Biome reconstructions for Japan
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
1
Biome reconstructions for Japan
(A)
2
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
14
C 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
14
C yr B.P. are likely to be
conservative estimates of the actual changes.
3 The biome distribution at 6000
14
C 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
14
C 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 glacial
maximum,
Biome reconstructions for Japan
(A)
3
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
14
C 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
Biome reconstructions for Japan
4
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
14
C 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.,
14
based on a linear age-depth model using the 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
14
C 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.
Biome reconstructions for Japan
(B)
5
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 cooltemperate 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
Biome reconstructions for Japan
6
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 coldtolerant 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
Biome reconstructions for Japan
7
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
Biome reconstructions for Japan
8
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
14
C 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
14
C 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
14
C 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
14
C 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.
Biome reconstructions for Japan
9
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 plateautype 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
Biome reconstructions for Japan
10
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 wellstudied (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
14
C 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.
Biome reconstructions for Japan
11
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/).
Biome reconstructions for Japan
(A)
12
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.
Biome reconstructions for Japan
13
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Morita, Y. (1990) Pollen analysis of Kurikigahara bog. Report of Scientific Research of Kurikigahara bog, pp.
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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)
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Tahara, Y. (1982) Palynological study on the Uenodai Archaeological site, Chiba Prefecture. The 4th Report of
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Biome reconstructions for Japan
20
Takahara, H. (1991) Palynological Study on the Forest History since the Last Glacial Period in the Kinki and
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Prefecture, western Japan. Japanese Journal of Palynology 39, 1-10. (in Japanese with English summary)
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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
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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)
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with English summary)
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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,
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Takeoka, M., Takahara, H. & Tanaka, Y. (1982) Palynological study of the Okameike moor in Soni plateau,
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Biome reconstructions for Japan
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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)
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Tsuji, S. & Suzuki, S. (1977) Pollen analysis of the Holocene Higata Formation in the North of the Kujukuri
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with English summary)
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southeastern area of Tohoku District. Japanese Journal of Palynology 33, 111-117. (in Japanese with
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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)
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Biome reconstructions for Japan
22
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.
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in Toyama Prefecture, central Japan, Journal of Phytogeography and Taxonomy 29, 40-50. (in Japanese
with English summary)
Biome reconstructions for Japan
23
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).
Biome reconstructions for Japan
24
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
Ushinohira*
Lat. Long.
(oN) (°E)
32.40 128.40
Elev.
(m)
50
Odanoike*
33.10 131.20
770
Karaike*
33.60 133.10
Ukishima-no-mori*
Ukishimanomori
Sample type
peat core top
modern
biome
WAMX
secondary warm temperate evergreen forest
Hatanaka, 1985
peat core top
WAMX
grassland
Hatanaka, 1982
1220
peat core top
TEDE
Chamaecyparis obtusa plantations
Yamanaka & Yamanaka,1977
33.70 136.00
5
peat core top
WAMX
warm temperate evergreen forest
Matsushita et al., 1988
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
Shoe*
33.80 131.00
5
peat core top
WAMX
Odai Nanatuike
34.20 136.10
1490
peat core top
Odai Masakigahra
34.20 136.10
1630
peat core top
Ubuka
34.50 131.60
390
peat core top
WAMX
Modern vegetation type
References
cool temperate mixed forest of Fagus & Abies
Yamanaka & Hamachiyo, 1981
archaeological site
Hatanaka, 1980
TEDE
Fagus & Abies forest
Takahara, 1997
COMX
subalpine conifer forest
Takahara, 1997
Cryptomeria japonica plantations
Hatanaka & Miyoshi, 1980
Ikenohira*
34.50 136.20
600
peat core top
WAMX
Makura
34.70 132.40
720
peat core top
TEDE
secondary forest of Pinus & deciduous Quercus
Matsuoka et al., 1983
cool temperate deciduous forest
Miyoshi & Hada, 1977
Oike
34.70 137.60
5
peat core top
Oochiba
34.80 137.50
4
peat core top
WAMX
secondary Pinus forest
Morita, unpub.
WAMX
secondary Pinus forest
Nose
35.00 135.40
190
Morita, unpub.
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
Yutorinuma
35.60 140.60
620
peat core top
TEDE
Ikenokochi
35.70 136.10
300
peat core top
Uenodai archaeological site* 35.70 140.10
5
Ofuke
35.70 135.20
550
Kamezaki
35.70 140.20
Kujukuri plain (YK-6)*
35.70 140.60
Kyoto Nonbara
25
Fagus forest
Hibino & Morita, unpub.
WAMX
secondary forest of Pinus, Castanea & Quercus
Takahara et al., 1995
peat core top
WAMX
paddy field
Tahara, 1982
peat core top
TEDE
secondary deciduous forest
Takahara, 1991
4
peat core top
WAMX
secondary deciduous forest
Uchiyama, 1994a
4.5
peat core top
WAMX
secondary deciduous forest
Tsuji & Suzuki, 1977
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
Iinogawa
38.50 141.30
7
peat core top
TEDE
Ryugahara moor*
39.10 140.10
1180
peat core top
TEDE & COMX
Kuzunori*
39.40 140.10
12
peat core top
Kurikigahara
39.90 140.90
1130
peat core top
Hachiman-Numa
40.00 140.90
1580
peat core top
Onuma
40.00 140.80
950
peat core top
Shibayachi*
40.30 140.60
90
Shirochiyama
40.50 140.80
Zenkojitai
40.50 140.90
26
secondary deciduous forest
Uchiyama, 1990
upper limit of cool temperate deciduous forest (Fagus forest)
Yamanaka, 1969
TEDE
paddy field
Tsuji, 1981
COMX
lower limit of subalpine coniferous forest
Morita, 1990
COCO
subalpine coniferous forest
Morita, 1984b, 1985d
TEDE
Fagus forest
Morita, 1984b, 1985d
peat core top
TEDE
secondary forest of Quercus, Pinus & Cryptomeria japonica
Hibino, 1991
1000
peat core top
TEDE
upper limit of Fagus forest
Morita & Aizawa, 1986
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
Biome reconstructions for Japan
27
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
Nakashima *
Lat.
(oN)
Long.
(oE)
Elev.
(m)
32.80
130.70
5
Sediment
type
Record
length (ka)
silt
?- >18
No. of
C dates
No. of
tephra
DC at
6000
14
C yr B.P.
1
1
2D**
14
DC at
References
18000 14C
yr B.P.
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
Makura
34.70
132.40
720
Tenpozan *
34.70
135.40
0
silt
6.3-?
0
1
2D**
n/a
Matsushita, 1992
silty clay
0->7.9
3
0
2C
n/a
Miyoshi & Hada, 1977
silt
?- >6.3
0
1
2D**
n/a
Furutani, 1979
Nakabori, 1981
Mizorogaike *
35.10
135.80
75
peat
0-14
0
6
2D**
n/a
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
Takahara, 1993
Kei *
35.60
134.80
2
Ikenokochi
35.70
136.10
Ofuke
35.70
Choshi Takagami *
35.70
Nazukari *
peat
0->25
1
2
2D**
n/a
silty clay
0-7.5
10
1
2D**
n/a
Maeda et al., 1989
300
peat
0-12
2
2
2D**
n/a
Takahara et al., 1995
135.20
550
peat
0->30
3
6
2D**
4C
Takahara, 1991
140.90
10
clay
?-8.4
9
0
1C
n/a
Matsushita, 1991
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
Biome reconstructions for Japan
28
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
Biome reconstructions for Japan
29
Table 3 Assignments of pollen taxa from Japan to the plant functional types used in the biomization procedure.
Abbr.
aa
af
Plant functional type
arctic/alpine dwarf shrub
arctic/alpine forb
ax
bec
bf
arctic/alpine fern or fern ally
boreal evergreen conifer
boreal forb
bs
bsc
ctc1
ctc2
ec
g
h
s
tf
boreal summergreen
boreal summergreen conifer
upper cool-temperate conifer
intermediate-temperate conifer
eurythermic conifer
grass
heath
sedge
temperate forb
ts
temperate summergreen
ts0
ts1
lower cool-temperate summergreen
cool-temperate summergreen
ts2
intermediate-temperate summergreen
ts3
warm-temperate summergreen
wtc
wte
warm-temperate conifer
warm-temperate broadleaved evergreen
wte1
cool-temperate broadleaved evergreen
Pollen taxa
Betula, Pinus subgenus Hyploxylon, Salix
Artemisia, Caryophyllaceae, Compositae, Cruciferae, Gentiana, Geranium,
Leguminosae, Liliaceae, Polygonaceae, Polygonum undiff., Polygonum
bistorta type, Ranunculaceae, Rosaceae, Stellaria, Thalictrum, Umbelliferae
Selaginella selaginoides
Abies, Picea, Pinus subgenus Hyploxylon
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
Alnaster, Alnus, Betula, Salix
Larix
Abies, Pinus subgenus Hyploxylon, Tsuga
Abies, Cryptomeria, Pinus subgenus Diploxylon, Sciadopitys, Tsuga
Cupressaceae, Pinus undiff.
Gramineae
Ericaceae, Ericales, Rhododendron
Cyperaceae
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, Polygonum
undiff., Ranunculaceae, Reynoutria, Rosaceae, Rumex, Sanguisorba,
Scabiosa, Stellaria, Thalictrum, Umbelliferae, Urticaceae
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
Fagus, Fagus japonica
Cercidiphyllum, Fagus, Fagus crenata, Myrica, Phellodentron, Pterocarya,
Tilia
Aesculus, Araliaceae,Carpinus tchonoskii, Castanea, CastaneaCastanopsis, Ilex, Parthenocissus
Alnaster, Celtis-Aphananthe, Celtis, Diospyros, Elaegnus, Ilex, Ligustrum,
Mallotus, Platycarya, Rhamnus, Zelkova
Podocarpus
Araliaceae, Aucuba, Camellia, Castanopsis, Castanea-Castanopsis,
Celastraceae, Euphorbiaceae, Ilex, Illicium, Ligustrum, Moraceae, Myrica,
Quercus subgenus Cyclobalanopsis, Skimmia, Symplocos
Ilex, Skimmia, Viscum
Biome reconstructions for Japan
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
30
Biome reconstructions for Japan
31
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
Biome reconstructions for Japan
32
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