Evidence for large-scale effects of competition: niche displacement in Canada lynx and bobcat

Determining the patterns, causes and consequences of character displacement is central to our understanding of competition in ecological communities. However, the majority of competition research has occurred over small spatial extents or focused on fine-scale differences in morphology or behaviour. The effects of competition on broad-scale distribution and niche characteristics of species remain poorly understood but critically important. Using range-wide species distribution models, we evaluated whether Canada lynx (Lynx canadensis) or bobcat (Lynx rufus) were displaced in regions of sympatry. Consistent with our prediction, we found that lynx niches were less similar to those of bobcat in areas of sympatry versus allopatry, with a stronger reliance on snow cover driving lynx niche divergence in the sympatric zone. By contrast, bobcat increased niche breadth in zones of sympatry, and bobcat niches were equally similar to those of lynx in zones of sympatry and allopatry. These findings suggest that competitively disadvantaged species avoid competition at large scales by restricting their niche to highly suitable conditions, while superior competitors expand the diversity of environments used. Our results indicate that competition can manifest within climatic niche space across species’ ranges, highlighting the importance of biotic interactions occurring at large spatial scales on niche dynamics.     

The niche, biogeography and species interactions

In this paper, I review the relevance of the niche to biogeography, and what biogeography may tell us about the niche. The niche is defined as the combination of abiotic and biotic conditions where a species can persist. I argue that most biogeographic patterns are created by niche differences over space, and that even ‘geographic barriers’ must have an ecological basis. However, we know little about specific ecological factors underlying most biogeographic patterns. Some evidence supports the importance of abiotic factors, whereas few examples exist of large-scale patterns created by biotic interactions. I also show how incorporating biogeography may offer new perspectives on resource-related niches and species interactions. Several examples demonstrate that even after a major evolutionary radiation within a region, the region can still be invaded by ecologically similar species from another clade, countering the long-standing idea that communities and regions are generally ‘saturated’ with species. I also describe the somewhat paradoxical situation where competition seems to limit trait evolution in a group, but does not prevent co-occurrence of species with similar values for that trait (called here the ‘competition–divergence–co-occurrence conundrum’). In general, the interface of biogeography and ecology could be a major area for research in both fields.     

Ecological Niche Transferability Using Invasive Species as a Case Study

Species distribution modeling is widely applied to predict invasive species distributions and species range shifts under climate change. Accurate predictions depend upon meeting the assumption that ecological niches are conserved, i.e., spatially or temporally transferable. Here we present a multi-taxon comparative analysis of niche conservatism using biological invasion events well documented in natural history museum collections. Our goal is to assess spatial transferability of the climatic niche of a range of noxious terrestrial invasive species using two complementary approaches. First we compare species’ native versus invasive ranges in environmental space using two distinct methods, Principal Components Analysis and Mahalanobis distance. Second we compare species’ native versus invaded ranges in geographic space as estimated using the species distribution modeling technique Maxent and the comparative index Hellinger’s I. We find that species exhibit a range of responses, from almost complete transferability, in which the invaded niches completely overlap with the native niches, to a complete dissociation between native and invaded ranges. Intermediate responses included expansion of dimension attributable to either temperature or precipitation derived variables, as well as niche expansion in multiple dimensions. We conclude that the ecological niche in the native range is generally a poor predictor of invaded range and, by analogy, the ecological niche may be a poor predictor of range shifts under climate change. We suggest that assessing dimensions of niche transferability prior to standard species distribution modeling may improve the understanding of species’ dynamics in the invaded range.     

The Geographic Distribution of a Tropical Montane Bird Is Limited by a Tree: Acorn Woodpeckers (Melanerpes formicivorus) and Colombian Oaks (Quercus humboldtii) in the Northern Andes

Species distributions are limited by a complex array of abiotic and biotic factors. In general, abiotic (climatic) factors are thought to explain species’ broad geographic distributions, while biotic factors regulate species’ abundance patterns at local scales. We used species distribution models to test the hypothesis that a biotic interaction with a tree, the Colombian oak (Quercus humboldtii), limits the broad-scale distribution of the Acorn Woodpecker (Melanerpes formicivorus) in the Northern Andes of South America. North American populations of Acorn Woodpeckers consume acorns from Quercus oaks and are limited by the presence of Quercus oaks. However, Acorn Woodpeckers in the Northern Andes seldom consume Colombian oak acorns (though may regularly drink sap from oak trees) and have been observed at sites without Colombian oaks, the sole species of Quercus found in South America. We found that climate-only models overpredicted Acorn Woodpecker distribution, suggesting that suitable abiotic conditions (e.g. in northern Ecuador) exist beyond the woodpecker’s southern range margin. In contrast, models that incorporate Colombian oak presence outperformed climate-only models and more accurately predicted the location of the Acorn Woodpecker’s southern range margin in southern Colombia. These findings support the hypothesis that a biotic interaction with Colombian oaks sets Acorn Woodpecker’s broad-scale geographic limit in South America, probably because Acorn Woodpeckers rely on Colombian oaks as a food resource (possibly for the oak’s sap rather than for acorns). Although empirical examples of particular plants limiting tropical birds’ distributions are scarce, we predict that similar biotic interactions may play an important role in structuring the geographic distributions of many species of tropical montane birds with specialized foraging behavior.     

Biology of gastrointestinal nematodes of ruminants

The development and survival of free-living stages of gastro-intestinal nematodes of small ruminants are influenced by several abiotic and biotic factors. Within the abiotic factors, most important are the environmental temperature and humidity. They regulate the development of larvae from eggs dispersed on the pasture by the animals faeces. Each parasite species that infect ruminants requires a different time to development, depending on temperature and humidity. Among trichostrongylids, Ostertagia, Teladorsagia and Nematodirus show a strong adaptation to low temperatures. Nematodirus larvae are able to survive to winter inside the egg shell. Temperature and humidity influence the distribution and survival of larvae on pasture. The larval third stage can migrate from faeces to pasture vegetation and they accumulate at the basis of vegetation where stay during the day or in the soil to avoid the desiccation. The forage species affects the migration of larvae on herbage too. Many biological factors contribute to disperse the larvae on the pasture. Dung burying beetles, coprophagous beetles and earthworms can greatly reduce the larvae of some trichostrongylids on pasture. They contribute to the spread of the faecal material on the pasture and allow the larval death as a consequence of drying.     

Scale dependence of temperature as an abiotic driver of species' distributions

Aim Scale dependence of patterns and processes remains one of the major unresolved problems in ecology. The responses of ecosystems to environmental stressors are reported to be strongly scale dependent, but projections of the effects of climate change on species' distributions are still restricted to particular scales and knowledge about scale dependence is lacking. Here we propose that the scale dependence of those species' niche dimensions related to climate change is strongly related to the strength of climatic cross‐scale links. More specifically, we hypothesize that the strong cross‐scale links between micro‐ and macroclimatic conditions are related to high cross‐scale similarity (low scale dependence) of species' realized temperature niches and, thus, species' spatial distributions. Location This study covers seven orders of magnitude of spatial scale, ranging from local‐scale (below a metre) and regional‐scale (kilometre) investigations in central European wetland ecosystems to continental‐scale (thousands of kilometres) studies of species' distributions. Methods We combined data on the spatial occurrence of species (vegetation records at local and regional scales, digitized distribution maps at the continental scale) with information about the corresponding temperature regime of vascular plant species occurring in environmentally stable wetland ecosystems characterized by strong cross‐scale links between micro‐ and macroclimatic conditions. Results We observed high cross‐scale similarity of the characteristics of species temperature niches across seven orders of magnitude of spatial scale. However, the importance of temperature as an abiotic driver decreased nonlinearly with decreasing scale, suggesting greater importance of additional (biotic) drivers of species' occurrence at small spatial scales. Main conclusions We report high cross‐scale similarity of realized temperature niches for species inhabiting ecosystems where small‐scale environmental noise is low and cross‐scale links between micro‐ and macroclimatic conditions are strong. By highlighting a strong relationship between abiotic and biotic cross‐scale similarity, our results will help to improve niche‐based species distribution modelling, one of the major assessment tools for determining the ecological effects of climate change.     

Integrating demography,dispersal and interspecific interactions into bird distribution models

Species’ ranges are primarily limited by the physiological (abiotic) tolerance of the species, described by their fundamental niche. Additionally, demographic processes, dispersal, and interspecific interactions with other species are shaping species distributions, resulting in the realised niche. Understanding the complex interplay between these drivers is vital for making robust biodiversity predictions to novel environments. Correlative species distribution models have been widely used to predict biodiversity response but also remain criticised, as they are not able to properly disentangle the abiotic and biotic drivers shaping species’ niches. Recent developments have thus focussed on 1) integrating demography and dispersal into species distribution models, and on 2) integrating interspecific interactions. Here, I review recent demographic and multi‐species modelling approaches and discuss critical aspects of these models that remain underexplored in general and in respect to birds, for example, the complex life histories of birds and other animals as well as the scale dependence of interspecific interactions. I conclude by formulating modelling guidelines for integrating the abiotic and biotic processes that limit species’ ranges, which will help to disentangle the complex roles of demography, dispersal and interspecific interactions in shaping species niches. Throughout, I pinpoint complexities of avian life cycles that are critical for consideration in the models and identify data requirements for operationalizing the different modelling steps.     

Habitat shifts of endangered species under altered climate conditions: importance of biotic interactions

Predicting changes in potential habitat for endangered species as a result of global warming requires considering more than future climate conditions; it is also necessary to evaluate biotic associations. Most distribution models predicting species responses to climate change include climate variables and occasionally topographic and edaphic parameters, rarely are biotic interactions included. Here, we incorporate biotic interactions into niche models to predict suitable habitat for species under altered climates. We constructed and evaluated niche models for an endangered butterfly and a threatened bird species, both are habitat specialists restricted to semiarid shrublands of southern California. To incorporate their dependency on shrubs, we first developed climate‐based niche models for shrubland vegetation and individual shrub species. We also developed models for the butterfly's larval host plants. Outputs from these models were included in the environmental variable dataset used to create butterfly and bird niche models. For both animal species, abiotic–biotic models outperformed the climate‐only model, with climate‐only models over‐predicting suitable habitat under current climate conditions. We used the climate‐only and abiotic–biotic models to calculate amounts of suitable habitat under altered climates and to evaluate species' sensitivities to climate change. We varied temperature (+0.6, +1.7, and +2.8 °C) and precipitation (50%, 90%, 100%, 110%, and 150%) relative to current climate averages and within ranges predicted by global climate change models. Suitable habitat for each species was reduced at all levels of temperature increase. Both species were sensitive to precipitation changes, particularly increases. Under altered climates, including biotic variables reduced habitat by 68–100% relative to the climate‐only model. To design reserve systems conserving sensitive species under global warming, it is important to consider biotic interactions, particularly for habitat specialists and species with strong dependencies on other species.     

Using ecological niche modelling to evaluate niche stability in deep time

Aim To investigate relative niche stability in species responses to various types of environmental pressure (biotic and abiotic) on geological time‐scales using the fossil record. Location The case study focuses on Late Ordovician articulate brachiopods of the Cincinnati Arch in eastern North America. Methods Species niches were modelled for a suite of fossil brachiopod species based on five environmental variables inferred from sedimentary parameters using GARP and Maxent . Niche stability was assessed by comparison of (1) the degree of overlap of species distribution models developed for a time‐slice and those generated by projecting niche models of the previous time‐slice onto environmental layers of a second time‐slice using GARP and Maxent , (2) Schoener’s D statistic, and (3) the similarity of the contribution of each environmental parameter within Maxent niche models between adjacent time‐slices. Results Late Ordovician brachiopod species conserved their niches with high fidelity during intervals of gradual environmental change but responded to inter‐basinal species invasions through niche evolution. Both native and invasive species exhibited similar levels of niche evolution in the invasion and post‐invasion intervals. Niche evolution was related mostly to decreased variance within the former ecological niche parameters rather than to shifts to new ecospace. Main conclusions Although the species examined exhibited morphological stasis during the study interval, high levels of niche conservatism were observed only during intervals of gradual environmental change. Rapid environmental change, notably inter‐basinal species invasions, resulted in high levels of niche evolution among the focal taxa. Both native and invasive species responded with similar levels of niche evolution during the invasion interval and subsequent environmental reorganization. The assumption of complete niche conservatism frequently employed in ecological niche modelling (ENM) analyses to forecast or hindcast species geographical distributions is more likely to be accurate for climate change studies than for invasive species analyses over geological time‐scales.     

Incidence of leaf mining in different vegetation types across rainfall,canopy cover and latitudinal gradients

Abstract Leaf miners are insects whose larval stages live between layers of leaf epidermis, feeding on mesophyll and lower epidermis to create mine‐like cavities. Little is known about the ecology or distribution of leaf miners in Australia. We investigated the incidence of leaf miners in relation to aridity, vegetation types, host plant taxonomy, leaf traits, canopy cover and latitude. We surveyed leaf miners at 15 sites in NSW, eastern Australia, situated along a rainfall gradient from 300 to 1700 mm per annum and a latitudinal gradient of 28°S to 33°S, within four vegetation types (mallee, heath, woodland and rainforest). Leaf mining was recorded from 36 plant species, 89% of which had no previous record of mining. The proportion of mined plant species at each site varied, but there was no significant difference between vegetation types. Leaf mining presence was positively correlated with both total leaf length and leaf thickness. No significant correlations were found between the proportion of mined species at a site and rainfall, latitude or foliar projected cover. We conclude that leaf mining is a widespread type of insect herbivory whose distribution patterns are more likely to be influenced by biotic than abiotic factors.     

Niche-habitat mechanisms and biotic interactions explain the coexistence and abundance of congeneric sandgrouse species

Ascertaining which niche processes allow coexistence between closely related species is of special interest in ecology. We quantified variations in the environmental niches and densities of two congeneric species, the pin-tailed and the black-bellied sandgrouse (Pterocles alchata and Pterocles orientalis) in allopatry and sympatry under similar abiotic, habitat and dispersal contexts to understand their coexistence. Using principal component analysis, we defined environmental gradients (niche dimensions) including abiotic, habitat and anthropogenic variables, and calculated niche breadth, position and overlap of both species in sympatry and allopatry. Additionally, sandgrouse density was modelled as a function of the niche dimensions and the density of the other species. We found evidence that each species occupies distinct environmental niches in sympatry and in allopatry. The black-bellied sandgrouse exploits a broader range of environmental conditions (wider niche breadth) while the pin-tailed sandgrouse reaches high densities where conditions seem to match its optimum. In sympatry, both species shift their niches to intermediate positions, indicating the importance of abiotic factors in setting coexistence areas. Environmental conditions determine regional densities of pin-tailed sandgrouse whereas biotic interactions explain the density of the black-bellied sandgrouse in areas with abiotic conditions similarly conducive for both species. Highly suitable areas for the pin-tailed sandgrouse fall beyond the upper thermal limit of the black-bellied sandgrouse, leading to niche segregation and low densities for the latter. Finally, local niche shift and expansion plus possible heterospecific aggregation allow the pin-tailed sandgrouse to thrive in a priori less favourable environments. This work provides insight into how different mechanisms allow species coexistence and how species densities vary in sympatry compared to allopatry as a result of environmental filtering and biotic interactions.     

Influence of soil and microclimate on species composition and grass encroachment in heath succession

Aims and Methods Mostly due to land use changes, European heathlands have become increasingly rare. In addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses and a loss in species diversity. Despite many investigations, information about the precise environmental parameters that determine the development and maintenance of heathland vegetation is still insufficient. In order to determine the environmental factors that control heath succession and grass encroachment, and to develop appropriate management schemes, we studied the influence of several soil and microclimate parameters on species composition and vegetation characteristics in five successional stages in a coastal heathland on the island of Hiddensee, north-east Germany, where the encroachment of Carex arenaria has become a major problem.Important findings We recorded the highest plant species richness in grey dune and birch forest plots, while the encroachment of C. arenaria let to a significant decline in plant species richness. The most important environmental factors influencing species richness and distribution of single species were microclimate, soil moisture, soil pH and the C/N ratio. While many studies reported the importance of differences in nutrient availability, we found no significant correlations between soil nutrient availability and vegetation pattern. Environmental conditions in dense C. arenaria stands, especially soil properties (e.g. soil pH), showed great differences in comparison to the other successional stages. However, no correlations between the encroachment of C. arenaria and single environmental factors were found. Our results show that not only soil nutrients are important abiotic factors in heaths but that also microclimate and soil moisture play an important role and that many factors are involved in heath succession and in the promotion of grass encroachment. Management plans for the conservation and restoration of heathlands should therefore focus on the specific site conditions and should take several abiotic and biotic factors into account.     

Environmental factors associated with the distribution of Anopheles arabiensis and Culex quinquefasciatus in a rice agro-ecosystem in Mwea, Kenya

Studies were conducted between May and June, 2006 to investigate the environmental factors affecting the distribution of An. arabiensis Patton and Culex quinquefasciatus Say in Mwea, Kenya. The sampling unit comprised all non-paddy aquatic habitats and ten randomly selected paddies and canals located within a 200 m radius from the periphery of the study site. Thirteen physico-chemical variables were recorded for each sampling site in each sampling occasion and a sample of mosquito larvae and other aquatic invertebrates collected. The non-paddy aquatic habitats identified included pools and marshes. Morphological identification of 1,974 mosquito larvae yielded four species dominated by Cx. quinquefasciatus (73.2%) and An. arabiensis (25.0%). Pools were associated with significantly higher Cx. quinquefasciatus larval abundance and less diversity of other aquatic invertebrates compared with other habitat types. In contrast, the abundance of An. arabiensis did not differ significantly among habitat types. Culex quinquefasciatus habitats had higher water conductivity and exhibited a higher abundance of other aquatic invertebrates than An. arabiensis habitats. Chi-square analysis indicated that the two species were more likely to coexist in the same habitats than would be expected by chance alone. Anopheles arabiensis larvae were positively associated with dissolved oxygen and adults of family Haliplidae and negatively associated with emergent vegetation and Heptageniidae larvae. Culex quinquefasciatus larvae were positively associated with dissolved oxygen, total dissolved solids, Chironomidae larvae, and Microvelidae adults and negatively associated with emergent vegetation. These findings suggest that both biotic and abiotic factors play a significant role in niche partitioning among Cx. quinquefasciatus and An. arabiensis, a factor that should be considered when designing an integrated vector control program.     

The role of climate for the range limits of parapatric European land salamanders

Abrupt range limits of parapatric species may serve as a model system to understand the factors that determine species’ range borders. Theory suggests that parapatric range limits can be caused by abiotic conditions along environmental gradients, biotic interactions or a combination of both. Geographic ranges of the parapatric salamanders, Salamandra salamandra and S. atra, meet in small contact zones in the European Alps and to date, the cause of parapatry and the restricted range of S. atra remain elusive. We combine multivariate approaches and climatic data analysis to explore niche differentiation among the two salamanders with respect to the available climatic environment at their contact zones. Our purpose is to evaluate whether climatic conditions explain the species’ sharp range limits or if biotic interactions may play a role for range delimitation. Analyses were carried out in three contact zones in Switzerland to assess possible geographic variation. Our results indicate that both species occur at localities with different climatic conditions as well as the presence of a strong climatic gradient across the species’ range limits. Although the species’ climatic niches differ moderately (with a wider niche breadth for S. atra), interspecific niche overlap is found. Comparisons among the contact zones confirm geographic variation in the species’ climatic niches as well as in the conditions within the geographically available space. Our results suggest that the change in climatic conditions along the recognized gradient represents a determining factor for species’ range limits within contact zones. However, our analyses of geographic variation in climatic conditions reveal that both salamander species can occur in a much wider range of conditions than observed within contact zones. This finding and the interspecific climatic niche overlap within each contact zone provides indirect evidence that biotic interactions (likely competition) between the two species may also determine their range limits.     

Using functional traits and phylogeny to understand local extinction risk in dragonflies and damselflies (Odonata)

Understanding the risk of local extinction of a species is vital in conservation biology, especially now when anthropogenic disturbances and global warming are severely changing natural habitats. Local extinction risk depends on species traits, such as its geographical range size, fresh body mass, dispersal ability, length of flying period, life history variation, and how specialized it is regarding its breeding habitat. We used a phylogenetic approach because closely related species are not independent observations in the statistical tests. Our field data contained the local extinction risk of 31 odonate (dragonflies and damselflies) species from Central Finland. Species relatedness (i.e., phylogenetic signal) did not affect local extinction risk, length of flying period, nor the geographical range size of a species. However, we found that closely related species were similar in hind wing length, length of larval period, and habitat of larvae. Both phylogenetically corrected (PGLS) and uncorrected (GLM) analysis indicated that the geographical range size of species was negatively related to local extinction risk. Contrary to expectations, habitat specialist species did not have higher local extinction rates than habitat generalist species nor was it affected by the relatedness of species. As predicted, species’ long larval period increased, and long wings decreased the local extinction risk when evolutionary relatedness was controlled. Our results suggest that a relatively narrow geographical range size is an accurate estimate for a local extinction risk of an odonate species, but the species with long life history and large habitat niche width of adults increased local extinction risk. Because the results were so similar between PGLS and GLM methods, it seems that using a phylogenetic approach does not improve predicting local extinctions.     

The genomic code: inferring Vibrionaceae niche specialization

The Vibrionaceae show a wide range of niche specialization, from free-living forms to those attached to biotic and abiotic surfaces, from symbionts to pathogens and from estuarine inhabitants to deep-sea piezophiles. The existence of complete genome sequences for closely related species from varied aquatic niches makes this group an excellent case study for genome comparison.     

Coexistence in a guild of filter feeding caddis larvae: do different instars act as different species?

Summary Guild structure and coexistence of filter feeding caddis larvae was studied in three lake-outlet streams in northern Finland. Natural stones were used as sampling units, and for each stone measurements of six microhabitat variables were taken. Species and species instar relationships in the multivariate microhabitat space were analyzed by a method based on Principal Components Analysis. Seven species of filtering caddis larvae were found at the study sites, but detailed analysis was restricted to the four most abundant ones (Polycentropus flavomaculatus, Hydropsyche angustipennis, H. pellucidula, H. saxonica). P. flavomaculatus was the microhabitat generalist of the guild, whereas all the hydropsychids were narrowly specialized in at least one larval instar. In all species, different instars utilized the microhabitat somewhat differentially. Moss content and Froude number were significantly different between instar microhabitats in all comparisons. Ontogenetic niche changes not only separated species instars from each other, but also profoundly changed the biotic as well as abiotic environment of the species at different phases of larval development. This may have important effects on the dynamics of interacting species and, consequently, on the guild structure. While some of the more classical models of coexistence may also be relevant for understanding coexistence in this guild of sessile stream insects, ontogenetic niche changes clearly need to be taken into consideration in future studies of guild structure and species interactions of filter feeding caddisfly larvae.     

Does post-fire abiotic habitat filtering create divergent plant communities in black spruce forests of eastern Canada?

We investigated the role of post-fire residual organic matter (ROM) thickness as a driver of community assembly in eastern Newfoundland. We hypothesized that if post-fire community assembly is predominantly controlled by ROM thickness (an abiotic habitat filter), then post-fire species composition and functional traits should correspond to the depth and distribution of ROM. However, if species interactions (biotic filter) are the primary constraints on community assembly, then post-fire species composition and their functional traits should be independent of the depth and distribution of ROM. We tested these predictions in three relatively mature plant communities, Kalmia angustifolia heath, black spruce (Picea mariana)-Kalmia shrub savannah and black spruce forest. Through pre-fire stand reconstruction, we found evidence that the three communities originated from black spruce forest. ROM thickness in heath was almost twice that of shrub savannah and six times more than forest, suggesting a gradient in fire severity. Distribution of ROM corresponded to patterns in vegetation dominance, where thick ROM (>2 cm) filtered out black spruce in favour of Kalmia. ROM thickness was a strong predictor of vegetation composition and function between heath and forest, but this was not found between the shrub savannah and forest. We attribute this to species interactions and allelopathy, which may have become important when ROM thickness was suitable for both seed (black spruce) and vegetative (Kalmia) regenerating species. Thus, priority effects or “who came first” may have lead to shrub savannah formation when ROM thickness was ~2 cm. We conclude that abiotic habitat filtering of thick ROM (>2 cm) on (primarily) species’ regeneration traits was the primary driver of community divergence from forest to heath and shrub savannah.     

Factors regulating the invasive success of an alien frog: a comparison of the ecology of the native and alien populations

We examined niche occupancy of Discoglossus pictus, an anuran recently established in Europe, comparing the niches of native (North Africa) and alien populations (south-western Europe) at two spatial scales to determine whether adaptive divergence had occurred between these two populations. Additionally, we determine whether the alien species showed a wider larvae niche and higher phenotypic variability compared with co-occurring anurans. We characterized the breeding habitats and the climatic space occupied by native and alien groups of populations of D. pictus and examined morphological traits of D. pictus and sympatric anuran larvae. Our results revealed no divergence in breeding habitat use between native and alien populations. A shift was observed between the realized niches occupied by the native and alien populations, but this shift might only reflect cryptic niche conservatism. The range of reproductive habitats selected by D. pictus was not wider than those of most native species. In the invaded range, D. pictus showed morphological overlap with some native species and broader phenotypic variability, but the adaptive advantages of this latter attribute were uncertain. Our results suggest that the invasive capacity of this species depends on favourable abiotic conditions rather than on its adaptive advantages over native anurans.     

Plant growth response to direct and indirect temperature effects varies by vegetation type and elevation in a subarctic tundra

There has been growing recent use of elevational gradients as tools for assessing effects of temperature changes on vegetation properties, because these gradients enable temperature effects to be considered over larger spatial and temporal scales than is possible through conventional experiments. While many studies have explored the direct effects of temperature, the indirect effects of temperature through its long‐term influence on soil abiotic or biotic properties remain essentially unexplored. We performed two climate chamber experiments using soils from a subarctic elevational gradient in Abisko, Sweden to investigate the direct effects of temperature, and indirect effects of temperature via soil legacies, on growth of two grass species. The soils were collected from each of two vegetation types (heath, dominated by dwarf shrubs, and meadow, dominated by graminoids and herbs) at each of three elevations. We found that plants responded to both the direct effect of temperature and its indirect effect via soil legacies, and that direct and indirect effects were largely decoupled. Vegetation type was a major determinant of plant responses to both the direct and indirect effects of temperature; responses to soils from increasing elevation were stronger and showed a more linear decline for meadow than for heath soils. The influence of soil biota on plant growth was independent of elevation, with a positive influence across all elevations regardless of soil origin for meadow soils but not for heath soils. Taken together, this means that responses of plant growth to soil legacy effects of temperature across the elevational gradient were driven primarily by soil abiotic, and not biotic, factors. These findings emphasize that vegetation type is a strong determinant of how temperature variation across elevational gradients impacts on plant growth, and highlight the need for considering both direct and indirect effects of temperature on plant responses to future climate change.