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.     

Late Quaternary biomes of Canada and the eastern United States

Pollen data have been used to construct biome maps for today, 6000 ¹⁴C yr bp and 18,000 ¹⁴C yr bp for Canada and the eastern United States. The inferred modern biome distributions agree well with independent reconstructions of North American vegetation prior to European settlement. Some discrepancies between the pollen data and the modern potential vegetation are caused by post‐settlement clearing of the landscape and the consequent increase of herbaceous types in the recent pollen record. Biome distributions at 6000 ¹⁴C yr bp reflected the warmer and drier conditions then prevalent in the continental interior, but the overall position of biomes was similar to that of today. The boreal treeline in North America was not significantly north of its present position, in contrast to the 100–200 km shift reported for Siberia. At the last glacial maximum (18,000 ¹⁴C yr bp ), steppe and tundra were prevalent in the Midwest and north‐western Canada, and coniferous forests and woodlands grew in eastern North America. The open vegetation at 18,000 ¹⁴C yr bp was probably due to drier conditions and/or lower concentrations of atmospheric CO₂. The composition and physical structure of biomes is not constant over time. Mid‐Holocene biomes were similar in structure to those of today, but shifts in the relative importance of individual plant functional types are large enough that the physical properties of biomes, such as albedo, canopy conductance and surface roughness, are likely to have varied even during the Holocene. Last glacial maximum biomes were structurally different from their modern counterparts. The biome maps therefore may obscure significant vegetational changes in space and time during the late Quaternary. The difference between the highest and next highest affinity scores for each sample measures how strongly affinity scores discriminate among biomes. For many biomes, the difference is not large, and affinity score ties are not uncommon, highlighting the importance of tie‐break procedures when using the biomization method.     

Biomes of western North America at 18,000, 6000 and 0 14C yr bp reconstructed from pollen and packrat midden data

A new compilation of pollen and packrat midden data from western North America provides a refined reconstruction of the composition and distribution of biomes in western North America for today and for 6000 and 18,000 radiocarbon years before present (¹⁴C yr bp ). Modern biomes in western North America are adequately portrayed by pollen assemblages from lakes and bogs. Forest biomes in western North America share many taxa in their pollen spectra and it can be difficult to discriminate among these biomes. Plant macrofossils from packrat middens provide reliable identification of modern biomes from arid and semiarid regions, and this may also be true in similar environments in other parts of the world. However, a weighting factor for trees and shrubs must be used to reliably reconstruct modern biomes from plant macrofossils. A new biome, open conifer woodland, which includes eurythermic conifers and steppe plants, was defined to categorize much of the current and past vegetation of the semiarid interior of western North America. At 6000 ¹⁴C yr bp , the forest biomes of the coastal Pacific North‐west and the desert biomes of the South‐west were in near‐modern positions. Biomes in the interior Pacific North‐west differed from those of today in that taiga prevailed in modern cool/cold mixed forests. Steppe was present in areas occupied today by open conifer woodland in the northern Great Basin, while in the central and southern Rocky Mountains forests grew where steppe grows today. During the mid‐Holocene, cool conifer forests were expanded in the Rocky Mountains (relative to today) but contracted in the Sierra Nevada. These differences from the forests of today imply different climatic histories in these two regions between 6000 ¹⁴C yr bp and today. At 18,000 ¹⁴C yr bp , deserts were absent from the South‐west and the coverage of open conifer woodland was greatly expanded relative to today. Steppe and tundra were present in much of the region now covered by forests in the Pacific North‐west.     

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.     

Pollen-based biome reconstruction for southern Europe and Africa 18,000 yr bp

Pollen data from 18,000 ¹⁴C yr bp were compiled in order to reconstruct biome distributions at the last glacial maximum in southern Europe and Africa. Biome reconstructions were made using the objective biomization method applied to pollen counts using a complete list of dryland taxa wherever possible. Consistent and major differences from present‐day biomes are shown. Forest and xerophytic woods/scrub were replaced by steppe, both in the Mediterranean region and in southern Africa, except in south‐western Cape Province where fynbos (xerophytic scrub) persisted. Sites in the tropical highlands, characterized today by evergreen forest, were dominated by steppe and/or xerophytic vegetation (cf. today’s Ericaceous belt and Afroalpine grassland) at the last glacial maximum. Available data from the tropical lowlands are sparse but suggest that the modern tropical rain forest was largely replaced by tropical seasonal forest while the modern seasonal or dry forests were encroached on by savanna or steppe. Montane forest elements descended to lower elevations than today.     

Last glacial maximum biomes reconstructed from pollen and plant macrofossil data from northern Eurasia

Pollen and plant macrofossil data from northern Eurasia were used to reconstruct the vegetation of the last glacial maximum (LGM: 18,000 ± 2000 ¹⁴C yr bp ) using an objective quantitative method for interpreting pollen data in terms of the biomes they represent ( Prentice et al., 1996 ). The results confirm previous qualitative vegetation reconstructions at the LGM but provide a more comprehensive analysis of the data. Tundra dominated a large area of northern Eurasia (north of 57°N) to the west, south and east of the Scandinavian ice sheet at the LGM. Steppe‐like vegetation was reconstructed in the latitudinal band from western Ukraine, where temperate deciduous forests grow today, to western Siberia, where taiga and cold deciduous forests grow today. The reconstruction shows that steppe graded into tundra in Siberia, which is not the case today. Taiga grew on the northern coast of the Sea of Azov, about 1500 km south of its present limit in European Russia. In contrast, taiga was reconstructed only slightly south of its southern limit today in south‐western Siberia. Broadleaved trees were confined to small refuges, e.g. on the eastern coast of the Black Sea, where cool mixed forest was reconstructed from the LGM data. Cool conifer forests in western Georgia were reconstructed as growing more than 1000 m lower than they grow today. The few scattered sites with LGM data from the Tien‐Shan Mountains and from northern Mongolia yielded biome reconstructions of steppe and taiga, which are the biomes growing there today.     

Pollen-based biome reconstructions for China at 0 and 6000 years

Biomization provides an objective and robust method of assigning pollen spectra to biomes so that pollen data can be mapped and compared directly with the output of biomgeographic models. We have tested the applicability of this procedure, originally developed for Europe, to assign modern surface samples from China to biomes. The procedure successfully delineated the major vegetation types of China. When the same procedure was applied to fossil pollen samples for 6000 years ago, the reconstructions showed systematic differences from present, consistent with previous interpretations of vegetation changes since the mid-Holocene. In eastern China, the forest zones were systematically shifted northwards, such that cool mixed forests displaced taiga in northeastern China, while broad-leaved evergreen forest extended c. 300 km and temperate deciduous forestc. 500–600 km beyond their present northern limits. In northwestern China, the area of desert and steppe vegetation was reduced compared to present. On the Tibetan Plateau, forest vegetation extended to higher elevations than today and the area of tundra was reduced. These shifts in biome distributions imply significant changes in climate since 6000 years ago that can be interpreted qualitatively as a response to orbital forcing and its secondary effects on the Asian monsoon.     

Pollen‐based biomes for Beringia 18,000, 6000 and 0 14C yr bp†

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.     

香果树属——秦岭北坡茜草科一新分布属

香果树一直被认为是中国亚热带中山或低山地区的落叶阔叶林或常绿、落叶混交林中的伴生树种,作者于2010～2011年在陕西周至秦岭北坡进行植物资源调查时,首次发现其野生种群分布,在暖温带的秦岭北坡发现香果树属植物——香果树,将中国该种自然分布区的纬度向北推移了0.5°。     

A biome classification of China based on plant functional types and the BIOME3 model

A biome classification for China was established based on plant functional types (PFTs) using the BIOME3 model to include 16 biomes. In the eastern part of China, the PFTs of trees determine mostly the physiognomy of landscape. Biomes range from boreal deciduous coniferous forest/woodland, boreal mixed forest/woodland, temperate mixed forest, temperate broad-leaved deciduous forest, warm-temperate broad-leaved evergreen/mixed forest, warm-temperate/cool-temperate evergreen coniferous forest, xeric woodland/scrub, to tropical seasonal and rain forest, and tropical deciduous forest from north to south. In the northern and western part of China, grass is the dominant PFT. From northeast to west and southwest the biomes range from moist savannas, tall grassland, short grassland, dry savannas, arid shrubland/steppe, desert, to alpine tundra/ice/polar desert. Comparisons between the classification introduced here and the four classifications which were established over the past two decades, i.e. the vegetation classification, the vegetation division, the physical ecoregion, and the initial biome classification have showed that the different aims of biome classifications have resulted in different biome schemes each with its own unique characteristics and disadvantages for global change study. The new biome classification relies not only on climatic variables, but also on soil factor, vegetation functional variables, ecophysiological parameters and competition among the PFTs. It is a comprehensive classification that using multivariables better expresses the vegetation distribution and can be compared with world biome classifications. It can be easily used in the response study of Chinese biomes to global change, regionally and globally.