Elevational gradients in bird diversity in the Eastern Himalaya: an evaluation of distribution patterns and their underlying mechanisms

Background

Understanding diversity patterns and the mechanisms underlying those patterns along elevational gradients is critically important for conservation efforts in montane ecosystems, especially those that are biodiversity hotspots. Despite recent advances, consensus on the underlying causes, or even the relative influence of a suite of factors on elevational diversity patterns has remained elusive.

Methods and Principal Findings

We examined patterns of species richness, density and range size distribution of birds, and the suite of biotic and abiotic factors (primary productivity, habitat variables, climatic factors and geometric constraints) that governs diversity along a 4500-m elevational gradient in the Eastern Himalayan region, a biodiversity hotspot within the world''s tallest mountains. We used point count methods for sampling birds and quadrats for estimating vegetation at 22 sites along the elevational gradient. We found that species richness increased to approximately 2000 m, then declined. We found no evidence that geometric constraints influenced this pattern, whereas actual evapotranspiration (a surrogate for primary productivity) and various habitat variables (plant species richness, shrub density and basal area of trees) accounted for most of the variation in bird species richness. We also observed that ranges of most bird species were narrow along the elevation gradient. We find little evidence to support Rapoport''s rule for the birds of Sikkim region of the Himalaya.

Conclusions and Significance

This study in the Eastern Himalaya indicates that species richness of birds is highest at intermediate elevations along one of the most extensive elevational gradients ever examined. Additionally, primary productivity and factors associated with habitat accounted for most of the variation in avian species richness. The diversity peak at intermediate elevations and the narrow elevational ranges of most species suggest important conservation implications: not only should mid-elevation areas be conserved, but the entire gradient requires equal conservation attention.     

Nonlinear Distribution Pattern of Hibernating Bats in Caves along an Elevational Gradient in Mountain (Carpathians,Southern Poland)

Background

Thermal gradients along changes in elevation in mountainous environments are reflected by different biotas. Although there have been studies of elevation variation in bat assemblages in summer, winter changes in the same gradients remain unknown.

Methodology/Principal Findings

The objective of this study was to document changes in the species composition of bats hibernating in caves along a temperate elevational gradient. We studied 70 caves between from 300 m to 1,930 m altitude along a slope of the Carpathian Mountains in southern Poland. We recorded changes in bats, including species richness, abundance, altitudinal distribution and dominance during consecutive winters between 2003 and 2009. Similarity of dominance of faunal structure was assessed by using the Bray-Curtis similarity index. We used the generalised additive model and rarefaction to study the variation in species richness, and generalized additive mixed models to examine the effect of abiotic factors on the qualitative and quantitative structure of bat assemblages. During 351 surveys we recorded 13,856 hibernating bats from 15 species. Species richness peaked around mid-elevation (1,100–1,400 m a.s.l.) with richness declining at both higher and lower elevations. Based on the results of a cluster analysis, we could distinguish among four altitudinal zones that differed in species richness and dominance structure.

Conclusions/Significance

This is the first study documenting changes in species richness and variation of structure of bats hibernating in caves along an elevational gradient. The most surprising and key finding is the fact that changes in the structure of assemblages of hibernating bats along the altitudinal gradient occurred in jumps, forming zones similar to those observed in the vegetation zones. Moreover, species richness and dominance structure of assemblages of hibernating bats in the mountains depended not only on location above sea level, but also on local geomorphologic conditions which strongly affected the microclimate of the caves.     

Elevational plant species richness patterns and their drivers across non-endemics,endemics and growth forms in the Eastern Himalaya

Despite decades of research, ecologists continue to debate how spatial patterns of species richness arise across elevational gradients on the Earth. The equivocal results of these studies could emanate from variations in study design, sampling effort and data analysis. In this study, we demonstrate that the richness patterns of 2,781 (2,197 non-endemic and 584 endemic) angiosperm species along an elevational gradient of 300–5,300 m in the Eastern Himalaya are hump-shaped, spatial scale of extent (the proportion of elevational gradient studied) dependent and growth form specific. Endemics peaked at higher elevations than non-endemics across all growth forms (trees, shrubs, climbers, and herbs). Richness patterns were influenced by the proportional representation of the largest physiognomic group (herbs). We show that with increasing spatial scale of extent, the richness patterns change from a monotonic to a hump-shaped pattern and richness maxima shift toward higher elevations across all growth forms. Our investigations revealed that the combination of ambient energy (air temperature, solar radiation, and potential evapo-transpiration) and water availability (soil water content and precipitation) were the main drivers of elevational plant species richness patterns in the Himalaya. This study highlights the importance of factoring in endemism, growth forms, and spatial scale when investigating elevational gradients of plant species distributions and advances our understanding of how macroecological patterns arise.     

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

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

Different responses of avian feeding guilds to spatial and environmental factors across an elevation gradient in the central Himalaya

Although elevational patterns of species richness have been well documented, how the drivers of richness gradients vary across ecological guilds has rarely been reported. Here, we examined the effects of spatial factors (area and mid‐domain effect; MDE) and environmental factors, including metrics of climate, productivity, and plant species richness on the richness of breeding birds across different ecological guilds defined by diet and foraging strategy. We surveyed 12 elevation bands at intervals of 300 m between 1,800 and 5,400 m a.s.l using line‐transect methods throughout the wet season in the central Himalaya, China. Multiple regression models and hierarchical partitioning were used to assess the relative importance of spatial and environmental factors on overall bird richness and guild richness (i.e., the richness of species within each guild). Our results showed that richness for all birds and most guilds displayed hump‐shaped elevational trends, which peaked at an elevation of 3,300–3,600 m, although richness of ground‐feeding birds peaked at a higher elevation band (4,200–4,500 m). The Normalized Difference Vegetation Index (NDVI)—an index of primary productivity—and habitat heterogeneity were important factors in explaining overall bird richness as well as that of insectivores and omnivores, with geometric constraints (i.e., the MDE) of secondary importance. Granivore richness was not related to primary production but rather to open habitats (granivores were negatively influenced by habitat heterogeneity), where seeds might be abundant. Our findings provide direct evidence that the richness–environment relationship is often guild‐specific. Taken together, our study highlights the importance of considering how the effects of environmental and spatial factors on patterns of species richness may differ across ecological guilds, potentially leading to a deeper understanding of elevational diversity gradients and their implications for biodiversity conservation.     

Processes at multiple scales affect richness and similarity of non‐native plant species in mountains around the world

Aim To investigate how species richness and similarity of non‐native plants varies along gradients of elevation and human disturbance. Location Eight mountain regions on four continents and two oceanic islands. Methods We compared the distribution of non‐native plant species along roads in eight mountainous regions. Within each region, abundance of plant species was recorded at 41–84 sites along elevational gradients using 100‐m² plots located 0, 25 and 75 m from roadsides. We used mixed‐effects models to examine how local variation in species richness and similarity were affected by processes at three scales: among regions (global), along elevational gradients (regional) and with distance from the road (local). We used model selection and information criteria to choose best‐fit models of species richness along elevational gradients. We performed a hierarchical clustering of similarity to investigate human‐related factors and environmental filtering as potential drivers at the global scale. Results Species richness and similarity of non‐native plant species along elevational gradients were strongly influenced by factors operating at scales ranging from 100 m to 1000s of km. Non‐native species richness was highest in the New World regions, reflecting the effects of colonization from Europe. Similarity among regions was low and due mainly to certain Eurasian species, mostly native to temperate Europe, occurring in all New World regions. Elevation and distance from the road explained little of the variation in similarity. The elevational distribution of non‐native species richness varied, but was always greatest in the lower third of the range. In all regions, non‐native species richness declined away from roadsides. In three regions, this decline was steeper at higher elevations, and there was an interaction between distance and elevation. Main conclusions Because non‐native plant species are affected by processes operating at global, regional and local scales, a multi‐scale perspective is needed to understand their patterns of distribution. The processes involved include global dispersal, filtering along elevational gradients and differential establishment with distance from roadsides.     

Spatiotemporal maintenance of flora in the Himalaya biodiversity hotspot: Current knowledge and future perspectives

Mountain ecosystems support a significant one‐third of all terrestrial biodiversity, but our understanding of the spatiotemporal maintenance of this high biodiversity remains poor, or at best controversial. The Himalaya hosts a complex mountain ecosystem with high topographic and climatic heterogeneity and harbors one of the world''s richest floras. The high species endemism, together with increasing anthropogenic threats, has qualified the Himalaya as one of the most significant global biodiversity hotspots. The topographic and climatic complexity of the Himalaya makes it an ideal natural laboratory for studying the mechanisms of floral exchange, diversification, and spatiotemporal distributions. Here, we review literature pertaining to the Himalaya in order to generate a concise synthesis of the origin, distribution, and climate change responses of the Himalayan flora. We found that the Himalaya supports a rich biodiversity and that the Hengduan Mountains supplied the majority of the Himalayan floral elements, which subsequently diversified from the late Miocene onward, to create today''s relatively high endemicity in the Himalaya. Further, we uncover links between this Miocene diversification and the joint effect of geological and climatic upheavals in the Himalaya. There is marked variance regarding species dispersal, elevational gradients, and impact of climate change among plant species in the Himalaya, and our review highlights some of the general trends and recent advances on these aspects. Finally, we provide some recommendations for conservation planning and future research. Our work could be useful in guiding future research in this important ecosystem and will also provide new insights into the maintenance mechanisms underpinning other mountain systems.     

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.     

Phylogenetic diversity,structure and diversification patterns of endemic plants along the elevational gradient in the Eastern Himalaya

ABSTRACT

Background: Several studies have documented the variation in species diversity patterns along elevational gradients in the Himalaya, but few have reported the evolutionary and biogeographic processes behind these patterns.

Aims: To understand whether evolutionary history and phylogeny have any role in structuring plant species communities along an elevational gradient in the Sikkim Himalaya.

Methods: We used data on endemic plant species occurrence from primary and secondary sources to construct family-level phylogenetic supertrees for different growth forms with the help of Phylomatic tool of Phylocom. These phylogenetic supertrees were used as a base for testing phylogenetic diversity (PD), niche conservatism, diversification time patterns and phylogenetic structure of various plant growth forms along an elevational gradient.

Results: PD was the highest at mid-elevations for all growth forms and PD had a significant positive correlation with endemic species richness. Species at mid-elevations were dominated by the ancestral/primitive taxa. There was phylogenetic clustering at higher elevations and phylogenetic overdispersion at lower and mid-elevations for the majority of the growth forms.

Conclusions: Time-for-speciation effect and niche conservatism along elevation (retention of niche-related ancestral elevational distribution over evolutionary time scale by species) together determine plant species diversity patterns in the Himalaya.     

Diversity and distribution of small mammals in the South American Dry Andes

The Andean mountain range has played an important role in the evolution of South American biota. However, there is little understanding of the patterns of species diversity across latitudinal and altitudinal gradients. In this paper, we examine the diversity of small mammals along the South Central Dry Andes (SCDA) within the framework of two contrasting hypotheses: (a) species richness decreases with increasing elevation and latitude; and (b) species richness peaks at altitudinal midpoints (mid‐domain). We explore the composition of the species pool, the impact of species–area relationships and the Rapoport effect (i.e. size of geographic ranges) along latitudinal and elevational gradients. First, we constructed a database of SCDA small mammals. Then, species richness patterns were analysed through generalized models, and species–area relationships were assessed by log–log regressions; the curvilinear method (c = S/Az) was use to compute richness corrected by area size. Lastly, the Rapoport effect was evaluated using the midpoint method. Our results show: (1) a richness of 67 small mammals along the SCDA, of which 36 are endemic; (2) a hump‐shaped pattern in species richness along elevation and latitudinal gradients; (3) a species–area relationship for both gradients; (4) endemic species corrected by area present a strong and positive relationship with elevation; (5) a Rapoport effect for the latitudinal ranges, but no effect across the elevational gradient; and (6) a major species turnover between 28° and 30° south latitude. This is the first study quantifying the diversity of small mammals encompassing the central Andean region. Overall, our macrogeographic analysis supports the previously postulated role of the Andes in the diversification of small mammals (i.e. in situ cladogenesis) and highlights some basic attributes (i.e. anatomy of geographic ranges; species–area relationships) when considering the consequences of climate change on biodiversity conservation of mountain ecosystems.     

Patterns of an elevational gradient affecting moths across the South Korean mountains: effects of geometric constraints,plants, and climate

We investigated elevational richness patterns of three moth groups (Erebidae, Geometridae, and Noctuidae) along four elevational gradients located on one northern and three southern mountains in South Korea, as well as the effects of plants and climatic factors on the diversity patterns of moths. Moths were collected with an ultraviolet light trap at 32 sites from May through October, 2013. Plant species richness and mean temperatures for January and June were acquired. Observed and estimated moth species richness was calculated and the diversity patterns with null models were compared. Species richness along four elevational gradients peaked at mid-elevations, whereas deviations occurred at elevations below mid-peak in the southern mountains and elevations higher than mid-peak on the northern mountain. Species richness curves of three moth groups also peaked at mid-elevations throughout South Korea. However, the species richness curves for Erebidae were positively skewed, indicating that a preference for lowlands, whereas curves of the Geometridae were negatively skewed, indicating a preference for highlands. The mid-peak diversity pattern between plants and moths on the Korean mountains showed an elevational breadth that overlapped between 800 and 900 m. Multiple regression analysis revealed that plant species richness and January mean temperature significantly influenced moth species richness and abundance. The rapid increase in mean annual temperature in the Korean peninsula and the unimodal elevational gradients of moths across the country suggest that an uphill shift in peak optimum elevation and changes in the highest peak of the curve will occur in the future.     

The mid‐domain effect and habitat complexity applied to elevational gradients: Moss species richness in a temperate semihumid monsoon climate mountain of China

The utility of elevational gradients as tools to test either ecological hypotheses and delineate elevation‐associated environmental factors that explain the species diversity patterns is critical for moss species conservation. We examined the elevational patterns of species richness and evaluated the effects of spatial and environmental factors on moss species predicted a priori by alternative hypotheses, including mid‐domain effect (MDE), habitat complexity, energy, and environment proposed to explain the variation of diversity. Last, we assessed the contribution of elevation toward explaining the heterogeneity among sampling sites. We observed the hump‐shaped distribution pattern of species richness along elevational gradient. The MDE and the habitat complexity hypothesis were supported with MDE being the primary driver for richness patterns, whereas little support was found for the energy and the environmental factors.     

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.     

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.     

喜马拉雅山脉种子植物多样性特征及分布格局

喜马拉雅山脉是全球著名的生物多样性热点地区之一。该研究对以往收集的喜马拉雅山脉南、北坡植物物种名录及其分布数据进行整合,借助在线数据库对分布数据进行补充与修订,最后整理并汇总了喜马拉雅山脉位于中国、印度、尼泊尔、不丹4国境内的种子植物分布情况,并在此基础上对科属特征、物种组成相似性、区系成分以及海拔梯度上物种分布格局进行分析,为该区域的生物多样性研究以及保护提供数据支撑。结果表明：(1)喜马拉雅山脉共分布有种子植物11 875种,隶属223科2 086属,其中包含7 906种草本植物(66.6%),2 583种灌木(21.8%)和1 386种乔木(11.7%)。(2)研究区涵盖物种数量位于前20的科有菊科(Asteraceae)、兰科(Orchidaceae)、禾本科(Poaceae)、豆科(Fabaceae)、杜鹃花科(Ericaceae)等科,共包含物种7 456种,约占喜马拉雅山脉植物种的62.8%;涵盖物种数量位于前20的属有杜鹃花属(Rhododendron)、报春花属(Primula)、马先蒿属(Pedicularis)、虎耳草属(Saxifraga)、薹草属(Carex)...     

秦岭南坡陕西洋县辖区哺乳动物物种多样性的空间分布格局

物种多样性的空间分布格局一直是生态学和生物地理学研究的一个热点问题。山地生态系统的生境异质性和物种多样性高, 适合研究物种多样性空间分布格局及其相关机制。2016年11月至2017年11月, 本研究选取秦岭南坡陕西洋县辖区作为研究区域, 采用样线法、红外相机法和笼捕/夹捕法, 系统分析了8目21科48种哺乳动物物种多样性的空间分布格局。研究结果发现秦岭南坡洋县辖区哺乳动物物种丰富度的空间分布格局大致是中南部低, 北部和东部高; 物种多样性指数大致是中南部和北部低, 东部高。啮齿类动物和非啮齿类动物的空间分布格局存在差异。哺乳动物物种丰富度和多样性指数的垂直分布格局都符合中峰模式, 但啮齿类动物和非啮齿类动物间存在差异。最优线性模型结果表明, 研究地区哺乳动物物种多样性的空间分布格局受到多种环境因素的共同影响。其中, 年均温与物种多样性的相关性最强, 在6个最优线性模型中贡献都是最大。综上, 秦岭南坡洋县辖区中高海拔区域的物种多样性较高, 应加强对中高海拔地区的保护, 以维系该区域较高的生物多样性。     

Phylogeography sheds light on the central–marginal hypothesis in a Mediterranean narrow endemic plant

Background and Aims

Understanding the factors that shape variation in genetic diversity across the geographic ranges of species is an important challenge in the effort to conserve evolutionary processes sustaining biodiversity. The historical influences leading to a central–marginal organization of genetic diversity have been explored for species whose range is known to have expanded from refugia after glacial events. However, this question has rarely been addressed for Mediterranean endemic plants of azonal habitats such as rocky slopes or screes. In this context, this comprehensive study examined molecular and field data from Arenaria provincialis (Caryophyllaceae), a narrow endemic plant of south-eastern France.

Methods

Across the whole geographic range, an investigation was made of whether high levels of abundance and genetic diversity (estimated from amplified fragment length polymorphism markers) are centrally distributed, to evaluate the relevance of the central–marginal hypothesis. Phylogeographic patterns inferred from chloroplast DNA (cpDNA) were used, applying Bayesian methods to test the influence of past biogeographic events. Multivariate analysis combining phylogeographic and ecological data was used to reveal the historical and ecological distinctiveness of populations.

Key Results

Despite the narrow distribution of A. provincialis, a high level of nucleotide variation is found within cpDNA loci, supporting its persistence throughout the Pleistocene period. The area characterized by the highest genetic diversity is centrally located. Structured phylogeography and Bayesian factor analysis supported the hypothesis that the central area of the distribution was the source of both westward and eastward migrations, probably during arid periods of the Pleistocene, and more recently was a crossroads of backward migrations. By contrast, the two areas located today at the range limits are younger, have reduced genetic diversity and are marginal in the ecological gradients.

Conclusions

This study highlights a case of strong population distinctiveness within a narrow range. Phylogeography sheds light on the historical role of the areas centrally situated in the distribution. The current range size and abundance patterns are not sufficient to predict the organization of genetic diversity.