ToTE: A global database on trees of the treeline ecotone.

Globally, treelines form a transition zone between tree-dominated forest downslope and treeless alpine vegetation upslope. Treelines represent the highest boundary of "tree" life form in high-elevation mountains and at high latitudes. Recently, treelines have been shifting upslope in response to climate warming, so it has become important to understand global tree diversity and treeline distributions. However, to the best of our knowledge, no global database on tree flora of treelines exists, which limits our capacity to undertake macroecological analyses. Here, for the first time, we present a global data set on the trees of the treeline ecotone, supported by an online ToTE database. We synthesized the database from 1202 studies published over the last 60 years (1962 to 2022) following the Preferred Reporting Items in Systematic Reviews and Meta-Analysis (PRISMA) protocol. We classified the tree species in the database into three categories: treeline tree (TL) species, near to treeline (NTL) tree species, and tree species with an upper montane range limit (TUMR). The ToTE Version-1 presents a total of 208 tree taxa, including 189 species, five subspecies, and 14 varieties, belonging to 54 genera and 26 families distributed across 34 mountain regions worldwide that either grow exactly at the treeline or have a range limit below the treeline. Of the total taxa, 155, 14, and 39 belong to TL, NTL, and TUMR, respectively. Genera such as Abies, Picea, Pinus, Larix, and Juniperus are more represented in the treeline tree category. On the other hand, Acer, Prunus, Populus, and Quercus have more representatives in the near to treeline category, whereas Erica, Nothofagus, and Polylepis contribute more tree species with an upper montane range limit. Furthermore, families such as Rosaceae and Pinaceae include trees that occur both at the treeline and with an upper montane range limit, whereas Sapindaceae includes trees that occur exclusively near to treeline. Our database also includes information on the global distribution patterns of treeline tree species richness across mountains and biomes. The mountains with the highest number of tree species are the Andes (39) followed by the Himalaya (37). Close to 67% of tree species show restricted distributions in different mountains, with the highest endemism in the Andes and the Himalaya. In terms of tree species distribution, Pinus sylvestris was widespread, with a distribution across nine mountain regions, followed by Picea glauca and Fagus sylvatica, both distributed across five mountain regions. In terms of species' distribution across biomes, the temperate biome harbors the highest treeline tree species richness (152 species), which may reflect the fact that the majority of studies are available from the temperate regions of the world. The remaining 56 species are distributed within five other biomes, with the least in dry tropical and subarctic (four species each). Furthermore, currently 40 treeline tree species fall under different International Union for Conservation of Nature threat categories. We anticipate that our database will help advance research on macroecological, biogeographic, evolutionary, climate-change, and conservation aspects of the treeline on a global scale. The data are released under a Creative Commons Attribution 4.0 international license. Please cite this data paper when the data are reused.

Different sets of traits determine transition of alien species along the invasion continuum.

Invasive alien species are currently considered as one of the dominant drivers of global environmental change. Till now, the majority of studies have focused on single or a few traits of alien species that facilitate their invasion. Also inclusion of all the traits which determine the transition of aliens along the different stages of invasion continuum (casual, naturalised and invasive) has remained largely overlooked. In this study, we collected a comprehensive trait dataset on 144 alien plant species of Kashmir Himalaya - a global biodiversity hotspot region. To test which traits of alien species, individually or in combination along with anthropogenic factors, determine their transition along the invasion continuum, we employed chi-square tests, boosted regression trees and phylogenetic methods. We found the perennial life span, longer residence time, greater number of introduced regions, and better seed dispersal mechanism were critical in determining the transition from casual to naturalised. The herbaceous growth form, therophyte Raunkiaer life-form, annual life span, achene fruit, longer residence time and broader introduced range were the species' traits determining transition from naturalised to invasive. Aliens introduced as ornamentals have more propensity to become naturalised; whereas aliens introduced unintentionally show overrepresentation at the invasive stage. Phylogeny alone showed mixed results indicating both clustering and dispersion; however, in combination with other traits, it plays a significant role in determining the stage of invasion. Overall, our study disentangles the individual and interactive roles of multiple traits that determine the transition of alien species' along the invasion continuum. Further, we foresee the potential applicability of our findings in designing robust invasion risk analysis protocols and stage-specific invasion management strategies in this Himalayan region, with learnings for elsewhere in the world.

Plant phenology shifts under climate warming: a systematic review of recent scientific literature.

Climate warming-driven temporal shifts in phenology are widely recognised as the foremost footprint of global environmental change. In this regard, concerted research efforts are being made worldwide to monitor and assess the plant phenological responses to climate warming across species, ecosystems and seasons. Here, we present a global synthesis of the recent scientific literature to assess the progress made in this area of research. To achieve this, we conducted a systematic review by following PRISMA protocol, which involved rigorous screening of 9476 studies on the topic and finally selected 215 studies for data extraction. The results revealed that woody species, natural ecosystems and plant phenological responses in spring season have been predominantly studied, with the herbaceous species, agricultural ecosystems and other seasons grossly understudied. Majority of the studies reported phenological advancement (i.e., preponement) in spring, followed by also advancement in summer but delay in autumn. Methodology-wise, nearly two -third of the studies have employed direct observational approach, followed by herbarium-based and experimental approaches, with the latter covering least temporal depth. We found a steady increase in research on the topic over the last decade with a sharp increase since 2014. The global country-wide scientific output map highlights the huge geographical gaps in this area of research, particularly in the biodiversity-rich tropical regions of the developing world. Based on the findings of this global synthesis, we identify the current knowledge gaps and suggest future directions for this emerging area of research in an increasingly warming world.

Functional trait diversity and aboveground biomass of herbaceous vegetation in temperate forests of Kashmir Himalaya.

Studying functional trait diversity can provide crucial clues about the adaptive survival strategies of regional plant species pool. Despite large-scale trait datasets available worldwide, the plant trait data from many biodiversity hotpot regions, like the Himalaya is still scarce. In this study, we aimed to investigate the plant functional traits and aboveground biomass of understory herbaceous vegetation in temperate forests of Overa-Aru wildlife sanctuary of Kashmir Himalaya. We also investigate how these functional traits correlate and what is the magnitude of trait-biomass relationship across the herbaceous species pool. For this, we conducted field sampling and measured leaf functional traits and aboveground biomass of 38 plant species in the study region during peak growing season (July-August) in the year 2021. The results revealed a significant interspecific trait variability among the species studied. We observed a high variability in leaf size and type spectra of the species, with nanophyll and simple leaf lamina, respectively, the most common types among the species studied. The correlation analysis revealed that plant height was positively correlated with aboveground biomass. The variation partitioning analysis revealed that the plant height explained the maximum fraction of variation in aboveground biomass, while least by specific leaf area. Overall, the findings from the present study provide useful insights in understanding trait-trait relationship and trait-environment interaction at the regional scale and can also help in recognizing adaptive functional traits of plant species that determine plant survival under the changing climate in this Himalayan region.

Diversity and utilization patterns of fodder resources in a Himalayan protected area.

In the Himalaya, collection of fodder from protected areas by local communities for sustenance of their livestock is an integral part of mountain farming systems. Here, we assess the diversity and utilization patterns of fodder resources collected by local communities from Overa-Aru wildlife sanctuary in Kashmir Himalaya. A multi-stage random sampling was employed to select sample villages (5) and households (81) for the survey. Data were collected through personal interviews of household heads administering an interview schedule and then analyzed using descriptive statistics and linear regression model. The results revealed that 74 fodder species were collected by the local communities from the sanctuary. The diversity and magnitude of fodder resource utilization varied across the study area. The fodder collection and utilization on per day and annual basis ranged from 25 to 90 and 2760 to 13,770 kg/household, respectively. Across the surveyed villages, the fodder was mainly collected by the females (60%). A positive but non-significant relationship was found between the number of persons collecting fodder and quantity of fodder collection/household/day, while a positive and significant relationship was found between the herd size and quantity of fodder collection/household/day. We also found a negative but non-significant relationship between the education status of respondents and the number of livestock holding. Based on our findings, we suggest policy and management interventions such as regulating livestock grazing, promoting plantation of frequently collected fodder species, and better use of agriculture byproducts to guide the sustainable management of fodder resources in this Himalayan protected area, with learning for elsewhere.

Sustaining Traditional Ethnomedicinal Knowledge and Protected Areas in Synergy: A Case Study of Overa-Aru Wildlife Sanctuary in Kashmir Himalaya.

The ethnic communities living near protected areas are critically dependent on medicinal plants for their healthcare. Although several ethnomedicinal studies are available from the Himalaya, quantitative studies focusing on protected areas in this region are largely overlooked. In this study, we aimed to document the ethnomedicinal plants of the Overa-Aru wildlife sanctuary in Kashmir Himalaya. We conducted a systematic field survey in the study area during 2020 - 2021 to collect primary data from 110 participants through semi-structured interviews and questionnaires. The data was analyzed using quantitative indices and Pearson's correlation. We recorded 64 plant species used against 8 human disease categories, including some first-time reports from the region. Asteraceae and Lamiaceae were the most dominant families and herbs were the dominant growth form. The most common plant part used were leaves, and the main method of preparation was decoction. Plant species with the highest relative frequency of citation was Achillea millifolium (0.83). Informant consensus factor for each disease category ranged from 0.94 to 0.97. We found a strong positive correlation between participants' age and citations (r = 0.85), but a negative correlation between the participants' education status and citations (r = - 0.11). Our results revealed that the younger generation possesses the least ethnomedicinal information. We recommend that the species with high use values and fidelity level merit priority in phytochemical and pharmaceutical studies. Our findings highlight that protected areas, besides their prime importance in biodiversity conservation, play a pivotal role in providing the healthcare provisioning ecosystem services to the ethnic communities.

From forest floor to tree top: Partitioning of biomass and carbon stock in multiple strata of forest vegetation in Western Himalaya.

The foremost role of forest vegetation in storing biomass and carbon (C) stock constitutes one of the main nature-based solutions to mitigate climate change. In this study, we aimed to quantify biomass and C stock partitioning in multiple vegetation strata (tree, shrub, herb, and ground floor layers) of major forest types in Jammu and Kashmir, Western Himalaya, India. We used a stratified random cluster sampling strategy to collect field data in 96 forest stands of 12 forest types (altitudinal range: 350 to 3450 m) in the study region. We evaluated the degree to which the carbon stock of the entire ecosystem was dependent on the multiple vegetation strata using the Pearson method. Across all the forest types, the average total ecosystem-level biomass was estimated to be 181.95 Mgha-1 (range: 60.64-528.98). Forest strata-wise, the maximum biomass of 172.92 Mgha-1 (range: 50.64-514.97) was found in the tree vegetation, followed by 5.58 Mgha-1 (range: 2.59-8.93) in understory vegetation (shrubs and herbaceous), and 3.44 Mgha-1 (range: 0.97 and 9.14) in the forest floor. The total ecosystem-level biomass showed a peak at mid-elevation coniferous forest types, whereas the lowest was observed in low-elevation broad leaved forest types. At the ecosystem-level, on average, the understory contributed 3% and the forest floor 2% to the total C stock across the forest types. The shrub layer contributed up to 80% of total understory C, with the herbaceous layer accounting for the remaining 20%. The ordination analysis clearly shows that anthropogenic and environmental variables significantly (p ≤ 0.002) influence the forest types' C stock in the region. Our findings have significant implications for conserving natural forest ecosystems and restoring degraded forest  landscapes in this Himalayan region, which in turn can lead to better carbon sequestration and climate mitigation outcomes.

Plant invasion shifts soil microbiome and physico-chemical attributes along an elevational gradient in Kashmir Himalaya.

Soil microbial communities, being situated at the interface of aboveground plant and belowground soil systems, can play a pivotal role in determining ecosystem response to the drivers of global environmental change, including invasive species. In mountains, invasive plants occurring along elevational gradients offer a unique natural experimental system to investigate the impact of invasions in determining patterns and relationships of soil microbial diversity and nutrient pools at much shorter spatial distances. Here, we studied the impact of a global plant invader, Leucanthemum vulgare, on the diversity of soil microbiome and physico-chemical attributes along an elevational gradient (1760-2880 m) in Kashmir Himalaya. We used Illumina MiSeq platform to characterize the soil microbiome in pair-wise invaded and uninvaded plots at four different sites along the gradient. We found a total of 1959 bacterial operational taxonomic units (OTUs) belonging to 152 species, and a relatively higher number of 2475 fungal OTUs belonging to 589 species. The α-diversity of soil microbiome showed a gradual increase from low to high elevation and differed significantly (p < 0.05) between the invaded and uninvaded plots. The ß-diversity revealed distinct microbiome clustering among the sampling sites. Plant invasion also altered soil physico-chemical attributes along the elevational gradient. Overall, our findings suggest that the L. vulgare-induced shifts in soil microbiome and nutrient pools may be a belowground self-reinforced mechanism to facilitate its successful invasion along the elevational gradient. Our study provides new insights into invasive plant-microbe relationships with wide implications for climate warming-driven elevational range shifts in mountains.

Alien flora causes biotic homogenization in the biodiversity hotspot regions of India.

Biotic homogenization by invasive alien species is one of the dominant drivers of global environmental change. However, little is known about the patterns of biotic homogenization in global biodiversity hotspots. Here, we fill this knowledge gap by studying the patterns of biotic homogenization and associated geographic and climatic correlates in Indian Himalayan Region (IHR). For this, we use a novel biodiversity database comprising 10,685 native and 771 alien plant species across 12 provinces of the IHR. The database was assembled by screening 295 and 141 studies published from 1934 to 2022 for natives and aliens, respectively. Our results revealed that each native species on average was distributed among 2.8 provinces, whereas the alien species in 3.6 provinces, thereby indicating wider distribution range of alien species in the IHR. The Jaccard's similarity index between the provinces was higher for alien species (mean = 0.29) as compared to natives (mean = 0.16). Addition of alien species pool has homogenized most of the provincial pairwise floras (89.4 %) across the IHR, with greater dissimilarity in their native floras. Our results revealed that the alien species have strong homogenization effect on the provincial floras, regardless of their differences in geographic and climatic distances. The biogeographic patterns of alien and native species richness in the IHR were better explained by a different set of climatic variables, the former by precipitation of driest month and the latter by annual mean temperature. Our study contributes to better understanding of the patterns of biotic homogenization in the IHR and its geographic and climatic correlates. Looking ahead, in an era of Anthropocene, we discuss the wide implications of our findings in guiding biodiversity conservation and ecosystem restoration in global hotspot regions.

Unravelling diversity, drivers, and indicators of soil microbiome of Trillium govanianum, an endangered plant species of the Himalaya.

In an era of global environmental change, conservation of threatened biodiversity and ecosystem restoration are formidable ecological challenges. The forest understory strata and the belowground soil environment including rhizospheric microbial communities, which are crucial for ecosystem functioning and overall forest biodiversity maintenance, have remained understudied. Here, we investigate the soil microbiome of Trillium govanianum - an endangered Himalayan Forest herb, to unravel the underground diversity, drivers, and potential indicators of the microbial community. We collected rhizospheric and bulk soil samples for microbiome and physicochemical analysis at three sites along an elevation gradient (2500-3300 m) in Kashmir Himalaya. Amplicon sequencing of 16 S rRNA and ITS was used to identify the bacterial and fungal soil microorganisms. We found significant differences in the structure and diversity of microbial community (bacterial and fungal) between the rhizosphere and bulk soil along the altitudinal gradient, and noticeable shifts in the nutrient level in dominant microbial phyla associated with T. govanianum. A significant difference between soil physicochemical parameters and increasing altitude suggests that microbial community structure is determined by altitude and soil type. Similarly, the microbial communities showed a significant (P < 0.05) correlation with soil physicochemical variables along the altitudinal gradient. The moisture content in bacterial and total organic carbon in fungal communities showed the most substantial impact on the physiochemical drivers. We also identify potential bacterial and fungal plant growth promoter indicator species in the soil microbiome of T. govanianum. Overall, our findings provide novel research insights that can be pivotal in designing integrated species recovery programs and long-term restoration plans for T. govanianum, with learnings for biodiversity conservation elsewhere.

Climate warming-driven phenological shifts are species-specific in woody plants: evidence from twig experiment in Kashmir Himalaya.

Experimental evidences in support of climate warming-driven phenological shifts are still scarce, particularly from the developing world. Here, we investigated the effect of experimental warming on flowering phenology of selected woody plants in Kashmir Himalaya. We selected the twigs of four congeneric pairs of temperate woody species (Prunus, Populus, Ulmus, Viburnum)-typical spring-flowering plants in the region. Using randomised block design, we monitored these winter dormant twigs in controlled growth chambers to study the effect of different temperature regimes (9, 17, 20 and 23 °C) and species identity on the patterns of phenological shifts. We observed a significant phenological shift in all the species showing preponement in the first flower out and senescence phases ranging from 0.56 to 3.0 and 0.77 to 4.04 days per degree increase in temperature, respectively. The duration of flowering phase in all the species showed a corresponding decrease along the gradient of increasing temperature, which was more driven by preponement of the flower senescence than the start of flowering. The patterns of phenological shifts were highly species-specific, and the magnitude of these shifts significantly varied in all the four pairs of congeneric species despite their phylogenetic similarity. Our study provides experimental support to the previous long-term observation and herbarium-based studies showing that the patterns of phenological shifts in response to global climate warming are likely to vary between species, even those belonging to same evolutionary stock. Our findings highlight that a one-size-fits-all strategy to manage the likely impacts of climate warming-induced phenological shifts will seldom succeed, and should instead be designed for the specific phenological responses of species and regions.

Patterns of floristic and functional diversity in two treeline ecotone sites of Kashmir Himalaya.

Globally, the treelines at higher elevations in mountains are reported to be advancing up-slope in response to recent climate warming. However, little is known about the treeline advancement in the Himalaya due to paucity of baseline vegetation data with which to compare, thus making their assessment and monitoring challenging. To fill this knowledge gap, the present study documented floristic and functional diversity of two treeline ecotone sites in Kashmir Himalaya. At each site, we conducted field sampling by laying five 20-m2 plots, with one at the highest limit (T0 plot), two plots below and two above the treeline and two nested subplots of 5-m2 for shrubs and five 1-m2 for herbs in each plot. We recorded 97 plant species belonging to 33 families from the two sites. We observed a considerable difference in species composition and distribution along the treeline ecotone. Majority of the species reported were perennial herbs. We observed a significant association of growth forms with the particular plots along the treeline ecotone. At both the sites, we recorded highest species richness at the T0 plot which was correlated well with the functional traits, thus indicating convergence of floristic and functional diversity at this transition zone. Interestingly, the T0 plot at both the sites showed maximum overlap of species with the plots above and below the treeline. In an era of climate warming, our study provides crucial baseline data that will facilitate assessment and monitoring of the Himalayan treelines.

Predicting shifts in distribution range and niche breadth of plant species in contrasting arid environments under climate change.

Arid environments face extreme risk from contemporary climate change; therefore, predicting the shifts in species distribution range and niche breadth in these environments assumes urgent research priority. Here we report the potential distribution and predict future distribution range of two model plant species typically representing contrasting environments across Asia and Africa: hot-arid Ephedra foliata and cold-arid E. gerardiana. We adopted a comparative modelling approach and used occurrence points from extensive field surveys, supplemented with herbaria records and publicly available distribution data. Our study reveals that currently an area of 8.797334 × 106 km2 (8.8%) is potentially suitable for E. foliata and nearly half 4.759326 × 106 km2 (4.8%) for E. gerardiana. Under future climate change scenarios, distribution range of E. foliata is predicted to expand but contract in E. gerardiana. Similarly, E. foliata showed broader niche breadth which is predicted to increase under B1 (0.097-0.125) and B2 (0.878-0.930) climatic change scenarios. In contrast, E. gerardiana had narrower niche breadth and expected to further decrease under B1 (0.081-0.078) and B2 (0.878-0.854). The most influential bioclimatic variable governing the potential distribution and niche breadth of E. foliata was the precipitation of warmest quarter, whereas that of E. gerardiana was temperature seasonality. The results from our study can help in developing potential indicator plant species for assessment and monitoring of distribution range shifts in response to changing climate in the arid environments.

Early Evidence of Shifts in Alpine Summit Vegetation: A Case Study From Kashmir Himalaya.

Under the contemporary climate change, the Himalaya is reported to be warming at a much higher rate than the global average. However, little is known about the alpine vegetation responses to recent climate change in the rapidly warming Himalaya. Here we studied vegetation dynamics on alpine summits in Kashmir Himalaya in relation to in situ measured microclimate. The summits, representing an elevation gradient from treeline to nival zone (3530-3740 m), were first surveyed in 2014 and then re-surveyed in 2018. The initial survey showed that the species richness, vegetation cover and soil temperature decreased with increasing elevation. Species richness and soil temperature differed significantly among slopes, with east and south slopes showing higher values than north and west slopes. The re-survey showed that species richness increased on the lower three summits but decreased on the highest summit (nival zone) and also revealed a substantial increase in the cover of dominant shrubs, graminoids, and forbs. The nestedness-resultant dissimilarity, rather than species turnover, contributed more to the magnitude of ß-diversity among the summits. High temporal species turnover was found on south and east aspects, while high nestedness was recorded along north and west aspects. Thermophilization was more pronounced on the lower two summits and along the northern aspects. Our study provides crucial scientific data on climate change impacts on the alpine vegetation of Kashmir Himalaya. This information will fill global knowledge gaps from the developing world.