Testing the Stress-Gradient Hypothesis at the Roof of the World: Effects of the Cushion Plant Thylacospermum caespitosum on Species Assemblages

Many cushion plants ameliorate the harsh environment they inhabit in alpine ecosystems and act as nurse plants, with significantly more species growing within their canopy than outside. These facilitative interactions seem to increase with the abiotic stress, thus supporting the stress-gradient hypothesis. We tested this prediction by exploring the association pattern of vascular plants with the dominant cushion plant Thylacospermum caespitosum (Caryophyllaceae) in the arid Trans-Himalaya, where vascular plants occur at one of the highest worldwide elevational limits. We compared plant composition between 1112 pair-plots placed both inside cushions and in surrounding open areas, in communities from cold steppes to subnival zones along two elevational gradients (East Karakoram: 4850–5250 m and Little Tibet: 5350–5850 m). We used PERMANOVA to assess differences in species composition, Friedman-based permutation tests to determine individual species habitat preferences, species-area curves to assess whether interactions are size-dependent and competitive intensity and importance indices to evaluate plant-plant interactions. No indications for net facilitation were found along the elevation gradients. The open areas were not only richer in species, but not a single species preferred to grow exclusively inside cushions, while 39–60% of 56 species detected had a significant preference for the habitat outside cushions. Across the entire elevation range of T. caespitosum, the number and abundance of species were greater outside cushions, suggesting that competitive rather than facilitative interactions prevail. This was supported by lower soil nutrient contents inside cushions, indicating a resource preemption, and little thermal amelioration at the extreme end of the elevational gradient. We attribute the negative associations to competition for limited resources, a strong environmental filter in arid high-mountain environment selecting the stress-tolerant species that do not rely on help from other plants during their life cycle and to the fact the cushions do not provide a better microhabitat to grow in.     

Transactions of the Botanical Society of Edinburgh

Background: Due to the dry continental climate, the mountains of eastern Ladakh are unglaciated up to 6200–6400 m, with relatively large areas of developed soils between 5600 and 6000 m covered by sparse subnival vegetation. However, there are no studies on the composition of plant assemblages from such extreme elevations, their microclimates, vertical distributions and adaptive strategies.

Aims: The subnival vegetation was described and the relationship between microclimate, species distribution and species functional traits was analysed.

Methods: In total, 481 vegetation samples from 91 permanent plots, a floristic database of Ladakh and extensive microclimate measurements were used. Measurements of 15 functional traits were made and their relationship with species distribution between 4600 and 6150 m was tested.

Results: The subnival zone was characterised by extreme diurnal temperature fluctuations, a short growing season (between 88 and 153 days) and low soil temperature during the growing season (between 2.9 °C and 5.9 °C). It hosted 67 species, mainly hemicryptophytes, and ranged from ca. 5600 m to the highest known occurrence of vascular plants in the region (6150 m). The most common plant families were Brassicaceae, Asteraceae, Poaceae, Fabaceae and Cyperaceae. Subnival specialists with narrow elevational ranges represented 42% of the flora; these species were shorter, had relatively higher water content and water-use efficiency and contained more nutrients and soluble carbohydrates than species with a wider elevational range.

Conclusions: The subnival vegetation of eastern Ladakh is dominated by generalist species with wide vertical ranges and not by high-elevation specialists. These findings, in view of the vast unglaciated areas available for range extension, suggest a relatively high resilience of the subnival flora to climate change in this region.     

Genetic differences in the elevational limits of native and introduced Lactuca serriola populations

Aim Differences in phenological timing might explain why populations of the annual Lactuca serriola reach higher elevational limits in a part of its introduced range than in its native range. I investigated (1) whether this difference in elevational limits has a genetic basis, (2) the importance of clinal genetic differentiation and phenotypic plasticity in phenology as responses to elevation in L. serriola, and whether these responses differ between regions, and (3) whether the realized temperature niche of L. serriola differs between the two regions. Location Plant material was collected in Canton Valais, Switzerland (native range) and the Wallowa Mountains, Oregon, USA (introduced range). The field experiment was conducted in Canton Grisons, Switzerland. Methods Plants from 20 populations collected along elevational gradients were grown in eight common gardens established at 200‐m elevational intervals (600–2000 m a.s.l.). The timing of phenological transitions was monitored and analysed with mixed‐effects models to determine differences in (1) elevational limits, and (2) clinal genetic differentiation and phenotypic plasticity as responses to elevation for plants from each region. The limits of the species along five temperature gradients were derived from generalized linear models using published occurrence data to quantify regional differences in the realized temperature niche. Results The limit of seed set (1400 m a.s.l.) was the same for plants of both regions. However, the limit of flowering, probably a better reflection of elevational limits in this study, was 400 m higher for plants from the introduced region due to their faster development. Native populations showed clines in development time with elevation consistent with expectations. However, these were weaker in introduced populations, the responses of which were rather characterized by phenotypic plasticity. Thus, although introduced populations grow at considerably cooler sites than in the native region, this is unlikely to have resulted from direct selection for tolerance of high‐elevation conditions. Main conclusions This study supports a genetic basis for differences in the elevational limits of L. serriola populations between two parts of its native and introduced range. Although it is not yet clear whether these differences evolved in the introduced range, these findings highlight the potential of alien species for gaining insights into niche evolution.     

Elevational patterns in the vascular flora of a highly diverse region in southern Mexico

We examined general and family-specific patterns of vascular plant richness along a large elevational gradient (0?C3,670?m a.s.l.), assessed the continuity of these patterns and analysed their potential underlying causes in a high diversity region of the Sierra Madre del Sur, Oaxaca, Mexico. We used a vascular plant database constructed previously. The gradient was divided into 18 200-m elevation belts. To examine elevational patterns of richness, we used both observed and estimated (interpolated) species richness, as well as genus and family observed richness, for each belt. A generalised linear model (GLM) was used to assess the effect of altitude on area-corrected species richness (standard area?=?100?km²), and a numerical classification of the elevational belts based on species richness was performed. Overall, richness at the three taxonomic levels decreased with elevation, but some individual families departed from this pattern. A sharp drop in species richness was observed at 1,800?m, and the dendrogram separated two elevational floristic groups at this elevation. The GLM revealed a significant negative effect of elevation on species richness. Despite this overall decreasing pattern for vascular plants along this extensive gradient, an examination of some family-specific patterns revealed the existence of other elevation?Cdiversity relationships, indicating taxon-specific responses to elevation. The most noticeable discontinuity in species richness, at ca. 1,800?m, is likely related to a critical temperature isocline.     

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.     

The dynamics of leaf extension in plants with diverse altitudinal ranges

Summary The dynamics of leaf extension in five species of Poa were studied with electronic auxanometers (LVDTs) along an elevational gradient from 600 to 3200 m in the Austrian Alps. Extension rates peak at midday at all elevations and rates at 20°C are almost twice as high at low elevation as compared with those from the highest sites. The low temperature threshold for leaf extension drops by 7 K over this range of elevation, with plants from the highest sites showing some extension around freezing point. Thus, there is a substantial adaptive adjustment in response of leaf extension to declining mean temperatures with increasing altitude, which is not paralleled by known altitudinal trends of photosynthetic responses in herbaceous plants.     

Heterogeneity in semi-natural grasslands: the importance of the elevational gradient

Heterogeneity patterns in semi-natural grasslands were studied within 20 sites at 4 different elevational levels ranging from 530 m a.s.1. at level 1 (southern boreal region) to 1230 m a.s.1. at level 4 (low alpine region) in Vang, southern Norway. Vegetational heterogeneity decreased with elevation for vascular plants. Cryptogam species increased in heterogeneity at mid-elevational levels (750 and 915 m a.s.1.), but heterogeneity decreased again at level 4. Mean values of soil nutrients and other soil variables (e.g. loss on ignition), and their heterogeneity increased with elevation, while pH showed a decrease for both parameters. The general increase in environmental heterogeneity in the soil was not reflected in vascular plant heterogeneity because soil variables explained 40 % variation at both level 1 and 4. However, the relative differences among levels suggest that vascular plant species patterns were more sensitive to soil heterogeneity at lower elevational levels. Soil variables were also relatively more important in structuring cryptogam species at higher elevations (level 3 and 4) compared to vascular plant species. The discrepancy between vascular plants and cryptogams are also shown in the relationship between heterogeneity and diversity. Gamma diversity decreased with decreasing heterogeneity of vascular plants, while the highest value of large scale cryptogam diversity corresponded with the lowest heterogeneity value at level 4. Alfa-diversity peaked at an intermediate level of heterogeneity for vascular plants while alfa-diversity of cryptogams decreased with increasing values of heterogeneity.     

Elevational Gradient of Vascular Plant Species Richness and Endemism in Crete – The Effect of Post-Isolation Mountain Uplift on a Continental Island System

Understanding diversity patterns along environmental gradients and their underlying mechanisms is a major topic in current biodiversity research. In this study, we investigate for the first time elevational patterns of vascular plant species richness and endemism on a long-isolated continental island (Crete) that has experienced extensive post-isolation mountain uplift. We used all available data on distribution and elevational ranges of the Cretan plants to interpolate their presence between minimum and maximum elevations in 100-m elevational intervals, along the entire elevational gradient of Crete (0–2400 m). We evaluate the influence of elevation, area, mid-domain effect, elevational Rapoport effect and the post-isolation mountain uplift on plant species richness and endemism elevational patterns. Furthermore, we test the influence of the island condition and the post-isolation mountain uplift to the elevational range sizes of the Cretan plants, using the Peloponnese as a continental control area. Total species richness monotonically decreases with increasing elevation, while endemic species richness has a unimodal response to elevation showing a peak at mid-elevation intervals. Area alone explains a significant amount of variation in species richness along the elevational gradient. Mid-domain effect is not the underlying mechanism of the elevational gradient of plant species richness in Crete, and Rapoport''s rule only partly explains the observed patterns. Our results are largely congruent with the post-isolation uplift of the Cretan mountains and their colonization mainly by the available lowland vascular plant species, as high-elevation specialists are almost lacking from the Cretan flora. The increase in the proportion of Cretan endemics with increasing elevation can only be regarded as a result of diversification processes towards Cretan mountains (especially mid-elevation areas), supported by elevation-driven ecological isolation. Cretan plants have experienced elevational range expansion compared to the continental control area, as a result of ecological release triggered by increased species impoverishment with increasing elevation.     

Linking species thermal tolerance to elevational range shifts in upland dung beetles

Climate warming has been proposed as the main cause of the recent range shifts seen in many species. Although species' thermal tolerances are thought to play a key role in determining responses to climate change, especially in ectotherms, empirical evidence is still limited. We investigate the connection between species' thermal tolerances, elevational range and shifts in the lower elevational limit of dung beetle species (Coleoptera, Aphodiidea) in an upland region in the northwest of England. We measured thermal tolerances in the laboratory, and used current and historical distribution data to test specific hypotheses about the area's three dominant species, particularly the species most likely to suffer from warming: Agollinus lapponum. We found marked differences between species in their minimum and maximum thermal tolerance and in their elevational range and patterns of abundance. Overall, differences in thermal limits among species matched the abundance patterns along the elevation gradient expected if distributions were constrained by climate. Agollinus lapponum abundance increased with elevation and this species showed lower maximum and minimum thermal limits than Acrossus depressus, for which abundance declined with elevation. Consistent with lower tolerance to high temperature, we recorded an uphill retreat of the low elevation limit of A. lapponum (177 m over 57 yr) in line with the increase in summer temperature observed in the region over the same period. Moreover, this species has been replaced at low and mid‐elevations by the other two warm‐tolerant species (A. depressus and Agrilinus ater). Our results provide empirical evidence that species' thermal tolerance constrains elevational ranges and contributes to explain the observed responses to climate warming. A mechanistic understanding of how climate change directly affects species, such as the one presented here, will provide a robust base to inform predictions of how individual species and whole assemblages may change in the future.     

Seed dispersal of a fleshy-fruited invasive shrub is affected by changes in the frugivorous bird assemblage along an elevational gradient

Seed dispersal by birds constitutes an essential mechanism for ornithochorous exotic plants to successfully invade a new system. New biotic associations with native birds might facilitate the upward spread of exotic plants from the foothills into the high mountains. However, environmental changes associated with elevation are known to drive changes in bird assemblages, and it is not clear how elevation changes impact the seed dispersal service of ornithochorous invaders. We evaluated changes in frugivorous bird assemblages of one of the exotic shrubs (Cotoneaster franchetii, Rosaceae) with the broadest elevation range among woody invaders in the Córdoba Mountains (Argentina). We quantified frugivory interactions (including absolute and proportional fruit consumption by seed dispersers, pulp consumers, and seed predators) using 4-h observations of focal C. franchetii shrubs distributed across low-elevation, mid-elevation, and high-elevation sites (700, 1100, and 1800 m a.s.l., respectively; 15 individuals per elevational band and one site per elevation). Seed disperser richness was highest at the low- and mid-elevation sites (three species vs. one at the high-elevation site), but proportional and absolute fruit consumption of C. franchetii was highest at the high-elevation site (39.1%, 88 seeds at high-elevation and 7.7%, 20 seeds at low-elevation). The Chiguanco Thrush (Turdus chiguanco, Turdidae) was the only seed disperser species found at the highest elevation site. Fruit consumption by seed dispersers was positively related to their abundance and elevation. In a high mountain system, a single abundant generalist seed disperser, rather than a high richness of seed disperser species, can uphold an effective dispersal service for an invasive ornithochorous shrub. This pattern may facilitate the spread of such plants across higher elevational ranges, thereby promoting the invasion of other exotic ornithochorous plants into upper elevations.     

Divergence of thermal physiological traits in terrestrial breeding frogs along a tropical elevational gradient

Critical thermal limits are thought to be correlated with the elevational distribution of species living in tropical montane regions, but with upper limits being relatively invariant compared to lower limits. To test this hypothesis, we examined the variation of thermal physiological traits in a group of terrestrial breeding frogs (Craugastoridae) distributed along a tropical elevational gradient. We measured the critical thermal maximum (CT_max; n = 22 species) and critical thermal minimum (CT_min; n = 14 species) of frogs captured between the Amazon floodplain (250 m asl) and the high Andes (3,800 m asl). After inferring a multilocus species tree, we conducted a phylogenetically informed test of whether body size, body mass, and elevation contributed to the observed variation in CT_max and CT_min along the gradient. We also tested whether CT_max and CT_min exhibit different rates of change given that critical thermal limits (and their plasticity) may have evolved differently in response to different temperature constraints along the gradient. Variation of critical thermal traits was significantly correlated with species’ elevational midpoint, their maximum and minimum elevations, as well as the maximum air temperature and the maximum operative temperature as measured across this gradient. Both thermal limits showed substantial variation, but CT_min exhibited relatively faster rates of change than CT_max, as observed in other taxa. Nonetheless, our findings call for caution in assuming inflexibility of upper thermal limits and underscore the value of collecting additional empirical data on species’ thermal physiology across elevational gradients.     

Patterns in diversity of anurans along an elevational gradient in the Western Ghats, South India

Aim To examine patterns in anuran species richness along an elevation gradient and identify factors that govern anuran species richness on a tropical elevational gradient. Location Sampling for anurans was carried out in Kalakad Mundanthurai Tiger Reserve (KMTR) in the southern Western Ghats, India. Methods Night‐time sampling for anuran species richness was carried out from 20 November 2004 to 20 April 2005, during the north‐east monsoon and dry seasons, using transects (50 × 2 m) and visual encounter surveys along the streams. The entire gradient was classified into thirteen 100‐m elevation zones. Sampling at the alpha (single drainage basin) level was carried out in the Chinnapul River drainage basin (40–1260 m a.s.l.) and at the gamma (landscape) level in four drainage basins. Additionally, published records were used to arrive at an empirical species richness (S) for the entire landscape. Mid‐Domain Null software was used to test for the possible influence of geometric constraints on anuran species at both the alpha and gamma levels. The influence of area under each elevation zone on empirical S was tested. The pattern in anuran species richness along the elevational gradient was investigated using: (1) species boundaries in each elevation zone and their habitat correlates, (2) abiotic factors as predictor variables, (3) mean snout vent lengths of anurans, and (4) correlation between the matrices of distance in the elevation zones based on microhabitat parameters and species composition. Cluster analysis on species presence–absence in the elevation zones was used to categorize the entire gradient into high, middle and low elevations. In these three elevation categories, pattern in composition of species was examined for endemism in Western Ghats–Sri Lanka biodiversity hotspot, uniqueness to an elevation zone, adaptations of adults and modes of breeding. Results Species richness at the alpha level increased linearly with elevation, while at the gamma level there were three peaks. Maximum species richness was observed at the highest elevation (1200 m) at both the alpha and the gamma levels. The observed patterns differed significantly from mid‐domain null predictions. The multi‐modal pattern in species richness was a consequence of overlapping species range boundaries. Soil temperature was the best single measure in explaining the majority of variation in species richness at the alpha level (r² = 0.846, P < 0.01). However, soil moisture was the best predictor when both the alpha and the gamma sites were pooled (r² = 0.774, P < 0.01). Anuran body size decreased with an increase in elevation. The highest proportions of endemic and unique species were found at high elevations (> 700 m). The proportion of arboreal anurans increased from low to high elevation. Anurans exhibiting direct development were predominantly found at high elevations. Main conclusions Geometric constraints did not influence anuran species richness along the elevational gradient. Overlapping range boundaries influenced species richness at the gamma level. Abiotic factors such as soil temperature and moisture influenced anuran species richness in the mountain range. The ‘Massenerhebung effect’ could be responsible for range restriction and endemism of anurans, differences in guilds and mode of reproduction. These findings highlight the importance of cloud forests for endemic anurans.     

Thermal constraints on embryonic development as a proximate cause for elevational range limits in two Mediterranean lacertid lizards

Local adaptation and range restrictions in alpine environments are central topics in biogeographic research with important implications for predicting impacts of global climate change on organisms. Temperature is strongly coupled to elevation and greatly affects life history traits of oviparous reptiles in mountain environments. Thus, species may encounter barriers for expanding their ranges if they are unable to adapt to the changing thermal conditions encountered along elevational gradients. We sought to determine whether thermal requirements for embryonic development provide a plausible explanation for elevational range limits of two species of lacertid lizards that have complementary elevational ranges in a Mediterranean mountain range (Psammodromus algirus is found at elevations below 1600 m and Iberolacerta cyreni is found at elevations above 1600 m). We combined experimental incubation of eggs in the laboratory with modelled estimates of nest temperature in the field. In both species, increasing temperature accelerated development and produced earlier hatching dates. The species associated with warmer environments (P. algirus) experienced an excessive hatching delay under the lowest incubation temperature. Moreover, newborns from eggs incubated at low temperatures showed poor body condition and very slow rates of postnatal growth. In contrast, eggs of the strictly alpine species I. cyreni exhibited shorter incubation periods than P. algirus that allowed hatching before the end of the active season even under low incubation temperatures. This was countered by lower reproductive success at higher temperatures, due to lower hatching rates and higher incidence of abnormal phenotypes. Elevational range limits of both species coincided well with threshold temperatures for deleterious effects on embryonic development. We suggest that incubation temperature is a major ecophysiological factor determining the elevational range limits of these oviparous lizards with predictable consequences for mountain distributions under future warmer climates.     

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.     

Support for the elevational Rapoport's rule among seed plants in Nepal depends on biogeographical affinities and boundary effects

As one of the most important hypotheses on biogeographical distribution, Rapoport's rule has attracted attention around the world. However, it is unclear whether the applicability of the elevational Rapoport's Rule differs between organisms from different biogeographical regions. We used Stevens’ method, which uses species diversity and the averaged range sizes of all species within each (100 m) elevational band to explore diversity‐elevation, range‐elevation, and diversity‐range relationships. We compared support for the elevational Rapoport's rule between tropical and temperate species of seed plants in Nepal. Neither tropical nor temperate species supported the predictions of the elevational Rapoport's rule along the elevation gradient of 100–6,000 m a.s.l. for any of the studied relationships. However, along the smaller 1,000–5,000 m a.s.l. gradient (4,300 m a.s.l. for range‐elevation relationships) which is thought to be less influenced by boundary effects, we observed consistent support for the rule by tropical species, although temperate species did not show consistent support. The degree of support for the elevational Rapoport's rule may not only be influenced by hard boundary effects, but also by the biogeographical affinities of the focal taxa. With ongoing global warming and increasing variability of temperature in high‐elevation regions, tropical taxa may shift upward into higher elevations and expand their elevational ranges, causing the loss of temperate taxa diversity. Relevant studies on the elevational Rapoport's rule with regard to biogeographical affinities may be a promising avenue to further our understanding of this rule.     

Richness and Composition of Vascular Plants and Cryptogams along a High Elevational Gradient on Buddha Mountain, Central Tibet

We explored patterns of plant species richness and composition along an elevational gradient (4,985?C5,685?m a.s.l.) on Buddha Mountain, 100?km northwest of Lhasa, Tibet. We recorded the presence of plants and lichens in 1-m² quadrats separated by 25-m elevational intervals (174 quadrats in 29 elevational bands) along a vertical transect with a SE aspect. We recorded 143 total species, including 107 angiosperms, 2 gymnosperms, 27 lichens, and 7 mosses. We measured stone cover in each quadrat, and soil pH, C, N and C/N ratio from two randomly located samples collected from 10-cm depth within each band. C, N and C/N decreased with elevation, stoniness increased and soil pH did not change with altitude. We employed detrended correspondence analysis (DCA), canonical correspondence analysis (CCA) and generalized linear models (GLMs) to assess the relationships of species richness and species composition to the environment. The first two axes of the CCA biplot explained 87.7% of total variation in the species-environment relationship, and 27.7% of total variance of species data. The first CCA axis is associated with elevation, while the second axis is related to soil pH and stone cover. We also compared patterns in species richness against expectations from species pools interpolated from the literature. Total species richness was relatively constant between 4,985 and 5,400?m a.s.l. and declined continuously above 5,400?m a.s.l. Similar declining patterns were observed for forbs and graminoids. Cushion plants and lichens abundance exhibited a unimodal relationship with altitude while shrubs declined monotonically. Except for lichens, models derived from our observations and the literature were quite similar in shape. The proportion of the species pool represented in each elevational band increased as a function of elevation for non-vascular plants, but decreased markedly for vascular plants. Thus, vascular plants are more likely to be constrained by dispersal at higher elevations, resulting in more local endemism, while the relatively easily-dispersed high-elevation cryptogams have little local differentiation. Our comparative approach demonstrates that complex scale-dependent differences between life forms may underlie the apparent simplicity of elevational gradients. Furthermore, elevational gradients summarized from distributional notes cannot be assumed to be proxies for elevational gradients on individual mountain slopes.     

Tree recruitment of European tree species at their current upper elevational limits in the Swiss Alps

Aim The physical and physiological mechanisms that determine tree‐line position are reasonably well understood, but explanations for tree species‐specific upper elevational limits below the tree line are still lacking. In addition, once these uppermost positions have been identified, questions arise over whether they reflect past expansion events or active ongoing recruitment or even upslope migration. The aims of this study were: (1) to assess current tree recruitment near the cold‐temperature limit of 10 major European tree species in the Swiss Alps, and (2) to rank species by the extent that their seedlings and saplings exceed the elevational limit of adult trees, possibly reflecting effects of the recent climate warming. Location Western and eastern Alps of Switzerland. Methods For each species, occurrences were recorded along six elevational transects according to three size classes from seedlings to adult trees in 25‐m‐elevation steps above and below their regional upper elevational limit. Two methods were used to compare upper elevational limits between seedlings, saplings and adults within species. First, we focused on the uppermost occurrence observed in each life stage for a given species within each studied region; and second, we predicted their upper distribution range using polynomial models fitted to presence/absence data. Results Species exhibited a clear ranking in their elevational limit. Regional differences in species limits (western versus eastern Swiss Alps) could largely be attributed to regional differences in temperature. Observed and predicted limits of each life stage showed that all species were represented by young individuals in the vicinity of the limit of adult trees. Moreover, tree recruitment of both seedlings and saplings was detected and predicted significantly beyond adult tree limits in most of the species. Across regions, seedlings and saplings were on average found at elevations 73 m higher than adult trees. Main conclusions Under current conditions, neither seed dispersal nor seedling establishment constitutes a serious limitation of recruitment at the upper elevational limits of major European trees. The recruits found beyond the adult limits demonstrate the potential for an upward migration of trees in the Alps in response to ongoing climate warming.     

Foliar C:N Ratio of Ferns along an Andean Elevational Gradient1

We studied the concentration of leaf N and C among 183 fern species along an elevational gradient at 1700 to 3400 m in humid montane forest in the Bolivian Andes at different levels of taxonomic resolution. For two species of Elaphoglossum sampled 8 and 14 times, respectively, there were no elevational trends. Similarly, a contrast of 22 species with wide elevational amplitudes sampled at their highest and lowest locations did not show any change in C or N contents, or in C:N ratios with elevation. At the community level, however, the mean values of C:N ratios for (a) all species found at a given elevation showed a significant decline with increasing elevation and (b) among epiphytic species, higher ratios (i.e., lower relative N content) than among terrestrial species at the same elevation. These trends were opposite to those of the upper soil layer, in which C:N ratios increased with elevation.     

Elevation‐richness pattern of vascular plants in wadis of the arid mountain Gebel Elba,Egypt

Mountains provide a unique opportunity to study drivers of species richness across relatively short elevation gradients. However, few studies have reported elevational patterns for arid mountains. We studied elevation‐richness pattern along an elevational gradient at the arid mountain Gebel Elba, south‐east of Egypt, expecting a unimodal richness pattern. We sampled 133 vegetation plots (10 × 10 m) in four wadis along an elevational gradient from 130 to 680 m which represents the transition from desert to mountain wadi systems. We used generalised additive models to describe the relationship between elevation and plant species richness. We found a strong increase in species richness and Shannon diversity at low elevations followed by a plateau at mid‐ to high elevations. When we analysed each tributary as a single gradient, no pattern was found. The analysed elevational gradient seems to be a major stress gradient in terms of temperature and water availability, exhibiting a trend of increasing species richness that changes to a plateau pattern; a pattern rarely observed for wadi systems in arid mountains. We discuss the observed pattern with the climatic stress hypothesis and the environmental heterogeneity hypothesis as possible explanations for the pattern.     

Changes to the elevational limits and extent of species ranges associated with climate change

The first expected symptoms of a climate change‐generated biodiversity crisis are range contractions and extinctions at lower elevational and latitudinal limits to species distributions. However, whilst range expansions at high elevations and latitudes have been widely documented, there has been surprisingly little evidence for contractions at warm margins. We show that lower elevational limits for 16 butterfly species in central Spain have risen on average by 212 m (± SE 60) in 30 years, accompanying a 1.3 °C rise (equivalent to c. 225 m) in mean annual temperature. These elevational shifts signify an average reduction in habitable area by one‐third, with losses of 50–80% projected for the coming century, given maintenance of the species thermal associations. The results suggest that many species have already suffered climate‐mediated habitat losses that may threaten their long‐term chances of survival.