Plant diversity and ecology on the Qinghai–Tibet Plateau

The Qinghai–Tibet Plateau (QTP) comprises a platform (sometimes called the Qinghai–Tibet Plateau sensu stricto), the Himalayas, and Hengduan Mountains (Liang et al.,2018; Mao et al.,2021). The latter two parts and adjacent highlands are also called the Pan-Himalaya. Numerous plants are distributed there with many endemic species, probably because of the high diverse landscapes created by continuous geological and climatic activities (Favre et al.,2015; Mao et al.,2021). As the well known biodiversity hotspot of the alpine plants in the world (Sun et al.,2017), many studies have been conducted on evolutionary origin and ecological adaptation of those species occurring in the QTP (e.g., Wen et al.,2014, 2019; Zhang et al.,2019). In the present special issue, we collected 15 related papers on this topic. Among them, two are invited reviews. Mao et al. (2021) provide a comprehensive review of evolutionary origin of species diversity on the QTP. Especially, they outlined major disputes and likely causes in this research topic, including circumscribing and naming the QTP, the QTP uplifts, dating of molecular phylogenetic trees, non-causal correlations between QTP uplifts and species diversification and the unified ice sheet. The authors also summarized genomic advancements related to high-altitude adaptation of both plants and animals. Tong et al. (2021) reviewed the reproductive strategies of animal-pollinated alpine plants on the QTP, involving pollination system, pollen limitation, self-pollination, and sexual system. In this region, 95.4% of animal-pollinated plants are pollinated by insects (i.e., bees, moths, butterflies, and flies) with only 4% by vertebrates (i.e., bats and birds). Self-pollination through self-compatibility shift from outcrossing has become an effective reproductive strategy to overcome pollen limitation in alpine plants. The other 13 research papers aimed to address origin and adaptation of alpine flora involving three major lines of evidence: genomics, ecology, and paleobotany. We hope that the collection of these papers will increase our understanding of the origin, speciation, and adaptation of alpine species on the QTP.     

Evolutionary history and underlying adaptation of alpine plants on the Qinghai–Tibet Plateau

As the highest and largest plateau in the world, the Qinghai–Tibet Plateau, with its numerous endemic species, is one of the important alpine biodiversity hotspots. Only recently have the evolutionary histories and underlying adaptations of these alpine plants become clear, through research mostly based on testable experiments and analyses. In this issue, we collected a total of 13 papers related to such aims. In addition, we selected a few published papers to highlight the major findings in the recent past. We also outlined the outlook and direction of future research.     

Phylogenomics and rapid diversification of the genus Eutrema on the Qinghai–Tibet Plateau and adjacent regions

Both geographic isolation and polyploidization are assumed to play an important role in driving species diversification. However, this is rarely illustrated through phylogenomic analyses. The genus Eutrema (Brassicaceae), which also includes the salt-resistant species, are distributed mainly in Asia with extensive species diversification in the Qinghai–Tibet Plateau (QTP) and adjacent regions. In this study, we revealed almost fully resolved backbone relationships of the genus with genome re-sequencing data for genomes of 168 individuals from 28 species. Phylogenetic analyses of both plastomes and single-copy nuclear genes from the whole genome recovered six well-supported clades with almost consistent relationships. The first two clades are mainly distributed in central China and central Asia, while the other four in the QTP and adjacent regions. All of them diversified within 12 million years. Within each clade, we recovered numerous conflicts in the interspecific relationships between nuclear and plastome phylogenies, likely suggesting hybridization and incomplete lineage sorting during species diversification. Our estimation of genome size and comparison of the number of the single-copy nuclear genes demonstrated frequent occurrences of polyploids in the genus. Except for an establishment of the backbone phylogeny, our phylogenomic analyses suggest that in addition to strong geographic isolation, polyploidization may have played an important role in species diversification of this genus.     

Survival and reproduction of plant species in the Qinghai–Tibet Plateau

The Qinghai–Tibet Plateau (QTP) is the highest and largest plateau in the world. It covers correspondingly wide geological, topographical, and climatic gradients, and thus hosts greater biodiversity than surrounding lowlands and other high elevation regions. Due to its extreme environmental and biological diversity, the QTP is an ideal region for studying adaptations of plant species under harsh environmental conditions at multiple evolutionary levels. Many recent ecological studies have revealed functions of distinctive morphological features of various plants in the region that improve their reproductive success. Examples include large and showy bracts, hairy inflorescences, and drooping flowers. Numerous other investigations have examined QTP plants' sexual systems, patterns of biomass allocations, and biotic interactions. This paper summarizes recent advances in understanding of morphological adaptations, plant–plant interactions, plant–pollinator interactions, floral color patterns, pollination adaptations, and resource allocation patterns of alpine plants of the QTP. The overall aim is to synthesize current knowledge of the general mechanisms of plant survival and reproduction in this fascinating region.     

Genetic diversity and population structure of Lamiophlomis rotata (Lamiaceae), an endemic species of Qinghai–Tibet Plateau

Lamiophlomis rotata (Lamiaceae), a perennial medicinal herb, is endemic to the Qinghai–Tibet Plateau. A total of 188 individuals from eight natural populations of L. rotata in Qinghai–Tibet Plateau (four from Tibet, two from Yunnan, and two from Qinghai) were analyzed using intersimple sequence repeats (ISSR) and randomly amplified polymorphic DNA (RAPD) techniques. Our results revealed that the level of genetic variation in L. rotata was relatively high (P = 94.85%, I = 0.440 ± 0.220, H _T = 0.289 ± 0.028). Three genetic groups corresponding to the three geographic regions were detected, suggesting significant geographic structure. Our results suggest that the highly structured geographic pattern found in L. rotata might represent diverging evolutionary processes associated with the uplifting of the Qinghai–Tibet Plateau and Quaternary climatic oscillations. These findings imply that as many populations as possible should be preserved in situ for the conservation of this species. Given their genetic variability and peripheral distribution, Qinghai and Yunnan populations should be assigned priority for conservation. Optimal harvesting strategies, domestication and tissue culture of L. rotata should be developed as soon as possible to guarantee its sustainable use.     

Phylogeography of an alpine aquatic herb Ranunculus bungei (Ranunculaceae) on the Qinghai–Tibet Plateau

Many phylogeographic studies of terrestrial plant species on the Qinghai–Tibet Plateau (QTP) have been carried out to elucidate the range shifts in response to climatic oscillations in the Quaternary. However, patterns of range shifts for aquatic plants following the climate change on the QTP are poorly understood. Here, we studied the historical range shifts of the aquatic herb Ranunculus bungei Steud. on the QTP using four chloroplast (cpDNA) non-coding spacers. We revealed low within-population genetic diversity (H_S = 0.052) and high interpopulation genetic differentiation (G_ST = 0.914; N_ST = 0.954). But the high population differentiation was not coupled with a distinct phylogeographic structure (N_ST > G_ST, P > 0.05). Phylogenetic analyses revealed two main cpDNA haplotype lineages and the split between these two lineages can be dated back to the late Tertiary (3.84–11.90 Ma). Two independent range expansions within the two intraspecific lineages at approximately 0.15–0.46 and 0.17–0.50 Ma were revealed. Our results suggested that R. bungei survived the Last Glacial Maximum and/or previous glacial periods on the QTP. Colonization or recolonization during the repeated range expansions may have replaced the early haplotypes and the pre-existing genetic structure and could explain the non-significant phylogeographical structure.     

Tree growth–climate relationships of Juniperus tibetica along an altitudinal gradient on the southern Tibetan Plateau

The southern Tibetan Plateau forms the ecotone between forest areas and alpine steppes and thus, tree growth is expect to react sensitive to climate variability in this semi-humid region. We sampled 328 increment cores from 169 trees at two study sites at four elevations along altitudinal transects from 4,000 to 4,500 m a.s.l. to evaluate elevation-dependent tree growth–climate relationships of Juniperus tibetica. Standard dendrochronological statistical parameters like mean inter-series correlation (Rbar), expressed population signal as well as signal-to-noise ratio is not significantly correlated to elevation. Mean segment lengths and average growth rates of the tree-ring series increase with elevation. Correlation and response function analysis with available climate data indicate that elevation has no significant effect on tree growth–climate relationships. Instead, local tree growth is mainly driven by common regional climatic signals as it is also indicated by significant correlations between all chronologies over their common period of A.D. 1550–2010. Moisture variability during April–June has the highest impact on tree growth, even close to the upper tree limit.     

Diversity and distribution of alkaliphilic psychrotolerant bacteria in the Qinghai–Tibet Plateau permafrost region

The Qinghai–Tibet Plateau represents a unique permafrost environment, being a result of high elevation caused by land uplift. And the urgency was that plateau permafrost was degrading rapidly under the current predicted climatic warming scenarios. Hence, the permafrost there was sampled to recover alkaliphilic bacteria populations. The viable bacteria on modified PYGV agar were varied between 10² and 10⁵ CFU/g of dry soil. Forty-eight strains were gained from 18 samples. Through amplified ribosomal DNA restriction analysis (ARDRA) and phylogenetic analyses, these isolates fell into three categories: high G + C gram positive bacteria (82.3%), low G + C gram positive bacteria (7.2%), and gram negative α-proteobacteria (10.5%). The strains could grow at pH values ranging from 6.5 to 10.5 with optimum pH in the range of 9–9.5. Their growth temperatures were below 37°C and the optima ranging from 10 to 15°C. All strains grew well when NaCl concentration was below 15%. These results indicate that there are populations of nonhalophilic alkaliphilic psychrotolerant bacteria within the permafrost of the Qinhai-Tibet plateau. The abilities of many of the strains to produce extracellular protease, amylase and cellulase suggest that they might be of potential value for biotechnological exploitation.     

Influences on mammals frequency of use of small bridges and culverts along the Qinghai–Tibet railway,China

Understanding the use of small bridges and culverts by wildlife to cross the Qinghai–Tibet railway will aid in the design of wildlife crossing structures for similar transportation infrastructure. From 2014 to 2016, 36 infrared cameras were placed inside 14 small bridges and 11 culverts along the Qinghai–Tibet railway to determine the structures’ effectiveness as wildlife passages. Thirteen species of mammals were found to use the small bridges and culverts to cross the railway. The crossing rates for all mammals were significantly higher for small bridges than for culverts. Tibetan antelope (Pantholops hodgsonii), Tibetan gazelle (Procapra picticaudata), kiang (Equus kiang), and wild yak (Bos mutus) preferred small bridges over culverts to cross the railway. In contrast, mountain weasel (Mustela altaica) and Asian badger (Meles leucurus) preferred culverts to cross the railway. The crossing rates of all mammals, particularly Tibetan gazelle and woolly hare, were positively influenced by structure width. Structure height had a positive influence on wild yak, but structure length had a negative influence on kiang. The distance to the highway had a positive influence on the crossing rates of all mammals, particularly wild yak and woolly hare. Human use of the structures had no influence on the crossings of most mammals except for common wolf. We suggest that road design schemes include large and open crossing structures to benefit most species with limitations on human activities near wildlife passages.     

High uncertainties detected in the wetlands distribution of the Qinghai–Tibet Plateau based on multisource data

Landscape and Ecological Engineering - Twenty wetland-related data products (including remote sensing datasets, compilation datasets and model simulation datasets) were collected to evaluate the...     

Genomic insights into speciation history and local adaptation of an alpine aspen in the Qinghai–Tibet Plateau and adjacent highlands

Natural selection serves as an important agent to drive and maintain interspecific divergence. Populus rotundifolia Griff. is an alpine aspen species that mainly occurs in the Qinghai–Tibet Plateau (QTP) and adjacent highlands, whereas its sister species, P. davidiana Dode, is distributed across southwest and central to northeast China in much lower altitude regions. In this study, we collected genome resequencing data of 53 P. rotundifolia and 42 P. davidiana individuals across their natural distribution regions. Our population genomic data suggest that the two species are well delimitated in the allopatric regions, but with hybrid zones in their adjacent region in the eastern QTP. Coalescent simulations suggest that P. rotundifolia diverged from P. davidiana in the middle Pleistocene with following continuous gene flow since divergence. In addition, we found numerous highly diverged genes with outlier signatures that are likely associated with high-altitude adaptation of these alpine aspens. Our finding indicate that Quaternary climatic changes and natural selection have greatly contributed to the origin and distinction maintenance of P. rotundifolia in the QTP.     

Alien dominance of the parasitoid wasp community along an elevation gradient on Hawai’i Island

Through intentional and accidental introduction, more than 100 species of alien Ichneumonidae and Braconidae (Hymenoptera) have become established in the Hawaiian Islands. The extent to which these parasitoid wasps have penetrated native wet forests was investigated over a 1,765 m elevation gradient on windward Hawai’i Island. For >1 year, malaise traps were used to continuously monitor parasitoid abundance and species richness in nine sites over three elevations. A total of 18,996 individuals from 16 subfamilies were collected. Overall, the fauna was dominated by aliens, with 44 of 58 species foreign to the Hawaiian Islands. Ichneumonidae was dominant over Braconidae in terms of both diversity and abundance, comprising 67.5% of individuals and 69.0% of species collected. Parasitoid abundance and species richness varied significantly with elevation: abundance was greater at mid and high elevations compared to low elevation while species richness increased with increasing elevation, with all three elevations differing significantly from each other. Nine species purposely introduced to control pest insects were found, but one braconid, Meteorus laphygmae, comprised 98.0% of this assemblage, or 28.3% of the entire fauna. Endemic species, primarily within the genera Spolas and Enicospilus, were collected almost exclusively at mid- and high-elevation sites, where they made up 22.1% and 36.0% of the total catch, respectively. Overall, 75.9% of species and 96.0% of individuals are inferred to parasitize Lepidoptera larvae and pupae. Our results support previous data indicating that alien parasitoids have deeply penetrated native forest habitats and may have substantial impacts on Hawaiian ecosystems.     

Biomass partitioning pattern of Rheum tanguticum on the Qinghai–Tibet Plateau was affected by water-related factors

To identify the patterns of aboveground biomass (M_A) and belowground biomass (M_B) partitioning of a tall perennial herbal plant, Rheum tanguticum Maxim. ex Balf. (R. tanguticum), we determined M_A, M_B, total biomass (M_T), and below- to aboveground biomass ratio (M_B/M_A) through three consecutive sampling campaigns during 2016–2018 on the Qinghai-Tibetan plateau. We then documented M_A, M_B, M_T, and M_B/M_A, and their log–log relationship with environmental factors using data from 47 sites. M_B/M_A showed a significant negative relationship with mean annual precipitation (MAP) and altitude but a positive relationship with mean annual temperature (MAT), soil total nitrogen content (TN), and soil humidity (SH). While M_T increased with altitude and decreased with MAT, TN, and SH, but the relationship between MAP and M_T was not significant. Reduced major axis analysis revealed a slope of the log–log relationship between M_A and M_B to be 1.16, supporting an allometric partitioning pattern of R. tanguticum. Furthermore, the scaling exponent revealed different changes to different environmental factors. Specifically, scaling exponent was sensitive to the gradient of MAP and SH, but not to the gradient of altitude, MAT, or TN. This indicated that the scaling exponent was affected by water availability.

     

Patterns of SOC and soil 13C and their relations to climatic factors and soil characteristics on the Qinghai–Tibetan Plateau

Background and aims

SOC inventory and soil δ¹³C were widely used to access the size of soil C pool and to indicate the dynamics of C input and output. The effects of climatic factors and soil physical characteristics and plant litter input on SOC inventory and soil δ¹³C were analyzed to better understand the dynamics of carbon cycling across ecosystems on the Qinghai-Tibetan Plateau.

Methods

Field investigation was carried out along the two transects with a total of 1,875 km in length and 200 km in width. Sixty-two soil profiles, distributed in forest, meadow, steppe, and cropland, were stratified sampled every 10 cm from 0 to 40 cm.

Results

Our result showed that SOC density in forest and meadows were much higher than in steppe and highland barley. In contrast, δ¹³C in forest and meadow were lower than in steppe and highland barley. Soil δ¹³C tended to enrich with increasing soil depth but SOC decline. SOC and δ¹³C (0–40 cm) were correlated with different climatic factors in different ecosystems, such that SOC correlated negatively with MAT in meadow and positively with MAP in steppe; δ¹³C correlated positively with MAT in meadow and steppe; and δ¹³C also tended to increase with increasing MAT in forest. Of the variation of SOC, 55.15 % was explained by MAP, pH and silt content and 4.63 % was explained by the interaction between MAT and pH across all the ecosystems except for the cropland. Meanwhile, SOC density explained 27.40 % of variation of soil δ¹³C.

Conclusions

It is suggested that different climatic factors controlled the size of the soil C pool in different ecosystems on the Tibetan Plateau. SOC density is a key contributor to the variation of soil δ¹³C.     

Trolox-equivalent antioxidant capacity and composition of five alpine plant species growing at different elevations on the Qinghai–Tibetan Plateau

Background: Alpine plants on the Qinghai–Tibetan Plateau are exposed to an extremely harsh environment, namely severe cold, strong ultraviolet radiation, hypoxia and low CO₂ partial pressure. These conditions are sources of oxidative stress, which increase in severity with increasing elevation.

Aims: To examine whether antioxidant capacity and chemical composition of alpine plants change with increasing elevation.

Methods: We measured the Trolox-equivalent antioxidant capacity (TEAC) and chemical composition of five alpine plant species at 3016, 3814 and 4621 m a.s.l.

Results: With increasing elevation: (1) the TEAC increased and total phenols and tannins tended to increase in two forb and two shrub species but not in a sedge species; (2) concentrations of protein and fat increased in all five plant species; (3) polyunsaturated fatty acids (PUFA) increased and (4) mineral concentrations tended to decrease, but trends were inconsistent.

Conclusions: We conclude that with increasing elevation, TEAC and total phenols and tannins increased which we interpreted as an adaptation to higher oxidative stress; and protein and fat contents increased to support high metabolic activity. The increase in PUFA and the trend for minerals to decrease with increasing elevation require further investigation.     

Range expansion,genetic differentiation,and phenotypic adaption of Hippophaë neurocarpa (Elaeagnaceae) on the Qinghai–Tibet Plateau

Plant species shift ranges in response to climate fluctuation over time. However, differentiation related to heterogeneity in space has been illustrated only rarely. Here we selected Hippophaë neurocarpa S. W. Liu & T. N. Ho (Elaeagnaceae), a shrub endemic to the Qinghai–Tibet Plateau, to exemplify this process. We characterized the genotypic, phenotypic, and climatic variations among 27 populations of this species sampled across its entire distribution. Genotype analyses revealed six highly differentiated groups that may have resulted from expansions in spatial range. Despite recent fluctuation in size, it is likely that most groups survived the last glacial maximum in situ. Instead of two previously described subspecies, we identified four morphotypes (stellate, peltate, and two additional types) that can be characterized based on leaf trichomes. This phenotypic subdivision is consistent with a climate gradient defined by temperature and precipitation, although similar phenotypes may derive from distinct genotypes. Collectively, we propose that the demographic history of H. neurocarpa is characteristic of an early spatial range expansion combined with a recent bottleneck, subsequently subdivided into multiple morphotypes following local adaption to heterogeneous climates.     

Fine-scale genetic diversity and genet dynamics of the fairy ring fungus Floccularia luteovirens on the Qinghai–Tibet plateau

The fungus Floccularia luteovirens is mainly distributed in the alpine meadows of the Qinghai–Tibet plateau. Its fruit bodies tend to form fairy rings with a visible stimulating zone. Our previous studies have investigated the large-scale genetic structure among wild populations of F. luteovirens, but the mechanisms underlying the current genotype distribution pattern remain unknown. The balance between sexual and asexual reproduction affects the establishment and structure of populations. Measuring genet size and density is an effective approach to investigating the reproduction strategies of this species. In the current study, 234 fruit bodies and 79 soil samples were collected from three sampling sites over 3y, revealing that F. luteovirens exhibits relatively large genets. Very few new genets were detected over the 3y, illustrating that this species relies more on vegetative growth and can persist for long periods underground as mycelia. Moreover, the underground genet data showed a close relationship with the above-ground genet data. Our study found limited fine-scale gene flow, contrary to our previous large-scale genetic study of F. luteovirens, the present study found limited fine-scale gene flow of this species. The commercial trade of F. luteovirens fruit bodies by humans probably enhances the gene flow over QTP.     

Better than nothing: Evolution of autonomous selfing under strong inbreeding depression in an alpine annual from the Qinghai–Tibet Plateau

In plants with infrequent pollinator services, the benefits of reproductive assurance could be eroded by severe ovule discounting and inbreeding depression (ID). However, it remains unclear how selfing evolves under complete pollinator failure and strong ID. We examined the mating system and ID under netting and robbing conditions in Comastoma pulmonarium (Turcz.) Toyok. (Gentianaceae), an alpine annual experiencing a high ratio of nectar robbery on the Qinghai–Tibet Plateau.Comastoma pulmonarium produced seeds via selfing at the study site. No pollinator was observed and thus the nectar was consumed exclusively by robbers. Inbreeding depression occurred in the life stages of seed mass and germination, and the cumulative ID was much higher than 0.5 under netting and robbing conditions. Overall, in comparison with netting, the magnitude of ID under robbing conditions tended to decrease. Our results suggested that selfing could assure reproduction for plants under complete pollinator failure and strong ID, supporting the “better than nothing” role of selfing and providing one of the few cases of the evolution of selfing under strong ID.     

Progressive changes in the structure and dynamics of the phytoplankton community along a pollution gradient in a lowland river — a multivariate approach

The phytoplankton of the River Lujan (Buenos Aires, Argentina) was studied for a period of 18 months, together with physical and chemical variables, in relation to a pollution gradient. 167 taxa were recorded within a seasonal succession characterized by dominance of diatoms with a brief summer green algae facies. A combination of several biotic indices and multivariate analysis was employed to assess the impact of pollution on the phytoplankton community. The biotic indices used were species diversity and richness, algal quotients (green algae/diatom ratio, Centrales/Pennales ratio) and the SD succession rate index. Multivariate procedures included cluster analysis and ordination by PCA of both species and samples, stepwise discriminant analysis and multiple discriminant analysis of variance (MANOVA). Results indicate that community dynamism is attenuated at the more polluted sites, concomitant with an increased predominance of a broad-tolerance algal assemblage, co-dominated by Cyclotella meneghiniana and Nitzschia stagnorum. The changes in the community structure and dynamics described herein involved alterations in the distribution and relative proportions of the algae, rather than modifications in the basic species composition. These changes may not be readily detectable by methods which over-simplify the ecological information, such as systems of indicator species and biotic indices, designed to assess the degree of pollution. The suitability of multivariate analysis and biotic indices in river phytoplankton studies is further discussed.