The effect of area on local and regional elevational patterns of species richness

Aim To calculate the degree to which differences between local and regional elevational species richness patterns can be accounted for by the effects of regional area. Location Five elevational transects in Costa Rica, Ecuador, La Réunion, Mexico and Tanzania. Methods We sampled ferns in standardized field plots and collated regional species lists based on herbarium and literature data. We then used the Arrhenius function S = cA^z to correct regional species richness (S) for the effect of area (A) using three slightly different approaches, and compared the concordance of local and regional patterns prior to and after accounting for the effect of area on regional richness using linear regression analyses. Results We found a better concordance between local and regional elevational species richness after including the effect of area in the majority of cases. In several cases, local and regional patterns are very similar after accounting for area. In most of the cases, the maximum regional richness shifted to a higher elevation after accounting for area. Different approaches to correct for area resulted in qualitatively similar results. Main conclusions The differences between local and regional elevational richness patterns can at least partly be accounted for by area effects, suggesting that the underlying causes of elevational richness patterns might be the same at both spatial scales. Values used to account for the effect of area differ among the different study locations, showing that there is no generally applicable elevational species–area relationship.     

Phylogenetically balanced evidence for structural and carbon isotope responses in plants along elevational gradients

We tested three hypotheses related to the functioning of mountain plants, namely their reproductive effort, leaf surface structure and effectiveness of CO₂ assimilation, using archive material from contrasting elevations. Analysis of elevational trends is at risk of suffering from two major biases: a phylogenetic bias (i.e. an elevational change in the abundance of taxonomic groups), and covariation of different environmental drivers (e.g. water, temperature, atmospheric pressure), which do not permit a mechanistic interpretation. We solved both problems in a subcontinental survey of elevational trends in key plant traits in the European Alps and the high Arctic (northern Sweden, Svalbard), using herbarium samples of 147 species belonging to the genera Carex, Saxifraga and Potentilla. We used both species and phylogenetically independent contrasts as data points. The analysis revealed enhanced reproductive efforts at higher elevation in insect-pollinated taxa (not in wind-pollinated taxa), no increase in leaf pubescence at high elevation (as is often assumed), and a strong correlation between ¹³C discrimination and elevation. Alpine taxa operate at a smaller mesophyll resistance to CO₂ uptake relative to diffusive resistance (stomata). By comparison with congeneric low altitude polar taxa (low temperature, but high atmospheric pressure), the response could be attributed to the elevational decline in atmospheric pressure rather than temperature (a mean increase in δ¹³C by 1.4‰ km^?1). The signal is consistent within and across genera and within species, suggesting rapid adjustment of leaf physiology to reduced partial pressure of CO₂. These results offer answers to long-debated issues of plant responses to high elevation life conditions.     

西藏东达山3种嵩草属植物气孔特征沿海拔的变化

嵩草属(Kobresia)植物是藏东南高山草甸的优势种和建群种,对该区畜牧业发展和维持生态系统平衡起着重要作用。选择西藏左贡县东达山为研究地点,从林线开始,海拔每升高约100m设置1个样带直至高山草甸分布边缘,共8个样带,调查各样带中物种的组成及盖度,并依据相对盖度和相对频度计算3种嵩草植物矮生嵩草(K.humilis)、线叶嵩草(K.capillifolia)和大花嵩草(K.macrantha)在群落中的重要值,同时取样观察它们叶片远、近轴面表皮细胞形态,测量气孔长度及保卫细胞宽度,计算气孔密度,探讨嵩草属植物对海拔梯度的适应性。结果表明:(1)3种嵩草属植物叶表皮细胞均呈波浪状,气孔器仅分布于远轴面,近轴面无气孔器分布。(2)3种嵩草属植物气孔密度沿海拔梯度的变化均呈单峰曲线分布格局,且在海拔4 537m样带处达到最大值,并表现为矮生嵩草(777.6个/mm2)线叶嵩草(476.4个/mm2)大花嵩草(414.3个/mm2)。(3)随海拔的增加,矮生嵩草和线叶嵩草气孔长度显著增大(P0.05),而保卫细胞宽度显著减小;但大花嵩草气孔长度随海拔的升高而显著减小,保卫细胞宽度基本保持不变。(4)矮生嵩草和线叶嵩草气孔密度、长度和保卫细胞宽度与海拔梯度均显著相关,气孔特征对海拔梯度变化的敏感程度高,与其在群落中重要值高的分布特征一致;而大花嵩草仅气孔密度和长度与海拔梯度显著相关,气孔特征对海拔梯度变化的敏感性低,与其在群落中重要值低的分布特征一致;嵩草属植物气孔密度、长度和保护细胞宽度与海拔梯度之间的相关性,反映出它们在海拔梯度上对生境的适应程度。可见,3种嵩草属植物气孔特征对海拔梯度上生境变化的适应性不同,从而影响它们在群落中的分布范围和物种优势度,其中矮生嵩草和线叶嵩草对环境变化敏感,而大花嵩草对环境变化相对不敏感;保卫细胞宽度与气孔长度同样对植物适应环境变化起重要作用。     

Functional diversity and composition of bryophyte water-related traits in Azorean native vegetation

Background: Functional trait-based approaches link species diversity patterns to ecosystem functioning. In the context of global change, understanding these links is vital for developing holistic biodiversity management strategies. Bryophytes, important ecosystem components owing to their biogeochemical functions, have not been the focus of many functional studies.

Aims: This is the first assessment of bryophyte functional diversity in the Azores archipelago, aiming to uncover multivariate trait richness and composition patterns along the elevational gradient on Terceira Island.

Methods: Based on five water acquisition and retention traits of leafy liverworts and mosses, we calculated functional diversity metrics within and among six bryophyte communities sampled along a 1021-m elevational transect.

Results: Trait composition differed significantly between coastal and mountain communities. Mosses presented inrolled leaves and ornamented leaf cells at low elevation but not at high elevation. These patterns were associated with an uphill shift from drier and warmer conditions to a moister and cooler environment.

Conclusions: Future climatic changes might affect bryophyte functional diversity patterns in Terceira Island, particularly for mosses. These results can be directly compared with those obtained for other archipelagos where the same protocol has been applied, allowing a joint assessment of insular vegetation functional diversity patterns.     

宝天曼自然保护区两种优势藓类植物C、N、P生态化学计量特征研究

该研究以宝天曼自然保护区两种优势藓类物种大羽藓(Thuidium cymbifolium)和无边提灯藓(Mnium immarginatum)为对象,测定和分析了不同海拔生境条件下苔藓植物绿色组织和土壤的C、N、P含量及其比值。结果表明:该区域内两种藓类植物绿色组织的C、N、P含量范围是46.81%~49.09%、0.21%~0.25%、0.02%~0.08%;不同元素在海拔间存在较大差异,具体表现为大羽藓的C、N含量在不同海拔间差异显著,无边提灯藓的N、P含量在不同海拔梯度间差异显著,但总体仅N元素随海拔的上升表现出显著下降的趋势;土壤的C、N、N∶P和C∶P比值总体上表现出随海拔升高而增加的变化趋势,而土壤P含量以及C∶N比值随海拔梯度的变化并未表现出一致的变化趋势;同时,随着土壤C含量增加,大羽藓C含量呈现上升趋势,无边提灯藓呈现下降趋势;随着土壤N元素含量的增加,两种藓类物种表现的趋势仍然与前者一致;随着土壤P元素含量的升高,大羽藓的N∶P显著下降,而无边提灯藓的N∶P则显著上升;此外,叶片N∶P均值为8.51,介于14和16之间,说明植物生长受N元素的限制。以上结果说明:两种藓类植物对土壤P元素利用上存在差异,导致两种藓类化学计量学特征的变化,且与全球尺度相比,该区域的藓类植物的生长共同受到了土壤N含量的限制作用。该研究结果为预测藓类植物营养元素的限制情况和生态适应机制提供了参考。     

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.     

Natives and non‐natives plants show different responses to elevation and disturbance on the tropical high Andes of Ecuador

The aim was to assess patterns of plant diversity in response to elevation and disturbance in a tropical mountain. The study area was located in north‐central portion of the Eastern Cordillera of the Ecuadorian Andes, on a road from 1,150 m a.s.l. (Osayacu) to 4,000 (Papallacta). Along a mountain road spanning a wide altitudinal gradient, at 20 elevations we sampled three plots: one at the roadside and two perpendicular to the roadside. The relationship between elevation and species richness was assessed using linear and quadratic regressions, the effect of disturbance on species richness was determined by ANCOVA and a t test with parameters obtained from quadratic equations. Similarity of species composition among the roadside and sites distant was evaluated with the Chao‐Jaccard and classic Jaccard similarity indices, the distribution of non‐native species according to their origin were analyzed with linear and quadratic regression. The native species showed a linearly monotonic decrease with elevation, whereas non‐natives showed a quadratic distribution. Disturbed areas had the greatest number of non‐native species and lower native species richness, showing also a high floristic similarity; less disturbed areas showed the opposite. The non‐native species of temperate origin were more numerous and showed unimodal elevational distribution, while species of tropical origin were few and decreased linearly with elevation. We conclude that in a tropical highland mountain range, native and non‐native plant species respond differently to elevation: native species exhibit a monotonically linear decrease, and non‐native species show a unimodal trend. Disturbance positively affects non‐native species showing higher richness and fewer species turnover. In addition, the non‐native species are located along of the elevational gradient in relation to their biogeographic origin.     

The vertical distribution of pronophiline butterflies (Nymphalidae, Satyrinae) along an elevational transect in Monte Zerpa (Cordillera de Mérida, Venezuela) with remarks on their diversity and parapatric distribution

Aim The distribution of neotropical butterflies of the tribe Pronophilini (Lepidoptera, Satyridae) was analysed with the aim of determining (i) the elevational ranges of distribution of each species (ii) the elevational gradient in diversity, and (iii) the existence of parapatric distributions of some closely related species pairs. Location The field study was carried out in the middle and upper section of the valley of the Río Albarregas, a tributary of the Río Chama on the southern slopes of the Serranía de la Culata, in the central part of the Cordillera de Mérida, Venezuela. Methods The material was collected along an elevational transect set on a trail leading from Merida to the Páramo de Los Conejos in the area known as Monte Zerpa. It consisted of a series of 32 collection sites set at every 25 m of altitude and covered an elevation from 2250 m to 3025 m. Results The ranges of Lymanopoda obsoleta, L. albocincta, L. zapatoza, Corades chelonis, C. pannonia, C. medeba, Mygona irmina, Pedaliodes epidipnis, P. japhleta, P. montagna and P. panyasis were all restricted to the lower half of the cloud forest zone and L. diezti, C. pax, P. polla, P. ornata and P. ferratilis were all restricted to the upper part. Two species, Erethris porphyria and Steroma bega, crossed the entire elevational range of the cloud forest zone.The biodiversity, as measured by Shannon’s index, attained a maximum at 2700 m, beyond which it gradually decreased. A parapatric distribution was confirmed for three pairs of closely related species: L. obsoleta and L. diezti, C. chelonis and C. pax and P. montagna and P. ferratilis. Main conclusions The peak in diversity at middle elevations seems to result from the overlapping ranges of species distributed over lower and upper parts of the cloud forest. Correlation of species composition at particular transect sites with elevation showed that increase in elevation was associated with an increase in species composition difference. Post‐mating isolation was suggested as a primary factor responsible for maintaining the parapatric distributions of species occupying the upper and lower zones and preventing their respective distributions from expanding.     

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.     

The third dimension of bat migration: evidence for elevational movements of Miniopterus natalensis along the slopes of Mount Kilimanjaro

Bats are important ecosystem service providers, and therefore most relevant for both lowland and highland habitats, particularly in the tropics. Yet, it is poorly understood to what extent they perform large-scale movements, especially movements along mountain slopes. Here, we studied the movement ecology of the potentially migratory species Miniopterus natalensis at Mount Kilimanjaro in Tanzania. We analysed stable isotope ratios of C (δ¹³C), N (δ¹⁵N) and H (δ²H) in keratin of sedentary frugivorous and insectivorous bats captured between 800 and 2,400 m above sea level to establish elevational gradients of stable isotope ratios in consumer tissues. We expected correlations between stable isotope ratios of the non-exchangeable portion of H in fur keratin and the elevation of capture site, but not necessarily for δ¹³C and δ¹⁵N. Yet, in bats of both feeding ensembles, we found δ¹⁵N of fur keratin to correlate positively with the elevation of capture sites but not δ²H. In frugivorous bats, δ¹³C increased with increasing capture elevation as well. By looking at intra-individual variation of δ¹³C and δ¹⁵N in fur keratin and wing membrane tissues of sedentary Rhinolophus cf. clivosus and of the potentially migratory species M. natalensis, we gathered evidence that M. natalensis migrates seasonally between low and high elevations along the slopes of Mount Kilimanjaro. Finally, based on an isoscape origin model we estimated that M. natalensis captured before and after the cold period at around 1,800 m above sea level originated from around 1,400 m a.s.l. or lower. Thus, we received convergent results in support of seasonal elevational movements of M. natalensis, probably in search for cold hibernacula at higher elevations of Mount Kilimanjaro.     

Aquatic insect β‐diversity is not dependent on elevation in Southern Rocky Mountain streams

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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.     

Genetic variation in leaf pigment,optical and photosynthetic function among diverse phenotypes of <Emphasis Type="Italic">Metrosideros polymorpha</Emphasis> grown in a common garden

Coordinated variation has been reported for leaf structure, composition and function, across and within species, and theoretically should occur across populations of a species that span an extensive environmental range. We focused on Hawaiian keystone tree species Metrosideros polymorpha, specifically, 13-year old trees grown (2–4 m tall) in a common garden (approximately 1 ha field with 2–3 m between trees) from seeds collected from 14 populations along an altitude–soil age gradient. We determined the genetic component of relationships among specific leaf area (SLA), the concentrations of nitrogen (N) and pigments (chlorophylls, carotenoids, and anthocyanins), and photosynthetic light-use efficiency. These traits showed strong ecotypic variation; SLA declined 35% with increasing source elevation, and area-based concentrations of N, Chl a + b and Car increased by 50, 109 and 96%, respectively. Concentrations expressed on a mass basis were not well related to source elevation. Pigment ratios expressed covariation that suggested an increased capacity for light harvesting at higher source elevation; Chl/N and Car/Chl increased with source elevation, whereas Chl a/b declined; Chl a/b was higher for populations on younger soil, suggesting optimization for low N supply. Parallel trends were found for the photosynthetic reactions; light-saturated quantum yield of photosystem II (Φ _PSII) and electron transport rate (ETR) increased with source elevation. Correlations of the concentrations of photosynthetic pigments, pigment ratios, and photosynthetic function across the ecotypes indicated a stoichiometric coordination of the components of the light-harvesting antennae and reaction centers. The constellation of coordinated morphological, biochemical and physiological properties was expressed in the leaf reflectance and transmittance properties in the visible and near-infrared wavelength region (400–950 nm), providing an integrated metric of leaf status among and between plant phenotypes.     

Speciation in a keystone plant genus is driven by elevation: a case study in New Guinean Ficus

Much of the world's insect and plant biodiversity is found in tropical and subtropical ‘hotspots’, which often include long elevational gradients. These gradients may function as ‘diversity pumps’ and contribute to both regional and local species richness. Climactic conditions on such gradients often change rapidly along short vertical distances and may result in local adaptation and high levels of population genetic structure in plants and insects. We investigated the population genetic structure of two species of Ficus (Moraceae) along a continuously forested elevational gradient in Papua New Guinea. This speciose plant genus is pollinated by tiny, species‐specific and highly coevolved chalcid wasps (Agaonidae) and represented by at least 73 species at our study gradient. We present results from two species of Ficus sampled from six elevations between 200 m and 2700 m a.s.l. (almost the entire elevational range of the genus) and 10 polymorphic microsatellite loci. These results show that strong barriers to gene flow exist between 1200 m and 1700 m a.s.l. Whereas lowland populations are panmictic across distances over 70 km, montane populations can be disjunct over 4 km, despite continuous forest cover. We suggest that the limited gene flow between populations of these two species of montane Ficus may be driven by environmental limitations on pollinator or seed dispersal in combination with local adaptation of Ficus populations. Such a mechanism may have wider implications for plant and pollinator speciation across long and continuously forested elevational gradients if generalist insect pollinators and vertebrate seed dispersers also form populations based on elevation.     

Fine‐scale determinants of butterfly species richness and composition in a mountain region

Aim Global patterns of species richness are often considered to depend primarily on climate. We aimed to determine how topography and land cover affect species richness and composition at finer scales. Location Sierra de Guadarrama (central Iberian Peninsula). Methods We sampled the butterfly fauna of 180 locations (89 in 2004, 91 in 2005) at 600–2300 m elevation in a region of 10800 km². We recorded environmental variables at 100‐m resolution using GIS, and derived generalized linear models for species density (number of species per unit area) and expected richness (number of species standardized to number of individuals) based on variables of topoclimate (elevation and insolation) or land cover (vegetation type, geology and hydrology), or both (combined). We evaluated the models against independent data from the alternative study year. We also tested for differences in species composition among sites and years using constrained ordination (canonical correspondence analysis), and used variation partitioning analyses to quantify the independent and combined roles of topoclimate and land cover. Results Topoclimatic, land cover and combined models were significantly related to observed species density and expected richness. Topoclimatic and combined models outperformed models based on land cover variables, showing a humped elevational diversity gradient. Both topoclimate and land cover made significant contributions to models of species composition. Main conclusions Topoclimatic factors may dominate species richness patterns in regions with pronounced elevational gradients, as long as large areas of natural habitat remain. In contrast, both topoclimate and land cover may have important effects on species composition. Biodiversity conservation in mountainous regions therefore requires protection and management of natural habitats over a wide range of topoclimatic conditions, which may assist in facilitating range shifts and alleviating declines in species richness related to climate change.     

Tree growth response to drought and temperature in a mountain landscape in northern Arizona, USA

Aim To understand how tree growth response to regional drought and temperature varies between tree species, elevations and forest types in a mountain landscape. Location Twenty‐one sites on an elevation gradient of 1500 m on the San Francisco Peaks, northern Arizona, USA. Methods Tree‐ring data for the years 1950–2000 for eight tree species (Abies lasiocarpa var. arizonica (Merriam) Lemm., Picea engelmannii Parry ex Engelm., Pinus aristata Engelm., Pinus edulis Engelm., Pinus flexilis James, Pinus ponderosa Dougl. ex Laws., Pseudotsuga menziesii var. glauca (Beissn.) Franco and Quercus gambelii Nutt.) were used to compare sensitivity of radial growth to regional drought and temperature among co‐occurring species at the same site, and between sites that differed in elevation and species composition. Results For Picea engelmannii, Pinus flexilis, Pinus ponderosa and Pseudotsuga menziesii, trees in drier, low‐elevation stands generally had greater sensitivity of radial growth to regional drought than trees of the same species in wetter, high‐elevation stands. Species low in their elevational range had greater drought sensitivity than co‐occurring species high in their elevational range at the pinyon‐juniper/ponderosa pine forest ecotone, ponderosa pine/mixed conifer forest ecotone and high‐elevation invaded meadows, but not at the mixed conifer/subalpine forest ecotone. Sensitivity of radial growth to regional drought was greater at drier, low‐elevation compared with wetter, high‐elevation forests. Yearly growth was positively correlated with measures of regional water availability at all sites, except high‐elevation invaded meadows where growth was weakly correlated with all climatic factors. Yearly growth in high‐elevation forests up to 3300 m a.s.l. was more strongly correlated with water availability than temperature. Main conclusions Severe regional drought reduced growth of all dominant tree species over a gradient of precipitation and temperature represented by a 1500‐m change in elevation, but response to drought varied between species and stands. Growth was reduced the most in drier, low‐elevation forests and in species growing low in their elevational range in ecotones, and the least for trees that had recently invaded high‐elevation meadows. Constraints on tree growth from drought and high temperature are important for high‐elevation subalpine forests located near the southern‐most range of the dominant species.     

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.     

Patterns and drivers of intraspecific variation in avian life history along elevational gradients: a meta‐analysis

Elevational gradients provide powerful natural systems for testing hypotheses regarding the role of environmental variation in the evolution of life‐history strategies. Case studies have revealed shifts towards slower life histories in organisms living at high elevations yet no synthetic analyses exist of elevational variation in life‐history traits for major vertebrate clades. We examined (i) how life‐history traits change with elevation in paired populations of bird species worldwide, and (ii) which biotic and abiotic factors drive elevational shifts in life history. Using three analytical methods, we found that fecundity declined at higher elevations due to smaller clutches and fewer reproductive attempts per year. By contrast, elevational differences in traits associated with parental investment or survival varied among studies. High‐elevation populations had shorter and later breeding seasons, but longer developmental periods implying that temporal constraints contribute to reduced fecundity. Analyses of clutch size data, the trait for which we had the largest number of population comparisons, indicated no evidence that phylogenetic history constrained species‐level plasticity in trait variation associated with elevational gradients. The magnitude of elevational shifts in life‐history traits were largely unrelated to geographic (altitude, latitude), intrinsic (body mass, migratory status), or habitat covariates. Meta‐population structure, methodological issues associated with estimating survival, or processes shaping range boundaries could potentially explain the nature of elevational shifts in life‐history traits evident in this data set. We identify a new risk factor for montane populations in changing climates: low fecundity will result in lower reproductive potential to recover from perturbations, especially as fewer than half of the species experienced higher survival at higher elevations.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.     

Ecological interactions among insect herbivores,ants and the host plant Baccharis dracunculifolia in a Brazilian mountain ecosystem

Insect–plant interactions occur in several ways and have considerable environmental and ecological importance. Many feeding strategies have evolved among herbivorous insects, with host–herbivore systems likely being influenced by trophobionts with ants. We investigated how these interactions vary across elevation gradients by evaluating the structure of the herbivorous insect community and ants associated with Baccharis dracunculifolia at three distinct elevations (800, 1100, and 1400 m a.s.l.) on a mountain in southeastern Brazil. Moreover, we evaluated the diversity and specialisation of interactions between herbivores and host plants along the elevational gradient. We sampled herbivores and ants on 60 plants at each elevation (totalling 180 plant individuals). Herbivore species composition differed among elevations, as did interaction diversity and specialisation. Richness and abundance of chewing insects increased with elevation, while β‐diversity among patches of the host plant was higher at the lowest elevation, probably due to the patchy occurrence of B. dracunculifolia. Richness and abundance of sap‐sucking insects were higher at the intermediate elevation, possibly due to local environmental conditions. We observed a positive relationship between ant and herbivore trophobiont richness on B. dracunculifolia. We found that interactions were more specialised and less diverse at higher elevations compared to the lowest elevation. Changes in vegetation and environmental variables shaped species distributions and their ecological interactions along the elevation gradient. Our study demonstrates that increased elevation changes the structure and patterns of interactions of the herbivore insect guilds associated with the host plant B. dracunculifolia. Ant effects depend on the context, the environment, and the species of ants involved, and are essential for the presence of insect trophobionts.