Habitat heterogeneity,temperature, and primary productivity drive elevational gradients in avian species diversity

AimAnticipating and mitigating the impacts of climate change on species diversity in montane ecosystems requires a mechanistic understanding of drivers of current patterns of diversity. We documented the shape of elevational gradients in avian species richness in North America and tested a suite of a priori predictions for each of five mechanistic hypotheses to explain those patterns.LocationUnited StatesMethodsWe used predicted occupancy maps generated from species distribution models for each of 646 breeding birds to document elevational patterns in avian species richness across the six largest U.S. mountain ranges. We used spatially explicit biotic and abiotic data to test five mechanistic hypotheses proposed to explain geographic variation in species richness.ResultsElevational gradients in avian species richness followed a consistent pattern of low elevation plateau‐mid‐elevation peak (as per McCain, 2009). We found support for three of the five hypotheses to explain the underlying cause of this pattern: the habitat heterogeneity, temperature, and primary productivity hypotheses.Main ConclusionsSpecies richness typically decreases with elevation, but the primary cause and precise shape of the relationship remain topics of debate. We used a novel approach to study the richness‐elevation relationship and our results are unique in that they show a consistent relationship between species richness and elevation among 6 mountain ranges, and universal support for three hypotheses proposed to explain the underlying cause of the observed relationship. Taken together, these results suggest that elevational variation in food availability may be the ecological process that best explains elevational gradients in avian species richness in North America. Although much attention has focused on the role of abiotic factors, particularly temperature, in limiting species’ ranges, our results offer compelling evidence that other processes also influence (and may better explain) elevational gradients in species richness.     

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.     

Current and Potential Future Elevational Distributions of Birds Associated with Pinyon–Juniper Woodlands in the Central Great Basin, U.S.A.

We examined the relationship of breeding birds to elevation across and within four adjacent mountain ranges in the central Great Basin, a cold desert in western North America. Data came from 7 years of point counts at elevations from 1,915 to 3,145 m. We focused on eight passerine species that in this region are associated frequently with Pinus monophylla–Juniperus spp. (pinyon–juniper) woodland. Mean elevation of species' presence differed significantly among mountain ranges for all species except Spizella passerina (Chipping Sparrow); all species except Spizella breweri (Brewer's Sparrow) occurred at the highest mean elevation in the Toquima Range. Observed patterns were consistent with the elevational distribution of pinyon–juniper woodlands that provide nesting and foraging habitat for these species. Across the Great Basin, driven in part by climate change, pinyon–juniper woodland is increasing in density and expanding its distribution at lower elevations. However, breeding habitat for species dependent on mature trees may not be available in expansion woodlands for several decades, and increased tree densities may have negative effects on bird species that are dependent on shrubs within open pinyon–juniper woodlands. Responses of individual species to elevation differed from the response of assemblage-level patterns. Responses to biotic and abiotic variables within guilds of birds are sufficiently diverse, and responses of individual species sufficiently heterogeneous, that one management strategy is unlikely to meet the needs of all species in the group.     

Human impacts on environment–diversity relationships: evidence for biotic homogenization from butterfly species richness patterns

Aim Broad‐scale spatial variation in species richness relates to climate and physical heterogeneity but human activities may be changing these patterns. We test whether climate and heterogeneity predict butterfly species richness regionally and across Canada and whether these relationships change in areas of human activity. Location Canada. Methods We modelled the ranges of 102 butterfly species using genetic algorithms for rule‐set production (GARP). We then measured butterfly species richness and potentially important aspects of human activity and the natural environment. These were included in a series of statistical models to determine which factors are likely to affect butterfly species richness in Canada. We considered patterns across Canada, within predominantly natural areas, human‐dominated areas and particular ecozones. We examined independent observations of butterfly species currently listed under Canada's endangered species legislation to test whether these were consistent with findings from statistical models. Results Growing season temperature is the main determinant of butterfly species richness across Canada, with substantial contributions from habitat heterogeneity (measured using elevation). Only in the driest areas does precipitation emerge as a leading predictor of richness. The slope of relationships between all of these variables and butterfly species richness becomes shallower in human‐dominated areas, but butterfly richness is still highest there. Insecticide applications, habitat loss and road networks reduce butterfly richness in human‐dominated areas, but these effects are relatively small. All of Canada's at‐risk butterfly species are located in these human‐dominated areas. Main conclusions Temperature affects butterfly species richness to a greater extent than habitat heterogeneity at fine spatial scales and is generally far more important than precipitation, supporting both the species richness–energy and habitat heterogeneity hypotheses. Human activities, especially in southern Canada, appear to cause surprisingly consistent trends in biotic homogenization across this region, perhaps through range expansion of common species and loss of range‐restricted species.     

Ecological niche modelling of montane mammals in the Great Basin, North America: examining past and present connectivity of species across basins and ranges

Aim The goal of this study was to determine the extent of suitable habitats across the basins and ranges of the Great Basin for 13 montane mammals in the present and during the Last Glacial Maximum (LGM). For all these mammal species, we test whether: (1) more suitable habitat was available in basin areas during the LGM; (2) suitable habitat shifted upwards in elevation between the LGM and the present; (3) more ranges have suitable habitat than are currently occupied; and (4) these species are currently restricted to suitable habitats at higher‐elevation range areas. We also examine whether and how much distributional response varies among these montane mammal species. Location The Great Basin of western North America. Methods We re‐examine the past and present distributions of 13 Great Basin montane mammals using ecological niche modelling techniques that utilize now widely available species occurrence data and new, fine‐scale past climatological GIS layers in the present and at the LGM. These methods provide an objective, repeatable means for visual comparison of past and present modelled distributions for species examined in previous biogeographical studies. Results Our results indicate greater areal and lower elevational suitable habitat in the LGM than at present for nearly all montane mammals, and that there is more suitable habitat at present than is currently occupied. Our results also show that lowland areas provide suitable dispersal routes between ranges for most of the montane mammals both at the LGM and at present. However, three of the 13 species have little to no predicted suitable habitat in the LGM near currently occupied ranges, in contrast to the pattern for the other 10. For these species, the model results support more recent long‐distance colonization. Main conclusions Our finding of suitable lowland dispersal routes in the present for most species supports and greatly extends similar findings from single‐species studies. Our results also provide a visually striking confirmation that changes in species distribution and colonization histories of Great Basin montane mammals vary in a fashion related to the tolerances and requirements of each of these species; this has previously been hypothesized but not rigorously tested for multiple montane mammals in the region.     

Elevational diversity gradients, biogeography and the structure of montane mammal communities in the intermountain region of North America

1  Distribution data were assembled for non-volant small mammals along elevational gradients on mountain ranges in the western U.S.A. Elevational distributions in the species-rich Uinta Mountains were compared to those on smaller mountain ranges with varying degrees of historical isolation from the Uintas.
2  For mountain ranges supporting the richest faunas, species richness is highest over a broad low- to mid-elevation zone and declines at both lower and higher elevations. Patterns on other mountain ranges are similar but reflect lower overall species richness.
3  A basic relationship between elevational and geographical distribution is apparent in the occurrence patterns of mammals on regional mountains. Faunas on mountains that have had low levels of historical isolation appear to be influenced by immigration rather than extinction. Species restricted to high elevations in the Uintas are poorly represented on historically isolated mountains and form a portion of local faunas shaped by extinction. Species occurring at lower elevations in the Uintas have better representation on isolated mountains and apparently maintain populations through immigration.
4  Several widespread species show substantial variation in maximum elevation records on different mountain ranges. This involves (1) an upward shift in habitat zones on small, isolated mountain ranges, allowing greater access by low-elevation species, and (2) expansion of certain low- and mid-elevation species into habitats normally occupied by absent high-elevation taxa.
5  Results indicate that montane mammal faunas of the intermountain region have been shaped by broad-scale historical processes, unique regional geography and local ecological dynamics. Parallel examples among mammals of the Philippine Islands suggest that such patterns may characterize many insular faunas.     

Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.     

Leap frog in slow motion: Divergent responses of tree species and life stages to climatic warming in Great Basin subalpine forests

In response to climate warming, subalpine treelines are expected to move up in elevation since treelines are generally controlled by growing season temperature. Where treeline is advancing, dispersal differences and early life stage environmental tolerances are likely to affect how species expand their ranges. Species with an establishment advantage will colonize newly available habitat first, potentially excluding species that have slower establishment rates. Using a network of plots across five mountain ranges, we described patterns of upslope elevational range shift for the two dominant Great Basin subalpine species, limber pine and Great Basin bristlecone pine. We found that the Great Basin treeline for these species is expanding upslope with a mean vertical elevation shift of 19.1 m since 1950, which is lower than what we might expect based on temperature increases alone. The largest advances were on limber pine‐dominated granitic soils, on west aspects, and at lower latitudes. Bristlecone pine juveniles establishing above treeline share some environmental associations with bristlecone adults. Limber pine above‐treeline juveniles, in contrast, are prevalent across environmental conditions and share few environmental associations with limber pine adults. Strikingly, limber pine is establishing above treeline throughout the region without regard to site characteristic such as soil type, slope, aspect, or soil texture. Although limber pine is often rare at treeline where it coexists with bristlecone pine, limber pine juveniles dominate above treeline even on calcareous soils that are core bristlecone pine habitat. Limber pine is successfully “leap‐frogging” over bristlecone pine, probably because of its strong dispersal advantage and broader tolerances for establishment. This early‐stage dominance indicates the potential for the species composition of treeline to change in response to climate change. More broadly, it shows how species differences in dispersal and establishment may result in future communities with very different specific composition.     

Dramatic response to climate change in the Southwest: Robert Whittaker's 1963 Arizona Mountain plant transect revisited

Models analyzing how Southwestern plant communities will respond to climate change predict that increases in temperature will lead to upward elevational shifts of montane species. We tested this hypothesis by reexamining Robert Whittaker's 1963 plant transect in the Santa Catalina Mountains of southern Arizona, finding that this process is already well underway. Our survey, five decades after Whittaker's, reveals large changes in the elevational ranges of common montane plants, while mean annual rainfall has decreased over the past 20 years, and mean annual temperatures increased 0.25°C/decade from 1949 to 2011 in the Tucson Basin. Although elevational changes in species are individualistic, significant overall upward movement of the lower elevation boundaries, and elevational range contractions, have occurred. This is the first documentation of significant upward shifts of lower elevation range boundaries in Southwestern montane plant species over decadal time, confirming that previous hypotheses are correct in their prediction that mountain communities in the Southwest will be strongly impacted by warming, and that the Southwest is already experiencing a rapid vegetation change.     

Species richness and phylogenetic diversity of seed plants across vegetation zones of Mount Kenya,East Africa

Mount Kenya is of ecological importance in tropical east Africa due to the dramatic gradient in vegetation types that can be observed from low to high elevation zones. However, species richness and phylogenetic diversity of this mountain have not been well studied. Here, we surveyed distribution patterns for a total of 1,335 seed plants of this mountain and calculated species richness and phylogenetic diversity across seven vegetation zones. We also measured phylogenetic structure using the net relatedness index (NRI) and the nearest species index (NTI). Our results show that lower montane wet forest has the highest level of species richness, density, and phylogenetic diversity of woody plants, while lower montane dry forest has the highest level of species richness, density, and phylogenetic diversity in herbaceous plants. In total plants, NRI and NTI of four forest zones were smaller than three alpine zones. In woody plants, lower montane wet forest and upper montane forest have overdispersed phylogenetic structures. In herbaceous plants, NRI of Afro‐alpine zone and nival zone are smaller than those of bamboo zone, upper montane forest, and heath zone. We suggest that compared to open dry forest, humid forest has fewer herbaceous plants because of the closed canopy of woody plants. Woody plants may have climate‐dominated niches, whereas herbaceous plants may have edaphic and microhabitat‐dominated niches. We also proposed lower and upper montane forests with high species richness or overdispersed phylogenetic structures as the priority areas in conservation of Mount Kenya and other high mountains in the Eastern Afro‐montane biodiversity hotspot regions.     

Thermophysiology,microclimates, and species distributions of lizards in the mountains of the Brazilian Atlantic Forest

Thermophysiological traits, particularly thermal tolerances and sensitivity, are key to understanding how organisms are affected by environmental conditions. In the face of ongoing climate change, determining how physiological traits structure species’ ranges is especially important in tropical montane systems. In this study, we ask whether thermal sensitivity in physiological performance restricts montane lizards to high elevations and excludes them from the warmer environments reported at low elevations. For three montane lizard species in the Brazilian Atlantic Forest, we collect thermophysiological data from lizards in the highest elevation site of each species’ distribution, and ask how well the individuals exhibiting those traits would perform across the Atlantic Forest. We use microclimatic and organism‐specific models to directly relate environmental conditions to an organism's body temperature and physiological traits, and estimate measures of thermophysiological performance. Our findings demonstrate that thermophysiological constraints do not restrict montane lizards to high elevations in this system, and thus likely do not determine the warm boundaries of these montane species’ distributions. Results also suggest that competition may be important in limiting the warm boundaries of the species’ ranges for two of the focal species. These experimental results suggest that caution should be used when claiming that physiology drives patterns of diversity and endemism within montane environments. They also highlight the importance of interdisciplinary experimental studies that bridge the fields of evolution and ecology to improve predictions of biological responses to future environmental shifts.     

Climate induced increases in species richness of marine fishes

Climate change has been predicted to lead to changes in local and regional species richness through species extinctions and latitudinal ranges shifts. Here, we show that species richness of fish in the North Sea, a group of ecological and socio-economical importance, has increased over a 22-year period and that this rise is related to higher water temperatures. Over eight times more fish species displayed increased distribution ranges in the North Sea (mainly small-sized species of southerly origin) compared with those whose range decreased (primarily large and northerly species). This increase in species richness can be explained from the fact that fish species richness in general decreases with latitude. This observation confirms that the interaction between large-scale biogeographical patterns and climate change may lead to increasing species richness at temperate latitudes.     

Islands in the sky or squeezed at the top? Ecological causes of elevational range limits in montane salamanders

Many animal and plant taxa reach their highest endemism and species richness in montane regions. The study of elevational range limits is central to understanding this widespread pattern and to predicting the responses of montane species to climate change. Yet, because large‐scale manipulations of the distributions of most species are difficult, the causes of species’ elevational range limits (e.g. competitive interactions, physiological specialization) are poorly understood. Here, we harness the power of new mechanistic approaches to dissect the factors that underlie the elevational replacement of two salamander species in the Appalachian Highlands. Our results challenge the long‐held idea that competitive interactions drive the lower elevational range limits of montane species and that physiological stress prevents low‐elevation species from expanding to high elevations. We show that physiological constraints drive the lower elevational range limit of the montane‐endemic species, Plethodon jordani. Conversely, we find that competition with P. jordani prevents the low‐elevation species, P. teyahalee, from expanding its range to include higher‐elevation habitats. These results are broadly consistent with the biogeography and behavior of other montane species, suggesting that similar mechanisms underlie patterns of elevational zonation across a variety of taxa and montane regions. To the extent that our findings are taxonomically and geographically widespread, these results challenge the idea that competitive release at species’ lower elevational range limits is driving the downslope range shifts exhibit by some montane taxa. Instead, our results raise the sobering possibility that even small changes in climate might cause erosion of the ranges of many high‐elevation species.     

Contrasting patterns of lichen functional diversity and species richness across an elevation gradient

Major environmental gradients co‐vary with elevation and have been a longstanding natural tool allowing ecologists to study global diversity patterns at smaller scales, and to make predictions about the consequences of climate change. These analyses have traditionally studied taxonomic diversity, but new functional diversity approaches may provide a deeper understanding of the ecological mechanisms driving species assembly. We examined lichen taxonomic and functional diversity patterns on 195 plots (200 m²) together with forest structure along an elevational gradient of 1000 m in a temperate low mountain range (Bohemian Forest, Germany). Along this elevation gradient temperature decreased and precipitation increased, two macroclimatic variables critical for lichens. Elevation was more important than forest structure in driving taxonomic and functional diversity. While species richness increased with elevation, functional diversity decreased and revealed that community patterns shift with elevation from random to clustered, reflecting selection for key shared traits. Higher elevations favored species with a complex growth form (which takes advantage of high moisture) and asexual reproductive mode (facilitating establishment under low temperature conditions). Our analysis highlights the need to examine alternative forms of diversity and opens the avenue for community predictions about climate change. For a regional scenario with increasing temperature and decreasing availability of moisture, we expect a loss of specialized species with a complex growth form and those with vegetative organs at higher elevations in low mountain ranges in Europe.     

Effects of spatial heterogeneity on butterfly species richness in Rocky Mountain National Park, CO, USA

We investigated butterfly responses to plot-level characteristics (plant species richness, vegetation height, and range in NDVI [normalized difference vegetation index]) and spatial heterogeneity in topography and landscape patterns (composition and configuration) at multiple spatial scales. Stratified random sampling was used to collect data on butterfly species richness from seventy-six 20 × 50 m plots. The plant species richness and average vegetation height data were collected from 76 modified-Whittaker plots overlaid on 76 butterfly plots. Spatial heterogeneity around sample plots was quantified by measuring topographic variables and landscape metrics at eight spatial extents (radii of 300, 600 to 2,400 m). The number of butterfly species recorded was strongly positively correlated with plant species richness, proportion of shrubland and mean patch size of shrubland. Patterns in butterfly species richness were negatively correlated with other variables including mean patch size, average vegetation height, elevation, and range in NDVI. The best predictive model selected using Akaike’s Information Criterion corrected for small sample size (AIC_c), explained 62% of the variation in butterfly species richness at the 2,100 m spatial extent. Average vegetation height and mean patch size were among the best predictors of butterfly species richness. The models that included plot-level information and topographic variables explained relatively less variation in butterfly species richness, and were improved significantly after including landscape metrics. Our results suggest that spatial heterogeneity greatly influences patterns in butterfly species richness, and that it should be explicitly considered in conservation and management actions.     

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.     

Distinguishing between signal and noise in faunal responses to environmental change

Aim We tested whether variation in snapshots of butterfly species composition and species richness taken from one to six years apart could be interpreted as an ecologically meaningful trend or whether they might merely reflect stochasticity. Location Field research was conducted in the Toquima Range and Shoshone Mountains, Lander and Nye counties, Nevada, USA. Methods We obtained data for 49 sites in the Toquima Range from 1996 to 2002 and 39 sites in the Shoshone Mountains from 2000 to 2002. Sites spanned the gradient of local topographic and climatic conditions in those mountain ranges. Data on species composition and species richness were based on comprehensive field inventories. We calculated similarity of species composition using the Jaccard index. We conducted one‐factor repeated‐measures analyses of variance to test whether the distribution of similarity of species composition and the distribution of mean species richness depended on the number of years between inventories. Results In both mountain ranges, much less of the difference in species composition was attributable to turnover of species composition within sites over time than to spatial differences among sites. Annual species richness in the Toquima Range was more variable than in the Shoshone Mountains, but again far less of the variation in species richness was attributable to year than to differences among sites. Main conclusions Despite the fact that desert ecosystems are not expected to be highly resilient to global environmental change, there may be a time lag between deterministic environmental changes and a detectable faunal response, even in taxonomic groups that are known to be sensitive to changes in climate and vegetation. Although information on species richness and similarity of species composition are among the most practical data to collect in managed landscapes, these measures may not be highly sensitive to environmental changes over the short to moderate term.     

21st century climate change threatens mountain flora unequally across Europe

Continental‐scale assessments of 21st century global impacts of climate change on biodiversity have forecasted range contractions for many species. These coarse resolution studies are, however, of limited relevance for projecting risks to biodiversity in mountain systems, where pronounced microclimatic variation could allow species to persist locally, and are ill‐suited for assessment of species‐specific threat in particular regions. Here, we assess the impacts of climate change on 2632 plant species across all major European mountain ranges, using high‐resolution (ca. 100 m) species samples and data expressing four future climate scenarios. Projected habitat loss is greater for species distributed at higher elevations; depending on the climate scenario, we find 36–55% of alpine species, 31–51% of subalpine species and 19–46% of montane species lose more than 80% of their suitable habitat by 2070–2100. While our high‐resolution analyses consistently indicate marked levels of threat to cold‐adapted mountain florae across Europe, they also reveal unequal distribution of this threat across the various mountain ranges. Impacts on florae from regions projected to undergo increased warming accompanied by decreased precipitation, such as the Pyrenees and the Eastern Austrian Alps, will likely be greater than on florae in regions where the increase in temperature is less pronounced and rainfall increases concomitantly, such as in the Norwegian Scandes and the Scottish Highlands. This suggests that change in precipitation, not only warming, plays an important role in determining the potential impacts of climate change on vegetation.     

Butterfly diversity and historical land cover change along an altitudinal gradient

Land cover and climate change are both major threats for biodiversity. In mountain ecosystems species have to adapt to fragmented habitats and harsh environmental conditions but so far, altitudinal effects in combination with land cover change have been rarely studied. The objective of this study was to determine the effects of altitude and historical land cover change on butterfly diversity. We studied species richness patterns of butterflies occuring in wetlands and other open habitats along an altitudinal gradient in a low mountain region (340–750 m a.s.l., Bavaria, Germany) with drastic loss of open habitats within the last 40–60 years. We recorded in 27 sites a total of 4,523 individuals of 49 butterfly species and five species of burnet moths. Species richness peaked at mid elevation and increased with patch size. Land cover change was most pronounced at high altitudes, but neither current open habitats, nor the historical loss of open habitats affected the species richness of butterflies. Neither open land specialized butterflies nor generalist and forest species were significantly affected by the loss of open habitats. However, increasing forest area in high altitudes reduces possible refuge open habitats for butterflies at their thermal distribution limits. This could lead to extinction of such butterfly species when temperatures further rise due to global warming.     

Comparative influence of spatial scale on beta diversity within regional assemblages of birds and butterflies

Aim We examined whether variation in species composition of breeding birds and resident butterflies in the Great Basin of North America depended on sampling grain (the smallest resolvable unit of study) and on the relative proximity of sampling units across the landscape. We also compared patterns between the two taxonomic groups with reference to their life‐history characteristics. Location Data for our analyses were collected from 1996 to 2003 in three adjacent mountain ranges in the central Great Basin (Lander and Nye counties, Nevada, USA): the Shoshone Mountains, Toiyabe Range and Toquima Range. Methods Data on species composition for both taxonomic groups were collecting using standard inventory methods for birds and butterflies in temperate regions. Data were compiled at three sampling grains, sites (average 12 ha), canyons (average 74 ha) and mountain ranges. For each sampling grain in turn, we calculated similarity of species composition using the Jaccard index. First, we investigated whether mean similarity of species composition among the three ranges differed as a function of the grain size at which data were compiled. Secondly, we explored whether mean similarity of species composition was greater for canyons within the same mountain range than for canyons within different mountain ranges. Thirdly, we examined whether mean similarity of species composition at the site level was different for sites within the same canyon, sites within different canyons in the same mountain range, and sites within canyons in different mountain ranges. We used a Bayesian model to analyse these comparisons. Results For both taxonomic groups, mean similarity of species composition increased as the sampling grain increased. The effect of spatial grain was somewhat greater for birds than for butterflies, especially when the intermediate sampling grain was compared with the smallest sampling grain. Similarity of species composition of butterflies at each sampling grain was greater than similarity of species composition of birds at the same grain. Mean similarity of species composition of both birds and butterflies at the canyon level and site level was affected by relative proximity of sampling locations; beta diversity increased as the relative isolation of sampling locations increased. Main conclusions The sensitivity of beta diversity to sampling grain likely reflects the effect of local environmental heterogeneity: as sampling grain increases, biotic assemblages appear more homogeneous. Although breeding birds in our study system have larger home ranges than resident butterflies, birds may have more specialized resource requirements related to vegetation structure and composition, especially at small sampling scales. The degree of variation in species composition of both taxonomic groups suggests that spatially extensive sampling will be more effective for drawing inferences about regional patterns of species diversity than intensive sampling at relatively few, smaller sites.