Showing papers in "Ecography in 2006"

Novel methods improve prediction of species' distributions from occurrence data

Abstract: Prediction of species' distributions is central to diverse applications in ecology, evolution and conservation science. There is increasing electronic access to vast sets of occurrence records in museums and herbaria, yet little effective guidance on how best to use this information in the context of numerous approaches for modelling distributions. To meet this need, we compared 16 modelling methods over 226 species from 6 regions of the world, creating the most comprehensive set of model comparisons to date. We used presence-only data to fit models, and independent presence-absence data to evaluate the predictions. Along with well-established modelling methods such as generalised additive models and GARP and BIOCLIM, we explored methods that either have been developed recently or have rarely been applied to modelling species' distributions. These include machine-learning methods and community models, both of which have features that may make them particularly well suited to noisy or sparse information, as is typical of species' occurrence data. Presence-only data were effective for modelling species' distributions for many species and regions. The novel methods consistently outperformed more established methods. The results of our analysis are promising for the use of data from museums and herbaria, especially as methods suited to the noise inherent in such data improve.

The effect of sample size and species characteristics on performance of different species distribution modeling methods

Abstract: Species distribution models should provide conservation practioners with estimates of the spatial distributions of species requiring attention. These species are often rare and have limited known occurrences, posing challenges for creating accurate species distribution models. We tested four modeling methods (Bioclim, Domain, GARP, and Maxent) across 18 species with different levels of ecological specialization using six different sample size treatments and three different evaluation measures. Our assessment revealed that Maxent was the most capable of the four modeling methods in producing useful results with sample sizes as small as 5, 10 and 25 occurrences. The other methods compensated reasonably well (Domain and GARP) to poorly (Bioclim) when presented with datasets of small sample sizes. We show that multiple evaluation measures are necessary to determine accuracy of models produced with presence-only data. Further, we found that accuracy of models is greater for species with small geographic ranges and limited environmental tolerance, ecological characteristics of many rare species. Our results indicate that reasonable models can be made for some rare species, a result that should encourage conservationists to add distribution modeling to their toolbox.

Post‐fire recovery of ant communities in Submediterranean Pinus nigra forests

Abstract: This study analyzes the variations in the structure and composition of ant communities in burned Pinus nigra forests in central Catalonia (NE Spain). Pinus nigra forests do not recover after fire, changing to shrublands and oak coppices. For this reason, we suggest that ant communities of burned P. nigra forests will change after fire, because the post-fire scenario, in particular with the increase of open areas, is different to the unburned one, and more favourable for some species than for others. In four locations previously occupied by P. nigra forests where different fires occurred 1, 5, 13 and 19 yr before the sampling, we sampled the structure and composition of ant communities with pitfall traps, tree traps and net sweeping in unburned plots and in plots affected by canopy and understory fire. The results obtained suggest that canopy and understory fire had little effect on the structure of ant communities. Thus, many variables concerning ant communities were not modified either by fire type (understory or canopy fire) or by time since fire. However, a number of particular species were affected, either positively or negatively, by canopy fire: three species characteristic of forest habitats decreased after fire, while eight species characteristic of open habitats increased in areas affected by canopy fire, especially in the first few years after fire. These differences in ant community composition between burned and unburned plots imply that the maximum richness is achieved when there is a mixture of unburned forests and areas burned with canopy fire. Moreover, as canopy cover in P. nigra forests burned with canopy fire is not completed in the period of time studied, the presence of the species that are characteristic of burned areas remains along the chronosequence studied, while the species that disappear after fire do not recover in the period of time considered. Overall, the results obtained indicate that there is a persistent replacement of ant species in burned P. nigra forests, as is also the case with vegetation.

Spatial patterns and competition of tree species in a Douglas-fir chronosequence on Vancouver Island

Abstract: While the successional dynamics and large-scale structure of Douglas-fir forest in the Pacific Northwest region is well studied, the fine-scale spatial characteristics at the stand level are still poorly understood. Here we investigated the fine-scale spatial structure of forest on Vancouver Island, in order to understand how the three dominant species, Douglas-fir, western hemlock, and western redcedar, coexist and partition space along a chronosequence comprised of immature, mature, and old-growth stands. We quantified the changes in spatial distribution and association of the species along the chronosequence using the scale-dependent point pattern analyses pair-correlation function g(r) and Ripley's L-function. Evidence on intra- and inter-specific competition was also inferred from correlations between nearest-neighbor distances and tree size. Our results show that 1) the aggregation of Douglas-fir in old-growth was primarily caused by variation in local site characteristics, 2) only surviving hemlock were more regular than their pre-mortality patterns, a result consistent with strong intra-specific competition, 3) inter-specific competition declined rapidly with stand age due to spatial resource partitioning, and (4) tree death was spatially randomly distributed among larger overstory trees. The study highlights the importance of spatial heterogeneity for the long-term coexistence of shade-intolerant pioneer Douglas-fir and shade-tolerant western hemlock and western redcedar.

Variations in species and functional plant diversity along climatic and grazing gradients

Abstract: Different components of biodiversity may vary independently of each other along environmental gradients giving insights into the mechanisms that regulate species coexistence. In particular, the functional diversity (FD) or the presence of rare or endemic species in natural assemblages do not necessarily increase with species diversity. We studied if different components of plant species diversity (species richness, Simpson diversity, evenness) varied similarly to FD (measured as a generalization of the Simpson index) and rarity along grazing intensity and climatic gradients. Plots under different sheep grazing regimes (high and low intensity, abandonment) were surveyed in five locations along a climatic gradient in north-eastern Spain, from semi-arid lowland to moist upland locations. Variation in species diversity, functional diversity and rarity followed different patterns. Species diversity was lowest in water-stressed environments (arid locations and southern aspects) and increased with grazing more makedly in humid locations. The FD was comparable between the most species-poor and species-rich locations and decreased with grazing in the moistest location, i.e. where species diversity markedly increased. The FD did not show a strong correlation with species richness nor with the Simpson index and less specious communities could show the highest functional diversity. The rarest species in the region were more frequently found in the abandoned areas, which held the lowest species diversity. Consequently, the mechanisms that enhance the diversity of species do not necessarily support a functional differentiation among those species or the maintenance of rare species in a region. We hypothesize that the degree of dependence of functional diversity on species diversity might be mostly related to the amplitude of the species' traits pool and on how species partition the niche space available.

Landscape level analysis of mountain pine beetle in British Columbia, Canada: spatiotemporal development and spatial synchrony within the present outbreak

Abstract: Aukema, B. H., Carroll, A. L., Zhu, J., Raffa, K. F., Sickley, T. A. and Taylor, S. W. 2006. Landscape level analysis of mountain pine beetle in British Columbia, Canada: spatiotemporal developments and spatial synchrony within the present outbreak. � Ecography 29: 427 � 441. Eruptive herbivores can exert profound landscape level influences. For example, the ongoing mountain pine beetle outbreak in British Columbia, Canada, has resulted in mortality of mature lodgepole pine over � /7 million ha. Analysis of the spatiotemporal pattern of spread can lend insights into the processes initiating and/or sustaining such phenomena. We present a landscape level analysis of the development of the current outbreak. Aerial survey assessments of tree mortality, projected onto discrete 12� /12 km cells, were used as a proxy for insect population density. We examined whether the outbreak potentially originated from an epicenter and spread, or whether multiple localized populations erupted simultaneously at spatially disjunct locations. An aspatial cluster analysis of time series from 1990 to 2003 revealed four distinct time series patterns. Each time series demonstrated a general progression of increasing mountain pine beetle populations. Plotting the geographical locations of each temporal pattern revealed that the outbreak occurred first in an area of westcentral British Columbia, and then in an area to the east. The plot further revealed many localized infestations erupted in geographically disjunct areas, especially in the southern portion of the province. Autologistic regression analyses indicated a significant, positive association between areas where the outbreak first occurred and conservation lands. For example, the delineated area of west-central British Columbia is comprised of three conservation parks and adjacent working forest. We further examined how population synchrony declines with distance at different population levels. Examination of the spatial dependence of temporal synchrony in population fluctuations during early, incipient years (i.e. 1990 � 1996) suggested that outbreaking mountain pine beetle populations are largely independent at scales � /200 km during non-epidemic periods. However, during epidemic years (i.e. 1999 � 2003), populations were clearly synchronous across the entire province, even at distances of up to 900 km. The epicentral pattern of population development can be used to identify and prioritize adjacent landscape units for both reactive and proactive management strategies intended to minimize mountain pine beetle impacts.

Plant species richness and environmental heterogeneity in a mountain landscape: effects of variability and spatial configuration

Abstract: The loss of biodiversity has become a matter of urgent concern and a better understanding of local drivers is crucial for conservation. Although environmental heterogeneity is recognized as an important determinant of biodiversity, this has rarely been tested using field data at management scale. We propose and provide evidence for the simple hypothesis that local species diversity is related to spatial environmental heterogeneity. Species partition the environment into habitats. Biodiversity is therefore expected to be influenced by two aspects of spatial heterogeneity: 1) the variability of environmental conditions, which will affect the number of types of habitat, and 2) the spatial configuration of habitats, which will affect the rates of ecological processes, such as dispersal or competition. Earlier, simulation experiments predicted that both aspects of heterogeneity will influence plant species richness at a particular site. For the first time, these predictions were tested for plant communities using field data, which we collected in a wooded pasture in the Swiss Jura mountains using a four-level hierarchical sampling design. Richness generally increased with increasing environmental variability and "roughness" (i.e. decreasing spatial aggregation). Effects occurred at all scales, but the nature of the effect changed with scale, suggesting a change in the underlying mechanisms, which will need to be taken into account if scaling up to larger landscapes. Although we found significant effects of environmental heterogeneity, other factors such as history could also be important determinants. If a relationship between environmental heterogeneity and species richness can be shown to be general, recently available high-resolution environmental data can be used to complement the assessment of patterns of local richness and improve the prediction of the effects of land use change based on mean site conditions or land use history.

Null Versus Neutral Models: What's The Difference?

Abstract: The neutral model posits that random variation in extinction and speciation events, coupled with limited dispersal, can account for many community properties, including the relative abundance distribution. There are important analogies between this model in ecology and a three-tiered hierarchy of models in evolution (Hardy Weinburg, drift, drift and selection). Because it invokes random processes and is used in statistical tests of empirical data, the neutral model can be interpreted as a specialized form of a null model. However, the application and interpretation of neutral models differs from that of standard null models in three important ways: 1) whereas most null models incorporate species-level constraints that are often associated with niche differences, the neutral model assumes that all species are functionally equivalent. 2) Null models are usually fit with constraints that are measured directly from the data set itself. In contrast, the neutral model requires parameters for speciation, extinction, and migration rates that are almost never measured directly, so their values must be guessed at or fitted. 3) Most important, null models are viewed as simple statistical descriptors: unspecified “random” forces generate variation in a simple model that excludes particular biological mechanisms (usually species interactions). Although the neutral model was originally framed as a null model, recent proponents of the neutral model have begun to treat it as a literal process-based description of community assembly. These differences lie at the heart of much of the recent controversy over the neutral model. If the neutral model is truly a process-based model, then its assumptions should be directly tested, and its predictions should be compared to those of an appropriate null model. Such tests are rarely informative, and most empirical data sets can be fit more parsimoniously to a simple log-normal distribution. Because unknown parameters in the neutral model must usually be guessed at or fit in ad-hoc ways, classical frequentist tests are compromised, and may be biased towards finding a good fit with the model. There has been little analysis of the potential for type I and type II errors in statistical tests of the neutral model. The neutral model has recently been proposed as a specific form of more general null models in biogeography (the mid-domain effect) and community ecology (species co-occurrence). In both cases, the neutral model is qualitatively, but not quantitatively, similar to the predictions of classic null models. However, because the important parameters in the neutral model can rarely be measured directly, it may be of limited value as a null hypothesis for empirical tests. Future progress may come from moving beyond dichotomous tests of neutral versus null models. Instead, the neutral model might be viewed as a mechanism that contributes to pattern along with other processes. Alternatively, the fit of data to the neutral model can be compared to the fit to other process-based models that are not based on neutrality assumptions. Finally, the neutral model can also be tested directly if its parameters can be estimated independently of the test data. However, these approaches may require more data than are often available. For these reasons, simple null model tests will continue to be important in the evaluation of the neutral model.

How does environmental variation influence body mass, body size, and body condition? Roe deer as a case study

Abstract: We tested the influence of population density and of drought intensity (measured as the Gaussen Index in spring and summer of the year of birth) on winter body mass, hind foot length, and body condition of roe deer fawns. Body mass decreased with increasing density and increased with increasing Gaussen Index in summer, in a similar way for both males and females. Hind foot length of males showed the same response. On the other hand, hind foot length of females decreased with increasing density only after dry summers, hence when environmental conditions were very harsh. Body condition was affected neither by density nor by drought intensity. Our results indicate that body mass and size are much better indicators of phenotypic quality than body condition in roe deer. The sex-specific responses of body size to environmental conditions could correspond to a differential allocation in favour of daughters by heavier than average roe deer mothers.

The mid-latitude biodiversity ridge in terrestrial cave fauna

Abstract: The world's obligate cave-dwelling fauna holds considerable promise for biogeographic analysis because it represents a large number of independent evolutionary experiments in isolation in caves and adaptation to subterranean life. We focus on seven north temperate regions of at least 2000 km2, utilizing more than 4300 records of obligate cave-dwelling terrestrial invertebrates. In North America, highest diversity was found in northeast Alabama while in Europe highest diversity was found in Ariege, France, and in southeast Slovenia. Based on these regions as well as more qualitative data from 16 other regions, we hypothesize that a ridge (ca 42°–46° in Europe and 34° in North America) of high biodiversity occurs in temperate areas of high productivity and cave density. This may reflect a strong dependence of cave communities on long term surface productivity (as reflected in actual evapotranspiration), because the subterranean fauna relies almost entirely on resources produced outside caves. This dependence may explain the unique biodiversity pattern of terrestrial cave invertebrates.

Species richness patterns and metapopulation processes – evidence from epiphyte communities in boreo‐nemoral forests

Abstract: suggested. We studied the relative importance of local environmental conditions and spatial aggregation of species richness of facultative and obligate epiphytic bryophytes and lichens within two old-growth forests in eastern Sweden. The effect of the local environment was analyzed using generalized linear models (GLM). We tested whether species richness was spatially structured by fitting variogram models to the residuals of the GLM. In addition, we analyzed the species-area relationship (area = tree diameter). Different environmental variables explained the richness of different species groups (bryophytes vs lichens, specialists vs generalists, sexual vs asexual dispersal). In most groups, the total variation explained by environmental variables was higher than the variation explained by the spatial model. Spatial aggregation was more pronounced in asexually than in sexually dispersed species. Bryophyte species richness was only poorly predicted by area, and lichen species richness was not explained by area at all. Spatial aggregation may indicate effects of dispersal limitation and metapopulation dynamics on community species richness. Our results suggest that species groups differ in habitat requirements and dispersal abilities; there were indications that presence of species with different dispersal strategies is linked to the age of the host tree. Separate analyses of the species richness of species groups that differ in the degree of habitat specialization and dispersal ability give insights into the processes determining community species richness. The poor species-area relationship, especially in lichens, may indicate species turnover rather than accumulation during the lifetime of the host tree. Epiphyte species extinctions may be mainly caused by deterministic processes, e.g. changes in habitat conditions as the host tree grows, ages and dies, rather than by stochastic population processes.

Assemblages of wood‐inhabiting fungi along the gradients of succession and naturalness in boreal pine‐dominated forests in Fennoscandia

Abstract: In boreal forests, the level of naturalness and the stage of succession explain most of the variation in forest structure within a particular forest type. Thus, these two factors should also have a major effect on species assemblages in forests, at least on species groups associated with wood. The present study is the first attempt to analyze empirically the simultaneous effects of forest succession and naturalness on wood-inhabiting fungi, a taxonomic group of special ecological importance. The study area was situated in eastern Finland, middle boreal zone. A total of 41 study plots were established in Pinus sylvestris forests representing three levels of forest naturalness: natural, seminatural and intensively managed forests. Five stages of succession were distinguished according to the age of the dominating tree layer ( 150 yr old), except in managed forests where only four stages were available. A total of 5328 records of 195 species of fungi were made. The first, open stage of succession was clearly the most species-rich period of succession in all levels of forest naturalness. In natural and seminatural forests, the first stage of succession was also very distinctive in its fungal composition, and thus of special value in protecting biodiversity in boreal forests. In the succession following the first stage, the level of naturalness had more effect on assemblages of fungi than did the stage of succession. Intensive forest management affects threatened species particularly. In conclusion, natural young stages of succession should also be included in the network of boreal forest reserves.

Habitat selection as a mechanism of resource partitioning in two cryptic bat species Pipistrellus pipistrellus and Pipistrellus pygmaeus

Abstract: Ecomorphological studies of bat communities often reveal the spatial and temporal coexistence of morphologically similar species, leading to suggestions that these communities are structured by non-deterministic processes. However, the diversification of echolocation call structure in bats allows for considerable morphological similarity while still permitting niche differentiation based on specialisation for prey type and habitat structure. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and P. pygmaeus, which are sympatrically distributed throughout their range, raises the question as to whether these two morphologically similar species partition resources in time and space. To test the hypothesis that the coexistence of these cryptic species is facilitated by differential habitat use, 14 P. pipistrellus, and 12 P. pygmaeus were radio-tracked from adjacent maternity roosts, in northeast Scotland, from May to September 2002/2003. The two species showed distinct habitat partitioning with P. pygmaeus foraging predominantly in riparian woodland and over water, and P. pipistrellus foraging along woodland edges and short isolated tree lines. Inter-specific overlap in habitat use was low and consequently foraging ranges were segregated spatially. The degree of habitat partitioning revealed in these species, which show considerable overlap in echolocation call parameters and functional morphology, suggests that morphological features, whilst useful in separating chiropteran species into coarse-grained foraging guilds, may not predict fine-grained ecological segregation.

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Abstract: Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca, at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus. After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.

Predicting patterns of plant species richness in megadiverse South Africa

Abstract: Using new tools (boosted regression trees) in predictive biogeography, with extensive spatial 23 distribution data for >19 000 species, we developed predictive models for South African plant species richness patterns. Further, biome level analysis explored possible functional determinants of country-wide regional species richness. Finally, to test model reliability independently, we predicted potential alien invasive plant species richness with an independent dataset. Amongst the different hypotheses generally invoked to explain species 30 diversity (energy, favorableness, topographic heterogeneity, irregularity and seasonality), results revealed topographic heterogeneity as the most powerful single explanatory variable for indigenous South African plant species richness. Some biome-specific responses were observed, i.e. two of the five analyzed biomes (Fynbos and Grassland) had richness best explained by the “species-favorableness” hypothesis, but even in this case, topographic heterogeneity was also a primary predictor. This analysis, the largest conducted on an almost exhaustive species sample in a species-rich region, demonstrates the preeminence of topographic heterogeneity in shaping the spatial pattern of regional plant species richness. Model reliability was confirmed by the considerable predictive power for alien invasive species richness. It thus appears that topographic heterogeneity controls species richness in two main ways: firstly, by providing an abundance of ecological niches in contemporary space (revealed by alien invasive species richness relationships) and secondly, by facilitating the persistence of ecological niches through time. The extraordinary richness of the South African Fynbos biome, a world-renowned hotspot of biodiversity with the steepest environmental gradients in South Africa, may thus have arisen through both mechanisms. Comparisons with similar regions of the world outside South Africa are needed to confirm the generality of topographic heterogeneity and favorableness as predictors of plant richness.

The relative importance of landscape and community features in the invasion of an exotic shrub in a fragmented landscape

Abstract: Although invasive plants are recognized as a major ecological problem, little is known of the relative importance of plant community characteristics versus landscape context in determining invasibility of communities. We determined the relative importance of community and landscape features of 30 woodlots in influencing the invasion of Lonicera maackii. We sampled woodlots using the point-quarter method and calculated canopy openness and basal areas and densities of shrub, sapling and tree species, as well as woody species richness. We used aerial photos and ArcView GIS to calculate landscape parameters from the same woodlots using a buffer distance of 1500 m. We used logistic and linear regression analyses to determine the community and landscape factors that best explain L. maackii presence and density. We also tested whether woodlot invasion by L. maackii begins at woodlot edges. Presence of L. maackii was significantly explained only by distance from the nearest town (logistic regression, p=0.017); woodlots nearer town were more likely to be invaded. Among invaded woodlots, density of L. maackii was positively related to the amount of edge in the landscape (partial R =0.592) and negatively related to total tree basal area (partial R 2 =0.134), number of native woody species (partial R 2 =0.054), and sapling shade tolerance index (partial R 2 =0.054). Lonicera maackii in woodlot interiors were not younger than those on the perimeters, leading us to reject the edge-first colonization model of invasion. Our findings reveal that landscape structure is of primary importance and community features of secondary importance in the invasion of L. maackii. This shrub is invading from multiple foci (towns) rather than an advancing front. Connectivity in the landscape (i.e. the number of corridors) did not promote invasion. However, edge habitat was important for invasion, probably due to increased propagule pressure. The community features associated with L. maackii invasion may be indicators of past disturbance.

Effects of an introduced crayfish, Procambarus clarkii, on the distribution of south-western Iberian amphibians in their breeding habitats

Abstract: The introduction of exotic species in aquatic habitats is one of the causes for the amphibian declines observed worldwide. In the 1970s, the red swamp crayfish Procambarus clarkii was introduced in the southwest Iberian Peninsula, where no native crayfish occur. In this study we assess the effect of P. clarkii presence in the breeding site distribution of each of the 13 southwest Iberian amphibians, while simultaneously accounting for the effects of potentially confounding habitat variables, as well as for the effects of the other large aquatic predators in the study area – predatory fish. Amphibian species richness was lower in places where P. clarkii was present than in places without P. clarkii, regardless of fish presence. After accounting for habitat variables and fish, crayfish presence was a negative predictor of the breeding probability for all urodeles (Pleurodeles waltl, Salamandra salamandra, Triturus boscai and T. marmoratus) and for two anurans (Pelobates cultripes and Bufo bufo). The majority of the species affected usually breed in temporary ponds without fish, but that may be colonized by the crayfish. The ongoing expansion of P. clarkii may eventually lead to a growing isolation of amphibian populations and ultimately to local extinctions and a permanent alteration of the amphibian communities in southwest Iberian Peninsula.

Effects of landscape composition and substrate availability on saproxylic beetles in boreal forests: a study using experimental logs for monitoring assemblages

Abstract: Intensive forestry practises in the Swedish landscape have led to the loss and fragmentation of stable old-growth habitats. We investigated relationships between landscape composition at multiple scales and the composition of saproxylic beetle assemblages in nine clear-cut, mature managed and old-growth spruce-dominated forest stands in the central boreal zone of Sweden. We set out fresh spruce and birch logs and created spruce snags in 2001–2002 to experimentally test the effects of coarse woody debris (CWD) type and forest management on the composition of early and late successional, and red-listed saproxylic beetle assemblages. We examined effects of CWD availability at 100 m, and landscape composition at 1 and 10 km on saproxylic beetle abundances. Additionally, we tested whether assemblage similarity decreased with increasing distance between sites. We collected beetles from the experimental logs using eclector and window traps in four periods during 2003. CWD was measured and landscape composition data was obtained from maps of remotely sensed data. The composition of saproxylic beetles differed among different CWD substrates and between clear-cuts and the older stand types, however differences between mature managed and old-growth forests were significant only for red-listed species. Assemblage similarities for red-listed species on clear-cuts were more different at greater distances apart, indicating that they have more localised distributions. CWD availability within 100 m of the study sites was rarely important in determining the abundance of species, suggesting that early successional saproxylic beetles can disperse further than this distance. At a larger scale, a large area of suitable stand types within both 1 and 10 km resulted in greater abundances in the study sites for several common and habitat-specific species. The availability of suitable habitat at scales of 1–10 km is thus likely to be important in the survival of many saproxylic species in forestry-fragmented areas.

Life history variation in an annual plant under two opposing environmental constraints along an aridity gradient

Abstract: Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species.

The significance of geographic range size for spatial diversity patterns in Neotropical palms

Abstract: We examined the effect of range size in commonly applied macroecological analyses using continental distribution data for all 550 Neotropical palm species (Arecaceae) at varying grain sizes from 0.5" to 5°. First, we evaluated the relative contribution of range-restricted and widespread species on the patterns of species richness and endemism. Second, we analysed the impact of range size on the predictive value of commonly used predictor variables. Species sequences were produced arranging species according to their range size in ascending, descending, and random order. Correlations between the cumulative species richness patterns of these sequences and environmental predictors were performed in order to analyse the effect of range size. Despite the high proportion of rare species, patterns of species richness were found to be dominated by a minority of widespread species (∼20%) which contained 80% of the spatial information. Climatic factors related to energy and water availability and productivity accounted for much of the spatial variation of species richness of widespread species. In contrast, species richness of range-restricted species was to a larger extent determined by topographical complexity. However, this effect was much more difficult to detect due to a dominant influence of widespread species. Although the strength of different environmental predictors changed with spatial scale, the general patterns and trends proved to be relatively stabile at the examined grain sizes. Our results highlight the difficulties to approximate causal explanations for the occurrence of a majority of species and to distinguish between contemporary climatic factors and history.

Relationship between tree size and insect assemblages associated with Anadenanthera macrocarpa

Abstract: Campos, R. I., Vasconcelos, H. L., Ribeiro, S. P., Neves, F. S. and Soares, J. P. 2006. Relationship between tree size and insect assemblages associated with Anadenanthera macrocarpa . � Ecology 29: 442 � 450. This study analyzed the effects of tree size, and correlated architectural tree characteristics, on the assemblages of ants and insect herbivores associated with Anadenanthera macrocarpa (Mimosaceae). The latter is a myrmecophilous tree species from the Atlantic rainforest in south-eastern Brazil. Ants and insect herbivores were collected in 30 individuals of A. macrocarpa , ranging from young individuals ( � / 3mi n height) to emergent trees (up to 40 m). Tree height was a strong indicator of other tree characteristics, including trunk diameter, crown height, crown volume, and number of bifurcations. Ants were collected using arboreal pitfall traps and beating, while insect herbivores with beating only. There was a significant increase in both abundance and species richness of ants and insect herbivores with an increase in tree height. In addition, tree height had a significant effect on the species composition of ants and insect herbivores. Assemblages of both taxa showed a nested organization pattern. The species found in small- and medium-sized trees, in general, consisted of a subset of the species found in the crowns and branches of larger, canopy or emergent trees. Thus, in A. macrocarpa , there was not a replacement of insect species with plant ontogeny. This finding is at variance with those conducted in tropical evergreen forests and which show a clear stratification between the understory and canopy insect faunas. Additional studies are needed to explain these contrasting patterns, but it is possible that differences in microclimate are involved. As the forest we studied is semi-deciduous, microclimatic gradients between the understory and the canopy habitat are probably less severe than in an evergreen forest, thus resulting in a lower turnover of species.

The Effects of Dispersal and Recruitment Limitation on Community Structure of Odonates in Artificial Ponds

Abstract: I examined the effects of isolation on the structure of both adult and larval dragonfly (Odonata: Anisoptera) communities forming at physically identical artificial ponds over two years. Isolation, whether measured by distance to the nearest source habitat or by connectivity to multiple sources, was significantly negatively related to the species richness of dragonflies observed at and collected in these ponds. These results indicate that dispersal and recruitment limitation acted as filters on the richness of communities at these artificial ponds. The richness of larval recruits in artificial ponds was lower than the richness of adult dispersers observed at ponds, and distance from a source habitat explained a greater fraction of the variation in larval than adult richness (83 and 50%, respectively). These results and a male biased sex-ratio in adults observed at artificial ponds suggest that isolated habitats may be more recruitment limited than observations of dispersers would suggest. A Mantel test indicated there was a spatial component to the composition of communities forming in tanks, and that distance between tanks and community dissimilarity (1-Jaccard's) were significantly positively related (r=0.52). This pattern suggests that their position with respect to alternative source environments influenced the composition of the communities that recruited into these ponds. These results provide further evidence of recruitment limitation in this system. Results from this study highlight the importance behaviorally limited dispersal may have in taxa morphologically capable of broad dispersal and suggest that the role of dispersal and recruitment limitation may be critical in shaping community structure across habitat gradients that include variation in habitat duration.

Spatial organisation and dynamics of the pine marten Martes martes population in Białowieza Forest (E Poland) compared with other European woodlands

Abstract: We studied factors affecting density and spacing patterns in the pine marten Martes martes population inhabiting temperate forests of Biatowieza National Park, eastern Poland. From 1985/1986 to 1995/1996 marten densities ranged from 3.63 to 7.57 individuals 10 km - 2 (mean 5.4) and were positively correlated with abundance of forest rodents in the previous year. The rate of marten population growth was inversely density-dependent and positively related to rodent density. Annual mortality rate averaged 0.384 and tended to be negatively related to marten densities. Mean annual home range of males (2.58 km 2 , SE =0.24) was larger than that of females (1.41 km2, SE=0.20). Seasonal home ranges also differed significantly between males and females. Both sexes held the smallest ranges in December-January. Female ranges increased in April-May, whereas those of males increased in June-September when they were mating. Fidelity of pine martens to their home ranges was very high. The mean shift between arithmetic centres of seasonal ranges was 0.25 km, and the ranges recorded in two consecutive seasons overlapped, on average, by 87-90%. We observed very little home range overlap between neighbouring male (mean 4-6%) or female (mean 6%) marten. Year round the neighbouring individuals of the same sex neither avoided nor attracted each other. Females attracted males only during the spring-summer mating season. A review of other studies has documented that winter severity and seasonal variation in ecosystem productivity were essential factors shaping the biogeographic variation in pine marten densities between 41° and 68°N. The density of marten populations increased in areas with mild winters and lower seasonality. Maximum population densities (indicative of habitat carrying capacity) were correlated with mean winter temperature. In Europe, male home ranges increased with decreasing forest cover in a study area, whereas female ranges varied positively with rodent abundance.

Emus as non‐standard seed dispersers and their potential for long‐distance dispersal

Abstract: Long-distance seed dispersal may have important consequences for species range, migration rates, metapopulation dynamics, and gene flow. Plants have evolved various adaptations for seed dispersal by standard agents, with typical dispersal distances associated with them. Seeds may also be dispersed by non-standard agents for which they do not show any apparent adaptation and may reach long distances. By sampling the droppings of emus Dromaius novaehollandiae at three localities in Western Australia, we investigated their potential to act as long-distance dispersers of seeds with adaptations for dispersal modes other than endozoochory, such as unassisted, ant, wind and exozoochory, for which they act as non-standard agents. Seventy-seven plant species with five types of dispersal syndromes were found in the 112 droppings analysed, with at least 68 having viable seeds. Although endozoochory was the most frequent syndrome, the presence of other syndromes was important in terms of number of species (61%) and seeds (50%). Estimates of species richness indicated that an increase in sampling effort would increase the number of species observed, especially among non-endozoochores. As a consequence of their long gut retention times and high mobility, emus can provide long-distance dispersal opportunities that may be especially relevant for species with dispersal modes of typically short distances (unassisted, ant). Our results suggest that the role of emus as non-standard agents for long-distance dispersal should be taken into account for understanding current geographic ranges, gene flow and metapopulation dynamics of some plant species, as well as for predicting their future responses to climate change and fragmentation.

Sex, isolation and fidelity: unbiased long-distance dispersal in a terrestrial amphibian

Abstract: Amphibians in general are considered poor dispersers and thus their dispersal curve should be dominated by short movements. Additionally, as male toads do not compete for females and sexual selection is by female choice, dispersal should be male-biased. Furthermore, since adults are site-loyal and polygynous, juveniles should move farther and faster than adults. We tested the hypotheses that dispersal would be limited and both sex- and age-biased in a population of Fowler's toads Bufo fowleri at Lake Erie, Ontario, Canada. Based on a mark-recapture study of 2816 toads, 1326 recaptured at least once, we found that although the toads did show high site fidelity, the dispersal curve was highly skewed with a significant "tail" where the maximum distance moved by an adult was 34 km. Dispersal was neither sex-biased nor age-biased despite clear theoretical predictions that dispersal should be biased towards males and juveniles. We conclude that the resource competition hypothesis of sex-biased dispersal does not predict dispersal tendencies as readily for amphibians as for mammals and birds. Toad dispersal only appears to be juvenile-biased because the juveniles are more abundant than the adults, not because they are the more active dispersers.

Combining richness and abundance into a single diversity index using matrix analogues of Shannon's and Simpson's indices

Abstract: Shannon's and Simpson's indices have been the most widely accepted measures of ecological diversity for the past fifty years, even though neither statistic accounts for species abundances across geographic locales ("patches") An abundant species that is endemic to a single patch can be as much of a conservation concern as a rare cosmopolitan species I extend Shannon's and Simpson's indices to simultaneously account for species richness and relative abundances - ie extend them to multispecies metacommunities - by making the inputs to each index a matrix, rather than a vector The Shannon's index analogue of diversity is mutual entropy of species and patches divided by marginal entropy of the individual geographic patches The Simpson's index analogue of diversity is a modification of mutual entropy, with the logarithm moved to the outside of the summation, divided by Simpson's index of the patches Both indices are normalized for number of patches, with the result being inversely proportional to biodiversity These methods can be extended to account for time-series of such matrices and average age-classes of each species within each patch, as well as provide a measure of spatial coherence of communities

Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects

Abstract: We studied frog biodiversity along an elevational gradient in the Hengduan Mountains, China. Endemic and non-endemic elevational diversity patterns were examined individually. Competing hypotheses were also tested for these patterns. Species richness of total frogs, endemics and non-endemics peaked at mid-elevations. The peak in endemic species richness was at higher elevations than the maxima of total species richness. Endemic species richness followed the mid-domain model predictions, and showed a nonlinear relationship with temperature. Water and energy were the most important variables in explaining elevational patterns of non-endemic species richness. A suite of interacting climatic and geometric factors best explained total species richness patterns along the elevational gradient. We suggest that the mid-domain effect was an important factor to explain elevational richness patterns, especially in regions with high endemism.

Scale-dependent effects of land use on plant species richness of mountain grassland in the European Alps

Abstract: Traditionally managed mountain grasslands in the Alps are species-rich ecosystems that developed during centuries of livestock grazing. However, changes in land use including fertilisation of well accessible pastures and gradual abandonment of remote sites are increasingly threatening this diversity. In five regions of the Swiss and French Alps we assessed the relationship between land use, soil resource availability, cover of the unpalatable species Veratrum album, species richness and vegetation composition of mountain grasslands across four spatial scales ranging from 1 to 1000 m2. Mean species richness and the increase in the number of species with increasing area were lower in intensively grazed, fertilised pastures than in traditional pastures or in abandoned pastures. Species composition of abandoned pastures differed from that of the other management types. Plant species richness was influenced by different factors at different spatial scales. At the 1 m2 scale, plant species richness was negatively related to soil nitrate and influenced by the cover of V. album, depending on land use: species richness and cover of V. album were negatively correlated in abandoned pastures, but positively correlated in fertilised grasslands. At the 1000 m2 scale, a negative effect of fertilization on richness was evident. These results indicate that at small scales species richness in mountain grasslands is determined by competition for light, which should be more important if nutrient availability is high, and by positive and negative interactions with unpalatable plants. In contrast, species richness at the large scale appears to be mainly influenced by land use. This result emphasizes the importance of studying such inter-relationships at multiple scales. Our study further suggests that the maintenance of the traditional land use scheme is crucial for the conservation of plant species richness of mountain pastures as both intensification and abandonment changed species composition and reduced plant species diversity.

Inter-specific competition from fallow deer Dama dama reduces habitat quality for the Italian roe deer Capreolus capreolus italicus

Abstract: Focardi, S., Aragno, P., Montanaro, P. and Riga, F. 2006. Inter-specific competitionfrom fallow deer Dama dama reduces habitat quality for the Italian roe deer Capreoluscapreolus italicus. Ecography 29: 407 417.The roe deer of Mediterranean habitats in the central and southern parts of Italyhas recently been recognised as a distinct subspecies, Capreolus capreolus italicus.A population of this endangered subspecies has been monitored in the Preserve ofCastelporziano, near Rome, since 1988. We observed an abrupt population decline in2000, which may severely threaten the sustainability of this population. We evaluatedthe hypothesis that competition by fallow deer may be a principal cause of this decline.By a new and innovative methodology, we modelled the spatial distribution of fallowdeer density (FDD) in the study area to show that 1) habitat quality for roe deer was aninverse function of FDD, 2) habitat apportionment between fallow and roe deerincreased as a function of FDD and by applying structural equation modelling 3)FDD was superior to habitat composition in explaining observed variations in homerange size and probably in habitat quality for roe deer. This analysis is the first todocument that inter-specific competition may influence the spatial behaviour of a deerspecies leading to poor phenotypic performance in the inferior competitor. Weconclude that the conservation of this relict population would benefit by reducingfallow deer numbers at Castelporziano and from other measures aimed to decrease thelevel of inter-specific competition.S. Focardi (stefano.focardi@infs.it), P. Aragno, P. Montanaro and F. Riga, IstitutoNazionale per la Fauna Selvatica, via Ca’ Fornacetta 9, IT-40064 Ozzano dell’ Emilia,Italy.

Null models for species co-occurrence patterns: assessing bias and minimum iteration number for the sequential swap

Abstract: The analysis of co-occurrence matrices is a common practice to evaluate community structure. The observed data are compared with a "null model", a randomised co-occurrence matrix derived from the observation by using a statistic, e.g. the C-score, sensitive to the pattern investigated. The most frequently used algorithm, "sequential swap", has been criticised for not sampling with equal frequencies thereby calling into question the results of earlier analysis. The bias of the "sequential swap" algorithm when used with the C-score was assessed by analysing 291 published presence-absence matrices. In 152 cases, the true p-value differed by > 5% from the p-value generated by an uncorrected "sequential swap". However, the absolute value of the difference was rather small. Out of the 291 matrices, there were only 5 cases in which an incorrect statistical decision would have been reached by using the uncorrected p-value (3 at the p <0.05 and 2 at the p <0.01 level), and in all 5 of these cases, the true p-value was close to the significance level. Our results confirm analytical studies of Miklos and Podani which show that the uncorrected swap gives slightly conservative results in tests for competitive segregation. However, the bias is very small and should not distort the ecological interpretation. We also estimated the number of iterations needed for the "sequential swap" to generate accurate p-values. While most authors do not exceed a number of 104 iterations, the suggested minimum number of swaps for 29 out of the 291 tested matrices is greater than 104. We recommend to use 30 000 "sequential swaps" if the required sample size is not assessed otherwise.