Mediterranean-Type Ecosystems: The Role of Nutrients

565 Pages·1983·27.289 MB·English

by  H. J. Deacon (auth.), Frederick J. Kruger, Professor Derek T. Mitchell, Professor Jennifer U. M. Jarvis (eds.)

Ecological Studies
Analysis and Synthesis
Edited by
W. D. Billings, Durham (USA) F. Golley, Athens (USA)
O. L. Lange, Wiirzburg (FRG) J. S. Olson, Oak Ridge (USA)
H. Remmert, Marburg (FRG)
Volume 43
Mediterranean-Type
Ecosystems
The Role of Nutrients
Edited by
F. 1. Kruger D. T. Mitchell 1. U. M. Jarvis
With 143 Figures
Springer-Verlag
Berlin Heidelberg New York Tokyo 1983
FREDERICK J. KRUGER
South African Forestry Research Institute
Department of Environment Affairs and Fisheries
Pretoria 0001, South Africa
Professor DEREK T. MITCHELL
Department of Botany
University of Cape Town
Rondebosch, South Africa
Professor JENNIFER U. M. JARVIS
Department of Zoology
University of Cape Town
Rondebosch, South Africa
ISBN-13: 978-3-642-68937-6 e-ISBN-13: 978-3-642-68935-2
DOl: 10.1007/978-3-642-68935-2
Library of Congress Cataloging in Publication Data. Main entry under title: Mediterranean-type ecosystems.
(Ecological studies; v. 43). Includes part of the proceedings of a symposium held at the University of Stellenbosch,
South Africa, Sept. 22-23, 1980. Bibliography: p. Includes indexes. I. Biogeochemical cycles - Congresses.
2. Mediterranean climate - Congresses. 3. Ecology - Congresses. I. Kruger, F. J. (Frederick J.), 1944-. II. Mitchell,
D. T. (Derek T.), 1942-. III. Jarvis, J. U. M. (Jennifer U. M.), 1939 -. IV. Series. QH344.M43. 1983. 574.5'222.
83-363.
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© by Springer-Verlag Berlin' Heidelberg 1983
Softcover reprint of the hardcover I st edition 1983
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Printing and binding: Briihlsche Universitiitsdruckerei, Giessen
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Preface
The theory of ecological convergence underlies the biogeographers' maps of
world biome-types. It also determines the degree to which ecological principles,
derived from research on particular populations, communities or ecosystems, are
generally valid, and hence also to what extent resource management principles are
general. To quote Di Castri and Mooney (1973): "In effect, in order to assess the
transfer of technology, it is essential to know to what extent information acquired
from studying one particular ecosystem is applicable to another ecosystem of the
same type but situated in a different location."
The five relatively small, isolated, mediterranean-climate zones of the earth,
each with its distinct fauna and flora, have provided the ideal testing grounds for
this theory. A heritage of precisely focused ecosystems research has resulted,
beginning with the international comparative analyses conducted by Specht
(l969a, b) but with antecedents in earlier studies in South Australia (Specht and
Rayson 1957, Specht 1973). Cody and Mooney (1978) reviewed the information
available at the time for the four zones excepting Australia and concluded that the
arrays of strategy-types to be found among the different biotas were so similar that
they could be explained only in terms of the convergence hypothesis; nevertheless,
evident differences in community organization and dynamics, especially phenol
ogy, required closer study of resource availability and resource-use patterns to
better explain relations between form and function overall, and to assess the degree
of convergence at higher levels of organization than the population.
Furthermore, the biogeographers' concept of a mediterranean-climate biome
type, established by Grisebach (1872) and Schimper (1903), has been challenged by
Specht (1979) and found untenable on the grounds that major sclerophyllous
shrublands, the Australian heathlands and South African fynbos, are determined
by nutrients, not by climate. Both of these vegetation formations occur within
mediterranean-climate zones but extend well beyond the boundaries of the
mediterranean-type climates as normally defined and in Australia occur extensively
within the tropics; their distribution correlates rather with soils exceptionally
poor in nutrients. Specht proposed that the floras of this vegetation had ancient
origins and evolved in edaphically determined oligotrophic habitats in the humid
tropical to sub-tropical environments of the Cretaceous and early Tertiary: the
physiological drought of the intermittently waterlogged soils preadapted these
plants to the seasonally arid climates that arose in the Tertiary, and the later
inception of mediterranean climates.
This book incorporates part of the proceedings of a conference called to assess
the role of nutrients in species and ecosystem convergence in mediterranean-
VI Preface
climate ecosystems. Specifically, the first aim was to draw the contrast between the
nutrient-poor shrublands of Australia and South Africa and the physiognomically
similar vegetations of richer soils. This comparison was to indicate how the
environmental contrasts are expressed in the adaptive responses of the biota as well
as in key ecosystem processes. The second question addressed, had two parts: why
is it that the biotas of these regions, especially the plants, are unusually rich in
species, and how is this richness maintained in resource-poor environments?
The symposium was held at the University of Stellenbosch, South Africa, from
22 to 23 September 1980 and was followed by a four-day workshop at Hermanus. It
was the third in a series of international conferences, beginning with the symposium
at the Universidad Austral de Chile, Valdivia, in March 1971 (see Di Castri and
Mooney 1973), and followed some time later by the meeting at Stanford University
in 1977 (Mooney and Conrad 1977). This series began with a critical overview of
information relevant to studies of convergence, with the emphasis essentially on the
Mediterranean Basin and the New World. The Stanford meeting focused on
applied ecological questions, principally the role of fire in resource management.
The Stellenbosch meeting saw a return to the question of convergence, though in
a wider context. These trends reflect the vigorous and encouraging interplay
between fundamental and applied research that has arisen in these zones.
These conferences have to a large degree been part of a continuous process of
development of integrated ecological research programmes involving comparative
studies of mediterranean-type ecosystems recently reviewed by Mooney (1981 a, b).
The papers from the Stellenbosch symposium included here are of two kinds:
invited review papers, and contributed papers that met the requirements of the
editorial panel; all papers considered for publication were subject to peer review
and revised accordingly, some extensively. The workshop proceedings are to be
published separately (Day 1983). Although at the outset the intention was to
provide a review of all major topics relevant to the conference theme, this proved
too ambitious. We found that certain topics were easily covered, and that
Australian heathlands and South African fynbos in particular seem inevitably to
get the greater attention, perhaps justly so. We therefore do not claim that this
book is comprehensive; nevertheless it introduces a new dimension and balance to
the subject and raises many novel questions. The international studies of vegetation
function in the mediterranean-type ecosystems of Chile and California (Mooney
1977, Miller 1981) provided the first truly synthetic comparative accounts of
ecosystem function. The Stellenbosch meeting proved a major step toward further
synthesis for several continents and these proceedings will hopefully help to
maintain the momentum for intercontinental comparative studies.
The text of 27 chapters is organized in six sections. The first, on the evolution
and characters of mediterranean-type ecosystems, proceeds from a review of the
evolutionary history of the systems, through major features of vegetation
environment patterns, to the question of detecting patterns in the geographic
distribution of plant forms. The second, on plant form and function, is aimed at
assessing our understanding of how environment is functionally linked to plant
forms. In the third, nutrient cycling is considered at the ecosystem level, while
plant nutrition is considered in the fourth. The fifth section contains eight chapters
that deal with community pattern and diversity, specifically how communities are
Preface VII
structured in these resource-poor environments, and how their dynamics con
tribute to maintenance of richness. The final section, on plant-animal interactions,
has two papers that concentrate on herbivory.
The symposium and workshop from which this volume proceeds was convened
by the Fynbos Biome Project, one of several large interdisciplinary cooperative
research projects of the National Programme for Environmental Sciences. The
support of the Council for Scientific and Industrial Research and the Department
of Environment Affairs and Fisheries is gratefully acknowledged.
Credit is owing to many for assistance in seeing this volume to the press. We
would especially like to acknowledge Suzanne Winkelman, then of the CSIR, for
assistance in editing copy during the early and crucial phases of the work. Minja
Martin contributed unstintingly in the latter stages. Margaret Orton undertook the
entire typography. For the support of these and other members of the section for
Cooperative Scientific Programmes of the Council for Scientific and Industrial
Research we are extremely grateful.
Pretoria and Cape Town, April 1983 FRED KRUGER
DEREK MITCHELL
JENNY JARVIS
References
Cody ML, Mooney HA (1978) Convergence versus nonconvergence in mediterranean
climate ecosystems. Annual Review of Ecology and Systematics 9:265-321
Day J (ed) (1983) Mineral nutrients in mediterranean ecosystems. Results of a work
shop in Hermanus, South Africa, September 1980 (in press)
Di Castri F, Mooney HA (eds) (1973) Mediterranean-type ecosystems: origin and structure.
Springer, Berlin Heidelberg New York, 405 pp
Grisebach AHR (1872) Die Vegetation der Erde nach ihrer klimatischen Anordnung.
W Engelmann, Leipzig
Miller PC (ed) (1981) Resource use by chaparral and matorral: a comparison of vegetation
function in two mediterranean-type ecosystems. Springer, Berlin Heidelberg New York,
455pp
Mooney HA (ed) (1977) Convergent evolution in Chile and California mediterranean
climate ecosystems. Dowden, Hutchinson and Ross, Stroudsberg, Pennsylvania, 224pp
Mooney HA (1982) Mediterranean-type ecosystems - research progress and oppurtunities.
South African Journal of Science 78:5-7
Mooney HA, Conrad CE (technical coordinators) (1977) Proceedings of the symposium
on the environmental consequences of fire and fuel management in mediterranean
ecosystems. USDA Forest Service General Technical Report WO-3, 498 pp
Schimper AFW (1903) Plant-geography upon a physiological basis. Clarendon Press,
Oxford, 824pp
Specht RL (1969a) A comparison of sclerophyllous vegetation characteristic of mediter
ranean-type climates in France, California and southern Australia. I. Structure,
morphology and succession. Australian Journal of Botany 17:277-292
Specht RL (1969b) A comparison of the sclerophyllous vegetation characteristic of
mediterranean-type climates in France, California, and southern Australia. II. Dry
matter, energy, and nutrient accumulation. Australian Journal of Botany 17:293-308
VIII Preface
Specht RL (1973) Structure and functional response of ecosystems in the mediterrranean
climate of Australia. In: Di Castri F, Mooney HA (eds) Mediterranean-type ecosystems:
origin and structure. Springer-Verlag, Berlin, pp 113-120
Specht RL (1979) Heathlands and related shrublands of the world. In: Specht RL (ed)
Ecosystems of the world, vol 9B. Heathlands and related shrublands. Part A:
Descriptive studies. Elsevier, Amsterdam, pp 1-18
Specht RL, Rayson P (1957) Dark Island heath (Ninety-Mile Plain, South Australia).
I. Definition of the ecosystem. Australian Journal of Botany 5:52-85
Contents
, Section I. Evolution and Character of Mediterranean-Type Ecosystems
1. The Comparative Evolution of Mediterranean-Type Ecosystems:
A Southern Perspective. H. J. DEACON . . . . . . . . . . . . . 3
2. Mediterranean-Type Heathlands and Sclerophyllous Shrublands of the
World: An Overview. R.L. SPECHT and E.J.MoLL . . . . . . 41
3. Ecological Control of the Vegetation of Southwestern Australia:
Moisture versus Nutrients. J. S. BEARD. . . . . . . . . . . . 66
4. Influence of Nutrients on the Composition and Distribution of Plant
Communities in Mediterranean-Type Ecosystems of Israel
A. RABINOVITCH-VIN . . . . . . . . . . . . . . . . . . . 74
5. Approaches to the Definition of Mediterranean Growth Forms
G.ORSHAN . . . . . . . . . . . . . . . . . . . . . . . 86
Section II. Plant Form and Function
6. Carbon-Gaining Capacity and Allocation Patterns of Mediterranean
Climate Plants. H. A. MOONEY . . . . . . . . . . . . . . . . . . 103
7. Moisture Regime and Nutrient Control of Seasonal Growth in
Mediterranean Ecosystems. R. L. SPECHT, E. J. MOLL, F. PRESSINGER, and
J. SOMMERVILLE. . . . . . . . . . . . . . . . . . . . . . . . . 120
8. Canopy Structure of Mediterranean-Type Shrubs in Relation to Heat
and Moisture. P. C. MILLER . . . . . . . . . . . . . . . . . . . 133
9. Carbon and Nitrogen Economy of Diplacus aurantiacus, a Californian
Mediterranean-Climate Drought-Deciduous Shrub. S. L. GULMON. . . 167
Section III. Nutrient Cycling
10. Nutrient Cycling in Australian Heath and South African Fynbos
R. H. GROVES . . . . . . . . . . . . . . . . . . . . . . . . . 179
11. Impact of Fire on Nutrient Cycles in Mediterranean-Type Ecosystems
with Reference to Chaparral. P. W. RUNDEL . . . . . . . . . . . . 192
12. Decomposition and Mineralization Processes in Mediterranean-Type
Ecosystems and in Heathlands of Similar Structure. D. J. READ and
D. T.MITCHELL . . . . . . . . . . . . . . . . . . . .. .208
13. The Effects of Different Fire Regimes on Soil Nutrient Levels in
Quercus coccifera Garrigue. L. TRABAUD . . . . . . . . . .. . 233
x Contents
Section IV. Plant Nutrition
14. Strategies for Maximizing Nutrient Uptake in Two Mediterranean
Ecosystems of Low Nutrient Status. B. B. LAMONT . . . . . . 246
15. The Effects of Fire on Photosynthesis in Chaparral Resprouts
w. C. OECHEL and S. J. HASTINGS. . . . . . . . . . . . . .274
16. Mineral Nutrient and Nonstructural Carbon Pools in Shrubs from
Mediterranean-Type Ecosystems of California and Chile. G. R. SHAVER 286
17. Comparative Phenology of Mediterranean-Type Plant Communities
J. KUMMEROW .. . .. ... . . . . . ... . . . . . . ... 300
Section V. Community Patterns and Diversity
18. Some Thoughts on Resource Competition and Diversity in Plant
Communities. D. TILMAN . . . . . . . . . . . . . . . . . . . . 322
19. On Alpha Diversity and the Richness of the Cape Flora: A Study in
Southern Cape Fynbos. W. BOND . . . . . . . . . . . . . .. . 337
20. Continental Diversity Patterns and Convergent Evolution in Bird
Communities. M. L. CODY . . . . . . . . . . . . . . . .. .357
21. Distribution and Species Diversity of Birds and Plants in Fynbos
Vegetation of Mediterranean-Climate Zone, South Africa
W. R. SIEGFRIED and T. M. CROWE . . . . . . . . . . . . . . 403
22. Plant Community Structure - Spatial Partitioning of Resources
W. E. WESTMAN . . . . . . . . . . . . . . . . . . . . .417
23. Plant Community Diversity and Dynamics in Relation to Fire
F. J. KRUGER. . . . . . . . . . . . . . . . . . . . . . .446
24. Mammal Species Diversity in Australian Heathlands: the Importance
of Pyric Succession and Habitat Diversity. B. J. Fox. . . . .. . 473
25. Animal Demography in Relation to Fire and Shortage of Food:
Some Indicative Models. A. E. NEWSOME and P. C. CATLING .. .490
Section VI. Plant-Animal Interactions
26. The Role of Sclerophyllous Leaves in Determining Insect Grazing
Damage. P. A. MORROW . . . . . . . . . . . . . . . . . . .509
27. Defoliation Patterns in Matorral Ecosystems. E. R. FUENTES and
J. ETCHEGARA Y. . . . . . . . . . . . . . . . . . . . .. . 525
SUbject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . 543
Contributors
BEARD, JOHN S. 6 Fraser Road, Applecross, Western Australia 6153,
Australia
BOND, WILLIAM Saasveld Forestry Research Station, Department of
Environment Affairs, Private Bag X6531, George 6530,
South Africa
CATLING, PETER C. CSIRO Division of Wildlife Research, P.O. Box 84,
Lyneham, ACT 2602, Australia
CODY, MARTIN L. Department of Biology, University of California, Los
Angeles, CA 90024, USA
CROWE, TIM M. Percy FitzPatrick Institute, University of Cape Town,
Private Bag, Rondebosch 7700, South Africa
DEACON, HILARY J. Department of Archaeology, University of Stellen
bosch, Stellenbosch 7600, South Africa
ETCHEGARAY, JULIA Instituto de Ciencias Biologicas, Pontificia Universidad
Catolica de Chile, Casilla 114-D, Santiago, Chile
Fox, BARRY J. School of Zoology, University of New South Wales,
P.O. Box I, Kensington 2033, Australia
FUENTES, EDUARDO R. Instituto de Ciencias Biologicas, Pontificia Universidad
Catolica de Chile, Casilla 114-D, Santiago, Chile
GROVES, RICHARD H. CSIRO Division of Plant Industry, P.O. Box 1600,
Canberra City, ACT 2601, Australia
GULMON, SHERRY L. Department of Biological Sciences, Stanford University,
Stanford, CA 94305, USA
HASTINGS, STEVEN J. Systems Ecology Research Group, San Diego State
University, San Diego, CA 92182, USA
KRUGER, FREDERICK J. South African Forestry Research Institute, P.O. Box
727, Pretoria 0001
KUMMEROW, JOCHEN Department of Botany, College of Sciences, San Diego
State University, San Diego, CA 92182, USA