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.     

南岭国家级自然保护区兰科植物物种多样性的海拔梯度格局

兰科是单子叶植物最大的科和被子植物中的第二大科.兰科植物物种多样性的海拔梯度格局因地域或坡向不同而呈不同趋势.南岭国家级自然保护区地处南岭山脉中段南坡,植物资源丰富.作者对南岭国家级自然保护区兰科植物进行了为期4年的调查,记录了其种类、数量、生活型、海拔、生境,分析了其区系成分,并依据IUCN(2001)濒危物种红色名录及保护区的具体情况对其濒危等级进行了评估.从海拔230-1,902m,每200 m分段,共划分为9个海拔段分析其垂直分布格局.结果显示:(1)实地记录兰科植物38属70种,大多数种类数量较少,乳阳管理处的种类最为丰富;(2)随着海拔的上升,总体兰科植物以及不同生活型的兰科植物物种数量均呈现"中间高度膨胀型"而峰值偏向低海拔的特点;(3)濒危物种主要集中在600-1,000 m人为活动稀少、环境好的小生境中;(4)800-1,000 m的区系成分最丰富,热带性质的兰花多数分布在海拔1,600 m以下;(5)相邻海拔段的兰科植物种类的相似性系数较高;海拔段相隔越远,相似性系数越小.小生境和水分条件对于兰科植物的生长非常重要,中海拔具有良好的水热条件,因此物种多样性最高;而低海拔地区的人为破坏使得该海拔段兰科植物物种的多样性显著降低.     

A comparison of altitudinal species richness patterns of bryophytes with other plant groups in Nepal, Central Himalaya

Aim To explore species richness patterns in liverworts and mosses along a central Himalayan altitudinal gradient in Nepal (100–5500 m a.s.l.) and to compare these patterns with patterns observed for ferns and flowering plants. We also evaluate the potential importance of Rapoport’s elevational rule in explaining the observed richness patterns for liverworts and mosses. Location Nepal, Central Himalaya. Methods We used published data on the altitudinal ranges of over 840 Nepalese mosses and liverworts to interpolate presence between maximum and minimum recorded elevations, thereby giving estimates of species richness for 100‐m altitudinal bands. These were compared with previously published patterns for ferns and flowering plants, derived in the same way. Rapoport’s elevational rule was assessed by correlation analyses and the statistical significance of the observed correlations was evaluated by Monte Carlo simulations. Results There are strong correlations between richness of the four groups of plants. A humped, unimodal relationship between species richness and altitude was observed for both liverworts and mosses, with maximum richness at 2800 m and 2500 m, respectively. These peaks contrast with the richness peak of ferns at 1900 m and of vascular plants, which have a plateau in species richness between 1500 and 2500 m. Endemic liverworts have their maximum richness at 3300 m, whereas non‐endemic liverworts show their maximum richness at 2700 m. The proportion of endemic species is highest at about 4250 m. There is no support from Nepalese mosses for Rapoport’s elevational rule. Despite a high correlation between altitude and elevational range for Nepalese liverworts, results from null simulation models suggest that no clear conclusions can be made about whether liverworts support Rapoport’s elevational rule. Main conclusions Different demands for climatic variables such as available energy and water may be the main reason for the differences between the observed patterns for the four plant groups. The mid‐domain effect may explain part of the observed pattern in moss and liverwort richness but it probably only works as a modifier of the main underlying relationship between climate and species richness.     

Elevational species richness patterns for vascular plants on Mount Kinabalu, Borneo

Aim  We quantify the elevational patterns of species richness for all vascular plants and some functional and taxonomic groups on a regional scale on a tropical mountain and discuss some possible causes for the observed patterns.
Location  Mount Kinabalu, Sabah, Borneo.
Methods  A data base containing elevational information on more than 28,000 specimens was analysed for vascular plant distribution, taking into account sampling effort. The total species richness pattern was estimated per 300-m elevational interval by rarefaction analyses. The same methods were also applied to quantify species richness patterns of trees, epiphytes, and ferns.
Results  Total species richness has a humped relationship with elevation, and a maximum species richness in the interval between 900 and 1200 m. For ferns and epiphytes the maximum species richness is found at slightly higher elevations, whereas tree species did not have a statistically significant peak in richness above the lowest interval analysed.
Main conclusions  For the first time a rigorous estimate of an elevational pattern in species richness of the whole vascular plant flora of a tropical mountain has been quantified. The pattern observed depends on the group studied. We discuss the differences between the groups and compare the results with previous studies of elevational patterns of species richness from other tropical areas. We also discuss the methods used to quantify the richness pattern and conclude that rarefaction gives an appropriate estimate of the species richness pattern.     

Elevational patterns and hierarchical determinants of biodiversity across microbial taxonomic scales

Microbial biogeography is gaining increasing attention due to recent molecular methodological advance. However, the diversity patterns and their environmental determinants across taxonomic scales are still poorly studied. By sampling along an extensive elevational gradient in subarctic ponds of Finland and Norway, we examined the diversity patterns of aquatic bacteria and fungi from whole community to individual taxa across taxonomic coverage and taxonomic resolutions. We further quantified cross‐phylum congruence in multiple biodiversity metrics and evaluated the relative importance of climate, catchment and local pond variables as the hierarchical drivers of biodiversity across taxonomic scales. Bacterial community showed significantly decreasing elevational patterns in species richness and evenness, and U‐shaped patterns in local contribution to beta diversity (LCBD). Conversely, no significant species richness and evenness patterns were found for fungal community. Elevational patterns in species richness and LCBD, but not in evenness, were congruent across bacterial phyla. When narrowing down the taxonomic scope towards higher resolutions, bacterial diversity showed weaker and more complex elevational patterns. Taxonomic downscaling also indicated a notable change in the relative importance of biodiversity determinants with stronger local environmental filtering, but decreased importance of climatic variables. This suggested that niche conservatism of temperature preference was phylogenetically deeper than that of water chemistry variables. Our results provide novel perspectives for microbial biogeography and highlight the importance of taxonomic scale dependency and hierarchical drivers when modelling biodiversity and species distribution responses to future climatic scenarios.     

Support for the elevational Rapoport's rule among seed plants in Nepal depends on biogeographical affinities and boundary effects

As one of the most important hypotheses on biogeographical distribution, Rapoport's rule has attracted attention around the world. However, it is unclear whether the applicability of the elevational Rapoport's Rule differs between organisms from different biogeographical regions. We used Stevens’ method, which uses species diversity and the averaged range sizes of all species within each (100 m) elevational band to explore diversity‐elevation, range‐elevation, and diversity‐range relationships. We compared support for the elevational Rapoport's rule between tropical and temperate species of seed plants in Nepal. Neither tropical nor temperate species supported the predictions of the elevational Rapoport's rule along the elevation gradient of 100–6,000 m a.s.l. for any of the studied relationships. However, along the smaller 1,000–5,000 m a.s.l. gradient (4,300 m a.s.l. for range‐elevation relationships) which is thought to be less influenced by boundary effects, we observed consistent support for the rule by tropical species, although temperate species did not show consistent support. The degree of support for the elevational Rapoport's rule may not only be influenced by hard boundary effects, but also by the biogeographical affinities of the focal taxa. With ongoing global warming and increasing variability of temperature in high‐elevation regions, tropical taxa may shift upward into higher elevations and expand their elevational ranges, causing the loss of temperate taxa diversity. Relevant studies on the elevational Rapoport's rule with regard to biogeographical affinities may be a promising avenue to further our understanding of this rule.     

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.     

Altitudinal biodiversity patterns of seed plants along Gongga Mountain in the southeastern Qinghai–Tibetan Plateau

The mechanisms underlying elevation patterns in species and phylogenetic diversity remain a central issue in ecology and are vital for effective biodiversity conservation in the mountains. Gongga Mountain, located in the southeastern Qinghai–Tibetan Plateau, represents one of the longest elevational gradients (ca. 6,500 m, from ca. 1,000 to 7,556 m) in the world for studying species diversity patterns. However, the elevational gradient and conservation of plant species diversity and phylogenetic diversity in this mountain remain poorly studied. Here, we compiled the elevational distributions of 2,667 native seed plant species occurring in Gongga Mountain, and estimated the species diversity, phylogenetic diversity, species density, and phylogenetic relatedness across ten elevation belts and five vegetation zones. The results indicated that species diversity and phylogenetic diversity of all seed plants showed a hump‐shaped pattern, peaking at 1,800–2,200 m. Species diversity was significantly correlated with phylogenetic diversity and species density. The floras in temperate coniferous broad‐leaved mixed forests, subalpine coniferous forests, and alpine shrublands and meadows were significantly phylogenetically clustered, whereas the floras in evergreen broad‐leaved forests had phylogenetically random structure. Both climate and human pressure had strong correlation with species diversity, phylogenetic diversity, and phylogenetic structure of seed plants. Our results suggest that the evergreen broad‐leaved forests and coniferous broad‐leaved mixed forests at low to mid elevations deserve more conservation efforts. This study improves our understanding on the elevational gradients of species and phylogenetic diversity and their determinants and provides support for improvement of seed plant conservation in Gongga Mountain.     

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)...     

Elevational gradient and human effects on butterfly species richness in the French Alps

We examined how butterfly species richness is affected by human impact and elevation, and how species ranges are distributed along the elevational gradient (200–2700 m) in the Isère Department (French Alps). A total of 35,724 butterfly observations gathered in summer (May–September) between 1995 and 2015 were analyzed. The number of estimated species per 100‐m elevational band was fitted to the elevational gradient using a generalized additive model. Estimations were also performed on a 500 m × 500 m grid at low altitude (200–500 m) to test for the human impact on species richness using generalized least squares regression models. Each species elevational range was plotted against the elevational gradient. Butterfly richness along the elevational gradient first increased (200–500 m) to reach a maximum of 150 species at 700 m and then remained nearly constant till a sharp decrease after 1900 m, suggesting that after some temperature threshold, only few specialized species can survive. At low elevation, urbanization and arable lands had a strongly negative impact on butterfly diversity, which was buffered by a positive effect of permanent crops. Butterfly diversity is exceptionally high (185 species) in this alpine department that represents less than 5% of the French territory and yet holds more than 70% of all the Rhopalocera species recorded in France. Both climate and habitat shape the distribution of species, with a negative effect of anthropization at low altitude and strong climatic constraints at high altitude.     

Spatial patterns of plant richness across treeline ecotones in the Pyrenees reveal different locations for richness and tree cover boundaries

Aim  To forecast the responses of alpine flora to the expected upward shift of treeline ecotones due to climatic warming, we investigated species richness patterns of vascular plants at small spatial scales across elevational transects.
Location  Richness patterns were assessed at local scales along the elevational gradient in two undisturbed treeline ecotones and one disturbed treeline ecotone in the Spanish Pyrenees.
Methods  We placed a rectangular plot (0.3–0.4 ha) in each treeline ecotone. We estimated and described the spatial patterns of plant richness using the point method and Moran's I correlograms. We delineated boundaries based on plant richness and tree cover using moving split windows and wavelet analysis. Then, to determine if floristic and tree cover boundaries were spatially related, overlap statistics were used.
Results  Plant richness increased above the forest limit and was negatively related to tree cover in the undisturbed sites. The mean size of richness patches in one of these sites was 10–15 m. Moving split windows and wavelets detected the sharpest changes in plant richness above the forest limit at both undisturbed sites. Most tree cover and plant richness boundaries were not spatially related.
Main conclusions  The upslope decrease of tree cover may explain the increase of plant richness across alpine treeline ecotones. However, the detection of abrupt richness boundaries well above the forest limit indicates the importance of local environmental heterogeneity to explain the patterns of plant richness at smaller scales. We found highly diverse microsites dominated by alpine species above the forest limit, which should be monitored to describe their response to the predicted upward shift of forests.     

喜马拉雅山哺乳动物物种多样性垂直分布格局

生物多样性的空间分布及其相关机制一直是生态学、生物地理学和保护生物学研究的热点问题。山地生态系统生境异质性和生物多样性高, 适合研究生物多样性空间分布及其相关机制。喜马拉雅山脉位于青藏高原南缘, 是全球生态热点区域。其地形复杂, 海拔落差大(100-8,844 m), 具有明显的垂直气候带。本研究通过整合野外调查和文献资料, 系统地分析了10目23科160属313种喜马拉雅山地区哺乳动物物种多样性的垂直分布格局, 发现该区域哺乳动物总体及其子集的物种多样性垂直分布格局都为左偏倚的中峰格局, 物种多样性在海拔900-1,400 m之间最高, 不同物种子集的物种多样性垂直分布格局的模式有所不同。UPGMA聚类分析表明, 喜马拉雅山地区哺乳动物群落沿海拔梯度可以划分为5个聚类簇(海拔100-1,500 m、1,500-2,000 m、2,000-3,000 m、3,000-4,200 m以及4,200-6,000 m的地区), 大致与该地区植被的垂直带分布相吻合。喜马拉雅山地区哺乳动物物种多样性在中低海拔最为丰富, 可能跟东洋界与古北界生物群扩散后的交汇地带相关。喜马拉雅山区贯通南北的沟谷是生物扩散和迁移的通道, 沟谷内水热资源较好, 气候稳定性高, 为高山生态系统内各种生物创造了栖息条件。综上, 喜马拉雅山沟谷地区是生物多样性热点地区, 也是生物扩散和交流关键的“生态走廊”, 应加强对喜马拉雅山沟谷地区的保护, 以维系该区域较高的生物多样性。     

An analysis of altitudinal behavior of tree species in Subansiri district,Eastern Himalaya

Plant species diversity and endemism demonstrate a definite trend along altitude. We analyzed the (i) pattern of tree diversity and its endemic subset (ii) frequency distribution of altitudinal range and (iii) upper & lower distributional limits of each tree species along altitudinal gradients in eastern Himalaya. The study was conducted in Subansiri district of Arunachal Pradesh. Data on the tree species (cbh ≥ 15 cm) were gathered every 200 m steps between 200 m and 2200 m gradients. Tree diversity demonstrated a greater variation along the gradients. A total of 336 species (of which 26 are endemic) were recorded belonging to 185 genera and 78 families. The alpha diversity demonstrated a decreasing pattern with two maxima (i.e., elevational peaks) along the gradients; one in 601–1000 m and the other in 1601–1800 m, corresponding to transition zones between tropical-subtropical and subtropical-temperate forests. Pattern diversity revealed a narrow range along the gradients. Frequency of altitudinal range was distributed between 1 and 41. Only one species (Altingia excelsa) showed widest amplitude, occurring over the entire range. Highest level of species turnover was found in 400–600 m step at lower elevational limit whereas for upper elevational limit, the highest turn over was recorded between 800 and 1000 m. Tree diversity decreased and its endemic subset increased along the gradients. Two maximas in tree diversity pattern correspond to forest transition zones with subtropical-temperate transition is narrower than tropical-subtropical. The pattern observed here could be attributed to varied microclimates or environmental heterogeneity. If altitudinal amplitude of a species is considered as an aspect of its niche breadth, it is clear from these results that niche breadth in these organisms is in fact independent of the diversity of the assemblage in which they occur. This analysis calls for detailed floristic studies to determine the breadth of changes between adjacent forest types and details of local species richness in high diversity areas.     

五台山亚高山土壤真菌海拔分布格局与构建机制

生物多样性的海拔分布格局与维持机制是生物多样性与生态系统功能研究的热点领域。尽管微生物驱动着地球上许多重要的生物地球化学循环,但与大型生物体相比,对微生物多样性海拔梯度分布格局知之甚少。运用Illumina Miseq高通量测序技术,全面分析了五台山亚高山生态系统（海拔2000-3058 m范围内）土壤真菌群落组成和多样性变化特征。结果表明,子囊菌门（Ascomycota）、担子菌门（Basidiomycota）、接合菌门（Zygomycota）、壶菌门（Chytridimycota）为主要的优势菌门。非度量多维尺度分析（NMDS）和相似性分析（ANOSIM）表明真菌群落组成和结构在海拔梯度上存在显著的差异（P<0.05）。典范对应分析（CCA）显示pH、植物丰富度、总碳含量与土壤真菌群落结构存在显著相关性（P<0.05）。局域海拔尺度上,土壤真菌多样性与植物多样性（α和β多样性）呈显著正相关关系（P<0.05）。方差分解分析（VPA）和偏Mantel分析表明土壤真菌群落构建过程中,环境因子和空间变量都起作用,并且环境因子占绝对的优势。土壤真菌群落之间的Bray-Curtis距离与海拔距离呈显著正相关关系（P<0.05）,说明环境选择是亚高山土壤真菌海拔分布格局的决定因素。总之,五台山亚高山沿海拔梯度土壤真菌群落结构和多样性产生明显的变化,群落构建主要由确定性过程和随机过程驱动,但确定性过程占主导地位。土壤pH、植物丰富度、总碳含量是影响土壤真菌群落结构的重要因素。     

Plant species richness, endemism, and genetic resources in Namibia

Namibia is a floristically diverse, arid to mesic country, with several highly distinct taxa. Including naturalized plants, there are about 4334 vascular plant species and infraspecific taxa within the country's borders, a substantial increase from the existing major reference work. Dominant families are the Poaceae (422species), Fabaceae (377), Asteraceae (385) and Mesembryanthemaceae (177). Freshwater algae and most other groups of lower plants remain poorly known. Concentrations of plant species richness are found in the Succulent Karoo biome, Kaokoveld, Otavi highland/Karstveld area, Okavango Basin, and Khomas highlands. Recent studies have led to a new estimate of 687 endemic plant species, defined as those contained wholly within Namibia's borders, amounting to about 17% of the Namibian flora. At least a further 275 species are Namib Desert endemics shared between the Kaokoveld and southern Angola (75spp.) and between the Succulent Karoo and northwestern South Africa (200spp.). Research on plant genetic resources is focused on species of potential or actual agricultural importance, such as pearl millet, Pennisetum glaucum, and cucurbits. Many wild plants have considerable genetic diversity and development potential. Primary threats to plant diversity fall in the category of poor land management and inappropriate development.     

广西九万山国家级自然保护区外来入侵和本地草本植物多样性垂直分布格局

外来入侵植物快速扩散、蔓延, 有些种类已进入到国家级自然保护区, 威胁保护区的生物多样性。本文基于九万山国家级自然保护区134个草本样方的调查数据, 对比分析了外来入侵草本和本地草本物种多样性的空间分布格局, 并阐明了气候因素对物种丰富度分布格局的影响。本研究共记录本地草本143种, 隶属于54科115属; 外来入侵草本16种, 隶属于7科13属。本地草本的物种丰富度和Shannon-Wiener指数随海拔上升显著增加(P < 0.05), 而外来入侵草本的物种丰富度和Shannon-Wiener指数随海拔升高呈下降趋势(P > 0.05)。本地草本和入侵草本的均匀度随海拔升高变化均不显著(P > 0.05)。在不同海拔梯度, 本地草本的Shannon-Wiener指数均显著高于外来入侵草本, 均匀度则相反。回归分析表明, 温度和降水量共解释了本地草本物种丰富度差异的41.6%, 其中温度解释率为21.4%, 降水量解释率为10.7%, 且温度的影响达到显著水平(P < 0.05)。温度和降水量对外来入侵草本的物种丰富度均无显著影响(P > 0.05), 单一因子解释率均较低, 分别为0.9%和0.4%。这些结果表明, 九万山国家级自然保护区外来入侵草本与本地草本物种多样性随海拔变化呈截然相反的空间分布格局, 且气候因素对两者分布格局的影响不同。外来入侵草本植物在海拔800-1,000 m地区具有相对多样的群落组成和较稳定的群落结构, 可能会威胁本地植物多样性, 建议加强此海拔区间外来入侵草本植物的管理。     

Elevational turnover in the composition of leaf miners and their interactions with host plants in Australian subtropical rainforest

Leaf miners are specialist herbivorous insects that are potentially vulnerable to environmental change because of their dependency on particular host plants. Little, however, is known about how climate affects the distribution of leaf miner communities and their interactions with host plants. Elevational gradients are useful tools for understanding how ecological communities respond to local clines in climate. Given that plant communities are known to undergo elevational turnover in response to changes in climatic conditions, we expect that leaf miner species will also change with elevation. We repeatedly hand collected leaf miners along three elevational gradients in subtropical rainforest in eastern Australia. Individual leaf miners were counted and identified to species, and their host plants were recorded. We tested if leaf miner species richness and the number of unique interactions among leaf miner and host plant species were affected by elevation. We also tested if the composition of leaf miner species and the composition of interactions between leaf miners and host plants showed a relationship with elevation. The rarefied number of unique leaf miner–host plant interactions significantly decreased with elevation, with a slight peak at approx. 700 m a.s.l., while neither rarefied or observed species richness (species density) of leaf miners nor observed numbers of unique interactions (interaction density) were significantly affected by elevation. The composition of leaf miner species and the composition of leaf miner–host plant interactions (occurrence of pairwise interactions) were significantly related to elevation. Elevational turnover in leaf miner species composition indicated that different species varied in their response to changes in biotic and/or abiotic conditions imposed by increasing elevation. Through our analyses, we identified four leaf miner species that may be locally vulnerable to climate change, as a result of their restricted elevational distribution and level of host specificity.