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

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

Elevational patterns of species richness and endemism for some important taxa in the Hengduan Mountains, southwestern China

We describe the elevational patterns of species richness and endemism of some important taxa in the Hengduan Mountains, southwest China. Species richness data came from publications, an online database, herbaria and field work. Species richness was estimated by rarefaction and interpolation. The Hengduan Mountains region was divided into a southern and northern subregion, and all species were assigned to four groups based on their distributional range within this region. The conditional autoregressive model (CAR) was used to relate species richness and explanatory variables. The elevational patterns of total, endemic and non-endemic species richness, at subregion and entire region scales, presented to be unimodal and peaked at similar elevations. Area size was strongly related with species richness, and was more powerful in explaining variation in species richness in the northern subregion than in the southern subregion. A single climatic variable (mean annual rainfall, potential evapotranspiration or moisture index) showed a weak relationship with the elevational pattern of species richness. Area and climatic variables together explained more than 67% of the variation in non-endemic richness, 53% in total richness, and 50% in endemic richness. There were three patterns of endemism at the generic level with increasing elevation: namely endemism increased, decreased, or peaked at middle elevations. All selected taxa have experienced rapid speciation and evolution within this region, which plays an important role in the uniform elevational patterns of total, endemic and non-endemic richness, and in the multiform elevational patterns of endemism. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects

We studied frog biodiversity along an elevational gradient in the Hengduan Mountains, China. Endemic and non-endemic elevational diversity patterns were examined individually. Competing hypotheses were also tested for these patterns. Species richness of total frogs, endemics and non-endemics peaked at mid-elevations. The peak in endemic species richness was at higher elevations than the maxima of total species richness. Endemic species richness followed the mid-domain model predictions, and showed a nonlinear relationship with temperature. Water and energy were the most important variables in explaining elevational patterns of non-endemic species richness. A suite of interacting climatic and geometric factors best explained total species richness patterns along the elevational gradient. We suggest that the mid-domain effect was an important factor to explain elevational richness patterns, especially in regions with high endemism.     

Elevational patterns of fern species assemblages and richness in central Japan

We conducted field surveys in 807 quadrats to evaluate the elevational belts, boundary and richness patterns of ferns and lycophytes in the temperate region of central Japan. We analysed fern species assemblages at 100 m elevational steps by cluster analysis and tested the number of upper and lower boundaries for elevational intervals against a null model of random distribution of elevational limits. We compared the pattern of fern species richness along the elevational gradients in central Japan with patterns in several locations to evaluate the fern flora in central Japan in relation to the rest of the world. We recorded 261 ferns species in total, which is one-third of the Japanese ferns. We found clear elevational boundaries of fern assemblages at 900 and 1,800 m and three fern elevational zones, which corresponded well to the elevational limits of forest types in central Japan. The pattern of fern species richness in central Japan was an asymmetric hump-shaped pattern that peaked close to the sea level, with the peak of local richness at lower elevations than that of regional richness. We found that the peak of fern species richness along the elevational gradient in Japan was located at lower elevations than that of fern elevational patterns in several locations around the world.     

Fern endemism and its correlates: contribution from an elevational transect in Costa Rica

We studied the distribution patterns of endemic ferns along an elevational gradient of 3400 m in Costa Rica, Central America. We related the endemism patterns of the whole species set and separated for life forms and microhabitats according to topography and environmental factors. Fern species were surveyed in 156 plots each with an area of 400 m², with up to five plots at every elevational step of 100 m. Global range size for every species was compiled from literature data, and species restricted to the mountain range from Costa Rica and adjacent western Panama were defined as endemic (24.5% of all species recorded). We found patterns of endemism rates mostly peaking at mid-elevation, but when separated for different life forms and microhabitats, some deviations from the overall pattern emerged. High constant humidity and reduced surface area were closely related to high levels of endemism. High humidity is discussed as a general predictor for high endemism rates in concert with highest overall richness. Restricted area of elevational belts, indicating a fragmented habitat, leads to a higher degree of population isolation and thus species differentiation. However, both interpretations were not fully supported by our data. Most importantly, endemism rates were fairly low on mountain tops that have the smallest available area in a topographically highly fragmented setting. In contrast, endemic species were more common than widespread species at the highest elevations. History and climatic shifts are assumed to play a role in this respect.     

Pollen representation of vegetational patterns along an elevational gradient

Abstract. Modern pollen assemblages from 16 small lakes (< 2.5 ha) and 11 moderate-sized lakes (4.5–19.3 ha) arrayed along an elevational gradient (300 to 1320 m) in the east-central Adirondack Mountains were studied to determine how well the pollen assemblages recorded patterns of forest composition along the gradient. Forest composition ranges from Pinus strobus/Tsuga/ hardwoods forests at low elevations through Tsuga/ hardwoods, hardwoods, and Picea/Abies forests to Abies-dominated forests at high elevations. Modern pollen percentages for 10 tree taxa were compared with lake elevation using scatter plots and correlation and regression analysis. Differential smoothing of vegetational patterns along the elevational gradient occurred among the 10 taxa owing to differences in pollen dispersibility, pollen production, and spatial pattern of taxon abundance in forests of the region. No differences were observed in pollen-elevation patterns between small and moderate-sized lakes. Pollen-elevation patterns were obscured for most taxa when the gradient was shortened (e.g. to 600–1320 m) owing to increased spatial smoothing by pollen dispersal. Design and interpretation of paleoecological studies of spatial gradients can be improved by careful attention to site spacing, gradient length, and gradient steepness in the context of pollen dispersal and representation models.     

Diversity patterns of vascular epiphytes along an elevational gradient in the Andes

Aim To document the elevational pattern of epiphyte species richness at the local scale in the tropical Andes with a consistent methodology. Location The northern Bolivian Andes at 350–4000 m above sea level. Methods We surveyed epiphytic vascular plant assemblages in humid forests in (a) single trees located in (b) 90 subplots of 400 m² each located in (c) 14 plots of 1 ha each. The plots were separated by 100–800 m along the elevational gradient. Results We recorded about 800 epiphyte species in total, with up to 83 species found on a single tree. Species richness peaked at c. 1500 m and declined by c. 65% to 350 m and by c. 99% to 4000 m, while forests on mountain ridges had richness values lowered by c. 30% relative to slope forests at the same elevations. The hump‐shaped richness pattern differed from a null‐model of random species distribution within a bounded domain (the mid‐domain effect) as well as from the pattern of mean annual precipitation by a shift of the diversity peak to lower elevations and by a more pronounced decline of species richness at higher elevations. With the exception of Araceae, which declined almost monotonically, all epiphyte taxa showed hump‐shaped curves, albeit with slightly differing shapes. Orchids and pteridophytes were the most species‐rich epiphytic taxa, but their relative contributions shifted with elevation from a predominance of orchids at low elevations to purely fern‐dominated epiphyte assemblages at 4000 m. Within the pteridophytes, the polygrammoid clade was conspicuously overrepresented in dry or cold environments. Orchids, various small groups (Cyclanthaceae, Ericaceae, Melastomataceae, etc.), and Bromeliaceae (below 1000 m) were mostly restricted to the forest canopy, while Araceae and Pteridophyta were well represented in the forest understorey. Main conclusions Our study confirms the hump‐shaped elevational pattern of vascular epiphyte richness, but the causes of this are still poorly understood. We hypothesize that the decline of richness at high elevations is a result of low temperatures, but the mechanism involved is unknown. The taxon‐specific patterns suggest that some taxa have a phylogenetically determined propensity for survival under extreme conditions (low temperatures, low humidity, and low light levels in the forest interior). The three spatial sampling scales show some different patterns, highlighting the influence of the sampling methodology.     

Regional variation in the historical components of global avian species richness

Aim Using a global data base of the distribution of extant bird species, we examine the evidence for spatial variation in the evolutionary origins of contemporary avian diversity. In particular, we assess the possible role of the timing of mountain uplift in promoting diversification in different regions. Location Global. Methods We mapped the distribution of avian richness at four taxonomic levels on an equal‐area 1° grid. We examined the relationships between richness at successive taxonomic levels (e.g. species richness vs. genus richness). We mapped the residuals from linear regressions of these relationships to identify areas that are exceptional in the number of lower taxa relative to the number of higher taxa. We use generalized least squares models to test the influence of elevation range and temperature on lower‐taxon richness relative to higher‐taxon richness. Results Peaks of species richness in the Neotropics were congruent with patterns of generic richness, whilst peaks in Australia and the Himalayas were congruent with patterns of both genus and family richness. Hotspots in the Afrotropics did not reflect higher‐taxon patterns. Regional differences in the relationship between richness at successive taxonomic levels revealed variation in patterns of taxon co‐occurrence. Species and genus co‐occurrence was positively associated with elevational range across much of the world. Taxon occurrence in the Neotropics was associated with a positive interaction between elevational range and temperature. Conclusions These results demonstrate that contemporary patterns of richness show different associations with higher‐taxon richness in different regions, which implies that the timing of historical effects on these contemporary patterns varies across regions. We suggest that this is due to dispersal limitation and phylogenetic constraints on physiological tolerance limits promoting diversification. We speculate that diversification rates respond to long‐term changes in the Earth's topography, and that the role of tropical mountain ranges is implicated as a correlate of contemporary diversity, and a source of diversification across avian evolutionary history.     

Biodiversity in South African fynbos and Mediterranean heathland

Abstract. The theory of convergence predicts that, given similar selective regimes, both present and past, unrelated ecological communities will show similar attributes. Mild Pleistocene climate, highly infertile soils, and similar fire regimes explain the remarkable convergence between mediterranean‐type vegetation from South Africa (fynbos) and Australia (kwongan). Heathlands in the Aljibe Mountains, at the western end of the Mediterranean basin, constitute a single vegetation type within the Mediterranean region. We studied the association between endemism and plant life form in a flora from environmentally similar areas of the South African Cape region (fynbos) and the Aljibe Mountains by contingency table analysis. We included two non‐acid, neighbouring areas to the latter region in the analysis as contrasts. We also compared the patterns of variation in three components of biodiversity (species richness, endemism level and taxonomic singularity) of fynbos and Aljibe heathland woody plant communities along similar soil fertility gradients by means of two‐way ANOVAs. At the regional (flora) level, our results show two common features in the biological aspects of endemism between the two regions: (1) edaphic endemism and (2) association of endemism with the shrub growth form. At the community level, we detected strong similarities in the patterns of variation of endemism and taxonomic singularity of woody communities from both regions along an ecological gradient related to soil fertility. We interpret these similarities, both at the regional and community levels, as suggestive of convergence between fynbos and Aljibe heathland.     

Vertebrate metacommunity structure along an extensive elevational gradient in the tropics: a comparison of bats,rodents and birds

Aim We evaluated the structure of metacommunities for each of three vertebrate orders (Chiroptera, Rodentia and Passeriformes) along an extensive elevational gradient. Using elevation as a proxy for variation in abiotic characteristics and the known elevational distributions of habitat types, we assessed the extent to which variation in those factors may structure each metacommunity based on taxon‐specific characteristics. Location Manu Biosphere Reserve in the Peruvian Andes. Methods Metacommunity structure is an emergent property of a set of species distributions across geographic or environmental gradients. We analysed elements of metacommunity structure (coherence, range turnover and range boundary clumping) to determine the best‐fit structure for each metacommunity along an elevational gradient comprising 13 250‐m elevational intervals and 58 species of rodent, 92 species of bat or 586 species of passerine. Results For each taxon, the environmental gradient along which the metacommunity was structured was highly correlated with elevation. Clementsian structure (i.e. groups of species replacing other such groups along the gradient) characterized rodents, with a group of species that was characteristic of rain forests and a group of species that was characteristic of higher elevation habitats (i.e. above 1500 m). Distributions of bats were strongly nested, with more montane communities comprising subsets of species at lower elevations. The structure of the passerine metacommunity was complex and most consistent with a quasi‐Clementsian structure. Main conclusions Each metacommunity exhibited a different structure along the same elevational gradient, and each structure can be accounted for by taxon‐specific responses to local environmental factors that vary predictably with elevation. The structures of rodent and bird metacommunities suggest species sorting associated with habitat specializations, whereas structure of the bat metacommunity is probably moulded by a combination of species‐specific tolerances to increasingly cold, low‐productivity environs of higher elevations and the diversity and abundance of food resources associated with particular habitat types.     

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.     

A taxonomic,functional, and phylogenetic perspective on the community assembly of passerine birds along an elevational gradient in southwest China

Integrating multiple facets of biodiversity to describe spatial and temporal distribution patterns is one way of revealing the mechanisms driving community assembly. We assessed the species, functional, and phylogenetic composition and structure of passerine bird communities along an elevational gradient both in wintering and breeding seasons in the Ailao Mountains, southwest China, in order to identify the dominant ecological processes structuring the communities and how these processes change with elevation and season. Our research confirms that the highest taxonomic diversity, and distinct community composition, was found in the moist evergreen broadleaf forest at high elevation in both seasons. Environmental filtering was the dominant force at high elevations with relatively cold and wet climatic conditions, while the observed value of mean pairwise functional and phylogenetic distances of low elevation was constantly higher than expectation in two seasons, suggested interspecific competition could play the key role at low elevations, perhaps because of relative rich resource result from complex vegetation structure and human‐induced disturbance. Across all elevations, there was a trend of decreasing intensity of environmental filtering whereas increasing interspecific competition from wintering season to breeding season. This was likely due to the increased resource availability but reproduction‐associated competition in the summer months. In general, there is a clear justification for conservation efforts to protect entire elevational gradients in the Ailao Mountains, given the distinct taxonomic, functional, and phylogenetic compositions and also elevational migration pattern in passerine bird communities.     

A taxonomic and phylogenetic perspective on plant community assembly along an elevational gradient in subtropical forests

亚热带森林植物群落沿海拔梯度的分类与系统发育研究 生物多样性沿海拔梯度的分布格局已受到广泛关注。然而,生物多样性格局沿海拔梯度的变异及其潜在机制尚不清楚。整合生物多样性的多维度信息为理解群落构建机制提供了新思路。本研究在我国东部亚热带森林沿海拔270–1470 m的梯度上设置了17个木本植物固定样地,分析了沿海拔梯度植物群落 构建的生态和进化驱动力。基于样地内物种出现(0–1数据)和多度信息,计算群落内被子植物的物种和系统发育alpha和beta多样性、系统发育结构等,并量化多样性指标与微气候和地形之间的关系。研究发现,不论多度加权与否,物种alpha多样性均沿海拔升高而增加,物种和系统发育的相似性随海拔距离的增加而呈衰减趋势。然而,多度加权与否会形成不同的系统发育alpha多样性格局。对于系统发育结构而言,沿海拔增加并无明显趋势。地形和微气候是多样性格局和系统发育结构的主要驱动力。与未考虑物种多度的多样性指标相比,多度加权的指标与坡度和胸高断面积相关性更高。这些结果表明,由局域物种多度介导的确定性过程对沿海拔梯度的植物群落构建具有一定影响。     

Contributions toward understanding the biodiversity of Passiflora in North America: Updates and a new combination from the Baja California Peninsula,Mexico and vicinity

The Baja California Peninsula and surrounding landmasses harbor an abundant flora in an otherwise harsh and arid environment. Of the many plant groups native to this peninsular and insular region, passionflowers (Passiflora, Passifloraceae) are represented by several conspicuous taxa that all belong to a single lineage, section Dysosmia. Basic questions remain regarding this group, particularly the taxonomic status among the Passiflora arida complex. Therefore, we investigated the claims of endemism, habitat characteristics, and taxon boundaries with in section Dysosmia in the Baja California region using extensive sampling of herbarium specimens and iNaturalist observations. We confirmed that only one of the native Passiflora taxa (P. fruticosa) was endemic to the Baja California Peninsula, with an additional taxon (P. palmeri) considered near‐endemic. Environmental data revealed significant distinctions between the habitats of many of the native taxa as well as within the P. arida complex, especially with respect to precipitation and temperature tolerances. Geometric morphometric analyses of leaf shape were largely not successful at separating taxa, indicating leaf shape may not be a good indicator of taxon identity in this particular group. Based on ecological differences and discrete macro‐ and micromorphological features, a varietal name is here synonymized and a new combination is proposed: Passiflora pentaschista.     

澜沧江流域水生昆虫群落分类多样性和功能多样性海拔格局的空间尺度依赖性

群落分类多样性和功能多样性的海拔格局研究, 是了解生物多样性空间分布现状、揭示多样性维持和变化机制的重要途径。当前对水生昆虫分类多样性和功能多样性沿海拔梯度分布格局, 及其尺度依赖性依旧缺乏深入研究。本文基于2013-2018年在云南澜沧江流域500-3,900 m海拔梯度共149个溪流点位的水生昆虫群落调查数据, 利用线性或二次回归模型探索并比较了局部尺度(点位尺度)和不同区域尺度(100 m、150 m、200 m、250 m海拔段)的分类多样性指数(物种丰富度指数、Simpson多样性指数和物种均匀度指数)和功能多样性指数(树状图功能多样性指数(dbFD)、Rao二次熵指数(RaoQ)和功能均匀度指数(FEve))的海拔格局。结果表明, 在局部尺度, 物种丰富度指数和dbFD指数沿海拔梯度均无显著分布特征, Simpson多样性指数、RaoQ指数、物种均匀度指数和FEve指数沿海拔梯度呈现U型或者单调递减趋势。在区域尺度, 随着区域海拔带宽度的增加, 物种丰富度指数沿海拔呈不显著的单调递减格局, 但dbFD指数沿海拔分布由U型转变为单调递减趋势; Simpson多样性指数和RaoQ指数沿海拔梯度由显著U型趋势转变为无显著分布特征; 物种均匀度指数沿海拔梯度无显著分布特征, 但FEve指数呈显著增加的海拔格局。综上, 群落分类多样性指数和功能多样性指数沿海拔梯度分布存在局部和区域尺度的空间差异, 但区域尺度下二者海拔格局随海拔带宽度的增加存在一定程度的一致性。     

Anthropogenic threats can have cascading homogenizing effects on the phylogenetic and functional diversity of tropical ecosystems

Determining the mechanisms that underlie species distributions and assemblages is necessary to effectively preserve biodiversity. This cannot be accomplished by examining a single taxonomic group, as communities comprise a plethora of interactions across species and trophic levels. Here, we examine the patterns and relationships among plant, mammal, and bird diversity in Madagascar, a hotspot of biodiversity and endemism, across taxonomic, phylogenetic, and functional axes. We found that plant community diversity and structure are shaped by geography and climate, and have significant influences on the taxonomic, phylogenetic and functional diversity of mammals and birds. Patterns of primate diversity, in particular, were strongly correlated with patterns of plant diversity. Furthermore, our findings suggest that plant and animal communities could become more phylogenetically and functionally clustered in the future, leading to homogenization of the flora and fauna. These results underscore the importance and need of multi‐taxon approaches to conservation, given that even small threats to plant diversity can have significant cascading effects on mammalian and avian community diversity, structure, and function.     

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.     

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.     

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

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