Table Of ContentEcology of Grasslands and Bamboolands in the World
Ecology of Grasslands
and Bamboolands in theWo rld
Edited by
Professor Dr. Makoto Numata, ChibajJapan
in collaboration with 15 scientists
With 102 Figures and 58 Tables
Dr. W. Junk bv Publishers The Hague-Boston-London 1979
First edition
All rights reserved
© VEB Gustav Fischer Verlag Jena, 1979
Softcover reprint of the hardcover I st edition 1979
Licence Nr. 261700/167/79
coedition by Dr. W. Junk b. v. Publishers, The Hague
ISBN-13: 978-94-009-9187-3 e-ISBN-13: 978-94-009-9185-9
001: 10.\007/978-94-009-9185-9
Contributors
Burgos, J. J.
Facultad de Agronomia y Veterinaria, Universidad de Buenos Aires, Buenos Aires,
Argentina
Calderon, Cleofe E.
Department of Botany, Smithonian Institution, Washington D.C. 20560, USA
Coaldrake, J. E.
CSIRO, Director A. A. Heath & Partners, 131 Leichhardt Street, Brisbane, Queens
land, Australia 4000
Coupland, R. T.
Department of Plant Ecology, University of Saskatchewan, Saskatoon, Canada
S7N OWO
Hayashi, Ichiroku
Kogen Experiment Centre, Tsukuba University, Sugadaira, Sanada-machi, Nagano
Pref., Japan
Innis, George S.
Department of Wildlife Science, Utah, State University Logan, Utah 84322, USA
Iwaki, Hideo
Biological Sciences, Tsukuba University, Sakura-mura, Ibaraki Pref., Japan
Kawanabe, Sukeo
Tropical Agricultural Center, Yatabe-machi, Ibaraki Pref., Japan
Knapp, Rii.diger
Botanical Institute, University Giellen, Federal Republic of Germany, Senckenberg
Str.17-25
Numata, Makoto
Labor. of Ecol., Fac. Sci., Chiba University, Yayoi-cho, Chiba, Japan
Papadakis, J.
Cordoba 4564, Buenos Aires, Argentina
Sochava, Victort
Institute of Geography of Siberia and the Far East, Academy of Sciences of USSR,
Siberian Department, Kievskaya 1, Irkoutsk-3, USSR
Soderstrom, Thomas S.
Department of Botany, Smithonian Institution, Washington D.C. 20560, USA
Soriano, A.
Facultad de Agronomia y Veterinaria, Univ. de Buenos Aires, Av. San Martin 4453,
Buenos Aires, Argentina
Valencia, Rafael F. J.
Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Buenos
Aires, Argentina
Wyk van, J. J. P.
Dept. of Biology, Eastern Montana College, Billings, Montana 59101, USA
Preface
In 1967 and again in 1970 and 1977 I consulted in Tokyo with Mrs. Schluter of VEB
Gustav Fischer Verlag, Jena, about the plan for a new book on the ecology of the
world's grasslands and bamboolands. Upon being requested to be the editor of this
new publication, two important books on the subject immediately came to mind,
a famous classic by J. W. Bews on "The World's Grasses" (1929) and a new book
"Grasses and Grasslands" edited by C. Barnard (1964). At the same time, the Inter
national Biological Program had begun, and I became responsible for the Japanese
Grassland Project under the sponsorship of the Ministry of Education. I personally
conducted the field study and synthesis on Japanese semi-natural grasslands the
results of which were published as "Ecological studies in Japanese Grasslands with
Special Reference to the IBP Area-Productivity of Terrestrial Communities" (JIBP
Synthesis Vol. 13, University of Tokyo Press, 1975). During this period, two books
on grassland ecology were published in Japanese, and International Synthesis Volumes
are in preparation for publication by the Cambridge University Press, including
two volumes on grasslands. In relation to grassland productivity studies, I attended
the Second Meeting of the PT Grassland Working Group of IBP held at Saskatoon
and Matador, Canada, in 1969 (the proceedings entitled "Grassland Ecosystems·
Reviews of Research" edited by R. T. Coupland and G. M. Van Dyne, 1970), and a
synthesis meeting at Fort Collins, Colorado, U.S.A. in 1972.
The plan of our new book was set at the beginning of 1971, and the publisher wrote
to the authors of each chapter requesting that they begin to prepare their manu
scripts. However, during the course of the work, there were some changes in the
authors from those originally planned. Dr. F. A. McClure of the Smithonian Institu
tion unfortunately passed away and Dr. T. R. Soderstrom agreed to write a chapter
on bamboos. It was difficult to find a suitable author to write on European grasslands,
however Prof. R. Knapp eventually accepted this role. A chapter on simulation
models of grasslands was added later.
At long last, this book has been published. In it, grasslands are noted as seral
stages in moist areas and as climax in dry areas. Some of them are left as they are
and not utilized while at the same time, many are used for grazing, mowing, etc.
Such grasslands are an object of study in animal ecology and ecosystems, however
the contents of this book are treated mainly from the standpoint of plant ecology
to avoid overlapping the work of the IBP. Bamboos as giant grasses are included
as a characteristic of the book. Though some tundra vegetations resemble grasslands,
tundra is not included in this book. There are numerous books on grasses and grass
lands written from the standpoint of agriculture, but very few are concerned with
ecological aspects. This book is aimed at an orthodox plant ecological view of the
grasslands and bamboolands of the world.
Chiba/Japan Makoto Numata
Contents
Preface .... 7
I. Introduction 11
II. Climate and Soil
1. Climate and Soils in the Grassland Areas of North America.
By R. T. Coupland . . . . . . . . . . . . . . . . . 15
2. Climate and Soils of Pasture Regions in South America.
By J. J. Burgos and R. F. J. Valencia. . . 20
3. Climate and Soils in Asian Grassland Areas.
By M. Numata. . . . . . . . . . . . . 35
4. Climate and Soils of the Grassland Distribution Area in the USSR.
By V. Sochav a. . . . . . . . . . . . ... . 43
5. Climate and Soils of Grassland Areas in Europe.
By R. Knapp . . . . . . . . . . . . . . . 49
6. Climates and Soils of Grassland Areas in Africa.
By J. Papadakis . . . . . . . . . . . . . . 57
7. Climates and Soils of Australasia in Relation to Grasslands.
By J. E. Coaldrake . . . . . . . . . . . . . . . . . . 69
ID. Grasses and Grasslands
1. Distribution of Grasses and Grasslands of North America.
By R. T. Coupland . . . . . . . . . . . . . . . . . 77
2. Distribution of Grasses and Grasslands of South America.
By R. Soriano . . . . . . . . . . . . . . . 84
3. Distribution of Grasses and Grasslands in Asia.
By M. Numata. . . . . . . . . . . . 92
4. Distribution of Grasslands in the USSR.
By V. Sochava . . . . . . . . . . . . 103
5. Distribution of Grasses and Grasslands in Europe.
By R. Knapp .............. . III
6. A General Account of the Grass Cover of Mrica.
By J. J. P. van Wyk . . . . . . . . 124
7. The Natural Grassland!' of Australasia.
By J. E. Coaldrake . . . . . . . . . 133
IV. Ecology, Phytosociology and Productivity of Grasses and Grasslands
1. The Autecology of Some Grassland Species.
By I. Hayashi . . . . . . . . . . . . . . . . . . . . . . 141
10 Contents
2. The Pattern of Temperature Response and Its Ecological Significance.
By S. Ka"\>anabe . . . . . . . . . . . . . . . . . . 153
3. Phytosociological Classification of Grassland Vegetation.
By R. Knapp . . . . . . . . . . . . . . . . . . 163
4. The Structure and Succession of Grassland Vegetation.
By M. Numata. . . . . . 182
5. Productivity of Grasslands.
By H. Iwaki. . . . . . . 197
6. Simulation Models of Grasslands and Grazing Lands.
By G. S. Innis . . . . . . . . . . . . . . . 211
V. Ecology and Phytosociology of Bamboo Vegetation
1. Distribution and Environment of the Bambusoideae.
By Th. R. Soderstrom and C1. E. Calderon. . . 223
2. Structure and Succession of Bamboo Vegetation.
By M. Numata. . . . . . . . . . . . . . . 237
3. The Relationship of Limiting Factors to the Distribution and Growth
of Bamboo.
By M. Numata . . . . . . . . . . . . . . . . . . . . . . . . . 258
Subject Index 277
Taxonomic Index 285
I. Introduction
M. Numata
There are many publications on management, use, improvement, maintenance, etc.
of pastures and meadows as the sites of animal husbandry. Grassland is a kind of
plant formation, biome, or ecosystem from the viewpoint of physiognomy, including
climax grasslands under grassland climate and seral grasslands under forest climate.
Grasslands in a wide sense include grasslands in a narrow sense, herblands, forblands,
savannas, etc. (Penfound, 1967). Such grasslands in a wide sense are classified into
many subdivisions from various standpoints as follows (Numata, 1969):
1) Grasslands as a botanical term
1.1) Local names: prairie, plain, steppe, pampa, campo, llano, savanna, veld, etc.
1.2) Referring to moisture conditions: mesophytic-, hygrophytic-, aquatic-, xerophytic grasslands,
etc.
1.3) Referring to salinity: fresh water-, salt water-, brackish water-(estuary-), coastal grasslands, etc.
1.4) Referring to substrata: loamy soil-, sandy soil-, gravelly soil-, peat grasslands, etc. including
rocky desert, coastal desert, etc.
1.5) Referring to plant succession: pioneer-, seral-, and climax grasslands.
1.6) Referring to physiognomy and floristic composition: grassland in a narrow sense, forbland,
herbland, Staudenfluren, bambooland, dwarf bambooland, sedgel and, rushland, vineland,
savanna, etc.
1.7) Referring to duration of life: annual-, perennial-, deciduous-, evergreen grasslands, etc.
1.8) Referring to height: high grass-(tall grass-), medium-height grass-(mid grass-), and short grass
grasslands
1.9) Referring to altitude and geomorphology: lowland-, mountain-, highland-, alpine-, summit-,
leeward-, windward grasslands, snow-patch carpet, alpine mat, etc.
2) Grasslands as an agricultural term
2.1) Administrative term: artificial-, natural-(wild-), improved grasslands, etc.
2.2) Referring to existence of biotic factors: natural-, semi-natural, and sown grasslands
2.3) Referring to a kind of biotic factor: meadow (mown grassland), pasture (grazed grassland),
abandoned field (old field), grasslands for thatching, etc.
2.4) Referring to degree of management: extensive-(range), improved-, intensive grasslands, etc.
2.5) Referring to soil fertility: fertile-, sterile-, barren grasslands, etc.
According to Ellenberg and Mueller-Dombois (1967), terrestrial (mainly meso
phytic) herbaceous communities are classified as follows:
A. Savannas and related grasslands
1. Mainly flat-leaved savannas
a. Woodland savanna
b. Tree savanna
c. Scrub savanna
d. Grass savanna
e_ Flood savanna
12 Introduction
2. Mainly narrow-leaved savannas
a. Tree savanna
b. Scrub savanna
c. Grass savanna
B. Steppes· and related grasslands
1. Tall grass steppes
a. Tall grass steppe with trees
b. Tall grass steppe with shrubs
c. Tall grass steppe without woody plants
2. Mid grass steppes
Subdivisions as in 1.
3. Short grass steppes
Subdivisions as in 1; furthermore,
d. Short grass steppe with suffrutescent plants
4. Forb-rich steppes
Subdivisions as in 1.
C. Meadows and pastures or related grasslands
1. Meadows and pastures below tree line
a.
Woodland pasture
b. Tree meadow (or pasture)
c. Scrub pasture (or meadow)
d. Grassy pasture without trees or shrubs
(1) Extensively grazed (Triftweide, not fertilized)
(2) Intensively grazed (Standweide or Umtriebsweide, fertilized)
e. Grassy meadow without trees or shrubs
(1) Litter meadow (Streuwiese, generally not fertilized)
(2) Hay meadow (Futterwiese, generally fertilized)
f. Sedge-rush meadow
g. Avalanche grassland
2. Pastures and meadows above mountain tree line
a. Closed alpine (or subpolar) mat
b. Alpine (or subnival) mat-patches
c. Snow bed formation
D. Forb vegetation
1. Mainly perennial forb communities
8. Forest margin herb formation
b. Tall forb (Hochstauden) formation
c. Bracken thicket (or heath)
d. Perennial forb formation on organic deposits at the flood lines
e. Perennial ruderal and clearing herb formation
f. Mcinly perennial weed formation on cultivated land
2. Mainly ephemeral forb communities
a. Tropical or subtropical ephemeral cloud desert formation
b. Ephemeral halophytic formation
c. Ephemeral ruderal and clearing forb formation
d. Mainly ephemeral weed formation on cultivated land
3. Episodical forb communities
&. Episodical desert forb formation (flowering desert)
b. Episodical formation on pond muds and similar sites
c. Episodical forb formation on organic deposits at the flood lines
d. Episodical river bed formation
Introduction 13
The term "grassland" is a general name of plant formation as mentioned above,
but "pasture" and "meadow" are special types of grassland used for grazing and
mowing.
In this book, grassland in general, or pasture and meadow will be discussed from
the ecological viewpoint. Such a direction has been found only in Bews (1929), Ellen
berg (1952), Barnard (1964), etc. We shall try a new approach to the worldwide
grasslands.
Moreover, the bamboolands are added to the grasslands in this book. Bamboos
are giant grasses distributed in limited areas in tropical and temperate regions. Dwarf
bamboos (sasa in Japanese) are included in the grassland in a narrow sense, but only
tall bamboos (take in Japanese) are treated in the chapters of bamboolands.
The editor believes that this is the first book on grasslands and bamboolands from
a truely ecological point of view, and acts as a textbook of grassland ecology.
References
Barnard, C. (Ed.) 1964. Grasses and Grasslands. London.
Bews, J. W. 1929. The World's Grasses. Their Differentiation, Distribution, Economics and Ecology.
London.
Ellenberg, H. 1952. Wiesen und Weiden und ihre standiirtliche Bewertung. Stuttgart.
-, and D. Mueller·Dombois. 1967. Tentative physiognomic·ecological classification of plant forma
tions of the earth. Ber. geobot. Inst. ETH, Stiftg. Rubel, ZUrich 37: 21-55.
Numata, M. 1969. A proposal to the study on productivity and conservation of grassland ecosystems.
Chiba.
- (Ed.) 1968. Studies on Productivity and Conservation of Grassland Ecosystems. Chiba.
Penfouncl, W. T. 1967. A physiognomic classification of vegetation in conterminous United States.
Bot. Rev. 33: 259-326.
