Showing papers in "Ecography in 2008"

Modeling of species distributions with Maxent: new extensions and a comprehensive evaluation

Abstract: Accurate modeling of geographic distributions of species is crucial to various applications in ecology and conservation. The best performing techniques often require some parameter tuning, which may be prohibitively time-consuming to do separately for each species, or unreliable for small or biased datasets. Additionally, even with the abundance of good quality data, users interested in the application of species models need not have the statistical knowledge required for detailed tuning. In such cases, it is desirable to use "default settings", tuned and validated on diverse datasets. Maxent is a recently introduced modeling technique, achieving high predictive accuracy and enjoying several additional attractive properties. The performance of Maxent is influenced by a moderate number of parameters. The first contribution of this paper is the empirical tuning of these parameters. Since many datasets lack information about species absence, we present a tuning method that uses presence-only data. We evaluate our method on independently collected high-quality presence-absence data. In addition to tuning, we introduce several concepts that improve the predictive accuracy and running time of Maxent. We introduce "hinge features" that model more complex relationships in the training data; we describe a new logistic output format that gives an estimate of probability of presence; finally we explore "background sampling" strategies that cope with sample selection bias and decrease model-building time. Our evaluation, based on a diverse dataset of 226 species from 6 regions, shows: 1) default settings tuned on presence-only data achieve performance which is almost as good as if they had been tuned on the evaluation data itself; 2) hinge features substantially improve model performance; 3) logistic output improves model calibration, so that large differences in output values correspond better to large differences in suitability; 4) "target-group" background sampling can give much better predictive performance than random background sampling; 5) random background sampling results in a dramatic decrease in running time, with no decrease in model performance.

Estimating space-use and habitat preference from wildlife telemetry data

Abstract: Management and conservation of populations of animals requires information on where they are, why they are there, and where else they could be. These objectives are typically approached by collecting data on the animals' use of space, relating these positional data to prevailing environmental conditions and employing the resulting statistical models to predict usage at other geographical regions. Technical advances in wildlife telemetry have accomplished manifold increases in the amount and quality of available data, creating the need for a statistical framework that can use them to make population-level inferences for habitat preference and space-use. This has been slow-in-coming because wildlife telemetry data are spatio-temporally autocorrelated, often unbalanced, presence-only observations of behaviourally complex animals, responding to a multitude of cross-correlated environmental variables. We review the evolution of regression models for the analysis of space-use and habitat preference and outline the essential features of a framework that emerges naturally from these foundations. This allows us to derive a relationship between usage of points in geographical space and preference of habitats in environmental space. Within this framework, we discuss eight challenges, inherent in the spatial analysis of telemetry data and, for each, we propose solutions that can work in tandem. Specifically, we propose a logistic, mixed-effects approach that uses generalized additive transformations of the environmental covariates and is fitted to a response data-set comprising the telemetry and simulated observations, under a case-control design. We apply this framework to a non-trivial case-study using satellite-tagged grey seals Halichoerus grypus from the east coast of Scotland. We perform model selection by cross-validation and confront our final model's predictions with telemetry data from the same, as well as different, geographical regions. We conclude that, despite the complex behaviour of the study species, flexible empirical models can capture the environmental relationships that shape population distributions.

Quaternary climate changes explain diversity among reptiles and amphibians

Abstract: It is widely believed that contemporary climate determines large-scale patterns of species richness. An alternative view proposes that species richness reflects biotic responses to historic climate changes. These competing ‘‘contemporary climate’’ vs ‘‘historic climate’’ hypotheses have been vigorously debated without reaching consensus. Here, we test the proposition that European species richness of reptiles and amphibians is driven by climate changes in the Quaternary. We find that climate stability between the Last Glacial Maximum (LGM) and the present day is a better predictor of species richness than contemporary climate; and that the 08C isotherm of the LGM delimits the distributions of narrow-ranging species, whereas the current 08C isotherm limits the distributions of wide-ranging species. Our analyses contradict previous studies of large-scale species richness patterns and support the view that ‘‘historic climate’’ can contribute to current species richness independently of and at least as much as contemporary climate.

Modelling species distributions without using species distributions: the cane toad in Australia under current and future climates

Abstract: Accurate predictions of the potential distribution of range-shifting species are required for effective management of invasive species, and for assessments of the impact of climate change on native species. Range-shifting species pose a challenge for traditional correlative approaches to range prediction, often requiring the extrapolation of complex statistical associations into novel environmental space. Here we take an alternative approach that does not use species occurrence data, but instead captures the fundamental niche of a species by mechanistically linking key organismal traits with spatial data using biophysical models. We demonstrate this approach with a major invasive species, the cane toad Bufo marinus in Australia, assessing the direct climatic constraints on its ability to move, survive, and reproduce. We show that the current range can be explained by thermal constraints on the locomotor potential of the adult stage together with limitations on the availability of water for the larval stage. Our analysis provides a framework for biologically grounded predictions of the potential for cane toads to expand their range under current and future climate scenarios. More generally, by quantifying spatial variation in physiological constraints on an organism, trait-based approaches can be used to investigate the range-limits of any species. Assessments of spatial variation in the physiological constraints on an organism may also provide a mechanistic basis for forecasting the rate of range expansion and for understanding a species’ potential to evolve at range-edges. Mechanistic approaches thus have broad application to process-based ecological and evolutionary models of range-shift.

Postglacial dispersal limitation of widespread forest plant species in nemoral Europe

Abstract: Climate is often singled out as the primary range limiting factor at large scales, while other environmental factors, notably soil, are thought to predominate at smaller scales. However, the postglacial migrational lag hypothesis controversially suggests that many species are strongly dispersal-limited and still expanding from their ice age refugia. We investigated the importance of postglacial migrational lag, climate, and soil as range determinants for 47 widespread forest plant species across nemoral Europe (47.0–60.0°N, west of 24.0°E) using regression modeling, information-theoretic model selection, multi-model inference, and variation partitioning. Migrational lag was represented by a measure of accessibility to recolonization from ice age refugia. Twelve species were largely ubiquitous and not analyzed further. For the remaining species, there was strong support for climate, soil, and accessibility to postglacial recolonization. Accessibility accounted for a small to moderate amount of variation, but its model-averaged regression coefficient was stronger than those for climate or soil for 11 species and the second or third strongest coefficient for an additional 10 species. The resulting odds ratios were greater than one for 33 out of 35 species, i.e. adjusted for climate and soil, prevalence for the far majority of the 35 non-ubiquitous species increased with increasing accessibility. There were no differences among growth forms in the importance of accessibility. In contrast, compared to non-boreal species, accessibility had little importance for species with wide boreal distributions, as expected from their more widespread, northern glacial distributions. In conclusion, even the ranges of many widespread forest plant species are probably still moderately to strongly limited by postglacial migrational lag. Therefore, although species ranges are also strongly influenced by climate, we cannot expect most forest plant species to closely track the expected 21st century climatic changes.

Movement of outbreak populations of mountain pine beetle: influences of spatiotemporal patterns and climate

Abstract: Insect outbreaks exert landscape-level influences, yet quantifying the relative contributions of various exogenous and endogenous factors that contribute to their pattern and spread remains elusive. We examine an outbreak of mountain pine beetle covering an 800 thousand ha area on the Chilcotin Plateau of British Columbia, Canada, during the 1970s and early 1980s. We present a model that incorporates the spatial and temporal arrangements of outbreaking insect populations, as well as various climatic factors that influence insect development. Onsets of eruptions of mountain pine beetle demonstrated landscape-level synchrony. On average, the presence of outbreaking populations was highly correlated with outbreaking populations within the nearest 18 km the same year and local populations within 6 km in the previous two years. After incorporating these spatial and temporal dependencies, we found that increasing temperatures contributed to explaining outbreak probabilities during this 15 yr outbreak. During collapse years, landscape-level synchrony declined while local synchrony values remained high, suggesting that in some areas host depletion was contributing to population decline. Model forecasts of outbreak propensity one year in advance at a 12 by 12 km scale provided 80% accuracy over the landscape, and never underestimated the occurrence of locally outbreaking populations. This model provides a flexible approach for linking temperature and insect population dynamics to spatial spread, and complements existing decision support tools for resource managers.

Detecting spatial hot spots in landscape ecology

Abstract: Hot spots are typically locations of abundant phenomena. In ecology, hot spots are often detected with a spatially global threshold, where a value for a given observation is compared with all values in a data set. When spatial relationships are important, spatially local definitions - those that compare the value for a given observation with locations in the vicinity, or the neighbourhood of the observation - provide a more explicit consideration of space. Here we outline spatial methods for hot spot detection: kernel estimation and local measures of spatial autocorrelation. To demonstrate these approaches, hot spots are detected in landscape level data on the magnitude of mountain pine beetle infestations. Using kernel estimators, we explore how selection of the neighbourhood size (τ) and hot spot threshold impact hot spot detection. We found that as T increases, hot spots are larger and fewer; as the hot spot threshold increases, hot spots become larger and more plentiful and hot spots will reflect coarser scale spatial processes. The impact of spatial neighbourhood definitions on the delineation of hot spots identified with local measures of spatial autocorrelation was also investigated. In general, the larger the spatial neighbourhood used for analysis, the larger the area, or greater the number of areas, identified as hot spots.

Relative importance of different dispersal vectors for small aquatic invertebrates in a rock pool metacommunity

Abstract: The extent and frequency of passive overland dispersal of freshwater invertebrates as well as the relative importance of different dispersal vectors is not well documented. Although anecdotal evidence subscribing the feasibility of individual vectors in various aquatic systems is abundant, dispersal rates have rarely been quantified for different vectors in one study system. Earlier studies also usually investigated dispersal potential rather than actual dispersal rates. In this study we have estimated passive dispersal rates of invertebrate propagules within a cluster of temporary rock pools via water, wind and amphibians in a direct way. Overflows after heavy rains mediated dispersal of a large number of propagules through eroded channels between pools, which were collected in overflow traps. Taking into account model based predictions of overflow frequency, this corresponds with average dispersal rates of 4088 propagules/channel yr−1. Wind dispersal rates as measured by numbers of propagules collected on sticky traps mounted between pool basins were very high (average dispersal rate: 649 propagules m−2 in one month) and were positively related to the proximity of source populations. Finally, invertebrate propagules were also isolated from the faeces of African clawed frogs Xenopus laevis caught from the pools (on average 368 propagules/frog). The combination of short distance wind and overflow dispersal rates likely explain the dominant species sorting and mass effect patterns observed in the metacommunity in a previous study. Amphibian mediated dispersal was much less important as the Xenopus laevis population was small and migrations very rare. Based on our own results and available literature we conclude that both vector and propagule properties determine local passive dispersal dynamics of freshwater invertebrates. Accurate knowledge on rates and vectors of dispersal in natural systems are a prerequisite to increase our understanding of the impact of dispersal on ecology (colonisation, community assembly, coexistence) and evolution (gene flow, local adaptation) in fragmented environments.

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Abstract: Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.

Determinants of species richness, endemism and turnover in European longhorn beetles

Abstract: This study assessed the diversity patterns of a large family of beetles, Cerambycidae, in Europe and tested the following hypotheses: 1) richness gradients of this hyperdiverse taxon are driven by water and energy variables; 2) endemism is explained by the same factors, but variation between areas also reflects post-glacial re-colonization processes; and 3) faunal composition is determined by the same climatic variables and, therefore, beta diversity (species turnover) is related to richness gradients. Species richness, endemism and beta diversity were modelled using inventories of 37 European territories, built from a database containing the distributions of 609 species. Area, spatial position, and nine topographical and climatic variables were used as predictors in regression and constrained analysis of principal coordinates modelling. Species richness was mostly explained by a temperature gradient, which produced a south-to-north decreasing richness gradient. Endemism followed the same pattern, but was also determined by longitudinal variation, peaking in the southwestern and southeastern corners of the continent. Faunal turnover was explained by an important purely spatial pattern and a spatially structured environmental gradient. Thus, contrary to other groups, cerambycid richness was mostly explained by environmental energy, but not by water availability. Endemism was concentrated in the Iberian and Greek peninsulas, but not in Italy. Thus, the latter area may have been the major source of post-glacial re-colonization for European longhorn beetles or, otherwise, a poor refuge during glaciations. Turnover patterns were independent of the richness gradient, because northern faunas are nested in southern ones. Turnover, in contrast to richness, was driven by both the independent effects of climate and geographic constraints that might reflect dispersal limitation or stochastic colonization events, suggesting that richness gradients are more environmentally deterministic phenomena than turnover patterns.

Why do mountains support so many species of birds

Abstract: Although topographic complexity is often associated with high bird diversity at broad geographic scales, little is known about the relative contributions of geomorphologic heterogeneity and altitudinal climatic gradients found in mountains. We analysed the birds in the western mountains of the New World to examine the two-fold effect of topography on species richness patterns, using two grains at the intercontinental extent and within temperate and tropical latitudes. Birds were also classified as montane or lowland, based on their overall distributions in the hemisphere. We estimated range in temperature within each cell and the standard deviation in elevation (topographic roughness) based on all pixels within each cell. We used path analysis to test for the independent effects of topographic roughness and temperature range on species richness while controlling for the collinearity between topographic variables. At the intercontinental extent, actual evapotranspiration (AET) was the primary driver of species richness patterns of all species taken together and of lowland species considered separately. In contrast, within-cell temperature gradients strongly influenced the richness of montane species. Regional partitioning of the data also suggested that range in temperature either by itself or acting in combination with AET had the strongest ‘‘effect’’ on montane bird species richness everywhere. Topographic roughness had weaker ‘‘effects’’ on richness variation throughout, although its positive relationship with richness increased slightly in the tropics. We conclude that bird diversity gradients in mountains primarily reflect local climatic gradients. Widespread (lowland) species and narrow-ranged (montane) species respond similarly to changes in the environment, differing only in that the richness of lowland species correlates better with broad-scale climatic effects (AET), whereas mesoscale climatic variation accounts for richness patterns of montane species. Thus, latitudinal and altitudinal gradients in species richness can be explained through similar climatic-based processes, as has long been argued.

Deep-sea nematode biodiversity in the Mediterranean basin : testing for longitudinal, bathymetric and energetic gradients

Abstract: The knowledge of the processes controlling the spatial distribution of species diversity is one of the main challenges of the present ecological research. Spatial patterns of benthic biodiversity in the deep sea are poorly known in comparison with other ecosystems and this limits our understanding of the mechanisms controlling the distribution and maintenance of high biodiversity in the largest ecosystems of our biosphere. Although the Mediterranean basin covers 7.5% of the global biodiversity. The high biogeographic complexity and the presence of steep ecological gradients contribute in making the Mediterranean a region of very high diversity. Here we report the results of an investigation on the patterns of nematode biodiversity in the deep-Mediterranean Sea, in relation with bathymetric, longitudinal and energetic gradients. Our results indicate that benthic biodiversity in the deep-Mediterranean decreases significantly with increasing depth. Moreover, at equally deep sites, nematode diversity decreased from the western to the eastern basin and longitudinal gradients were evident when comparing sites at 4000-m depth, with 3000-m depth. The analysis of the available energy (measured as labile organic matter content of the sediments) suggests that biodiversity patterns are not controlled by the amounts of food resources, but instead bio-availability is the key factor. A more detailed analysis revealed an extremely high deep-sea beta-diversity (turnover diversity), both among sites at different depths as well as at similar depths of different longitude or within the same basin. This new finding has not only important implications on the estimates of the overall regional diversity (gamma diversity), but also suggests the presence of high biogeographic complexity in the deep benthic domain of the Mediterranean Sea.

Representing genetic variation as continuous surfaces: An approach for identifying spatial dependency in landscape genetic studies

Abstract: Landscape genetics, an emerging field integrating landscape ecology and population genetics, has great potential to influence our understanding of habitat connectivity and distribution of organisms. Whereas typical population genetics studies summarize gene flow as pairwise measures between sampling localities, landscape characteristics that influence population genetic connectivity are often continuously distributed in space. Thus, there are currently gaps in both the ability to analyze genotypic data in a continuous spatial context and our knowledge of expected of landscape genetic structure under varying conditions. We present a framework for generating continuous ‘‘genetic surfaces’’, evaluate their statistical properties, and quantify statistical behavior of landscape genetic structure in a simple landscape. We simulated microsatellite genotypes under varying parameters (time since vicariance, migration, effective population size) and used ancestry (q) values from STRUCTURE to interpolate a genetic surface. Using a spatially adjusted Pearson’s correlation coefficient to test the significance of landscape variable(s) on genetic structure we were able to detect landscape genetic structure on a contemporary time scale (]5 generations post vicariance, migration probability 50.10) even when population differentiation was minimal (FST ]0.00015). We show that genetic variation can be significantly correlated with geographic distance even when genetic structure is due to landscape variable(s), demonstrating the importance of testing landscape influence on genetic structure. Finally, we apply genetic surfacing to analyze an empirical dataset of black bears from northern Idaho USA. We find black bear genetic variation is a function of distance (autocorrelation) and habitat patch (spatial dependency), consistent with previous results indicating genetic variation was influenced by landscape by resistance. These results suggest genetic surfaces can be used to test competing hypotheses of the influence of landscape characteristics on genetic structure without delineation of categorical groups.

Dynamic occupancy models reveal within-breeding season movement up a habitat quality gradient by a migratory songbird

Abstract: The timing of settlement decisions likely influences the quality of breeding site choices.This is particularly the case in migratory birds, because the conditions that enhance breeding success are often not apparent upon arrival after migration. A strategy that addresses this problem is to adjust settlement decisions when reliable information becomes available. We used a new indirect method – dynamic site occupancy modeling – to estimate apparent movement of black-throated blue warblers Dendroica caerulescens among sites within a breeding season. Because individuals should disperse to sites that maximize their fitness, we hypothesized that warblers would move up a habitat quality gradient when opportunities arose. For our study species, that would involve moving into sites with greater shrub density and at higher elevation within northern hardwoods forest, as these two features are positively correlated with reproduction and apparent survival in this species. Although the probability of site occupancy in our study landscape remained consistent throughout the breeding season (range: 0.66–0.69), occupancy models revealed substantial support for apparent movement of individuals within the breeding season. The mean probability of emigration from a point count site was 0.21 (±0.03 SE), and the mean probability of immigration to a site not previously occupied was 0.51 (±0.05 SE). The spatial distribution of this movement was a function of habitat quality. A portion of the black-throated blue warbler population appears to arrive on the breeding grounds and settle initially in sub-optimal habitat, moving subsequently into high quality densely shrubbed habitat at higher elevations. This modeling approach provides a new means to test hypotheses about habitat selection and movement by using presence–non-detection data.

Lichens on dead wood: species‐substrate relationships in the epiphytic lichen floras of the Pacific Northwest and Fennoscandia

Abstract: Dead wood is an important habitat feature for lichens in forest ecosystems, but little is known about how many and which lichens are dependent on dead wood. We reviewed substrate use by epiphytic lichens in the combined floras of Fennoscandia and the Pacific Northwest of North America based on literature and herbarium data and analyzed substrate affinity relative to life form, reproductive mode and major phylogenetic group within the floras. A total of 550 (43%) of the 1271 epiphytic species in the combined floras use wood, and 132 species (10%) are obligately associated with dead wood in one or both regions. Obligate and facultative wood-dwelling guilds in the two floras were strongly similar in terms of internal guild structure in each region, but differ somewhat in species composition, while the bark-dwelling guild differs strongly in both. Most obligate dead wood users are sexually reproducing crustose lichens. The largest numbers of species are associated with forest structural features such as logs and snags that have been greatly reduced by forest practices. Conservation of lichens inhabiting wood requires greater attention to crustose lichen species and the development of conservation strategies that look beyond numbers and volumes of dead wood and consider biologically meaningful dead wood structure types.

A unified index to measure ecological diversity and species rarity

Abstract: Several indices have been created to measure diversity, and the most frequently used are the Shannon-Wiener (H) and Simpson (D) indices along with the number of species (S) and evenness (E). Controversies about which index should be used are common in literature. However, a generalized entropy (Tsallis entropy) has the potential to solve part of these problems. Here we explore a family of diversity indices (S q ; where q is the Tsallis index) and evenness (E q ), based on Tsallis entropy that incorporates the most used indices. It approaches S when q = 0, H when q → 1 and gives D when q = 2. In general, varying the value of the Tsallis index (q), S q varies from emphasis on species richness (q 1). Similarly, E q also works as a tool to investigate diversity. In particular, for a given community, its minimum value represents the maximum deviation from homogeneity (E q = 1) for a particular q (herein named q*). It is remarkable that our analysis indicates that q* and its corresponding evenness, E q *, are negatively affected by S when using simulated data. They may represent an index related to species rarity. Furthermore, S q* (i.e. the value of S q for a specific q*) is positively affected by richness that is an important property of any diversity index. In general, our findings indicate that the indices H, D, S, S q* , E and E q* are only part of a whole set of possibilities. In addition, the ecological properties of E q* and S q* , proposed here for the first time, show promise in ecology.

Demographics in an alpine reindeer herd: effects of density and winter weather

Abstract: We examined how population density, winter weather, snow conditions, and 2 large-scale climatic indices (North Atlantic Oscillation, NAO, and Arctic Oscillation, AO) influenced demography (reproduction and mortality) in an alpine herd of semi-domesticated reindeer Rangifer tarandus between 1959 and 2000 in Finnish Lapland. The herd lived on heavily grazed lichen pastures, with winter densities between 0.8 and 3.9 individuals km -2 . Icing conditions occurred every 7th yr, on an average, and decreased reproductive rate (calves/females) by 49%. In general linear models icing remarkably increased the fit of snow models to reproductive rate. Incorporation of an interaction term between icing and the snow depth index provided better fit than a model without interaction. Delayed snowmelt decreased reproductive rate. For the day of snowmelt, however, the model without interaction was better than the interaction model. These 3 models provided the best fit to the data and accounted for 51-54% of the variation in reproductive rate. Winter mortality was related to density and large-scale climatic indices, but not to local winter weather except a slight increase in mortality during an icing winter. The best model for winter mortality, including reindeer density and NAO, accounted for 26% of variation in mortality. Three factors may be involved explaining weak density dependence or the lack of such dependence; climate change scenarios that predict higher winter temperature, more frequent thawing-freezing periods, and deeper snow would be expected to decrease reproductive rate and increase winter mortality of reindeer and thus to reduce profitability of reindeer husbandry. In contrast, early springs would be advantageous for reindeer in the short term.

Exploring arboreal ant community composition and co-occurrence patterns in plantations of oil palm Elaeis guineensis in Borneo and Peninsular Malaysia

Abstract: The oil palm, Elaeis guineensis, is a native of West Africa and is now extensively grown in south-east Asia. In 2005, the crop covered four million ha of Malaysia. We examined the arboreal ant fauna in two oil palm plantations that had been farmed with integrated pest management practices for at least 10 yr. Having cut and examined palm fronds from 595 and 503 palms in Borneo and Peninsular Malaysia respectively, we studied diversity in ant assemblages, analysed composition of ant mosaics by stepwise use of null-models, and explored the influence of exotic vs native species on community structure. Altogether we sampled 52 species of canopy ants, of which 23 species were shared between Borneo and Peninsular Malaysia. Seventeen species in Borneo and 24 species in Peninsular Malaysia accounted for 95% of all species occurrences (SOCs). Fortysix percent of all SOCs belonged to 12 tramp ant species with pan-tropical distributions. Forty percent of all SOCs were exotic ants. Six dominant species were found in Borneo, five in the Peninsula. The three most abundant species, Anoplolepis gracilipes, Oecophylla smaragdina and Technomyrmex albipes were dominant in both sites. Three dominants were invasive species that were found on more palms than the native group. Dominant tramp species coexisted with non-dominants more often and tolerated more species of non-dominant ants than native dominants. Null model analyses indicated that species were arranged in mosaics at both the Peninsular Malaysia and Borneo sites. Dominant species showed much less co-occurrence than expected by chance, pointing towards species segregation (even in presence of introduced, invasive species). Similar results were obtained for all species. Non-dominant ants were either positively associated with their dominants, or co-occurred randomly in the presence of dominant species. In contrast, ant assemblages on palms without dominants showed species segregation.

Spider assemblages in winter oilseed rape affected by landscape and site factors

Abstract: Spiders are an abundant and diverse group of generalist predators in arable fields. Knowledge on what landscape and site factors affect this group can be valuable for efforts to reduce biodiversity loss in agricultural landscapes and can have implications for natural pest control. We investigated the impact of landscape and site factors on epigeic spiders in 29 winter oilseed rape fields (Brassica napus, OSR) embedded in differently structured landscapes in an agricultural region east of Vienna (Austria). Landscape factors included proportions of non-crop areas, woody areas and fallows, lengths of road-side strips and hedges, and landscape diversity at different spatial scales (r =250-2000 m). Site factors included OSR stand density, soil index, soil cultivation intensity, nitrogen fertilisation level, OSR vegetation cover in late autumn, and insecticide applications. Data were analysed using regression, ordination, and variation partitioning. Different characteristics of spider assemblages responded to different landscape factors at different spatial scales. Observed species richness showed the strongest positive reponse to proportions of woody areas at rather small scale (radius 500 m), but the relation remained significant up to the 1250 m radius. Standardised species richness was positively related to non-crop area at the smallest scale (radius 250 m). Activity density was positively related to length of road-side strips with maximum effects at large scale (radius 1750 m) and non-crop area (radius 750 m). Site characteristics (stand density, insecticide applications, and late autumn ground cover) and landscape factors (woody areas and fallows at radius 500 m) were similarly important for explaining species composition. We interpret the scale-dependency of relations as the result of differences in dispersal power of the studied spider species. These results demonstrate the important, scale dependent influence of natural and semi-natural habitats on spider assemblages in arable fields.

Where do seedlings go? A spatio-temporal analysis of seedling mortality in a semi-arid gypsophyte

Abstract: Studies of seedling population dynamics often focus on survival because it provides an integrated measure of seedling performance. However, this approach involves a substantial loss of information because survival is the net result of a wide range of mechanisms. The present study overcomes these shortcomings by investigating spatial and temporal patterns in the causes of plant mortality in a population of Helianthemum squamatum seedlings. We use new point pattern analyses based on K functions combined with a new null model (‘‘independent labeling’’). A total of 871 seedlings of H. squamatum were mapped and regularly monitored over an 18-month period. More than 60% of seedlings died during this period. Causes of mortality were spatially structured, and these structures shifted through time. Small differences in either the time of emergence or the environment surrounding H. squamatum seedlings had profound influences on their fate. Seedlings emerging late in the season under the canopy of adult plants died from drought more often than expected, whereas those emerging earlier in the same microsite survived more than expected. The identity of neighbors also affected the spatio-temporal dynamics of mortality causes. Our results show that seedling-adult interactions cannot be easily predicted from simple models, and that the time of seedling emergence, its age and the identity of its neighbors determine the sign and the spatial scale of these interactions. The new methods introduced in this article open an avenue for the detailed analyses of the spatio-temporal dynamics of plant mortality and can help to disentangle the complexity of biotic interactions along environmental severity gradients. The ‘‘stress gradient hypothesis’’ (SGH) predicts that the relative importance of facilitation and competition should vary inversely along gradients of abiotic stress, with facilitation increasing in importance over competition as abiotic stress increases (Bertness and Callaway 1994). Most research testing this hypothesis has been conducted on

Likely responses of the Antarctic benthos to climate-related changes in physical disturbance during the 21st century, based primarily on evidence from the West Antarctic Peninsula region.

Abstract: Disturbance has always shaped the evolution and ecology of organisms and nowhere is this more apparent that on the iceberg gouged continental shelves of the Antarctic Peninsula (AP). The vast majority of currently described polar biodiversity occurs on the Southern Ocean shelf but current and projected climate change is rapidly altering disturbance intensities in some regions. The AP is now amongst the fastest warming and changing regions on earth. Seasonal sea ice has decreased in time and extent, most glaciers in the region have retreated, a number of ice shelves have collapsed, and the surface waters of the seas west of the AP have warmed. Here, we review the influences of disturbance from ice, sedimentation, freshening events, wave action and humans on shallow water benthic assemblages, and suggest how disturbance pressures will change during the 21st century in the West Antarctic Peninsula (WAP) and Scotia Arc region. We suggest that the intensity of ice scouring will increase in the region over the next few decades as a result of decreased winter sea ice periods and increased ice loading into coastal waters. Thus, the most frequently disturbed environment on earth will become more so, which will lead to considerable changes in community structure and species distributions. However, as ice fronts retreat past their respective grounding lines, sedimentation and freshening events will become relatively more important. Human presence in the region is increasing, through research, tourism, and resource exploitation, which represents a considerable threat to polar biodiversity over the next century. Adapting to or tolerating multiple, changing environmental stressors will be difficult for a fauna with typically slow generation turnovers that has evolved largely in isolation. We suggest that intensifying acute and chronic disturbances are likely to cause significant changes in ecosystem structure, and probably a considerable loss of polar marine biodiversity, over relatively short timescales.

Demography of alpine red squirrel populations in relation to fluctuations in seed crop size

Abstract: Vertebrate population dynamics, social organisation and space use often are closely associated with the distribution of critical resources, such as food. Tree squirrels are ideal models to study these relationships, since both key demographic parameters (reproduction, survival and dispersal) and spatio-temporal variation in food supplies (measured as seed-crop size) can be reliably estimated. In this paper we test the following two predictions underlying the association between annual food abundance and demography in six alpine red squirrel populations, both with and without time-lag effects: 1) between-season and between-year fluctuations in survival rate, population density and increase parallel those in food availability; and 2) individuals follow a resource tracking strategy and increase in density mainly the year after a rich seed-crop. Red squirrels occurred at higher densities in Scots pine forest, characterised by stable seed-crops, than in Norway spruce with more abundant but more variable seed crops. Fluctuations in numbers were positively correlated with food availability, measured as annual conifer seed-crop sizes. Overall, adult survival rates were higher than those of subadults, and survival substantially fluctuated between seasons and years. Autumn densities and rates of population increase (summer-autumn) were strongly correlated with the same year's autumn seed-crop, while correlations with the previous year's seed-crop (time-lag models) were either weak (population density) or absent (population increase). Results of this paper show that fluctuations in red squirrel densities in habitats with strong temporal variation in seed production are more closely linked with food availability than in more stable habitats. In addition, in the Alpine conifer forests squirrel population sizes, in autumn, increase in synchrony with food resources, eliminating the population lag normally present when resources are produced in pulses.

Fruit tracking between sites and years by birds in Mediterranean wintering grounds

Abstract: J. L. Telleri´a (telleria@bio.ucm.es), A. Ramirez and J. Pe´rez-Tris, Dept of Zoology, Faculty of Biology, Univ. Complutense, ES-28040 Madrid,Spain.Knowledge of the ability of birds to track spatiotemporal variation in fruit distribution is essential for understandingplant-frugivore interactions. Arguably, although total fruit availability sets an upper limit to the number of birds that canexploit a habitat patch, not all species can equally distribute abundance according to variation in fruit resources. Toexplore this, we studied bird and fruit abundance in 1999 2005 in Mediterranean scrublands and woodlands of southernSpain. We analysed whether changes of fruit abundance in eight different sites during six winters could predict numericalchanges of a set of frugivorous passerines of the area (blackcap Sylvia atricapilla, Sardinian warbler S. melanocephala, robinErithacus rubecula, song thrush Turdus philomelos and blackbird T. merula). We also investigated if all frugivores togethertracked fruits better than individual species, thereby supporting a shared use of resources. Results showed strong inter-specific differences. Only the most abundant species (blackcaps and robins) tracked the spatial patterning of food despitestrong differences in the use of space (vagrant and territorial, respectively). This suggests plastic behaviour of territorialrobins, with individuals changing from strictly territorial to wandering, a flexibility that would favour between-sitenumerical arrangements according to food resources. Annual changes in bird numbers were independent of theavailability of fruits, except for blackcaps, an abundant vagrant bird that tracked inter-winter changes in fruit abundance.The abundance of blackcaps fitted the spatiotemporal patterning of fruit resources better than the whole guild offrugivorous birds, inconsistent with the idea that these species track together the changing availability of fruit resources.

Integrating life history and physiology to understand latitudinal size variation in a damselfly

Abstract: Our understanding of latitudinal life history patterns may benefit by jointly considering age and mass at maturity and growth rate. Additional insight may be gained by exploring potential constraints through pushing growth rates to their maximum and scoring physiological cost-related variables. Therefore, we reared animals of a univoltine Spanish and Belgian population and of a semivoltine Swedish population of the damselfly Enallagma cyathigerum (spanning a latitude gradient of ca 2350 km) in a common environment from the eggs until adult emergence and exposed them to a transient starvation period to induce compensatory growth. Besides age and mass at maturity and growth rate we also scored investment in energy storage (i.e. triglycerides) and immune function (i.e. total activity of phenoloxidase). At emergence, body mass was greater in Spain and Sweden and lower in Belgium, suggesting a genetic component for the U-shaped latitudinal pattern that was found also in a previous study based on field-collected adults. The mass difference between univoltine populations can be explained by the shorter development time in the Belgian population, and this despite a higher growth rate, a pattern consistent with undercompensating countergradient variation. In line with the assumed shorter growth seasons, Belgian and Swedish animals showed higher routine growth rates and compensatory growth after transient starvation. Despite a strong link with metabolic rates (as measured by oxygen consumption) populations with higher routine growth rates had no lower fat content and had higher immune function (i.e. immune function decreased from Sweden to Spain), which was unexpected. Rapid compensatory growth did, however, result in a lowered immune function. This may contribute to the absence of perfect compensating countergradient variation in the Belgian population and the lowest routine growth rates in the Spanish population. Our results underscore the importance of integrating key life historical with physiological traits for understanding latitudinal population differentiation.

Decreased resistance and increased tolerance to native herbivores of the invasive plant Sapium sebiferum

Abstract: Release from natural enemies may favor invasive plants evolving traits associated with reduced herbivore-resistance and faster-growth in introduced ranges. Given a genetic trade-off between resistance and tolerance, invasive plants could also become more tolerant to herbivory than conspecifics in the native range. We conducted a field common garden study in the native range of Sapium sebiferum using seeds from native Chinese populations and invasive North American populations to compare their growth and herbivory resistance. We also performed a cage-pot experiment to compare their resistance and tolerance to Bikasha collaris beetles that are specialist feeders on S. sebiferum trees in China. Results of the common garden study showed that Sapium seedlings of invasive populations relative to native populations were more frequently attacked by native herbivores. Growth and leaf damage were significantly higher for invasive populations than for native populations. Growth of invasive populations was not significantly affected by insecticide spray, but insecticide spray benefited that of native populations. In the bioassay trial, beetles preferentially consumed leaf tissue of invasive populations compared to native populations when beetles had a choice between them. Regression of percent leaf damage on biomass showed that invasive populations tolerated herbivory more effectively than native populations. Our results suggest that S. sebiferum from the introduced range had lower resistance but higher tolerance to specialist herbivores. Both defense strategies could have evolved as a response to the escape from natural enemies in the introduced range.

Evolutionary species–area curves as revealed by single‐island endemics: insights for the inter‐provincial species–area relationship

Abstract: The species–area relationship (SAR) between different biological provinces is one of the most interesting, but least explored aspects of the well-known species–area pattern. Following the usage that a biological province is a region whose species have for the most part evolved within it, rather than immigrating from somewhere else, we propose that islands can be considered equivalent to biological provinces for single island endemic species (SIEs). Hence, analyses of the relationships between numbers of SIEs and island area can be used as model systems to explore the form of inter-provincial SARs. We analyzed 13 different data sets derived from 11 sources, using the power (log–log) model. Eleven of the SIE–area relationships were statistically significant, explaining high proportions of the variance in SIE numbers (R2 0.57–0.95). The z-values (slopes) of the statistically significant SIE–area relationships ranged from 0.47 to 1.13, with a mean value of 0.80 (SD±0.24). All the island systems in which SIE represent >50% of species exhibited z-values for the SARs of native species higher than those deemed typical of archipelagic SARs. The SIE–area slopes are quite similar to those previously calculated for inter-provincial SARs, indicating that, when speciation becomes the dominant process adding to the species richness of assemblages, high z-values should be anticipated, regardless of the biogeographical scale of the study system.

RangeModel: tools for exploring and assessing geometric constraints on species richness (the mid‐domain effect) along transects

Abstract: RangeModel is a computer application that offers animated demonstrations of the mechanism behind the mid-domain effect. The program also provides analytical tools for the assessment of geometric constraints in empirical datasets for one-dimensional domains (transects). The mid-domain effect (MDE) is the increasing overlap of species ranges towards the center of a shared, bounded domain due to geometric boundary constraints in relation to the distribution of range sizes, producing a peak or plateau of species richness towards the center of the domain. Domains may be spatial, temporal, or functional. RangeModel is a stand-alone, graphical-interface, freeware application for PC and Mac OS platforms.

Predicting the spread of an invasive plant: combining experiments and ecological niche model

Abstract: Rapid evolutionary adjustments to novel environments may contribute to the successful spread of invasive species, and can lead to niche shifts making range dynamics unpredictable. These effects might be intensified by artificial selection in the course of breeding efforts, since many successful plant invaders were deliberately introduced and cultivated as ornamentals. We hypothesized that the invasion success of Buddleja davidii, the ornamental butterfly bush, is facilitated by local adaptation to minimum temperatures and thus, exhibits unpredictable range dynamics. To assess the potential effects of adaptive evolution and artificial selection on the spread of B. davidii, we combined a common garden experiment investigating local adaptation to frost, with ecological niche modelling of the species’ native and invasive ranges. We expected that populations naturalized in sub-continental climate are less susceptible to frost than populations from oceanic climate, and that the invasive range does not match predictions based on climatic data from the native range. Indeed, we revealed significant variation among invasive B. davidii populations in frost resistance. However, frost hardiness was not related to geographic location or climatic variables of the populations’ home site, suggesting that invasive B. davidii populations are not locally adapted to minimum temperatures. This is in line with results of our ecological niche model that did not detect a niche shift between the species’ native range in China, and its invasive range in Europe and North America. Furthermore, our niche model showed that the potential invasive range of B. davidii is still not completely occupied. Together with the frost resistance data obtained in our experiment, the results indicate that climatic conditions are currently not limiting the further spread of the species in Europe and North America.

How can we preserve and restore species richness of pollinating insects on agricultural land

Abstract: During recent decades, concern about the loss of biodiversity on agricultural land has increased, and semi-natural grasslands have been highlighted as critical habitats. Temperate European agricultural landscapes require distinct and appropriate management to prevent further impoverishment of the flora and fauna. This is especially urgent for pollinating insects that provide important ecosystem services. Our aim was to examine how species richness of three important groups of pollinating insects; solitary bees, butterflies and burnet moths are related to different farm characteristics, and if there are any differences between these three groups. A further aim was to test if red-listed species are related to any farm characteristics. Species richness of solitary bees, butterflies and burnets was measured on all seminatural grasslands at 16 farms in a forest-dominated area of 50 km2 in southern Sweden, using systematic transect walks in April to September 2003 (only butterflies and burnets) and 2005. Species richness of solitary bees and butterflies was intercorrelated, both before and after controlling for the area of semi-natural grassland. Species richness of solitary bees increased with the area of semi-natural grassland. After controlling for the effect of the area of semi-natural grassland species richness was strongly positively related with the density of the plant Knautia arvensis and negatively related with the proportion of grazed grassland. The results were similar for solitary bees and butterflies. The number of red-listed solitary bees was positively related to the proportion of meadows with late harvest(after mid-July) and decreased with increased farm isolation. The number of burnet species (all red-listed) was positively related to vegetation height, flower density and the proportion of meadows with late harvest on a farm. Areas with a high density of K. arvensis and with traditional hay-meadow with late harvest present, harbour most species. Promoting traditional hay-meadows, late extensive grazing and the herb K. arvensis, people managing agricultural biodiversity can encompass high species richness of pollinating insects and support red-listed species. Further, we suggest that the density of K. arvensis at a farm can be used as a biodiversity indicator, at least for pollinating insects. (Less)