Fynbos : ecology, evolution, and conservation of a megadiverse region

413 Pages·2014·33.85 MB·English

by  Allsopp, Nicky, Colville, Jonathan F., Verboom, G. Anthony

Fynbos

Fynbos
Ecology, Evolution,
and Conservation
of a Megadiverse Region
EDITED BY
Nicky Allsopp
South African Environmental Observation Network (SAEON),
Fynbos Node, South  Africa
Jonathan F. Colville
Kirstenbosch Research Centre
South African National Biodiversity Institute, South Africa
G. Anthony Verboom
Bolus Herbarium and Department of Biological Sciences
University of Cape Town, South Africa
CONSULTING EDITOR
Richard M. Cowling
Department of Botany
Nelson Mandela Metropolitan University, South Africa
1
Fynbos: Ecology, Evolution, and Conservation of a Megadiverse Region. Edited by Nicky Allsopp,
Jonathan F. Colville and G. Anthony Verboom. © Oxford University Press 2014.
Published 2014 by Oxford University Press.
3
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Preface
We have used the term Fynbos in the title of this book. the evolutionary sphere has perhaps been most promi-
But what is fynbos? In a strict sense, fynbos refers to a nent, significant advances have also been made in the
narrowly defined vegetation type found in the e xtreme fields of conservation science, pollination biology,
southwestern corner of Africa, the Cape Floristic Region invasive plant biology, palaeoecology, and palaeoan-
(CFR). The term also has broader meaning, however, thropology (modern human behaviour evolved along
being used colloquially to refer to any fine-leaved vege- the Cape coast some 160 000 years ago). Moreover, a
tation found in this region, and even being used by non- growing recognition of the physiographic and floristic
biologists to refer more loosely to the sun-drenched, affinities of the CFR with the neighbouring succulent
mediterranean-type environments of the Cape region. karoo region has led many workers to broaden the
Although the material presented in the present geographical scope of their thinking to embrace the
volume addresses a diversity of vegetation types (e.g. entire GCFR.
renosterveld, subtropical thicket, succulent karoo), Lessons from the Cape region (CFR and GCFR)
besides fynbos, which occur in the Greater Cape Flo- offer much of value for understanding the origin,
ristic Region (GCFR; encompassing the CFR and the maintenance, and conservation of biological diversity
adjacent succulent karoo region), we have entitled it in general. Besides offering an independent view on
Fynbos for two reasons. First, most informed biologists megadiversity from that provided by tropical ecosys-
are familiar with the term and immediately associate it tems, the region offers valuable alternative perspec-
with the extremely species- and endemic-rich vegeta- tives for understanding and testing ecological theory,
tion which forms the backbone of the renowned Cape much of which has been developed in the context of
flora. Second, while a title referring to the GCFR might the Northern Hemisphere systems which first emerged
have been technically more appropriate, we felt that in the wake of receding glacial ice sheets 12 000 years
such a title would be cumbersome and consequently ago. The Cape r egion presents a contrary perspective,
lack punch. Besides, much of the material p resented both on account of the greater importance of fire and
does focus on fynbos vegetation, making this an appro- nutrients as ecological drivers, and on account of its
priate point of convergence and departure throughout deep evolutionary history, undoubtedly a product of
the book. Overall, this book represents an interdis- its greater environmental stability at least since the
ciplinary account of current knowledge relating to Early Pliocene. Such contrasts to accepted theory offer
the ecology, evolution, and management of biologi- the grist to challenge and refine the models we use to
cal diversity in the GCFR, although some themes are understand the natural world and how it will respond
addressed more broadly than others, both with regard to global change.
to geographical scope (e.g. GCFR versus CFR) and the The purpose of the present volume is to document
range of vegetation types addressed. Fynbos, however, advances in our understanding of the biodiversity,
occupies centre stage throughout. evolution and ecology of the Cape region since the
The CFR has intrigued biologists for centuries, last major synthesis of the topic, The ecology of fynbos:
achieving iconic status as a locus of megadiversity and nutrients, fire and diversity,1 was published more than
a place to study the ecological underpinnings of mas- two decades ago. We can think of at least two compel-
sive evolutionary radiations. Scholars, both local and ling justifications for a new synthesis and interpreta-
from overseas, have made great advances, over the tion: first, to capture the insights of a new generation
past two decades, in unravelling the ecological and
evolutionary complexities of the CFR, especially that 1 Cowling, R.M. (1992). The ecology of fynbos: nutrients, fire
of the species-rich fynbos vegetation. While progress in and diversity. Oxford University Press, Cape Town.
v
vi PREfa CE
of biologists, and, second, to incorporate the large chapters. First and foremost, the editors wish to thank
body of knowledge that has accumulated in the two Richard Cowling for providing the inspiration behind
decades since the publication of the first volume. With this book. We also thank Richard for providing ongoing
regard to the latter, huge developments in comput- advice on editorial matters and for his willingness to
ing power and analytical methodologies, as well as serve as a ‘third reviewer’ on several chapter drafts. We
the rapid availability of DNA sequence data for many are also extremely grateful to Michelle Bradshaw, who
Cape plant lineages and spatially comprehensive digi- performed an outstanding job as copy editor, bring-
tal plant and animal datasets, have made it possible ing a rigour to this task which we as editors would
to implement analyses that the authors of the first not have been able to do. We also owe an enormous
book could only dream about. In addition, it is hoped vote of thanks to a large corps of individuals, some
that the present synthesis will facilitate the urgent anonymous, who generously gave their time to review
development of management policies and guidelines either the initial book proposal or the component chap-
geared towards addressing the biodiversity challeng- ters: Sally Archibald, Nigel Barker, Mark Botha, John
es posed by a ccelerating global change. The present Compton, Richard Cowling, Helen de Klerk, Barend
book is aimed at an international audience of schol- Erasmus, Karen Esler, Pat Holmes, Peter Goldblatt,
ars interested in the understanding of evolution and Gavin Gouws, Susan Harrison, Andrew Latimer, David
ecology of the GCFR biota. As such, our objective has Le Maitre, John Manning, Michael Meadows, Eugene
been to produce a book that is comprehensible by the Moll, Fernando Ojeda, Anibal Pauchard, Craig Peter,
informed, but not necessarily specialist, reader. Tony Rebelo, David Richardson, Robert Ricklefs, Brad
The chapters included in this book fall into four broad Ripley, Philip Rundel, Erik Veneklaas, Sandi Willows-
themes. The first of these, represented by C hapters 1–4, Munro, Sarah Wurtz, Colin Yates. Finally, we thank the
relates to environmental context, comprising a ccounts following organizations for financial support, C.A.P.E.
of the vegetation of the GCFR (Chapter 1), the historical and the South African National Biodiversity Institute,
and contemporary physical environment ( Chapter 2), and our families, colleagues and institutions for their
and variation in fire regime and fire management (Chap- support during the writing and editing of this book.
ter 3). Also included in this section is an evaluation of Throughout this book we have sought to standard-
the general validity of the GCFR as a biogeographic ize terminology relating to the description of veg-
unit (Chapter 4). The second theme, represented by etation. Where left uncapitalized, the terms ‘fynbos’,
Chapters 5–8, relates to the historical development of ‘renosterveld’, ‘succulent karoo’, ‘forest’, and ‘thicket’
the Cape biota, with chapters exploring the Cenozoic (including ‘strandveld’) are used to refer to the prin-
assembly of the flora (Chapter 5), diversification pro- cipal vegetation types as found within the GCFR
cess (Chapter 6), phylogeographic pattern and its deter- (Fig 1.1). Similarly, ‘desert’, ‘Nama karoo’, ‘grassland’,
minants (Chapter 7), and the palaeoecological context and ‘savanna’ are used to refer to vegetation types of
of the emergence of modern humans in the r egion adjacent regions outside of the GCFR. Where capital-
(Chapter 8). A third theme, presented in Chapters ized, however, the terms ‘Fynbos’, ‘Forest’, ‘Albany
9–11, is centred on ecological process, specifically that Thicket’, ‘Succulent Karoo’, ‘Nama-Karoo’, ‘Grassland’,
underpinning community structure and richness. Top- ‘Desert’, and ‘Savanna’ refer explicitly to the formal
ics covered here include the assembly and functioning biomes, as defined by Mucina and Rutherford in The
of plant communities (Chapter 9), biotic i nteractions vegetation of South Africa, Lesotho and Swaziland. In some
(Chapter 10), and the influence of plant form and func- instances (e.g. grassland), traditional vegetation types
tion (Chapter 11). The fourth major theme, represented and b iomes are approximately equivalent, but in others
by Chapters 12–15, relates to h uman i mpacts and their they are not. Most relevant, in the context of this book, is
management, the major issues a ddressed here b eing the inclusion of renosterveld and thicket vegetation, in
biological invasions (Chapter 12), climate change addition to fynbos, within the Fynbos Biome. This ren-
(Chapter 13), c onservation (Chapter 14), and people ders the terms fynbos and Fynbos non-equivalent. In
and sustainability (Chapter 15). Finally, Chapter 16 contrast to all of these terms, which refer to vegetational
provides a global perspective, setting the climate and entities, the terms CFR and GCFR are geographical.
biodiversity of the GCFR in the context of the world’s To assist readers unfamiliar with the geography of the
other Mediterranean-type ecosystems. GCFR, a map indicating key localities and landscape fea-
Invariably, a project of this scope relies on the sup- tures referred to in the book is provided in the beginning
port and contributions of many individuals, some of the book (Figure A). This is accompanied, for reference,
of whom are not listed as authors or co-authors on by a geological timescale for the Cenozoic (Figure B).
Figure A a map of the Greater Cape floristic Region showing features and places referred to in the text
h
Sub-era Period Epoch Sub-epoc Ma
0.01
QUAT. PLEISTOCENE
1.8
Late
PLIOCENE 3.6
Early
5.3
Late
E
EN 11.6
G
EO MIOCENE Middle
N
16.0
Early
23.0
Late
OLIGOCENE 28.1
RY Early
A
RTI 33.9
E
T Late
37.8
E`
N
E Middle
G
O EOCENE
E
A
L
A
P 47.8
Early
56.0
Late
PALAEOCENE
61.6 Figure B a geological time scale for the Cenozoic Era. The Holocene is indicated by hatching. ages
Early are from Walker, J.D., Geissman, J.W., Bowring, S.a., and Babcock, L.E., compilers. (2012). Geologic
66.0 Time Scale, version 4.0. Geological Society of america, doi: 10.1130/2012.CTS004R3C.
Contents
List of Contributors xi
1 Vegetation types of the Greater Cape Floristic Region 1
Nicola G. Bergh, G. anthony Verboom, Mathieu Rouget, and Richard M. Cowling
2 Landscapes, rock types, and climate of the Greater Cape Floristic Region 26
Peter L. Bradshaw and Richard M. Cowling
3 Drivers, ecology, and management of fire in fynbos 47
Tineke Kraaij and Brian W. van Wilgen
4 Floristic and faunal Cape biochoria: do they exist? 73
Jonathan f. Colville, alastair J. Potts, Peter L. Bradshaw, G. John Measey, Dee Snijman,
Mike D. Picker, Şerban ProcheŞ, Rauri C. K. Bowie, and John C. Manning
5 Cenozoic assembly of the Greater Cape flora 93
G. anthony Verboom, H. Peter Linder, félix forest, Vera Hoffmann, Nicola G. Bergh,
and Richard M. Cowling
6 Speciation and extinction in the Greater Cape Floristic Region 119
allan G. Ellis, G. anthony Verboom, Timotheus van der Niet, Steven D. Johnson,
and H. Peter Linder
7 The shifting landscape of genes since the Pliocene: terrestrial
phylogeography in the Greater Cape Floristic Region 142
Krystal a. Tolley, Rauri C.K. Bowie, G. John Measey, Benjamin W. Price,
and félix forest
8 Stone Age people in a changing South African Greater Cape Floristic Region 164
Curtis W. Marean, Hayley C. Cawthra, Richard M. Cowling, Karen J. Esler, Erich fisher,
antoni Milewski, alastair J. Potts, Elzanne Singels, and Jan De Vynck
9 The assembly and function of Cape plant communities in a changing world 200
Jasper a. Slingsby, David D. ackerly, andrew M. Latimer, H. Peter Linder, and anton Pauw
10 Biotic interactions 224
Bruce anderson, Nicky allsopp, allan G. Ellis, Steven D. Johnson, Jeremy J. Midgley, anton Pauw,
and James G. Rodger
11 Plant ecophysiological diversity 248
Michael D. Cramer, adam G. West, Simon C. Power, Robert Skelton, and William D. Stock
ix