Biome reconstructions for Japan 1 - Bristol · Biome reconstructions for Japan 2 (A) ABSTRACT 1 A...

Biome reconstructions for Japan 1

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

Hikaru Takahara1, Shinya Sugita2,3,4 Sandy P. Harrison5,6, Norio Miyoshi7, Yoshimune Morita8 and Takashi

Uchiyama9

1 University Forest, Kyoto Prefectural University, Kyoto 606, Japan.2 Department of Ecology, Evolution and Behavior, University of Minnesota, St. Paul, Minnesota 55108, USA.3 School of Ecology, Lund University, Sölvegatan 37, S-22362 Lund, Sweden.4 Department of Forest Resources, College of Agriculture, Ehime University, Matsuyama 790-8566, Japan.5 Dynamic Palaeoclimatology, Lund University, Box 117, S-22100 Lund, Sweden.6 Max Planck Institute for Biogeochemistry, Box 100164, D-07701 Jena, Germany.7 Department of Biosphere-Geosphere System Science. Faculty of Informatics, Okayama University of Science,

Okayama 700-0005, Japan.8 Research Insitute of Natural Science, Okayama University of Science, Okayama 700-0005, Japan.9 Department of Elementary Education, Chiba Keizai College, Chiba 263-0021, Japan.

Address for correspondance: Dr. H. Takahara, University Forest, Kyoto Prefectural University, Kyoto 606,

Japan (Fax: +81 75 703 5680, email: [email protected])

Ms. for: Journal of Biogeography, BIOME 6000 special issue

1 March, 2000


(A) ABSTRACT

1 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 B.P. 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.

2 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 B.P. are likely to be

conservative estimates of the actual changes.

3 The biome distribution at 6000 14C yr B.P. was rather similar to today, suggesting that changes in the

bioclimate of Japan have been small since the mid-Holocene.

4 At 18,000 14C yr B.P. 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.

Key words: pollen data, plant functional types, biomes, vegetation changes, Japan, mid-Holocene, last glacialmaximum,


(A) INTRODUCTION

Reconstruction of past vegetation patterns using palaeoecological data has generally relied on subjective

interpretations by individual investigators. Differences in the regional flora and vegetation classification schemes

have also made it difficult to compare maps of reconstructed vegetation. However, the recent development of an

objective method of biome reconstruction based on plant functional types (PFTs) rather than species (Prentice et

al., 1996; Prentice & Webb, 1998) has paved the way for producing palaeovegetation maps that are comparable

between different regions and continents. The resulting palaeovegetation maps are an important source of

information to validate simulations made with atmospheric general circulation models (e.g. Harrison et al., 1998;

Jolly et al., 1998) and coupled atmosphere-vegetation models (e.g. Texier et al., 1997).

Although the area of Japan is only 3.8 x 105 km2, it has a diverse vegetation and flora, covering sub-tropical to

subalpine environments (Yoshioka, 1973; Numata, 1974; Kira et al., 1976). Since there is high precipitation

throughout the year (ca 1,700 mm yr-1 on average), the potential vegetation is forest in most of Japan. Most of

the lowland area has never been glaciated (Suzuki, 1962; Ono, 1984), resulting in a relatively large number of

sites with sediment records back to the last glacial maximum (LGM, ca 18,000 14C yr B.P.) and beyond (e.g.

Yasuda, 1982; Miyoshi & Yano, 1986; Takahara & Takeoka, 1986, 1992b). Thus, Japan is one of the key areas

contributing to the reconstruction of global palaeovegetation patterns and climate at the LGM. Furthermore,

because of its geographical location, palaeo-records from Japan document changes in both the location and

strength of air masses in the northern Pacific and Siberia, and of the Asian monsoons, from the LGM to present.

Although the vegetation history of Japan since the LGM has been studied in detail over the last several decades

(Tsukada, 1988; and many others) and the density of studied sites is one of the highest in the world, the majority

of the records have not been accessible to the international community, and discussion of the implications of

palaeo-records from Japan in a global context has been limited.

In this paper, we apply the biomization method (Prentice et al., 1996) to reconstruct the biomes and vegetation

of Japan at 0, 6000, and 18,000 14C yr B.P., and discuss the implications of the results for our understanding of

vegetation and climate changes during the Late Quaternary. The main objectives of the paper are (1) to test the

biomization method (Prentice et al., 1996) using modern pollen/vegetation data from Japan, and (2) to map

pollen-based biomes assignments at 6000 and 18,000 14C yr B.P. at individual sites.

(A) DATA AND METHODS

(B) Pollen data for 0, 6000, 18,000 14C yr B.P.

In order to obtain a sufficient geographical coverage of pollen sites sampling all the major vegetation types, we

have used two forms of pollen data for both the modern and fossil data sets: (a) raw pollen counts; and (b) pollen

percentages digitized from published pollen diagrams. Raw pollen counts appear to provide a better

discrimination between non-arboreal biomes (Jolly et al., 1998; Yu et al., this issue). However, Prentice et al.,

(1996) showed that the use of raw pollen counts and digitized pollen data together does not affect the


biomization results for the forested regions of Europe, and a similar conclusion was reached by Tarasov et al.

(1998) for Russia. The vegetation of Japan has almost always been dominated by forests, even during the LGM

(Tsukada, 1988), so it is reasonable to use both forms of data here.

Modern pollen assemblages were extracted for 94 sites (Table 1). Raw pollen counts were obtained for 54 sites,

and digitized data were used for the rest. Of the 94 pollen assemblages, 93 were the top samples from a sediment

profile; the remaining assemblage was collected from the water/sediment interface of a lake. Descriptions of the

surrounding vegetation were obtained from the primary sources of the pollen data, i.e. from individuals who

provided raw pollen counts or from the original papers which included pollen diagrams.

The data set for 6000 14C yr B.P. consists of 58 sites (Table 2), and includes raw pollen counts from 31 sites and

digitized pollen percentages from 27 sites. At 34 sites, sediment chronologies were estimated by linear

interpolation between the 14C-dated horizons, and the nearest sample to 6000 14C yr B.P., provided it fell within

the window 6000 ± 500 14C yr B.P., was selected for inclusion in the data set. For the remaining 24 sites, the

widely-distributed Kikai-Akahoya ash (K-Ah), which is dated to 6300 14C yr B.P. (Machida & Arai, 1978,

1983), was used as a time marker for the selection of the 6000 14C yr B.P. samples. At these sites, the selected

pollen assemblage was from the sample immediately below the tephra and thus represents the vegetation at the

time the tephra was deposited. The use of the 6300 14C yr B.P. sample was considered preferable to sampling

above the tephra layer for two reasons. In practical terms, the transition from the tephra to the overlying

sediment is usually indistinct, and it is thus difficult to use the upper limit of the tephra as a dating horizon.

Secondly, although both the effects of volcanic emissions on forests and the recovery process of forest from

tephra-related damages are poorly understood (Saito, 1977; Franklin, 1988), a significant event like the

deposition of the K-Ah ash is likely to have caused local, short-term changes to both vegetation structure and

function. Pollen assemblages from such intervals will not therefore be representative of the long-term mean

climate and vegetation of the region.

Pollen assemblages at 18,000 14C yr B.P. were extracted for 15 sites (Table 2). Raw pollen counts were used at 5

sites and digitized data at 10 sites. All the selected pollen assemblages were dated 18,000 ± 2,000 14C yr B.P.,

based on a linear age-depth model using the 14

C-dated horizons. The quality of the dating control for 18,000 14C

yr B.P. samples varies (Table 2), but 8 sites have a dating control of 6D/3C or better according to the COHMAP

dating-control terminology (Webb, 1985; Yu & Harrison, 1995).

Pollen percentages for the 0, 6000, and 18,000 14C yr B.P. data were calculated based on the total sum,

excluding Alnus, a highly localized plant taxon mostly grown in marshy environments in Japan, aquatics

(Alisma, Haloragis, Iridaceae, Lysichiton, Lythrum, Menyanthes, Myriophyllum, Nuphar, Nymphaea,

Potamogeton, Rotala, Sagittaria, Sparganium, Trapa and Typha), generic monolete or trilete spores, Sphagnum,

Lycopodium spp., Gleichenia, Ophiloglossaceae, Osmunda, and unknown or unidentifiable grains.


(B) Biomization procedure

The biomization procedure is described by Prentice et al. (1996) and Prentice & Webb (1998). Biomization

requires (1) the assignment of individual pollen taxa to plant functional types (PFTs); (2) the specification of the

set of PFTs that can occur in each biome; (3) the calculation of the affinity score of a given pollen assemblage to

every biome; and (4) the assignment of the pollen assemblage to the biome for which it has the largest affinity

score. In cases where the affinity score for two or more biomes is equal, a tie-breaking rule is applied to

determine the biome attributed to the sample, following Prentice et al. (1996).

The pollen taxon-PFT matrix for Japanese plants (Table 3) was prepared based on our knowledge of the ecology

and biology of the individual plants, and on the descriptions of the flora and vegetation given in Kira & Yoshino

(1967), Yoshioka (1973), Numata (1974), Kira et al. (1976), Yamanaka (1979), Hattori & Nananishi (1985) and

Satake (1989). The matrix includes over 135 individual pollen taxa.

We adopted the PFT classification used for China (Yu et al., 1998; Yu et al., this issue), except for the cool-

temperate conifer (ctc) and temperate summergreen (ts) PFTs. We divided the cool-temperate conifer PFT into a

cool (ctc1) and an intermediate (ctc2) variant. Kira & Yoshino (1967) have shown that temperate conifers

occupy two distinct climate regions in Japan. One group, including Abies homolepis, Taxus cuspidata, Tsuga

diversifolia, Pinus parviflora, and Thuja standishii, occurs in subalpine and upper cool temperate forests, and

does not grow in warm temperate forests. The other group, including Abies firma, Tsuga sieboldii, Sciadopitys

verticillata, Cryptomeria japonica, Chamaecyparis obtusa, C. pisifera, Torreya nucifera, Pseudotsuga japonica

and Thujopsis dolabrata, grows in both the lower cool temperate and upper warm temperate forests. We

therefore define the former group of temperate conifers as ctc1, and the latter as ctc2, for the Japanese

biomization.

We add a lower cool-temperate summergreen (ts0) variant to the temperate summergreen (ts) PFT (Tables 3 and

4). This PFT comprises Fagus japonica, which is confined to temperate forests, where the warm-temperate

evergreen broadleaved taxa (e.g. Quercus subgenus Cyclobalanopsis and Castanopsis) do not occur. Definitions

of the other variants (ts1, ts2 and ts3) of the temperate summergreen PFT are consistent with those used in China

(Yu et al., 1998; Yu et al., this issue).

Two species of Picea (i.e. P. bicolor and P. polita) grow in the cool temperate forests in southwestern Japan,

although their ranges are extremely limited (Kira & Yoshino, 1967). We initially assigned Picea to ctc1 and

ctc2, as well as to the boreal evergreen conifer (bec) PFT. However, the biomization results were poor for both

the modern and the 6000 14C yr B.P. cases. In the final pollen taxon-PFT matrix, we assigned Picea only to bec

(Table 3).

The biome-PFT matrix (Table 4) was created from a look-up table (Table 5), which translates the vegetation

zones of Japan (Yoshioka, 1973; Fig. 1) into the nine biome types (as defined in Prentice et al., 1992, and

Prentice et al., 1996) which occur there. Biomes are assigned in the order they appear in Table 4. We created a


new biome (temperate conifer forest: TECO) to represent the warm temperate conifer forest defined by

Yoshioka (1973) (Fig. 1). This forest type, which is mainly characterized by ctc2 and ts0, occurs in areas where

the temperature of the coldest month (MTCO) is too low for warm-temperate evergreen broadleaved trees (wte)

but summer temperatures are too high for cool-temperate summergreen trees (ts1) such as Fagus crenata

(Numata, 1974; Kira et al., 1976; Hattori & Nakanishi, 1985). Kira et al. (1976) indicate that this forest type is

unique to Japan, a part of southern China and the montane region of the southern Himalaya. Its distribution in

Japan is rather limited because of heavy anthropogenic impacts.

The cold deciduous forest and taiga biomes include boreal summergreen taxa such as Larix and Betula (Prentice

et al., 1992, 1996). The subalpine conifer forests and the subarctic mixed broadleaved deciduous/coniferous

forests in Japan lack Larix gmelinii, a typical boreal summergreen conifer in eastern Eurasia. Although L.

leptolepis grows in the cool-temperate and subalpine zones in central Japan, this species does not occur in

Hokkaido, the northern most island of Japan today (Asakawa et al., 1981) and must therefore be less cold-

tolerant than L. gmelinii. We therefore conclude that cold deciduous forest and taiga (sensu Prentice et al., 1992,

1996) do not exist in modern Japan.

(A) RESULTS

(B) Predicted vs observed modern biomes

The pollen-derived biome map (Fig. 2b) shows patterns which mostly correspond to the observed patterns in

vegetation distribution (Figs 1 and 2a). Biomes at 51 sites, 54% of the 94 sites in the modern pollen data set are

correctly predicted by the biomization method (Table 6). Temperate deciduous forest is correctly predicted most

often (27 correct predictions out of 34 actual occurrences), broadleaved evergreen/warm mixed forest is second

(21 out of 32), and cool mixed forest is third (3 out of 10). The single tundra site in Hokkaido is predicted as

cool mixed forest. All cool conifer forest sites are incorrectly assigned either to cool mixed forest (7 sites) or

temperate deciduous forest (7 sites). All three temperate conifer sites are classified as temperate deciduous

forests. These systematic mismatches are well-depicted in the latitude-altitude diagram (Fig. 3b) and can be

explained as follows:

• The observed tundra site (142.9o E, 43.6o N, 1735 m alt.) occurs only ca 300 m above tree line in Hokkaido.

In such a situation, arboreal pollen from cool conifer and cool mixed forests below tree line, carried upwards

by orographic winds, apparently masks the local signal of tundra vegetation in the pollen assemblage (e.g.

Tsukada, 1958).

• All of the fourteen observed cool conifer forest sites (35.9°N- 40.0°N) are located 200-600 m above the

boundary between the cool-temperate and subalpine zones (Fig. 3a). The seven cool mixed forest sites, which

are misclassified as temperate deciduous forests, are also located close to the boundary with temperate

deciduous forest between 42.3°N- 44.1°N. Again, pollen carried by upward orographic winds is most likely

present in the pollen assemblages from these sites, influencing the biome predictions.

• The six observed temperate deciduous forest sites that are incorrectly assigned to temperate conifer forest are

mostly located close to the broadleaved evergreen/warm mixed forest zone. Pollen assemblages at the six


sites include large amounts of warm-temperate summergreen (ts3) taxa, affecting the biome prediction.

• Broadleaved evergreen/warm mixed forest is correctly predicted at 22 of the 32 sites where this biome

occurs. However, seven sites are incorrectly assigned to temperate deciduous forest, two sites to temperate

conifer forest and one site to tundra (Table 6). The sites incorrectly assigned to temperate deciduous and

temperate conifer forests are mostly located in the upper part of the broadleaved evergreen/warm mixed

forest zone, where temperate summergreen components or intermediate-temperate conifers (ctc2), especially

Cryptomeria japonica, are more abundant. Although broadleaved evergreen taxa grow throughout this zone,

they are less abundant at the upper, or northern, part of the range limits, partially caused by human impacts

(Numata, 1974) and partially because of interactions with temperate deciduous taxa influenced by the

bioclimate conditions (Hattori & Nahanishi, 1985).

• The pollen assemblage at one site in the broadleaved evergreen/warm mixed forest zone (134.0°E, 35.4°N,

640 m alt.) is heavily dominated by local Gramineae pollen, resulting in the prediction of tundra.

• There is no obvious explanation for why the three temperate conifer sites are all assigned to temperate

deciduous forest by the biomization method. However, they are all located within a limited area by the Sea of

Japan (134.5°E - 135.9°E and 35.3°N- 35.4°N) and may therefore be an atypical representation of the biome.

More modern samples from a wider geographical range are required to assess how well the biomization

method can effectively discriminate temperate conifer forest from other biomes.

The biomization of the modern data set defines some limits on the likely accuracy of the biome assignments

when the method is applied to the 6000 and 18,000 14C yr B.P. data sets:

• When broadleaved evergreen/warm mixed forest is assigned to the pollen sample, the biome prediction is

rather conservative but reliable.

• When temperate deciduous forest is assigned, the predicted biome is less certain; the actual biome could also

be cool conifer, cool mixed, temperate deciduous, temperate conifer or broadleaved evergreen/warm mixed

forest.

• Tundra will likely be assigned when local non-arboreal pollen types, especially Gramineae and Cyperaceae,

are abundant; in the modern case, such an assignment may result either from heavy anthropogenic impact or

the inclusion of sites that represent local, azonal conditions; at 6000 and 18,000 14C yr B.P. it will occur if

sites are not carefully selected to represent the regional vegetation rather than to sample a local phenomenon.

• Further validation of biomes that are more widely distributed in Japan, such as temperate conifer forest, cool

mixed forest and cool conifer forest, is necessary to determine how well we can reconstruct these biomes in

the past.

(B) Mid-Holocene biomes

The predicted distribution of biomes at 6000 14C yr B.P. (Figs 2c and 3c) shows that the northern limit of the

broadleaved evergreen/warm mixed forest was ca 36°N. There are no sites beyond the modern northern limit of

this biome (Fig. 2a). However, one site (35.2°N, 134.1°E, 970 m alt.) assigned to this biome is located above the

modern boundary between the broadleaved evergreen/warm mixed and cool temperate deciduous forest zones

(Fig. 3c). The modern biomization indicates that the prediction of broadleaved evergreen/warm mixed forest is


conservative but realistic, suggesting that this biome may have occurred at higher elevations in central Japan

during the mid-Holocene. Temperate conifer forests are assigned to four sites located at the modern northern

and/or upper limits of the broadleaved evergreen/warm mixed forest zone. Given that we are unable to predict

temperate conifer forest in the modern biomization, and that modern assemblages from the temperate deciduous

forest sites are frequently assigned to temperate conifer forest, it is unlikely that the temperate conifer forest was

more widely distributed than today. The temperate deciduous forest sites are confined to south of ca 42°N today

(Figs 2a and 3a), but two sites at ca 44°N are classified as temperate deciduous forest at 6000 14C yr B.P.

Temperate deciduous forests are predicted at elevations up to ca 500 m higher than today in central Japan.

However, both the latitudinal and elevational extent of the biome is overestimated in the modern biomization,

and thus it is difficult to assess the significance of the apparent shift in this biome between today and 6000 14C yr

B.P. There is no apparent change in the extent of cool mixed forest between today and 6000 14C yr B.P. The

realism of the prediction of tundra at a single site (36.6°N, 137.6°E, 2440 m alt.) in central Japan is difficult to

judge. The pollen assemblage is dominated by Cyperaceae, rather than by arctic/alpine forbs, and could

represent an atypical, local environment. This site is, however, located near the modern timberline, and if our

prediction is right, the timberline would not have shifted at 6000 14C yr B.P. Further investigation is clearly

needed to clarify the location of timberline at 6000 14C yr B.P.

(B) Last glacial maximum biomes

In contrast to the situation at 6000 14C yr B.P., the predicted distribution of biomes at 18,000 14C yr B.P. was

significantly different from today (Figs 2d and 3d). Taiga is predicted at lowland sites in Hokkaido and

northeastern Honshu, and cool mixed forest is predicted at lowland and high elevation sites elsewhere. The only

exception to this pattern is a single site (35.4°N, 134.6°E, 610 m alt.) at relatively low elevations in central

Japan, which is classified as temperate deciduous forest because of the low abundance of Picea and relatively

high abundance of Betula, Quercus subgenus Lepidobalanus and Ulmus-Zelkova, temperate summergreen (ts)

taxa, in the 18,000 14C yr B.P. assemblage.

(A) DISCUSSION AND CONCLUSIONS

(B) The biomization method

The biomization procedure has been shown to work rather well in a number of different regions and across a

range of climates and vegetation types (Prentice et al., 1996; Prentice & Webb, 1998; Jolly et al., 1998; Tarasov

et al., 1998; Yu et al., 1998; papers in this issue). However, the focus in other regions has been on mapping at a

sub-continental scale, and the majority of sites used were from non-mountainous regions. Japan is the first region

where it has been necessary to examine how well the biomization procedure works in complex mountain

topography. Our application of the biomization method to modern pollen data from Japan resulted in the correct

classification of over half of the samples, which we consider an acceptable match between the observed and

predicted biomes. Even more encouragingly, most of the mismatches appear to reflect systematic biases that can

be understood in terms of the nature of pollen transport in mountainous terrain.


The application of the biomization procedure to derive vegetation maps assumes that the majority of pollen in

the assemblages comes primarily from the regional vegetation zone (Prentice et al., 1996). In lowland or plateau-

type environments this assumption would hold true for most of the sites typically sampled for pollen analysis.

The biomization of the modern Japanese data clearly shows that sites in mountainous terrain, particularly those

near the lower altitudinal limits of a given biome, can be apparently misclassified rather frequently. Nearly 25%

of the modern sites are misclassified to the biome which occupies the altitudinal vegetation zone lower than the

one in which the site actually occurs. This reflects the fact that the areal extent of individual biomes in

mountainous terrain is limited, and that pollen coming from biomes at lower elevations overwhelms the pollen

coming from within individual biomes. In addition, differences in pollen dispersal characteristics particularly

affect pollen assemblages in higher elevations. For example, Quercus pollen, a dominant pollen type in

temperate deciduous forests, can be transported by wind much more effectively than pollen of Picea and Abies

(Prentice, 1988; Sugita, 1993) which are major components of cool mixed and cool conifer forests. Daytime

upward orographic wind could also enhance over-representation of pollen types from lower down the mountain

(e.g. Tsukada, 1958). The misclassification of pollen assemblages from sites in mountainous terrain has been

seen in other regions (e.g. Prentice et al., 1996; Jolly et al., 1998) However, the biomization of modern samples

from Japan has demonstrated that this is a systematic bias and is not confined to the definition of upper treelines.

This systematic bias could be reduced by using basin size as a site-selection criterion (Prentice, 1985; Sugita,

1993, 1994), so as to include only those sites which sample the vegetation at the spatial scale appropriate to

reconstruct changes over relatively short distances along an altitudinal gradient.

The systematic over-representation of temperate deciduous forest in the modern biomization is not confined to

the upper limit of the biome. Temperate deciduous forest is also predicted at lower elevation sites where the

observed vegetation is temperate conifer forest or even broadleaved evergreen/warm mixed forest. This results in

a misclassification of 11% of the sites in the modern pollen data. In part, this misclassification reflects the poor

taxonomic resolution in the pollen data sets. For example, there are important differences in the bioclimatic

limits of temperate summergreen taxa (e.g. Fagus crenata vs. F. japonica, Carpinus tschonoskii vs. C. japonica,

Quercus subgenus Cyclobalanopsis vs. Q. subgenus Lapidobalanus) and cool-temperate conifer taxa (e.g. Tsuga

sieboldii vs. T. diversifolia). Further improvement in pollen identification of these taxa (e.g. Takahara &

Takeoka, 1992b) would make it possible to distinguish temperate deciduous forest from temperate conifer forest

more reliably. The inclusion of macrofossil data, which generally has a better taxonomic resolution, should also

result in an improved discrimination between biomes (Jolly et al., 1998; Thompson & Anderson, this issue). It is

possible that the use of raw pollen counts, rather than digitized data, at all sites would also result in improved

discrimination between temperate deciduous forest and other forest biomes, by allowing minor taxa with a more

specific distribution to influence the biome affiliation.

In summary, improvements to the application of the biomization procedure for mapping the vegetation of

mountainous regions such as Japan are likely to be achieved by: (a) applying a more rigorous set of site-selection

criteria in order to exclude sites which are sampling either mostly local, atypical vegetation or an area which is

inappropriately large for mountainous terrain; (b) using macrofossil data in conjunction with pollen data in order


to improve the taxonomic resolution of the assemblages used for biomization; and (c) using raw pollen counts at

all sites in order to improve the range of species taken into account in the biomization procedure. Finally,

although the climate constraints on the distribution of most conifers and broadleaved trees in Japan are well-

studied (e.g. Kira 1977a, 1977b; Kira & Yoshino 1967), better bioclimate information is required in order to

assign many other species to specific PFTs more objectively (Prentice et al., 1992, 1996).

(B) Vegetation and climate of Japan at 6000 14C yr B.P.

Our biomization suggests that the vegetation distribution at 6000 14C yr B.P. was rather similar to present. On

the basis of the pollen record from a single site, the broadleaved evergreen/warm mixed forest may have been

present at higher elevations in the mountains of central Japan. However, the northern limit of the biome was

apparently similar to present. The modern biomization indicates that broadleaved evergreen/warm mixed forest

is only predicted when it is actually present. If it is true that this biome occurred at higher elevations than today,

it might be expected that other forest biomes (including temperate deciduous and temperate conifer forests)

would also have occurred at higher elevations. However, there does not seem to be any robust evidence for a

northward or upward expansion of individual biomes in Japan at 6000 14C yr B.P.

Previous reconstructions of mid-Holocene vegetation changes in Japan have suggested that broadleaved

evergreen/warm mixed forest was less extensive than today (Tsukada, 1988; Tsuji, 1989) while temperate

deciduous forests (Takahara et al., 1995, 1997) and temperate conifer forests (Tsukada, 1982, 1986; Takahara &

Takeoka, 1992a, 1992b; Takahara, 1994) were more extensive. Although our biome reconstructions may be

consistent with the proposed extention of temperate deciduous and conifer forests, the evidence for such an

extension does not appear to be particularly strong. Furthermore, our reconstruction does not show any evidence

of a reduction in the extent of broadleaved evergreen/warm mixed forests.

The lack of strong evidence for any latitudinal shift in biomes is in stark contrast to the evidence from mainland

China (Yu et al., this issue). The northern limit of broadleaved evergreen/warm mixed forest in China was at

35oN - 36oN at 6000 14C yr B.P., ca 200 km further north than today. Temperate deciduous forest occurred as far

north as ca 48oN, i.e. 800 km north of its present limit, in the zone occupied today by cool mixed forest and

taiga. Yu et al. (this issue) argue that these latitudinal shifts imply that the winters at 6000 14C yr B.P. were

warmer than present and, since this is contrary to the direct response to insolation changes (Berger, 1978), must

reflect an indirect climatic response acting through a change in atmospheric and/or oceanic circulation patterns.

Our results, which show that there is no northward shift in broadleaved evergreen/warm mixed and temperate

deciduous forests in Japan, make it impossible that winter warming in eastern China can be explained by a

change in oceanic circulation.

(B) Vegetation and climate of Japan at the last glacial maximum

Our biomization shows that taiga and cool mixed forests occurred in Japan at the LGM. Broadleaved

evergreen/warm mixed forests were not recorded, and temperate deciduous forest only occurred at a single site.


The reconstruction of taiga and cool mixed forests in northern and central Japan is consistent with previous

studies (e.g. Tsukada, 1985; Kamei et al., 1981). However, earlier studies have suggested that southwestern

Japan was covered by temperate conifer forests at the LGM (Tsukada, 1983, 1985; Takahara & Takeoka,

1992b). We do not predict temperate conifer forests at any site. However, the affinity score of the temperate

conifer biome was identical to the score for temperate deciduous forest at the single site we classify as temperate

deciduous forest (Ohnuma: 134.6°E, 35.4oN, 610 m alt.), and this site was only attributed to temperate deciduous

forest on the basis of the tie-breaking rule (Prentice et al., 1996). Thus, temperate conifer forest may have been

present in southwestern Japan at the LGM. More detailed palaeoecological studies including macrofossil

analysis, will be necessary to determine whether temperate conifer forests occurred in southwestern Japan at the

LGM.

The dominance of taiga and cool mixed forests in Japan at 18 ka reflects a significant, and most likely year

round, cooling. This is consistent with the year-round cooling implied by vegetation change in China (Yu et al.,

this issue) and in Beringia (Edwards et al., this issue). Our results are also consistent with the expected climate

changes due to the presence of large ice sheets in the northern hemisphere (Manabe & Broccoli, 1985; Kutzbach

& Guetter, 1986; Harrison et al., 1992; Kutzbach et al., 1998). The southward expansion of taiga in Japan was

larger than in lowland China, although the southward expansion of taiga in China may have been partly limited

by aridity. This suggests that the global signal of cooling may have been amplified by regional influences,

specifically conditions in the Sea of Japan (Kamei et al., 1981; Oba et al., 1991, 1995; Tada, 1997). The partial

or complete closure of the Korean and Tsushima Straits (Kamei et al., 1981; Japan Association for Quaternary

Research, 1987) would have blocked the flows of the Tsushima Current producing significant cooling in

adjacent land areas, including Japan. Whether the magnitude of this effect is sufficient to explain why taiga

occurred further south than in China could be investigated using a mesoscale climate model (e.g. Giorgi et al.,

1993a, 1993b).

(A) ACKNOWLEDGEMENTS

The biomization described here was begun at the BIOME 6000 regional workshop for Beringia and Japan

(October 15th - 29th, 1997, Lund Sweden), which was funded by the International Geosphere-Biosphere

Programme (IGBP) through the IGBP Data and Information System (IGBP-DIS), the Global Analysis,

Interpretation and Modelling (GAIM) task force, and the PAst Global changES (PAGES) core project. Sugita

was supported by a guest scientist grant from the Swedish Natural Science Research Council (NFR). Technical

assistance with data compilation from F. Dobos is greatly acknowledged. We are grateful to John Dodson, Yugo

Ono, Colin Prentice and Dominique Jolly for discussion and comments on earlier versions of this paper, and to

Ben Smith, who created the programme BIOMISE which was used here to carry out the biomization procedure.

This paper is a contribution to BIOME 6000, to TEMPO (Testing Earth System Models, with

Palaeoenvironmental Observations) and to the international Palaeoclimate Modelling Intercomparison Project

(PMIP). This paper is also a contribution to the Japanese Pollen Data Base. The Japanese data are available from

the BIOME 6000 website (http://www.bgc-jena.mpg.de/bgc_prentice/).


(A) BIOPIC

The Japanese Pollen Database. All the raw pollen counts compiled and used for this paper are included in the

Japanese Pollen Database, which started in 1995 with a two-year Grant-in-Aid from the Ministry of Education,

Science, Sports and Culture of Japan to the International Center for Japanese Studies (coordinator: Y. Yasuda)

and partially supported by IGBP PAGES of Japan (coordinator: Y. Ono). The Japanese Pollen Database

archives pollen and macrofossils from Japan, along with site-specific information and chronological data, in

machine-readable form. The database is under development by H. Takahara at Kyoto Prefectural University, Y.

Yasuda at International Center for Japanese Studies, and Y. Ono at Hokkaido University.


REFERENCES

Asakawa, S., Katsuta, M. & Yokoyama, T. (1981) Seeds of woody plants in Japan: Gymnospermae. Japan

Forest Tree Breeding Association, Tokyo. (in Japanese)

Berger, A. (1978) Long term variations of daily insolation and Quaternary climate changes. Journal of

Atmospheric Sciences 35, 2362-2376.

Edwards, M.E., Anderson, P.M., Brubaker, L.B., Ager, T., Andreev, A.A., Bigelow, N.H., Cwynar, L.C., Eisner,

W.R., Harrison, S.P., Hu, F.-S., Jolly, D., Lozhkin, A.V., McDonald, G.M., Mock, C.J., Ritchie, J.C., Sher,

A.V., Spear, R.W., Williams, J. & Yu, G. (this issue) Pollen-based biomes for Beringia 18,000, 6000 and 014C yr B.P. Journal of Biogeography.

Franklin, J.F. (1988) Pacific Northwest forest. North American terrestrial vegetation (ed. by M.G. Barbour &

W.D. Billings), pp. 103-130. Cambridge University Press, Cambridge.

Giorgi, F., Marinucci, M.R. & Bates, G.T. (1993a) Development of a second-generation regional climate model

(RegCM2). Part 1: Boundary-layer and radiative transfer processes. Monthly Weather Review 121, 2794-

2813.

Giorgi, F., Marinucci, M.R. & Bates, G.T. (1993b) Development of a second-generation regional climate model

(RegCM2). Part 2: Convective processes and assimilation of lateral boundary conditions. Monthly Weather

Review 121, 2814-2832.

Harrison, S.P., Prentice, I.C. & Bartlein, P.J. (1992) Influence of insolation and glaciation on atmospheric

circulation in the North Atlantic sector: implications of general circulation model experiments for the Late

Quaternary climatology of Europe. Quaternary Science Reviews 11, 283-299.

Harrison, S.P., Jolly, D., Laarif, F., Abe-Ouchi, A., Dong, B., Herterich, K., Hewitt, C., Joussaume, S.,

Kutzbach, J.E., Mitchell, J., de Noblet, N. & Valdes, P. (1998) Intercomparison of simulated global

vegetation distribution in response to 6kyr B.P. orbital forcing. Journal of Climate 11, 2721-2742.

Hattori, T. & Nakanishi, S. (1985) On the distributional limits of the lucidophyllous forest in the Japanese

Archipelago. Botanical Magazine, Tokyo 98, 317-333.

Japan Association for Quaternary Research (ed.) (1987) Quaternary maps of Japan. University Tokyo Press,

Tokyo.

Jolly, D., Prentice, I.C., Bonnefille, R., Ballouche, M.B., Brenac, P., Buchet, C., Burney, D., Cazet, J.-P.,

Cheddadi, R., Edorh, T., Elenga, H., Elmoutaki, S., Guiot, J., Laarif, F., Lamb, H., Lezine, A.-M., Maley,

J., Mbenza, M., Peyron, O., Reille, M., Reynaud-Farrera, I., Riollet, G., Ritchie, J.C., Roche, E., Scott, L.,

Ssemmanda, I., Straka, H., Umer, M., Van Campo, E., Vilimumbalo, S., Vincens, A. & Waller, M. (1998)

Biome reconstruction from pollen and plant macrofossil data for Africa and the Arabian peninsula at 0 and

6 ka. Journal of Biogeography 25, 1007-1028.

Kamei, T. & Research Group for the Biogeography from WÜRM GLACIAL (1981) Fauna and flora of the

Japanese Islands in the Last Glacial Time. Daiyonki-Kenkyu (Quaternary Research) 20, 191-205. (in

Japanese with English summary)

Kikuchi, T. (1985) Vegetation zones. Organisms of Japan, a country of mountains (ed. by M. Horikoshi & J.

Aoki). Iwanami-shoten, Tokyo.

Kira, T. (1977a) Climatological interpretation of Japanese vegetation zones. Vegetation science and


environmental protection (ed. by A. Miyawaki & R. Tüxen), pp. 21-30. Maruzen, Tokyo.

Kira, T. (1977b) Forest Vegetation of Japan 1.1 Introduction. Primary productivity of terrestrial communities-

JIBP Synthesis 16 (ed. by T. Shidei & T. Kira), pp. 1-9. University of Tokyo Press, Tokyo.

Kira, T. & Yoshino, M. (1967) Thermal distribution of Japanese conifer species. Contributions in Honor of Dr.

Kinji Imanishi on the occasion of his sixtieth birthday, Vol. 1, Natural history-ecological studies (ed. by

M. Morisita & T. Kira), pp. 133-161. Chuo-Koron Sha, Tokyo. (in Japanese)

Kira, T., Shidei, T., Numata, M. & Yoda, K. (1976) Vegetation of Japan - its ordination in the global system of

vegetation. Kagaku 46, 235-247. ( in Japanese)

Kutzbach, J.E. & Guetter, P.J. (1986) The influence of changing orbital parameters and surface boundary

conditions on climate simulations for the past 18000 years. Journal of the Atmospheric Sciences 43, 1726-

1759.

Kutzbach, J.E., Gallimore, R., Harrison, S.P., Behling, P., Selin, R. & Laarif, F. (1998) Climate and biome

simulations for the past 21,000 years. Late Quaternary climates: data syntheses and model experiments.

(ed. by T. Webb III), Quaternary Science Reviews 17, 473-506.

Machida, H. & Arai, F. (1978) Akahoya ash - a Holocene widespread tephra erupted from the Kikai Caldera,

south Kyushu, Japan. Daiyonki-Kenkyu (Quaternary Research) 17, 143-163. (in Japanese with English

summary)

Machida, H. & Arai, F. (1983) Widespread late Quaternary tephras in Japan with special reference to

archaeology. Daiyonki-Kenkyu (Quaternary Research) 22, 133-148. (in Japanese with English summary)

Manabe, S. & Broccoli, A.J. (1985) The influence of continental ice sheets on the climate of an ice age. Journal

of Geophysical Research 90, 2167-2190.

Miyoshi, N. & Yano, N. (1986) Late Pleistocene and Holocene vegetational history of the Ohnuma Moor in the

Chugoku Mountains, western Japan. Review of Palaeobotany and Palynology 46, 355-376.

Numata, M. (1971) Ecological interpretation of vegetational zonation of high mountains, particularly in Japan

and Taiwan. Landschaftsökologie der Hochgebirge Eurasiens (ed. by C. Troll), pp. 288-299.

Erdwissenshaftliche Forschung.

Numata, M. (ed.) (1974) The flora and vegetation of Japan. Kodansha, Tokyo.

Oba, T., Kato, M., Kitazato, H., Koizumi, I., Omura, A., Sakai, T. & Takayama, T. (1991) Palaeoenvironmental

changes in the Japan Sea during the last 85,000 years. Paleoceanography 6, 499-518.

Oba, T., Murayama, M., Matsumoto, E. & Nakamura, T. (1995) AMS- 14C ages of Japan Sea cores from the Oki

Ridge. Daiyonki-Kenkyu (Quaternary Research) 34, 289-296. (in Japanese with English summary)

Ono, Y. (1984) Last glacial paleoclimate reconstructed from glacial and periglacial landforms in Japan.

Geographical Review of Japan (Series B) 57, 87-100.

Prentice, I.C. (1985) Pollen representation, source area, and basin size: toward a unified theory of pollen

analysis. Quaternary Research 23, 76-86.

Prentice, I.C. (1988) Records of vegetation in time and space: the principles of pollen analysis. Vegetation

history (ed. by B. Huntley & T. Webb III), pp. 17-42. Kluwer Academic Publishers, Dordrecht.

Prentice, I.C. & Webb III, T. (1998) BIOME 6000: reconstructing global mid-Holocene vegetation patterns

from palaeoecological records. Journal of Biogeography 25, 997-1005.

Prentice, I.C., Cramer, W., Harrison, S.P., Leemans, R., Monserud, R.A. & Solomon, A.M. (1992) A global


biome model based on plant physiology and dominance, soil properties and climate. Journal of

Biogeography 19, 117-134.

Prentice, I.C., Guiot, J., Huntley, B., Jolly, D. & Cheddadi, R. (1996) Reconstructing biomes from

palaeoecological data: a general method and its application to European pollen data at 0 and 6 ka. Climate

Dynamics 12, 185-194.

Saito, K. (1977) Volcanic zone. Distribution and environments of plant communities (ed. by K. Ishizuka), pp.

307-394. Asakura-shoten, Tokyo. (in Japanese)

Satake, Y. (1989) Wild flowers of Japan. Woody plants I and II. Heibun-Sha, Tokyo.

Sugita, S. (1993) A model of pollen source area for an entire lake surface. Quaternary Research 39, 239-244.

Sugita, S. (1994) Pollen representation of vegetation in Quaternary sediments: theory and method in patchy

vegetation. Journal of Ecology 82, 881-897.

Suzuki, H. (1962) Southern limit of peri-glacial landforms at low level and the climatic classification of the latest

ice age in Japan. Geographical Review of Japan 35, 67-76. (in Japanese with English summary)

Tada, R. (1997) Paleoenvironmental changes in and around the Japan Sea since the Last Glacial Period.

Daiyonki-Kenkyu (Quaternary Research) 36, 287-300. (in Japanese with English summary)

Takahara, H. (1994) Vegetation history since the last glacial period in the Kinki & the eastern Chugoku regions,

western Japan. Bulletin of Kyoto Prefectural University, Forestry 38, 89-112. (in Japanese with English

summary)

Takahara, H. & Takeoka, M. (1986) Vegetational changes since the last glacial maximum around the

Hatchodaira Moor, Kyoto, Japan. Japanese Journal of Ecology 36, 105-116. (in Japanese with English

summary)

Takahara, H. & Takeoka, M. (1992a) Postglacial vegetation history around Torishima, Fukui Prefecture, Japan.

Ecological Research 7, 79-85.

Takahara, H. & Takeoka, M. (1992b) Vegetation history since the last glacial period in the Mikata lowland, the

Sea of Japan area, Western Japan. Ecological Research 7, 371-386.

Takahara, H., Nishida, S. & Takemura, K. (1995) Vegetation history since the late-glacial period around the

Ikenokochi Moor, Fukui Prefecture, Japan. Japanese Journal of Palynology 41, 99-108. (in Japanese with

English summary)

Takahara, H. Fujiki, T., Miyoshi, N. & Nishida, S. (1997) Vegetation history since the middle Holocene around

Orogatawa Moor, Okayama Prefecture, Japan. Japanese Journal of Palynology 43, 97-106. (in Japanese

with English summary)

Tarasov, P.E., Webb III, T., Andreev, A.A., Afanas'eva, N.B., Berezina, N.A., Bezusko, L.G., Blyakharchuk,

T.A., Bolikhovskaya, N.S., Cheddadi, R., Chernavskaya, M.M., Chernova, G.M., Dorofeyuk, N.I., Dirksen,

V.G., Elina, G.A., Filimonova, L.V., Glebov, F.Z., Guiot, J., Gunova, V.S., Harrison, S.P., Jolly, D.,

Khomutova, V.I., Kvavadze, E.V., Osipova, I.R., Panova, N.K., Prentice, I.C., Saarse, L., Sevastyanov,

D.V., Volkova, V.S. & Zernitskaya, V.P. (1998) Present-day and mid-Holocene biomes reconstructed from

pollen plant macrofossil data from the Former Soviet Union and Mongolia. Journal of Biogeography 25,

1029-1053.

Texier, D., de Noblet, N., Harrison, S.P., Haxeltine, A., Joussaume, S., Jolly, D., Laarif, F., Prentice, I.C. &

Tarasov, P.E. (1997) Quantifying the role of biosphere-atmosphere feedbacks in climate change: a coupled


model simulation for 6000 yr B.P. and comparison with palaeodata for Northern Eurasia and northern

Africa. Climate Dynamics 13, 865-881.

Thompson, R. & Anderson, K. (this issue) Biomes of western North America at 18,000, 6000 and 0 14C yr B.P.

reconstructed from pollen and packrat midden data. Journal of Biogeography.

Tsuji, S. (1989) Paleoenvironmental reconstruction in the dissected valley in prehistory: Holocene

paleoenvironment at the Akayama site in Kawaguchi, central Kanto Plain, Japan. Daiyonki-Kenkyu

(Quaternary Research) 27, 331-356. (in Japanese with English summary)

Tsukada, M. (1958) Untersuchungen über das Verhältniss zwischen fem Pollengehalt der Oberflächenproben

und der Vegetation des Hochlandes Shiga. Journal of Institute of Polytechnics, Osaka City University,

Series D 9, 235-249.

Tsukada, M. (1982) Cryptomeria japonica: glacial refugia and late glacial and post glacial migration. Ecology

63, 1091-1105.

Tsukada, M. (1983) Vegetation and climate during the Last Glacial Maximum in Japan. Quaternary Research

19, 212-235.

Tsukada, M. (1985) Map of vegetation during the Last Glacial Maximum in Japan. Quaternary Research 23,

369-381.

Tsukada, M. (1986) Altitudinal and latitudinal migration of Cryptomeria japonica for the past 20,000 years in

Japan. Quaternary Research 26, 135-152.

Tsukada, M. (1988) Japan. Vegetation history (ed. by B. Huntley & T. Webb III), pp. 459-518. Kluwer

Academic Publishers, Dordrecht.

Webb III, T. (1985) A Global Paleoclimate Data Base for 6000 yr B.P. DOE/eV/10097-6, pp. 155. US

Department of Energy, Washington DC.

Yamanaka, T. (1979) Forest vegetation of Japan. Tsukiji-Shokan, Tokyo. (in Japanese)

Yasuda, Y. (1982) Pollen analytical study of the sediment from the Lake Mikata in Fukui prefecture, central

Japan. Daiyonki-Kenkyu (Quaternary Research) 21, 255-271. (in Japanese with English summary)

Yoshioka, K. (1973) Plant geography. Kyoritsu-Shuppan, Tokyo. (in Japanese)

Yu, G. & Harrison, S.P. (1995) Lake status records from Europe: Data base documentation. NOAA

Paleoclimatology Publications Series Report 3, 451 pp.

Yu, G., Prentice, I.C., Harrison, S.P. & Sun, X. (1998) Pollen-based biome reconstructions for China at 0 ka and

6 ka. Journal of Biogeography 25, 1055-1069.

Yu, G., Chen, X., Ni, J., Cheddadi, R., Guiot, J., Han, H., Harrison, S.P., Huang, C., Ke, M., Kong, Z., Li, S., Li,

W., Liew, P., Liu, G., Liu, J., Liu, Q., Liu, K., Prentice, I.C., Qui, W., Ren, G., Song, C., Sugita, S., Sun,

X., Tang, L., van Campo, E., Xia, Y., Xu, Q., Yan, S., Yang, X., Zhao, J. & Zheng, Z. (this issue)

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

Journal of Biogeography.

REFERENCES FOR THE DATA SETS

Choi, K. & Hibino, K. (1985) Palynological study of the Miyatokooyachi moor in Fukushima Prefecture. Report

of Miyagi Agricultural College 33, 81-82. (in Japanese)


Endo, K., Kosugi, M., Matsushita, M., Miyaji, N., Hishida, R. & Takano, T. (1989) Holocene environmental

history in and around the Paleo-Nagareyama Bay, Central Kanto Plain. Daiyonki-Kenkyu (Quaternary

Research) 28, 61-77. (in Japanese with English summary)

Furutani, M. (1979) Studies on the forest history in the Osaka area since Wurm glacial age in Japan. Daiyonki-

Kenkyu (Quaternary Research) 18, 121-141. (in Japanese with English summary)

Hatanaka, K. (1982) Palynological study of the sediments from Odano-ike Moor in Ooita Prefecture. Journal of

Faculty of Literature, Kitakyushu University (Series B) 15 , 113-119. (in Japanese with English summary)

Hatanaka, K. (1985) Palynological studies on the vegetational succession since the Würum Glacial Age in

Kyushu and adjacent areas. Journal of Faculty of Literature, Kitakyushu University (Series B) 18, 29-71.

Hatanaka, K. & Miyoshi, N. (1980) History of the late Pleistocene and Holocene vegetation in the Ubuka Basin,

southwestern Japan. Japanese Journal of Ecology 30, 239-244. (in Japanese with English summary)

Hibino, K. (1991) Sediments and pollen analysis of Shibayachi bog. Report of Scientific Research for the

Vegetation of Shibayachi Bog, pp. 63-74. (in Japanese)

Hibino, K & Sasaki, M. (1982) Palynological studies in the northwestern part of Nagano Prefecture, Central

Honshu, Japan. Report of Miyagi Agricultural College 30, 93-101. (in Japanese with English summary)

Hibino, K. & Horie, S. (1991) Pollenanalytische Forschungen über Vegetationsveränderungen nach der letzten

Eiszeit im Kizaki-See, Präfektur Nagano. Die Geschichte des Biwa-Sees in Japan (ed. by S. Horie), pp.

285-295. Universitatsverlag Wagner, Innsbruck.

Hibino, K., Morita, Y., Miyagi, T. & Yagi, H. (1991) Palynological study on the vegetational change during the

last 120,000 years B.P. in Kawadoi Basin, Yamagata Prefecture, Japan. Report of Miyagi Agricultural

College 39, 35-49. (in Japanese with English summary)

Igarashi, Y., Igarashi, T., Daimaru, H., Yamada, O., Miyagi, T., Matsushita, K. & Hiramatsu, K. (1993)

Vegetation history of Kenbuchi Basin and Furano Basin in Hokkaido, north Japan, since 32,000 yrs BP.

Daiyonki-Kenkyu (Quaternary Research) 32, 89-105. (in Japanese with English summary)

Ishida, S., Kawada, K. & Miyamura, M. (1984) Geology of the Hikone-Seibu District. Quadrangle series scale

1:50,000 Kyoto (11) No. 17. Geological Survey of Japan, Tokyo. (in Japanese)

Iwauchi, A. & Hase, Y. (1992) Vegetational changes after the Last Glacial Age in the Kumamoto Plain and Aso

Caldera areas. Japanese Journal of Palynology 38, 116-133. (in Japanese with English summary)

Kanauchi, A., Tahara, Y., Sugihara, S. & Koaze, T. (1988) Ages and pollen analysis of cores from the

Yashimagahara peat bog, central Japan. Abstract of the Annual Meeting, Japan Association for Quaternary

Research 18, 156-157. (in Japanese)

Kanauchi, A., Sugihara, S. & Koaze, T. (1989a) Stratigraphy and pollen analysis of the Yashimagahara bog,

Nagano Prefecture and Ominemuma bog, Gunma Prefecture. Abstract of the Spring Meeting, Association

of Japanese Geographers. (in Japanese with English summary)

Kanauchi, A., Tahara, Y., Nakamura, J. & Sugihara, S. (1989b) Stratigraphy and pollen analysis of the boring

core at Lake Ippekiko (Numa-Ike), Ito City, Shizuoka Prefecture, Japan. Daiyonki-Kenkyu (Quaternary

Research) 28, 27-34. (in Japanese)

Kiyonaga, J. (1990) Pollen analysis of Holocene sediments from lowland along the Kashio River, southwestern

part of Yokohama, Japan. Daiyonki-Kenkyu (Quaternary Research) 29, 351-360. (in Japanese with English

summary)


Maeda, Y., Nakai, N., Matsumoto, E., Nakamura, T., Kusunoki, S., Matsushima, Y., Sato, H., Matsubara, A.,

Kumano, S., Kuromi, M., Nukada, M., Aoki, T., Furuta, N., Kobayashi, T., Matsui, J., Kawahara, N. &

Yamashita, H. (1989) Holocene paleoenvironmental changes in the Kei lowland, eastern part of the San-in

coast. The Journal of Ritsumeikan Geographical Society 1, 1-19. (in Japanese)

Matsuda, I. (1983) Pollen analytical study in Shari region I. Totsuru Mire. Scientific Report of Shiretoko

Museum 5, 77-93. (in Japanese)

Matsuoka, K., Nishida, S., Kanehara, M. & Takemura, K. (1983) Pollen analysis of Holocene sediments

obtained from the Muro mountain, central part of Kii Peninsula, Japan. Daiyonki-Kenkyu (Quaternary

Research) 22, 1-10. (in Japanese with English summary)

Matsushita, M. (1991) Holocene vegetation history of the Takagami lowland on Choshi Peninsula, Central

Japan. Japanese Journal of Ecology 41, 19-24. (in Japanese with English summary)

Matsushita, M. (1992) Fossil pollen and palaeoenvironment of Tarumi-Hyuga archaeological site. Report of 1st,

3rd and 4th Scientific Research of Tarumi-Hyuga Archaeological Site, pp. 155-198. The Board of

Education, Kobe City. (in Japanese)

Matsushita, M., Maeda, Y., Matsumoto, E. & Matsushima, Y. (1988) Vegetation history during Holocene at

Shingu on Kii Peninsula and Muroto Point, south-eastern Japan, with special reference to development of

Castanopsis forest. Japanese Journal of Ecology 38, 1-8. (in Japanese with English summary)

Miyagi, T., Hibino, K. & Kawamura, T. (1979) Processes of hillslope denudation and environmental changes

during the Holocene around Sendai, Northeast Japan. Daiyonki-Kenkyu (Quaternary Research) 18, 143-

154. (in Japanese with English summary)

Miyagi, T., Hibino, K., Kawamura, T. & Nakagami, K. (1981) Hillslope development under changing

environment since 20,000 years B.P. in northeast Japan. The Science Report of Tohoku University, 7th

Series (Geography) 31, 1-14.

Miyoshi, N. (1983) Vegetation history in the Chugoku region in the late Pleistocene based on palynological

studies. Vegetation of Japan: Chugoku (ed. by A. Miyawaki), pp. 82-89. Shibundo, Tokyo. (in Japanese)

Miyoshi, N. (1989) Vegetational history of the Hosoike Moor in the Chugoku Mountains, western Japan during

the late Pleistocene and Holocene. Japanese Journal of Palynology 35, 27-42.

Miyoshi, N. & Hada, Y. (1977) Pollen analysis studies of moor sediments in Chugoku, Japan. IV. Makura moor

(Hiroshima Prefecture). Japanese Journal of Ecology 27, 285-290. (in Japanese with English summary)

Miyoshi, N. & Yano, N. (1986) Late Pleistocene and Holocene vegetational history of the Ohnuma Moor in the

Chugoku Mountains, western Japan. Review of Palaeobotany and Palynology 46, 355-376.

Morita, Y. (1981) The relationship between the distributional patterns of surface pollen and vegetation in

Hakkoda Mountains. Japanese Journal of Ecology 31, 317-328. (in Japanese with English summary)

Morita, Y. (1984a) Preliminary palynological studies of the moors in the uplands in Hokkaido. Ecological

Review 20, 237-240.

Morita, Y. (1984b) The relationship between the modern pollen spectra and vegetation on the subalpine zone in

Northeast Japan. Daiyonki-Kenkyu (Quaternary Research) 23, 197-208. (in Japanese with English

summary)

Morita, Y. (1984c) The vegetational history of the subalpine zone in northeast Japan. I. The Azuma Mountain.

Japanese Journal of Ecology 34, 347-356. (in Japanese with English summary)


Morita, Y. (1985a) Palynological study of the deposits from the Uryu-Numa Moor in Mt. Shokanbetsu,

Hokkaido. Annals of Tohoku Geographer Association 37, 166-172. (in Japanese with English summary)

Morita,Y. (1985b) The vegetational history of the subalpine zone in the Zao Mountains, northeast Japan.

Japanese Journal of Palynology 31, 1-5. (in Japanese with English summary)

Morita, Y. (1985c) Pollen diagrams of some peat moors in the subalpine zone in the Shinshu District, Japan.

Ecological Review 20, 301-307.

Morita, Y. (1985d) The vegetational history of the subalpine zone in northeast Japan. II. The Hachimantai

Mountains. Japanese Journal of Ecology 35, 411-420. (in Japanese with English summary)

Morita, Y. (1987a) The vegetational history of the subalpine zone in northeast Japan. III. The Hakkoda

Mountains. Japanese Journal of Ecology 37, 107-117. (in Japanese with English summary)

Morita, Y. (1987b) Palynological study of Tomizawa archaeological site. Report of the 15th Excavation of

Tomizawa Archaeological Site, pp. 439-460. (in Japanese)

Morita, Y. (1990) Pollen analysis of Kurikigahara bog. Report of Scientific Research of Kurikigahara bog, pp.

23-43. Department of Nature Conservation, Iwate Prefecture. (in Japanese)

Morita, Y. & Aizawa, S. (1986) Pollen-analytical study on the vegetational history of the subalpine zone in the

northern Tohoku District. Annals of Tohoku Geographer Association 38, 24-31. (in Japanese with English

summary)

Nakabori, K. (1981) Pollen record from Mizorogaike Pond. Report of Scientific Research of Mizorogaike Pond,

pp. 163-180. Department of Culture and Tourism, Kyoto City. (in Japanese)

Nakamura, J. (1968) Palynological aspects of the Quaternary in Hokkaido. V. Pollen succession and climatic

change since the upper Pleistocene. Research Report of Kochi University 17, 39-51. (in Japanese with

English summary)

Nakamura, J. (1971) Palynological evidence for recent destruction of natural vegetation IV. Swamps of Noda

and Itakuranuma. Annual Report, JIBP-CT(P) of the Fiscal Year 1971, pp. 90-95.

Sakaguchi, Y. (1987) Japanese prehistoric culture flourished in forest-grassland mixed areas. Bulletin of the

Department of Geography, University of Tokyo 19, 1-19.

Suzuki, M., Yoshikawa, M., Endo, K. & Takano, T. (1993) Environmental changes during the last 32,000 years

in the Sakuragawa Lowland, Ibaraki Prefecture. Daiyonki-Kenkyu (Quaternary Research) 32, 195-208. (in

Japanese with English summary)

Tahara, Y. (1980) Palynological study on the sediments of Higashiterayama, Chiba. Bulletin of the Biological

Society of Chiba 29, 27-33. (in Japanese)

Tahara, Y. (1982) Palynological study on the Uenodai Archaeological site, Chiba Prefecture. The 4th Report of

the Uenodai Archaeological site, Chiba, pp. 137-146. The Board of Education, Chiba City. (in Japanese)

Tahara, Y. (1984) Pollen analysis at the Yatsu archaeological site, Chiba Prefecture. Yatsu Archaeological Site,

pp. 560-572. The Board of Education, Chiba City. (in Japanese)

Tahara, Y. & Nakamura, J. (1997) Palynological study on an origin of rice crop in Chiba Prefecture. Annual

Report of the Origin and Spread of Rice Crop. Report of the Grant-in-Aid for Scientific Research (the

fiscal year 1996). (in Japanese)

Takahara, H. (1985) Forest changes around Nose, Osaka Prefecture. Abstract of Annual Meeting, Kansai.

Japanese Forestry Society 36, 141-144. (in Japanese)


Takahara, H. (1991) Palynological Study on the Forest History since the Last Glacial Period in the Kinki and

the eastern Chugoku regions, western Japan. Ph.D. thesis, Kyoto Prefectural University, Kyoto. (in

Japanese)

Takahara, H. (1993) Vegetation history since the last glacial period around the Yamakado Moor, Shiga

Prefecture, western Japan. Japanese Journal of Palynology 39, 1-10. (in Japanese with English summary)

Takahara, H. (1997) Palynological study on disturbance history and its impact on old growth forests. Report of

the Grant-in-Aid for General Scientific Research (C)(2)(06660196). (in Japanese)

Takahara, H. & Takeoka, M. (1980) Studies on the distribution of the natural stand of Sugi (Cryptomeria

japonica D. Don) in the districts along the Sea of Japan (III) Changes of the forests in the neighbourhood

of the Sugawara moor, Tottori Prefecture. Transactions of the 91st Meeting of the Japanese Forestry

Society, pp. 293-294. (in Japanese)

Takahara, H. & Takeoka, M. (1986) Vegetational changes since the last glacial maximum around the

Hatchodaira Moor, Kyoto, Japan. Japanese Journal of Ecology 36, 105-116. (in Japanese with English

summary)

Takahara, H. & Takeoka, M. (1987) Changes of the forest around Nonbara on the Tango peninsula, Kyoto

Prefecture, Japan - On the natural forest of Sugi (Cryptomeria japonica). Journal of Japanese Forestry

Society 69, 215-220. (in Japanese with English summary)

Takahara, H. & Takeoka, M. (1992a) Postglacial vegetation history around Torishima, Fukui Prefecture, Japan.

Ecological Research 7, 79-85.

Takahara, H. & Takeoka, M. (1992b) Vegetation history since the last glacial period in the Mikata lowland, the

Sea of Japan area, Western Japan. Ecological Research 7, 371-386.

Takahara, H., Yamaguchi, H. & Takeoka, M. (1989) Forest changes since the late glacial period in the Hira

mountains of the Kinki region, Japan. Jounal of Japanese Forestry Society 71, 223-231.

Takahara, H., Nishida, S. & Takemura, K. (1995) Vegetation history since the late-glacial period around the

Ikenokochi Moor, Fukui Prefecture, Japan. Japanese Journal of Palynology 41, 99-108. (in Japanese with

English summary)

Takahara, H., Fujiki, T., Miyoshi, N. & Nishida, S. (1997) Vegetation history since the middle Holocene around

Orogatawa Moor, Okayama Prefecture, Japan. Japanese Journal of Palynology 43, 97-106. (in Japanese


Takahashi, N. & Igarashi, Y. (1986) Origin and vegetational succession of upland bogs in the Daisetsuzan

Mountains, central Hokkaido (II). Daiyonki-Kenkyu (Quaternary Research) 25, 113-128. (in Japanese with

English summary)

Takeoka, M. & Takahara, H. (1983) Changes of the forests around the Ukishima moor at Shinguu, Wakayama

Prefecture, based on pollen analysis. Transactions of the 94th Meeting of the Japanese Forestry Society,

383-385. (in Japanese)

Takeoka, M., Takahara, H. & Tanaka, Y. (1982) Palynological study of the Okameike moor in Soni plateau,

Nara. Science Report of Kyoto Prefectural University, Agriculture 34, 51-57. (in Japanese with English

summary)

Tsuda, M. (1990) Pollen analysis of core from the Ooahara Bog at Nyugasa-yama in Nagano Prefecture.

Daiyonki-Kenkyu (Quaternary Research) 29, 439-446. (in Japanese)


Tsuji, S. (1980) Data of the flora of northern Tohoku region since the Last Glacial Maximum. Abstracts of the

Annual Meeting, Japan Association for Quaternary Research 10, 22-23. (in Japanese)

Tsuji, S. (1981) Late Holocene pollen assemblages in the lowlands of Akita Region, northern Japan. Annals of

Tohoku Geographer Association 33, 81-88. (in Japanese with English summary)

Tsuji, S. & Suzuki, S. (1977) Pollen analysis of the Holocene Higata Formation in the North of the Kujukuri

Coastal Plain, Chiba Prefecture, Japan. Daiyonki-Kenkyu (Quaternary Research) 16, 1-12. (in Japanese


Tsuji, S., Minaki, M. & Kosugi, M. (1986) The history of Morinji-numa and its surroundings in central Japan.

The Educational Board of Tatebayashi City, Gunma Prefecture. (in Japanese)

Uchiyama, T. (1987) Palynological studies of some alluvial sediments in a temperate ecotone in Japan (I)

southeastern area of Tohoku District. Japanese Journal of Palynology 33, 111-117. (in Japanese with

English summary)

Uchiyama, T. (1990) Palynological studies of alluvial sediments in the mid-temperate zone of Japan. (II)

northeastern area of the Tohoku district. Japanese Journal of Palynology 36, 17-32. (in Japanese with

English summary)

Uchiyama, T. (1994a) Palynological studies in the deforestation of England and Japan. Bulletin of Elementary

Education, Chiba Keizai College 17, 3-35. (in Japanese with English summary)

Uchiyama, T. (1994b) Vegetation history in the Osado region. The vegetation history in the Osado mountains.

Report of the Grant-in-Aid for General Scientific Research ((C) 03660149, the fiscal year of 1993) (ed. by

M. Tanaka), pp. 38-45. (in Japanese)

Yamaguchi, H., Takahara, H. & Takeoka, M. (1987) Changes of the forests in the Hira Mountains, Shiga

Prefecture (1) pollen analysis of the Sugiyaike Moor. Bulletin of Kyoto Prefectural University, Forestry 31,

6-15. (in Japanese with English summary)

Yamanaka, M. (1969) Palynological studies of peat moors in Mt. Chokai. Ecological Review 17, 203-208.

Yamanaka, M. (1978) Vegetational history since the late Pleistocene in northeast Japan. I. Comparative studies

of the pollen diagrams in the Hakkoda Mountains. Ecological Review 19, 1-36.

Yamanaka, M. (1984) Palynological study of the Quaternary deposits in Kochi City. Research Report of Kochi

University 32, 151-160. (in Japanese with English summary)

Yamanaka, M. & Hamachiyo, M. (1981) Palynological studies of the Holocene deposits from the Kannarashi-ike

bog in the Shikoku Mountains. Memoirs of Faculty of Science, Kochi University (Series D) 2, 11-18.

Yamanaka, M., Saito, K. & Ishizuka, K. (1973) Historical and ecological studies of Abies mariesii on Mt.

Gassan, the Dewa mountains, northeast Japan. Japanese Journal of Ecology 23, 171-185.

Yamanaka, M., Ishizuka, S. & Sugawara, K. (1990) Palynological studies of Quaternary sediments in northeast

Japan. VII. Shiriya-zaki moor in Shimokita Peninsula. Ecological Review 22, 33-47.

Yamanaka, T. & Yamanaka, M. (1977) The vegetation and the Holocene deposits of the Karaike Bog, Kochi

Prefecture. Research Report of Kochi University 26, 17-30. (in Japanese with English summary)

Yamanoi, T. (1986) Establishment and history of natural environment of the Yamagata Basin. Environmental

studies of basins in the Tokou region. Report of the Special Research Fund of Yamagata University (the

fiscal years of 1983, 1984 and 1985), pp. 47-86. (in Japanese)

Yasuda, Y. (1982) Pollen analytical study of the sediment from the Lake Mikata in Fukui prefecture, central


Japan. Daiyonki-Kenkyu (Quaternary Research) 21, 255-271. (in Japanese with English summary)

Yoshii, R. (1988) Palynological study of the bog deposits from the Murodo-daira, Mt. Tateyama, central Japan.

Japanese Journal of Palynology 34, 43-53. (in Japanese with English summary)

Yoshii, R. & Fujii, S. (1981) Palynogical study of the bog deposits from the Midagahara Plateau, Mt. Tateyama

in Toyama Prefecture, central Japan, Journal of Phytogeography and Taxonomy 29, 40-50. (in Japanese



TABLE AND FIGURE CAPTIONS

Table 1. Characteristics of the surface pollen sample sites. Site names with asterisks (*) indicate digitized data.

Biome codes are given in Table 4.

Table 2. Characteristics of the 6000 and 18,000 14C yr B.P. pollen sites. Site names with asterisks (*) indicate

digitized data. Dating control (DC) codes are based on the COHMAP dating control scheme (Webb, 1985; Yu &

Harrison, 1995). For site with continuous sedimentation (indicated by a C after the numeric code), the dating

control is based on bracketing dates where 1 indicates that both dates are within 2000 years of the selected

interval, 2 indicates one date within 2000 years and the other within 4000 years, 3 indicates both within 4000

years, 4 indicates one date within 4000 years and the other within 6000 years, 5 indicates both dates within 6000

years, 6 indicates one date within 6000 years and the other within 8000 years, and 7 indicates bracketing dates

more than 8000 years from the selected interval. For sites with discontinuous sedimentation (indicated by D after

the numeric code), 1 indicates a date within 250 years of the selected interval, 2 a date within 500 years, 3 a date

within 750 years, 4 a date within 1000 years, 5 a date within 1500 years, 6 a date within 2000 years, and 7 a date

more than 2000 years from the selected interval. DC estimates marked with a double asterisk (**) are based on

the K-Ah tephra, which is radiocarbon dated to 6300 14C yr B.P.

Table 3. Assignments of pollen taxa from Japan to the plant functional types used in the biomization procedure.

Table 4. Assignment of plant functional types to biomes from Japan.

Table 5. Correspondence between the vegetation zones (Yoshioka, 1973) and biomes of Japan.

Table 6. Observed vs. predicted biomes using the modern pollen data set.

Figure 1. Modern vegetation map of Japan, modified from Yoshioka (1973).

Figure 2. Modern biomes (a) observed at individual sites, and (b) predicted from modern surface pollen samples

(including duplicates from the same site), compared with fossil-pollen based biome distributions at (c) 6000 14C

yr B.P. and (d) 18,000 14C yr B.P.

Figure 3. Biomes along elevational gradient from south to north in Japan, (a) observed at individual modern

sites, (b) predicted from modern surface pollen samples, compared with fossil-pollen based biome distributions

at (c) 6000 14C yr B.P. and (d) 18,000 14C yr B.P. The lines (1), (2) and (3) in Figures 3a and 3b represent the

timber line, the boundary between the cool temperate deciduous forest and the subalpine cool mixed/cool conifer

forest, and the boundary between broadleaved evergreen/warm mixed forest and cool temperate deciduous

forest, respectively (modified from Numata, 1971; Kikuchi, 1985).


Table 1 Characteristics of the surface pollen sample sites. Site names with asterisks (*) indicate digitized data. Biome codes are given in Table 4.

Site Name Lat.(oN)

Long.(°E)

Elev.(m)

Sample type modernbiome

Modern vegetation type References

Ushinohira* 32.40 128.40 50 peat core top WAMX secondary warm temperate evergreen forest Hatanaka, 1985

Odanoike* 33.10 131.20 770 peat core top WAMX grassland Hatanaka, 1982

Karaike* 33.60 133.10 1220 peat core top TEDE Chamaecyparis obtusa plantations Yamanaka & Yamanaka,1977

Ukishima-no-mori* 33.70 136.00 5 peat core top WAMX warm temperate evergreen forest Matsushita et al., 1988

Ukishimanomori 33.70 136.00 7 peat core top WAMX evergreen broadleaved forest Takeoka & Takahara, 1983

Kannarashiike* 33.80 133.20 1600 peat core top TEDE cool temperate mixed forest of Fagus & Abies Yamanaka & Hamachiyo, 1981

Shoe* 33.80 131.00 5 peat core top WAMX archaeological site Hatanaka, 1980

Odai Nanatuike 34.20 136.10 1490 peat core top TEDE Fagus & Abies forest Takahara, 1997

Odai Masakigahra 34.20 136.10 1630 peat core top COMX subalpine conifer forest Takahara, 1997

Ubuka 34.50 131.60 390 peat core top WAMX Cryptomeria japonica plantations Hatanaka & Miyoshi, 1980

Ikenohira* 34.50 136.20 600 peat core top WAMX secondary forest of Pinus & deciduous Quercus Matsuoka et al., 1983

Makura 34.70 132.40 720 peat core top TEDE cool temperate deciduous forest Miyoshi & Hada, 1977

Oike 34.70 137.60 5 peat core top WAMX secondary Pinus forest Morita, unpub.

Oochiba 34.80 137.50 4 peat core top WAMX secondary Pinus forest Morita, unpub.

Nose 35.00 135.40 190 peat core top WAMX secondary Pinus forest Takahara, 1985

Naganoyama 35.00 137.50 550 peat core top WAMX secondary Pinus forest Morita, unpub.

Mizorogaike * 35.10 135.80 75 peat core top WAMX secondary Pinus & deciduous Quercus forest Nakabori, 1981

Hatchodaira C 35.20 135.80 810 peat core top TEDE cool temperate deciduous forest Takahara & Takeoka, 1986

Sugiyaike 35.20 135.90 950 peat core top TEDE cool temperate deciduous forest Yamaguchi et al., 1987

Sonemuma * 35.20 136.20 86 peat core top WAMX paddy field Ishida et al., 1984

Yakumogahara 35.30 135.90 910 peat core top TECO Fagus forest with Cryptomeria japonica Takahara et al., 1989

Chojidani 35.30 135.80 637 peat core top TECO Fagus forest with Cryptomeria japonica Takahara, 1997

Hosoike 35.40 134.10 970 peat core top TEDE Fagus forest & Cryptomeria japonica plantations Miyoshi, 1989

Koseinuma 35.40 134.50 1470 peat core top TECO Cryptomeria japonica forest Takahara et al., unpub.

Kanai site* 35.40 139.50 9 peat core top WAMX paddy fields Kiyonaga, 1990

Ohnuma 35.40 134.60 610 peat core top TEDE cool temperate deciduous forest Miyoshi & Yano, 1986

Sugawara 35.40 134.00 640 peat core top WAMX secondary Pinus & deciduous forest Takahara & Takeoka, 1980

Sugano* 35.50 134.40 340 peat core top WAMX secondary Pinus forest & Cryptomeria plantation Miyoshi, 1983

Iwaya 35.50 135.90 20 peat core top WAMX secondary forest of Pinus densiflora Takahara & Takeoka, 1992b

Hanawa* 35.60 140.20 11.8 peat core top WAMX paddy field Tahara, 1984

Yamakado Moor 35.60 136.10 300 peat core top WAMX secondary forest of Pinus & Quercus Takahara, 1993

Higashiterayama* 35.60 140.10 5 peat core top WAMX secondary forest, paddy field Tahara, 1980

Biome reconstructions for Japan 25Yutorinuma 35.60 140.60 620 peat core top TEDE Fagus forest Hibino & Morita, unpub.

Ikenokochi 35.70 136.10 300 peat core top WAMX secondary forest of Pinus, Castanea & Quercus Takahara et al., 1995

Uenodai archaeological site* 35.70 140.10 5 peat core top WAMX paddy field Tahara, 1982

Ofuke 35.70 135.20 550 peat core top TEDE secondary deciduous forest Takahara, 1991

Kamezaki 35.70 140.20 4 peat core top WAMX secondary deciduous forest Uchiyama, 1994a

Kujukuri plain (YK-6)* 35.70 140.60 4.5 peat core top WAMX secondary deciduous forest Tsuji & Suzuki, 1977

Kyoto Nonbara 35.70 135.10 160 peat core top WAMX secondary deciduous forest Takahara & Takeoka, 1987

Horinouchi* 35.80 139.90 22 peat core top WAMX paddy fields Tahara & Nakamura, 1997

Tanohara 35.90 137.50 2000 peat core top COCO subalpine coniferous forest Morita, 1985c

Ippekiko* 35.90 139.10 187 peat core top WAMX secondary forest of Pinus & deciduous Quercus Kanauchi et al., 1989b

Noda swamp* 36.00 139.90 5 peat core top WAMX no natural vegetation Nakamura, 1971

Noda * 36.00 139.90 5 peat core top WAMX no natural vegetation Sakaguchi, 1987

Shirakoma moor 36.10 138.40 2130 peat core top COCO subalpine coniferous forest Morita, 1985c

Yashimagahara* 36.10 138.20 1630 peat core top TEDE & COMX upland meadow Kanauchi et al., 1989a

Morinjinuma* 36.20 139.50 8 peat core top WAMX grassland Tsuji et al., 1986

Itakuranuma* 36.20 139.60 10 peat core top WAMX n/a Nakamura, 1971

Kojonuma* 36.20 139.60 8 peat core top WAMX paddy fields Tsuji et al., 1986

Krakemi* 36.50 137.90 950 peat core top TEDE deciduous Quercus forest Hibino & Sasaki, 1982

Lake Kizaki* 36.60 137.80 764 lake mud TEDE deciduous Quercus forest Hibino & Horie, 1991

Midagahara site L* 36.60 137.60 1890 peat core top COCO subalpine conifer forest Yoshii & Fujii, 1981

Hijikura* 36.80 137.90 980 peat core top TEDE deciduous Quercus forest Hibino & Sasaki, 1982

Tarosan 36.80 139.50 2300 peat core top COCO subalpine coniferous forest Morita, unpub.

Oze-nakatashiro 36.90 139.20 1400 peat core top TEDE upper limit of Fagus forest Morita, unpub.

Oze-shimotashiro 36.90 139.30 1400 peat core top TEDE upper limit of Fagus forest Morita, unpub.

Miyatokooyachi moor* 37.30 139.50 850 peat core top TEDE cool temperate deciduous forest (Fagus forest) Choi & Hibino, 1985

Akaiyachi 37.50 140.00 520 peat core top TEDE secondary forest of Quercus & Pinus Morita, unpub.

Hoshojiri* 37.60 140.10 530 peat core top TEDE & WAMX secondary forest of Quercus & Pinus Miyagi et al., 1981

Haranomachi 37.60 141.00 10 peat core top TEDE secondary deciduous forest Uchiyama, 1987

Yachidaira 37.70 140.20 1500 peat core top COMX lower limit of subalpine coniferous forest Morita, 1984c

Babayachi 37.80 140.10 1450 peat core top TEDE upper limit of Fagus forest Morita, 1984c

Tojuro 37.80 140.20 1800 peat core top COCO subalpine coniferous forest Morita, 1984c

Tozugawa 38.10 138.40 12 peat core top TEDE secondary deciduous forest Uchiyama, 1994b

Zao no.4* 38.10 140.50 1690 peat core top COCO upper limit of subalpine conifer forest Morita, 1985b

Nenoshiroishi* 38.40 140.80 270 peat core top TEDE secondary forest of deciduous Quercus & Castanea Miyagi et al., 1979

Ishinomaki 38.50 141.40 2 peat core top TEDE secondary deciduous forest Uchiyama, 1990

Nembutsugahara* 38.50 140.10 1100 peat core top TEDE cool temperate deciduous forest (Fagus forest) Yamanaka, 1973

Biome reconstructions for Japan 26Iinogawa 38.50 141.30 7 peat core top TEDE secondary deciduous forest Uchiyama, 1990

Ryugahara moor* 39.10 140.10 1180 peat core top TEDE & COMX upper limit of cool temperate deciduous forest (Fagus forest) Yamanaka, 1969

Kuzunori* 39.40 140.10 12 peat core top TEDE paddy field Tsuji, 1981

Kurikigahara 39.90 140.90 1130 peat core top COMX lower limit of subalpine coniferous forest Morita, 1990

Hachiman-Numa 40.00 140.90 1580 peat core top COCO subalpine coniferous forest Morita, 1984b, 1985d

Onuma 40.00 140.80 950 peat core top TEDE Fagus forest Morita, 1984b, 1985d

Shibayachi* 40.30 140.60 90 peat core top TEDE secondary forest of Quercus, Pinus & Cryptomeria japonica Hibino, 1991

Shirochiyama 40.50 140.80 1000 peat core top TEDE upper limit of Fagus forest Morita & Aizawa, 1986

Zenkojitai 40.50 140.90 950 peat core top TEDE Fagus forest Morita, unpub.

Oseyachi 40.60 140.90 1250 peat core top COCO subalpine coniferous forest Morita, 1987a

Komagaminekita 40.60 140.90 1250 peat core top COCO subalpine coniferous forest Morita, 1987a

Komaganenishi 40.60 140.90 1300 peat core top COCO subalpine coniferous forest Morita, 1987a

Sarukura 40.60 140.90 1300 peat core top COCO subalpine coniferous forest Morita, 1987a

Yabitsuyachi 43 40.60 140.90 1080 peat core top COMX subalpine coniferous forest Morita, 1981

Takadayachi 44 40.60 140.90 1050 peat core top COMX subalpine coniferous forest Morita, 1987a

Takadayachi 43 40.60 140.90 1050 peat core top COMX subalpine coniferous forest Morita, 1987a

Shimokenashi 40.70 140.90 1050 peat core top COMX lower limit of subalpine coniferous forest Morita, 1987a

Ogawara 40.70 141.30 7 peat core top TEDE paddy field Morita, unpub.

Tashiro moor* 40.90 140.90 550 peat core top TEDE secondary cool temperate deciduous forest forest Yamanaka, 1978

Shiriyazaki* 41.40 141.50 10 peat core top TEDE secondary Quercus & Thujopsis forest Yamanaka et al., 1990

Orochigahara 42.90 141.10 980 peat core top COCO subalpine coniferous forest Morita, 1984a

Ponchubetsudake* 43.60 142.90 1735 peat core top TUND Pinus pumila shrubs Takahashi & Igarashi, 1986

Uryu-Numa 43.70 141.60 850 peat core top COCO Betula ermanii forest Morita, 1985a

Chippubetsu bog* 43.80 142.00 40 peat core top COMX secondary Quercus forest Nakamura, 1968

Ukishima 44.00 142.90 870 peat core top COCO subalpine coniferous forest Morita, 1984a

Kenbuchi Basin* 44.10 142.40 135 peat core top COMX cool temperate deciduous forest Igarashi et al., 1993


Table 2 Characteristics of the 6000 and 18,000 14C yr B.P. pollen sites. Site names with asterisks (*) indicate digitized

data. Dating control (DC) codes are based on the COHMAP dating control scheme (Webb, 1985; Yu & Harrison, 1995).

For site with continuous sedimentation (indicated by a C after the numeric code), the dating control is based on bracketing

dates where 1 indicates that both dates are within 2000 years of the selected interval, 2 indicates one date within 2000

years and the other within 4000 years, 3 indicates both within 4000 years, 4 indicates one date within 4000 years and the

other within 6000 years, 5 indicates both dates within 6000 years, 6 indicates one date within 6000 years and the other

within 8000 years, and 7 indicates bracketing dates more than 8000 years from the selected interval. For sites with

discontinuous sedimentation (indicated by D after the numeric code), 1 indicates a date within 250 years of the selected

interval, 2 a date within 500 years, 3 a date within 750 years, 4 a date within 1000 years, 5 a date within 1500 years, 6 a

date within 2000 years, and 7 a date more than 2000 years from the selected interval. DC estimates marked with a double

asterisk (**) are based on the K-Ah tephra, which is radiocarbon dated to 6300 14C yr B.P.

Site Lat.(oN)

Long.(oE)

Elev.(m)

Sedimenttype

Recordlength (ka)

No. of14C dates

No. oftephra

DC at6000

14C yr B.P.

DC at18000 14C

yr B.P.

References

Nakashima * 32.80 130.70 5 silt ?- >18 1 1 2D** n/a Iwauchi & Hase, 1992

Maruike * 33.60 133.60 0 silt 0->10 0 1 2D** n/a Yamanaka, 1984

Ukishima-no-mori * 33.70 136.00 5 clay 0->6.3 1 1 2D** n/a Matsushita et al., 1988

Okameike 34.50 136.20 700 clay peat 0->10 1 1 2D** n/a Takeoka et al., 1982

Ubuka 34.50 131.60 390 silt 0->16 2 0 2D 6D Hatanaka & Miyoshi, 1980

Ikenohira * 34.50 136.20 600 peaty clay 0->11 3 2 2D** n/a Matsuoka et al., 1983

Tarumi-Hyuga site * 34.60 135.10 1 silt 6.3-? 0 1 2D** n/a Matsushita, 1992

Makura 34.70 132.40 720 silty clay 0->7.9 3 0 2C n/a Miyoshi & Hada, 1977

Tenpozan * 34.70 135.40 0 silt ?- >6.3 0 1 2D** n/a Furutani, 1979

Mizorogaike * 35.10 135.80 75 peat 0-14 0 6 2D** n/a Nakabori, 1981

Hatchodaira B 35.20 135.80 810 peat 0->25 1 2 3D n/a Takahara & Takeoka, 1986

Sonemuma * 35.20 136.20 86 peat 0-12 4 2 2D** n/a Ishida et al., 1984

Orogatawa 35.30 133.70 680 peaty clay 0-6.5 0 1 2D** n/a Takahara et al., 1997

Yakumogahara B 35.30 135.90 910 peat 0-14 2 1 2D** n/a Takahara et al., 1989

Hosoike 35.40 134.10 970 muck 0->34 2 3 2D** 2C Miyoshi, 1989

Ohnuma 35.40 134.60 610 peat & clay 0-19 6 0 2C 1C Miyoshi & Yano, 1986

Sugawara 35.40 134.00 640 peat 0->6.3 1 1 2D** n/a Takahara & Takeoka, 1980

Iwaya 35.50 135.90 20 peat 0->30 4 3 2D** 1D Takahara & Takeoka, 1992b

Torihama 35.60 135.90 2 peaty clay 0-20 0 3 2D** n/a Takahara & Takeoka, 1992a

Lake Mikata * 35.60 135.90 0 peat 0->42 8 1 n/a 2C Yasuda, 1982

Yamakado 35.60 136.10 300 peat 0->25 1 2 2D** n/a Takahara, 1993

Kei * 35.60 134.80 2 silty clay 0-7.5 10 1 2D** n/a Maeda et al., 1989

Ikenokochi 35.70 136.10 300 peat 0-12 2 2 2D** n/a Takahara et al., 1995

Ofuke 35.70 135.20 550 peat 0->30 3 6 2D** 4C Takahara, 1991

Choshi Takagami * 35.70 140.90 10 clay ?-8.4 9 0 1C n/a Matsushita, 1991

Nazukari * 35.80 139.90 5 silt 1.5->6 9 8 1D n/a Endo et al., 1989

Ooahara * 35.90 138.20 1800 peat 0-25 2 2 n/a 4C Tsuda, 1990

Yashimagahara * 36.10 138.20 1630 peat 0-12 4 5 2D** n/a Kanauchi et al., 1988, 1989a

Shimo-Oshima * 36.10 140.10 30 peat 16->25 5 6 n/a 1C Suzuki et al., 1993

Karakemi * 36.50 137.90 950 peat 0->8 1 2 6D n/a Hibino & Sasaki, 1982

Midagahara site L * 36.60 137.60 1890 peat 0->6.3 0 1 2D** n/a Yoshii & Fujii, 1981

Murododaira site L * 36.60 137.60 2440 peat 0->6.3 3 3 2D** n/a Yoshii, 1988

Lake Kizaki * 36.60 137.80 764 lake seds 0-25 3 3 1C 6C Hibino & Horie, 1991


Oze-nakatashiro 36.90 139.20 1400 peat 0->7.3 1 2 3C n/a Morita, unpub.

Oze-shimotashiro 36.90 139.30 1400 peat 0->8.5 1 3 3C n/a Morita, unpub.

Miyatokooyachi moor * 37.20 139.50 850 peat 0->7 1 0 3D n/a Choi & Hibino, 1985

Akaiyachi 37.50 140.00 520 peat 0->6.5 2 0 4C n/a Morita, unpub.

Hoshojiri * 37.60 140.10 530 peat 0->25 6 1 2C 4C Miyagi et al., 1981

Yachidaira 37.70 140.20 1500 peat 0->6.3 0 2 2D** n/a Morita, 1984c

Babayachi 37.80 140.10 1450 peaty clay 0->6 0 2 2D** n/a Morita, 1984c

Tojuro 37.80 140.20 1800 peaty clay 0->6.4 1 2 1D n/a Morita, 1984c

Kawadoi * 38.10 140.30 300 peat 0-120 6 14 n/a 3C Hibino et al., 1991

Tomizawa 38.20 140.90 10 silty clay 0-6.0 4 1 1D n/a Morita, 1987b

Nenoshiroishi * 38.40 140.80 270 peat 0-7 7 0 1C n/a Miyagi et al., 1979

Ukinuma * 38.50 140.40 86 peat 10->36 4 1 n/a 6C Yamanoi, 1986

Kuzunori * 39.40 140.10 12 silt 0->5.5 1 0 4D n/a Tsuji, 1981

Onuma 40.00 140.80 950 peat 0->2.5 0 2 4C n/a Morita, 1985d

Shibayachi * 40.30 140.60 90 organic clay 0->9.5 1 0 7D n/a Hibino, 1991

Takadayachi 44 40.60 140.90 1050 peat 0->4.5 0 3 1C n/a Morita, 1987a

Oseyachi 40.60 140.90 1250 peat 0->4.5 0 3 1C n/a Morita, 1987a

Komagaminekita 40.60 140.90 1250 peat 0->4.5 0 3 1C n/a Morita, 1987a

Sarukura 40.60 140.90 1300 peat 0->4.5 0 3 1C n/a Morita, 1987a

Komaganenishi 40.60 140.90 1300 peat 0->4.5 0 4 1C n/a Morita, 1987a

Yabitsuyachi 40.60 140.90 1080 peat 0->6.9 5 4 1C n/a Morita & Aizawa, 1986

Takadayachi43 40.60 140.90 1050 peat 0->4.5 0 3 1C n/a Morita, 1987a

Shimokenashi 40.70 140.90 1050 peat 0->4.5 0 3 1C n/a Morita, 1987a

Dekijima * 40.90 140.30 0 peat ?- >25 2 3 n/a 6C Tsuji, 1980

Tashiro moor * 40.90 140.90 550 peat 0-12 1 7 1D n/a Yamanaka, 1978

Furano Basin * 43.40 142.40 173 peat 0-32 5 1 6D 7C Igarashi et al., 1993

Ponchubetsudake * 43.60 142.90 1735 peat 0-7.5 2 3 2C n/a Takahashi & Igarashi, 1986

Uryu-Numa 43.70 141.60 850 peat 0-9.5 1 1 4C n/a Morita, 1985a

Chippubetsu bog * 43.80 142.00 40 peat/clay 0->24 1 0 7D n/a Nakamura, 1968

Totsuru mire * 43.90 144.60 5 silty clay 0->5.6 2 0 3D n/a Matsuda, 1983

Kenbuchi * 44.10 142.40 135 clay 0-32 5 0 4D 1C Igarashi et al., 1993


Table 3 Assignments of pollen taxa from Japan to the plant functional types used in the biomization procedure.

Abbr. Plant functional type Pollen taxaaa arctic/alpine dwarf shrub Betula, Pinus subgenus Hyploxylon, Salixaf arctic/alpine forb Artemisia, Caryophyllaceae, Compositae, Cruciferae, Gentiana, Geranium,

Leguminosae, Liliaceae, Polygonaceae, Polygonum undiff., Polygonumbistorta type, Ranunculaceae, Rosaceae, Stellaria, Thalictrum, Umbelliferae

ax arctic/alpine fern or fern ally Selaginella selaginoidesbec boreal evergreen conifer Abies, Picea, Pinus subgenus Hyploxylonbf boreal forb Aconitum, Alium, Artemisia, Caryophyllaceae, Compositae, Cruciferae,

Epilodium, Gentiana, Geranium, Leguminosae, Liliaceae, Polygonaceae,Polygonum undiff., Polygonum bistorta type, Ranunculaceae, Rosaceae,Rumex, Sanguisorba, Scabiosa, Scheuchzeria, Stellaria, Thalictrum,Umbelliferae

bs boreal summergreen Alnaster, Alnus, Betula, Salixbsc boreal summergreen conifer Larixctc1 upper cool-temperate conifer Abies, Pinus subgenus Hyploxylon, Tsugactc2 intermediate-temperate conifer Abies, Cryptomeria, Pinus subgenus Diploxylon, Sciadopitys, Tsugaec eurythermic conifer Cupressaceae, Pinus undiff.g grass Gramineaeh heath Ericaceae, Ericales, Rhododendrons sedge Cyperaceaetf temperate forb Aconitum, Alium, Artemisia, Cardamine, Caryophyllaceae,

Chenopodiaceae, Chenopodiaceae-Amaranthaceae, Cichorioideae,Compositae, Coptis, Cruciferae, Epilodium, Filipendula, Gentiana,Geranium, Humulus, Hygrophila, Impatiens, Labiatae, Leguminosae,Liliaceae, Lyshimachia, Patrinia, Plantago, Polygonaceae, Polygonumundiff., Ranunculaceae, Reynoutria, Rosaceae, Rumex, Sanguisorba,Scabiosa, Stellaria, Thalictrum, Umbelliferae, Urticaceae

ts temperate summergreen Acanthopanax, Acer, Alnus, Betula, Carpinus, Carpinus-Ostrya,Celastraceae, Clematis, Cleyera, Corylus, Cornus, Euonymus, Fraxinus,Hamamelis, Juglans-Pterocarya, Juglandaceae, Leguminosae, Moraceae,Prunus, Quercus subgenus Lepidobalanus, Rhus, Rosaceae, Salix, Sorbus,Symplocos, Ulmus-Zelkova, Ulmus, Viburnum, Vitis, Weigela

ts0 lower cool-temperate summergreen Fagus, Fagus japonicats1 cool-temperate summergreen Cercidiphyllum, Fagus, Fagus crenata, Myrica, Phellodentron, Pterocarya,

Tiliats2 intermediate-temperate summergreen Aesculus, Araliaceae,Carpinus tchonoskii, Castanea, Castanea-

Castanopsis, Ilex, Parthenocissusts3 warm-temperate summergreen Alnaster, Celtis-Aphananthe, Celtis, Diospyros, Elaegnus, Ilex, Ligustrum,

Mallotus, Platycarya, Rhamnus, Zelkovawtc warm-temperate conifer Podocarpuswte warm-temperate broadleaved evergreen Araliaceae, Aucuba, Camellia, Castanopsis, Castanea-Castanopsis,

Celastraceae, Euphorbiaceae, Ilex, Illicium, Ligustrum, Moraceae, Myrica,Quercus subgenus Cyclobalanopsis, Skimmia, Symplocos

wte1 cool-temperate broadleaved evergreen Ilex, Skimmia, Viscum


Table 4 Assignment of plant functional types to biomes from Japan.

Biome Code Plant functional types

tundra TUND aa, af, ax, bf, g, h, s

cold deciduous forest CLDE bf, bs, bsc, ec, h

taiga TAIG bec, bf, bs, bsc, ec, h

cold mixed forest CLMX bf, bs, ctc1, ec, h

cool conifer forest COCO bec, bf, bs, ctc1, ec, h

cool mixed forest COMX bec, bf, bs, ctc1, ctc2, ec, h, tf, ts, ts1

temperate deciduous forest TEDE bf, bs, ctc1, ctc2, ec, h, tf, ts, ts0, ts1, ts2, wte1

broadleaved evergreen/warm mixed forest WAMX ctc2, ec, h, ts, ts, ts0, ts2, ts3, wtc, wte, wte1

temperate conifer forest TECO ctc2, ec, h, tf, ts, ts0, ts2, ts3, wtc, wte1


Table 5 Correspondence between the vegetation zones (Yoshioka, 1973) and biomes of Japan.

Modern vegetation zone Biome Code

alpine vegetation tundra TUND

not present cold deciduous forest CLDE

not present taiga TAIG

not present cold mixed forest CLMX

subalpine conifer forest cool conifer forest COCO

subalpine deciduous broadleaved thicket cool mixed forest COMX

subarctic mixed forest no equivalent

cool temperate broadleaved deciduous forest temperate deciduous forest TEDE

warm temperate broadleaved evergreen forest broadleaved evergreen/warm mixed forest WAMX

warm temperate conifer forest temperate conifer forest TECO


Table 6 Observed vs predicted biomes using the modern pollen data set.

Biomes predicted using modern pollen data set

TUND COCO COMX TEDE WAMX TECO total

TUND 0 0 1 0 0 0 1

Observed COCO 0 0 7 7 0 0 14

Modern COMX 0 0 3 7 0 0 10

Biomes TEDE 0 0 0 27 1 6 34

WAMX 1 0 0 8 21 2 32

TECO 0 0 0 3 0 0 3

total 1 0 11 52 22 8 94

Date post:	14-Mar-2020
Category:	Documents
Upload:	others
View:	2 times
Download:	0 times

Biome reconstructions for Japan 1 - Bristol · Biome reconstructions for Japan 2 (A) ABSTRACT 1 A...

Documents