Showing papers in "Ecography in 2004"

Presence-absence versus presence-only modelling methods for predicting bird habitat suitability

Abstract: Habitat suitability models can be generated using methods requiring information on species presence or species presence and absence. Knowledge of the predictive performance of such methods becomes a critical issue to establish their optimal scope of application for mapping current species distributions under different constraints. Here, we use breeding bird atlas data in Catalonia as a working example and attempt to analyse the relative performance of two methods: the Ecological Niche factor Analysis (ENFA) using presence data only and Generalised Linear Models (GLM) using presence/absence data. Models were run on a set of forest species with similar habitat requirements, but with varying occurrence rates (prevalence) and niche positions (marginality). Our results support the idea that GLM predictions are more accurate than those obtained with ENFA. This was particularly true when species were using available habitats proportionally to their suitability, making absence data reliable and useful to enhance model calibration. Species marginality in niche space was also correlated to predictive accuracy, i.e. species with less restricted ecological requirements were modelled less accurately than species with more restricted requirements. This pattern was irrespective of the method employed. Models for wide-ranging and tolerant species were more sensitive to absence data, suggesting that presence/absence methods may be particularly important for predicting distributions of this type of species. We conclude that modellers should consider that species ecological characteristics are critical in determining the accuracy of models and that it is difficult to predict generalist species distributions accurately and this is independent of the method used. Being based on distinct approaches regarding adjustment to data and data quality, habitat distribution modelling methods cover different application areas, making it difficult to identify one that should be universally applicable. Our results suggest however, that if absence data is available, methods using this information should be preferably used in most situations.

Effects of restricting environmental range of data to project current and future species distributions

Abstract: We examine the consequences of restricting the range of environmental conditions over which niche-based models are developed to project potential future distributions of three selected European tree species to assess first, the importance of removing absences beyond species known distributions (‘‘naughty noughts’’) and second the importance of capturing the full environmental range of species. We found that restricting the environmental range of data strongly influenced the estimation of response curves, especially towards upper and lower ends of environmental ranges. This induces changes in the probability values towards upper and lower environmental boundaries, leading to more conservative scenarios in terms of changes in distribution projections. Using restricted data analogous to not capturing the fun species’ environmental range, reduces strongly the combinations of environmental conditions under which the models are calibrated, and reduces the applicability of the models for predictive purposes. This may generate unpredictable effects on the tails of the species response curves, yielding spurious projections into the future provided that probability of occurrence is not set to zero outside the environmental limits of the species. Indeed, as the restricted data does not capture the whole of the response curve, projections of future species distributions based of ecological niche modelling may be only valid if niche models are able to approach the complete response curve of environmental predictors.

Modelling species distributions in Britain: a hierarchical integration of climate and land-cover data

Abstract: modelling framework for studying the combined effects of climate and land-cover changes on the distribution of species is presented. The model integrates land-cover data into a correlative bioclimatic model in a scale-dependent hierarchical manner, whereby Artificial Neural Networks are used to characterise species' climatic requirements at the European scale and land-cover requirements at the British scale. The model has been tested against an alternative non-hierarchical approach and has been applied to four plant species in Britain: Rhynchospora alba, Erica tetralix, Salix herbacea and Geranium sylvaticum. Predictive performance has been evaluated using Cohen's Kappa statistic and the area under the Receiver Operating Characteristic curve, and a novel approach to identifying thresholds of occurrence which utilises three levels of confidence has been applied. Results demonstrate reasonable to good predictive performance for each species, with the main patterns of distribution simulated at both 10 km and 1 km resolutions. The incorporation of land-cover data was found to significantly improve purely climate-driven predictions for R. alba and E. tetralix, enabling regions with suitable climate but unsuitable land-cover to be identified. The study thus provides an insight into the roles of climate and land-cover as determinants of species' distributions and it is demonstrated that the modelling approach presented can provide a useful framework for making predictions of distributions under scenarios of changing climate and land-cover type. The paper confirms the potential utility of multi-scale approaches for understanding environmental limitations to species' distributions, and demonstrates that the search for environmental correlates with species' distributions must be addressed at an appropriate spatial scale. Our study contributes to the mounting evidence that hierarchical schemes are characteristic of ecological systems.

A local nearest-neighbor convex-hull construction of home ranges and utilization distributions

Abstract: We describe a new method for estimating the area of home ranges and constructing utilization distributions (UDs) from spatial data. We compare our method with bivariate kernel and alpha-hull methods, using both randomly distributed and highly aggregated data to test the accuracy of area estimates and UD isopleth construction. The data variously contain holes, corners, and corridors linking high use areas. Our method is based on taking the union of the minimum convex polygons (MCP) associated with the k-1 nearest neighbors of each point in the data and, as such, has one free parameter k. We propose a minimum spurious hole covering (MSHC) rule for selecting k and interpret its application in terms of type I and type II statistical errors. Our MSHC rule provides estimates within 12% of true area values for all 5 data sets, while kernel methods are worse in all cases: in one case overestimating area by a factor of 10 and in another case underestimating area by a factor of 50. Our method also constructs much better estimates for the density isopleths of the UDs than kernel methods. The alpha-hull method does not lead directly to the construction of isopleths and also does not always include all points in the constructed home range. Finally we demonstrate that kernel methods, unlike our method and the alpha-hull method, does not converges to the true area represented by the data as the number of data points increase.

Effects of habitat complexity on ant assemblages

Abstract: We investigated responses of ant communities to habitat complexity, with the aim of assessing complexity as a useful surrogate for ant species diversity. We used pitfall traps to sample ants at twenty-eight sites, fourteen each of low and high habitat complexity, spread over ca 12 km in Sydney sandstone ridge-top woodland in Australia. Ant species richness was higher in low complexity areas, and negatively associated with ground herb cover, tree canopy cover, soil moisture and leaf litter. Ant community composition was affected by habitat complexity, with morphospecies from the genera Monomorium, Rhytidoponera and Meranoplus being the most significant contributors to compositional differences. Functional group responses to anthropogenic disturbance may be facilitated by local changes in habitat complexity. Habitat complexity, measured as a function of differences in multiple strata in forests, may be of great worth as a surrogate for the diversity of a range of arthropod groups including ants.

Potential impact of climatic change on the distribution of forest herbs in Europe

Abstract: The aim of this study is to evaluate the possible consequences of climate change on a representative sample of forest herbs in Europe. A fuzzy climatic envelope was used to predict the location of suitable climatic conditions under two climatic change scenarios. Expected consequences in terms of lost and gained range size and shift in distribution for 26 forest herbs were estimated. These results were combined in an Index of Predicted Range Change for each species. Finally, the effects of habitat fragmentation for potential dispersal routes were evaluated and options for management on a European scale are discussed. Generally, a good agreement of the estimated suitability under the present climate and the observed current distribution was observed. However, species vary a lot in the degree to which they occupy the presently climatically suitable areas in Europe. Many species are absent from large areas with suitable climate and thus could be said to have poor range-filling capacity. A general change in location (range centroid) of the total suitable area was observed: The total suitable area will on average move strongly northwards and moderately eastwards under the relatively mild B1 scenario and more strongly so under the A2 scenario. The required average minimum migration rate per year to track the potential range shift is 2.1 km under the B1 scenario and 3.9 km under the A2 scenario. Moderate losses in the total suitable area in Europe are predicted for most species under both scenarios. However, the predicted changes are very variable, with one species (Actaea erythrocarpa) experiencing total range elimination in Europe (A2 scenario) while the total suitable area is predicted to show large increases for other species. The species that are predicted to experience the greatest proportional losses in their climatically suitable area within their presently realised range tend to have northern or eastern range centroids. The Index of Predicted Range Change roughly divides the species studied in four groups: One species face a high risk of extinction; eight species are expected to experience moderate to severe threat of extinction; 11 species are not considered at risk and, finally, six species may actually benefit from global warming. An analysis of potential migration routes shows the importance of maintaining and, if possible, improving the network of forest throughout Europe to make migration possible. It is also suggested to closely monitor the status of boreal and subalpine species that are most threatened by global warming. Finally it is recommended that special concern should be given to increased protection and restoration of forest habitats in southern montane areas for their crucial long-term importance for the maintenance of European plant diversity.

‘Latitude’ and geographic patterns in species richness

Abstract: FORUM is a lighter channel of communication between readers and contributors; it aims to stimulate discussion and debate, particularly by presenting new ideas and by suggesting alternative interpretations to the more formal research papers published in ECOGRAPHY and elsewhere. A lighter prose is encouraged and no summary is required. Contributions should be concise and to the point, with a relatively short bibliography. Formal research papers, however short, will not be considered.

Biological trait composition of European stream invertebrate communities: assessing the effects of various trait filter types

Abstract: Understanding the action of filters on the biological trait composition of communities is constrained by the multitude of filter types (e.g. abiotic vs biotic, actual vs historical) that may cause changes of a multitude of traits (e.g. small vs large body size, short vs long life cycle) at a multitude of spatial scales (e.g. continent vs landscape vs local site). Using published data on the as natural as possible abundances and 11 biological traits (described through 63 categories) of 254 European stream invertebrate genera, we assessed how already available knowledge can serve to identify the importance of the action of different types of trait filters at two spatial scales. Therefore, we analysed observed and simulated abundance-weighted trait compositions at the local scale of 384 running water sites (RWS) and at the landscape scale of 14 large biogeographical regions (LER). Actual abiotic filters acted significantly and independently of the taxonomic richness on the invertebrate traits at the RWS- and LER-scale, whereas biotic filters had no significant effect. Evidence for the action of historical trait filters across Europe was only weak at both scales. Size, reproductive cycle, respiration and locomotion technique, feeding habits and vulnerability to disturbance responded to altitude and stream width of the RWS according to existing views about the effects of riparian, physiological, interstitial or disturbance controls of these traits. These controls acted independently on trait categories that did not co-occur within the genera, because correlations of size categories with other trait categories were higher in the abundance-weighted trait array (across communities) than in the original trait array (across genera). Overall, many of the 63 trait categories were scarcely affected by the trait filters considered in this study. Therefore, we briefly discuss potential effects of continental filters and of stream system-specific, local physical filters, as the latter should produce similar trait patterns on a global scale. Our study suggests that analyses of the currently available knowledge can simplify the complicated hypothetical framework on trait filter actions, which sharpens the focus on future research needs.

Benthic diatom communities in boreal streams: community structure in relation to environmental and spatial gradients

Abstract: An important goal for community ecology is the characterization and prediction of changes in community patterns along environmental gradients. We aimed to identify the major environmental correlates of diatom distribution patterns in boreal running waters. We classified 197 stream sites based on their diatom flora. Direct ordination methods were then used to identify the key environmental determinants of this diatom-based stream typology. Finally, we tested whether a regional classification scheme based on terrestrial landscapes (ecoregions) provides a reasonable framework for a regional grouping of streams based on their diatom flora. Two-way indicator species analysis produced 13 site groups, which were primarily separated by chemical variables, mainly conductivity, total P and water colour. In partial CCA, the environmental and spatial factors accounted for 38% and 24%, respectively, of explained variation in community composition. A high proportion (almost 40%) of variation explained by the combined effect (spatially-structured environmental) indicated that diatom communities of boreal streams incorporate a strong spatial component. At the level of subecoregions, classification strength was almost equally strong for all sites as for near-pristine reference sites only. Procrustes analysis indicated that spatial factors and patterns in diatom community structure were strongly concordant. Our data support the argument that diatom communities are strongly spatially structured, with distinctly different communities in different parts of the country. Because of the strong spatial patterns of community composition, bioassessment programs utilising lotic diatoms would clearly benefit from regional stratification. A combination of regional stratification and the prediction of assemblage structure from local environmental features might provide the most robust framework for diatom-based assessment of the biological integrity of boreal streams.

Bergmann's rule and the mammal fauna of northern North America

Abstract: animal body mass might show such spatial variation, and indeed whether it is a general feature of animal assemblages, is currently unclear. We discuss reasons for this uncertainty, and use our conclusions to direct an analysis of Bergmann's rule in the mammals in northern North America, in the communities of species occupying areas that were covered by ice at the last glacial maximum. First, we test for the existence of Bergmann's rule in this assemblage, and investigate whether small- and large-bodied species show different spatial patterns of body size variation. We then attempt to explain the spatial variation in terms of environmental variation, and evaluate the adequacy of our analyses to account for the spatial pattern using the residuals arising from our environmental models. Finally, we use the results of these models to test predictions of different hypotheses proposed to account for Bergmann's rule. Bergmann's rule is strongly supported. Both small- and large-bodied species exhibit the rule. Our environmental models account for most of the spatial variation in mean, minimum and maximum body mass in this assemblage. Our results falsify predictions of hypotheses relating to migration ability and random colonisation and diversification, but support predictions of hypotheses relating to both heat conservation and starvation resistance.

Vertebrate herbivores and ecosystem control: cascading effects of faeces on tundra ecosystems

Abstract: We tested the hypothesis that large herbivores manipulate their own food supply by modifying soil nutrient availability. This was investigated experimentally the impact of faeces on grasses, mosses and soil biological properties in tundra ecosystems. For this, we increased the density of reindeer Rangifer tarandus platyrhynchus faeces and studied the response of a tundra system on Spitsbergen to this single faecal addition treatment for four subsequent years. From the third year onwards faecal addition had unambiguously enhanced the standing crop of grasses, as evidenced by an increase in both shoot density and mass per shoot. Although reindeer grazing across experimental plots was positively related to the abundance of grasses in anyone year, the increase in grass abundance in fouled plots failed to result in greater grazing pressure in those plots. Faecal addition enhanced soil microbial biomass C and N, particularly under wet conditions where faecal decay rates were greatest, whilst grasses appeared to benefit from faeces under dry conditions. Whilst growth of grasses and soil microbial biomass were stimulated by faecal addition, the depth of the extensive moss layer that is typical of tundra ecosystems was significantly reduced in fouled plots four years after faecal addition. The greatest reduction in moss depth occurred where fouling increased soil microbial biomass most, suggesting that enhanced decomposition of moss by a more abundant microbial community may have caused the reduced moss layer depth in fouled plots. Our field experiment demonstrates that by the production of faeces alone, vertebrate herbivores greatly impact on both above- and belowground components of tundra ecosystems and in doing so manipulate their own food supply. Our findings verify the assertion that grazing is of fundamental importance to tundra ecosystem productivity, and support the hypothesis that herbivory is instrumental in promoting grasses whilst suppressing mosses. The widely observed inverse relationship between grass and moss abundance in the field may therefore reflect the long history of plant-herbivore interactions in tundra ecosystems.

Arthropod biodiversity after forest fires: winners and losers in the winter fire regime of the southern Alps

Abstract: Since prehistoric times, natural and man made fires have been important factors of natural disturbance in many forest ecosystems, like those on the southern slopes of the Alps. Their effect on scarce, endangered or stenotopic species and on the diversity of invertebrate species assemblages which depend on a mosaic of successional habitat stages, is controversially discussed. In southern Switzerland, in a region affected by regular winter fires, we investigated the effect of the fire frequency on a large spectrum of taxonomic groups. We focussed on total biodiversity, taxonomic groups specific to certain habitat types, and on scarce and endangered species. Overall species richness was significantly higher in plots with repeated fires than in the unburnt control sites. Plots with only one fire in the last 30 yr harboured intermediate species numbers. Fire frequency had a significantly positive effect on species richness of the guilds of interior forest species and forest edge specialists. Species of open landscape, open forests and interior forests were not influenced by fire frequency. A positive effect of fire on species richness was observed for ground beetles (Carabidae), hoverflies (Syrphidae), bees and wasps (Hymenoptera aculeata, without Formicidae), and spiders (Araneae). True bugs (Heteroptera), lacewings (Neuroptera) and the saproxylic beetle families Cerambycidae, Buprestidae and Lucanidae showed positive trends, but no statistically significant effects of fire on species numbers or/and abundances. Negative effects of fire on species numbers or/and abundances were found only for isopods and weevils (Curculionidae). A compromise for forest management is suggested, which considers the risk of damage by fire to people and goods, while avoiding the risk of damage to biodiversity by imitating the effects of sporadic fires and providing a mosaic forest with open gaps of different successional stages.

Landscape effects on butterfly assemblages in an agricultural region

Abstract: We examined the butterfly fauna at 62 sites in southeastern Sweden within a region exhibiting high variation in the landscape surrounding the studied grasslands. The landscape varied from an intensively-managed agricultural landscape with a large amount of open fields to a landscape with a high amount of deciduous forest/semi-natural grassland. We made 12 179 observations of 57 species of butterflies. The amount of neighbouring deciduous forest/semi-natural grassland, with >25% tree and bush cover, was the most important environmental factor explaining the variation in the butterfly assemblages. Landscape analyses at three different spatial scales showed that the variation in butterfly assemblages could be explained only at the largest scale (radius 5000 m) and not at the smaller ones (radii 500 and 2000 m). Logistic regressions were used to predict presence/absence of butterfly species. Our study indicated that there may be critical thresholds for the amount of habitat at the landscape scale for several butterfly species as well as for species richness. For Melitaea athalia, there was a sharp increase in occupancy probability between 3 and 10% deciduous forests/semi-natural grasslands at the 5000-m scale. For 12 other species, the value for 50% probability of occurrence varied between 2 and 12% deciduous forest/semi-natural grassland. Species which had high occupancy probabilities in landscapes with a low amount of surrounding deciduous forests/semi-natural grasslands were significantly more mobile than others. Our study highlights the importance of applying a landscape perspective in conservation management, and that single-patch management might fail in maintaining a diverse butterfly assemblage.

Analysis of determinants of mammalian species richness in South America using spatial autoregressive models

Abstract: Classically, hypotheses concerning the distribution of species have been explored by evaluating the relationship between species richness and environmental variables using ordinary least squares (OLS) regression. However, environmental and ecological data generally show spatial autocorrelation, thus violating the assumption of independently distributed errors. When spatial autocorrelation exists, an alternative is to use autoregressive models that assume spatially autocorrelated errors. We examined the relationship between mammalian species richness in South America and environmental variables, thereby evaluating the relative importance of four competing hypotheses to explain mammalian species richness. Additionally, we compared the results of ordinary least squares (OLS) regression and spatial autoregressive models using Conditional and Simultaneous Autoregressive (CAR and SAR, respectively) models. Variables associated with productivity were the most important at determining mammalian species richness at the scale analyzed. Whereas OLS residuals between species richness and environmental variables were strongly autocorrelated, those from autoregressive models showed less spatial autocorrelation, particularly the SAR model, indicating its suitability for these data. Autoregressive models also fit the data better than the OLS model (increasing R2 by 5-14%), and the relative importance of the explanatory variables shifted under CAR and SAR models. These analyses underscore the importance of controlling for spatial autocorrelation in biogeographical studies.

Contrasting effects of habitat quantity and quality on moth communities in fragmented landscapes

Abstract: Habitat loss is commonly identified as a major threat to the loss of global biodiversity. In this study, we expand on our previous work by addressing the question of how lepidopteran species richness and composition vary among remnants of North American eastern deciduous forest located within agricultural or pastoral landscapes. Specifically, we tested the relative roles of habitat quantity (measured as stand area and percent forest in the greater landscape) and habitat quality (measured as tree species diversity) as determinants of moth species richness. We sampled > 19 000 individuals comprising 493 moth species from 21 forest sites in two forested ecoregions. In the unglaciated Western Allegheny Plateau, the species richness of moths with woody host plants diminished as forest stand size and percent forest in the landscape decreased, but the total species richness and abundance of moths were unaffected by stand size, percent forest in the landscape, or tree species diversity. In contrast, the overall species richness and abundance of moths in the glaciated North Central Tillplain were affected primarily by tree species diversity and secondarily by forest size. Higher tree species diversity may reduce species loss from smaller forest stands, suggesting that small, diverse forests can support comparable numbers of species to those in less diverse, large stands. Smaller forests, however, contained a disproportionate number of moth species that possess larvae known to feed on herbaceous vegetation. Thus, although woody plant feeding moths are lost from forests with changes in stand area, new species appear capable of recolonizing smaller fragments from the surrounding habitat matrix. Our study further suggests that when species replacement occurs, local patch size and habitat quality may be more important than landscape context in determining the community structure of forest Lepidoptera.

Primary productivity and species richness: relationships among functional guilds, residency groups and vagility classes at multiple spatial scales

Abstract: One of the major determinants of species richness is the amount of energy available, often measured as primary productivity. Heterogeneity of environmental variables has also been found to influence species richness. Predicting species distributions across landscapes and identifying areas that have high species richness, or vulnerable groups of species, is useful for land management. Remotely sensed data may help identify such areas, with the Normalized Difference Vegetation Index (NDVI) providing an estimate of primary productivity. We examined the relationship between maximum productivity (NDVI), heterogeneity of productivity, and species richness of birds and butterflies at multiple spatial scales. We also explored relationships between productivity, functional guilds and residency groups of birds, and vagility classes of butterflies. Positive linear relationships between maximum NDVI and number of functional guilds of birds were found at two spatial scales. We also found positive linear relationships between maximum NDVI and species richness of neotropical migrant birds at two scales. Heterogeneity of NDVI, by contrast, was negatively associated with number of functional guilds of birds and species richness of resident birds. Maximum NDVI was associated with species richness of all butterflies and of the most vagile butterflies. No association was found between heterogeneity of NDVI and species richness of butterflies. In the Great Basin, where high greenness and availability of water correspond to areas of high species richness and maximum NDVI, our results suggest that NDVI can provide a reliable basis for stratifying surveys of biodiversity, by highlighting areas of potentially high biodiversity across large areas. Measures of heterogeneity of NDVI appear to be less useful in explaining species richness.

Patterns of resource tracking by avian frugivores at multiple spatial scales: two case studies on discordance among scales

Abstract: Scaling is relevant for the analysis of plant-frugivore interaction, since the ecological and evolutionary outcomes of seed dispersal depend on the spatial and temporal scale at which frugivory patterns emerge. We analyse the relationship between fruit abundance and frugivore activity at local and landscape spatial scales in two different systems composed, respectively, by the bird-dispersed woody plants Juniperus communis and Bursera fagaroides, and their frugivore assemblages. We use a hierarchical approach of nested patchiness of fruit-resource, where patches are defined by individual plants within site, at the local scale, and by sites within region, at the landscape scale. The structure of patches is also described in terms of contrast (differences in fruit availability among patches) and aggregation (spatial distribution of patches). For J. communis, frugivore activity was positively related to fruit availability at the landscape scale, this pattern seldom emerging at the local scale; conversely, B. fagaroides showed a general trend of positive local pattern that disappeared at the landscape scale. These particular trends might be partially explained by differences in contrast and aggregation. The strong contrast among plants within site together with a high aggregation among sites would promote the B. fagaroides pattern to be only local, whereas in J. communis, low aggregation among sites within region would favour a sharp landscape-scale pattern. Both systems showed discordant patterns of fruitresource tracking among consecutive spatial scales, but the sense of discordance differed among systems. These results, and the available multi-scale frugivory data, suggest that discordance among successive scales allows to link directly frugivory patterns to resource-tracking mechanisms acting at particular scales, resulting, thus, more informative than concordance observational data, in which landscape patterns might result from accumulated effect of local mechanisms. In this context, we propose new methodological approaches for a better understanding of the hierarchical behavioural mechanisms underpinning the multi-scale resource tracking by frugivores.

Distribution pattern of the epiphyte Neckera pennata on three spatial scales – importance of past landscape structure, connectivity and local conditions

Abstract: We tested which factors explain the distribution pattern of the epiphytic moss Neckera pennata on three spatial scales using the framework of generalized linear models. First, we tested which factors explained its occurrence in forest stands in a 2500 ha landscape. At this scale, we also tested the effect of the historic landscape structure. We recorded its occurrence in all suitable stands. The occurrence probability increased with increasing present quantity of Acer platanoides, and with increasing present and past quantity of Fraxinus excelsior. The probability also increased with increasing connectivity to occupied stands. However, the connectivity to stands present in 1977 (recorded from infra-red aerial photographs) explained more of the variation. This suggests that the regional metapopulation size of N. pennata has decreased during the past decades, and that its present distribution pattern reflects the age of the remaining stands, and the distribution of past dispersal sources in the landscape. Second, we tested which factors explained the occurrence and abundance on individual trees in three forest stands. Neckera pennata mainly occurred on Acer and Fraxinus stems. The most important variable in explaining occurrence probability was connectivity to surrounding occupied trees, which probably reflects the restricted dispersal range in this species. The abundance on occupied trees was also explained by this variable. The occurrence probability and abundance also increased with increasing tree diameter, probably reflecting the time that a tree has been available for colonization and the time since colonization, respectively. The occurrence probability and abundance furthermore decreased on strongly leaning (and deteriorating) trees. The occurrence probability increased with increasing bark roughness, probably reflecting increasing suitability regarding bark chemistry and moisture. Third, we tested its vertical distribution on occupied trees. The main distribution was below 1.6 m.

Elevational gradients, area and tropical island diversity: an example from the palms of New Guinea

Abstract: The factors causing spatial variation in species richness remain poorly known. In this study, factors affecting species richness of palms (Palmae/Arecaceae) were studied along the elevational gradient of New Guinea. Interpolated elevational ranges were calculated from a database of all known collections for 145 species in 32 genera. The amount of land area at different elevations greatly affects the species richness gradient. If assessed in equal-elevation bands species richness appears to decline monotonically, but when assessed in equal-area bands species richness shows a pronounced mid-elevation peak, due to the large proportion of lowlands in New Guinea. By randomising species ranges within the total elevational gradient for palms and accounting for area, we found the mid-elevation peak to be consistent with a mid-domain effect caused by the upper and lower limits to palm distribution. Our study illustrates the importance of accounting for area in macroecological studies of richness gradients and introduces a novel yet simple method for doing this through the use of equal-area bands. Together, the effect of area and the mid-domain effect explain the majority of variation in species richness of New Guinea palms. We support calls for the multivariate assessment of the mid-domain effect on an equal footing with other potential explanations of species richness.

Environmental and spatial variables influence the composition of frog assemblages in sub-tropical eastern Australia

Abstract: In community ecology, contrasting theories suggest that the distribution and abundance of species, and thus the composition of assemblages, are influenced by i) environmental gradients, or ii) contagious biotic processes such as predation, competition, dispersal and disease. In the former case, sites with similar environments would tend to support similar assemblages, while in the latter, geographically proximate sites would tend to support more similar assemblages than widely separated sites. I investigated the relative influence of environmental variables and spatial position on the composition of frog assemblages at forest streams in sub-tropical eastern Australia using redundancy analysis (RDA) and partial RDA. Data on the maximum abundance of the frog species at 65 survey sites were transformed such that RDA would yield the Hellinger distance between sites. The following analysis identified 11 environmental variables that explained 45% of the variation in the abundance of species at the survey sites (the species matrix), as a proportion of total variance. The geographic co-ordinates of the survey sites accounted for 12%, while the environmental and spatial variables combined accounted for 47% of the variation in the species matrix. Partial redundancy analysis indicated that of the explained variation, 74% was purely environmental, 5% was purely spatial and 21% was spatial environmental variation. This study is the first to quantify the relative influence of environmental and spatial variables on the composition of amphibian assemblages. It provides support for both the environmental control model and the biotic control model of species' distributions and assemblage composition, although environmental variables appear to have the greater effect at this scale of analysis.

Plant responses to livestock grazing frequency in an Australian temperate grassland

Abstract: Livestock grazing is often thought to enhance native plant species co-existence in remnant grasslands but may also favour exotic invaders. Recommendations for appropriate grazing strategies are needed, for which an understanding of the response of plant species is necessary. We explored the response of plant species and plant functional groups to grazing in temperate grassland of the Monaro Tablelands of south-east Australia by comparing species abundance in adjacent areas that differed in livestock grazing regime (minimal, infrequent and frequent). We also examined whether species with similar responses to grazing share certain traits and consider whether these traits might provide a useful method of assessing grazing impact. At the scale measured (0.25 m 2- ), an infrequent grazing regime maximised plant species co-existence in these grasslands due to widespread invasion by exotic plant species at infrequent grazing intensity. Many native species declined in abundance when grazing frequency increased from minimal to infrequent. Annuals invaded under infrequent grazing while perennials declined most strongly under high frequency grazing. Low levels of grazing apparently reduce cover and create sites suitable for seed recruitment whereas more frequent grazing reduces the persistence of perennials. While there was a tendency for native species to be more susceptible to grazing impact than exotics, plant traits, in particular longevity (perennial, annual) provided a better prediction of the response of plants to grazing. Although a few native plant species persisted at high grazing frequency, even infrequent livestock grazing may not be appropriate for the conservation of many native perennial grassland species. Targeted reductions in grazing frequency may be necessary to enable the long-term coexistence of grazing susceptible species.

Lasting effects of conditions at birth on moose body mass

Abstract: Whether cohort effects can be retrieved in adult phenotypes depends on the possibility for individuals to compensate for a good or bad start in life. This ability to compensate may itself depend on the environment and on individual sex. In large polygynous ungulates, male reproductive success relies more on body size than the reproductive success of females, which makes them more sensitive to a bad start in life. Based on current theories of life history evolution and sexual selection, we tested the following predictions in a moose population: 1) cohort effects and year effects occur in both male and female adult body mass, but due to 2) compensatory growth, cohort effects tend to fade away with the individual's age; and 3) males are more sensitive to cohort effects than females. In support of the first prediction, we found that density and climate during the year-of-birth and the year-of-harvest affected moose body mass in both sexes. However, the magnitude of the effects of environmental conditions at birth on adult body mass decreased with increasing age, but less so in males than in females. Thus, as expected based on our third prediction, environmental conditions early in life were more influential on adult body mass of males compared to females. Such a result supports the existence of sex-specific life history tactics, with males maximising growth rate and females rather trading-off growth for reproduction.

Evaluating alternative data sets for ecological niche models of birds in the Andes

Abstract: Ecological niche modeling (ENM) is an effective tool for providing innovative insights to questions in evolution, ecology and conservation. As environmental datasets accumulate, modelers need to evaluate the relative merit of different types of data for ENM. We used three alternative environmental data sets: climatic data, remote-sensing data (Normalized Difference Vegetation Index), and elevation data, to model the distribution of six bird species of the genus Grallaria in the Ecuadorian Andes. We assessed the performance of models created with each environmental data set and all possible combinations by comparing the geographic predictions of our models with detailed maps developed by expert ornithologists. Results varied depending on the specific measure of performance. Models including climate variables performed relatively well across most measures, whereas models using only NDVI performed poorly. Elevation based models were relatively good at predicting most sites of expected occurrence but showed a high over-prediction error. Combinations of data sets usually increased the performance of the models, but not significantly. Our results highlight the importance of including climatic variables in ENM and the simultaneous use of various data sets when possible. This strategy attenuates the effects of specific variables that decrease model performance. Remote-sensing data, such as NDVI, should be used with caution in topographically complex regions with heavy cloud-cover. Nonetheless, remote-sensing data have the potential to improve ENM. Finally, we suggest a priori designation of modeling purposes to define specific performance measures accordingly.

Beta diversity and latitude in North American mammals: testing the hypothesis of covariation

Abstract: Several hypotheses attempt to explain the latitudinal gradient of species diversity, but some basic aspects of the pattern remain insufficiently explored, including the effect of scales and the role of beta diversity. To explore such components of the latitudinal gradient, we tested the hypothesis of covariation, which states that the gradient of species diversity should show the same pattern regardless of the scale of analysis. The hypothesis implies that there should be no gradients of beta diversity, of regional range size within regions, and of the slope of the species-area curve. For the fauna of North American mammals, we found contrasting results for bats and non-volant species. We could reject the hypothesis of covariation for non-volant mammals, for which the number of species increases towards lower latitudes, but at different rates depending on the scale. Also, for this group, beta diversity is higher at lower latitudes, the regional range size within regions is smaller at lower latitudes, and z, the slope of the species-area relationship is higher at lower latitudes. Contrarily bats did not show significant deviations from the predictions of the hypothesis of covariation: at two different scales, species richness shows similar trends of increase at lower latitudes, and no gradient can be demonstrated for beta diversity, for regional range size, or for the slopes of the species-area curve. Our results show that the higher diversity of non-volant mammals in tropical areas of North America is a consequence of the increase in beta diversity and not of higher diversity at smaller scales. In contrast, the diversity of bats at both scales is higher at lower latitudes. These contrasting patterns suggest different causes for the latitudinal gradient of species diversity in the two groups that are ultimately determined by differences in the patterns of geographic distribution of the species.

Associations of garden birds with gradients in garden habitat and local habitat

Abstract: Habitat associations of 40 bird species were analysed using data from a survey of 12892 garden sites throughout the United Kingdom between 1995 and 2002 Gradients in 20 garden habitat variables and 25 variables describing the surrounding local habitat were derived from Detrended Correspondence Analysis The main garden axis explained 133% of variation in the data and described a gradient from sites containing and bounded by high levels of vegetation cover to open sites bounded by walls and fences The main local axis explained 299% of variation in the data and described a gradient from rural to urban habitats Higher probability of occurrence was detected in larger sites in 25 species and in smaller sites in four (urban-associated) species Many species (22 out of 40) were significantly associated with the local gradient: seven species showed the highest probability of occurrence in sites within more urbanised habitats Other species were most likely to occur in sites within rural or suburban habitats Only five species showed a significant association with garden habitat gradient, four of these being most likely to occur in sites with highest vegetation cover These associations were similar when controlling for garden size and for food provision in gardens The results imply that the likelihood of many species occurring in gardens is dependent on the surrounding local habitat rather than the garden habitat However, survey participants were likely to have “bird-friendly” gardens which may have reduced the variation in garden habitat across the sample An assessment of the representativeness of survey sites is needed to further understand habitat associations of garden birds

Rarefaction method for assessing plant species diversity on a regional scale

Abstract: 544.In national conservation plans, it is necessary to comparatively assess species pools ofdifferent regions and monitor their changes over time. Two specific problems arise: i)species diversity must be standardized per area, because regions differ in size, and ii) thediversity measure should take into account how common or rare the species are on theregional scale. We used the rarefaction method combined with a fitting procedure tocalculate the expected number of species E(S). The method takes into account thenonlinearity of species and area, as well as how common or rare each species is andallows analysis of species groups’ contribution to total species diversity. The slopeparameter of the fitted power function is used as an indicator of species turnover, andthus, of b-diversity. For the analysis, Switzerland was divided into seven biogeographicregions (256

Environmental correlates of mammal species richness in South America: effects of spatial structure, taxonomy and geographic range

Abstract: Although some consensus exists regarding the positive synergism between energy and heterogeneity in increasing species diversity, the role of environmental variability remains controversial. We examine how these factors interact to explain spatial variation in mammal species richness in South America. After taking into account the effects of spatial autocorrelation and area, elevation variability and energy mainly drive spatial variation in mammal species richness. The effect of environmental variability is less important. When different taxonomic groups of mammals are analyzed separately, three ways emerge whereby energy and heterogeneity interact to promote species richness. Heterogeneity may have no effect on species richness, habitat heterogeneity and energy availability contribute independently to species richness, or heterogeneity increases in importance with an increase in energy availability. The partition of species into range size quartiles shows that habitat heterogeneity and temporal instability in the resource supply account for the species richness pattern in the narrowest- ranging species. Habitat heterogeneity is significant also for intermediate ranging species but not for the widest-ranging species. Energy alone drives the species richness pattern in the latter species. The interplay between ecology and biogeographic history may ultimately explain these differences given that narrow- and wide-ranging species show distinct biogeographic patterns, and different taxonomic groups also unequally represent them.

Butterfly species richness in mainland Portugal: predictive models of geographic distribution patterns

Abstract: A three-step protocol described elsewhere is used to obtain a map of butterfly species density in Portugal on a 50 x 50 km grid. First, all available faunistic information was compiled and analysed to explore the historic patterns of butterfly sampling in Portugal, and to determine which grid cells are sufficiently prospected to produce reliable estimates of species richness. Then, we relate the estimated species richness scores from these areas to a set of environmental and spatial variables by means of General Linear Models, obtaining a function to extrapolate of species density scores to the rest of Portugal. Finally, the model is validated, results explored and outliers identified and deleted. Any spatial autocorrelation remaining in the residuals is examined. Lastly, model parameters are recalculated in absence of deleted outliers, and the resulting function is used to predict species richness scores throughout mainland Portugal. A highly-predictive function based on some variables previously related to butterfly composition at macro-scale, such as number of sunny days per year, temperature or environmental heterogeneity, was obtained. However, in Portugal those variables are highly spatially structured along a steep latitudinal gradient, leading to difficulty in ascertaining if the latitudinal gradient detected by our analysis is due to macroecological or historic effects. Information on European and Iberian butterfly assemblages and causal processes are discussed in the light of the patterns observed. Then, previous information obtained on Portuguese scarabs is added to identify conservation areas, biogeographically important for both insect groups. Finally, the main drawbacks and advantages of this approach to mapping biodiversity for conservation are discussed briefly.

The region effect on mesoscale plant species richness between eastern Asia and eastern North America

Abstract: 136.The greater number of plant species in temperate eastern Asia compared to easternNorth America has been ascribed to both local environment and regionalcharacteristics, but the relative contributions of each have not been resolved. In thisanalysis, we related species richness of flowering plants in mesoscale floras (B

Geographical variation in host specificity of fleas (Siphonaptera) parasitic on small mammals: the influence of phylogeny and local environmental conditions

Abstract: The evolution of host specificity remains a central issue in the study of host-parasite relationships. Here we tackle three basic questions about host specificity using data on host use by fleas (Siphonaptera) from 21 geographical regions. First, are the host species exploited by a flea species no more than a random draw from the locally available host species, or do they form a taxonomically distinct subset? Using randomization tests, we showed that in the majority of cases, the taxonomic distinctness (measured as the average taxonomic distances among host species) of the hosts exploited by a flea is no different from that of random subsets of hosts taken from the regional pool. In the several cases where a difference was found, the taxonomic distinctness of the hosts used by a flea was almost always lower than that of the random subsets, suggesting that the parasites use hosts within a narrower taxonomic spectrum than what is available to them. Second, given the variation in host specificity among populations of the same flea species, is host specificity truly a species character? We found that host specificity measures are repeatable among different populations of the same flea species: host specificity varies significantly more among flea species than within flea species. This was true for both measures of host specificity used in the analyses: the number of host species exploited, and the index measuring the average taxonomic distinctness of the host species and its variance. Third, what causes geographical variation in host specificity among populations of the same flea species? In the vast majority of flea species, neither of our two measures of host specificity correlated with either the regional number of potential host species or their taxonomic distinctness, or the distance between the sampled region and the center of the flea's geographical range. However, in most flea species host specificity correlated with measures of the deviation in climatic conditions (precipitation and temperature) between the sampled region and the average conditions computed across the flea's entire range. Overall, these results suggest that host specificity in fleas is to a large extent phylogenetically constrained, while still strongly influenced by local environmental conditions.