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.

Pollen, plant macrofossil and charcoal records for palaeovegetation reconstruction in the Mediterranean-Black Sea Corridor since the Last Glacial Maximum

New reconstructions of changing vegetation patterns in the Mediterranean-Black Sea Corridor since the Last Glacial Maximum are being produced by an improved biomisation scheme that uses both pollen and plant macrofossil data, in conjunction. Changes in fire regimes over the same interval will also be reconstructed using both microscopic and macroscopic charcoal remains. These reconstructions will allow a diagnosis of the interactions between climate, fire and vegetation on millennial timescales, and will also help to clarify the role of coastline and other geomorphic changes, salinity and impacts of human activities in this region. These new data sets are being produced as a result of collaboration between the Palynology Working Group (WG-2) within the IGCP-521 project and the international Palaeovegetation Mapping Project (BIOME 6000). The main objective of this paper is to present the goals of this cooperation, methodology, including limitations and planned improvements, and to show the initial results of some applications.

Climate and CO2 controls on global vegetation distribution at the last glacial maximum: analysis based on palaeovegetation data, biome modelling and palaeoclimate simulations.

The global vegetation response to climate and atmospheric CO2 changes between the last glacial maximum and recent times is examined using an equilibrium vegetation model (BIOME4), driven by output from 17 climate simulations from the Palaeoclimate Modelling Intercomparison Project. Features common to all of the simulations include expansion of treeless vegetation in high northern latitudes; southward displacement and fragmentation of boreal and temperate forests; and expansion of drought-tolerant biomes in the tropics. These features are broadly consistent with pollen-based reconstructions of vegetation distribution at the last glacial maximum. Glacial vegetation in high latitudes reflects cold and dry conditions due to the low CO2 concentration and the presence of large continental ice sheets. The extent of drought-tolerant vegetation in tropical and subtropical latitudes reflects a generally drier low-latitude climate. Comparisons of the observations with BIOME4 simulations, with and without consideration of the direct physiological effect of CO2 concentration on C3 photosynthesis, suggest an important additional role of low CO2 concentration in restricting the extent of forests, especially in the tropics. Global forest cover was overestimated by all models when climate change alone was used to drive BIOME4, and estimated more accurately when physiological effects of CO2 concentration were included. This result suggests that both CO2 effects and climate effects were important in determining glacial-interglacial changes in vegetation. More realistic simulations of glacial vegetation and climate will need to take into account the feedback effects of these structural and physiological changes on the climate.

Palaeovegetation (Communications arising): diversity of temperate plants in east Asia

The exceptionally broad species diversity of vascular plant genera in east Asian temperate forests, compared with their sister taxa in North America, has been attributed to the greater climatic diversity of east Asia, combined with opportunities for allopatric speciation afforded by repeated fragmentation and coalescence of populations through Late Cenozoic ice-age cycles1. According to Qian and Ricklefs1, these opportunities occurred in east Asia because temperate forests extended across the continental shelf to link populations in China, Korea and Japan during glacial periods, whereas higher sea levels during interglacial periods isolated these regions and warmer temperatures restricted temperate taxa to disjunct refuges. However, palaeovegetation data from east Asia2, 3, 4, 5, 6 show that temperate forests were considerably less extensive than today during the Last Glacial Maximum, calling into question the coalescence of tree populations required by the hypothesis of Qian and Ricklefs1.

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.

Impact of a drier Early-Mid-Holocene climate upon Amazonian forests

This paper uses a palaeoecological approach to examine the impact of drier climatic conditions of the Early-Mid-Holocene (ca 8000-4000 years ago) upon Amazonia's forests and their fire regimes. Palaeovegetation (pollen data) and palaeofire (charcoal) records are synthesized from 20 sites within the present tropical forest biome, and the underlying causes of any emergent patterns or changes are explored by reference to independent palaeoclimate data and present-day patterns of precipitation, forest cover and fire activity across Amazonia. During the Early-Mid-Holocene, Andean cloud forest taxa were replaced by lowland tree taxa as the cloud base rose while lowland ecotonal areas, which are presently covered by evergreen rainforest, were instead dominated by savannahs and/or semi-deciduous dry forests. Elsewhere in the Amazon Basin there is considerable spatial and temporal variation in patterns of vegetation disturbance and fire, which probably reflects the complex heterogeneous patterns in precipitation and seasonality across the basin, and the interactions between climate change, drought- and fire susceptibility of the forests, and Palaeo-Indian land use. Our analysis shows that the forest biome in most parts of Amazonia appears to have been remarkably resilient to climatic conditions significantly drier than those of today, despite widespread evidence of forest burning. Only in ecotonal areas is there evidence of biome replacement in the Holocene. From this palaeoecological perspective, we argue against the Amazon forest 'dieback' scenario simulated for the future.

Palynological differentiation between genera of the Moraceae family and implications for Amazonian palaeoecology

The Moraceae family is one of the most abundant and ecologically important families in Neotropical rainforests and is very well-represented in Amazonian fossil pollen records. However, difficulty in differentiating palynologically between the genera within this family, or between the Moraceae and Urticaceae families, has limited the amount of palaeoecological information that can be extracted from these records. The aim of this paper is to analyse the morphological properties of pollen from Amazonian species of Moraceae in order to determine whether the pollen taxonomy of this family can be improved. Descriptive and morphometric methods are used to identify and differentiate key pollen types of the Moraceae (mulberry) and Urticaceae (nettle) families which are represented in Amazonian rainforest communities of Noel Kempff Mercado National Park (NKMNP), Northeast Bolivia. We demonstrate that Helicostylis, Brosimum, Pseudolmedia, Sorocea and Pourouma pollen can be identified in tropical pollen assemblages and present digital images of, and a taxonomic key to, the Moraceae pollen types of NKMNP. Indicator species, Maquira coriacea (riparian evergreen forest) and Brosimum gaudichaudii (open woodland and upland savanna communities), also exhibit unique pollen morphologies. The ability to recognise these ecologically important taxa in pollen records provides the potential for much more detailed and reliable Neotropical palaeovegetation reconstructions than have hitherto been possible. In particular, this improved taxonomic resolution holds promise for resolving long-standing controversies over the interpretation of key Amazonian Quaternary pollen records.

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.

Evaluation of coupled ocean-atmosphere simulations of northern hemisphere extratropical climates in the mid-Holocene

We have used the BIOME4 biogeography–biochemistry model and comparison with palaeovegetation data to evaluate the response of six ocean–atmosphere general circulation models to mid-Holocene changes in orbital forcing in the mid- to high-latitudes of the northern hemisphere. All the models produce: (a) a northward shift of the northern limit of boreal forest, in response to simulated summer warming in high-latitudes. The northward shift is markedly asymmetric, with larger shifts in Eurasia than in North America; (b) an expansion of xerophytic vegetation in mid-continental North America and Eurasia, in response to increased temperatures during the growing season; (c) a northward expansion of temperate forests in eastern North America, in response to simulated winter warming. The northward shift of the northern limit of boreal forest and the northward expansion of temperate forests in North America are supported by palaeovegetation data. The expansion of xerophytic vegetation in mid-continental North America is consistent with palaeodata, although the extent may be over-estimated. The simulated expansion of xerophytic vegetation in Eurasia is not supported by the data. Analysis of an asynchronous coupling of one model to an equilibrium-vegetation model suggests vegetation feedback exacerbates this mid-continental drying and produces conditions more unlike the observations. Not all features of the simulations are robust: some models produce winter warming over Europe while others produce winter cooling. As a result, some models show a northward shift of temperate forests (consistent with, though less marked than, the expansion shown by data) and others produce a reduction in temperate forests. Elucidation of the cause of such differences is a focus of the current phase of the Palaeoclimate Modelling Intercomparison Project.

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.

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.

Land surface feedbacks and palaeomonsoons in northern Africa

We ran a sequence of climate model experiments for 6000 years ago, with land-surface conditions based on a realistic map of palaeovegetation, lakes and wetlands, to quantify the effects of land-surface feedbacks in the Saharan region. Vegetation-induced albedo and moisture flux changes produced year-round warming, forced the monsoon to 17°–25°N two months earlier, and shifted the precipitation belt ≈300 km northwards compared to the effects of orbital forcing alone. The addition of lakes and wetlands produced localised changes in evaporation and precipitation, but caused no further extension of the monsoon belt. Diagnostic analyses with biome and continental hydrology models showed that the combined land-surface feedbacks, although substantial, could neither maintain grassland as far north as observed (≈26°N) nor maintain Lake “MegaChad” (330,000 km²).

Vegetation and soils feedbacks on the response of the African monsoon response to orbital forcing in early to middle Holocene

Fossil pollen, ancient lake sediments and archaeological evidence from Africa indicate that the Sahel and Sahara regions were considerably wetter than today during the early to middle Holocene period, about 12,000 to 5,000 years ago1–4. Vegetation associated with the modern Sahara/Sahel boundary was about 5° farther north, and there were more and larger lakes between 15 and 30° N. Simulations with climate models have shown that these wetter conditions were probably caused by changes in Earth's orbital parameters that increased the amplitude of the seasonal cycle of solar radiation in the Northern Hemisphere, enhanced the land-ocean temperature contrast, and thereby strengthened the African summer monsoon5–7. However, these simulations underestimated the consequent monsoon enhancement as inferred from palaeorecords4. Here we use a climate model to show that changes in vegetation and soil may have increased the climate response to orbital forcing. We find that replacing today's orbital forcing with that of the mid-Holocene increases summer precipitation by 12% between 15 and 22° N. Replacing desert with grassland, and desert soil with more loamy soil, further enhances the summer precipitation (by 6 and 10% respectively), giving a total precipitation increase of 28%. When the simulated climate changes are applied to a biome model, vegetation becomes established north of the current Sahara/Sahel boundary, thereby shrinking the area of the Sahara by 11% owing to orbital forcing alone, and by 20% owing to the combined influence of orbital forcing and the prescribed vegetation and soil changes. The inclusion of the vegetation and soil feedbacks thus brings the model simulations and palaeovegetation observations into closer agreement.