REPORTS
Chinese Science Bulletin 2003 Vol. 48 No. 3 291 295
analogue to reconstruct the past vegetations.
Surface pollen and vegetation
reconstruction from central
and northern mountains of
Taiwan
1 Modern climate and vegetation
YU Ge1, LIEW Pingmei2, XUE Bin1 & LI Zhengyi2
1. Nanjing Institute of Geography & Limnolog of the Chinese Academy
of Sciences, Nanjing 210008, China;
2. Department of Geology, Taiwan University, Taibei 106-17, Taiwan,
China
Abstract Surface pollen from various montane vegetations in Shalixian Mt. and Lepei Mt. of central and northern
Taiwan have been explored and analyzed. The pollen data
were used to simulate vegetation types by using the biomisation technique. Computing three matrixes of pollen taxa,
plant functional types and combined biomes, vegetation
types at each site have been finally defined through the fuzz
selections of a tie-breaking rule. The results show that surface pollen data can simulate the subtropical Castanopsis and
Cyclobalanopsis forests, alpine warm-temperate Quercus and
Quercus-Pinus forests, and alpine temperate/cool-temperate
conifers of Tsuga-Picea and Picea-Abies forests. Simulated
elevations of the forests are similar to actual locations of the
forests. This study can add vertical surface pollen data for
extending investigation of various vegetation types in China,
provide PFT schemes and vegetation types in low-latitude
and montane areas, and be applied as modern analogues for
fossil pollen in order to reconstruct Quaternary vegetations.
Keywords: surface pollen, vertical vegetation, Taiwan mountains.
Late-Quaternary vegetations have been changed
sysmatically in the Northern Hemisphere. Although
vegetations in low latitudes were not more significantly
changed in geographical shift than that in mid-high latitudes, climate and vegetation changes in tropic mountains
have shown their strong singles of the Earth dynamics [1,2].
Pollen data have been used to simulate the past vegetation
of China and showed the spatial shift of vegetation
zones[3,4] . However, there was little work that involves in
the vertical vegetations; this is mainly due to lack of pollen data. Taiwan Island is located in low latitudes of the
tropic and subtropical area. High mountains with large
elevation differences and Asian winter/summer monsoons
have played important roles on evolution of the montane
climate-vegetations with 3-dimensional types, which have
developed vertical climate-vegetation zones from tropic to
cold desert[4]. The plant diversity and montane vegetation
complex of Taiwan have provided an idea area to study
low-latitude pollen records and to carry on biome simulations. Climate-vegetation changes in low latitudes were
shown mainly in vertical changes from low to high elevations, which can provide a scientific basis of the modern
Chinese Science Bulletin Vol. 48 No. 3 February 2003
Taiwan Island is the largest island of China. The area
is 36000 km2 and the geographical location is between 25
21 N. Although the latitudes span ca. 4 , the difference of elevation is more than 4000 m that have produced
various vertical climate zones. According to analysis of
climate data from 200 meteorological stations in Taiwan[5],
the correlation relationship between temperature and elevation is very significant, and temperature is the largest
weight in all climate elements. Different montane tropic,
subtropical, temperate, and alpine cold vegetations have
occurred with increase of elevations in Taiwan mountains.
The different annual temperatures in the mountains have
controlled distributions of different vegetations. From low
to high elevations in central Taiwan, there are Ficus-Machilus forest, Machilus-Castanopsis forest, Cyclobalanopsis-Trochdendron forest, Quercus forest, Alnus forest,
Chamecyparis forest, Pinus and Pinus-Quercus forest,
Tsuga-Picea forest, Abies forest, Juniperus forest, Rhododendron shrubland, and bamboo grove [6] (see Table 1 as
the summary).
2 Data and method
Two sets of pollen samples were collected from
various montane vegetations in Shalixian Mt. of central
Taiwan and Lepei Mt. of northern Taiwan. Shalixian Mt.
is located between 23°28 — 23°33 N and 120°53 120°59 E,
where its south end is at Yushan Mt. (3952 m), reaching
eastward to Beifeng Mt. (3850 m), westwards to Xishan
Mt. (3528 m) and Qioanshan Mt. (3236 m), northwards to
Lanxi of Dazhushan Mt. (2853 m) and convergent to the
Shalixian River. The lowest valley in the catchment is at
500 m that valley opens to the north, and the highest point
at the Yushan Mt. The elevation difference between two
places reaches 2752 m. 26 pollen samples were collected
from forest/shrubland soils at the elevations between 750
and 3400 m. Elevation of Ficus- Machilus forest is below
500 m. Elevation of Machilus- Castanopsis forest is between 500 1500 m, and Cyclobalanopsis-Quercu forest
between 1500 2500 m. Elevation of Tsuga-Picea forest
is above 2500 m, Abies forest above 3100 m, and Juniperus conifer and Rhododendron shrubland above 3600 m.
The Leperi Mt. is located in 24°05 N and 121°02 E. The
highest point is at 1582 m and the lowest valley is between 350 700 m, which elevations difference is ca.
1200 m. There is deciduous Quercus forest above 1200 m,
Machilus-Cas- tanopsis forest below 1200 m, and artificially planting cryptomeria forest below 1100 m in the
mountain. 36 pollen samples were collected from different
soils of the forest/shrubland. The geographical locations
and elevations of two sets of pollen samples are shown in
Fig. 1.
291
REPORTS
Table 1
Elevation/m
>3600
Annual
temperature/
<5
3100 3600
2500 3100
5 8
8 11
2000 2500
11 14
1500 2000
14 17
500 1500
17 23
<500
>23
Montane climate-vegetation zones in Taiwan
Vegetation zone
Dominant plant
alpine vegetation
bamboo grove,
Miscanthus
Juniperus, Rhododendron, Pinus
Abies
Tsuga, Picea
Major plant
Yushania niitakayamensis
Climate zone
Miscanthus
subarctic
J. squamata, Rh. pseudochrysanthemum,
P. taiwanensis, P. morrisonicols
fir forest
A. kawskamii
cold-temperature
Homlock-spruce forest
Tsuga chinensis, Picea morrisonicola,
cool-temperature
Pinus armandii mastersiana
deciduous oak forest,
Quercus,
temperate
Ch. formosanansis, Ch. taiwania, Cunningupper layer
Alnus, Chamecyparis, hamia, Taxua, Tsuga, Picea;
Pinus
Pseudotsuga wilsoniana, Calocadrus formosana;
Quercus spinoaa miyabei, Q. variabilis,
Alnus formosana;
Acer, Juglans, Ulmus, Carpinus, Platycarya
evergreen oak forest,
Cyclobalanopsis,
Cyc. morii, Cyc. stenophylloides, Cas. warm-temperate
lower layer
Trochdendron
cariesii, Cyc. longinux, Cyc. gilva, Litsea,
Lithocarpus;
Pistacia,
Rhus,
Trema,
Sapindus,
Aphananthe, Koelreuteria, Hacranga,
Zelkova, Fraxinus, Albizia, Liquidambar,
Glochidion, Celtis, Sapium
Machilus-Castanopsis Machilus, Castanopsis M. japonica, M. kusanoi, Ficus, Turpihia,
subtropical
Lagerstroemia, C. hystrix, C. kawakamii,
forest
Schima superba, Engeihardtia,
Lithocarpus, tree fern
tropical
Ficus-Machilus forest Ficus, Machilus
Ficus, Machilus Laportea, Bischfia, Acacia
confusa, Leucaena, Jeucocephala, Vitex
negundo, Clerodendron, Dodonaes
Fig. 1. Geographical locations of two sites in Taiwan and elevations of the pollen samples. 1, 36 samples in Lepei Mt., 2, 26 samples in Shalixian Mt.
292
Chinese Science Bulletin Vol. 48 No. 3 February 2003
REPORTS
The method we used in the present work is the biomisation technique[7 ]. This method was originally developed from Europe, and has been applied in China and
validated by modern Chinese vegetations[3,4]. The method
used in Hainan Island has proved that it enables simulating the sub-scale vegetation types in tropical/subtropical
areas[8]. Assigning Plant Functional Types (PFTs) helps to
classify plant ecology for treating pollen taxa. Certain
basic physiognomic characteristics generally appear in
functional-type classifications, namely life-form (arboreal/nonarboreal), leaf-form (broad/needle), phenology
(evergreen/summer green), and criteria controlled by climate (warm/cold, dry/wet, etc.). We assign PFTs
broadleaved evergreen, broad-leaved deciduous, evergreen
conifer, forb/shrub and fern. Climate index is one important key to assigning the PFTs, while temperature is a
major control element in Taiwan mountains. On the basis
of Taiwan annual temperature extents (Table 1),
broadleaved deciduous has been divided into tropical
evergreen, subtropical evergreen, southern warmtemperate evergreen, warm-temperate evergreen, and
cool-temperate evergreen. Broadleaved deciduous has
Vegetation type
Broadleaved
evergreen
Broadleaved
deciduous
been further assigned to cool temperate summer green,
intermediate temperate summer green, southern warmtemperate summer green, and temperate summer green.
Evergreen conifer is divided into warm-temperate conifer,
temperate conifers, cool temperate conifer, boreal evergreen conifer, and eurythermic conifer. For the forb/shrub,
there are tropical and subtropical evergreen forbs/shrub,
temperate forbs/shrub, arctic-alpine forbs/shrub, steep
forb/shrub, and desert forb/shrub differently (Table 2).
Based on PFTs, we further combine the PFTs into vegetation defined by these known criteria largely controlled by
climate. Although there are different classifications of
global biomes, the biomisation adapted 14 zonal vegetation types without non-zonal vegetations (such as swamp
and mangrove) or artificial coverage (such as farmland,
economic forest)[7]. As regional assigning and mapping,
there were not Savanna or Mediterranean vegetations in
China so that there were 12 vegetation types applied [3].
The present work has produced 10 montane vegetation
types according to the local vegetation character of Taiwan (see Table 3).
Table 2 PFTs in Taiwan and the pollen taxa included
Pollen taxa
PFT code
te
tropical evergreen
Cassia, Ficus, Helicia, Homalanthus, Leguminosae, Medinilla, Palmae,
Piperaceae, Randia, Trema
subtropical evergreen
Kleinhovia, Leguminosae, Medinilla, Syzygium, Terminalia
tr
southern warm-temperate eve rgreen
Castanopsis, Lithocarpus
wte2
warm-temperate evergreen
Capparis, Acanthaceae, Actinidia, Anacardiaceae, Aralia, Araliaceae,
wte
Bischofia, Bredia, Cyclobalanopsis, Elaeocarpus, Engelhardtia,
Glochidion, Koelreuteria, Macaranga, Maesa, Mallotus, Meliaceae,
Moraceae, Oleaceae, Pasania, Rhamnaceae, Rubiaceae, Rutaceae,
Saurauja, Schefflera, Sycopsis, Symplocaceae, Symplocos, Zanthoxyllum, Trochodendron
cool-temperate evergreen
Ilex, Ligustrum
wte1
cool-temperate summe rgreen
Alnus, Carpinus,Ulmus
ts1
PFT
intermediate-temperate summer green
Carya, Celtis, Diospyros, Elaeagnus, Fagus, Juglans, Liquidambar,
Myrica, Pterocarya, Quercus_(deciduous), Urticaceae
southern warm-temperate summer green
Rhus, Sapium, Taxillus, Zelkova
temperate summer green
Acanthopanax, Acer, Clematis, Fraxinus, Osmanthus
Evergreen conifer eurythermic conifer
Pinus_(diploxylon), Juniperus
warm-temperate conifer
Chamaecyparis, Cryptomeria, Keteleeria, Podocarpus
temperate conifers
Taxodiaceae
cool temperate conifer
Picea, Tsuga
boreal evergreen conifer
Abies
Forb and shrub tropical and subtropical eve rgreen forbs/shrub Arabis, Araceae, Davallia, Evolvalus, Gesneriaceae, Gymnosporium,
Hedryotis, Hiptage, Impatiens, Litosanthes, Microlepia, Strobilanthes
temperate forbs/shrub
Euphorbia, Aconitum, Actinostema, Chenopodium, Convolvulaceae,
Crawfurdia, Cucurbitaceae, Cuscuta, Galium, Humulus, Labiatae,
Paraphlomis, Ranunculaceae, Rhododendron, Scrophulariaceae, Solanaceae, Umbelliferae, Valeriana
arctic-alpine forbs/shrub
Ericaceae, Compositae, Cruciferae, Polygonum, Rhododendron,
Thalictrum, Salix
desert forb/shrub
Tamarix
steppe forb/shrub
Artemisia, Justicia
Fern
tree ferns
Cyatheaceae
undifferentiated ferns
Dryopteris, Lepisorus, Lycopodium, Polypodiaceae, Pteris, Selaginella
Others
grass
Gramineae
sedge
Cyperaceae
Chinese Science Bulletin Vol. 48 No. 3 February 2003
ts2
ts3
ts
ec
wtc
tc
ctc
bec
tef
tf
af
df
sf
tx
x
g
s
293
REPORTS
Table 3 Montane biomes in Taiwan and combinations of the PFTs (PFT code as in Table 2)
Vegetation zone
PFTs included
Tropical evergreen forest
te + tx + tef
Subtropical evergreen forest
tr + tx + wte + ts3+ tef
Southern warm-temperate e vergreen forest
wte1 + wte2 + ts3 + tef + tf
Warm-temperate mixed conifer and deciduous forest
wte1 + wtc + tc + ec+ ts3 + ts2 + ts1 + ts + tf
Cool-temperate conifer forest
ctc + ec + h + g
Cold-temperate conifer forest
bec + ec + h + g
Alpine conifer forest
ec + af + g
Alpine shrub land
tf + sf + af + ax + g
Alpine tundra
af + s + g
Alpine desert
af + df + g
The biomisation procedure is briefly described as the
following: (1) each pollen taxon is assigned to one or
more PFTs on the basis of the known biology of the plants.
This first step produces a relation between PFTs and pollen taxa (a PFT × taxon matrix). (2) By knowing which
PFTs occur in each biome, we then derive a biome ×
PFT matrix. (3) Both matrices (PFT × taxon and biome
× PFT) are combined to yield a biome × taxon matrix.
(4) Affinity scores are then calculated for all pollen. (5)
Finally the pollen samples are each assigned to the biome
with which they have maximum affinity. The only additional rule required is a tie-breaking rule, to deal with
cases where a pollen sample has exactly equal affinity
with more than one biome. Through the biomisation computing for each pollen sample, each biome can be defined
finally.
3 Results and discussion
The results of simulated vegetations are plotted in
Fig. 2. From low elevation to high elevation, we have
simulated tropic and subtropical evergreen forests, warmtemperate evergreen and deciduous forests, mixed forest,
temperate and cold conifer forest. Castanopsis-dominant
forest occurs between 700 1530 m in the Shalixian Mt.
and between 540 1100 m in the Lepei Mt. These simulations are closed to elevation distributions of the actual
Castanopsis forest over Shalixian Mt. 1) and Lepei Mt.2).
Cyclobalanopsi-dominant forest occurs between 1780
2150 m in Shalixian Mt. and between 700 1500 m in the
Lepei Mt. Quercus-dominant temperate deciduous forest
in the Shalixian Mt. was simulated between 2250 2500
m, cool-temperate conifer of Tsuga- Picea forest between
2600 2800 m, and cold temperate conifer of Abiesdominant forest above 3400 m. These simulated biomes
and their elevation distributions can be compared very
well with the corresponding natural forests in the Shalixian Mt.
Surface pollen from Taiwan mountains show their
complex species and weak dominant-taxa, indicating the
plant diversity of tropic vegetation. These pollen characteristics are quite similar to surface pollen from Hainan
Island[7]. We did not find pollen Machilus neither in samples of the Ficus-Machilus forest nor in samples of the
Machilu-Castanopsis forest. Even there is not Machilus
pollen information from airborne pollen records[10]. This
mismatch phenomenon between Machilus pollen and
Machilus plant is mainly due to its very thin pollen-wall
and less-preserved. The limit of pollen Machilus made us
mainly rely on pollen Ficus to treat the Ficus-Machilus
forest, and rely on pollen Castanopsis to treat the Machilus-Castanopsis forest.
Castanopsis and Cyclobalanopsis are species in both
subtropical and war-temperate broadleaved evergreens,
which were treated the PFT as warm-temperate evergreen
(wte)[3,10]. In Taiwan vegetations, they are two dominant
plants in subtropical evergreen Machilus-Castanopsis
forest and warm-temperate evergreen CyclobalanopsisTrochdendron forest respectively, which are growing at
500 1500 m and 1500 2000 m and controlled by annual temperature 17 23
and 14 17
differently.
Therefore, on basis of known PFT wte, we added a new
PFT of wte2 (southern warm-temperate evergreen) and
assigned Castanopsis as wte2. In this case, we can distinguish wte2 of Castanopsis from wte of Cyclobalanopsis
and simulate subtropical evergreen Castanopsis-dominant
forest and warm-temperate evergreen Cyclobalanopsisdominant forest separately. The results showed that this
design is correct.
Due to human activities, natural vegetation below
500 m has been largely disturbed so that pollen data cannot show the zonal vegetation well. Compared with vertical vegetation zones, the surface pollen from two mountains can well-simulate vegetation types above 500 m,
including the subtropic Castanopsis and Cyclobalanopsis
1) Zhong Nianjun, Ecology and preservation in Shalixian area of Central Taiwan, Ph. D. thesis, Institute of Forestry, Taiwan University, 1994, 3
15.
2) Xu Yuemei, Constructions and components of warm-temperate rain forest from the Lepei Mt. in northern Taiwan, MS. Thesis, Institute of
Botany, Taiwan University, 1991, 2 8.
294
Chinese Science Bulletin Vol. 48 No. 3 February 2003
REPORTS
Fig. 2. Boimisation-simulated vegetation from Taiwan mountains: (a) Lepei Mt., (b) Shalixian Mt. The dash lines show the lower boundaries of
simulated vegetation and actual vegetation in order to compare. Codes for boimisation-simulated vegetation are: 1, tropical evergreen forest; 2, subtropical evergreen forest; 3, southern warm-temperate evergreen forest; 4, warm-temperate mixed conifer and deciduous forest; 5, cool-temperate conifer forest; 6, cold-temperate conifer forest; 7, alpine conifer forest; 8, Alpine shrub land; 9, alpine tundra; 10, alpine desert.
forests, alpine warm-temperate Quercus and QuercusPinus forests, and alpine temperate/cool-temperate conifers of Tsuga-Picea and Picea-Abies forests. Simulated
forest locations and elevations are similar to each actual
forest. This work can extend surface pollen coverage of
China into Taiwan areas, provide the PFT and biome
schemes in tropic/subtropical mountains, improve the
modern analogues of biomistion in vertical vegetations
in order to reconstruct the past biome environments.
Acknowledgements This work was supported by Nanjing Institue of
Geography & Limnology (Grant No. CXNIGLAS-A02-06) and Department of Geology, Taiwan University.
References
1.
2.
3.
Liew, P. M., Kuo, C. M., Huang, S. Y. et al., Vegetation change
and terrestrial carbon storage in eastern Asia during the Last Glacial Maximum as indicated by a new pollen record from central
Taiwan, Global and Planetary Change, 1998(16-17): 85 95.
Farrera, I., Harrison, S. P., Prentice, I. C. et al., Tropical climates
at the last glacial maximum: a new synthesis of terrestrial palaeoclimate data, . Vegetation, lake-levels and geochemistry, Climate
Dynamics, 1999, 15: 823 856.
Yu, G., Chen, X. D., Ni, J. et al., Palaeovegetation of China: a
Chinese Science Bulletin Vol. 48 No. 3 February 2003
4.
5.
6.
7.
8.
9.
10.
pollen data-based synthesis for the mid-Holocene and last glacial
maximum, Journal of Biogeography, 2000, 27: 635 664.
Members of China Quaternary Pollen Data Base, Pollen based
biome reconstruction at Middle Holocene (6 kaBP) and Last Glacial Maximum (18 kaBP) in China, Acta Botanica Sinica (in Chinese), 2000, 42(11): 1201 1209.
Su, H. J., Studies on the climate and vegetation types of the natural forest in Taiwan ( ): Analysis of the variation in climate factors, Quarterly Journal of Chinese Forest, 1984a, 17(3): 1 14.
Su, H. J., Studies on the climate and vegetation types of the natural forest in Taiwan ( ): Altitudinal variation zones in relation to
temperature gradient, Quarterly Journal of Chinese Forest, 1984b,
17(4): 57 73.
Prentice, I. C., Guiot, J., Huntley, B. et al., Reconstructing biomes
from palaeoecological data: a general method and its application
to European pollen data at 0 and 6 ka, Climate Dynamics, 1996,
12(12): 185 194.
Yu, G., Han, H. Y., Surface pollen and tropical vegetation simulations on Hainan Island, China, Marine Geology & Quaternary,
Geology (in Chinese), 1998, 18 (3): 103 111.
Yang, Y. –L., Chen, S. -H., An investigation of airborne pollen in
Taipei City, Taiwan, 1993 1994, Journal of Plant Research, 1998,
111: 501 508.
Yu, G., Prentice, C. I., Harrison, P. S. et al., Pollen-based biome
reconstructions for China at 0 ka and 6 ka, Journal of Biogeogr aphy, 1998, 25: 1055 1069.
(Received September 6, 2002)
295