Climate-Driven Plant Response and Resilience on the Tibetan Plateau in Space and Time: A Review

Climate change variation on a small scale may alter the underlying processes determining a pattern operating at large scale and vice versa. Plant response to climate change on individual plant levels on a fine scale tends to change population structure, community composition and ecosystem processes and functioning. Therefore, we reviewed the literature on plant response and resilience to climate change in space and time at different scales on the Tibetan Plateau. We report that spatiotemporal variation in temperature and precipitation dynamics drives the vegetation and ecosystem function on the Tibetan Plateau (TP), following the water–energy dynamics hypothesis. Increasing temperature with respect to time increased the net primary productivity (NPP) on most parts of the Tibetan Plateau, but the productivity dynamics on some parts were constrained by 0.3 °C decade−1 rising temperature. Moreover, we report that accelerating studies on plant community assemblage and their contribution to ecosystem functioning may help to identify the community response and resilience to climate extremes. Furthermore, records on species losses help to build the sustainable management plan for the entire Tibetan Plateau. We recommend that incorporating long-term temporal data with multiple factor analyses will be helpful to formulate the appropriate measures for a healthy ecosystem on the Tibetan Plateau.publishedVersio

A pan-Himalayan test of predictions on plant species richness based on primary production and water-energy dynamics

Spatial variation in plant species diversity is well-documented but an overarching first-principles theory for diversity variation is lacking. Chemical energy expressed as Net Primary Production (NPP) is related to a monotonic increase in species richness at a macroscale and supports one of the leading energy-productivity hypotheses, the More individuals Hypothesis. Alternatively, water-energy dynamics (WED) hypothesizes enhanced species richness when water is freely available and energy supply is optimal. This theoretical model emphasises the amount and duration of photosynthesis across the year and therefore we include the length of the growing season and its interaction with precipitation. This seasonal-WED model assumes that biotemperature and available water represent the photosynthetically active period for the plants and hence, is directly related to NPP, especially in temperate and alpine regions. This study aims to evaluate the above-mentioned theoretical models using interpolated elevational species richness of woody and herbaceous flowering plants of the entire Himalayan range based on data compiled from databases. Generalized linear models (GLM) and generalized linear mixed models (GLMM) were used to analyse species richness (elevational gamma diversity) in the six geopolitical sectors of the Himalaya. NPP, annual precipitation, potential evapotranspiration (derived by the Holdridge formula), and length of growing season were treated as the explanatory variables and the models were evaluated using the Akaike Information Criterion (AIC) and explained deviance. Both precipitation plus potential evapotranspiration (PET), and NPP explain plant species richness in the Himalaya. The seasonal-WED model explains the species richness trends of both plant life-forms in all sectors of the Himalayan range better than the NPP-model. Despite the linear precipitation term failing to precisely capture the amount of water available to plants, the seasonal-WED model, which is based on the thermodynamical transition between water phases, is reasonably good and can forecast peaks in species richness under different climate and primary production conditions.publishedVersio

Weighted average regression and environmental calibration as a tool for quantifying climate-driven changes in vegetation

Aims: Studies of the climatic responses of plant assemblages via vegetation-based environmental reconstructions by weighted averaging (WA) regression and calibration are a recent development in modern vegetation ecology. However, the performance of this technique for plot-based vegetation datasets has not been rigorously tested. We assess the estimation accuracy of the WA approach by comparing results, mainly the root mean square error of prediction (RMSEP) of WA regressions for six different vegetation datasets (total species, high-frequency species and low-frequency species as both abundance and incidence) each from two sites. Methods: Vegetation-inferred environment (plot elevation) calibrated over time is used to quantify the elevational shift in species assemblages. Accuracy of the calibrations is assessed by comparing the linear regression models developed for estimating elevational shifts. The datasets were also used for the backward predictions to check the robustness of the forward predictions. Important Findings: WA regression has a fairly high estimation accuracy, especially with species incidence datasets. However, estimation bias at the extremes of the environmental gradient is evident with all datasets. Out of eight sets (each set with a model for total species, low-frequency species and high-frequency species) of WA regression models, the lowest RMSEPs are produced in the four models based on the total species datasets and in three models based on the high-frequency species only. The inferred environment mirrored the estimation precision of the WA regressions, i.e. precise WA regression models produced more accurate calibrated environmental estimates, which, in turn, resulted in regression models with a higher adjusted r2 for estimating the elevational shift in the species assemblages. Reliable environmental estimates for plot-based datasets can be achieved by WA regression and calibration, although the edge effect may be evident if species turnover is high along an extensive environmental gradient. Species incidence (0/1) data may improve the estimation accuracy by minimizing any potential census and field estimation errors that are more likely to occur in species abundance datasets. Species data processing cannot guarantee the most reliable WA regression models. Instead, generally optimal estimations can be achieved by using all the species with a consistent taxonomy in the training and reconstruction datasets.acceptedVersio

Rate-of-change analysis in palaeoecology revisited: a new approach

Dynamics in the rate of compositional change (rate-of-change; RoC), preserved in paleoecological sequences, are thought to reflect changes due to exogenous (climate and human forcing) or endogenous (local dynamics and biotic interactions) drivers. However, changes in sedimentation rates and sampling strategies can result in an uneven distribution of time intervals and are known to affect RoC estimates. Furthermore, there has been relatively little exploration of the implications of these challenges in quantifying RoC in paleoecology. Here, we introduce R-Ratepol – an easy-to-use R package – that provides a robust numerical technique for detecting and summarizing RoC patterns in complex multivariate time-ordered stratigraphical sequences. First, we compare the performance of common methods of estimating RoC and detecting periods of high RoC (peak-point) using simulated pollen-stratigraphical data with known patterns of compositional change and temporal resolution. In addition, we propose a new method of binning with a moving window, which shows a more than 5-fold increase in the correct detection of peak-points compared to the more traditional way of using individual levels. Next, we apply our new methodology to four representative European pollen sequences and show that our approach also performs well in detecting periods of significant compositional change during known onsets of human activity, early land-use transformation, and changes in fire frequency. Expanding the approach using R-Ratepol to open-access paleoecological datasets in global databases, such as Neotoma, will allow future paleoecological and macroecological studies to quantify major changes in biotic composition or in sets of abiotic variables across broad spatiotemporal scales.publishedVersio

A guide to the processing and standardization of global palaeoecological data for large-scale syntheses using fossil pollen

Aim: Palaeoecological data are crucial for comprehending large-scale biodiversity patterns and the natural and anthropogenic drivers that influence them over time. Over the last decade, the availability of open-access research databases of palaeoecological proxies has substantially increased. These databases open the door to research questions needing advanced numerical analyses and modelling based on big-data compilations. However, compiling and analysing palaeoecological data pose unique challenges that require a guide for producing standardized and reproducible compilations. Innovation: We present a step-by-step guide of how to process fossil pollen data into a standardized dataset compilation ready for macroecological and palaeoecological analyses. We describe successive criteria that will enhance the quality of the compilations. Though these criteria are project and research question-dependent, we discuss the most important assumptions that should be considered and adjusted accordingly. Our guide is accompanied by an R-workflow—called FOSSILPOL—and corresponding R-package—called R-Fossilpol—that provide a detailed protocol ready for interdisciplinary users. We illustrate the workflow by sourcing and processing Scandinavian fossil pollen datasets and show the reproducibility of continental-scale data processing. Main Conclusions: The study of biodiversity and macroecological patterns through time and space requires large-scale syntheses of palaeoecological datasets. The data preparation for such syntheses must be transparent and reproducible. With our FOSSILPOL workflow and R-package, we provide a protocol for optimal handling of large compilations of fossil pollen datasets and workflow reproducibility. Our workflow is also relevant for the compilation and synthesis of other palaeoecological proxies and as such offers a guide for synthetic and cross-disciplinary analyses with macroecological, biogeographical and palaeoecological perspectives. However, we emphasize that expertise and informed decisions based on palaeoecological knowledge remain crucial for high-quality data syntheses and should be strongly embedded in studies that rely on the increasing amount of open-access palaeoecological data.publishedVersio

Exploring spatio-temporal patterns of palynological changes in Asia during the Holocene

Historical legacies influence present-day ecosystem composition and dynamics. It is therefore important to understand the long-term dynamics of ecosystems and their properties. Analysis of ecosystem properties during the Holocene using fossil pollen assemblages provides valuable insights into past ecosystem dynamics by summarising so-called pollen-assemblage properties (PAPs). Using 205 fossil pollen data-sets (records), we quantify eight PAPs [pollen-taxonomic richness, diversity, evenness, pollen-compositional turnover, pollen-compositional change, and rate of pollen-compositional change (RoC)] for the Asian continent at different spatial scales (in individual records, within and across climate-zones, and within the continent) and time (temporal patterns over the past 12,000 years). Regression tree (RT) partitioning of the PAP-estimates using sample-age as a sole predictor revealed the “change-point(s)” (time or sample-age of major change in a PAP). We estimated the density of RT and multivariate regression tree (MRT) change-points in 1,000-year time bins during the Holocene. Pollen-compositional turnover (range of sample scores along the first DCCA axis) and change (number of MRT partitions) in each record reveal gradual spatial variation across latitude and a decline with longitude eastward. Temporally, compositional turnover declines linearly throughout the Holocene at all spatial scales. Other PAPs are heterogeneous across and within spatial scales, being more detectable at coarser scales. RT and MRT change-point density is broadly consistent in climate-zones and the continent, increasing from the early- to mid-Holocene, and mostly decrease from the mid-Holocene to the present for all PAPs. The heterogenous patterns in PAPs across the scales of study most likely reflect responses to variations in regional environmental conditions, anthropogenic land-use, and their interactions over space and time. Patterns at the climate-zone and continental scales indicate a gradual but congruent decline in major PAPs such as compositional turnover, rate of compositional change, and major temporal compositional changes (MRT) during the Holocene, especially during recent millennia, suggesting that vegetation in Asia has become progressively more homogenous. Data properties (e.g., spatial distribution of the records, distribution of samples within the records, and data-standardisation and analytical approaches) may also have partly influenced the results. It is critically important to evaluate the data properties and the approaches to data standardisation and summarisation.publishedVersio

Approaches to pollen taxonomic harmonisation in Quaternary palynology

Pollen taxonomic harmonisation involves the standardisation of the nomenclature of pollen and fern spores with similar morphotypes at the determination level that is common to all grains or spores with that morphotype within the pollen sequence(s) of interest. Such harmonisation is required prior to subsequent investigations such as numerical analysis, comparing, mapping, synthesis, and environmental reconstruction involving several pollen sequences. Here we present two approaches to harmonisation. These are a ‘top-down’ and a ‘bottom-up’ approach. The bottom-up approach is preferred. It is based on the concept of the regional pollen flora for the sequence(s) in the spatial area(s) of study. We present bottom-up harmonisation tables for the continental or sub-continental scales developed for the Humans on Planet Earth (HOPE) project. The tables are for North America, Latin America, Europe, Asia (three parts), and Indo-Pacific. These harmonisations are project-specific and sequence-specific, relating to the geographical area and to the sequences in the area under consideration, both of which are linked to the research questions being addressed. A new bottom-up harmonisation with a consistent taxonomic level and nomenclature is needed when additional sequences or areas are added. However, the HOPE tables can serve as a starting point for further research involving multi-sequence analyses or syntheses.publishedVersio

GrassPlot v. 2.00 – first update on the database of multi-scale plant diversity in Palaearctic grasslands

Abstract: GrassPlot is a collaborative vegetation-plot database organised by the Eurasian Dry Grassland Group (EDGG) and listed in the Global Index of Vegetation-Plot Databases (GIVD ID EU-00-003). Following a previous Long Database Report (Dengler et al. 2018, Phyto- coenologia 48, 331–347), we provide here the first update on content and functionality of GrassPlot. The current version (GrassPlot v. 2.00) contains a total of 190,673 plots of different grain sizes across 28,171 independent plots, with 4,654 nested-plot series including at least four grain sizes. The database has improved its content as well as its functionality, including addition and harmonization of header data (land use, information on nestedness, structure and ecology) and preparation of species composition data. Currently, GrassPlot data are intensively used for broad-scale analyses of different aspects of alpha and beta diversity in grassland ecosystems

Spatiotemporal dynamics of plant assemblages under changing climate and land-use regimes in central Nepal Himalaya

The trajectories of vegetation and landscape dynamics have been redirected globally by climate change and land-use change. The drivers and mechanisms of the spatiotemporal changes are likely scale dependent and are most probably confounded. Himalayan landscapes are under-explored and it is particularly crucial to understand the mechanisms and trajectories of the changes they are experiencing and their scale relationship for an effective management of these landscapes. This thesis synthesises four case studies that have documented spatiotemporal changes in plant assemblages driven by climate and land-use change over the last two decades, assessed the relationship between alpha diversity (plot-based species richness) and gamma diversity (regional species richness) with a focus on scale, and tested the performance of weighted averaging (WA) regression and calibration for quantifying the elevational changes of species assemblages. Studies were conducted in temperate, subalpine, and alpine vegetation at two locations in central Nepal. Scale sensitivity of the elevational species richness was assessed by treating the alpha diversity of different plant life-forms at different grain sizes and areas used for gamma estimation as response variables in a generalised linear model [Paper I]. Systematic changes in temporally resurveyed assemblages were analysed by ordinations and attributed to climate and land-use change using regression analyses [Papers II, III]. WA regression and calibration technique was evaluated by comparing the models for different types of temporal datasets [Paper IV]. Elevational gamma diversity can significantly predict alpha diversity and the relationship is largely scale invariant, although it is slightly less so for woody species. At a regional level, climate warming is a major driver of the demonstrated spatiotemporal changes, i.e. thermophilisation of plant assemblages. However, landuse change may confound or counteract the climatic effects at a local or landscape level. WA regression and calibration predicts fairly accurately the elevation of the plot-based vegetation assemblages. Use of species incidence data may improve the accuracy, but species data processing cannot guarantee more accurate calibrations. To conclude, the systematic spatiotemporal changes in plant assemblages over the last two decades in central Nepal are significant, are largely irrespective of spatial scale, and are most likely related to interactions and feedback mechanisms between climate change and land-use change at different spatiotemporal scales. Taxonomic, census, and sampling accuracy are crucial in the analyses of temporal changes, especially by environmental reconstructions

Does tree canopy closure moderate the effect of climate warming on plant species composition of temperate Himalayan oak forest?

Question: We ask if there are significant changes in plant species composition after 20 yr of documented temperature increase (~1 °C), and if the temporal changes are different in closed temperate oak forest compared to semi-open canopy. Answers to these questions may indicate if climate warming and/or canopy closure, controlled by land-use regime, is the main driver of any documented compositional changes in the forest between 1993 and 2013. Location: Phulchoki Mountain, central Himalaya (Nepal). Methods: We resampled 64 plots of 100 m2 after 20 yr and recorded all terrestrial vascular plants and percentage canopy cover in each sample plot. We analysed the compositional changes in terms of species abundance, frequency and spatial translocation in relation to atmospheric temperature and canopy cover using univariate and multivariate statistics. Results: We find clear changes in the species composition, with abundance of almost half of the studied species having increased and one-quarter of the species having decreased over the past two decades. The changes vary among life forms: trees increased, whereas decreasing species were mainly herbaceous or shrubs. Similarly, shade-tolerant species increased, whereas those adapted to open habitat decreased. The compositional changes are mainly explained through the increased regional temperature, with a significant buffering effect of tree canopy cover. A significant increase in low-elevation (warm-adapted) species is detected, but this thermophilization is only found in the semi-open forest. Conclusions: Fine-scale temporal changes in the temperate oak forests are mainly driven by macroclimate warming, although change in land-use regime also has a profound direct effect by governing canopy closure that, in turn, moderates the effect of climate warming. Tree canopy cover governs the relative abundances of different species based on their adaptive characteristics, such as shade tolerance and life-form traits. The magnitude of the temporal vegetation change would therefore be dependent on both the degree of regional climate warming and forest canopy closure