Part I An Introduction to Angola’s Biomes and Ecoregions 
1 Opportunities in and Approaches to the Study of Angolan 
Ecology ...................................................... 5 
1.1 The Science and Practice of Ecology ........................ 6 
1.2 The Structure of This Introduction to Terrestrial Ecology ....... 11 
1.3 How to Use This Book: Interconnections and Recurrent 
Messages ............................................... 13 
References .................................................... 13 
2 Biomes: Concepts, Characteristics and Terminology .............. 15 
2.1 Biodiversity, Biogeography and Centres of Endemism ......... 17 
2.2 Biome and Ecoregion Definition and Characterisation ......... 19 
2.3 The Biomes of Africa in Outline ............................ 21 
2.4 Classification of Angola’s Biomes and Ecoregions ............ 26 
References .................................................... 40 
3 Profiles of Angola’s Biomes and Ecoregions ...................... 43 
3.1 Guineo-Congolian Rain Forest Biome and Forest/Savanna 
Mosaic ................................................. 43 
3.2 Afromontane Forest and Grassland Biome ................... 47 
3.3 Mesic Savanna Biome .................................... 49 
3.4 Arid Savanna Biome ...................................... 59 
3.5 Namib Desert Biome ..................................... 62 
3.6 Mangrove Biome ......................................... 65 
3.7 Angolan Escarpment Zone ................................. 67 
References .................................................... 68 
Part II Determinants of Pattern: Conditions, Resources and 
Disturbance 
4 Landscapes: Geology, Geomorphology and Hydrology ............ 73 
4.1 Geology: A Brief History of the Earth and of Angola .......... 78
4.2 Landscapes, Landscape Ecology and Geomorphology ......... 83 
4.3 Hydrology: Wetlands, Water Towers, Rivers, Floodplains 
and Estuaries ............................................ 88 
References .................................................... 92 
5 Solar Energy, Temperature and Rainfall ......................... 95 
5.1 Where Life Begins: Solar Radiation and Photosynthesis ........ 96 
5.2 Seasonality of Day Length and Temperature .................. 98 
5.3 Large-Scale Drivers of Climate: Oceanic and Atmospheric 
Circulation .............................................. 99 
5.4 Local Influences: Maritime, Continental, Altitudinal 
and Aspect .............................................. 104 
5.5 Water Cycle Processes: Precipitation 
and Evapotranspiration .................................... 105 
5.6 Regional Climates of Angola ............................... 106 
5.7 Climate as a Determinant of Vegetation Patterns .............. 111 
5.8 El Niño, La Niña, Droughts and Floods ...................... 118 
References .................................................... 124 
6 Soil, Water and Nutrients ...................................... 127 
6.1 Soil Structure ............................................ 128 
6.2 Development of Soil: Weathering and Pedogenesis ............ 130 
6.3 Soil Water Relations ...................................... 132 
6.4 Soil Chemistry and Nutrient Status ......................... 135 
6.5 Soils and Tree Growth .................................... 137 
6.6 Soil Classification ........................................ 140 
6.7 Key Soil Groups of Angola ................................ 140 
References .................................................... 147 
7 The Ecological Role of Fire ..................................... 149 
7.1 The Ancient History of Fire in Africa ....................... 150 
7.2 Fire Regimes and Pyromes ................................ 151 
7.3 Fire Intensity ............................................ 154 
7.4 Fire Season .............................................. 155 
7.5 Fire Frequency ........................................... 157 
7.6 Impacts of Fire on Soils ................................... 157 
7.7 Plant Responses to Fire ................................... 159 
7.8 Responses of Animals to Fire .............................. 160 
References .................................................... 163 
8 Herbivory: Mammalian Grazers and Browsers .................. 167 
8.1 The Evolution of African Mammalian Herbivores 
and of Arid/Eutrophic Savannas ............................ 168 
8.2 The Herbivomes of Africa ................................. 173 
8.3 Patterns of Impacts of Browsers and Grazers ................. 176 
References .................................................... 181
Part III Ecological Concepts and Ecosystem Processes 
9 General Concepts in Ecology ................................... 185 
9.1 The Evolutionary Basis of Ecology ......................... 193 
9.1.1 Species Richness, Evenness and Diversity ............ 196 
9.2 Endemism, Threatened Species and Hotspots ................. 200 
9.3 Life History Patterns: r-and K-Selection Strategies ........... 202 
9.4 The Concepts of the Ecological Niche and the Guild .......... 203 
9.5 Herbivory, Predation, Parasitism and Mutualism .............. 205 
9.6 Theories of Island Biogeography and of Metapopulations ...... 209 
References .................................................... 212 
10 Ecosystem Processes and Dynamics in Mesic Savannas ........... 215 
10.1 Food Chains, Food Webs and Trophic Levels ................. 219 
10.2 The Photosynthetic Pathways of Angolan Plants .............. 221 
10.3 Primary Production in Mesic Savannas ...................... 222 
10.4 Standing Crop Biomass and Canopy Cover in Mesic 
Savannas ................................................ 226 
10.5 Herbivory and Primary Consumption in African Savannas ...... 226 
10.6 Decomposition and Nutrient Cycling ........................ 230 
10.7 Savanna/Forest Dynamics: Fire as the Driver of Alternative 
Stable States ............................................. 236 
References .................................................... 245 
11 Adaptations to Life in the Namib Desert ......................... 249 
11.1 Water: The Currency of Life in Desert Environments .......... 250 
11.2 Adaptations to the Heat of the Desert ........................ 256 
11.3 Competition for Food in a Low Productivity Environment ...... 259 
11.4 Defence Mechanisms: Camouflage, Deceit, Mimicry, 
Spines, Poisons and Social Behaviour ....................... 260 
11.5 Reproduction and Survival in Desert Environments ............ 265 
References .................................................... 273 
Part IV Ecological Features of Angolan Biomes and Ecoregions 
12 The Guineo-Congolian Rain Forest Biome ....................... 279 
12.1 Definitition and Distribution ............................... 284 
12.2 Climate and Microclimate: The Roles of Fog (Cacimbo) 
and of Shade ............................................ 286 
12.3 Physiognomic Structure ................................... 288 
12.4 Forest Canopy Growth Cycles: Gap-Phase Dynamics .......... 289 
12.5 Floristic Composition ..................................... 293 
12.6 Faunal Composition ...................................... 297 
12.7 Rain Forest Mammals ..................................... 297 
12.8 Interconnections Between Plants and Animals ................ 300 
References .................................................... 302
13 The Afromontane Forest and Montane Grassland Biome .......... 305 
13.1 Afromontane Forests (Ecoregion 4) ......................... 306 
13.2 Afromontane Grasslands (Ecoregion 5) ...................... 308 
13.3 Faunal Composition of Afromontane Forests 
and Grasslands ........................................... 310 
References .................................................... 312 
14 The Mesic Savanna Biome ..................................... 313 
14.1 Definition and Distribution ................................ 315 
14.2 Landscapes, Soils, the Catena Concept, Termitaria 
and Underground Forests .................................. 317 
14.3 Climate and Seasonality ................................... 331 
14.4 Floristic Composition, Physiognomic Structure 
and Phenology ........................................... 332 
14.5 Faunal Composition of Mesic Savannas ..................... 334 
14.6 Mammals of the Angolan Mesic Savannas ................... 335 
References .................................................... 335 
15 The Arid Savanna Biome ...................................... 339 
15.1 Angolan Mopane Woodlands (Ecoregion 12) ................. 340 
15.2 Namib Savanna Woodlands (Ecoregion 13) .................. 347 
15.3 Coastal Arid Savannas (Ecoregion 14) ....................... 350 
15.4 Faunal Composition of Arid Savannas ....................... 357 
References .................................................... 358 
16 The Namib Desert Biome ...................................... 361 
16.1 The Age and Evolution of the Namib Desert ................. 362 
16.2 Landscapes and Soils ..................................... 365 
16.3 Climate: The South Atlantic Anticyclone, the Benguela 
Current and Wind ........................................ 369 
16.4 Floristic Composition and Physiognomy of the Angolan 
Namib Desert Ecoregion .................................. 371 
16.5 Faunal Composition of the Namib Desert .................... 374 
References .................................................... 381 
17 The Mangrove Biome .......................................... 383 
17.1 Definition and Distribution ................................ 383 
17.2 Floristic Composition and Physiognomy ..................... 385 
17.2.1 Mangrove Adaptations to Waterlogged Soil ........... 387 
17.2.2 Adaptations to Changes in Salinity .................. 388 
17.2.3 Reproductive Adaptations .......................... 388 
17.2.4 Seagrass Meadows ................................ 389 
17.3 Faunal Composition of Mangrove and Seagrass Meadow 
Ecosystems .............................................. 390 
References .................................................... 391
18 The Angolan Escarpment Zone ................................. 393 
18.1 Sectors of the Angolan Escarpment Zone .................... 394 
18.2 Sectors of the Angolan Escarpment Zone .................... 396 
18.3 Vertebrate Fauna of the Escarpment Zone .................... 398 
References .................................................... 403 
Part V Synopsis and Conclusions 
19 Key Elements of Angolan Terrestrial Ecology .................... 407 
19.1 The Big Picture: Global Tectonic and Climatic Forces 
that Have Shaped Angola’s Biomes and Biota ................ 408 
19.2 Contemporary Drivers of Ecosystem Structure 
and Function in Angola ................................... 410 
19.3 Ecological Patterns at African and Angolan Scales ............ 414 
19.4 Patterns at Landscape Scale in Angola ....................... 415 
19.5 Ecological Concepts and Theories Relevant to Conserving 
Angolan Species ......................................... 418 
19.6 Conclusions: Why Ecology? An Opportunity for Young 
Angolan Students ........................................ 419 
References .................................................... 420 
Appendix: Names of Vertebrate Species Mentioned in This Volume .... 423 
Glossary of Ecological Terms ....................................... 435