Elevational Ranges of Montane Birds and Deforestation in the Western Andes of Colombia

Deforestation causes habitat loss, fragmentation, degradation, and can ultimately cause extinction of the remnant species. Tropical montane birds face these threats with the added natural vulnerability of narrower elevational ranges and higher specialization than lowland species. Recent studies assess the impact of present and future global climate change on species’ ranges, but only a few of these evaluate the potentially confounding effect of lowland deforestation on species elevational distributions. In the Western Andes of Colombia, an important biodiversity hotspot, we evaluated the effects of deforestation on the elevational ranges of montane birds along altitudinal transects. Using point counts and mist-nets, we surveyed six altitudinal transects spanning 2200 to 2800m. Three transects were forested from 2200 to 2800m, and three were partially deforested with forest cover only above 2400m. We compared abundance-weighted mean elevation, minimum elevation, and elevational range width. In addition to analysing the effect of deforestation on 134 species, we tested its impact within trophic guilds and habitat preference groups. Abundance-weighted mean and minimum elevations were not significantly different between forested and partially deforested transects. Range width was marginally different: as expected, ranges were larger in forested transects. Species in different trophic guilds and habitat preference categories showed different trends. These results suggest that deforestation may affect species’ elevational ranges, even within the forest that remains. Climate change will likely exacerbate harmful impacts of deforestation on species’ elevational distributions. Future conservation strategies need to account for this by protecting connected forest tracts across a wide range of elevations.     

Spatially heterogeneous impact of climate change on small mammals of montane California

Resurveys of historical collecting localities have revealed range shifts, primarily leading edge expansions, which have been attributed to global warming. However, there have been few spatially replicated community-scale resurveys testing whether species'' responses are spatially consistent. Here we repeated early twentieth century surveys of small mammals along elevational gradients in northern, central and southern regions of montane California. Of the 34 species we analysed, 25 shifted their ranges upslope or downslope in at least one region. However, two-thirds of ranges in the three regions remained stable at one or both elevational limits and none of the 22 species found in all three regions shifted both their upper and lower limits in the same direction in all regions. When shifts occurred, high-elevation species typically contracted their lower limits upslope, whereas low-elevation species had heterogeneous responses. For high-elevation species, site-specific change in temperature better predicted the direction of shifts than change in precipitation, whereas the direction of shifts by low-elevation species was unpredictable by temperature or precipitation. While our results support previous findings of primarily upslope shifts in montane species, they also highlight the degree to which the responses of individual species vary across geographically replicated landscapes.     

Range dynamics of small mammals along an elevational gradient over an 80‐year interval

One expected response to observed global warming is an upslope shift of species elevational ranges. Here, we document changes in the elevational distributions of the small mammals within the Ruby Mountains in northeastern Nevada over an 80‐year interval. We quantified range shifts by comparing distributional records from recent comprehensive field surveys (2006–2008) to earlier surveys (1927–1929) conducted at identical and nearby locations. Collector field notes from the historical surveys provided detailed trapping records and locality information, and museum specimens enabled confirmation of species' identifications. To ensure that observed shifts in range did not result from sampling bias, we employed a binomial likelihood model (introduced here) using likelihood ratios to calculate confidence intervals around observed range limits. Climate data indicate increases in both precipitation and summer maximum temperature between sampling periods. Increases in winter minimum temperatures were only evident at mid to high elevations. Consistent with predictions of change associated with climate warming, we document upslope range shifts for only two mesic‐adapted species. In contrast, no xeric‐adapted species expanded their ranges upslope. Rather, they showed either static distributions over time or downslope contraction or expansion. We attribute these unexpected findings to widespread land‐use driven habitat change at lower elevations. Failure to account for land‐use induced changes in both baseline assessments and in predicting shifts in species distributions may provide misleading objectives for conservation policies and management practices.     

Climate change induced elevational range shifts of Himalayan tree species

Global warming may force montane species to shift upward to keep pace with their shifting climate niche. How species differences in such distribution shifts depend on their elevational positions, elevation-dependent warming rates, and other environmental constraints, or plant functional traits is poorly understood. Here, we analyzed for 137 Himalayan tree species how distribution shifts vary with elevational niche positions, environmental constraints, and their functional traits. We developed ecological niche models using MaxEnt by combining species survey and botanical collections data with 19 environmental predictors. Species distributions were projected to 1985 and 2050 conditions, and elevational range parameters and distribution areas were derived. Under the worst-case RCP 8.5 scenario, species are predicted to shift, on average, 3 m/year in optimum elevation, and have 33% increase in distribution area. Highland species showed faster predicted elevational shifts than lowland species. Lowland and highland species are predicted to expand in distribution area in contrast to mid-elevation species. Tree species for which species distribution models are driven by responses to temperature, aridity, or soil clay content showed the strongest predicted upslope shifts. Tree species with conservative trait values that enable them to survive resource poor conditions (i.e., narrow conduits) showed larger predicted upslope shifts than species with wide conduits. The predicted average upslope shift in maximum elevation (8 m/year) is >2 times faster than the current observations indicating that many species will not be able to track climate change and potentially go extinct, unless they are supported by active conservation measures, such as assisted migration.     

Exploring the role of physiology and biotic interactions in determining elevational ranges of tropical animals

Tropical mountains contain some of the world’s richest animal communities as a result of high turnover of species along elevational gradients. We describe an approach to study the roles of biotic and abiotic factors in establishing elevational ranges, and to improve our ability to predict the effects of climate change on these communities. As a framework we use Hutchinson’s concept of the fundamental niche (determined by the match between the physical environment and the organism’s physiological and biophysical characteristics) and realized niche (the subset of the fundamental niche determined by biotic interactions). Using tropical birds as an example, we propose a method for estimating fundamental niches and discuss five biotic interactions that we expect to influence distributions of tropical montane animals: predation, competition, parasites and pathogens, mutualisms, and habitat associations. The effects of biotic factors on elevational ranges have been studied to some extent, but there is little information on physiological responses of tropical montane animals. It will be necessary to understand all of these ecological constraints in concert to predict current and future elevational ranges and potential threats to montane species. Given the importance of tropical mountains as global biodiversity hotspots, we argue that this area of research requires urgent attention.     

A weak upward elevational shift in the distributions of breeding birds in the Italian Alps

Aim To test whether bird assemblages are shifting upwards in their elevational distribution in synchrony with current climate warming and/or habitat changes. Location A gradient of elevation in the Italian Alps (Alta Valsessera, Piedmont). Methods We used data from two recent atlas surveys performed on a 1 × 1 km grid at an 11‐year interval (1992–94 and 2003–05). We modelled the elevational gradient of avifaunal composition, using a sample‐based approach, in an effort to detect evidence for an upward elevational shift of bird zonation. Changes in species richness were controlled for. The results from this analysis were compared with those obtained using a species‐based approach. Changes in climate and landscape between the two surveys were assessed using local meteorological data and Corine Land Cover maps, respectively. Results We detected small avifaunal changes between the two surveys: (1) mean elevations increased for the majority of species, but the average change was not significantly different from zero; (2) the species richness increased, but this was mainly due to an increase in sampling effort; and (3) a change in species composition was detected, which was at the limit of significance and corresponded on average to a 29‐m upward elevational shift in the distribution of the avifauna. The shift was the same for open land and forest bird communities. During the same period, the mean temperature increased by c. 1 °C in the area, and a slight trend towards vegetation closure by woody plants was detected. Main conclusions The use of fine‐scale breeding bird atlases in mountainous regions, together with ordination methods, provides a sensitive tool to test and measure elevational shifts in species ranges, but the results have to be interpreted carefully. In our case, the observed elevational shift in the distributions of the avifauna cannot unambiguously be attributed to climate warming. This shift is smaller than expected from the regional increase in temperature, which raises the question of how closely bird distributions match climate change.     

Insects Overshoot the Expected Upslope Shift Caused by Climate Warming

Along elevational gradients, climate warming may lead to an upslope shift of the lower and upper range margin of organisms. A recent meta-analysis concluded that these shifts are species specific and considerably differ among taxonomic lineages. We used the opportunity to compare upper range margins of five lineages (plants, beetles, flies, hymenoptera, and birds) between 1902–1904 and 2006–2007 within one region (Bavarian Forest, Central Europe). Based on the increase in the regional mean annual temperature during this period and the regional lapse rate, the upslope shift is expected to be between 51 and 201 m. Averaged across species within lineages, the range margin of all animal lineages shifted upslope, but that of plants did not. For animals, the observed shifts were probably due to shifts in temperature and not to changes in habitat conditions. The range margin of plants is therefore apparently not constrained by temperature, a result contrasting recent findings. The mean shift of birds (165 m) was within the predicted range and consistent with a recent global meta-analysis. However, the upslope shift of the three insect lineages (>260 m) exceeded the expected shift even after considering several sources of uncertainty, which indicated a non-linear response to temperature. Our analysis demonstrated broad differences among lineages in their response to climate change even within one region. Furthermore, on the considered scale, the response of ectothermic animals was not consistent with expectations based on shifts in the mean annual temperature. Irrespective of the reasons for the overshooting of the response of the insects, these shifts lead to reorganizations in the composition of assemblages with consequences for ecosystem processes.     

Bumblebee community homogenization after uphill shifts in montane areas of northern Spain

Widespread alterations in species distribution and abundance as a result of global environmental change include upwards and polewards shifts driven by local extinctions in the south or at lower elevations and colonizations of newly available habitat elements in the north or at higher elevations. Although cumulative changes on patterns of community composition are also expected, studies following a community-level approach are still scarce. Here, we estimate changes in abundance and distribution of bumblebee (Bombus spp.) species over two decades along an elevational gradient to test whether these changes entailed concomitant alterations on patterns of community composition. Bumblebee species showed an overall trend to shift uphill their upper- or lower-elevational boundaries, resulting in narrower elevational ranges from one period to another, coincident with a regional warming of ca. 0.9 °C. Changes in elevational ranges were, however, mainly related to retractions of the lower limit of species distribution, rather than to variations in their upper elevational limit. Species turnover was associated with colonization and extinction events and also with variability in the relative abundance of short-, medium- and long-tongued species along the elevational gradient. Extinctions were especially relevant at medium elevations, while only communities at higher elevations had a positive net outcome between colonization and extinction events. The combination of these effects resulted in the homogenization of bumblebee assemblages, especially between medium and upper elevations. The changes reported in our study strongly match with predictions of global change driving elevational shifts in species distribution and provide the first evidence of elevational changes in bumblebees at both species and community level.     

Systematics, speciation and biogeography of the dwarf chameleons (Brookesia; Reptilia, Squamata, Chamaeleontidae) of northern Madagascar

Brookesia dwarf chameleons, endemic to Madagascar, were surveyed at the following localities in northern Madagascar (north of 16°S): Montagne d'Ambre, Ankarana, Manongarivo, Tsaratanana, Marojejy and Masoala. A total of 15 species occur in this region. Six new species are described and five new synonyms are identified. The genus Brookesia , the most speciose chamaeleontid genus in Madagascar, contains 23 species.
Almost all the northern Brookesia species are restricted to rainforest and occupy a relatively narrow elevational range. Although the northern rainforests represent just one-third of the total rainforest and about 5% of the total island area, 65% of the Brookesia species occur in this region, and 52% are endemic to the northern rainforest. Five new biogeographic regions of the northern rainforest are identified based on centres of Brookesia endemicity: Montagne d'Ambre, Northwest, Tsaratanana, Northeast and East. Speciation is thought to have been facilitated in the north through geographic isolation, with the Tsaratanana mountain range and the dry forests south of Montagne d'Ambre forming barriers to dispersal, and the Tsaratanana mountains acting as a centre of isolation. The fragmented distribution of several Brookesia species of low altitude rainforest suggests a period in Madagascar's history when the climate was wetter and low altitude rainforest much more widespread.     

An elevational shift in butterfly species richness and composition accompanying recent climate change

The geographic ranges of many species have shifted polewards and uphill in elevation associated with climate warming, leading to increases in species richness at high latitudes and elevations. However, few studies have addressed community‐level responses to climate change across the entire elevational gradients of mountain ranges, or at warm lower latitudes where ecological diversity is expected to decline. Here, we show uphill shifts in butterfly species richness and composition in the Sierra de Guadarrama (central Spain) between 1967–1973 and 2004–2005. Butterfly communities with comparable species compositions shifted uphill by 293 m (± SE 26), consistent with an upward shift of approximately 225 m in mean annual isotherms. Species richness had a humped relationship with elevation, but declined between surveys, particularly at low elevations. Changes to species richness and composition primarily reflect the loss from lower elevations of species whose regional distributions are restricted to the mountains. The few colonizations by specialist low‐elevation species failed to compensate for the loss of high‐elevation species, because there are few low‐elevation species in the region and the habitat requirements of some of these prevent them from colonizing the mountain range. As a result, we estimated a net decline in species richness in approximately 90% of the region, and increasing community domination by widespread species. The results suggest that climate warming, combined with habitat loss and other drivers of biological change, could lead to significant losses in ecological diversity in mountains and other regions where species encounter their lower latitudinal‐range margins.     

Modelling the responses of Andean and Amazonian plant species to climate change: the effects of georeferencing errors and the importance of data filtering

Aim Species distribution models are a potentially powerful tool for predicting the effects of global change on species distributions and the resulting extinction risks. Distribution models rely on relationships between species occurrences and climate and may thus be highly sensitive to georeferencing errors in collection records. Most errors will not be caught using standard data filters. Here we assess the impacts of georeferencing errors and the importance of improved data filtering for estimates of the elevational distributions, habitat areas and predicted relative extinction risks due to climate change of nearly 1000 Neotropical plant species. Location The Amazon basin and tropical Andes, South America. Methods We model the elevational distributions, or ‘envelopes’, of 932 Amazonian and Andean plant species from 35 families after performing standard data filtering, and again using only data that have passed through an additional layer of data filtering. We test for agreement in the elevations recorded with the collection and the elevation inferred from a digital elevation model (DEM) at the collection coordinates. From each dataset we estimate species range areas and extinction risks due to the changes in habitat area caused by a 4.5 °C increase in temperature. Results Amazonian and Andean plant species have a median elevational range of 717 m. Using only standard data filters inflates range limits by a median of 433 m (55%). This is equivalent to overestimating the temperature tolerances of species by over 3 °C – only slightly less than the entire regional temperature change predicted over the next 50–100 years. Georeferencing errors tend to cause overestimates in the amount of climatically suitable habitat available to species and underestimates in species extinction risks due to global warming. Georeferencing error artefacts are sometimes so great that accurately predicting whether species habitat areas will decrease or increase under global warming is impossible. The drawback of additional data filtering is large decreases in the number of species modelled, with Andean species being disproportionately eliminated. Main conclusions Even with rigorous data filters, distribution models will mischaracterize the climatic conditions under which species occur due to errors in the collection data. These errors affect predictions of the effects of climate change on species ranges and biodiversity, and are particularly problematic in mountainous areas. Additional data filtering reduces georeferencing errors but eliminates many species due to a lack of sufficient ‘clean’ data, thereby limiting our ability to predict the effects of climate change in many ecologically important and sensitive regions such as the Andes Biodiversity Hotspot.     

A test of concordance in community structure between leafhoppers and grasslands in the central Tien Shan Mountains

Leafhoppers and related Auchenorrhynchous Hemiptera (AH) are among the most diverse grassland herbivores, and many have been linked inexorably to grassland vegetation through diet and shelter for millions of years. Are AH–plant communities in natural grasslands tightly integrated, how does the interaction differ across major ecological gradients, and do habitat or environmental factors explain the most variance in AH community structure? These questions have implications for the conservation of biodiversity and in evaluating effects of a warming climate. Using grasslands of the central Tien Shan Mountains as a natural laboratory, we examine whether AH species assemblages are concordant with vegetation in terms of community structure using closely associated species-level samples. Data were recorded from a nearly 3000-m elevation gradient crossing four arid and three montane grassland vegetation classes. We found elements of AH–plant community classification and structure to be closely correlated except for at the arid–montane habitat transition where a small group of widespread AH species were significant indicators for vegetation classes in both major grassland types. AH species richness and abundance are positively correlated with plant species density and percent cover and, correspondingly, peak at mid-elevations in association with montane grasslands. While overall elevation (and covariate mean annual temperature) explains the most variance in AH species assemblages, the sum total of habitat factors explain more variance than environmental factors when arid and montane grasslands were examined separately, but environmental factors are co-equal with habitat factors when the grassland types are combined. Unexplained variance in the AH community assemblages, attributable to individualistic species responses to environmental and habitat factors, slightly exceeds the total accounted for by the model.     

Upslope Range Shifts of Andean Dung Beetles in Response to Deforestation: Compounding and Confounding Effects of Microclimatic Change

Warmer, and sometimes drier, conditions associated with global climate change are driving many species to shift poleward and/or upslope. I hypothesized that microclimatic changes related to deforestation cause similar shifts for forest species persisting within degraded landscapes. This appears to be the first study to examine this novel hypothesis. I examined elevational distributions of dung beetle communities along parallel intact and disturbed elevational gradients from 290 to 3450 m asl in the Andes of southeastern Peru. Deforested sites were consistently warmer and drier than forested sites. To maintain the same ambient temperature as in forest, species in a deforested landscape would need to shift on average 489±59 m upslope. Dung beetle species showed a mean upslope range shift of 132±64 m (maximum=743 m) in the deforested landscape. Eight species occurred farther upslope in the degraded landscape, while none shifted downslope. In addition to upper range limit expansions, six species shifting upslope also showed range contractions or population declines at their lower range boundary. High elevation and disturbance‐tolerant species did not show range shifts. These findings suggest that land‐use change may both confound and compound the influence of global climate change on biodiversity. Synergies between habitat degradation and climate change could more than double previous range shift projections for this century, leading to unexpectedly rapid changes in biodiversity, especially for sensitive organisms such as tropical insects. On the other hand, range shifts caused by habitat degradation may be mistakenly attributed to global climate change. Abstract in Spanish is available in the online version of this article.     

Current Analogues of Future Climate Indicate the Likely Response of a Sensitive Montane Tropical Avifauna to a Warming World

Among birds, tropical montane species are likely to be among the most vulnerable to climate change, yet little is known about how climate drives their distributions, nor how to predict their likely responses to temperature increases. Correlative models of species’ environmental niches have been widely used to predict changes in distribution, but direct tests of the relationship between key variables, such as temperature, and species’ actual distributions are few. In the absence of historical data with which to compare observations and detect shifts, space-for-time substitutions, where warmer locations are used as analogues of future conditions, offer an opportunity to test for species’ responses to climate. We collected density data for rainforest birds across elevational gradients in northern and southern subregions within the Australian Wet Tropics (AWT). Using environmental optima calculated from elevational density profiles, we detected a significant elevational difference between the two regions in ten of 26 species. More species showed a positive (19 spp.) than negative (7 spp.) displacement, with a median difference of ∼80.6 m across the species analysed that is concordant with that expected due to latitudinal temperature differences (∼75.5 m). Models of temperature gradients derived from broad-scale climate surfaces showed comparable performance to those based on in-situ measurements, suggesting the former is sufficient for modeling impacts. These findings not only confirm temperature as an important factor driving elevational distributions of these species, but also suggest species will shift upslope to track their preferred environmental conditions. Our approach uses optima calculated from elevational density profiles, offering a data-efficient alternative to distribution limits for gauging climate constraints, and is sensitive enough to detect distribution shifts in this avifauna in response to temperature changes of as little as 0.4 degrees. We foresee important applications in the urgent task of detecting and monitoring impacts of climate change on montane tropical biodiversity.     

Elevational zonation of afrotropical forest bird communities along a homogeneous forest gradient

Aim This study analyses the distribution and abundance of birds from a forested tropical gradient in order to determine whether elevationally distinct communities are detectable in this habitat. Location An avifaunal census was carried out on a single transect within the tropical forest of the Udzungwa Mountains in the Eastern Arc, Tanzania, covering a range in elevation from 300 to 1850 m. Methods Two complementary data sets on forest birds were analysed, encompassing (1) data derived from standardized 20‐ha spot‐mapping censuses performed at nine elevations over 175‐m intervals from 400 to 1800 m a.s.l., and (2) all observations of birds binned into 32 data points at 50‐m intervals, from 300 to 1850 m a.s.l. The degree of zonation in the avian community along the elevational gradient was examined using the chronological clustering method, an agglomerative hierarchical clustering method that can be carried out with a range of similarity indices. Results The chronological clustering analysis of the data set based on standardized spot‐mapping revealed a clearly defined boundary at c. 1200 m a.s.l., separating lowland from montane communities. Most bird species could be categorized as belonging to one of these two communities. The data set based on all observations revealed a number of potential secondary boundaries, although these boundaries delimited the entire elevational ranges of individual species in only relatively few cases. Main conclusions In contrast to previously published studies, we find evidence of an elevational zonation of distinct communities within a seemingly homogeneous habitat. Although similar boundaries have been assumed to arise as a result of vegetational ecotones, or because of interspecific competition, these mechanisms are poorly corroborated. We suggest that the causes of patterns of zonation are not well understood, and that the interplay among species distributions, species richness, and environmental factors needs more consideration. The chronological clustering method is proposed as an appropriate tool for studying these specific patterns.     

Distributional change and conservation on the Andean flank: a palaeoecological perspective

Aim To review the insights that palaeoecology can offer on the threat posed to Andean communities by global climate change. Location The geographical focus is the eastern flank of the tropical Andes, with particular reference to Peru. Method The article presents a synthetic review of the problem. Results Species‐rich communities of the eastern Andean flank are threatened both by development and climate change. If, as predicted, the cloudbase and frost line lifts 600 m elevation this century, there will be a substantial loss of cloud forest habitat. Palaeoecology provides insights on the location and nature of past ecotones, the continuity of niche availability, and estimates for rates and modes of migration. With further warming and drying of lower montane regions, upslope migration of species will be individualistic: a new equilibrium with the altered climate is unlikely to be attained. The zone of agriculture will move upslope faster than the migrating trees and so landscape conversion will negate the ability of some species to respond to the new conditions. Conservation of the lower reaches of modern cloud forest is advocated as a means to limit this foreseeable extinction event. Main Conclusions Climate change will induce changes in plant and animal communities. Human disturbance will invade climatically marginal agricultural lands at the pace of climate change. Rapid migratory response such as that witnessed at the onset of the Holocene may not be evident as climates warm, because Holocene climatic stability has reduced outlying upslope populations that may have served as expansion nuclei. Conservation must aim to maintain plant and animal niches, rather than particular communities of species.     

Climate change,extinction, and Sky Island biogeography in a montane lizard

Around the world, many species are confined to “Sky Islands,” with different populations in isolated patches of montane habitat. How does this pattern arise? One scenario is that montane species were widespread in lowlands when climates were cooler, and were isolated by local extinction caused by warming conditions. This scenario implies that many montane species may be highly susceptible to anthropogenic warming. Here, we test this scenario in a montane lizard (Sceloporus jarrovii) from the Madrean Sky Islands of southeastern Arizona. We combined data from field surveys, climate, population genomics, and physiology. Overall, our results support the hypothesis that this species' current distribution is explained by local extinction caused by past climate change. However, our results for this species differ from simple expectations in several ways: (a) their absence at lower elevations is related to warm winter temperatures, not hot summer temperatures; (b) they appear to exclude a low‐elevation congener from higher elevations, not the converse; (c) they are apparently absent from many climatically suitable but low mountain ranges, seemingly “pushed off the top” by climates even warmer than those today; (d) despite the potential for dispersal among ranges during recent glacial periods (~18,000 years ago), populations in different ranges diverged ~4.5–0.5 million years ago and remained largely distinct; and (e) body temperatures are inversely related to climatic temperatures among sites. These results may have implications for many other Sky Island systems. More broadly, we suggest that Sky Island species may be relevant for predicting responses to future warming.     

Distinct responses and range shifts of lizard populations across an elevational gradient under climate change

Ongoing climate change has profoundly affected global biodiversity, but its impacts on populations across elevations remain understudied. Using mechanistic niche models incorporating species traits, we predicted ecophysiological responses (activity times, oxygen consumption and evaporative water loss) for lizard populations at high-elevation (<3600 m asl) and extra-high-elevation (≥3600 m asl) under recent (1970–2000) and future (2081–2100) climates. Compared with their high-elevation counterparts, lizards from extra-high-elevation are predicted to experience a greater increase in activity time and oxygen consumption. By integrating these ecophysiological responses into hybrid species distribution models (HSDMs), we were able to make the following predictions under two warming scenarios (SSP1-2.6, SSP5-8.5). By 2081–2100, we predict that lizards at both high- and extra-high-elevation will shift upslope; lizards at extra-high-elevation will gain more and lose less habitat than will their high-elevation congeners. We therefore advocate the conservation of high-elevation species in the context of climate change, especially for those populations living close to their lower elevational range limits. In addition, by comparing the results from HSDMs and traditional species distribution models, we highlight the importance of considering intraspecific variation and local adaptation in physiological traits along elevational gradients when forecasting species' future distributions under climate change.     

Winners and losers: How the elevational range of breeding birds on Alps has varied over the past four decades due to climate and habitat changes

Climate warming and habitat transformation are widely recognized as worrying threatening factors. Understanding the individual contribution of these two factors to the change of species distribution could be very important in order to effectively counteract the species range contraction, especially in mountains, where alpine species are strongly limited in finding new areas to be colonized at higher elevations. We proposed a method to disentangle the effects of the two drivers of range change for breeding birds in Italian Alps, in the case of co‐occurring climate warming and shrub and forest encroachment. For each species, from 1982 to 2017, we related the estimated yearly elevational distribution of birds to the correspondent overall average of the daily minimum temperatures during the breeding season and the estimated amount of shrubs and forest cover. Using a hierarchical partitioning approach, we assessed the net contribution (i.e., without the shared effect) of each driver. Both temperature and shrub and forest cover showed a positive trend along the time series and resulted the most likely causes of the significant elevational displacement for 21 of the 29 investigated birds. While shrub and forest cover was found to be an important driver of the expansion of forest bird range toward higher elevations, the effect of temperature on favouring the colonization of previously climatically unsuitable forests at higher elevations was not negligible. Shrub and forest expansion resulted the main driver of the range contraction for edge and open habitat species, which suffered a distribution shrinkage at their lower elevational boundary. In light of climate warming, these results highlighted how the net range loss for edge and open habitat species, caused by shrub and forest encroachment consequent to land abandonment, should be counteracted by implementing proper conservation management strategies and promoting sustainable economic activities in rangeland areas.