Can Rapoport's rule explain tree species richness along the Himalayan elevation gradient,Nepal?

Rapoport's rule applied to an elevation gradient predicts a positive correlation between elevation ranges and elevation. This is supposed to be caused by the increasing magnitude of the climatic extremes at higher elevations, and thus, it is deduced that species richness should decrease with increasing elevation. The distribution of 614 tree species was used to test Rapoport's elevational rule along a gradient from 100 to 4300 m a.s.l., in the Nepalese Himalaya. The relationship between species richness and elevation was analysed by using generalized linear models (GLM). Generalized additive models (GAM) were used to examine the relationship between elevational range and the elevational mid-point of a species along the gradient. The widest elevation ranges are observed at mid-elevations, and narrow elevation ranges are observed at both ends of the gradient. This does not support Rapoport's elevation rule, as proposed by Stevens. There is a peak in species richness between 900 and 1000 m, and not in the tropical lowland as projected by Rapoport's elevation rule.     

Fern species richness along a central Himalayan elevational gradient, Nepal

Aim The study explores fern species richness patterns along a central Himalayan elevational gradient (100–4800 m a.s.l.) and evaluates factors influencing the spatial increase and decrease of fern richness. Location The Himalayas stretch from west to east by 20°, i.e. 75–95° east, and Nepal is located from 80 to 88° east in this range. Methods We used published data of the distribution of ferns and fern allies to interpolate species elevational ranges. Defining species presence between upper and lower elevation limit is the basis for richness estimates. The richness pattern was regressed against the total number of rainy days, and gradients that are linearly related to elevation, such as length of the growing season, potential evapotranspiration (PET, energy), and a moisture index (MI = PET/mean annual rainfall). The regressions were performed by generalized linear models. Results A unimodal relationship between species richness and elevation was observed, with maximum species richness at 2000 m. Fern richness has a unimodal response along the energy gradients, and a linear response with moisture gradients. Main conclusions The study confirms the importance of moisture on fern distributions as the peak coincides spatially with climatic factors that enhance moisture levels; the maximum number of rainy days and the cloud zone. Energy‐related variables probably control species richness directly at higher elevations but at the lower end the effect is more probably related to moisture.     

The microbial gene diversity along an elevation gradient of the Tibetan grassland

Tibet is one of the most threatened regions by climate warming, thus understanding how its microbial communities function may be of high importance for predicting microbial responses to climate changes. Here, we report a study to profile soil microbial structural genes, which infers functional roles of microbial communities, along four sites/elevations of a Tibetan mountainous grassland, aiming to explore the potential microbial responses to climate changes via a strategy of space-for-time substitution. Using a microarray-based metagenomics tool named GeoChip 4.0, we showed that microbial communities were distinct for most but not all of the sites. Substantial variations were apparent in stress, N and C-cycling genes, but they were in line with the functional roles of these genes. Cold shock genes were more abundant at higher elevations. Also, gdh converting ammonium into urea was more abundant at higher elevations, whereas ureC converting urea into ammonium was less abundant, which was consistent with soil ammonium contents. Significant correlations were observed between N-cycling genes (ureC, gdh and amoA) and nitrous oxide flux, suggesting that they contributed to community metabolism. Lastly, we found by Canonical correspondence analysis, Mantel tests and the similarity tests that soil pH, temperature, NH₄⁺–N and vegetation diversity accounted for the majority (81.4%) of microbial community variations, suggesting that these four attributes were major factors affecting soil microbial communities. On the basis of these observations, we predict that climate changes in the Tibetan grasslands are very likely to change soil microbial community functional structure, with particular impacts on microbial N-cycling genes and consequently microbe-mediated soil N dynamics.     

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)。     

Variation in plant species richness of different life forms along a subtropical elevation gradient in the Himalayas, east Nepal

Aim To explore the variation in species richness along a subtropical elevation gradient, and evaluate how climatic variables explain the richness of the different life forms such as trees, shrubs, climbers, herbs and ferns. Location The study was made in a subtropical to warm temperate region in the south‐eastern part of Nepal, between 100 and 1500 m above sea level (a.s.l.). Methods The number of species was counted in six plots (50 × 20 m) in each of the 15 100 m elevation bands covering the main physiognomic structures along an imaginary transect. Each species recorded was assigned to a life form. Potential evapotranspiration (PET, i.e. energy), mean annual rainfall (MAR), and their ratio (MI = moisture index) were evaluated as explanatory variables by means of generalized linear models (GLM). Each variable was tested individually, and in addition MAR and PET were used to test the water‐energy dynamics model for each life form. Results The richness of herbaceous species, including herbaceous climbers, was unrelated to any of the climate variables. PET was strongly negatively correlated with elevation, and the following relationships were found between increasing PET and richness: (i) shrubs, trees and total species (sum of all life forms) showed unimodal responses (ii) ferns decreased monotonically, and (iii) woody climbers increased monotonically. Richness of all woody groups increased monotonically with MAR and MI. The water‐energy dynamics model explained 63% of the variation in shrubs, 67% for trees and 70% for woody species combined. Main conclusions For the various herbaceous life forms (forbs, grasses, and herbaceous climbers) we found no significant statistical trends, whereas for woody life forms (trees, shrubs, and woody climbers) significant relationships were found with climate. E.M. O’Brien's macro‐scale model based on water‐energy dynamics was found to explain woody species richness at a finer scale along this elevational‐climatic gradient.     

贺兰山不同海拔典型植被带土壤微生物多样性

土壤微生物多样性在海拔梯度的分布格局研究近年来受到和植物动物一样的重视程度,但是干旱风沙区微生物多样性在海拔梯度上的多样性分布规律尚未揭示。本研究以处于干旱风沙区的贺兰山不同海拔的六个典型植被带（荒漠草原带、山地旱生灌丛带、温性针叶林带、针阔混交林带、寒温性针叶林带和亚高山草甸带）土壤为研究对象,利用Biolog微平板法和磷脂脂肪酸甲酯法(FAMEs)系统研究微生物多样性群落特征以及在不同植被带分布规律。结果表明：土壤微生物功能多样性随海拔增加发生变化,且微生物群落结构存在显著差异。Biolog分析显示土壤微生物群落代谢活性依次是：亚高山草甸>寒温性针叶林>针阔混交林>温性针叶林>山地旱生灌丛>荒漠草原,随海拔的升高土壤微生物群落物种丰富度指数（H）和均匀度指数（E）总体上均表现出增大的趋势,差异显著（P＜0.05）;FAMEs分析表明不同海拔的微生物区系发生了一定程度的变化,寒温性针叶林土壤微生物磷酸脂肪酸生物标记的数量和种类均最高,且细菌、真菌特征脂肪酸相对含量也最高;土壤微生物群落结构多样性次序是：寒温性针叶林带>针阔混交林带>温性针叶林带>亚高山草甸>山地旱生灌丛>荒漠草原。本研究结果表明贺兰山海拔梯度的微生物多样性分布规律不同于已有的植物多样性“中部膨胀”研究结果,这说明在高海拔地区有更多的适合该生境的微生物存在,这对维持干旱风沙区的生态系统功能稳定性具有重要意义。     

小秦岭森林群落数量分类、排序及多样性垂直格局

采用分层取样的方法,沿小秦岭林区海拔梯度设立56块20 m×20 m样地,用多元回归树(MRT)方法对小秦岭森林群落进行分类,采用除趋势对应分析(DCA)进行排序,用广义可加模型(GAM)研究不同生活型物种多样性沿海拔梯度分布格局。结果表明:(1)56个样地进行MRT分类,经交叉验证并依据植物群落分类和命名原则,本区植物群落可分为5类;(2)样方DCA排序明确地揭示各群落类型生境分布范围,较好地反映小秦岭自然保护区森林群落与环境因子的关系;(3)不同生活型物种多样性指数随海拔梯度变化发生一定的波动,且呈现不同的多样性格局:丰富度指数中,乔木层呈显著的单峰分布格局,灌木层在中海拔段呈明显下降趋势,草本层随着海拔的升高总体呈下降趋势;Shannon-Wiener多样性指数中,不同生活型物种随海拔变化趋势与物种丰富度变化趋势大体相同;不同生活型物种的均匀度指数随海拔变化趋势较平缓。     

环境梯度下蒙古栎群落的物种多样性特征

通过样地法研究了东北地区处于不同经度、纬度和海拔的 13个地点蒙古栎群落的物种丰富度、Gini指数、PIE指数、Shannon指数和 Pielou指数 ,利用相关和回归的统计方法分析了不同地点物种的丰富度指数、Simpson多样性指数和 Shannon多样性指数与各地所处的经度、纬度和海拔的关系。结果发现 :不仅不同地点 (较大尺度 )的物种丰富度和多样性指数均有差异 ,即使在相同的地点 (较小尺度 ) ,物种丰富度及多样性指数也有差异 ,有时还具有很大的差异 ,呈现空间异质性分布的特征 ;因为影响这些多样性指数的环境因子更加复杂 ,不仅受经度、纬度和海拔的影响 ,也受地形、群落的年龄、干扰史等多种生态因子影响。不同地点的物种丰富度与海拔和纬度都具有明显的相关性 (p<0 .0 5 ) ,物种丰富度随海拔和纬度的升高而降低 ,依据显著度的大小可以推测物种丰富度与海拔的相关性比与纬度的相关性更密切 ;蒙古栎群落不同类群的植物种的丰富度具有不同的分布格局 ,木本植物的丰富度与当地纬度具有明显的相关性 (p<0 .0 5 ) ,而与所在地的海拔没有显著的关系 (p>0 .0 5 ) ,而草本植物受海拔的影响更显著 (p<0 .0 5 ) ,而与纬度之间没有显著的关系 (p>0 .0 5 )。群落的 Gini指数、PIE指数、Shannon指数和Pielou指数未发现与海     

Mammalian species richness and morphological complexity along an elevational gradient in the arid south-west

Aim We examined the relationship between species richness and morphological complexity of terrestrial mammal communities along an elevational gradient. Location The gradient is in the Sonoran Desert in Southern California and extends from a sand dune habitat near sea level to coniferous forest ending at >2600 m. Methods Morphological diversity, characterized by both size and shape of coexisting mammal species, was estimated within and between sites from projections of variables on principal components axes. Similarities among species were calculated as Euclidean distances. To tease apart size and shape, we constructed two principal component analyses: one based on log-transformed original measurements, the other on log-transformed proportional shape variables. To test whether species number accounted for the morphological diversity at each site we designed two null models. The models generated were random communities generated from the forty-two-species pool. Indices of morphological diversity for real communities were compared with the results of 500 simulations of each null model. Results Species richness varied along the gradient, peaking in the mid-elevation agave-ocotillo habitat. Morphological diversity of shapes and sizes correlated strongly with species richness. Locomotor, tooth, and skull traits were all important in distinguishing among species. Main conclusions Two important patterns emerged: (1) diversity of both sizes and of shapes of species within communities correlated positively with species number, and both sets of variables behaved similarly across this gradient; (2) the most species rich sites were not composed of specialists on these best places, but rather, a community of species derived from overlapping faunal groups.     

Genetic and phenotypic variation,dispersal limitation and reproductive success in the invasive herb Eschscholzia californica along an elevation gradient in central Chile

Background: Reductions of genetic diversity and phenotypic changes in invasive plants are often observed to occur at high elevations. Genetic/phenotypic changes of invasive plants along elevation help to understand mechanisms of the presumed resistance of mountain ecosystems to invasion.

Aims: To assess genetic variability and phenotypic plasticity along an elevation gradient of Eschscholzia californica in the Andes, central Chile.

Methods: Eleven microsatellites were used to describe the genetic structure and the allelic diversity individuals, distributed at three elevations and two sites. We assessed the number of flowers per plant, floral biomass, leaf area, number of leaves, vegetative biomass and plant height of plants at each elevation.

Results: Genetic diversity as genetic structure did not decrease with elevations. Plant height and flower numbers decreased while leaf number and vegetative biomass increased with elevation. The ratio of the number of flowers to vegetative biomass, decreased significantly with elevation.

Conclusions: Strong genetic differences among elevations and similar genetic diversity along elevation do not suggest dispersal limitation to higher elevation. Reduction of reproductive and vegetative traits concomitantly with an increase of the reproductive cost suggests reproductive stress with increasing elevation, reducing the invasiveness of this species to higher elevation.     

Distribution of vascular plant species richness and endemic richness along the Himalayan elevation gradient in Nepal

Aim Species richness and endemic richness vary along elevation gradients, but not necessarily in the same way. This study tests if the maxima in gamma diversity for flowering plants and the endemic subset of these plants are coherent or not. Location The study was conducted in Nepal, between 1000 and 5000 m a.s.l. Methods We used published data on distribution and elevational ranges of the Nepalese flora to interpolate presence between maximum and minimum elevations. Correlation, regression and graphical analyses were used to evaluate the diversity pattern between 1000 and 5000 m a.s.l. Results The interval of maximum species endemic to Nepal or the Himalayas (3800–4200 m) is above the interval of maximum richness (1500–2500 m). The exact location of maximum species density is uncertain and its accuracy depends on ecologically sound estimates of area in the elevation zones. There is no positive statistically significant correlation between log‐area and richness (total or endemic). Total richness is positively correlated with log‐area‐adjusted, i.e. estimated area adjusted for the degree of topographic heterogeneity. The proportion of endemic species increases steadily from low to high elevations. The peak in endemism (c. 4000 m) corresponds to the start of a rapid decrease in species richness above 4000 m. This may relate to the last glacial maximum (equilibrium line at c. 4000 m) that penetrated down to 2500–3000 m. This dynamic hard boundary may have caused an increase in the extinction rate above 4000 m, and enhanced the probability of isolation and facilitated speciation of neoendemics, especially among genera with a high proportion of polyploids. Main conclusions The results reject the idea of corresponding maxima in endemic species and species richness in the lowlands tentatively deduced from Stevens’ elevational Rapoport effect. They confirm predictions based on hard boundary theory, but hard‐boundaries should be viewed as dynamic rather than static when broad‐scale biogeographical patterns with a historical component are being interpreted.     

Epiphytic bryophyte diversity and range distributions along an elevational gradient in Marojejy,Madagascar

Describing spatial variation in species richness and understanding its links to ecological mechanisms are complementary approaches for explaining geographical patterns of richness. The study of elevational gradients holds enormous potential for understanding the factors underlying global diversity. This paper investigates the pattern of species richness and range-size distribution of epiphytic bryophytes along an elevational gradient in Marojejy National Park, northeast Madagascar. The main objectives are to describe bryophyte species composition and endemism in Marojejy National Park, to describe the species richness and distribution patterns of epiphytic bryophytes along an elevational gradient from 250 m to 2050 m and to evaluate the explanatory value of environmental variables for the observed patterns. Bryophyte samples were collected following a nested design with four hierarchical levels: elevational belts, plots, quadrats, and microplots. In total, 254 epiphytic bryophyte species were recorded, comprising 157 liverworts and 97 mosses. Twenty-three of these are endemic to Madagascar. Species richness exhibits a hump-shaped pattern along the elevational gradient, peaking at 1,250 m. Eighty-seven percent of the total recorded species have a range distribution lower than 1,000 m, at which point 36% are restricted to these single elevations. Our results suggest that mean temperature, relative humidity, and vapor pressure deficit play important roles in shaping the richness pattern observed in this study. While the liverwort richness pattern did not correlate to vapor pressure deficit and responded only weakly to relative humidity, the richness pattern shown by mosses correlates well with mean temperature, relative humidity, and vapor pressure deficit.     

羌塘高寒草地物种多样性与生态系统多功能关系格局

传统的生物多样性-生态系统功能研究大多侧重于单一生态系统功能与物种多样性的关系,忽略了生态系统的重要价值在于其能够同时提供多种功能或服务,即生态系统的多功能性。基于藏北羌塘高寒草地样带调查数据,选取植被地上生物量、地下生物量、土壤全氮、硝态氮及铵态氮含量、土壤全磷含量、土壤有机碳储量等7个与植物生长、养分循环、土壤有机碳蓄积相关的参数来表征生态系统多功能性。采用上述参数转换为Z分数后的平均值计算多功能性指数(M)。分析了不同生物多样性指数与生态系统多功能指数的关系以及年降水量和年均温度对物种多样性和生态系统多功能性指数的影响。结果表明,物种丰富度指数与生态系统多功能性之间呈极显著的正相关关系,Shannon-wiener和Simpson物种多样性指数也与多功能性指数间呈显著的正相关,但多功能性指数与Pielou均匀度指数没有表现出明显的相关关系。物种丰富度与表征植物生长、养分循环以及土壤有机碳蓄积的生态系统功能指数间也均呈极显著的正相关关系。降水格局显著影响羌塘高原物种丰富度和生态系统多功能指数,二者均随年降雨量的增加而显著增加,但物种多样性指数并未与年降水量呈现显著相关关系。研究强调了群落物种丰富度即群落物种数量对维持生态系统多功能性的重要意义,这意味着由于人类活动导致的物种丧失可能会给藏北高寒草地生态系统多功能和生态服务带来更为严重的后果。就退化草地恢复或草地可持续管理而言,在藏北羌塘地区,本地植物种的物种丰富度恢复和维持应作为重要目标之一。     

Non-native species distribution along the elevation gradient in the western Italian Alps

Abstract

In this work the occurrence of non-native species was recorded along the elevation gradient in the Alps, in order to establish their distribution pattern, their current altitudinal limits and to elucidate which species are presumably adapted to higher elevations. Plots were located along the course of rivers in five valleys from 100 to 2100 m a.s.l. Sixty-eight non-natives were recorded in the study area. The proportion of invasives was found to be much higher in the study area then in the whole administrative region. The number of non-natives per plot decreased strongly with increasing elevation. The great majority (94%) of the non-native species grows at the lowest elevations, while only 6% survive up to 1500 m and none was found above this limit. Results were interpreted considering the factors driving the invasion process (disturbance, the native communities' resistance to invasion, propagule pressure, climatic conditions) and plant traits with particular respect to pre-adaptation to the harsh climate, which increases progressively with elevation. Results confirm that the Alps are not immune from invasion, at least up to medium elevation.     

Relationships between alpha diversity of plant species in bloom and climatic variables across an elevation gradient

This study analyzes a 20-year record of flowering observations collected near Tucson, Arizona, USA. In contrast to traditional phenological records, this dataset is a record of all species observed in bloom collected in five segments of approximately 1 mile (1.61 km) in length across a 4,158-ft (1,200-m) elevation gradient. The data showed differing seasonal and interannual patterns, demonstrating the influence of climatic factors and elevation on flowering. Miles at higher elevations showed bloom peaks in summer, consistent with temperate and montane communities. Conversely, lower miles demonstrated two distinct flowering seasons, typical of the surrounding Sonoran Desert. Interannual fluctuations in total species observed in bloom were not consistent across the 5 miles (c. 8 km), suggesting that these communities respond to different flowering cues. Consistent with documented flowering triggers in semi-arid systems, the alpha diversity of species in bloom at lower elevations in this study was strongly influenced by precipitation. Upper elevation bloom numbers were heavily influenced by temperature, correspondent with bloom triggers in temperate and montane systems. In general, different life forms exhibited similar bloom triggers within the study miles, believed to be a function of shallow soils. Multivariate community analyses showed that anomalous climate conditions yielded unique seasonal bloom compositions. Over the course of the study, average summer temperature showed an upward trend; the number of species in bloom in summer (July–October) in the highest mile (1,940–2,210 m) demonstrated a concurrent increasing trend. Community analysis suggested a gradual shift in the composition of species in bloom in this mile over the study period.     

The mid-domain effect in ectomycorrhizal fungi: range overlap along an elevation gradient on Mount Fuji,Japan

Mid-domain effect (MDE) models predict that the random placement of species'' ranges within a bounded geographical area leads to increased range overlap and species richness in the center of the bounded area. These models are frequently applied to study species-richness patterns of macroorganisms, but the MDE in relation to microorganisms is poorly understood. In this study, we examined the characteristics of the MDE in richness patterns of ectomycorrhizal (EM) fungi, an ecologically important group of soil symbionts. We conducted intensive soil sampling to investigate overlap among species ranges and the applicability of the MDE to EM fungi in four temperate forest stands along an elevation gradient on Mount Fuji, Japan. Molecular analyses using direct sequencing revealed 302 EM fungal species. Of 73 EM fungal species found in multiple stands, 72 inhabited a continuous range along the elevation gradient. The maximum overlap in species range and the highest species richness occurred at elevations in the middle of the gradient. The observed richness pattern also fit within the 95% confidence interval of the mid-domain null model, supporting the role of the MDE in EM fungal richness. Deviation in observed richness from the mean of the mid-domain null estimation was negatively correlated with some environmental factors, including precipitation and soil C/N, indicating that unexplained richness patterns could be driven by these environmental factors. Our results clearly support the existence of microbial species'' ranges along environmental gradients and the potential applicability of the MDE to better understand microbial diversity patterns.