Contextualized niche shifts upon independent invasions by the dung beetle <Emphasis Type="Italic">Onthophagus taurus</Emphasis>

The historical contingencies of biological invasions may have important consequences for final invasion outcomes. Here, we characterize the variations in the realized niche during the invasions of the bull-headed dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae) from its native Mediterranean range following accidental (Eastern North America) as well as deliberate (Western North America, Western Australia, and Eastern Australia) releases into novel, exotic ranges approximately 50 years ago. Specifically, we examined whether the climatic responses of exotic O. taurus have diverged from those characterizing their native range, and if so, to what degree and in what dimensions. We found that when compared to the native range, all exotic populations exhibited similar overlap proportions regardless of invasion history. However, more detailed analysis of climatic niche features showed that all three deliberately established populations were characterized by overall similar climatic niche features, whereas the accidentally-established Eastern North American populations have undergone significant changes in their climatic niche. Specifically, when analog climates were considered on the background of each pairwise range comparison, accidentally-established Eastern North American populations showed a different climatic niche expansion than their deliberately introduced Australian or Western North American counterparts, in particular towards colder and more humid climates. We discuss our results in the context of the widely divergent introduction histories of O. taurus in Australia and North America, and highlight the possible roles of contrasting propagule sizes, disparate genetic profiles and variances, adaptive processes and invadable landscapes in shaping invasion outcomes in the different exotic ranges.     

Patterns of niche filling and expansion across the invaded ranges of an Australian lizard

Studies of realized niche shifts in alien species typically ignore the potential effects of intraspecific niche variation and different invaded‐range environments on niche lability. We incorporate our detailed knowledge of the native‐range source populations and global introduction history of the delicate skink Lampropholis delicata to examine intraspecific variation in realized niche expansion and unfilling, and investigate how alternative niche modelling approaches are affected by that variation. We analyzed the realized niche dynamics of L. delicata using an ordination method, ecological niche models (ENMs), and occurrence records from 1) Australia (native range), 2) New Zealand, 3) Hawaii, 4) the two distinct native‐range clades that were the sources for the New Zealand and Hawaii introductions, and 5) the species’ global range (including Lord Howe Island, Australia). We found a gradient of realized niche change across the invaded ranges of L. delicata: niche stasis on Lord Howe Island, niche unfilling in New Zealand (16%), and niche unfilling (87%) and expansion (14%) in Hawaii. ENMs fitted to native‐range data generally identified suitable climatic conditions at sites where the species has established non‐native populations, whereas ENMs based on native‐range source clades and non‐native populations had lower spatial transferability. Our results suggest that the extent to which realized niches are maintained during invasion does not depend on species‐level traits. When realized niche shifts are predominately due to niche unfilling, fully capturing species’ responses along climatic gradients by basing ENMs on native distributions may be more important for accurate invasion forecasts than incorporating phylogenetic differentiation, or integrating niche changes in the invaded range.     

Climatic niche evolution in the Andean genus <Emphasis Type="Italic">Menonvillea</Emphasis> (Cremolobeae: Brassicaceae)

The study of how climatic niches change over evolutionary time has recently attracted the interest of many researchers. Different methodologies have been employed principally to analyze the temporal dynamics of the niche and specially to test for the presence of phylogenetic niche conservatism. Menonvillea, a genus of Brassicaceae including 24 species, is distributed primarily along the Andes of Argentina and Chile, with some taxa growing in southern Patagonia and others in the Atacama Desert and the Chilean Matorral. The genus is highly diversified morphologically but also presents a remarkably wide ecological range, growing from the high Andean elevations, to the dry coastal deserts in Chile, or the Patagonia Steppe in Argentina. In this study, we used molecular phylogenies together with climatic data to study climatic niche evolution in the genus. The results show that the main climatic niche shifts in Menonvillea occurred between the sections Cuneata-Scapigera and sect. Menonvillea throughout the Mid-Late Miocene, and associated with the two main geographical distribution centers of the genus: the highlands of the central-southern Andes and the Atacama Desert-Chilean Matorral, respectively. Climatic niches in these lineages were mainly differentiated by the aridity and potential evapotranspiration, the minimum temperatures of the coldest month, and the temperature annual range and seasonality. Niche evolution in Menonvillea deviated from a Brownian motion process, with most of the climatic dimension best-fitting to an Ornstein-Uhlenbeck model of multiple adaptive peaks. Our results also indicated that higher aridity levels and lower annual temperature ranges were associated with the evolution of the annual habit, as exemplified by the distribution of sect. Menonvillea. Finally, the results suggested that climatic niche evolution in Menonvillea exhibited some degree of phylogenetic niche conservatism, fundamentally within the two main lineages (sect. Menonvillea and sects. Cuneata-Scapigera).     

Unresolved native range taxonomy complicates inferences in invasion ecology: <Emphasis Type="Italic">Acacia dealbata</Emphasis> Link as an example

Elaborate and expensive endeavours are underway worldwide to understand and manage biological invasions. However, the success of such efforts can be jeopardised due to taxonomic uncertainty. We highlight how unresolved native range taxonomy can complicate inferences in invasion ecology using the invasive tree Acacia dealbata in South Africa as an example. Acacia dealbata is thought to comprise two subspecies based on morphological characteristics and environmental requirements within its native range in Australia: ssp. dealbata and spp. subalpina. Biological control is the most promising option for managing invasive A. dealbata populations in South Africa, but it remains unknown which genetic/taxonomic entities are present in the country. Resolving this question is crucial for selecting appropriate biological control agents and for identifying areas with the highest invasion risk. We used species distribution models (SDMs) and phylogeographic approaches to address this issue. The ability of subspecies-specific and overall species SDMs to predict occurrences in South Africa was also explored. Furthermore, as non-overlapping bioclimatic niches between the two taxonomic entities may translate into evolutionary distinctiveness, we also tested genetic distances between the entities using DNA sequencing data and network analysis. Both approaches were unable to differentiate the two putative subspecies of A. dealbata. However, the SDM approach revealed a potential niche shift in the non-native range, and DNA sequencing results suggested repeated introductions of different native provenances into South Africa. Our findings provide important information for ongoing biological control attempts and highlight the importance of resolving taxonomic uncertainties in invasion ecology.     

Acorns of invasive Northern Red Oak (<Emphasis Type="Italic">Quercus rubra</Emphasis>) in Europe are larval hosts for moths and beetles

In their first phase of expanding into new areas, invasive plants often take advantage of the inability of existing herbivores and pathogenic species to exploit them. However, in the longer term local enemies may adapt to using these invasive species as a food source. This study assesses the use of mature acorns of two oak species in Europe (the native Pedunculate Oak Quercus robur and the invasive Northern Red Oak Quercus rubra) by moths Cydia fagiglandana and Cydia splendana and beetles Curculio spp. We show that acorns of invasive oak species can be equally attractive to C. splendana but only partially so to C. fagiglandana where infestation rates where significantly lower (approximately half) compared to the native oak. The infestation by Curculio beetles of Northern Red Oak was marginal, less than 1% of the rate in the native oak species. The larval final weights did not differ significantly between host species, but emergence of C. splendana and Curculio spp. took significantly longer in acorns of Northern Red Oak. It is likely that C. fagiglandana and C. splendana have increased their niche breadths by exploiting invasive oak species and avoiding competition with the Curculio weevils. Furthermore, the occurrence of Northern Red Oak could stabilize food resources during years when native oak species have poor acorn crops.     

The invasion of the azooxanthellate coral <Emphasis Type="Italic">Tubastraea</Emphasis> (Scleractinia: Dendrophylliidae) throughout the world: history,pathways and vectors

In this review, we describe the history, pathways and vectors of the biological invasion of the azooxanthellate coral Tubastraea (Scleractinia: Dendrophylliidae) throughout the world. In order to do so we consulted previous reports in the literature and also compiled new unpublished information on the distribution of the three species of Tubastraea which have been reported as non indigenous species, both within their native and non-native ranges and also on vectors, and where cryptogenic. We combine these data with historical aspects of marine vectors in order to get insights into how Tubastraea species have successfully spread around the world, established and invaded and where future studies would be best focused. T. coccinea and T. tagusensis are recognized as being highly invasive and are causing significant environmental, economic, and social impacts requiring management actions. The third species, T. micranthus so far only reported outside its native range on oil platforms, may have similar potential for negative impact. The vectors of introduction of Tubastraea may have changed throughout history and the biological invasion of these invasive corals may reflect changing practices, demands and legislation in shipping activities over the years. Today it is clear that these corals are fouling organisms strongly associated with oil and gas platforms worldwide which are thus primary vectors for new introductions.     

Ecological niche differentiation between native and non-native shrimps in the northern Baltic Sea

Invasions of non-native species are modifying global biodiversity but the ecological mechanisms underlying invasion processes are still not well understood. A degree of niche separation of non-native and sympatric native species can possibly explain the success of novel species in their new environment. In this study, we quantified experimentally and in situ the environmental niche space of caridean shrimps (native Crangon crangon and Palaemon adspersus, non-native Palaemon elegans) inhabiting the northern Baltic Sea. Field studies showed that the non-native P. elegans had wider geographical range compared to native species although the level of habitat specialization was similar in both Palaemon species. There were clear differences in shrimp habitat occupancy with P. elegans inhabiting lower salinity areas and more eutrophicated habitats compared to the native species. Consequently, the non-native shrimp has occupied large areas of the northern Baltic Sea that were previously devoid of the native shrimps. Experiments demonstrated that the non-native shrimp had higher affinity to vegetated substrates compared to native species. The study suggests that the abilities of the non-native shrimp to thrive in more stressful habitats (lower salinity, higher eutrophication), that are sub-optimal for native shrimps, plausibly explain the invasion success of P. elegans.     

Community Impacts of Prosopis juliflora Invasion: Biogeographic and Congeneric Comparisons

We coordinated biogeographical comparisons of the impacts of an exotic invasive tree in its native and non-native ranges with a congeneric comparison in the non-native range. Prosopis juliflora is taxonomically complicated and with P. pallida forms the P. juliflora complex. Thus we sampled P. juliflora in its native Venezuela, and also located two field sites in Peru, the native range of Prosopis pallida. Canopies of Prosopis juliflora, a native of the New World but an invader in many other regions, had facilitative effects on the diversity of other species in its native Venezuela, and P. pallida had both negative and positive effects depending on the year, (overall neutral effects) in its native Peru. However, in India and Hawaii, USA, where P. juliflora is an aggressive invader, canopy effects were consistently and strongly negative on species richness. Prosopis cineraria, a native to India, had much weaker effects on species richness in India than P. juliflora. We carried out multiple congeneric comparisons between P. juliflora and P. cineraria, and found that soil from the rhizosphere of P. juliflora had higher extractable phosphorus, soluble salts and total phenolics than P. cineraria rhizosphere soils. Experimentally applied P. juliflora litter caused far greater mortality of native Indian species than litter from P. cineraria. Prosopis juliflora leaf leachate had neutral to negative effects on root growth of three common crop species of north-west India whereas P. cineraria leaf leachate had positive effects. Prosopis juliflora leaf leachate also had higher concentrations of total phenolics and L-tryptophan than P. cineraria, suggesting a potential allelopathic mechanism for the congeneric differences. Our results also suggest the possibility of regional evolutionary trajectories among competitors and that recent mixing of species from different trajectories has the potential to disrupt evolved interactions among native species.     

Echoes of the whispering land: interacting roles of vicariance and selection in shaping the evolutionary divergence of two <Emphasis Type="Italic">Calceolaria</Emphasis> (Calceolariaceae) species from Patagonia and Malvinas/Falkland Islands

A key to understanding the origin and identity of young species lays on the knowledge of the Quaternary climatic oscillations’ effect on gene flow and vicariance. Even though the effect of climatic fluctuations is relatively well understood for southern hemisphere plant species, little is known about their effect on the evolutionary histories of species from mainland and islands. Thus, we investigated whether Quaternary climate-driven fluctuations translated into lineage divergence and speciation, followed or not by climatic niche differentiation, in two allopatric plant species, Calceolaria uniflora and C. fothergillii from Patagonia and Malvinas/Falkland islands, respectively. We sampled the range of both species, and sequenced two chloroplastic (cpDNA; trnS–trnG and trnH–psbA), and one single copy “anonymous” non-coding nuclear region (nDNA). We performed phylogeographic and dating analyses, and adjusted spatio-temporal diffusion models. We complemented molecular evidence with climatic niche differentiation analyses and species paleo-distribution projections. A species coalescent reconstruction based on multi-locus data retrieved both species as monophyletic. Estimates from cpDNA indicated the species diverged during the Great Patagonian Glaciation. Chloroplast and nuclear DNA showed east–west distribution of the main genetic groups but with contrasting spatial genetic diversity. The spatio-temporal diffusion analyses showed that between 1–0.8 Mya and 570 Kya the lineage leading to C. fothergillii diverged from C. uniflora and arrived to the islands. Climatic niche projections hindcasted range expansions during glaciations, and contractions during the interglacial periods. Comparisons of climatic niches between the two study species indicated that temperature variables show evidence of niche conservatism while precipitation regimes supported niche divergence, even when considering the background environmental divergence. Our study indicates that glacial fluctuations affected the mainland/islands connections favouring speciation mediated not only by isolation, but also by climatic niche differentiation.     

Climatic Similarity of Extant and Extinct <Emphasis Type="Italic">Dasypus</Emphasis> Armadillos

The similar geographic distributions of an extinct (Dasypus bellus) and an extant (D. novemcinctus) armadillo species have long been of interest to scholars because of the unresolved phylogeny. The relationship between the two species has been investigated through morphological and phylogenetic studies, whereas the ecological perspective has been overlooked, the importance of which is more and more acknowledged in speciation events. Here, we used ecological niche models to study the climatic niche similarity of three species of Dasypus (D. bellus, D. novemcinctus, and D. kappleri) and provide new insights on the relationship among them. The climatic niche similarity was compared in two ways: hindcast of ecological niche models based on occurrences and climatic layers, and direct niche boundary comparison along bioclimatic axes. The fossil records of D. bellus were not predicted suitable by the ecological niche models of the two extant armadillos. The direct comparison of niche boundary showed that D. bellus lived in colder and relative dryer climates, with high temperature variation and low precipitation variation. Our results did not support the previously assumed ecological similarity of D. bellus and D. novemcinctus based on their geographic distributions and emphasized the possibility of a cold adapted characteristic of the life history of D. bellus.     

Comparison of trophic function between the globally invasive crayfishes <Emphasis Type="Italic">Pacifastacus leniusculus</Emphasis> and <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

Impacts of invasive species may manifest most strongly if these organisms are highly distinct functionally from the native species they often replace. Yet, should we expect functional differences between native and invasive species of generalist organisms like freshwater crayfish? Some existing evidence has pointed to native and invasive crayfish species as ecologically equivalent. We contribute to this literature by comparing the trophic niches of the globally invasive crayfishes Pacifastacus leniusculus and Procambarus clarkii, by applying carbon and nitrogen stable isotope analyses to replicated allopatric (alone) and sympatric (together) lake populations in western Washington State, USA, where P. clarkii has been recently introduced and P. leniusculus is presumed native. Our study corrected for potential inherent differences in lake food webs as a consequence of lake abiotic or biotic characteristics using random effects in linear mixed effects models. We found that although overall trophic niche size or area of these species was not significantly different, P. leniusculus was significantly higher in trophic position than P. clarkii when also accounting for the effects of body size, sex, and lakes as random effects. This pattern of increased trophic position of P. leniusculus over P. clarkii was conserved over time in one sympatric lake for which we had data over multiple years. Cumulatively, our findings point to trophic differences between the globally cosmopolitan crayfishes P. leniusculus and P. clarkii, particularly when accounting for the ways that ecosystem context can affect food web structure of communities and the trophic resources available to these consumers.     

Life history responses to variations in temperature by the marine amphipod <Emphasis Type="Italic">Eogammarus possjeticus</Emphasis> (Gammaridae) and their implications for productivity in aquaculture

The invasion of tilapia can result in substantial impacts on native communities. Thus, understanding the spatial dynamics of invasions may help prevent future introductions and mitigate impacts. This study estimated the environmentally suitable areas for occurrence of eight tilapia species (genera Coptodon, Oreochromis, Pelmatolapia, and Sarotherodon) in the Americas and their invasive potential using Ecological Niche Models (ENMs). The United States is the most invaded country, receiving all tilapia species. In South America, the southeast and south regions of Brazil are highlighted as the areas where two species are concentrated. The ENMs predicted that all tilapia species have high invasive potential in the Americas, and despite having more tilapias in North America, South and Central Americas are more susceptible to tilapia invasion. All South American basins were predicted to harbor tilapia species that have not yet arrived on the subcontinent. Our study evidences the need to implement management measures and governmental policies in the Americas to deal with problems caused by tilapia introductions. In North America, the focus is on the control of tilapia populations and in Central and South America priority should be given to contention of introduction processes.     

Parasites of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>: current research and perspectives

Harmonia axyridis (Coleoptera: Coccinellidae) has been introduced widely for biological control of agricultural pests. Harmonia axyridis has established in four continents outside of its native range in Asia and it is considered an invasive alien species (IAS). Despite a large body of work on invasion ecology, establishment mechanisms of IAS and their interactions with natural enemies remain open questions. Parasites, defined as multicellular organisms that do not directly kill the host, could potentially play an important role in regulating host populations. This study presents a review of the parasites of H. axyridis, discussing their distributions and effects on host populations across the host’s native and invasive range. These parasites are: Hesperomyces virescens Thaxt. fungi, Coccipolipus hippodamiae (McDaniel and Morrill) mites, and Parasitylenchus bifurcatus Poinar and Steenberg nematodes.     

Current status of American bullfrog, <Emphasis Type="Italic">Lithobates catesbeianus</Emphasis>, invasion in Uruguay and exploration of chytrid infection

The American bullfrog Lithobates catesbeianus is an invasive species that can strongly affect native amphibian communities through competition, predation, or introduction of diseases. This frog has invaded multiple areas in South America, for which niche models predict suitable environments across much of the continent. This paper reveals the state of the invasion of this species in Uruguay and its possible relationship with the chytrid pathogenic fungus, Batrachochytrium dendrobatidis. Surveys at invaded sites were conducted from 2007 to 2015, identified two populations undergoing recent range expansion (one of them exponential), two populations that failed to establish, and a new record in an urban area of the capital city, Montevideo. In all the analysed feral populations, chytridiomycosis was found. Our data suggest that the invasion of L. catesbeianus in Uruguay is at an early stage, with very localized populations, which might allow for the implementation of cost-effective management plans, with eradication constituting a plausible option.     

Displacement and replacement in real time: <Emphasis Type="Italic">Polistes dominula</Emphasis>’<Emphasis Type="Italic">s</Emphasis> impact on <Emphasis Type="Italic">P. fuscatus</Emphasis> in the northeastern U.S.

Two major challenges in studying the impacts of exotic invasive species on native species are identifying mechanisms of displacement and replacement and the lack of long-term population studies in these systems. A solution for the first is to study invasive and native congeners that occupy the same niche. A solution for the second is to study many populations for one year instead of one population for many years. We studied the invasion biology of the invasive European paper wasp Polistes dominula and its native congener the Northern paper wasp P. fuscatus, two species which compete for similar resources. We tracked the demography of the two wasps at sites in the northeastern United States. We found that the survival of P. dominula to the reproductive period in August was three times that of P. fuscatus, across all sites. The reproductive output of P. fuscatus declined in direct proportion to the percentage of P. dominula nests at the site. P. fuscatus nests at uninvaded sites had three times the nest cells of those at the most invaded sites. These findings suggest a positive feedback cycle in the establishment of P. dominula, in which the invasive wasp drives population declines in the native that in turn allow P. dominula to further establish. This system provides an example of a possible extinction vortex caused by competitive exclusion of a species by its invasive congener.     

Habitat fragmentation provides a competitive advantage to an invasive tree squirrel, <Emphasis Type="Italic">Sciurus carolinensis</Emphasis>

Changes in the composition of biological communities can be elicited by competitive exclusion, wherein a species is excluded from viable habitat by a superior competitor. Yet less is known about the role of environmental change in facilitating or mitigating exclusion in the context of invasive species. In these situations, decline in a native species can be due to the effects of habitat change, or due to direct effects from invasive species themselves. This is summarized by the “driver-passenger” concept of native species loss. We present a multi-year study of tree squirrels that tested the hypothesis that tree canopy fragmentation, often a result of human development, influenced the replacement of native western gray tree squirrels (Sciurus griseus) by non-native eastern gray tree squirrels (Sciurus carolinensis). We tested this hypothesis along a continuum of invasion across three study sites in central California. We found that within the developed areas of the University of California at Santa Cruz campus and city of Santa Cruz, S. carolinensis excluded S. griseus from viable habitat. The competitive advantage of S. carolinensis may be due to morphological and/or behavioral adaptation to terrestrial life in fragmented hardwood forests. We classify S. carolinensis as a “driver” of the decline of native S. griseus in areas with high tree canopy fragmentation. Future habitat fragmentation in western North America may result in similar invasion dynamics between these species. Our study warrants consideration of existing and predicted interactions between potentially invasive species that co-occur with native species where land use change is proposed.     

Deep rooting and global change facilitate spread of invasive grass

Abiotic global change factors, such as rising atmospheric CO₂, and biotic factors, such as exotic plant invasion, interact to alter the function of terrestrial ecosystems. An invasive lineage of the common reed, Phragmites australis, was introduced to North America over a century ago, but the belowground mechanisms underlying Phragmites invasion and persistence in natural systems remain poorly studied. For instance, Phragmites has a nitrogen (N) demand higher than native plant communities in many of the ecosystems it invades, but the source of the additional N is not clear. We exposed introduced Phragmites and native plant assemblages, containing Spartina patens and Schoenoplectus americanus, to factorial treatments of CO₂ (ambient or +300 ppm), N (0 or 25 g m^?2 year^?1), and hydroperiod (4 levels), and focused our analysis on changes in root productivity as a function of depth and evaluated the effects of introduced Phragmites on soil organic matter mineralization. We report that non-native invasive Phragmites exhibited a deeper rooting profile than native marsh species under all experimental treatments, and also enhanced soil organic matter decomposition. Moreover, exposure to elevated atmospheric CO₂ induced a sharp increase in deep root production in the invasive plant. We propose that niche separation accomplished through deeper rooting profiles circumvents nutrient competition where native species have relatively shallow root depth distributions; deep roots provide access to nutrient-rich porewater; and deep roots further increase nutrient availability by enhancing soil organic matter decomposition. We expect that rising CO₂ will magnify these effects in deep-rooting invasive plants that compete using a tree-like strategy against native herbaceous plants, promoting establishment and invasion through niche separation.     

Let’s eat out,there’s crayfish for dinner: American bullfrog niche shifts inside and outside native ranges and the effect of introduced crayfish

In invasion ecology, niche width has been recognized as a crucial factor for the outcome of an invasion. A common characteristic of successful invaders seems to be a broad niche width, and their impact on native communities may increase with increasing niche size. Overall, successful invader predators are predicted to shift their niche width by broadening it from native to invaded conditions. The scarcity of ecological studies examining invasive species in their native ranges prevents researchers from knowing if the prevalence of generalist invaders represents conservatism of broad native-range niches or instead niche shifts as a result of different processes acting in the invaded areas. Here we reviewed literature on trophic niche of the predatory invader American bullfrog (Lithobates catesbeianus) in both native and invaded ranges. We used statistical and graphic tools to analyse possible shifts in dietary niche width and the effect of introduced crayfish on the feeding strategy of L. catesbeianus. Globally, our results indicate that food sources used by the species differed in native and invaded sites, with a narrower trophic niche width in invaded areas. However, this pattern was disrupted by the occurrence of introduced crayfish that represents the major driver of the observed niche-width variation. Our data shed light on possible complications in interpreting and predicting patterns of biological invasions due to the interaction among species from different trophic levels that apparently disrupt general patterns that are likely bound to be idiosyncratic and complex.     

Unveiling the factors shaping the distribution of widely distributed alpine vertebrates,using multi-scale ecological niche modelling of the bat <Emphasis Type="Italic">Plecotus macrobullaris</Emphasis>

Several alpine vertebrates share a distribution pattern that extends across the South-western Palearctic but is limited to the main mountain massifs. Although they are usually regarded as cold-adapted species, the range of many alpine vertebrates also includes relatively warm areas, suggesting that factors beyond climatic conditions may be driving their distribution. In this work we first recognize the species belonging to the mentioned biogeographic group and, based on the environmental niche analysis of Plecotus macrobullaris, we identify and characterize the environmental factors constraining their ranges. Distribution overlap analysis of 504 European vertebrates was done using the Sorensen Similarity Index, and we identified four birds and one mammal that share the distribution with P. macrobullaris. We generated 135 environmental niche models including different variable combinations and regularization values for P. macrobullaris at two different scales and resolutions. After selecting the best models, we observed that topographic variables outperformed climatic predictors, and the abruptness of the landscape showed better predictive ability than elevation. The best explanatory climatic variable was mean summer temperature, which showed that P. macrobullaris is able to cope with mean temperature ranges spanning up to 16C. The models showed that the distribution of P. macrobullaris is mainly shaped by topographic factors that provide rock-abundant and open-space habitats rather than climatic determinants, and that the species is not a cold-adapted, but rather a cold-tolerant eurithermic organism. P. macrobullaris shares its distribution pattern as well as several ecological features with five other alpine vertebrates, suggesting that the conclusions obtained from this study might be extensible to them. We concluded that rock-dwelling and open-space foraging vertebrates with broad temperature tolerance are the best candidates to show wide alpine distribution in the Western Palearctic.