Palaeovegetation in China during the late Quaternary: Biome reconstructions based on a global scheme of plant functional types

Two previous reconstructions of palaeovegetation across the whole of China were performed using a simple classification of plant functional types (PFTs). Now a more explicit, global PFT classification scheme has been developed, and a substantial number of additional pollen records have become available. Here we apply the global scheme of PFTs to a comprehensive set of pollen records available from China to test the applicability of the global scheme of PFTs in China, and to obtain a well-founded reconstruction of changing palaeovegetation patterns. A total of 806 pollen surface samples, 188 mid-Holocene (MH, 6000 14C yr BP) and 50 last glacial maximum (LGM, 18,000 14C yr BP) pollen records were used to reconstruct vegetation patterns in China, based on a new global classification system of PFTs and a standard numerical technique for biome assignment (biomization). The biome reconstruction based on pollen surface samples showed convincing agreement with present potential natural vegetation. Coherent patterns of change in biome distribution between MH, LGM and present are observed. In the MH, cold and cool-temperate evergreen needleleaf forests and mixed forests, temperate deciduous broadleaf forest, and warm-temperate evergreen broadleaf and mixed forest in eastern China were shifted northward by 200–500 km. Cold-deciduous forest in northeastern China was replaced by cold evergreen needleleaf forest while in central northern China, cold-deciduous forest was present at some sites now occupied by temperate grassland and desert. The forest–grassland boundary was 200–300 km west of its present position. Temperate xerophytic shrubland, temperate grassland and desert covered a large area on the Tibetan Plateau, but the area of tundra was reduced. Treeline was 300–500 m higher than present in Tibet. These changes imply generally warmer winters, longer growing seasons and more precipitation during the MH. Westward shifts of the forest–shrubland–grassland and grassland–desert boundaries imply greater moisture availability in the MH, consistent with a stronger summer monsoon. During the LGM, in contrast, cold-deciduous forest, cool-temperate evergreen needleleaf forest, cool mixed forests, warm-temperate evergreen broadleaf and mixed forest in eastern China were displaced to the south by 300–1000 km, while temperate deciduous broadleaf forest, pure warm-temperate evergreen forest, tropical semi-evergreen and evergreen broadleaf forests were restricted or absent from the mainland of southern China, implying colder winters than present. Strong shifts of temperate xerophytic shrubland, temperate grassland and desert to the south and east in northern and western China and on the Tibetan Plateau imply drier conditions than present.

Ecophysiological and bioclimatic foundations for a global plant functional classification

Question: What plant properties might define plant functional types (PFTs) for the analysis of global vegetation responses to climate change, and what aspects of the physical environment might be expected to predict the distributions of PFTs? Methods: We review principles to explain the distribution of key plant traits as a function of bioclimatic variables. We focus on those whole-plant and leaf traits that are commonly used to define biomes and PFTs in global maps and models. Results: Raunkiær's plant life forms (underlying most later classifications) describe different adaptive strategies for surviving low temperature or drought, while satisfying requirements for reproduction and growth. Simple conceptual models and published observations are used to quantify the adaptive significance of leaf size for temperature regulation, leaf consistency for maintaining transpiration under drought, and phenology for the optimization of annual carbon balance. A new compilation of experimental data supports the functional definition of tropical, warm-temperate, temperate and boreal phanerophytes based on mechanisms for withstanding low temperature extremes. Chilling requirements are less well quantified, but are a necessary adjunct to cold tolerance. Functional traits generally confer both advantages and restrictions; the existence of trade-offs contributes to the diversity of plants along bioclimatic gradients. Conclusions: Quantitative analysis of plant trait distributions against bioclimatic variables is becoming possible; this opens up new opportunities for PFT classification. A PFT classification based on bioclimatic responses will need to be enhanced by information on traits related to competition, successional dynamics and disturbance.

TRY: a global database of plant traits

Plant traits – the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs – determine how primary producers respond to environmental factors, affect other trophic levels, influence ecosystem processes and services and provide a link from species richness to ecosystem functional diversity. Trait data thus represent the raw material for a wide range of research from evolutionary biology, community and functional ecology to biogeography. Here we present the global database initiative named TRY, which has united a wide range of the plant trait research community worldwide and gained an unprecedented buy-in of trait data: so far 93 trait databases have been contributed. The data repository currently contains almost three million trait entries for 69 000 out of the world's 300 000 plant species, with a focus on 52 groups of traits characterizing the vegetative and regeneration stages of the plant life cycle, including growth, dispersal, establishment and persistence. A first data analysis shows that most plant traits are approximately log-normally distributed, with widely differing ranges of variation across traits. Most trait variation is between species (interspecific), but significant intraspecific variation is also documented, up to 40% of the overall variation. Plant functional types (PFTs), as commonly used in vegetation models, capture a substantial fraction of the observed variation – but for several traits most variation occurs within PFTs, up to 75% of the overall variation. In the context of vegetation models these traits would better be represented by state variables rather than fixed parameter values. The improved availability of plant trait data in the unified global database is expected to support a paradigm shift from species to trait-based ecology, offer new opportunities for synthetic plant trait research and enable a more realistic and empirically grounded representation of terrestrial vegetation in Earth system models.

Impact of land cover uncertainties on estimates of biospheric carbon fluxes

Large-scale bottom-up estimates of terrestrial carbon fluxes, whether based on models or inventory, are highly dependent on the assumed land cover. Most current land cover and land cover change maps are based on satellite data and are likely to be so for the foreseeable future. However, these maps show large differences, both at the class level and when transformed into Plant Functional Types (PFTs), and these can lead to large differences in terrestrial CO2 fluxes estimated by Dynamic Vegetation Models. In this study the Sheffield Dynamic Global Vegetation Model is used. We compare PFT maps and the resulting fluxes arising from the use of widely available moderate (1 km) resolution satellite-derived land cover maps (the Global Land Cover 2000 and several MODIS classification schemes), with fluxes calculated using a reference high (25 m) resolution land cover map specific to Great Britain (the Land Cover Map 2000). We demonstrate that uncertainty is introduced into carbon flux calculations by (1) incorrect or uncertain assignment of land cover classes to PFTs; (2) information loss at coarser resolutions; (3) difficulty in discriminating some vegetation types from satellite data. When averaged over Great Britain, modeled CO2 fluxes derived using the different 1 km resolution maps differ from estimates made using the reference map. The ranges of these differences are 254 gC m−2 a−1 in Gross Primary Production (GPP); 133 gC m−2 a−1 in Net Primary Production (NPP); and 43 gC m−2 a−1 in Net Ecosystem Production (NEP). In GPP this accounts for differences of −15.8% to 8.8%. Results for living biomass exhibit a range of 1109 gC m−2. The types of uncertainties due to land cover confusion are likely to be representative of many parts of the world, especially heterogeneous landscapes such as those found in western Europe.

Last glacial vegetation of northern Eurasia

In order to investigate the potential role of vegetation changes in megafaunal extinctions during the later part of the last glacial stage and early Holocene (42–10 ka BP), the palaeovegetation of northern Eurasia and Alaska was simulated using the LPJ-GUESS dynamic vegetation model. Palaeoclimatic driving data were derived from simulations made for 22 time slices using the Hadley Centre Unified Model. Modelled annual net primary productivity (aNPP) of a series of plant functional types (PFTs) is mapped for selected time slices and summarised for major geographical regions for all time slices. Strong canonical correlations are demonstrated between model outputs and pollen data compiled for the same period and region. Simulated aNPP values, especially for tree PFTs and for a mesophilous herb PFT, provide evidence of the structure and productivity of last glacial vegetation. The mesophilous herb PFT aNPP is higher in many areas during the glacial than at present or during the early Holocene. Glacial stage vegetation, whilst open and largely treeless in much of Europe, thus had a higher capacity to support large vertebrate herbivore populations than did early Holocene vegetation. A marked and rapid decrease in aNPP of mesophilous herbs began shortly after the Last Glacial Maximum, especially in western Eurasia. This is likely implicated in extinction of several large herbivorous mammals during the latter part of the glacial stage and the transition to the Holocene.

Pollen-based reconstructions of Japanese biomes at 0, 6000 and 18,000 14C yr bp

A biomization method, which objectively assigns individual pollen assemblages to biomes ( Prentice et al., 1996 ), was tested using modern pollen data from Japan and applied to fossil pollen data to reconstruct palaeovegetation patterns 6000 and 18,000 14C yr bp Biomization started with the assignment of 135 pollen taxa to plant functional types (PFTs), and nine possible biomes were defined by specific combinations of PFTs. Biomes were correctly assigned to 54% of the 94 modern sites. Incorrect assignments occur near the altitudinal limits of individual biomes, where pollen transport from lower altitudes blurs the local pollen signals or continuous changes in species composition characterizes the range limits of biomes. As a result, the reconstructed changes in the altitudinal limits of biomes at 6000 and 18,000 14C yr bp are likely to be conservative estimates of the actual changes. The biome distribution at 6000 14C yr bp was rather similar to today, suggesting that changes in the bioclimate of Japan have been small since the mid-Holocene. At 18,000 14C yr bp the Japanese lowlands were covered by taiga and cool mixed forests. The southward expansion of these forests and the absence of broadleaved evergreen/warm mixed forests reflect a pronounced year-round cooling.

Responses of leaf traits to climatic gradients: adaptive variation versus compositional shifts

Dynamic global vegetation models (DGVMs) typically rely on plant functional types (PFTs), which are assigned distinct environmental tolerances and replace one another progressively along environmental gradients. Fixed values of traits are assigned to each PFT; modelled trait variation along gradients is thus driven by PFT replacement. But empirical studies have revealed "universal" scaling relationships (quantitative trait variations with climate that are similar within and between species, PFTs and communities); and continuous, adaptive trait variation has been proposed to replace PFTs as the basis for next-generation DGVMs. Here we analyse quantitative leaf-trait variation on long temperature and moisture gradients in China with a view to understanding the relative importance of PFT replacement vs. continuous adaptive variation within PFTs. Leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC) and nitrogen content of dry matter were measured on all species at 80 sites ranging from temperate to tropical climates and from dense forests to deserts. Chlorophyll fluorescence traits and carbon, phosphorus and potassium contents were measured at 47 sites. Generalized linear models were used to relate log-transformed trait values to growing-season temperature and moisture indices, with or without PFT identity as a predictor, and to test for differences in trait responses among PFTs. Continuous trait variation was found to be ubiquitous. Responses to moisture availability were generally similar within and between PFTs, but biophysical traits (LA, SLA and LDMC) of forbs and grasses responded differently from woody plants. SLA and LDMC responses to temperature were dominated by the prevalence of evergreen PFTs with thick, dense leaves at the warm end of the gradient. Nutrient (N, P and K) responses to climate gradients were generally similar within all PFTs. Area-based nutrients generally declined with moisture; Narea and Karea declined with temperature, but Parea increased with temperature. Although the adaptive nature of many of these trait-climate relationships is understood qualitatively, a key challenge for modelling is to predict them quantitatively. Models must take into account that community-level responses to climatic gradients can be influenced by shifts in PFT composition, such as the replacement of deciduous by evergreen trees, which may run either parallel or counter to trait variation within PFTs. The importance of PFT shifts varies among traits, being important for biophysical traits but less so for physiological and chemical traits. Finally, models should take account of the diversity of trait values that is found in all sites and PFTs, representing the "pool" of variation that is locally available for the natural adaptation of ecosystem function to environmental change.

Characterization of a highly biodiverse floodplain meadow using hyperspectral remote sensing within a plant functional trait framework

We assessed the potential for using optical functional types as effective markers to monitor changes in vegetation in floodplain meadows associated with changes in their local environment. Floodplain meadows are challenging ecosystems for monitoring and conservation because of their highly biodiverse nature. Our aim was to understand and explain spectral differences among key members of floodplain meadows and also characterize differences with respect to functional traits. The study was conducted on a typical floodplain meadow in UK (MG4-type, mesotrophic grassland type 4, according to British National Vegetation Classification). We compared two approaches to characterize floodplain communities using field spectroscopy. The first approach was sub-community based, in which we collected spectral signatures for species groupings indicating two distinct eco-hydrological conditions (dry and wet soil indicator species). The other approach was “species-specific”, in which we focused on the spectral reflectance of three key species found on the meadow. One herb species is a typical member of the MG4 floodplain meadow community, while the other two species, sedge and rush, represent wetland vegetation. We also monitored vegetation biophysical and functional properties as well as soil nutrients and ground water levels. We found that the vegetation classes representing meadow sub-communities could not be spectrally distinguished from each other, whereas the individual herb species was found to have a distinctly different spectral signature from the sedge and rush species. The spectral differences between these three species could be explained by their observed differences in plant biophysical parameters, as corroborated through radiative transfer model simulations. These parameters, such as leaf area index, leaf dry matter content, leaf water content, and specific leaf area, along with other functional parameters, such as maximum carboxylation capacity and leaf nitrogen content, also helped explain the species’ differences in functional dynamics. Groundwater level and soil nitrogen availability, which are important factors governing plant nutrient status, were also found to be significantly different for the herb/wetland species’ locations. The study concludes that spectrally distinguishable species, typical for a highly biodiverse site such as a floodplain meadow, could potentially be used as target species to monitor vegetation dynamics under changing environmental conditions.

Pollen-based reconstructions of biome distributions for Australia, Southeast Asia and the Pacific (SEAPAC region) at 0,6000 and 18,000 14C yr B.P.

Aim  This paper documents reconstructions of the vegetation patterns in Australia, Southeast Asia and the Pacific (SEAPAC region) in the mid-Holocene and at the last glacial maximum (LGM). Methods  Vegetation patterns were reconstructed from pollen data using an objective biomization scheme based on plant functional types. The biomization scheme was first tested using 535 modern pollen samples from 377 sites, and then applied unchanged to fossil pollen samples dating to 6000 ± 500 or 18,000 ± 1000 14C yr bp. Results  1. Tests using surface pollen sample sites showed that the biomization scheme is capable of reproducing the modern broad-scale patterns of vegetation distribution. The north–south gradient in temperature, reflected in transitions from cool evergreen needleleaf forest in the extreme south through temperate rain forest or wet sclerophyll forest (WSFW) and into tropical forests, is well reconstructed. The transitions from xerophytic through sclerophyll woodlands and open forests to closed-canopy forests, which reflect the gradient in plant available moisture from the continental interior towards the coast, are reconstructed with less geographical precision but nevertheless the broad-scale pattern emerges. 2. Differences between the modern and mid-Holocene vegetation patterns in mainland Australia are comparatively small and reflect changes in moisture availability rather than temperature. In south-eastern Australia some sites show a shift towards more moisture-stressed vegetation in the mid-Holocene with xerophytic woods/scrub and temperate sclerophyll woodland and shrubland at sites characterized today by WSFW or warm-temperate rain forest (WTRF). However, sites in the Snowy Mountains, on the Southern Tablelands and east of the Great Dividing Range have more moisture-demanding vegetation in the mid-Holocene than today. South-western Australia was slightly drier than today. The single site in north-western Australia also shows conditions drier than today in the mid-Holocene. Changes in the tropics are also comparatively small, but the presence of WTRF and tropical deciduous broadleaf forest and woodland in the mid-Holocene, in sites occupied today by cool-temperate rain forest, indicate warmer conditions. 3. Expansion of xerophytic vegetation in the south and tropical deciduous broadleaf forest and woodland in the north indicate drier conditions across mainland Australia at the LGM. None of these changes are informative about the degree of cooling. However the evidence from the tropics, showing lowering of the treeline and forest belts, indicates that conditions were between 1 and 9 °C (depending on elevation) colder. The encroachment of tropical deciduous broadleaf forest and woodland into lowland evergreen broadleaf forest implies greater aridity. Main conclusions  This study provides the first continental-scale reconstruction of mid-Holocene and LGM vegetation patterns from Australia, Southeast Asia and the Pacific (SEAPAC region) using an objective biomization scheme. These data will provide a benchmark for evaluation of palaeoclimate simulations within the framework of the Palaeoclimate Modelling Intercomparison Project.

Palaeovegetation of China: a pollen data-based synthesis for the mid-Holocene and last glacial maximum.

Pollen data from China for 6000 and 18,000 14C yr bp were compiled and used to reconstruct palaeovegetation patterns, using complete taxon lists where possible and a biomization procedure that entailed the assignment of 645 pollen taxa to plant functional types. A set of 658 modern pollen samples spanning all biomes and regions provided a comprehensive test for this procedure and showed convincing agreement between reconstructed biomes and present natural vegetation types, both geographically and in terms of the elevation gradients in mountain regions of north-eastern and south-western China. The 6000 14C yr bp map confirms earlier studies in showing that the forest biomes in eastern China were systematically shifted northwards and extended westwards during the mid-Holocene. Tropical rain forest occurred on mainland China at sites characterized today by either tropical seasonal or broadleaved evergreen/warm mixed forest. Broadleaved evergreen/warm mixed forest occurred further north than today, and at higher elevation sites within the modern latitudinal range of this biome. The northern limit of temperate deciduous forest was shifted c. 800 km north relative to today. The 18,000 14C yr bp map shows that steppe and even desert vegetation extended to the modern coast of eastern China at the last glacial maximum, replacing today’s temperate deciduous forest. Tropical forests were excluded from China and broadleaved evergreen/warm mixed forest had retreated to tropical latitudes, while taiga extended southwards to c. 43°N.

Pollen-based biomes for Beringia 18,000, 6000 and 0 14C yr B.P.

The objective biomization method developed by Prentice et al. (1996) for Europe was extended using modern pollen samples from Beringia and then applied to fossil pollen data to reconstruct palaeovegetation patterns at 6000 and 18,000 14C yr bp. The predicted modern distribution of tundra, taiga and cool conifer forests in Alaska and north-western Canada generally corresponds well to actual vegetation patterns, although sites in regions characterized today by a mosaic of forest and tundra vegetation tend to be preferentially assigned to tundra. Siberian larch forests are delimited less well, probably due to the extreme under-representation of Larix in pollen spectra. The biome distribution across Beringia at 6000 14C yr bp was broadly similar to today, with little change in the northern forest limit, except for a possible northward advance in the Mackenzie delta region. The western forest limit in Alaska was probably east of its modern position. At 18,000 14C yr bp the whole of Beringia was covered by tundra. However, the importance of the various plant functional types varied from site to site, supporting the idea that the vegetation cover was a mosaic of different tundra types.

Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa.

BIOME 6000 is an international project to map vegetation globally at mid-Holocene (6000 14C yr bp) and last glacial maximum (LGM, 18,000 14C yr bp), with a view to evaluating coupled climate-biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site-based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method’s skill in reconstructing present-day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south-western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 14C yr bp in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial-interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now-arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes. Major changes in physical land-surface conditions, shown by the palaeobiome data, have implications for the global climate. The data can be used directly to evaluate the output of coupled atmosphere-biosphere models. The data could also be objectively generalized to yield realistic gridded land-surface maps, for use in sensitivity experiments with atmospheric models. Recent analyses of vegetation-climate feedbacks have focused on the hypothesized positive feedback effects of climate-induced vegetation changes in the Sahara/Sahel region and the Arctic during the mid-Holocene. However, a far wider spectrum of interactions potentially exists and could be investigated, using these data, both for 6000 14C yr bp and for the LGM.

Improved simulation of fire–vegetation interactions in the Land surface Processes and eXchanges dynamic global vegetation model (LPX-Mv1)

The Land surface Processes and eXchanges (LPX) model is a fire-enabled dynamic global vegetation model that performs well globally but has problems representing fire regimes and vegetative mix in savannas. Here we focus on improving the fire module. To improve the representation of ignitions, we introduced a reatment of lightning that allows the fraction of ground strikes to vary spatially and seasonally, realistically partitions strike distribution between wet and dry days, and varies the number of dry days with strikes. Fuel availability and moisture content were improved by implementing decomposition rates specific to individual plant functional types and litter classes, and litter drying rates driven by atmospheric water content. To improve water extraction by grasses, we use realistic plant-specific treatments of deep roots. To improve fire responses, we introduced adaptive bark thickness and post-fire resprouting for tropical and temperate broadleaf trees. All improvements are based on extensive analyses of relevant observational data sets. We test model performance for Australia, first evaluating parameterisations separately and then measuring overall behaviour against standard benchmarks. Changes to the lightning parameterisation produce a more realistic simulation of fires in southeastern and central Australia. Implementation of PFT-specific decomposition rates enhances performance in central Australia. Changes in fuel drying improve fire in northern Australia, while changes in rooting depth produce a more realistic simulation of fuel availability and structure in central and northern Australia. The introduction of adaptive bark thickness and resprouting produces more realistic fire regimes in Australian savannas. We also show that the model simulates biomass recovery rates consistent with observations from several different regions of the world characterised by resprouting vegetation. The new model (LPX-Mv1) produces an improved simulation of observed vegetation composition and mean annual burnt area, by 33 and 18% respectively compared to LPX.