko (Hemidactylus turcicus) in terms of commonly applied climate variables in ENMs. We analyse which variables are more conserved versus relaxed (i.e. subject to niche shift). Furthermore, we study the predictive power of different sets of climate variables. Location The Mediterranean region and North America. Methods We developed models using Maxent and various subsets of variables out of 19 bioclimatic layers including: (1) two subsets comprising almost all variables excluding only highly collinear ones; (2) two subsets with minimalistic variable sets of water availability and energy measures; (3) two subsets focused on temperature-related parameters; (4) two subsets with precipitation-related parameters; and (5) one subset comprising variables combining temperature and precipitation characteristics. Occurrence data from the native Mediterranean range were used to predict the potential introduced range in North America and vice versa. Degrees of niche similarity and conservatism were assessed using both Schoener's index and Hellinger distances. The significance of the results was tested using null models. Results The degree of niche similarity and conservatism varied greatly among the predictors and variable sets applied. Shifts observed in some variables could be attributed to active habitat selection while others apparently reflected background effects. Main conclusions The study was based on comprehensive occurrence data from all regions where Hemidactylus turcicus is present in Europe and North America, providing a robust foundation. Our results clearly indicate that the degree of conservatism of niches in H. turcicus largely varies among predictors and variable sets applied. Therefore, the extent of niche conservatism of variables applied should always be tested in ENMs. This has an important impact on studies of biological invasion, impacts of climate change and niche evolution.

Seedling recruitment, survival and facilitation in alpine Pinus uncinata tree line ecotones. Implications and potential responses to climate warming

Abstract: Aims Alpine tree line ecotones are harsh environments where low temperatures constrain tree regeneration and growth. However, the expected upward shift of tree line ecotones in response to climate warming has not been ubiquitous. The lack of coupling between tree line dynamics and climate warming might be explained by factors other than climate variation that determine seedling recruitment in these ecotones. We want to assess how the availability of suitable habitat for establishment and the effects of facilitation on seedling survival and growth affect tree recruitment within tree line ecotones and modulate their responses to climate. Location We evaluate the relevance of these factors for Pinus uncinata tree line ecotones in the Catalan Pyrenees (north-east Spain) and Andorra. Methods We analysed the microhabitat of naturally established seedlings in rectangular plots at the tree line ecotone, assessing the habitat type and the proximity to potentially protective elements that may improve microsite conditions. We tested whether krummholz individuals influence regeneration at the tree line by performing a transplantation field experiment to evaluate the extent of facilitation on seedling survival and growth in height. A total of 820 seedlings were transplanted at different distances and orientations (resulting in 12 positions) from krummholz mats and monitored over 2 years. Results Safe sites for P. uncinata recruits consisted of sparse vegetation covering bare soil, gravel or litter, and close to protective elements that may ameliorate microsite conditions. The field experiment showed that directional positive interactions enhance seedling survival and growth, altering the spatial patterns of recruit survivorship, especially during harsh winter conditions (shallow and irregular snowpack). Main conclusions Our results suggest that scarce availability of safe sites and uneven facilitation by krummholz control seedling recruitment patterns within alpine tree line ecotones. Such constraints may distort or counter the response of tree line ecotones to climate warming at local and regional scales.

Predation has a greater impact in less productive environments: variation in roe deer, Capreolus capreolus, population density across Europe

Abstract: Aim We aimed to describe the large-scale patterns in population density of roe deer Caprelous capreolus in Europe and to determine the factors shaping variation in their abundance.

Interacting effects of changes in climate and forest cover on mortality and growth of the southernmost European fir forests

Abstract: Aims The combined effects of changes in climate and land use on tree mortality and growth patterns have rarely been addressed. Relict tree species from the Mediterranean Basin serve as appropriate models to investigate these effects, since they grow in climatically stressed areas which have undergone intense cover changes. The aim is to use climate, aerial photographs, stand structure and radial-growth data to explain the mortality and historical patterns of growth of Abies pinsapo in the area where this relict species was first protected. Location Sierra de las Nieves, West Baetic Range, southern Spain. Methods We assessed variations of tree cover in A. pinsapo forests through image analyses of aerial photographs spanning the last 50 years. We sampled 31 stands to assess current altitudinal patterns of forest structure and mortality. We evaluated the relationships between radial growth and regional climate using linear models in three sites at different elevations. Results Regional warming and a decrease in precipitation were detected. Forest tree cover increased at all elevations from 1957 until 1991, but it afterwards decreased below 1100 m. Currently, the likelihood of tree mortality increases downwards and is associated with dense, closed stands with a low living basal area. In contrast to previous droughts, a sharp synchronized reduction in tree growth, not fully accounted for in linear climate‐growth models, occurred at low elevations in 1994‐95, but not upwards. It was preceded by a weakening of the negative association between low-elevation growth and water deficit since the late 1970s. Conclusions The intense densification of A. pinsapo forests following strict protection measures in the late 1950s enhanced the vulnerability of climate-sensitive A. pinsapo forests to recent drier conditions. Such abrupt land-use changes help to explain recent patterns of mortality and growth decline in low-elevation A. pinsapo forests.

Beta diversity in relation to dispersal ability for vascular plants in North America

Abstract: Aim To test the hypothesis that plant species with a higher dispersal ability have a lower beta diversity. Location North America north of Mexico. Method Propagules of pteridophytes (ferns and their allies) are more vagile than propagules of spermatophytes (gymnosperms and angiosperms), and thus pteridophytes have a higher dispersal ability than do spermatophytes. The study area was divided into 71 geographical units distributed in five latitudinal zones. Species lists of pteridophytes and spermatophytes were compiled for each geographical unit. Three measures of beta diversity were used: β sim , which is one minus the Simpson index of similarity, β slope , which is the slope of the relationship between Simpson index and geographical distance, and β 0.5-distance , which is the distance that halves the similarity from its initial value. Results Average β sim is higher for spermatophytes than for pteridophytes, regardless of whether the data are analysed for the whole continent or for latitudinal zones. Average β sim decreases with increasing latitude for both spermatophytes and pteridophytes. The difference in average β sim between the two plant groups increases with increasing latitude, indicating that beta diversity decreases with increasing latitude faster for pteridophytes than for spermatophytes. When the Simpson index is regressed against geographical distance, the regression slope ( β slope ) is steeper for spermatophytes than for pteridophytes, and the slope decreases with increasing latitude for both plant groups. Similarly, β 0.5-distance was shorter for spermatophytes than for pteridophytes in each latitudinal zone and increased with increasing latitude for both plant groups. The results of the analyses using the three different measures of beta diversity are consistent.

Past and future range shifts and loss of diversity in dwarf willow (Salix herbacea L.) inferred from genetics, fossils and modelling

Abstract: Aim Climate change may cause loss of genetic diversity. Here we explore how a multidisciplinary approach can be used to infer effects of past climate change on species distribution and genetic diversity and also to predict loss of diversity due to future climate change. We use the arctic-alpine plant Salix herbacea L. as a model.

The global distribution of frugivory in birds

Abstract: Aim To examine patterns of avian frugivory across clades, geography and environments. Location Global, including all six major biogeographical realms (Afrotropics, Australasia, Indo-Malaya, Nearctic, Neotropics and Palaearctic). Methods First, we examine the taxonomic distribution of avian frugivory within orders and families. Second we evaluate, with traditional and spatial regression approaches, the geographical patterns of frugivore species richness and proportion. Third, we test the potential of contemporary climate (water–energy, productivity, seasonality), habitat heterogeneity (topography, habitat diversity) and biogeographical history (captured by realm membership) to explain geographical patterns of avian frugivory. Results Most frugivorous birds (50%) are found within the perching birds (Passeriformes), but the woodpeckers and allies (Piciformes), parrots (Psittaciformes) and pigeons (Columbiformes) also contain a significant number of frugivorous species (9–15%). Frugivore richness is highest in the Neotropics, but peaks in overall bird diversity in the Himalayan foothills, the East African mountains and in some areas of Brazil and Bolivia are not reflected by frugivores. Current climate explains more variance in species richness and proportion of frugivores than of non-frugivores whereas it is the opposite for habitat heterogeneity. Actual evapotranspiration (AET) emerges as the best single climatic predictor variable of avian frugivory. Significant differences in frugivore richness and proportion between select biogeographical regions remain after differences in environment (i.e. AET) are accounted for. Main conclusions We present evidence that both environmental and historical constraints influence global patterns of avian frugivory. Whereas water–energy dynamics possibly constrain frugivore distribution via indirect effects on food plants, regional differences in avian frugivory most likely reflect historical contingencies related to the evolutionary history of fleshy fruited plant taxa, niche conservatism and past climate change. Overall our results support an important role of co-diversification and environmental constraints on regional assembly over macroevolutionary time-scales.

Penguins from space: faecal stains reveal the location of emperor penguin colonies

Abstract: Aim To map and assess the breeding distribution of emperor penguins ( Aptenodytes forsteri ) using remote sensing. Location Pan-Antarctic. Methods Using Landsat ETM satellite images downloaded from the Landsat Image Mosaic of Antarctica (LIMA), we detect faecal staining of ice by emperor penguins associated with their colony locations. Emperor penguins breed on sea ice, and their colonies exist in situ between May and December each year. Faecal staining at these colony locations shows on Landsat imagery as brown patches, the only staining of this colour on sea ice. This staining can therefore be used as an analogue for colony locations. The whole continental coastline has been analysed, and each possible signal has been identified visually and checked by spectral analysis. In areas where LIMA data are unsuitable, freely available Landsat imagery has been supplemented. Results We have identified colony locations of emperor penguins at a total of 38 sites. Of these, 10 are new locations, and six previously known colony locations have been repositioned (by over 10 km) due to poor geographical information in old records. Six colony locations, all from old or unconfirmed records, were not found or have disappeared. Main conclusions We present a new pan-Antarctic species distribution of emperor penguins mapped from space. In one synoptic survey we locate extant emperor penguin colonies, a species previously poorly mapped due to its unique breeding habits, and provide a vital geographical resource for future studies of an iconic species believed to be vulnerable to future climate change.

Plants intertwine fluvial landform dynamics with ecological succession and natural selection : a niche construction perspective for riparian systems

Abstract: Aim To contribute to the development of a macroevolutionary framework for riparian systems, reinforcing conceptual linkages between earth surface processes and biological and ecological processes. Location Riparian systems. Methods Literature review leading to an original proposition for perceiving the functioning of riparian systems in a new and different way. Results Riparian systems provide diverse landforms, habitats and resources for animals and plants. Certain organisms, defined as ‘ecosystem engineers’, significantly create and modify the physical components of riparian systems. Many studies have highlighted such engineering effects by animals on riparian systems, but the identification and understanding of the effects and responses of plants within fluvial corridors have emerged only recently. The modulation of matter, resources and energy flows by engineering plants helps establish characteristic sequences of fluvial landform creation and maintenance associated with synergetic ecological successions. We relate this process to the concept of niche construction, developed mainly by evolutionary biologists. Feedbacks between adaptive responses of riparian plants to flow regime and adjusting effects on biostabilization and bioconstruction are discussed in the context of niche construction at the scale of ecological succession and the evolution of organisms. Main conclusions Our conceptualization forges an integrated approach for understanding vegetated fluvial systems from a macroevolutionary perspective, for elucidating riparian ecosystem dynamics and potentially for establishing long-term stream conservation and restoration strategies.

The latitudinal gradient of beta diversity in relation to climate and topography for mammals in North America

Abstract: Aim Spatial turnover of species, or beta diversity, varies in relation to geographical distance and environmental conditions, as well as spatial scale. We evaluated the explanatory power of distance, climate and topography on beta diversity of mammalian faunas of North America in relation to latitude. Location North America north of Mexico. Methods The study area was divided into 313 equal-area quadrats (241 × 241 km). Faunal data for all continental mammals were compiled for these quadrats, which were divided among five latitudinal zones. These zones were comparable in terms of latitudinal and longitudinal span, climatic gradients and elevational gradients. We used the natural logarithm of the Jaccard index (ln J ) to measure species turnover between pairs of quadrats within each latitudinal zone. The slope of ln J in relation to distance was compared among latitudinal zones. We used partial regression to partition the variance in ln J into the components uniquely explained by distance and by environmental differences, as well as jointly by distance and environmental differences. Results Mammalian faunas of North America differ more from each other at lower latitudes than at higher latitudes. Regression models of ln J in relation to distance, climatic difference and topographic difference for each zone demonstrated that these variables have high explanatory power that diminishes with latitude. Beta diversity is higher for zones with higher mean annual temperature, lower seasonality of temperature and greater topographic complexity. For each latitudinal zone, distance and environmental differences explain a greater proportion of the variance in ln J than distance, climate or topography does separately. Main conclusions The latitudinal gradient in beta diversity of North American mammals corresponds to a macroclimatic gradient of decreasing mean annual temperature and increasing seasonality of temperature from south to north. Most of the variance in spatial turnover is explained by distance and environmental differences jointly rather than distance, climate or topography separately. The high predictive power of geographical distance, climatic conditions and topography on spatial turnover could result from the direct effects of physical limiting factors or from ecological and evolutionary processes that are also influenced by the geographical template.

Macroecology meets macroevolution: evolutionary niche dynamics in the seaweed Halimeda

Abstract: Aim Because of their broad distribution in geographical and ecological dimensions, seaweeds (marine macroalgae) offer great potential as models for marine biogeographical inquiry and exploration of the interface between macroecology and macroevolution. This study aims to characterize evolutionary niche dynamics in the common green seaweed genus Halimeda, use the observed insights to gain understanding of the biogeographical history of the genus and predict habitats that can be targeted for the discovery of species of special biogeographical interest. Location Tropical and subtropical coastal waters. Methods The evolutionary history of the genus is characterized using molecular phylogenetics and relaxed molecular clock analysis. Niche modelling is carried out with maximum entropy techniques and uses macroecological data derived from global satellite imagery. Evolutionary niche dynamics are inferred through application of ancestral character state estimation. Results A nearly comprehensive molecular phylogeny of the genus was inferred from a six-locus dataset. Macroecological niche models showed that species distribution ranges are considerably smaller than their potential ranges. We show strong phylogenetic signal in various macroecological niche features. Main conclusions The evolution of Halimeda is characterized by conservatism for tropical, nutrient-depleted habitats, yet one section of the genus managed to invade colder habitats multiple times independently. Niche models indicate that the restricted geographical ranges of Halimeda species are not due to habitat unsuitability, strengthening the case for dispersal limitation. Niche models identified hotspots of habitat suitability of Caribbean species in the eastern Pacific Ocean. We propose that these hotspots be targeted for discovery of new species separated from their Caribbean siblings since the Pliocene rise of the Central American Isthmus.

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients.

Abstract: Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km2) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km2) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R2= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R2= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low-altitude areas than in cold, high-altitude areas where human population density is low.

Characterizing and predicting species distributions across environments and scales: Argentine ant occurrences in the eye of the beholder

Abstract: Aim Species distribution models (SDMs) or, more specifically, ecological niche models (ENMs) are a useful and rapidly proliferating tool in ecology and global change biology. ENMs attempt to capture associations between a species and its environment and are often used to draw biological inferences, to predict potential occurrences in unoccupied regions and to forecast future distributions under environmental change. The accuracy of ENMs, however, hinges critically on the quality of occurrence data. ENMs often use haphazardly collected data rather than data collected across the full spectrum of existing environmental conditions. Moreover, it remains unclear how processes affecting ENM predictions operate at different spatial scales. The scale (i.e. grain size) of analysis may be dictated more by the sampling regime than by biologically meaningful processes. The aim of our study is to jointly quantify how issues relating to region and scale affect ENM predictions using an economically important and ecologically damaging invasive species, the Argentine ant ( Linepithema humile ). Location California, USA.

Pervasive effects of dispersal limitation on within- and among-community species richness in agricultural landscapes

Abstract: Aim To determine whether the effect of habitat fragmentation and habitat heterogeneity on species richness at different spatial scales depends on the dispersal ability of the species assemblages and if this results in nested species assemblages. Location Agricultural landscapes distributed over seven temperate Europe countries covering a range from France to Estonia. Methods We sampled 16 local communities in each of 24 agricultural landscapes (16 km(2)) that differ in the amount and heterogeneity of semi-natural habitat patches. Carabid beetles were used as model organisms as dispersal ability can easily be assessed on morphological traits. The proximity and heterogeneity of semi-natural patches within the landscape were related to average local (alpha), between local (beta) and landscape (gamma) species richness and compared among four guilds that differ in dispersal ability. Results For species assemblages with low dispersal ability, local diversity increased as the proximity of semi-natural habitat increased, while mobile species showed an opposite trend. Beta diversity decreased equally for all dispersal classes in relation to proximity, suggesting a homogenizing effect of increased patch isolation. In contrast, habitat diversity of the semi-natural patches affected beta diversity positively only for less mobile species, probably due to the low dispersal ability of specialist species. Species with low mobility that persisted in highly fragmented landscapes were consistently present in less fragmented ones, resulting in nested assemblages for this mobility class only. Main conclusions The incorporation of dispersal ability reveals that only local species assemblages with low dispersal ability show a decrease of richness as a result of fragmentation. This local species loss is compensated at least in part by an increase in species with high dispersal ability, which obscures the effect of fragmentation when investigated across dispersal groups. Conversely, fragmentation homogenizes the landscape fauna for all dispersal groups, which indicates the invasion of non-crop habitats by similar good dispersers across the whole landscape. Given that recolonization of low dispersers is unlikely, depletion of these species in modern agricultural landscapes appears temporally pervasive.

Can the tropical conservatism hypothesis explain temperate species richness patterns? An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini

Abstract: Aim A latitudinal gradient in species richness, defined as a decrease in biodiversity away from the equator, is one of the oldest known patterns in ecology and evolutionary biology. However, there are also many known cases of increasing poleward diversity, forming inverse latitudinal biodiversity gradients. As only three processes (speciation, extinction and dispersal) can directly affect species richness in areas, similar factors may be responsible for both classical (high tropical diversity) and inverse (high temperate diversity) gradients. Thus, a modified explanation for differential species richness which accounts for both patterns would be preferable to one which only explains high tropical biodiversity.

Climate and history explain the species richness peak at mid‐elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions

Abstract: Aim We studied elevational species richness patterns of Schizothorax fishes and identified the roles of ecological and evolutionary factors in shaping the patterns of elevational diversity. Location The Tibetan Plateau and its adjacent regions. Methods We assembled distribution and altitude data for all Schizothorax species using the literature. We merged ecological and evolutionary approaches to test the relationships between species richness and ecological factors (climate, area, the mid-domain effect) or evolutionary factors (diversification rates and time of colonization). Results We found that species richness of Schizothorax fishes peaked at mid-elevations. Rainfall, area, the mid-domain effect and diversification rate were weak predictors of the richness pattern. Temperature showed a nonlinear relationship with species richness. Temperature and time of colonization were the most important variables in explaining the elevational diversity pattern. Main conclusion Our findings indicate that the time-for-speciation effect and niche conservatism play important roles in variation of species richness.

Invasion success of alien plants: do habitat affinities in the native distribution range matter?

Abstract: Aim: To assess how habitat affinities in the native distribution range influence the invasion success of 282 central European neophytes (alien plants introduced after ad 1500). -- Location: Czech Republic. -- Methods: Classification trees were used to determine which native habitats donate the most alien species, the correspondence between habitats occupied by species in their native and invaded distribution ranges, and invasion success of species originating from different habitats. -- Results: The species most likely to naturalize in Central Europe are those associated with thermophile woodland fringes in their native range (81%), cultivated areas of gardens and parks (75%) and broad-leaved deciduous woodlands (72%). The largest proportions of invasive species recruit from those that occur on riverine terraces and eroded slopes, or grow in both deciduous woodland and riverine scrub. When the relative role of habitats in the native range is assessed as a determinant of the probability that a species will become invasive in concert with other factors (the species residence time, life history, region of origin), the direct effect of habitat is negligible. However, the effect of native habitats on patterns of invasions observed in central Europe is manifested by large differences in the numbers of species they supply to the invaded region. More than 50 neophytes were recruited from each of the following habitats: dry grasslands, ruderal habitats, deciduous woodland, inland cliffs, rock pavements and outcrops, and tall-herb fringes and meadows. -- Main conclusions: Casual species recruit from a wider range of habitats in their native range than they occupy in the invaded range; naturalized but not invasive species inhabit a comparable spectrum of habitats in both ranges, and successful invaders occupy a wider range of habitats in the invaded than in the native range. This supports the idea that the invasive phase of the process is associated with changes in biological features that allow for extension of the spectrum of habitats invaded.

Effects of species traits on the genetic diversity of high‐mountain plants: a multi‐species study across the Alps and the Carpathians

Abstract: To test the influence of various species traits, elevation and phylogeographical history on the genetic diversity of high-mountain plants in the Alps and Carpathians. The regular sampling grid comprised the whole range of the European Alps and the Carpathians. Twenty-two high-mountain plant species were exhaustively sampled and their genetic diversity was assessed with amplified fragment length polymorphisms (AFLPs). ANOVAs were used to check for relationships between species traits and species genetic diversity, and to test whether genetic diversity was influenced by altitude and phylogeographical history (i.e. Alps versus Carpathians). In both mountain systems, species dispersed and pollinated by wind showed higher genetic diversity than species with self or insect pollination, and with animal- or gravity-dispersed seeds. Only in the Alps did altitudinal range size affect species genetic diversity significantly: species with narrow altitudinal ranges in the highest vegetation belts had significantly higher genetic diversity than those expanding over wide altitudinal ranges. Genetic diversity was species specific and significantly higher in the Alps than in the Carpathians, but it was not influenced by elevation. Wind pollination and wind dispersal seem to foster high genetic diversity. However, species traits are often associated and their effects on genetic diversity cannot be clearly disentangled. As genetic diversity is species specific, comparisons across species need to be interpreted with care. Genetic diversity was generally lower in the Carpathians than in the Alps, due to higher topographical isolation of alpine habitats in the Carpathians and this mountain massif's divergent phylogeographical history. Elevation did not influence genetic diversity, challenging the long-held view of decreasing genetic diversity with increasing elevation in mountain plants.

Threshold response of Madagascar's littoral forest to sea‐level rise

Abstract: Aim Coastal biodiversity hotspots are globally threatened by sea-level rise. As such it is important to understand how ecosystems resist, respond and adapt to sea-level rise. Using pollen, geochemistry, charcoal and diatom records in conjunction with previously published palaeoclimatic records, we investigated the mechanism, interactions and ecosystem response and resilience of Madagascar’s littoral forest to late Holocene sea-level rise. Location Sediment sequences were collected along the south-east coast of Madagascar in two adjacent habitats in Mandena; the highly diverse littoral forest fragment and species-poor Erica -matrix. Methods We used a multi-proxy approach to investigate the relative influence of environmental changes on the littoral ecosystem. We reconstructed past vegetation and fire dynamics over the past 6500 years at two sites in the littoral forest using fossil pollen and macrofossil charcoal contained in sedimentary sequences. Alongside these records we reconstructed past marine transgressions from the same sedimentary sequences using geochemical analyses, and a salinity and drought index through the analysis of fossil diatoms. Results Our findings indicated that it was the synergistic effect of sea-level rise coupled with rainfall deficits that triggered a threshold event with a switch from two types of littoral forest (an open Uapaca forest and a closed littoral forest fragment) to an Erica‐Myrica heath/grassland occurring in approximately less than 100 years. Resilience to sea-level rise differed in the two adjacent habitats, suggesting that the littoral forest fragment was more resilient to the impacts of sea-level change and aridity than the open Uapaca woodland. Conclusions We demonstrated that the littoral ecosystem was influenced by late Holocene sea-level rise and climatic desiccation. While climate change-integrated conservation strategies address the effects of climate change on species distribution and dispersal, our work suggests that more attention should be paid to the impacts of interactive climatic variables that affect ecosystem thresholds.

Origin of the lichen–spruce woodland in the closed‐crown forest zone of eastern Canada

Abstract: Aim We investigate the timing and factors responsible for the transformation of closed-crown forests into lichen‐spruce woodlands. Location The study area extends between 70 ° and 72 ° W in the closed-crown forest zone from its southern limit near 47 ° 30 ′ N to its northern limit at the contact with the lichen‐spruce woodland zone around 52 ° 10 ′ N. A total of 24 lichen‐spruce woodlands were selected. Methods Radiocarbon dating of charcoals at mineral soil contact and within the organic horizons allowed the principal factors causing the degradation of the closedcrown forest to be identified, i.e. light fires, successive fires and the occurrence of a spruce budworm epidemic followed by a fire. Results Charcoals dated in the organic horizon were less than 200 years old, suggesting a recent transformation of the closed-crown forest following surface fires. Before their transformation into lichen‐spruce woodlands, stands were occupied by old, dense forests that originated from fires dating back to 1000 yr bp . The radiocarbon dating of charcoals in the organic horizon indicated that several stands burned twice in less than 50 years, while others burned shortly after a spruce budworm epidemic. Light fires are frequent within the lichen‐spruce woodlands according to multiple charcoal layers found within the organic matter horizon. Main conclusions While closed-crown forests are predicted to expand under climate warming, compound disturbances diminish the natural regeneration of the closed-crown forests in the south and favour the expansion of lichen‐spruce woodlands. As black spruce germinates on mineral soils, surface fires accentuate the expansion of the lichen‐spruce woodlands southward. Under global warming, warmer springs will lead to earlier low-intensity fires that do not remove as much organic matter, and hence prevent conditions suitable for black spruce regeneration. Also, spruce budworm reduces seed production for a certain time. The occurrence of fire during this period is critical for regeneration of black spruce.

Combining spatial and phylogenetic eigenvector filtering in trait analysis

Abstract: Aim To analyse the effects of simultaneously using spatial and phylogenetic information in removing spatial autocorrelation of residuals within a multiple regression framework of trait analysis. Location Switzerland, Europe. Methods We used an eigenvector filtering approach to analyse the relationship between spatial distribution of a trait (flowering phenology) and environmental covariates in a multiple regression framework. Eigenvector filters were calculated from ordinations of distance matrices. Distance matrices were either based on pure spatial information, pure phylogenetic information or spatially structured phylogenetic information. In the multiple regression, those filters were selected which best reduced Moran’s I coefficient of residual autocorrelation. These were added as covariates to a regression model of environmental variables explaining trait

Changes in the onset of spring growth in shrubland species in response to experimental warming along a north–south gradient in Europe

Abstract: Aim To test whether the onset of spring growth in European shrublands is advanced in response to the warmer conditions projected for the next two decades by climate models, and, if there is a change, whether it differs across Europe. Location The studied sites spanned a broad north–south European gradient with average annual temperatures (8.2–15.6 °C) and precipitation (511–1427 mm). Methods 'Bud break' was monitored in eight shrub and grass species in six European sites under control and experimentally warmer conditions generated by automatic roofs covering vegetation during the night. Results Species responsive to increased temperatures were Vaccinium myrtillus and Empetrum nigrum in Wales, Deschampsia flexuosa in Denmark, Calluna vulgaris in Netherlands, Populus alba in Hungary and Erica multiflora in Spain. Although the acceleration of spring growth was the commonest response to warming treatments, the responses at each site were species specific and year dependent. Under experimental warming 25% of cases exhibited a significantly earlier onset of the growing season and 10% had a significantly delayed onset of vegetative growth. No geographical gradient was detected in the experimental warming effects. However, there was a trend towards a greater dominance of phenological advances with more intense the warming treatments. Above 0.8 °C warming, only advancements were recorded. Main conclusions Our results show that warmer temperatures projected for the next decades have substantial potential effects on the phenology of the spring growth of dominant species in different European shrublands, with a dominant trend towards advancements the more intense the warming is. However, our study also demonstrates the overall difficulties of applying simple predictive relationships to extrapolate the effects of global change on phenology. Various combinations of environmental factors occur concurrently at different European sites and the interactions between different drivers (e.g. water and chilling) can alter phenology significantly.