Patterns of species diversity and phylogenetic structure of vascular plants on the Qinghai‐Tibetan Plateau

Large-scale patterns of species richness and the underlying mechanisms regulating these patterns have long been the central issues in biogeography and macroecology. Phylogenetic community structure is a result of combined effects of contemporary ecological interactions, environmental filtering, and evolutionary history, and it links community ecology with biogeography and trait evolution. The Qinghai-Tibetan Plateau provides a good opportunity to test the influence of contemporary climate on shaping species richness because of its unique geological history, cold climate, and high biodiversity. In this study, based on high-resolution distributions of ˜9000 vascular plant species, we explored how species richness and phylogenetic structure of vascular plants correlate with climates on the highest (and species rich) plateau on the Earth. The results showed that most of the vascular plants were distributed on the eastern part of the plateau; there was a strong association between species richness and climate, even after the effects of habitat heterogeneity were controlled. However, the responses of richness to climate remarkably depended on life-forms. Richness of woody plants showed stronger climatic associations than that of herbaceous plants; energy and water availability together regulated richness pattern of woody plants; whereas water availability predominantly regulated richness pattern of herbaceous plants. The phylogenetic structure of vascular species clustered in most areas of the plateau, suggesting that rapid speciation and environment filtering dominated the assembly of communities on the plateau. We further propose that biodiversity conservation in this area should better take into account ecological features for different life-forms and phylogenetic lineages.     

Exploring the community phylogenetic structure along the slope aspect of subalpine meadows in the eastern Qinghai–Tibetan Plateau,China

Exploring the community assembly has been important for explaining the maintenance mechanisms of biodiversity and species coexistence, in that it is a central issue in community ecology. Here, we examined patterns of the community phylogenetic structure of the subalpine meadow plant community along the slope gradient in the Qinghai–Tibetan Plateau of China. We surveyed all species and constructed the phylogenetic tree of the plant community based on data from the Angiosperm Phylogeny Group III. We selected the net relative index (NRI) and evaluated the community phylogenetic structure along the five slope plants communities. We found that the phylogenetic structure varied from phylogenetic clustering to phylogenetic overdispersion with the slope aspect from north to south. In the north slope, the community phylogenetically cluster indicated that the limiting similarity played a leading role in the community assembly and the maintenance of biodiversity. Community phylogenetic overdispersion in the east, southeast, and south slopes indicated that habitat filtration was the driving force for community assembly. The NRI index of the northeast slope was close to zero, implying random dispersion. But it may be driven by the neutral process or limiting similarity, in that the community assembly process was the result of a combination of several ecological factors and thus required further study.     

Phylogeographic structure of Hippophae tibetana (Elaeagnaceae) highlights the highest microrefugia and the rapid uplift of the Qinghai‐Tibetan Plateau

The uplift of the Qinghai‐Tibetan Plateau (QTP) dramatically changed the topography and climate of Asia and affected the biodiversity of the plateau and its adjacent areas. However, the effects of the uplift on the dispersal, differentiation and adaptation of plants remain a puzzle when the date and processes of the uplift cannot be determined with certainty and the impacts of the Quaternary glaciations on plants on the QTP are unknown. To clarify the relationships among plants on the QTP with the plateau uplift and the Quaternary glaciations, the cpDNA trnT‐trnF regions of 891 individuals from 37 populations of Hippophae tibetana, endemic to the QTP, were sequenced in the present study. A total of 50 haplotypes were found and a strong phylogeographic structure was revealed (N_ST = 0.854, G_ST = 0.611, N_ST > G_ST, P < 0.01). The results show that three main lineages of the present populations of H. tibetana occupy the western, the middle, and the eastern geographical range, respectively, and their divergence time dates back to 3.15 Ma before present. Of 50 haplotypes, 33 (66%) are private haplotypes, which are restricted to single populations. These private haplotypes are scattered throughout the present geographical range of H. tibetana and originated from multiple differentiations in many lineages during more than 1.0 Ma period, strongly suggesting that multiple microrefugia of H. tibetana existed throughout the present geographical range during the last glacial maximum (LGM) and even earlier glaciations. Additionally, the average elevation of present populations is over 4500 m in the west and the equilibrium‐line of glaciers in the LGM was 500–300 m lower than present in the major interior part of the plateau suggesting that at most sites in the west, LGM microrefugia of H. tibetana may have been above 4000 m above sea level, the highest of all known refugia. Moreover, the divergence times among and within the three lineages and their distinct distributions as well as dispersal barriers support the theory of the recent and rapid uplift of the QTP. The rapid uplift of the plateau within the last 3.4 Ma and the associated environmental changes may have affected the dispersal and differentiation of H .tibetana and shaped its phylogeographic structure.     

青藏高原东部近林线紫果云杉原始林的群落结构与物种组成

原始云杉林是青藏高原东部林区近林线森林的重要类型之一,但关于其林分结构与多样性的科学认识到目前为止仍然是贫乏的.为了揭示云杉原始林的层次结构及其相互关系、维管植物组成特点,选择四川省壤塘县北部日柯沟近林线的紫果云杉(Picea purpurea)原始林为对象,调查了6个40 m×50 m样地的乔木层结构,并在样地内采用机械布点的方法设置180个小样方,调查了林下灌木层、草本层和苔藓层结构以及灌草层物种组成.结果显示:(1)云杉原始林为复层异龄林,乔木层分化明显,可划分为4个亚层;幼树密度不高但集中分布于林窗及其边缘(144±93株/ha);(2)林下灌木盖度与物种丰富度低,层片结构简单,呈明显的斑块状分布;而草本层与苔藓层发达,盖度与物种丰富度均较高;(3)6个样地共发现维管植物124种,隶属于30科68属.其中草本109种,灌木15种,以温带区系成分占优势,特有性缺乏;生活型以地面芽植物为主体(61.29%).综合分析表明,该林分为藓类云杉林,群落结构完整、稳定,自然更新能力强,与本区域内亚高山或高山林线森林具有相似的物种组成和多样性特点,但群落结构则明显不同.     

The role of the uplift of the Qinghai‐Tibetan Plateau for the evolution of Tibetan biotas

Biodiversity is unevenly distributed on Earth and hotspots of biodiversity are often associated with areas that have undergone orogenic activity during recent geological history (i.e. tens of millions of years). Understanding the underlying processes that have driven the accumulation of species in some areas and not in others may help guide prioritization in conservation and may facilitate forecasts on ecosystem services under future climate conditions. Consequently, the study of the origin and evolution of biodiversity in mountain systems has motivated growing scientific interest. Despite an increasing number of studies, the origin and evolution of diversity hotspots associated with the Qinghai‐Tibetan Plateau (QTP) remains poorly understood. We review literature related to the diversification of organisms linked to the uplift of the QTP. To promote hypothesis‐based research, we provide a geological and palaeoclimatic scenario for the region of the QTP and argue that further studies would benefit from providing a complete set of complementary analyses (molecular dating, biogeographic, and diversification rates analyses) to test for a link between organismic diversification and past geological and climatic changes in this region. In general, we found that the contribution of biological interchange between the QTP and other hotspots of biodiversity has not been sufficiently studied to date. Finally, we suggest that the biological consequences of the uplift of the QTP would be best understood using a meta‐analysis approach, encompassing studies on a variety of organisms (plants and animals) from diverse habitats (forests, meadows, rivers), and thermal belts (montane, subalpine, alpine, nival). Since the species diversity in the QTP region is better documented for some organismic groups than for others, we suggest that baseline taxonomic work should be promoted.     

An evaluation of government‐recommended stocking systems for sustaining pastoral businesses and ecosystems of the Alpine Meadows of the Qinghai‐Tibetan Plateau

China introduced the “Retire Livestock and Restore Grassland” policy in 2003. It was strengthened in 2011 by additional funding for on‐farm structures. On the Qinghai‐Tibetan Plateau (QTP), fences were erected, livestock excluded from degraded areas, rotational stocking introduced, nighttime shelters were built, forages grown, and seed sown. However, the effectiveness of these actions and their value to Tibetan herders has been questioned. We conducted a sheep stocking experiment for 5 years in an Alpine Meadow region of the QTP to evaluate stocking options recommended by Government. Cold and warm season stocking each at three rates (0, 8, and 16 sheep/ha) and continuous stocking at 0 and 4 sheep/ha were compared. We measured live weights of sheep, plant species richness and evenness, root biomass and carbon (C), nitrogen (N) and phosphorus (P) contents of the 0–10 cm of soil. We found that resting grassland from stocking during the warm season for later cold season stocking significantly reduced plant species richness and evenness and root biomass but not soil C, N, and P. During cold season stocking, live weights of sheep declined whether at a stocking rate of 8 or 16 per ha. In contrast, sheep continuously stocked on grassland at 4 per ha gained weight throughout both the warm and cold seasons and plant species richness and evenness were maintained. Warm season stocking at 8 and 16 sheep/ha increased plant species richness and root biomass but reduced plant species evenness. Resting these alpine grasslands from stocking in the warm season has adverse consequences for plant conservation. Fencing from stocking in the warm season is not justified by this study; all grassland should be judiciously stocked during the warm season to maintain plant species richness. Neither resting nor stocking during the cold season appears to have any adverse consequences but sheltering and in‐door feeding of sheep during the cold season may be more profitable than cold season stocking with use of open nighttime yards.     

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.     

The Pleistocene demography of an alpine juniper of the Qinghai‐Tibetan Plateau: tabula rasa,cryptic refugia or something else?

Aim Numerous palaeoecological and genetic studies have shown that different tree species responded in very different ways to Pleistocene climatic oscillations. Some were forced into small refugia far from their current range, while others were able to survive in small refugia close to, or even within, their current natural range. In this study we examine the Pleistocene demography of a juniper species (Juniperus przewalskii, Cupressaceae) from the Qinghai‐Tibetan Plateau. Location The Qinghai‐Tibetan Plateau (QTP). Methods Eight nuclear loci were sequenced in 141 individuals from 20 natural populations distributed across the entire natural range of J. przewalskii, and coalescent analysis was used to test demographic hypotheses. Results The overall nucleotide diversity in the sample was low (π_sil = 0.0029), with few rare alleles and pronounced population genetic structure (F_ST = 0.181). We detected a division previously found using chloroplast DNA markers: all segregating sites in populations from the central part of the QTP appear to be a subset of those found around the edge of the plateau, confirming the relatively young age of the former. In contrast to the middle Pleistocene bottlenecks detected in boreal tree species, the coalescent‐based analyses failed to reject the standard neutral model for the juniper species considered here. Main conclusions Juniperus przewalskii did not undergo marked changes in population sizes during the Pleistocene, although this species seems to have experienced recent, post‐glacial expansion. This finding is largely consistent with the limited number of previous studies on conifer species of the QTP, but contradicts findings of studies on boreal species. These findings have wide implications for understanding plant species’ responses to past climatic oscillations on the high‐elevation QTP.     

Species composition and structure characteristics of the major montane shrub communities in the west slope of Helan Mountains,Nei Mongol,China

灌丛是贺兰山最主要的植被组分, 但关于该地区灌丛群落的专门研究并不多见。为了解该地区灌丛资源的分布及生长状况, 该研究对内蒙古贺兰山国家级自然保护区内主要的山地灌丛群落进行样方调查, 通过分析样方数据, 量化描述了各群落的基本特征。结果表明: 贺兰山自然保护区沿海拔梯度主要有准噶尔栒子群系(Form. Cotoneaster soongoricus)、紫丁香群系(Form. Syringa oblata)、小叶金露梅群系(Form. Potentilla parvifolia)、虎榛子群系(Form. Ostryopsis davidiana)、小叶忍冬群系(Form. Lonicera microphylla)、银露梅群系(Form. Potentilla glabra)、山生柳群系(Form. Salix oritrepha)和鬼箭锦鸡儿群系(Form. Caragana jubata) 8种主要的山地灌丛群系。     

Comparative phylogeography of five avian species: implications for Pleistocene evolutionary history in the Qinghai‐Tibetan plateau

Pleistocene climate fluctuations have shaped the patterns of genetic diversity observed in extant species. In contrast to Europe and North America where the effects of recent glacial cycles on genetic diversity have been well studied, the genetic legacy of the Pleistocene for the Qinghai‐Tibetan (Tibetan) plateau, a region where glaciation was not synchronous with the North Hemisphere ice sheet maxima, remains poorly understood. Here, we compared the phylogeographical patterns of five avian species on the Qinghai‐Tibetan plateau by three mitochondrial DNA fragments: the Tibetan snow finch (Montifringilla adamsi), the Blanford’s snow finch (Pyrgilauda blanfordi), the horned lark (Eremophila alpestris), the twite (Carduelis flavirostris) and the black redstart (Phoenicurus ochruros). Our results revealed the three species mostly distributed on the platform region of the plateau that experienced population expansion following the retreat of the extensive glaciation period (0.5–0.175 Ma). These results are at odds with the results from avian species of Europe and North America, where population expansions occurred after Last Glacial Maximum (LGM, 0.023–0.018 Ma). A single refugium was identified in a restricted semi‐continuous area around the eastern margin of the plateau, instead of multiple independent refugia for European and North American species. For the other two species distributed on the edges of the plateau (the twite and black redstart), populations were maintained at stable levels. Edge areas are located on the eastern margin, which might have had little or no ice cover during the glaciation period. Thus, milder climate may have mitigated demographic stresses for edge species relative to the extremes experienced by platform counterparts, the present‐day ranges of which were heavily ice covered during the glaciation period. Finally, various behavioural and ecological characteristics, including dispersal capacities, habitat preference and altitude specificity along with evolutionary history might have helped to shape different phylogeographical structures appearing in these five species.     

Age‐mediation of tree‐growth responses to experimental warming in the northeastern Tibetan Plateau

The trajectory of tree‐growth response to climate warming may be related to attributes like tree age. However, age‐mediation of temperature sensitivity of tree growth has received little attention. This study aimed to determine how age affects tree growth in a future warmer world. In a 2‐year ecosystem warming experiment in the northeastern Tibetan Plateau of China, we explored the response of Qinghai spruce saplings at two life stages to two warming levels. Our results indicated a significant interaction between warming and age for sapling growth of Qinghai spruce. In high‐level warming scenario, the experiment increased growing season air temperatures by approximately 1.0°C and annual growing degree‐days by 38%. In response, warmed saplings lengthened the growing season by 10 days on average and increased the final shoot length to a maximum of 104% compared to control groups. Comparison of age classes revealed that old saplings exhibited significantly higher temperature sensitivity than young saplings. This performance may be caused by the differences in adaptive strategy to the asymmetric warming occurring during the whole day. Increased daytime temperature was expected to significantly enhance leaf photosynthesis, whereas lack of obvious nighttime warming would effectively restrict autotrophic respiration, thus resulting in the higher growth rate of old saplings compared with young saplings. Moreover, lack of nighttime warming rendered young saplings to be still in high stresses of freezing injury at low temperatures. These findings highlight the need for additional research on the effects of further climate anomalies on tree species during their ontogenetic processes.     

藏北高寒草甸群落结构与物种组成对增温与施氮的响应

气候变暖和氮沉降增加作为全球环境问题,将严重影响陆地生态系统的结构与功能.研究发现,近几十年来青藏高原增温显著,其中冬季升温最明显.而已有的研究更多关注全年增温,对冬季增温研究较少.本文基于高寒草甸地区增温和氮素添加影响研究的不足,在青藏高原高寒草甸区开展模拟增温和氮添加试验,研究长期增温与氮添加对高寒草甸群落结构与物种组成的影响.试验布设于2010年7月,地点在西藏当雄高寒草甸区,共有3种增温方式:对照、全年增温、冬季增温;每种增温处理下设置5个氮素添加梯度:0、10、20、40、80 kg N·hm^-2·a^-1,系统研究气候变暖与氮添加对高寒草甸生态系统群落结构与物种组成的影响.结果表明: 2012—2014年,增温与施氮处理均显著影响群落总盖度:全年增温处理降低了群落总盖度;在不施氮处理下,冬季增温降低了群落盖度,但在施氮处理下,随着氮剂量的提高群落盖度逐渐升高.增温与施氮对不同功能群植物的影响不同,增温处理降低了禾草与莎草植物盖度,而施氮提高了禾草植物盖度.相关分析表明,植被群落总盖度与生长旺盛期土壤含水量呈正相关关系,推测在降雨较少的季节增温导致的土壤含水量降低是群落盖度降低的主要原因.半干旱区高寒草甸土壤水分主要受降雨的调控,未来气候变化情景下,降雨时空格局的改变会显著影响植被群落盖度及组成,且大气氮沉降的增加对植被群落的影响也依赖于降雨条件的变化.     

贺兰山西坡主要山地灌丛群落的基本特征

灌丛是贺兰山最主要的植被组分, 但关于该地区灌丛群落的专门研究并不多见。为了解该地区灌丛资源的分布及生长状况, 该研究对内蒙古贺兰山国家级自然保护区内主要的山地灌丛群落进行样方调查, 通过分析样方数据, 量化描述了各群落的基本特征。结果表明: 贺兰山自然保护区沿海拔梯度主要有准噶尔栒子群系(Form. Cotoneaster soongoricus)、紫丁香群系(Form. Syringa oblata)、小叶金露梅群系(Form. Potentilla parvifolia)、虎榛子群系(Form. Ostryopsis davidiana)、小叶忍冬群系(Form. Lonicera microphylla)、银露梅群系(Form. Potentilla glabra)、山生柳群系(Form. Salix oritrepha)和鬼箭锦鸡儿群系(Form. Caragana jubata) 8种主要的山地灌丛群系。     

Mountains too high and valleys too deep drive population structuring and demographics in a Qinghai–Tibetan Plateau frog Nanorana pleskei (Dicroglossidae)

Pleistocene glacial–interglacial climatic oscillations greatly shaped the current genetic structure of many species. However, geographic features may influence the impact of climatic cycling. Distinct geographic and environmental characters between northern and southern parts of the eastern Qinghai–Tibetan Plateau (EQTP) facilitate explorations into the impacts of geographic features on species. The northern parts of EQTP contain large areas of marsh, and the environment is rather homogeneous. In contrast, the southern EQTP harbors complex alpine valleys and a much more heterogeneous setting. We evaluate DNA sequence variation from both the mitochondrial and nuclear genomes in Nanorana pleskei, a species endemic to the EQTP. Hypothesis testing on the evolutionary history of N. pleskei indicates that northern populations can disperse freely, but alpine valleys isolate southern populations. Demographic histories between northern and southern populations also differ. Northern populations appear to have experienced population expansions, while southern frogs exhibit a far more stable demographic history. By combining climatic analyses and species' distribution models, our study suggests that geographic and environmental features drive the differences between the northern and southern EQTP.     

Phylogeography of the genus Dasiphora (Rosaceae) in the Qinghai‐Tibetan Plateau: divergence blurred by expansion

Plateau uprisings and climatic oscillations are considered to have caused extensive allopatric divergences that account for the rich species diversity of the Qinghai‐Tibetan Plateau (QTP). However, secondary contact during range shifts in the Quaternary glacial cycles or inter‐uplift stages may have restored the gene flow between species and so counteracted these divergences, particularly in rapidly‐adapting dominant elements. We tested this hypothesis by determining the phylogeographical history of Dasiphora (Rosaceae), a genus of two species that are widely distributed on the QTP and co‐exist in numerous localities. We sequenced two chloroplast DNA fragments (rbcL, trnT‐L) for 559 individuals from 87 populations. Bayesian methods were used to identify phylogenetic relationships and to estimate divergence times. Demographic histories were inferred using neutrality tests, mismatch distribution analysis, and coalescent simulation. A total of 112 haplotypes that clustered into three major groups were identified. The formation of these groups and their subgroups was dated to between the Pliocene and the late Pleistocene. In addition, we found that some groups underwent multiple extensive expansions. Species‐specific haplotypes were identified for each species, although these haplotypes phylogenetically intermixed. These results suggest that recent plateau uplifts and climatic oscillations might have caused the deep divergences observed within this genus. However, later range expansions probably blurred these divergences and possible species boundaries. Our results shed new light on the complex evolutionary history of the QTP alpine plants. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 777–788.     

Ecological traits influence the phylogenetic structure of bird species co‐occurrences worldwide

The extent to which species’ ecological and phylogenetic relatedness shape their co‐occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co‐occurrence patterns are linked – after accounting for regional effects – to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co‐occurrences of carnivorous, migratory and cold‐climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life‐history traits beyond the climatic niche. They further offer a novel species‐oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.