Effects of topographic and edaphic conditions on alpine plant species distribution along a slope gradient on Mount Norikura,central Japan

This study examined the effects of topographic and edaphic conditions on alpine plant species distribution along a slope gradient on Mt. Norikura (3026 m a.s.l.), central Japan. Topographic and edaphic factors investigated at 40 plots were: slope inclination, ground surface texture, soil water content and soil inorganic nitrogen concentration (NO₃-N, NH₄-N). The topographic and edaphic factors changed with slope positions: slope inclination was steeper, soil texture was coarser, and soil water and inorganic nitrogen concentration decreased with increasing slope position. Five vegetation types were located along the slope gradient and related to two factor-groups: (1) changes in soil water, NH₄-N, slope inclination along the slope gradient, and (2) ground surface texture. A tall herbaceous plant community developed at the low slope position, near tarns, with fine soil surface texture, high soil water and NH₄-N, while Dicentra peregrina dominated on an unstable rubble slope near the ridge top. The distribution of each species was predictable from the two factor-groups. Although the five vegetation types were related to the two factor-groups, responses to the two factor-groups differed among species, even within the same vegetation type. Therefore, this study showed that the topography of the terrain largely regulated alpine plant distribution by affecting edaphic conditions, and that global warming may alter species composition by changing edaphic conditions.     

Effects of climate on the radial growth of tree species in the upper and lower distribution limits of an altitudinal ecotone on Mount Norikura,central Japan

Tree-ring width chronologies were developed for Abies veitchii, Betula ermanii and Betula platyphylla var. japonica in their altitudinal ecotone (approximately 1600m a.s.l.) on Mount Norikura, central Japan, to determine what climatic conditions affect the growth of tree species in the upper and lower distribution limits of an altitudinal ecotone. This altitude was the lower distribution limit for A.veitchii and B.ermanii in the subalpine zone, and was the upper distribution limit for B.platyphylla var. japonica in the montane zone on Mount Norikura. Tree-ring widths of the two Betula species and A.veitchii were positively correlated with the August precipitation of the current and previous years, respectively. Precipitation in August (the hottest month) was reduced compared with other months during summer. Tree-ring width of B.platyphylla var. japonica showed no correlation with temperatures in any month in its upper distribution limit. In contrast, tree-ring widths of B.ermanii and A.veitchii were negatively correlated with the August temperatures of the current and previous years, respectively, at the lower distribution limit of these species. Therefore, the two Betula species and A.veitchii responded to climatic conditions of the current and previous years, respectively. The present study also suggests that a water deficit in August reduces growth of these three species in this altitudinal ecotone, irrespective of the upper or lower distribution limits, and that a high August temperature is more detrimental to the growth of A.veitchii and B.ermanii in their lower distribution limits. Thus, the three species with different altitudinal distributions examined in the present study responded differently to climatic conditions in this altitudinal ecotone on Mount Norikura.     

青藏高原高寒草甸物种多样性的海拔梯度分布格局及对地上生物量的影响

植物物种多样性在海拔梯度上的变化规律以及物种多样性与生产力的关系是生态学研究的热点, 至今还没有得出一般性规律。本文以青海省海南藏族自治州贵德县的拉脊山(36°21′ N, 101°27′ E, 海拔3,389-3,876 m)和果洛藏族自治州的玛沁县军牧场山体(34°22′ N, 100°30′ E, 海拔4,121-4,268 m)为研究对象, 对植物高度、盖度、地上生物量和物种多样性随海拔高度的变化进行调查和统计分析, 以探讨青藏高原高寒草甸的物种多样性和地上生物量在海拔梯度上的变化规律及两者的关系。结果表明: (1)两条山体样带上地上生物量与物种多样性随海拔的变化规律一致: 随着海拔的升高, 地上生物量线性降低; Shannon-Wiener指数、Simpson指数和物种丰富度都呈单峰曲线, 在中间海拔最大, 而Pielou指数随海拔的升高线性增加。结合目前针对青藏高原高寒草甸的研究数据, 发现物种丰富度随海拔高度的变化均呈单峰曲线, 说明随着海拔的升高物种多样性先升高后降低可能是青藏高原物种多样性分布的普遍规律。(2)地上生物量与物种多样性的关系在两条山体样带上表现一致: 地上生物量随Shannon- Wiener指数、Simpson指数和Pielou指数的升高而线性降低, 但与物种丰富度不相关。综合两条山体样带所有样方数据, 发现地上生物量与Shannon-Wiener指数、Simpson指数不相关, 而随物种丰富度的升高线性增加。结合目前在青藏高原的相关研究数据, 发现地上生物量与物种丰富度呈S型曲线(logistic model)。     

Changes in plant biomass and species composition of alpine Kobresia meadows along altitudinal gradient on the Qinghai-Tibetan Plateau

Alpine Kobresia meadows are major vegetation types on the Qinghai-Tibetan Plateau. There is growing concern over their relationships among biodiversity, productivity and environments. Despite the im-portance of species composition, species richness, the type of different growth forms, and plant bio-mass structure for Kobresia meadow ecosystems, few studies have been focused on the relationship between biomass and environmental gradient in the Kobresia meadow plant communities, particularly in relation to soil moisture and edaphic gradients. We measured the plant species composition, her-baceous litter, aboveground and belowground biomass in three Kobresia meadow plant communities in Haibei Alpine Meadow Ecosystem Research Station from 2001 to 2004. Community differences in plant species composition were reflected in biomass distribution. The total biomass showed a de-crease from 13196.96±719.69 g/m2 in the sedge-dominated K. tibetica swamp to 2869.58±147.52 g/m2 in the forb and sedge dominated K. pygmaea meadow, and to 2153.08±141.95 g/m2 in the forbs and grasses dominated K. humilis along with the increase of altitude. The vertical distribution of below-ground biomass is distinct in the three meadow communities, and the belowground biomass at the depth of 0-10 cm in K. tibetica swamp meadow was significantly higher than that in K. humilis and K. pygmaea meadows (P<0.01). The herbaceous litter in K. tibetica swamp was significantly higher than those in K. pygnaeca and K. humilis meadows. The effects of plant litter are enhanced when ground water and soil moisture levels are raised. The relative importance of litter and vegetation may vary with soil water availability. In the K. tibetica swamp, total biomass was negatively correlated to species richness (P<0.05); aboveground biomass was positively correlated to soil organic matter, soil moisture, and plant cover (P<0.05); belowground biomass was positively correlated with soil moisture (P<0.05). However, in the K. pygnaeca and K. humilis meadow communities, aboveground biomass was posi-tively correlated to soil organic matter and soil total nitrogen (P<0.05). This suggests that the distribu-tion of biomass coincided with soil moisture and edaphic gradient in alpine meadows.     

Morphological variations of the Solidago virgaurea L. complex along an elevational gradient on Mt Norikura,central Japan

Morphological variations in vegetative and reproductive organs in the Solidago virgaurea complex were examined for eight elevations between 1600 and 2400 m a.s.l. in Japan. The rosette diameter and stem height were lower at higher elevations. The stem diameter at any stem height was greater at higher elevations, suggesting that the S. virgaurea complex increases mechanical stiffness against strong wind at high elevations. The number of flower heads at any stem height was less at high elevations (2000–2400 m a.s.l.) than at low elevations (1600–1900 m a.s.l.). Leaf nitrogen concentration did not change along the elevational gradient. Leaf mass per area (LMA) tended to decrease with increasing elevation, except for 2400 m a.s.l. The decrease of LMA would contribute to maintaining a positive carbon balance at high elevations. The number of involucral rays of flower heads was mostly four at low elevations (1600–1900 m a.s.l.) and three at high elevations (2000–2400 m a.s.l.). The number of involucral scales and diameter of flower heads were greater at high elevations (2000–2400 m a.s.l.) than at low elevations (1600–1900 m a.s.l.). Therefore, the S . virgaurea complex is suggested to adapt to high elevations that have cool temperature, a short growing season and strong wind conditions by changing its vegetative and reproductive traits.     

Geomorphological disturbance affects ecological driving forces and plant turnover along an altitudinal stress gradient on alpine slopes

The altitudinal gradient is considered as a stress gradient for plant species because the development and fitness of plant communities tend to decrease as a result of the extreme environmental conditions present at high elevations. Abiotic factors are predicted to be the primary filter for species assemblage in high alpine areas, influencing biotic interactions through both competition for resources and positive interactions among species. We hypothesised that the relative importance of the ecological driving forces that affect the biotic interactions within plant communities changes along an elevation gradient on alpine debris slopes. We used multiple gradient analyses of 180 vegetation plots along an altitudinal range from ~1,600 to 2,600 m and single 100 m-bands in the Adamello-Presanella Group (Central Alps) to investigate our hypothesis; we measured multiple environmental variables related to different ecological driving forces. Our results illustrate that resource limitations at higher elevations affect not only the shift from competition to facilitation among species. A geomorphological disturbance regime along alpine slopes favours the resilience of the high-altitude species within topographic/geomorphological traps. An understanding of the ecological driving forces and positive interactions as a function of altitude may clarify the mechanisms underlying plant responses to present and future environmental changes.     

Patterns and origin of intraspecific functional variability in a tropical alpine species along an altitudinal gradient

Background : Intraspecific functional variability (IFV) along altitudinal gradients is a powerful proxy to infer the responses of plants to abrupt environmental changes. We envisage that IFV shows distinctive patterns in tropical and extratropical alpine regions.

Aims : To characterise the patterns and explore the origin of IFV in a tropical alpine species in a context of upward range extension.

Methods : We examined variations in a series of plant functional traits in Lasiocephalus ovatus, inside and outside a nurse plant along a 600 m altitudinal gradient in the Ecuadorian Andes, and we studied its genetic variability.

Results : More conservative traits were developed at higher elevation, in contrast to extratropical alpine plants, which commonly develop opportunistic traits in response to late snowmelt close to their upper altitudinal limit. The presence of nurse cushions did not alter this trend. Increasing genetic distance along the gradient suggested that IFV might be partly genetically induced.

Conclusions : Our data combined with existing literature in tropical alpine environments lead the way to a stimulating scientific challenge: determining if patterns of plant altitudinal distribution in tropical alpine areas in response to climate change are predictable from patterns described in extratropical alpine areas.     

Flowering phenology and reproduction of the Solidago virgaurea L. complex along an elevational gradient on Mt Norikura,central Japan

This study examined the flowering phenology and reproductive traits of the Solidago virgaurea complex at four elevations in the subalpine zone in Japan using a bagging experiment. Flowering started earlier at higher elevations. Syrphid flies mainly visited flowers of the S. virgaurea complex, and the number of visits was considerably lower at the lowest elevation than at the three other elevations. Although the number of seeds per individual did not differ among the four elevations, total achene weight per individual was lower at the lowest elevation than at the three other elevations. The weight of an achene and seed germination rate of the control were much greater at higher elevations than at the lowest elevation. The weight of an achene and germination rate for the control were as low as the bagging treatment at the lowest elevation with infrequent flower visitors, which indicates that the S. virgaurea complex is a facultative outcrosser. The S. virgaurea complex is thought to produce seeds during a short growing season at high elevations by starting to flower earlier, and the large seed size is advantageous for seedling establishment at high elevations. Therefore, elevational changes in flowering phenology and reproductive traits are thought to be an adaptation to the short growing season at high elevations.     

不同海拔高度高寒草甸光能利用效率的遥感模拟

利用植被光合模型模拟了藏北高原3个海拔高度(4300,4500 m和4700 m)的高寒草甸生态系统的光能利用效率.海拔4500 m的光能利用效率均值(0.47 g C/MJ)显著高于海拔4300 m(0.38 g C/MJ)和4700 m(0.35 g C/MJ),而海拔4300 m和4700 m两者间差异不显著.相关分析和多重逐步回归分析表明,影响每个海拔光能利用效率季节变化的主要因子为空气温度,相对湿度以及地表水分指数,这3个因子共同解释了99％以上的光能利用效率的季节变化,其中空气温度的贡献最大,相对湿度的贡献次之,地表水分指数的贡献最小,这说明在3个海拔的任何一个海拔高度,温度对光能利用效率季节变化的胁迫作用大于水分对光能利用效率季节变化的胁迫作用.多重逐步线性回归分析表明,生长季节均土壤含水量是决定生长季节均光能利用效率沿海拔高度分布的主导因子.单因子线性回归分析表明,地表水分指数可以定量化高寒嵩草草甸生态系统水分状况,它同时可以反应土壤水分、近地表空气湿度以及生态系统植被含水量状态.因此,在高寒嵩草草甸生态系统,用地表水分指数反应生态系统尺度水分对光能利用效率的胁迫作用是可行的.     

Rotifer species richness along an altitudinal gradient in the Alps

Aim Biodiversity patterns along altitudinal gradients are less studied in aquatic than terrestrial systems, even though aquatic sites provide a more homogeneous environment independent of moisture constraints. We studied the altitudinal species richness pattern for planktonic rotifers in freshwater lakes and identified the environmental predictors for which altitude is a proxy. Location Two hundred and eighteen lakes of Trentino–South Tyrol (Italy) in the eastern Alps; lakes covered 98% (range 65–2960 m above sea level) of the altitudinal gradient in the Alps. Methods We performed: (1) linear regression between species richness and altitude to evaluate the general pattern, (2) multiple linear regression between species richness and environmental predictors excluding altitude to identify the most important predictors, and (3) linear regression between the residuals of the best model of step (2) and altitude to investigate any additional explanatory power of altitude. Selection of environmental predictors was based on limnological importance and non‐parametric Spearman correlations. We applied ordinary least squares regression, generalized linear, and generalized least squares modelling to select the most statistically appropriate model. Results Rotifer species richness showed a monotonic decrease with altitude independent of scale effects. Species richness could be explained (R²= 51%) by lake area as a proxy for habitat diversity, reactive silica and total phosphorus as proxies for productivity, water temperature as a proxy for energy, nitrate as a proxy for human influence and north–south and east–west directions as covariates. These predictors completely accounted for the species richness–altitude pattern, and altitude had no additional effect on species richness. Main conclusions The linear decrease of species richness along the altitudinal gradient was related to the interplay of habitat diversity, productivity, heat content and human influence. These factors are the same in terrestrial and aquatic habitats, but the greater environmental stability of aquatic systems seems to favour a linear pattern.     

Analysis of the vertical distribution of epiphytic lichens on Pinus nigra (Mount Olympos,Greece) along an altitudinal gradient

Epiphytic lichen vegetation was studied in 10 sites along an altitude gradient from 750 to 1510 m on NW-facing slopes of Mount Olympos, Greece to assess the main spatial heterogeneity of microhabitats affecting communities and species composition. Community structure along the gradient was studied by using multivariate techniques. The critical factor for spatial heterogeneity seems to be the height at which the lichen community develops on the tree trunks. Changes in the community structure of the epiphytic lichen vegetation were also detected along the altitudinal gradient and the altitude of 1200 m is considered to be an ecotone. A comparative study of epiphytic lichen communities on Mt. Olympos and in the Thessaloniki area revealed species indicators of air pollution.     

Main vegetation types and plant species diversity along an altitudinal gradient of Al Baha region,Saudi Arabia

Plant species composition patterns and vegetation types were investigated along Elevational Gradients in Al Baha region, Saudi Arabia. Sandy plain, wadis, drainage lines, rocky outcrops, hills and fallow lands occur over a wide geographic range encompassing variation in plant species and communities among these different ecological sites. To provide a quantitatively based classification of the vegetation we used Multi Variant Statistical Package (MVSP) software, followed by the re-arrangement of a matrix of the similar plant species in rows and similar sample sites in columns. Plant density and environmental variables were measured and recorded in each quadrat. Two-way indicator species analysis and Canonical Correspondence Analysis (CCA) were used to analyze the relationships between vegetation and environmental variables, while Arc Map was used to analyze the pattern of plant species density. A total of 59 sample plots (25 × 25 m), stratified, randomly-placed relevés were collected in Al Baha region, along a cross section running from south-west to north-west. About 190 plant species belonging to 59 families were recognized. This study showed that these plant species formed 15 vegetation types that primarily correspond mainly to different combinations of elevation, and topography. The study concluded that this research has provided the first quantitative and systematic survey of the vegetation in Al Baha region.     

Effects of temperature on photosynthesis of two morphologically contrasting plant species along an altitudinal gradient in the tropical high Andes

The effects of temperature on photosynthesis of a rosette plant growing at ground level, Acaena cylindrostachya R. et P., and an herb that grows 20–50 cm above ground level, Senecio formosus H.B.K., were studied along an altitudinal gradient in the Venezuelan Andes. These species were chosen in order to determine – in the field and in the laboratory – how differences in leaf temperature, determined by plant form and microenvironmental conditions, affect their photosynthetic capacity. CO₂ assimilation rates (A) for both species decreased with increasing altitude. For Acaena leaves at 2900 m, A reached maximum values above 9 μmol m⁻² s⁻¹, nearly twice as high as maximum A found at 3550 m (5.2) or at 4200 m (3.9). For Senecio leaves, maximum rates of CO₂ uptake were 7.5, 5.8 and 3.6 μmol m⁻² s⁻¹ for plants at 2900, 3550 and 4200 m, respectively. Net photosynthesis-leaf temperature relations showed differences in optimum temperature for photosynthesis (A _o.t.) for both species along the altitudinal gradient. Acaena showed similar A _o.t. for the two lower altitudes, with 19.1°C at 2900 m and 19.6°C at 3550 m, while it increased to 21.7°C at 4200 m. Maximum A for this species at each altitude was similar, between 5.5 and 6.0 μmol m⁻² s⁻¹. For the taller Senecio, A _o.t. was more closely related to air temperatures and decreased from 21.7°C at 2900 m, to 19.7°C at 3550 m and 15.5°C at 4200 m. In this species, maximum A was lower with increasing altitude (from 6.0 at 2900 m to 3.5 μmol m⁻² s⁻¹ at 4200 m). High temperature compensation points for Acaena were similar at the three altitudes, c. 35°C, but varied in Senecio from 37°C at 2900 m, to 39°C at 3550 m and 28°C at 4200 m. Our results show how photosynthetic characteristics change along the altitudinal gradient for two morphologically contrasting species influenced by soil or air temperatures. Received: 5 July 1997 / Accepted: 25 October 1997     

Local adaptation and cogradient selection in the alpine plant, Poa hiemata, along a narrow altitudinal gradient

Alpine environments are particularly susceptible to environmental changes associated with global warming but there is potential for alpine plants to adapt to warming if local adaptation occurs and gene flow allows genotypes adapted to low altitudes to colonize higher altitude sites. Here we examine the adaptive potential of a common alpine grass, Poa hiemata, within the restricted alpine habitat of Australian mountains, across a narrow altitudinal gradient replicated in three areas. Grasses at high altitude sites had shorter leaf lengths and larger circumferences than those at lower sites. Transplant experiments with clonal material and plants grown from seed indicated that these differences were partly genetic, with environmental and genetic factors both contributing to the differences between altitudes. Differences in altitudinal forms were also evident in a common garden experiment. Plants showed a home-site advantage in terms of survival. A fitness analysis indicated that at high altitude sites, selection favored plants with short leaves and larger circumferences, whereas these traits were selected in the opposite direction at the low altitude sites. These findings indicate cogradient selection and potential for both plastic and genotypic shifts in response to climate change in P. hiemata.     

Short-term variation in species richness across an altitudinal gradient of alpine summits

In response to climate warming, high altitude alpine vegetation may be replaced by typically lower altitude species, as species re-assemble and migrate to new areas. However, empirical evidence showing vegetation change in response to climate warming is largely unavailable for Australian alpine areas. Here, we examine changes in species richness with respect to climate and altitude over a 7?year period at a range of spatial scales in a re-survey of five alpine summits that are part of the Global Observation Research Initiative in Alpine Environments monitoring network. Eighty species were recorded in 2011 across all summits, an increase of 6 species since 2004. Mean species richness increased at the whole-of-summit scale from 45 to 50 species (about 12?%). At this scale, the rate of species richness increase was almost one new species per year, with 15 new species recorded at one summit. Here, shrub and graminoid species showed the largest increases. At the smaller spatial scales, changes in species richness were less pronounced. Turnover at the species and community level was typically moderate at all spatial scales and on all summits. The strength and direction of species richness change (the difference in species richness between the two sample periods, +/?) was not related to altitude nor variation in climate. Future re-surveys of the summits will confirm whether these short-term variations in species richness, particularly increases in shrubs, are indeed signals of longer-term trends and interactions with a changing climate.     

Unimodal pattern of soil hydrophobicity along an altitudinal gradient encompassing Mediterranean,temperate, and alpine ecosystems

Background and Aims

Soil water repellency (SWR, i.e. the reduced affinity for water due to the presence of hydrophobic coatings on soil particles) has relevant hydrological implications on the rate of water infiltration, surface runoff, and overland flow. Here, we test how SWR varies along a 2490 m altitudinal gradient encompassing six ecosystems including Mediterranean, Temperate, and Alpine vegetation types.

Methods

Water repellency, measured by the Molarity of an Ethanol Droplet (MED) test, was quantified in 80 soil samples collected for 16 different elevations. Soil quality was assessed by measuring soil texture, pH, organic carbon, salinity, and nutrient availability.

Results

SWR showed a unimodal pattern along the 2490 m transect, peaking at intermediate elevations. Unexpectedly, SWR was the highest under broad-leaf deciduous forests, and the lowest under evergreen, sclerophyllous Mediterranean vegetation types. The soil organic carbon content, and the pH were the main determinants of water repellency, showing respectively a positive, and a negative correlation with the SWR. In contrast, soil texture and salinity resulted unrelated to the SWR.

Conclusions

With this study we demonstrated a linkage between SWR, vegetation type and soil pH and organic carbon content along the elevation gradient. Further studies are needed to explicitly evaluate the impact SRW on erosion risk at catchment scale in the context of climatic change.

     

高黎贡山种子植物物种丰富度沿海拔梯度的变化

物种丰富度沿海拔梯度的分布格局成为生物多样性研究的热点。为探讨中尺度区域物种丰富度沿海拔梯度的分布,本文以高黎贡山为研究对象,利用该地区的地方植物志资料,结合通过GIS生成的区域数字高程模型(DEM)数据,分析了该区域全部种子植物和乔木、灌木、草本三种生活型种子植物物种丰富度的垂直分布格局以及物种密度沿海拔梯度的变化特征。结果表明：(1)全部种子植物和不同生活型植物物种丰富度随着海拔的升高呈现先增加后减小的趋势,最大值出现在海拔1500—2000m的范围;(2)物种密度与海拔也呈现单峰曲线关系;(3)物种丰富度和物种密度分布格局的形成主要受海拔所反映的水、热状况组合以及物种分布的边界影响。     

Analysis of the distribution of epiphytic lichens within homogeneousFagus sylvatica stands along an altitudinal gradient (Mount Olympos,Greece)

Epiphytic lichen vegetation onFagus sylvatica sas studied in 4 sites along an altitudinal gradient from 930 to 1500 m on SE facing slopes of Mount Olympos (Greece). The crucial factor determining the spatial heterogeneity of epiphytic lichens onF. sylvatica is the altitude and not the height on the trunk at which lichen community is established. 17 out of 26 taxa are confined to a particular elevation range, while another three are clearly ubiquitous in their distribution. The number of lichen species at breast height is higher than at the base of the trunks. The results were compared with those gathered earlier in an analogous study on the vertical distribution of epiphytic lichens onPinus nigra along an altitudinal gradient from 750 to 1510 m of the same mountain. Comparison suggests that spatial heterogeneity of epiphytic lichens onF. sylvatica is different from the one onP. nigra.     

Performance of two alpine plant species along environmental gradients in an alpine meadow ecosystem in central Tibet

Understanding how biotic interactions and abiotic conditions affect plant performance is important for predicting changes in ecosystem function and services in variable environments. We tested how performances of Astragalus rigidulus and Potentilla fruticosa change along gradients of biotic interactions (represented by plant species richness, abundance of the dominant plant species Kobresia pygmaea, and herbivory intensity) and abiotic conditions (represented by elevation, aspect, and slope steepness) across a semi-arid landscape in central Tibet. Redundancy analyses showed that the biotic variables explained 30 and 39 % of the variation in overall performance of A. rigidulus (P = 0.03) and P. fruticosa (P = 0.01), respectively. Abiotic variables did not contribute significantly to variation among A. rigidulus populations. Plant size decreased with species richness in both species and was larger on south- rather than north-facing slopes. Reproductive effort for both species was significantly negatively related to the abundance of K. pygmaea and both species had larger reproductive effort on south- rather than north- and west-facing slopes. The proportion of biomass allocated to sexual reproduction in P. fruticosa was negatively correlated with K. pygmaea abundance and herbivory intensity. The population density of P. fruticosa was positively related to elevation, species richness, and K. pygmaea abundance. We conclude that plant performance at a local scale was more strongly related to biotic than abiotic conditions, but different components of plant performance responded differently to predictor variables and the responses were species-specific. These findings have important implications for rangeland management under changing environmental conditions.