BackgroundPhylogeny-based assessments of Holocene vegetation are rare. They are important for understanding eco-evolutionary dynamics of species assembly under environmental changes and ever-increasing anthropogenic pressure.AimsTo reveal any spatio-temporal variation in the phylogenetic dispersion (PD, mean phylogenetic distance of co-occurring taxa) in angiosperm pollen assemblages in mainland Europe and North America and their relationship with regional climate and anthropogenic influence.MethodsUsing 12,250 and 14,898 fossil pollen assemblages from Europe and North America, respectively, we analysed family level spatio-temporal variation in Holocene PD by hierarchical generalised linear models and tested its concordance with Holocene climate and anthropogenic influence by Procrustes randomisation tests.ResultsHolocene PD exhibited a reverse-humped pattern along latitudinal gradients in both continents. A longitudinal pattern in Holocene PD in Europe was not pronounced. However, in North America, it declined irregularly along the gradient from west to east. PD in both continents varied significantly during the early- and late-Holocene in concordance with Holocene climate and anthropogenic influences. Complex fine-scale spatio-temporal variations in PD were also pronounced.ConclusionProfound and complex Holocene eco-evolutionary changes in angiosperm assemblages in Europe and North America correspond largely to changes in extrinsic and intrinsic drivers and mechanisms of ecological changes. Future environmental changes of similar or a greater magnitude may drive further changes in the phylogenetic structure of the assemblages. A more detailed study of their eco-evolutionary dynamics requires detailed spatio-temporal assemblage data with precise taxonomy and census, and a detailed and robust species-level phylogeny of the species pool.

Abstract Spatio-temporal assessment of phylogenetic diversity gradients during the Holocene (past 12,000 years) provides an opportunity for a deeper understanding of the dynamics of species co-occurrence patterns under environmental fluctuations. Using two robust metrics of phylogenetic dispersion (PD) and 99 fossil pollen sequences containing 6557 samples/assemblages, we analyse spatio-temporal variation in PD of angiosperms and its relationship with Holocene climate in central Asia. Overall, PD throughout the Holocene decreases linearly with increasing latitude, except for a rise in mean nearest taxon distance from ca. 25 to 35° N. This indicates that phylogenetically divergent taxa decrease progressively with increasing latitude, leaving more phylogenetically closely related taxa in the assemblages, thereby increasing phylogenetic relatedness among the co-occurring taxa. The latitudinal gradient of PD has not been consistent during the Holocene, and this temporal variation is concordant with the Holocene climate dynamics. In general, profound temporal changes in the latitudinal PD toward higher latitudes implies that the major environmental changes during the Holocene have driven considerable spatio-temporal changes in the phylogenetic assembly of high-latitude angiosperm assemblages. Our results suggest that environmental filtering and the tendency of taxa and lineages to retain ancestral ecological features and geographic distributions (phylogenetic niche conservatism) are the main mechanisms underlying the phylogenetic assembly of angiosperms along the climate-latitudinal gradient. Ongoing environmental changes may pose future profound phylogenetic changes in high-latitude plant assemblages, which are adapted to harsh environmental conditions, and therefore are phylogenetically less dispersed (more conservative or clustered).

Controversies exist regarding the extent of past human influence on terrestrial ecosystems and the relative importance of human versus climatic factors in shaping Holocene vegetation. However, there has been no systematic examination of these issues at a global scale. Here we integrate palaeoecological, archaeological, and palaeoclimate data to assess the influence of humans and climate in driving patterns of past vegetation during the early and middle Holocene (8500 - 2000 years before present) as recorded by pollen-assemblage properties. We quantify and summarise the patterns of change in different properties from individual records to a global scale and assess the relative importance of humans and/or climate in influencing them. Additionally, we assess whether the internal relationships among these properties changed through time. While we find evidence that humans have high localized impact on vegetation dynamics in various regions and times periods, when considered globally, humans appear to be a secondary factor, during the early-to-mid Holocene. Our results underscore the value of merging multidisciplinary palaeodata to provide a holistic understanding of millennia-spanning relationships between humans, climate, and vegetation across different spatial scales.

Fossil pollen data are valuable for understanding past spatio-temporal variations in plant diversity. However, deducing robust patterns of fossil pollen-assemblage properties is challenging due to factors related to pollen production, dispersal, and fossilisation; sediment sampling; and nomenclature and taxonomy of the fossil pollen grains reflecting the vegetation that produced them. If the composition pattern of the most recent fossil pollen assemblage concords with those in modern or surface pollen assemblages from the same location(s), the pattern can be regarded as robust. Using recent (50-500 years) fossil pollen and surface pollen assemblages of angiosperms compiled from the same geographic area of Central Asia, we analyse spatial patterns in compositional turnover and phylogenetic dispersion. These are widely used to quantify different aspects of plant diversity. Despite differences in the data regarding spatial coverage, number of pollen records, and number of pollen taxa, the spatial pattern in the modern and fossil pollen-assemblage properties across climate zones is broadly similar. Especially, the latitudinal patterns of pollen-assemblage properties are strikingly similar between both data types. However, the degree of concordance varies according to the diversity metric being analysed, and the magnitudes of the metric may differ between both data types. When the data are compiled, processed, and standardised following consistent criteria and analysed using appropriate metrics and methods, spatial patterns in pollen-assemblage properties of fossil pollen data are as robust as those in surface pollen data (similar to modern vegetation). Therefore, regional patterns of compositional turnover and phylogenetic dispersion can be confidently inferred from older fossil assemblages through 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.

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.

AimPalaeoecological 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. InnovationWe 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 ConclusionsThe 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.

Global vegetation over the past 18,000 years has been transformed first by the climate changes that accompanied the last deglaciation and again by increasing human pressures; however, the magnitude and patterns of rates of vegetation change are poorly understood globally. Using a compilation of 1181 fossil pollen sequences and newly developed statistical methods, we detect a worldwide acceleration in the rates of vegetation compositional change beginning between 4.6 and 2.9 thousand years ago that is globally unprecedented over the past 18,000 years in both magnitude and extent. Late Holocene rates of change equal or exceed the deglacial rates for all continents, which suggests that the scale of human effects on terrestrial ecosystems exceeds even the climate-driven transformations of the last deglaciation. The acceleration of biodiversity change demonstrated in ecological datasets from the past century began millennia ago.

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.

AbstractDynamics in the rate of compositional change (rate-of-change; RoC) of biotic or abiotic assemblages preserved in palaeoecological sequences, are thought to reflect changes due to exogenous drivers such as climate and human forcing as well as endogenous factors linked to local dynamics and biotic interactions. 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 palaeoecology.Here, we introduce R-Ratepol – an easy-to-use R package – that provides a robust numerical technique for detecting and summarising 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 palaeoecological and macroecological studies to quantify major changes in biotic composition or in sets of abiotic variables across broad spatio-temporal scales.

Questions Species-area relationships (SARs) are fundamental for understanding biodiversity patterns and are generally well described by a power law with a constant exponent z. However, z-values sometimes vary across spatial scales. We asked whether there is a general scale dependence of z-values at fine spatial grains and which potential drivers influence it. Location Palaearctic biogeographic realm. Methods We used 6,696 nested-plot series of vascular plants, bryophytes and lichens from the GrassPlot database with two or more grain sizes, ranging from 0.0001 m(2) to 1,024 m(2) and covering diverse open habitats. The plots were recorded with two widespread sampling approaches (rooted presence = species "rooting" inside the plot; shoot presence = species with aerial parts inside). Using Generalized Additive Models, we tested for scale dependence of z-values by evaluating if the z-values differ with gran size and tested for differences between the sampling approaches. The response shapes of z-values to grain were classified by fitting Generalized Linear Models with logit link to each series. We tested whether the grain size where the maximum z-value occurred is driven by taxonomic group, biogeographic or ecological variables. Results For rooted presence, we found a strong monotonous increase of z-values with grain sizes for all grain sizes below 1 m(2). For shoot presence, the scale dependence was much weaker, with hump-shaped curves prevailing. Among the environmental variables studied, latitude, vegetation type, naturalness and land use had strong effects, with z-values of secondary peaking at smaller grain sizes. Conclusions The overall weak scale dependence of z-values underlines that the power function generally is appropriate to describe SARs within the studied grain sizes in continuous open vegetation, if recorded with the shoot presence method. When clear peaks of z-values occur, this can be seen as an expression of granularity of species composition, partly driven by abiotic environment.

The Apuan Alps are one of the most peculiar mountain chain in the Mediterranean, being very close to the coastline and reaching an elevation of almost 2000 m. Based on published flora, we investigated the distribution of plant species richness along the whole elevational gradient of this chain considering: (i) all species, (ii) endemic versus alien species; and (iii) functional groups of species based on Raunkiær life forms (RLF). Generalized Linear Models (GLMs) were used to analyse richness patterns along the elevational gradient, and elevational richness models versus the area of the elevational belts were fitted to test the effect of surface area. Our results showed decreasing species richness with increasing elevation. In contrast, endemic species richness increased along the elevational gradient. Alien species were mainly distributed at low elevations, but this result should be taken with caution since we used historical data. Species life forms were not equally distributed along the elevation gradient: chamaephytes and hemicryptophytes were the richest groups at high elevations, while therophytes showed highest species richness at low elevations. Our findings suggest that in the Apuan Alps there is a major elevational gradient in species composition that could reflect plant evolutionary history. Furthermore, we highlight the key role of published floras as a relevant source of biodiversity data.

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.

Aims: Understanding fine- grain diversity patterns across large spatial extents is fundamental for macroecological research and biodiversity conservation. Using the GrassPlot database, we provide benchmarks of fine- grain richness values of Palaearctic open habitats for vascular plants, bryophytes, lichens and complete vegetation (i.e., the sum of the former three groups). Location: Palaearctic biogeographic realm. Methods: We used 126,524 plots of eight standard grain sizes from the GrassPlot database: 0.0001, 0.001, 0.01, 0.1, 1, 10, 100 and 1,000 m 2 and calculated the mean richness and standard deviations, as well as maximum, minimum, median, and first and third quartiles for each combination of grain size, taxonomic group, biome, region, vegetation type and phytosociological class. Results: Patterns of plant diversity in vegetation types and biomes differ across grain sizes and taxonomic groups. Overall, secondary (mostly semi- natural) grasslands and natural grasslands are the richest vegetation type. The open- access file ”GrassPlot Diversity Benchmarks” and the web tool “GrassPlot Diversity Explorer” are now available online (https://edgg.org/datab ases/Grass landD ivers ityEx plorer) and provide more insights into species richness patterns in the Palaearctic open habitats. Conclusions: The GrassPlot Diversity Benchmarks provide high- quality data on species richness in open habitat types across the Palaearctic. These benchmark data can be used in vegetation ecology, macroecology, biodiversity conservation and data quality checking. While the amount of data in the underlying GrassPlot database and their spatial coverage are smaller than in other extensive vegetation- plot databases, species recordings in GrassPlot

This study explores the relationship between the species composition of lichen and vascular plant species with microtopography at fine scale. We conducted our study in Hardengervidda National Park, Norway. Specifically, we aim to test whether the species richness of different plant life-forms peaks at middle of the microtopography gradient, and then explain the observed patterns with an aid of snow cover gradient along microtopography and snow cover. We sampled 69 species of vascular plants and lichens in 151 plots of 4 m(2) along 23 transects during summer on Tronsbu, Sandhaug and Besso. Detrended correspondence analysis (DCA) was performed to explore how microtopographical gradient was related to the variation in the species composition. One-way ANOVA was performed to test the microtopographic variability in species richness. Afterwards, generalized linear model (GLM) was used to reveal species richness patterns along the snow cover gradient. The first axis in DCA represents the complex gradient from snow free ridge to wet snowbed habitats and the second axis represents a gradient from acidic to calcareous sites. Lichens species richness is greater in ridge than in snowbeds, while all other life forms follow the opposite trend. Species richness for total plant species, vascular plant species and herbaceous plant species increased with increase in weighted average snow indicator value (WASI), whilst species richness for lichen species declined substantially towards the maximum WASI value. In contrast, species richness for dwarf shrub species showed a unimodal relationship with WASI. This study shows that liquid water availability provides a good potential explanation for species composition and richness in mountains, which is controlled by snow cover and prevalent wind direction.

Ethnopharmacological relevance: Neopicrorhiza scrophulariiflora (Pennell) Hong is important medicinal plant that is native to the eastern Himalayas and Hengduan mountains in China. It is also distributed in Nepal, north east India, Bhutan and northern Myanmar. Plant parts are traditionally used against different kinds of diseases and various compounds present in different plant parts are also effective against many diseases. Thus, N. scrophulariiflora has a high potential to maintain human health. Aim of the review: Although N. scrophulariiflora is very important and widely studied plant species but there is no comprehensive up-to-date review of published and unpublished literature. So, in the present article we have compiled and critically commented on the botanical characteristics, traditional uses, plant growth and cultivation, micropropagation, conservation status, secondary metabolites, pharmacology and toxicity of the plant. Materials and methods: Extensive literature searches both electronic online databases (Google Scholar, Scopus, Springer Link, Web of Science, ScienceDirect, ResearchGate, PubMed, ChemSpider, USPTO, Google patents and Espacenet) and library visits in Nepal were carried out to collect the literature on information published prior to April 2019. Results: N. scrophulariiflora was traditionally used for 82 ailments/diseases. There are 124 major phytochemicals extracted from the plant. Several compounds are effective in bioactivity. Pharmacologically, the plant is proved to be anti-atherosclerotic, antidiabetic and anti-inflammatory in-vivo studies, and antimicrobial, antimalarial, antioxidative, hepatoprotective, immunomodulatory and nerve growth factor potentiating from in-vitro studies. Renal improvement activities were confirmed from both in-vivo and in-vitro studies. Toxicological tests and a single clinical trial in human beings have supported the notion that the plant is not poisonous but beneficial for curing wide ranges of diseases. Conclusion: N. scrophulariiflora is valuable medicinal plant that can serve as promising source of non-harmful and potential medicinal herbal remedies for human beings.

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, Phytocoenologia 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.

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. 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. 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 r(2) 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.