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Clarifying the relationship between distribution patterns of organisms and geological events is critical to understanding the impact of environmental changes on organismal evolution. Quercus sect. Heterobalanus is now distributed across the Himalaya-Hengduan Mountains ( HHM) and warm lowland in East China, yet how the distribution patterns of this group changed in response to the HHM uplift remains largely unknown. This study examines the effect of tectonic events in the HHM region on the oaks, providing a biological perspective on the geological history of this region. Fifty-six populations of Quercus sect. Heterobalanus were genotyped using four chloroplast DNA regions and nine nuclear simple sequence repeat loci to assess population structure and diversity, supplemented by molecular dating and ancestral area reconstructions. The underlying demographic dynamics were compared using ecological niche models of the species distributions during the last glacial maximum and the present. These analyses illustrate that Quercus sect. Heterobalanus diversified as the HHM uplifted and climatic cooling during the mid-Miocene, colonizing the cold habitats from warm broadleaf mixed forests. Lineages in cold highlands and warm lowlands have diverged as a consequence of local adaptation to diverging climates since the late Miocene. Our results suggest that continuous uplift of the HHM in the late Miocene to early Pliocene accompanied by simultaneous cooling triggered the differentiation of oaks. The biogeography of Quercus sect. Heterobalanus illuminates the geological events responsible for the modern-day HHM. [ABSTRACT FROM AUTHOR]

The Hengduan Mountains are the core of the Sino-Himalayan Floristic Region. Rosa soulieana Crép. is an important wild rose species that is widely distributed in the Hengduan Mountains. To provide better future utilization of this wild rose, and also to add some possible proof of the effect of geomorphological and ecological characteristics of the Hengduan Mountains on the current spatial distribution and genetic diversity of local species, the genetic diversity and genetic structure of 556 individuals from 37 populations of R. soulieana were studied using fluorescent amplified fragment length polymorphisms (AFLPs). R. soulieana showed a moderately high level of genetic diversity and a high level of genetic differentiation at the species level. The total percentage of polymorphic loci, total heterozygosity (Ht), Shannon index (I), and heterozygosity value within populations (Hs) were 97.8%, 0.253, 0.339, and 0.139, respectively. More than half of the total genetic variation (54.0%) occurred within populations, and the overall gene differentiation coefficient (Gst) was 0.451. The genetic differentiation among populations was positively and significantly correlated with geographic distance. The neighbor-joining cluster and the Bayesian analysis divided all the populations and individuals into 3 groups, and did not support the morphology based intraspecific varieties. The results confirmed that the ancient R. soulieana of the third group survived in northwestern Yunnan and Yalongjiang valley and then moved upnorth along the valley. The spatial distribution of the other two groups was the result of allopatric divergence due to long period of adaptation to the different climatic conditions of its distribution at either side of the Yalongjiang River. [ABSTRACT FROM AUTHOR]

Aim The Himalaya-Hengduan Mountain (HHM) biodiversity hotspot including the 'sky islands' of Southwest China harbour exceptional plant diversity and endemicity at subnival summits (most of them exceeding 4300 m a.s.l.). This study is the first using a comparative phylogeographical framework to gain insights into the temporal origin of this highly fragmented subnival flora, and the historical factors shaping its genetic architecture as exemplified by four perennial herbs. Location Himalaya-Hengduan Mountains, China. Methods Based on nuclear and/or chloroplast (cp) DNA sequences for each of the four studied species, we performed AMOVA and mismatch distributional analyses to assess molecular structure, diversity and demographic history in relation to current and last glacial distributions using ecological niche modelling (ENM). Time-calibrated phylogenetic reconstructions of cpDNA data were used to infer species-specific stem and crown ages. Results Our time estimates suggest that these species originated during the Late Pliocene or early-to-mid Pleistocene, whereas their onset of diversification generally falls into the mid-Pleistocene. All four species exhibited island-like population genetic structures, with all of them showing signatures of recent population growth and/or spatial expansion based on cpDNA. By contrast, ENM indicated that species broad-scale distributions remained fairly stable over the last glacial/post-glacial cycle. Main conclusions The temporal origin of the four subnival HHM species is likely associated with tectonic changes in the region, while their near-simultaneous onset of diversification during the 'Naynayxungla Glaciation' (0.72-0.50 Ma) could reflect initial population divergence through climate-induced habitat fragmentation. Despite a rather stable distributional history, geographical population isolation and localized range expansion/contractions likely resulted in significant genetic structure and differentiation over the last glacial/ post-glacial cycle. Overall, the present results are strongly indicative of shared evolutionary histories and phylogeographical structures among subnival plants from the 'sky island system' of the HHM region. [ABSTRACT FROM AUTHOR]

Quaternary climatic factors have played a significant role in population divergence and demography. Here we investigated the phylogeography of Osteomeles schwerinae, a dominant riparian plant species of the hot/warm-dry river valleys of the Hengduan Mountains (HDM), Qinling Mountains (QLM) and Yunnan-Guizhou Plateau (YGP). Three chloroplast DNA (cpDNA) regions (trnD-trnT, psbD-trnT, petL-psbE), one single copy nuclear gene (glyceraldehyde 3-phosphate dehydrogenase; G3pdh), and climatic data during the Last Interglacial (LIG; c. 120–140 ka), Last Glacial Maximum (LGM; c. 21 ka), and Current (c. 1950–2000) periods were used in this study. Six cpDNA haplotypes and 15 nuclear DNA (nDNA) haplotypes were identified in the 40 populations of O. schwerinae. Spatial Analysis of Molecular Variance, median-joining networks, and Bayesian phylogenetic trees based on the cpDNA and nDNA datasets, all suggested population divergence between the QLM and HDM-YGP regions. Our climatic analysis identified significant heterogeneity of the climatic factors in the QLM and HDM-YGP regions during the aforementioned three periods. The divergence times based on cpDNA and nDNA haplotypes were estimated to be 466.4–159.4 ka and 315.8–160.3 ka, respectively, which coincide with the time of the weakening of the Asian monsoons in these regions. In addition, unimodal pairwise mismatch distribution curves, expansion times, and Ecological Niche Modeling suggested a history of population expansion (rather than contraction) during the last glaciation. Interestingly, the expansion times were found being well consistent with the intensification of the Asian monsoons during this period. We inferred that the divergence between the two main lineages is probably caused by disruption of more continuous distribution because of weakening of monsoons/less precipitation, whilst subsequent intensification of the Asian monsoons during the last glaciation facilitated the expansion of O. schwerinae populations. [ABSTRACT FROM AUTHOR]

The Hengduan Mountains is the core of the Sino-Himalayan Floristic Region. Different views have been postulated from phylogeographic studies in understanding the historical biological events shaping present-day distributions and species diversities in this area. We analyzed the phylogeography of Rosa soulieana, a widely distributed rose species in dry and semi-dry valleys of the Hengduan Mountains of southwestern China using two cpDNA intergenic spacers. 9 cpDNA haplotypes were confirmed in 650 individuals from 39 populations, with the most dominant one existing in 36 populations. The relationship among the cpDNA haplotypes did not reflect the morphology-based intra-specific taxonomy of different varieties. There was very low cpDNA genetic diversity and relatively high population differentiation combined with lack of phylogeographic structure. The populations from Northwestern Yunnan, Southeastern Tibet and upper Yalongjiang River embraced most of the species genetic diversity, but with contribution of different historical effects. Northwestern Yunnan and upper Yalongjiang River appear to be the species refugia and centers of adaptive diversification, while Southeastern Tibet rather represents the species 'melting pot' in the Quaternary climate oscillations. [ABSTRACT FROM AUTHOR]

Questions What is the community-level consequence of biotic processes, in particular the importance of facilitation, in determining patterns of diversity in alpine plant communities of the Himalayan Hengduan Mountains? Does facilitation intensity change with environmental severity, and are these changes due to environmental severity or nurse trait effects? Location Eleven alpine plant communities dominated by cushion plants in the Himalayan Hengduan Mountains, within the mountain system of south-central China. Methods We determined plant species richness and abundance in habitats created by cushions and cushion-free areas, and assessed the cushion effects on species richness and abundance with rarefaction curves and the relative interaction index, respectively. We examined the relationship between cushion effects on diversity and habitat severity and tested if changes in the net cushion effects along the severity gradient were due to a change in the performance of species without cushions (environmental severity effect) and/or with cushions (nurse trait-mediated effect). Results The presence of cushion plants increased species richness and abundance in most studied plant communities. The net facilitation effect of cushions increased with increasing habitat severity, an effect that could be attributed mainly to a reduction in species richness in cushion-free areas with increasing severity. However, the changes in magnitude of facilitation of particular cushion species growing in high and low severity habitats could be attributed to both environmental severity and neighbour trait effects. Despite the overall positive effect of cushion plants on plant species richness, not all cushion species contributed to higher species richness. The deviation of individual cushion species from the general pattern of increasing facilitation with increasing severity indicated the species specificity of facilitation depends on the traits of nurse species. Conclusion The presence of cushion plants generally increased species richness in alpine plant communities of the Himalayan Hengduan Mountains, with the importance of the cushion effect increasing with habitat severity due to a buffering effect by cushions of the negative effect of habitat severity on species richness observed in cushion-free areas. This indicates the pivotal role of facilitative interactions among plant species in supporting high diversity in these severe environments. [ABSTRACT FROM AUTHOR]

Chromosome numbers and the morphology of members of 24 populations representing 15 species of the Sino-Himalayan endemic genus Cyananthus (Campanulaceae) were studied using karyological and numerical taxonomic techniques. The chromosome numbers of these taxa are 2 n = 10, 12, 14, 24, 28, and the basic chromosome numbers for the genus are x = 5, 6, 7. All species except C. inflatus Hook. f. & Thoms. and C. microphyllus Edgew. were examined for the first time and a new ploidy level (tetraploidy) is reported in the genus. Chromosome measurement data were analyzed using cluster analysis. The relationships between three sections within Cyananthus and evolutionary trends within the genus are discussed in the light of karyological data. The cytological data suggest that Cyananthus is a relatively primitive genus in Campanulaceae and there is significant division within the genus, and polyploidization may have played an important role in the chromosome evolution and speciation of Cyananthus in the Himalayan-Hengduan Mountains. [ABSTRACT FROM AUTHOR]

There have been few studies of the alpine subnival belt flora in the Hengduan Mountains (HM), which host remarkable biodiversity. To extend knowledge of this flora, we examined published florae, herbarium specimens, and field observations (and material) collected by both ourselves and others. In total, 942 seed plant species have been recorded in the belt, representing 168 genera and 48 families. Twenty-four large families (with ≥10 species) are present, represented by 873 species (92.68% of the total). These include Asteraceae, Saxifragaceae, and Brassicaceae (146, 82, and 71 species, respectively). There are also 27 large genera (represented by ≥10 species), collectively contributing 587 species (62.31%) to the flora, including Saxifraga, Corydalis, and Saussurea (75, 55, and 49 species, respectively). Areal elements represented by the highest numbers of genera are the North temperate, Sino-Himalaya, and Old World temperate elements (39.88%, 14.29%, and 12.50%, respectively), while the Hengduan Mountains, Sino-Himalaya, and Qinghai-Tibetan Plateau elements are most species-rich (355, 281, and 161 species, respectively), collectively accounting for 84.61% species of the known flora. Of these, 295 species (31.32%) are endemic to the Sino-Himalayan alpine subnival belt and 151 (16.03%) strictly endemic to the alpine subnival belt of the HM. These findings indicate that the flora is young, strongly differentiated, probably developed as a result of the plateau's uplift, and speciation has been accelerated by the harsh environment and strong heterogeneity of niches. [ABSTRACT FROM AUTHOR]

Plant sexual systems can be affected by various environmental factors, and thus show a range of morphological differences. The associations between plant sexual systems and morphology are unique, for example, the subnival belt of the Hengduan Mountains has harsh climatic conditions, but hosts remarkable biodiversity. However, these associations have received surprisingly little attention. We analyzed the associations in this region, and compared the pattern of sexual systems with that of other floras, and we also reviewed the breeding system of 36 alpine species. Information on plant sexual systems and these traits were mainly based on published information, herbarium collections, and field observations. The results showed that there was a high proportion of hermaphroditic species and a low proportion of dioecious and monoecious species. Hermaphroditism was strongly associated with showy floral displays, specialist entomophily, and perennial forbs. Dioecy was strongly associated with inconspicuous, pale-colored flowers, anemophily, and shrubs, whereas monoecy was strongly associated with inconspicuous, pale-colored flowers, generalist entomophily, and perennial grasses. Although no association was found between sexual systems and fruit type, plants with dry fruits (98.1%) were predominant in our study region. With respect to the plant breeding systems in the alpine region of southwest China, we predicted that pollination service might be particularly low; most of the plants lacked enough outcrossing pollinators. Our data revealed that 97.1% of the hermaphroditic species were self-compatible, and 88.2% of them showed autonomous or facilitated selfing. The plant sexual systems identified might serve to optimize the probability of pollination and thus ensure reproductive success. [ABSTRACT FROM AUTHOR]

To better understand the elevational pattern of phylogenetic structure shown by alpine taxa and the underlying causes, we analyzed the phylogenetic structure of each elevational belt of alpine plants in the Hengduan Mountains Region, measured by net related index ( NRI) and net nearest taxon index ( NTI). We found both the indices of phylogenetic diversity indicated that alpine plants tended to show phylogenetic overdispersion at low elevational belts, implying that the distribution of alpine plants in these belts was mainly determined by interspecific competition. Alpine plants at higher elevational belts tended to phylogenetic clustering indicated by NRI, and NTI revealed phylogenetic clustering at the belts between 4300 m and 5500 m, which presumably suggested environment filtering and rapid speciation. Above 5500 m, NTI indicated that the phylogenetic structure became random again, perhaps due to the low intensity of filtering and the large distances between plants at the top of the scree slopes. We concluded that phylogenetic structure was, indeed, influenced by the environmental filter, interspecies interaction, rapid speciation during the uplift of the Qinghai-Tibet Plateau, and distance between plants. [ABSTRACT FROM AUTHOR]

Phylogeographic studies on alpine plants endemic to the Hengduan Mountains of the southeastern Qinghai-Tibet Plateau are still limited in number. In this study, we used sequence variation of one nuclear gene ( ncpGS, which encodes the chloroplastic glutamine synthetase) and in two chloroplast DNA segments to investigate the phylogeographic structure and population demographic history of Cyananthus delavayi, a narrow-range species endemic to this region. We identified eight chlorotypes and 16 nuclear genotypes in a survey of 10 populations sampled throughout the range of the species. The results of both phylogenetic and network analyses suggested that the genealogical relationships of both chlorotypes and nuclear genotypes showed a clear geographical correlation. High total genetic diversity, low levels of within-population diversity, and strong population differentiation (chloroplast DNA: hT= 0.827, hS= 0.087, NST= 0.899, GST= 0.895; nuclear DNA: hT= 0.910, hS= 0.348, NST= 0719, GST= 0.618) were identified. Based on the mismatch distribution analyses, no evidence of recent demographic population expansion was found for this species. Nested clade analyses of both chlorotypes and nuclear genotypes indicated that restricted gene flow resulting from isolation by distance and allopatric fragmentation were likely to have been the major processes that shaped their present-day spatial distribution. Our dating of the genetic divergences between three major geographic lineages suggested that the largest glaciation of the early Quaternary developed in the Qinghai-Tibet Plateau and mountainous isolation may have together led to deep intraspecific vicariance within this species. [ABSTRACT FROM AUTHOR]

The article presents a study which investigated chromosome numbers and karyomorphology of eight species of Chaetoseri and two Stenoseries species from the Hengduan Mountain of Southwest China. A close relationship between the two genera is noted based on cytological data of chromosomal numbers and karyotypes. According to the study, polyploidization might played a minor role in their evolutionary speciation in the Hengduan Mountains region.

Several members of the subgenus Amphilaena (Saussurea; Asteraceae) growing at high elevations in the Himalayan region have semi-translucent bracts covering their inflorescences, the function of which is not fully understood. We investigated the thermal benefits of these bracts in Saussurea velutina in situ in the Hengduan Mountains, SW China, and compared the reflectance spectra and anatomical characters of bracts and rosette leaves in the laboratory. After removal of bracts in the field the average inflorescence daytime temperature was 2.5 °C colder than in control plants with intact bracts. Manually opened treatment inflorescences grew slower and set significantly fewer seeds than control inflorescences with bracts intact. The mesophyll of bracts is a single cell layer with a lower reflectance in the infrared range than rosette leaves. We show that the semi-translucent bracts in S. velutina function to ensure warmer temperatures for floral and seed development at high elevations in the Hengduan Mountains. [ABSTRACT FROM AUTHOR]

This paper aimed to explore the division of the southern and northern Hengduan Mountains based on gradients in species similarity and richness, and to analyze species richness in each sub-region. The Hengduan Mountain region was divided into nine latitudinal belts using one degree of latitude to define the belt after which distribution of seed plants within each latitudinal belt was recorded. Latitudinal patterns of species similarity were measured using the Jaccard similarity index for each pair of adjacent latitudinal belts. Non-metric multidimentional scaling (NMDS) was also used to analyze the similarity in species composition among the nine latitudinal belts. The latitudinal pattern of species similarity and the NMDS ordination both showed a great change in species composition across the 29°N latitudinal line, essentially dividing the Hengduan Mountain region into southern and northern sub-regions. Species richness, shown by the c-value of the species-area power function, and species-area ratio along a latitudinal gradient both showed a sharp decrease across the latitudinal belt from 29°0′ to 29°59′N. The southern sub-region occupied 40% of the total area of the Hengduan Mountain region, but contained more than 80% of all the seed plants in the region. The higher species richness and endemism in the southern sub-region showed it to be the core of the Hengduan biodiversity hotspot, a result not unexpected because of the greater extremes of topography and wider diversity of habitats in the southern portion. [ABSTRACT FROM AUTHOR]

Several members of the vascular plant genus Saussurea, which are found at elevations exceeding 5000 m, have dense layers of woolly trichomes on their leaves, bracts, and inflorescences, the function of which is not fully understood. Here we explore the thermal benefits of pubescence in Saussurea medusa, both in situ in the Chinese Hengduan Mountains and under controlled conditions. Mean daytime inflorescence temperature was 5.9 K above air temperature. Pubescence removal revealed that most of this warming is not related to pubescence but to radiant warming of the compact inflorescence itself (4.1 K warming in shaved plants, i.e. 1.8 K less). The effect of pubescence on nighttime radiative cooling was negligible. Our data indicated that the functional role of pubescence in these high-elevation plants cannot be attributed solely to producing warming tissues, but may also include other functions such as water repellency and reflection of short peaks of high radiation. These other functions must be specific to a small group of species, given that most other high elevation taxa do not exhibit this woolly character. [ABSTRACT FROM AUTHOR]

The karyotypes of six species of Solms-laubachia and one each of Parrya and Desideria (Brassicaceae) from the Himalayan-Hengduan Mountains (Tibet, Sichuan and Yunnan provinces) were investigated. Chromosome counts of S. xerophyta, S. lanata, S. platycarpa and D. baiogoinensis are reported for the first time and found to be diploid with 2n = 14. Diploid counts were also made of S. retropilosa, a species previously known to be a tetraploid. The karyotype formulae of D. baiogoinensis, S. eurycarpa, S. pulcherrima, S. retropilosa, S. xerophyta and S. platycarpa are 2n = 14 = 6 m + 6sm + 2st, whereas those of S. lanata and P. nudicaulis are 2n = 14 = 6 m + 2sm + 6st and 2n = 14 = 8 m + 4sm + 2st, respectively. Karyotypes and counts of S. eurycarpa and S. pulcherrima are in agreement with prior studies. The relationships within Solms-laubachia and among the three genera are discussed. Speciation within Solms-laubachia may have been the result of primarily intrachromosomal variation at the diploid level, and the genus probably evolved within the Himalayan-Hengduan Mountains instead of migrating from other regions. [ABSTRACT FROM AUTHOR]

The Hengduan Mountains (HDM) biodiversity hotspot exhibits exceptional alpine plant diversity. Here, we investigate factors driving intraspecific divergence within a HDM alpine species Salix brachista (Cushion willow), a common component of subnival assemblages. We produce a high-quality genome assembly for this species and characterize its genetic diversity, population structure and pattern of evolution by resequencing individuals collected across its distribution. We detect population divergence that has been shaped by a landscape of isolated sky island-like habitats displaying strong environmental heterogeneity across elevational gradients, combined with population size fluctuations that have occurred since approximately the late Miocene. These factors are likely important drivers of intraspecific divergence within Cushion willow and possibly other alpine plants with a similar distribution. Since intraspecific divergence is often the first step toward speciation, the same factors can be important contributors to the high alpine species diversity in the HDM. Exceptional alpine plant diversity exists in the Hengduan Mountains. Here, through genome assembly and population genomics studies, the authors find notable intraspecific divergence among Cushion willow populations isolated by the sky island-like habitats and consider it contributes to speciation and biodiversity. [ABSTRACT FROM AUTHOR]