The Karoo: Ecological Patterns and Processes

, by Edited by W. Richard J. Dean , Suzanne Milton

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ISBN: 9780521554503 | 0521554500
Cover: Hardcover
Copyright: 6/28/1999

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The succulent and Nama-karoo form part of the arid south-western zone of Africa, a vast region of rugged landscapes and low treeless vegetation. Studies of this unique biome have yielded fascinating insights into the ecology of its flora and fauna. This book, originally published in 1999, is the first to synthesise these studies, presenting information on biogeographic patterns and life processes, form and function of animals and plants, foraging ecology, landscape-level dynamics and anthropogenic influences. Detailed analyses of the factors distinguishing the biota of the Karoo from that of other temperate deserts are given and generalisations about semi-arid ecosystems challenged. The ideas expounded, the ecological principles reviewed, and the results presented are relevant to all those working in the extensive arid and semi-arid regions of the world.

List of contributors

xvi

Foreword

xix

Preface

xxi

Acknowledgements

xxiii

Part One Biogeographic patterns and the driving variables

(86)

S.J. Milton

W.R.J. Dean

The climate of the karoo -- a functional approach

(14)

P.G. Desmet

R.M. Cowling

Introduction

(1)

A general overview of the climate of the karoo

(6)

Precipitation

(1)

Rainfall

(1)

Fog

(1)

Dew

(3)

Reliability of sources of moisture

(1)

El Nino in southern Africa

(1)

Temperature

(1)

Cloud and light

(1)

Wind

(1)

Berg winds

(1)

The weather systems of the karoo

(2)

Fine-weather conditions

(1)

Subtropical anticyclones

(1)

Coastal lows

(1)

Tropical disturbances

(1)

Easterly waves and lows

(1)

Subtropical lows

(1)

Temperate disturbances

(1)

Westerly waves

(1)

Cut-off lows

(1)

Southerly meridional flow

(1)

Ridging anticyclones

(1)

West-coast troughs

(1)

Cold fronts

(1)

Other rain-producing systems

(1)

Thunderstorms

(1)

Development of the continuous high-pressure cell

(1)

An ecological interpretation of the weather patterns of the karoo: the role of climate in understanding vegetation patterns

(4)

The model

(1)

Discussion: vegetation--climate relationships

(1)

Succulent karoo

(2)

Nama-karoo

(1)

Desert

(1)

Conclusions

(1)

Soils of the arid south-western zone of Africa

(10)

M.K. Watkeys

Introduction

(1)

Controlling factors on soil variability

(3)

General factors

(1)

Geology and geomorphology of the karoo

(2)

Climate of the karoo

(1)

Soils of the karoo

(5)

Region 1. West coast

(1)

Region 2. Namaqualand, Bushmanland and Korannaland

(1)

Region 3. The Great Karoo (north)

(1)

Region 4. Great Karoo (south)

(1)

Region 5. Little Karoo

(1)

Region 6. The Great Escarpment

(2)

Discussion of the soil pattern

(1)

Acknowledgements

(2)

Palaeoenvironments

(15)

M.E. Meadows

M.K. Watkeys

Introduction

(1)

Long-term geological and biological history

(5)

Pre-Gondwana (3400--1000 Ma)

(2)

Gondwana assembling (800--250 Ma)

(2)

Gondwana assembled (250--155 Ma)

(1)

Gondwana breaking up (155 Ma to present)

(1)

Tertiary historical biogeography

(2)

Palaeoenvironmental indications during the Tertiary

(2)

The Banke flora

(1)

Geomorphological development

(1)

The Quaternary period including the Holocene

(5)

Fluctuating climates of the Quaternary

(1)

Florisbad

(1)

Aliwal North

(1)

Pan sites of the karoo

(1)

Valley deposits of the karoo

(2)

Other deposits

(1)

Climate change or human impact?

(1)

The karoo as a dynamic landscape

(2)

Plant biogeography, endemism and diversity

(15)

R.M. Cowling

C. Hilton-Taylor

Introduction

(1)

Biogeographical background

(4)

Biogeographical units and delimitation

(2)

Composition and characteristics of floras

(1)

Selective regimes

(1)

Endemism

(2)

Patterns

(1)

Taxonomic aspects

(1)

Biological aspects

(1)

Habitat aspects

(1)

Local diversity

(3)

Theory

(1)

Patterns

(1)

Explanations

(1)

Differentiation diversity

(1)

Regional diversity

(3)

Theory

(1)

Patterns

(1)

Explanations

(1)

General discussion

(2)

Phytochoria compared

(1)

Global comparisons

(1)

Diversification in an arid world

(1)

Acknowledgements

(1)

Biogeography, endemism and diversity of animals in the karoo

(30)

C.J. Vernon

Introduction

(1)

Environmental influences

(1)

Seasonal movements

(1)

Evolution

(1)

Origins of the karoo fauna

(7)

Gondwana

(1)

Palaearctic links and the mussel fauna

(3)

Drainage basins and the fish fauna

(1)

Rivers as dispersal routes

(1)

Afro-montane zone

(1)

The arid corridor and other biogeographic patterns

(1)

The Cradock corridor

(1)

The Transkei gap

(1)

Marine regression

(1)

Savanna--fynbos links

(1)

Endemicity and distribution patterns -- Nama-karoo

(12)

Invertebrates

(2)

Vertebrates

(4)

Endemicity and distribution patterns -- succulent karoo

(1)

Invertebrates

(2)

Vertebrates

(3)

Anthropogenic factors influencing biogeography

(1)

Conclusions

(9)

Part Two Form and function

(92)

B.G. Lovegrove

Form and function in perennial plants

(16)

G.F. Midgley

F. van der Heyden

Introduction

(1)

Growth form abundance, distribution and diversity

(1)

Distribution and abundance of photosynthetic types

(1)

Water

(8)

Succulence

(3)

Succulent vs. non-succulent shrubs

(2)

Minutism

(1)

Grasses

(1)

Desiccation tolerance

100

(1)

Root architecture

100

(2)

Temperature

102

(1)

Herbivory

103

(2)

Comparing shrubs with grasses

103

(1)

Shrub functional types

104

(1)

Nutrient availability

105

(1)

Conclusions

106

(1)

Acknowledgements

106

(1)

Functional aspects of short-lived plants

107

(16)

M.W. van Rooyen

Introduction

107

(1)

Seedbanks

108

(2)

Mechanisms for seed longevity

108

(1)

Spatial and temporal distribution of seed

108

(2)

Germination

110

(3)

After-ripening

110

(1)

Polymorphism

110

(1)

Selective response mechanisms to specific environments

111

(1)

Endogenous rhythm

112

(1)

Parental effects

112

(1)

Vegetative growth

113

(1)

Establishment

113

(1)

Growth rate

113

(1)

Photosynthetic pathway

114

(1)

Reproductive growth

114

(2)

Timing of reproduction

114

(1)

Phenotypic plasticity

115

(1)

Seed dispersal

116

(1)

Biotic interactions

117

(5)

Disturbance

117

(1)

Competition

118

(2)

Functional classification into guilds

120

(2)

Conclusions

122

(1)

Acknowledgements

122

(1)

Plant reproductive ecology

123

(22)

K.J. Esler

Introduction

123

(1)

General phenology

124

(1)

Phenology

124

(1)

Flowering phenology

124

(3)

Timing of flowering

124

(1)

Triggers for flowering

125

(1)

Landscape patterns of flowering

126

(1)

Pollination biology

127

(3)

Flowering phenology and competition for pollinators

127

(1)

Insect pollination

128

(1)

Other forms of pollination

129

(1)

Pollinator abundance and seed set

129

(1)

Associations among modes of pollination, seed dispersal and establishment

129

(1)

Pre-dispersal hazards

130

(1)

Pollen loss

130

(1)

Florivory

130

(1)

Resource availability

130

(1)

Seed production

131

(1)

Seed numbers (per plant)

131

(1)

Seed numbers (landscape level)

131

(1)

Dispersal

132

(3)

Dispersal mechanisms and habitat

132

(1)

Short-distance dispersal and seed retention

132

(1)

Dispersal in Mesembryanthema

133

(1)

Dispersal distances

133

(2)

Dispersal and seed polymorphism

135

(1)

Other forms of dispersal

135

(1)

Seedbanks

135

(1)

Soil seedbanks

135

(1)

Canopy seedbanks

135

(1)

Spatial distribution of seeds in seedbanks

135

(1)

Correspondence between the seedbank and vegetation

136

(1)

Post-dispersal hazards

136

(1)

Germination

137

(2)

Germination and life-histories

137

(1)

Germination and intrinsic factors

138

(1)

Germination and extrinsic factors

139

(1)

Seedling establishment

139

(3)

Timing of recruitment

139

(1)

Size of recruitment events

140

(1)

Microsites and recruitment

140

(1)

Disturbance and recruitment

141

(1)

Mycorrhizal status and recruitment

142

(1)

Seeding mortality

142

(1)

Seed vs. vegetative reproduction

143

(1)

Concluding remarks and future research

143

(1)

Acknowledgements

144

(1)

Animal form and function

145

(19)

B.G. Lovegrove

Introduction

145

(1)

Water balance

145

(3)

Water loss

146

(1)

Water gain

147

(1)

Thermal balance

148

(4)

Thermal tolerance

148

(1)

Heat gain

149

(2)

Heat loss

151

(1)

Energy balance

152

(1)

Basal metabolic rates

152

(1)

Circadian metabolic rhythms

152

(1)

Torpor

153

(1)

Retreat

153

(3)

Short-term retreat

154

(1)

Burrowing adaptations

154

(1)

Circadian rhythms of retreat

155

(1)

Nest shelter retreats

155

(1)

Long-term retreat (diapause, migration and nomadism)

156

(1)

Sociality and co-operative breeding

156

(2)

Predator--prey adaptations

158

(2)

Behaviour

158

(1)

Crypsis and colouration

159

(1)

Mimicry

159

(1)

Morphology and body shape

160

(1)

Reproduction

160

(4)

Cues and Zeitgeber for reproduction

160

(2)

The brown locust

162

(2)

Animal foraging and food

164

(15)

W.R.J. Dean

S.J. Milton

Introduction

164

(1)

Dependable resources

164

(6)

Below-ground foragers

165

(1)

Herbivores

165

(1)

Carnivores

165

(1)

Above-ground foragers

165

(1)

Detritivores

165

(1)

Herbivores

166

(1)

Termitivores

167

(2)

Formicivores

169

(1)

Other insectivores

169

(1)

Parasitoids and parasites

169

(1)

Seasonal resources

170

(5)

Opportunism and diet switching

170

(2)

Storage and caching of food

172

(1)

Seasonally active and short-lived animals

173

(2)

Unpredictable resources scattered in time and space

175

(1)

Granivores

175

(1)

Herbivores

176

(1)

Food webs

176

(3)

Part Three Dynamics

179

(60)

P.A. Novellie

Population level dynamics

183

(25)

S.J. Milton

R.A.G. Davies

G.I.H. Kerley

Introduction

183

(1)

Structure and dynamics of plant populations

183

(8)

Ephemerals and annuals

183

(1)

Grasses

184

(2)

Succulents

186

(1)

Non-succulent dwarf shrubs

187

(3)

Shrubs and trees

190

(1)

Principal drivers of karoo plant populations

191

(1)

Invertebrates

191

(3)

Brown locust Locustana pardalina

191

(2)

Other insects

193

(1)

General patterns in invertebrate populations

193

(1)

Reptiles and birds

194

(4)

Tortoises

194

(1)

Spatial and temporal patterns in bird abundance

194

(2)

Resident, raptorial birds

196

(1)

Resident, omnivorous birds

196

(2)

Local migrants and nomads

198

(1)

Mammals

198

(8)

Lagomorphs, rodents and shrews

198

(1)

Rock hyrax Procavia capensis

199

(1)

Additions to rock hyrax populations

200

(2)

Losses to rock hyrax populations

202

(1)

Key factors affecting rock hydrax populations

203

(1)

Large herbivores

204

(1)

Primates and carnivores

205

(1)

Conclusions

206

(2)

Community patterns and dynamics

208

(16)

A.R. Palmer

P.A. Novellie

J.W. Lloyd

Introduction

208

(1)

Vegetation patterns: landscape scale

209

(2)

Topo-moisture gradients, rainfall seasonality and uncertainty

209

(1)

Soil types, depth, stoniness, texture and chemistry

210

(1)

Vegetation patterns: local scale

211

(4)

Nutrient enrichment patterns

211

(3)

Nutrient transport patterns

214

(1)

Photosynthetic patterns

214

(1)

Diversity patterns in the landscape

215

(1)

Plant community dynamics

215

(5)

Rainfall variability

215

(1)

Fire

215

(1)

Grazing

215

(1)

Mechanisms for grazing-induced change

215

(4)

Implications of grazing-induced change

219

(1)

Reversal of grazing-induced changes

219

(1)

Time scales in plant community dymanics

220

(1)

Animal community dynamics

220

(2)

Indigenous ungulates

220

(1)

Invertebrates

221

(1)

Termites and ants

222

(1)

Conclusions

222

(2)

Modelling populations and community dynamics in karoo ecosystems

224

(7)

K. Kellner

J. Booysen

Introduction

224

(1)

Models to predict the outbreak of problem animals

224

(1)

Models to study animal population dynamics

225

(2)

Production and drought prediction models

227

(1)

Models for vegetation dynamics and sustainable rangeland management

228

(2)

Conclusions

230

(1)

Spatially explicit computer simulation models -- tools for understanding vegetation dynamics and supporting rangeland management

231

(8)

F. Jeltsch

T. Wiegand

C. Wissel

Introduction

231

(1)

Simulation models

232

(4)

Vegetation dynamics of a shrub ecosystem in the karoo

232

(1)

Shrub encroachment in the north-western karoo/southern Kalahari

233

(2)

Analysing a sophisticated grazing strategy at a ranch level

235

(1)

How do the models contribute to the understanding and management of karoo vegetation?

236

(3)

Part Four Human impacts

239

(50)

W.R. Siegfried

The environment

239

(1)

The people

240

(1)

Impacts

240

(3)

Hunters and herders in the karoo landscape

243

(14)

A.B. Smith

Introduction

243

(1)

Human adaptation to arid lands

243

(2)

Early humans in the karoo

245

(3)

The later Stone Age

248

(7)

The Cape west coast

248

(1)

Namaqualand

249

(1)

Northern Namaqualand and the Namib

250

(1)

Bushmanland and the Middle Orange River

251

(1)

Cape Fold Belt (west) and Tanqua Karoo

251

(2)

Cape Fold Belt (south) and Little Karoo

253

(1)

The Great Karoo

253

(2)

Rock art

255

(1)

Khoisan/settler interactions

255

(1)

Conclusions

256

(1)

Historical and contemporary land use and the desertification of the karoo

257

(17)

M.T. Hoffman

B. Cousins

T. Meyer

A. Petersen

H. Hendricks

Introduction

257

(1)

Communal lands

257

(6)

Historical background

257

(3)

Current land use practices and their determinants

260

(3)

Commercial agriculture

263

(7)

Settlement and historical land use practices

263

(2)

Current land use practices

265

(5)

Desertification of the karoo

270

(2)

A brief history of the karoo desertification debate

270

(1)

Hypothesis 1: The precolonial eastern karoo was a stable and extensive `sweet' grassland

271

(1)

Hypothesis 2: An altered and less-productive karoo vegetation is expanding into more productive grassland environments

272

(1)

Conclusions

272

(2)

Alien plant invaders of the karoo: attributes, impacts and control

274

(15)

S.J. Milton

H.G. Zimmermann

J.H. Hoffmann

Introduction

274

(1)

Invasibility of karoo vegetation types

274

(2)

Attributes of plants invasive in the karoo

276

(4)

General trends

276

(2)

Moist habitats

278

(1)

Dryland habitats

278

(1)

South-western areas:succulent karoo and Little Karoo

278

(1)

Northern and central areas:Kalahari and Bushmanland

279

(1)

Eastern areas: Great, Lower and Eastern Nama-karoo

279

(1)

Impacts on the karoo ecosystem and economy

280

(3)

Woody plants

280

(1)

Cactaceae

281

(1)

Herbaceous plants

282

(1)

Control

283

(1)

Prosopis

283

(1)

Cactaceae

283

(1)

Comparison with other arid and semi-arid systems

284

(2)

Invasive taxa and life forms

285

(1)

Karoo species as invaders elsewhere

285

(1)

Future scenarios and research needs

286

(1)

Acknowledgements

287

(2)

Part Five Comparisons

289

(30)

Comparison of ecosystem processes in the Nama-karoo and other deserts

291

(12)

W.G. Whitford

Introduction

291

(1)

Physical environment and primary productivity

291

(3)

Precipitation patterns

291

(1)

Topographic similarities

292

(1)

Water redistribution

292

(1)

Primary productivity

292

(1)

Shrub morphology

293

(1)

Decomposition and nutrient cycling

294

(6)

Soil organisms and soil processes

294

(1)

Termites

295

(1)

Buried litter and roots

296

(1)

Role of soil biota

296

(1)

Mineralization and immobilization

297

(1)

Rhizosphere processes

298

(1)

Soil aggregates

298

(1)

Soil perturbation

299

(1)

Granivory and herbivory

299

(1)

Historical changes and desertification

300

(2)

Acknowledgements

302

(1)

The succulent karoo in a global context: plant structural and functional comparison with North American winter-rainfall deserts

303

(11)

K.J. Esler

P.W. Rundel

R.M. Cowling

Introduction

303

(1)

Selective regime

304

(2)

Plant form and community structure

306

(1)

Community phenology

307

(2)

A Namaqualand--Namib domain (succulent karoo) model

309

(1)

Plant function

310

(3)

Population structure and turnover

313

(1)

Conclusions

313

(1)

The karoo: past and future

314

(5)

S.J. Milton

W.R.J. Dean

Current perceptions

314

(2)

Patterns

314

(1)

Processes

315

(1)

Uncertainties and contradictions

316

(1)

Directions for future research

317

(2)

References

319

(50)

Index

369

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The Karoo: Ecological Patterns and Processes

Summary

Table of Contents

Supplemental Materials

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The Karoo: Ecological Patterns and Processes

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