Global change and terrestrial ecosystems in monsoon Asia

190 Pages·1996·6.251 MB·English

by  M. Ohsawa (auth.), T. Hirose, B. H. Walker (eds.)

Global change and
terrestrial ecosystems
in monsoon Asia
Edited by
T. IDROSE and B. H. WA LKER
Reprinted from Vegetatio Volume 121
Springer-Science+Business Media, B.V.
Library of Congress Cataloging-in-Publication Data
Global change and terrestrial ecosystems in monsoon Asia I edited by
T. Hirose and B.H. Walker.
p. cm -- (Tasks for vegetation seienee : v. 33)
Ine ludes bibl iographleal referenees.
ISBN 978-94-010-4152-2 ISBN 978-94-011-0343-5 (eBook)
DOI 10.1007/978-94-011-0343-5
,. Forest eeology--Asia. 2, Forest mieraclimatalagy--Asia.
3. CI imatie ehanges--Asia. 1. Hirase, T. CTadaki) II. Walker, B.
H. (Brian Harrison), 1940- III. Series.
OK341 .058 1995
581.5'2642'095--dc20 95-34330
ISBN 978-94-010-4152-2
Printed on acid-free paper
All Rights Reserved
© 1996 Springer Science+Business Media Dordrecht
Originally published by Kluwer Academic Publishers in 1996
Softcover reprint ofthe hardcover Ist edition 1996
No part of the material protected by this copyright notice may be reproduced or
utilized in any form or by any means, electronic or mechanical,
including photocopying, recording or by any information storage and
retrieval system, without written permission from the copyright owner.
Table of contents
Editorial by T. Hirose and B. H. Walker vii
M. Ohsawa, Latitudinal comparison of altitudinal changes in forest structure, leaf-type, and species
richness in humid monsoon Asia 3
S.Yu. Grishin, The boreal forests of north-eastern Eurasia 11
T. Nakashizuka & S. !ida, Composition, dynamics and disturbance regime of temperate deciduous
forests in monsoon Asia 23
N. Ruangpanit, Tropical seasonal forests in monsoon Asia: With emphasis on continental southeast Asia 31
S. Riswan & L. Hartanti, Human impacts on tropical forest dynamics 41
G.W. Koch, P.M. Vitousek, W.L. Steffen & B.H. Walker, Terrestrial transects for global change
research 53
R.F. Sage, J. Santrucek & DJ. Grise, Temperature effects on the photosynthetic response of C plants to
3
long-term CO enrichment 67
2
E.-D. Schulze, R. Leuning & F.M. Kelliher, Environmental regulation of surface conductance for
evaporation from vegetation 79
K. Kikuzawa, The basis for variation in leaf longevity of plants 89
E.O. Box, Factors determining distributions of tree species and plant functional types 101
T. Kohyama & N. Shigesada, A size-distribution-based model of forest dynamics along a latitudinal
environmental gradient 117
Y. Iwasa, T. Kubo & K. Sato, Maintenance offorest species diversity and latitudinal gradient 127
Guofan Shao, H.H. Shugart & T.M. Smith, A role-type model (rope) and its application in assessing
climate change impacts on forest landscapes 135
K. Nakane & Nam-Juu Lee, Simulation of soil carbon cycling and carbon balance following clear-
cutting in a mid-temperate forest and contribution to the sink of atmospheric CO 147
2
P.G. Jarvis, The role of temperate trees and forests in CO fixation 157
2
G. Esser, Contribution of monsoon Asia to the carbon budget of the biosphere, past and future 175
Abstracts 189
Vegetatio 121: 1-2, 1995.
T. Hirose and B.H. Walker (eds). Global change and terrestrial ecosystems in monsoon Asia
©1995 Kluwer Academic Publishers.
Editorial
The world's terrestrial ecosystems are being subjected to global environmental change of an unprecedented scale,
both in their rate and in their geographical extent [Global Change and Terrestrial Ecosystems (GCTE), The
Operational Plan, IGBP Report 21, 1992]. An international workshop on 'Global Change Impacts on Terrestrial
Ecosystems in Monsoon Asia' was held in Tokyo, Japan, on 4-6 September 1993 to launch a research project
TEMA (Global Change Impacts on Terrestrial Ecosystems in Monsoon Asia). Twenty papers were presented at
the workshop and 16 papers are included in this volume (the other 4 papers are in abstract form). TEMA was
coordinated by the TEMA SSC of the Japan Subcommittee for GCTE and accepted as a contribution to Core
Research of GCTE in December 1992. The objectives of TEMA are (1) to predict the effects of elevated CO and
2
climatic change on the distribution and structure of forests in monsoon Asia and (2) to determine the associated
feedback effects to the global carbon cycle.
The TEMA project is based on the environmental gradient along a transect in monsoon Asia from boreal forests,
through cool and warm temperate forests, to tropical rain forests. This transect includes two high priority areas of
GCTE: one is the boreal forest region, which it is suggested will change most significantly in response to global
warming, and the other is tropical rain forests, which are endangered by rapid changes in land use under high
population pressure. Humid climate prevails over monsoon Asia. High precipitation with more or less conspicuous
rainy seasons is characteristic to this area. Under this climate the predominant vegetation is forests, which are
distributed without any intervening arid zone. Air temperature is the most important factor that determines the
type of forest vegetation along this transect. Another characteristic of this area is a range of high mountains. Air
temperature changes altitudinally as well as latitudinally and forest types change zonally with the temperature
gradient. The pattern of vegetation arrangement along the environmental gradient established under current climate
conditions forms a basis for the present study of global change impacts in monsoon Asia. TEMA has four research
components: (1) Screening of key species with respect of the response to global change. (2) Modelling of forest
structure as an integration of functional types. (3) Biogeographical analysis of the distribution and structure of
forest ecosystems in monsoon Asia. (4) Modelling of the carbon cycle of forest ecosystem in monsoon Asia. These
research components correspond more or less to the four sections of this volume.
The TEMA workshop was sponsored by the Science Council of Japan, Ministry of Education, Science and
Culture, Center for Global Environmental Research of the National Institute for Environmental Studies, Waseda
University and Aeon Group Foundation. The organizers gratefully acknowledge their support. Special thanks are
due to Makoto and Junko Nikkawa for their contributions in coordinating funding and administration.
T. Hirose and B. H. Walker
2
Reviewers
P. Chesson
D. L. DeAngelis
H. Diemont
1. R. Evans
R. M. Gifford
K. Harrison
K. Hikosaka
T. Hirose
M. Ishibashi
y. Iwasa
P. G. Jarvis
D. W. Johnson
S. Kojima
M. 1. Lechowicz
A. Makino
P. F. Maycock
T. Oikawa
K. Ono
1. Terashima
B. H. Walker
R. W. Wein
M. 1. A. Werger
F. 1. Woodward
Vegetatio 121: 3-10, 1995.
T. Hirose and B.H. Walker (eds). Global change and terrestrial ecosystems in monsoon Asia 3
©1995 Kluwer Academic Publishers.
Latitudinal comparison of altitudinal changes in forest structure, leaf-type,
and species richness in humid monsoon Asia
M.Ohsawa
Laboratory of Ecology, Chiba University, Yayoicho, Chiba 263, Japan
Accepted 13 February 1995
Key words: East Asia, Leaf size, Mountain vegetation, Temperature sum, Tree height, Tropical mountain, Vegetation
template
Abstract
A new template for mountain vegetation zonation along latitudinal gradients is proposed for examining geographical
pattern of various forest attributes in humid monsoon Asia. The contrasting temperature regime in tropical and
temperate mountains, i.e., the former is a non-seasonal, temperature-sum controlled environment, and the latter is
a seasonal, low temperature limiting environment, leads to different altitudinal patterns of tree height distribution
and species richness. In the tropical mountains, both tree height and species richness decrease steeply, and the
tree height often stepwise. The decline of tree height and species diversity in the temperate mountains is far less
pronounced except near the forest limit. Both trends are explained by their temperature regime.
Introduction are arranged along inclined parallels, i.e. high on the
mountains at low latitudes and low at higher latitudes
In southeast to east Asia, humid mountain forest (Ohsawa 1993a).
extends from the equatorial zone to the northern forest Such a basic structure of forest zonation in east
limit. This condition provides us with the best oppor Asia is largely determined by the present climatic con
tunity to study forest distribution in relation to tem ditions, particularly temperature (Wolfe 1979; Ohsawa
perature gradients along altitudes as well as latitudes 1995). In the present paper, some attributes of the forest
without a marked drought stress. Geographical patterns communities at different coordinates on the template
of forest zonation along the coordinates of latitude and are examined and compared in order to understand the
altitude has been described and examined by several underlying climatic rules of geographical organization
authors (cf. Wolfe 1979; Ohsawa 1990, 1993a). The of the vegetation in the humid Asian mountains.
most striking feature in latitudinal changes of altitudi
nal zonation in east Asia is the switch in zonation type
from tropical to temperate at ca. 20-30oN (Ohsawa Data
1993a). In the tropical type of zonation, all forest zones
from lowland to the forest limit are represented by ever Climatic data were collected from various stations
green trees, while in the temperate type, the dominants in southeast to east Asia (Ohsawa 1990). The sum
are successively replaced by different leaf-types from of monthly mean temperatures above 5 °C was used
evergreen broad-leaved trees at low altitude, to decid as the temperature sum, commonly used as "warmth
uous broad-leaved trees at middle and needle-leaved index" in Japan (Kira 1948; Tuhkanen 1980; Ohsawa
trees at high altitude. The configuration of each altitu 1990).
dinal zone is also different in both types of zonation. In Floristic and structural data on various humid
the tropical type, all the zones are tiered, or arranged forests in the southeast and east Asian mountains
horizontally parallel to latitude, while in the temperate were collected from different sources in the litera
type above the transition zone, the altitudinal zones ture (Brown 1919; Ohsawa 1983, 1984; Ohsawa et
4
al. 1973, 1985; Irikura 1984; Kitayama 1992). Only
data based on quadrat sampling were included in the
analysis.
TRANSITION ZONE
'S
I
Results ZONATION TYPE TROPICAL TYPE TEMPERATE TYPE
5000
The mountain-vegetation template for humid
monsoon Asia 4000 1------..1
Upper Montane
Various templates depicting vegetation distribution 3000
have been proposed to understand the interrelation
ships between formation or ecosystem types and their 2000 Lower Montane
environmental factors on a regional or worldwide scale
1000f------+____.
(Troll 1948; Dansereau 1957; Holdridge 1967; Whit
lowland
taker 1975 etc.). Most of these use the two axes of
oL--~_~_~L_J__~~~ _ _"
temperature and precipitation, or the triangle with an 10°8 10 20 30 40
additional factor of humidity that is derived from the Tropical __________ LATITUDINAL ZONATION
above two factors. Of these, the most notable and use Subtropical
ful one for understanding the pattern of mountain veg Cool-temperate
etation is Troll's diagram (Troll 1948). The diagram Cold-temperate
well describes the different patterns of vegetation zona 10 20 30 40 50
LATITUDE
tion on humid mountain chains from the Northern to
Southern hemisphere, and also indicates the difference
between tropical and temperate zonation in the North Fig. 1. Mountain-vegetation template for southeast and east Asia.
The boundary of zones are drawn with some modification from data
ern hemisphere. Troll's diagram helped to establish
on several high mountains in the region. Detailed explanation can
the fact that there occur changes in leaf-type at the for be found in Ohsawa (1990, 1993a).
est limit from the conifer type in the temperate to the
evergreen broad-leaved in the tropical zone.
Ohsawa (1990) has explained the controlling fac limit at ca. 20oN, and from that point onwards, due
tors of this latitudinal change in the leaf-type at the to winter coldness, the evergreen broad-leaved trees
forest limit by distinguishing between two temperature cannot reach the forest limit any longer.
determinants, summer heat and winter coldness. The On the basis of these facts, Ohsawa (1993a) pro
summer heat or the temperature sum (warmth index) of posed a new scheme for the mountain vegetation zona
15°C· months controls the altitude of the forest limit. tion in humid southeast to east Asia, which is proposed
This temperature sum condition correlates closely with to be a mUltipurpose mountain-vegetation template
the warmest month temperature of 10 ° C proposed by (Fig. 1). To draw a picture of latitudinal change of alti
Koppen (1884) for temperate latitudes, but it deviates tudinal zonation for the humid mountains of the globe,
towards low and high monthly mean temperatures of Troll (1948) had to combine mountain chains from
the warmest month in non-seasonal tropical climates the different continents, such as Spitzbergen, Europe,
and in strongly seasonal high latitudes, respectively Himalaya and Mexico, due to lack of data from even
(Ohsawa 1990). On the other hand, the winter coldness the Northern hemisphere. The southeast to east Asian
or the coldest month temperature controls the leaf-type mountains, on the contrary, provide relatively continu
at the forest limit. The coldest month temperature of 0 ous mountain chains from the equatorial region to the
to -1°C controls the upper limit of evergreen-broad northern forest limit in Siberia. Here the template can
leaved trees (cf. Ohsawa 1990), but in the equatorial be used to examine the geographical pattern in forest
region the coldest month temperature of 0 to -1°C attributes along altitudinal and latitudinal temperature
occur at ca. 4700 m, high above the forest limit, and has gradients in a continuous series.
no effect on the distribution of evergreen broad-leaved The framework of the mountain-vegetation tem
trees. The temperature curve of 0 to -1°C drops with plate suggests the importance of increasing seasonality
higher latitudes; it crosses the altitude of the forest in temperature for the differentiation of the two moun-
5
tain zonation types in the Northern hemisphere. It is (1967). The temperature sum (WI) has a significant
well known that, in temperate mountains the increas linear relationship with the potential evapotranspira
ing winter coldness limits the plant distribution to high tion value (PET) at least in the range of 15 to 200
altitudes or to the north (Iversen 1944; Woodward °C . months in which tropical to temperate forests are
1988; Ohsawa 1990). In tropical mountains, howev developed (Fang & Yoda 1989). If there is no signifi
er, the altitudinal distribution of forest is controlled by cant drought condition throughout a year, PET nearly
the temperature sum (Ohsawa 1993b). Thus the rel equals the actual evapotranspiration (AET). Frequent
ative importance of the two temperature conditions, ly the available energy for plants is indicated by AET,
i.e., temperature sum and winter temperature, shift at and a linear relationship has been found between AET
around 20-30oN where the annual range of tempera and net above-ground productivity (Rosenzweig 1968;
ture is ca. 10-20 °C. This template is largely based Currie 1991). Thus the temperature sum also shows a
on continuous mountain chains, and thus it is appro linear correlation with AET and net productivity. The
priate to overlay various community attributes on the amount of energy expressed as temperature sum along
template. This will provide us with further insights on an altitudinal gradients has a much steeper inclination
the interrelationships between climatic conditions, dis in tropical mountains than in temperate ones (Fig. 3).
tribution of formation types, and various community Therefore, for the same increase in altitude, energy
attributes. or the productivity decreases more in tropical moun
tains than in temperate ones. Kitayama (1992) stated
Latitudinal comparison of altitudinal changes in tree that the tropical zonation distinguished on the basis of
height the temperature sum (Kira's WI) is more compressed,
than the zonation in Koppen's system. This is solely
The most simple, easily available, and relatively accu due to the lack of seasonality in the temperature, i.e.,
rate indicator of forest biomass is the maximum tree gradients in temperature sum are steeper than those in
height when the forest is in closed condition. Tree monthly mean temperature. In the tropical zonation,
height is highest in the tropical lowland and decreases high-altitude forests have dropped the overstory trees
at both high altitudes and high latitudes. The altitudinal of the previous lower zone (cf. Ohsawa 1991). This
change in maximum tree height of the forest shows a may suggest that energy shortage cannot support taller
different pattern in the tropical and temperate moun trees in forests at successively higher altitude. A simi
tains of southeast and east Asia (Fig. 2). In the trop lar idea has been proposed as one of the causes of the
ical zonation, tree height decreased step-wise, i.e., it forest limit, emphasizing the structural features at the
changed by nearly half from one zone to the next high boundary (Stevens & Fox 1991).
er one. On the other hand, in the temperate zonation, In temperate mountains, however, the controlling
though the physiognomy or leaf-type changed in each factor of the forest type is not energy but rather winter
zone from evergreen to deciduous and then to conifer coldness. Since the summer temperature in temper
ous, the forest has similar height from the foothill to ate mountains is high enough to support high produc
high altitudes, except very near the forest limit where tivity, the decreasing rate of the temperature sum is
the scrub was found dominating (Fig. 2). low compared with that in tropical mountains (Fig.
These facts clearly indicate that the controlling fac 3). Therefore, the forest mass expressed as tree height
tors of tropical zonation are different from those of shows little change and only leaf-type which expresses
temperate zonation. Two important factors may keep adaptation to winter coldness, changes drastically with
the forest short, (i) reduced biomass production and (ii) decreasing altitude (Fig. 2). However, of all bound
a high rate of population turnover. The latter, however, aries in the humid temperate mountain zonation, only
is not plausible under the low temperature conditions the forest limit is controlled by summer temperature or
at high altitudes. Tropical mountains have no season, the temperature sum (Ohsawa 1990). The forest lim
and therefore, many of the structural and dynamic it is the only "structural" boundary, namely between
features of the forest, including productivity, forest forest and grassland, in the temperate zonation, while
biomass, or stratification, etc., are controlled by the in the tropical zonation every boundary represents a
year-round temperature condition as a whole, which is "structural" change of the top layer of the forest (Fig.
often well expressed, e.g., by the temperature sum (WI, 2).
a threshold of 5°C), or by the biotemperature (a lower
threshold of 0 and a higher of 30°C) of Holdridge
6
TROPICAL TEMPERATE
30
5000 () 30~
! 20 alt. 1200 m ()
CMT = -1 C E::I
elt. 2500 m
~" 10 ______________ olt.3500m
4000 I'" 0 4000 1>'"7'-T--T'<,,", .11. BOO ..
0" . olt. 1&00 m
-.......~ Temperature sum = 15 C·months lit. 2450 m
E 3000 0 :\0 0.)= 3000
Ev (Mi) 0
Q)
§'C --- - - - -~ • ~ Temperature sum __ ~ Temperature sum = 15 C'months
0;::-..Q
85 C'months o 00 " '-0
<I: 2000 Evergreen ~ 2000
(Notophyll) •• 0 Conifer 8~
- - - - - - - - --0-0 Absolute minimum = -40 C
-------------• --.---- •• CMT = -6 - -8' C
Deciduous , ~
1000 • 1000 • \ ~
- - - - - - - - - - - -:t'-Absolute minimum = -15 C
Evergreen (Mesophyll) Evergreen . :, CMT = -1 C
(Notophyll) ..
0~0----------~20~----------4-0------L--
20 40
Height, m Height, m
Fig. 2. Decreasing trend in tree height in tropical and temperate zonation. Data on tropical mountains, open circles for Mount Kerinci (after
Ohsawa et al. 1985), and closed circles for Mount Kinabalu (after Kitayama 1992), and on temperate mountains: open circles for Mount
Fuji (Ohsawa 1984), and closed circles for Yakushima (Irikura 1984). Some additional explanations are Bv (Mi) for evergreen trees with
microphyllous leaves. eMT: monthly mean temperature of the coldest month. Yearly courses of monthly mean temperature at critical altitudes
are inset in the figure.
Change in leaf size 1979) might be the best term to indicate the habitat
conditions, in general, which lead to microphyllous
As has been reported by Richards (1952), Grubb (1974) forest. Whatever might be the case, notophyllous trees
etc., with increasing altitude leaf size of tropical moun are competitively superior to microphyllous trees both
tain forest decreases from mesophyllous in lowland, to in their initial growth rate and also in the maximum
notophyllous in lower montane, and to microphyllous attainable tree height in favourable, mesic sites. And,
in upper montane forest. This tendency is, however, not only the conditions which suppress the notophyllous
only encountered on high mountains, but also on vari trees can accommodate microphyllous trees to dom
ous small peaks often covered by clouds (cloud forest), inate (Ohsawa 1993b). The change in leaf-size with
at windy ridge tops, etc. (Whitmore 1984). The micro altitude in tropical mountains is a clear phenomenon
phyllous forest roughly coincides with the cloud forest which can be illustrated on the template, but the pat
in tropical upper montane zone (Ohsawa 1995). At the tern still needs further ecophysiological and commu
northern latitudinal limit of tropical montane forests nity ecological study.
or the subtropical warm-temperate forests in southern
Japan, the change from notophyllous to microphyllous Altitudinal patterns of species richness
evergreen broad-leaved forest can often be observed
along the successional series, in the stratification of a Many hypotheses have been proposed to explain the
well developed forest, and along stress gradients such gradational pattern of species richness along various
as ridge-to-valley, windy mountain top to foothill, etc. environmental gradients including latitude, longitude,
(Ohsawa 1993b). Therefore, the change is caused not altitude, and other specific factors, such as tempera
only by the decrease in temperature sum but is also ture, humidity, etc. (Iwasa et al. 1993; Huston 1994).
affected by various other factors. Stress (sensu Grime There are many reports on a decreasing pattern of
7
5000 incomplete data, show a strong contrast between trop
ical and temperate mountains. The tropical mountains,
including Mt. Kerinci, Mt. Kinabalu, and Mt. Maquil
4000 ing show a very steep decrease in species diversity
Forest Limit with increasing of altitude. In contrast, the temperate
mountains show rather small changes in species diver
E
3000 sity along altitudinal gradients.
(l) The similarity in the distributional patterns of
-0
:::l species diversity and of tree height along altitudinal
~ \
«'.i=j ::\ gradients in both tropical and temperate mountains
2000 Mt. Kinabalu brings me to generate a hypothesis considering the
\
\ (60 5') common factor that controls both biomass distribution
and the pattern in species diversity. Currie (1991) has
1000 shown a strong relationship between tree-species rich
\ Mt. Fuji
ness and AET. Adams & Woodward (1989) have clear
\ (350 21 ')
'\ ly demonstrated a close linear relationship between
tree species richness and primary productivity in the
0
northern temperate zone. Thus, the energy or produc
0 100 200 300
tivity hypothesis seems again valid to explain the dif
Temperature sum (WI), , C' month ferent patterns in species diversity in tropical and tem
perate mountains, and the common factor of energy
Fig. 3. Altitudinal changes in temperature sum (WI) in a tropical
seems essential to understand the structural differences
mountain (Mount Kinabalu at 6° 5'), and in a temperate mountain
(Mount Fuji at 35° 21'). between tropical and temperate mountains. The steep
depression in species diversity with altitude on tropi
cal mountains as compared to temperate mountains, is
species richness along elevational gradients, howev probably caused by the steep decline in energy in the
er only few hypotheses have so far dealt with the cause former (Fig. 3). In temperate mountains, the increasing
of the altitudinal change in species richness (Gentry altitude gives rise mainly to the shortening of growing
1988). Fig. 4 shows the template with species rich period or the lowering of winter temperature rather than
ness data, expressed as the diversity index = (s-1 )/Iog to an energy shortage. It does not affect species diversi
n, where s is the number of species and n the num ty nor tree height very much along altitudinal gradient.
ber of tree individuals (Margalef 1968). The patterns This issue, however, can only be verified by comparing
of species diversity greatly differ between tropical and biodiversity as well as forest structure along altitudi
temperate mountains. The highest diversity, in the trop nal gradients in tropical mountains, because the non
icallowlands decrease steeply towards the forest limit seasonal environment results in a gradational change
and towards higher latitudes. In temperate mountains, in terms of energy but not seasonal change in temper
lowland species diversity is already rather low and the ature.
decrease with altitude is not as prominent as in trop Large differences in biomass productivity between
ical mountains. Fig. 5 shows the altitudinal changes temperate and tropical regions can often be ascribed
in species diversity in each mountain. Most of the solely to differences in the length of the growing season
data include high altitude forests up to the forest limit. (Ryabchikov 1975; Rodin et al. 1975). This suggests
In some mountains, unfortunately, no sites have been that the productive efficiency does not differ between
sampled at low altitudes. However, e.g. the lowest sam the tropical and temperate regions. The shorter grow
pling point at 1600 m at Mount Kerinci has a similar ing season in temperate mountains inevitably is less
diversity as that at the same altitude on Mount Kina productive and may give rise fo shorter tree heights and
balu, and both are mountains of a similar scale with the decreased species richness. Jordan & Murphy (1978),
peak of the former at 3800 m and the latter at 4101 m. however, hypothesized that the productive potential is
Mount Maquiling, though having a rather high, similar higher in temperate regions than in tropical regions
diversity as the other tropical mountain in its foothills, due to a higher efficiency in temperate trees, as they
has a peak of 1050 m and is too small to compare. It can had adapted to the limited amount of solar radiation.
be used only for lowland species diversity. Even such This issue is still open for discussion, and only detailed