Evolutionary Diversity Patterns in the Cape Flora of South Africa

  • Félix ForestEmail author
  • Jonathan F. Colville
  • Richard M. Cowling


The Greater Cape Floristic Region of South Africa is a relatively small area found in the south-western corner of the African continent. It is characterised by a Mediterranean-type climate and harbours an incredible floral diversity with almost 11,500 species of plants, of which more than three-quarters are found nowhere else in the world. The floristic composition of the region is very distinctive with families not dominant in other floras, such as Iridaceae, Aizoaceae, Ericaceae, Proteaceae and Restionaceae. In addition, small-leaved, sclerophyllous low shrubs and geophytes are by far the predominant growth forms with a low proportion of tree and annual species present. Understandably, this flora has attracted a lot of interest, and many have pondered what environmental and ecological factors could be responsible for its richness and uniqueness and the resulting biodiversity patterns observed today. In this chapter, the current state of knowledge of large-scale evolutionary patterns, investigated using phylogenetic diversity metrics, within the Cape region are reviewed. As well as published works, we report on ongoing studies examining biodiversity patterns using phylogenetic diversity, phylogenetic endemism and phylobetadiversity, both in plants and animals. We also review how these phylogeny-based approaches have been used to address applied questions (e.g. biodiversity surrogates, phytogeographical delimitations).



We thank Krystal Tolley for providing information on the current state of reptile phylogenetic diversity patterns. This chapter is derived, in part (i.e. Fig. 9.1), from an article published in the Transactions of the Royal Society of South Africa on 2 February 2017, available online:


  1. Allsopp N, Colville JF, Verboom GA (2014) Fynbos: ecology evolution and conservation of a megadiverse region. In: Allsopp N, Colville JF, Verboom GA (eds) Fynbos: ecology, evolution, and conservation of a megadiverse region. Oxford University Press, OxfordCrossRefGoogle Scholar
  2. Bar-Matthews M, Marean CW, Jacobs Z, Karkanas P, Fisher EC, Herries AIR, Brown K, Williams HM, Bernatchez J, Ayalon A, Nilssen PJ (2010) A high resolution and continuous isotopic speleothem record of paleoclimate and paleoenvironment from 90 to 53 ka from pinnacle point on the south coast of South Africa. Quat Sci Rev 29:2131–2145. Scholar
  3. Bayer MB (1984) The cape flora and the Karoo - a winter rainfall biome versus a fynbos biome. Veld Flora 70:17–19Google Scholar
  4. Bergh NG, Verboom GA, Rouget M, Cowling RM (2014) Vegetation types of the greater cape floristic region. In: Allsopp N, Colville JF, Verboom GA (eds) Fynbos: ecology, evolution, and conservation of a Megadiverse region. Oxford University Press, Oxford, pp 1–25Google Scholar
  5. Bond P, Goldblatt P (1984) Plant of the Cape flora: a descriptive catalogue. J South Afr Bot 13 p 1–455Google Scholar
  6. Born J, Linder HP, Desmet P (2007) The greater cape floristic region. J Biogeogr 34:147–162CrossRefGoogle Scholar
  7. Bradshaw PL, Colville JF, Linder HP (2015) Optimising regionalisation techniques: identifying centres of endemism in the extraordinarily endemic-rich cape floristic region. PLoS One 10:e0132538. Scholar
  8. Braschler B, Chown SL, Gaston KJ (2012) The Fynbos and succulent Karoo biomes do not have exceptional local ant richness. PLoS One 7:e31463. Scholar
  9. Braun K, Marean CW, Bar-Matthews M, Zahn R, Cowling RM, Matthews A, Ayalon A, Dyez K, Karkanas P, Fisher E, Zilberman T (Submitted) A climate and environment record dating between ~444 and 41 ka from pinnacle point (south coast, South Africa) as derived from speleothem stable isotopic compositions. Quat Sci RevGoogle Scholar
  10. Britton MN, Hedderson TAJ, Verboom GA (2014) Topography as a driver of cryptic speciation in the high-elevation cape sedge Tetraria triangularis (Boeck.) C. B. Clarke (Cyperaceae: Schoeneae). Mol Phylogenet Evol 77:96–109. Scholar
  11. Buerki S, Callmander MW, Bachman S, Moat J, Labat JN, Forest F (2015) Incorporating evolutionary history into conservation planning in biodiversity hotspots. Philos Trans R Soc B 370(1662):20140014. Scholar
  12. Colville JF, Potts AJ, Bradshaw PL, Measey GJ, Snijman DA, Picker MD, Procheş Ş, Bowie RCK, Manning JC (2014) Floristic and faunal cape biochoria: do they exist? In: Allsopp N, Colville JF, Verboom GA (eds) Fynbos: ecology, evolution, and conservation of a megadiverse region. Oxford University Press, Oxford, pp 73–92CrossRefGoogle Scholar
  13. Colville JF, Forest F, Beale CM, Altwegg R, Huntley B, Bradshaw PL, Cowling RM (In prep) High contemporary and evolutionary plant diversity linked to stability and ecological heterogeneityGoogle Scholar
  14. Cowling RM (1987) Fire and its role in coexistence and speciation in gondwanan shrublands. S Afr J Sci 83:106–112Google Scholar
  15. Cowling RM, Holmes PM (1992) Endemism and speciation in a lowland flora from the cape floristic region. Biol J Linn Soc 47:367–383CrossRefGoogle Scholar
  16. Cowling RM, Lombard AT (2002) Heterogeneity, speciation/extinction history and climate: explaining regional plant diversity patterns in the cape floristic region. Divers Distrib 8:163–179CrossRefGoogle Scholar
  17. Cowling RM, Holmes PM, Rebelo AG (1992) Plant diversity and endemism. In: Cowling RM (ed) Ecology of Fynbos. Nutrients, fire and diversity. Oxford University Press, Cape Town, pp 69–112Google Scholar
  18. Cowling RM, Procheş Ş, Partridge TC (2009) Explaining the uniqueness of the cape flora: incorporating geomorphic evolution as a factor for explaining its diversification. Mol Phylogenet Evol 51:64–74CrossRefPubMedGoogle Scholar
  19. Cowling RM, Bradshaw PL, Colville JF, Forest F (2017) Levyns’ law: explaining the evolution of a remarkable longitudinal gradient in cape plant diversity. Trans R Soc S Afr 72:184–201. Scholar
  20. Cox CB (2001) The biogeographic regions reconsidered. J Biogeogr 28:511–523CrossRefGoogle Scholar
  21. Daru BH, Van der Bank M, Davies TJ (2015) Spatial incongruence among hotspots and complementary areas of tree diversity in southern Africa. Divers Distrib 21:769–780CrossRefGoogle Scholar
  22. Daru BH, Van der Bank M, Maurin O, Yessoufou K, Schaefer H, Slingsby JA, Davies TJ (2016) A novel phylogenetic regionalization of phytogeographical zones of southern Africa reveals their hidden evolutionary affinities. J Biogeogr 43:155–166. Scholar
  23. Daru BH, Van der Bank M, Davies TJ (2018) Unravelling the evolutionary origins of biogeographic assemblages. Divers Distrib 24:313–324CrossRefGoogle Scholar
  24. Davies TJ, Barraclough TG, Chase MW, Soltis PS, Soltis DE, Savolainen V (2004) Darwin’s abominable mystery: insights from a supertree of the angiosperms. Proc Natl Acad Sci U S A 101:1904–1909. Scholar
  25. Dynesius M, Jansson R (2000) Evolutionary consequences of changes in species’ geographical distributions driven by Milankovitch climate oscillations. Proc Natl Acad Sci U S A 97:9115–9120CrossRefPubMedPubMedCentralGoogle Scholar
  26. Endrödy-Younga S (1978) Coleoptera. In: Werger MJA (ed) Biogeography and ecology of southern Africa. Dr W. Junk bv Publishers, The Hague, pp 799–820Google Scholar
  27. Engelbrecht FA, Marean CW, Cowling RM, Potts AJ, Engelbrecht C, Nkoana R, O’Neill D, Fisher E, Shook e FJ, Neumann F, Scott L, Thatcher M, McGregor J, Van der Merwe J, Dedekind Z, Difford M (Submitted) Scaling last glacial maximum climate over southern Africa. Quat Sci RevGoogle Scholar
  28. Faith DP (2008) Threatened species and the potential loss of phylogenetic diversity: conservation scenarios based on estimated extinction probabilities and phylogenetic risk analysis. Conserv Biol 22:1461–1470. Scholar
  29. Faith DP, Walker PA (1996) Environmental diversity: on the best possible use of surrogate data for assessing the relative biodiversity of sets of areas. Biodivers Conserv 5:399–415CrossRefGoogle Scholar
  30. Forest F, Grenyer R, Rouget M, Davies TJ, Cowling RM, Faith DP, Balmford A, Manning JC, Procheş Ş, van der Bank M, Reeves G, Hedderson TAJ, Savolainen V (2007) Preserving the evolutionary potential of floras in biodiversity hotspots. Nature 445:757–760CrossRefPubMedGoogle Scholar
  31. Giliomee JH (2003) Insect diversity in the cape floristic region. Afr J Entomol 41:237–244Google Scholar
  32. Goldblatt P (1978) Analysis of the flora of southern-Africa - its characteristics, relationships, and origins. Ann Mo Bot Gard 65:369–436CrossRefGoogle Scholar
  33. Goldblatt P, Manning JC (2000) Cape plants: a conspectus of the cape flora of South Africa, Strelitzia 9. National Botanical Institute of South Africa and Missouri Botanical Garden, Pretoria and St. LouisGoogle Scholar
  34. Goldblatt P, Manning JC (2002) Plant diversity of the cape region of southern Africa. Ann Mo Bot Gard 89:281–302CrossRefGoogle Scholar
  35. Good R (1974) The geography of the flowering plants, 4th edn. Longman Green, LondonGoogle Scholar
  36. Graham CH, Fine PVA (2008) Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time. Ecol Lett 11:1265–1277CrossRefPubMedGoogle Scholar
  37. Grantham HS, Pressey RL, Wells JA, Beattie AJ (2010) Effectiveness of biodiversity surrogates for conservation planning: different measures of effectiveness generate a kaleidoscope of variation. PLoS One 5:e0011430. Scholar
  38. Guerin GR, Lowe AJ (2015) Mapping phylogenetic endemism in R using georeferenced branch extents. SoftwareX 3-4:22–26CrossRefGoogle Scholar
  39. Hanson JO, Rhodes JR, Riginos C, Fuller RA (2017) Environmental and geographic variables are effective surrogates for genetic variation in conservation planning. Proc Natl Acad Sci 114:12755–12760CrossRefPubMedGoogle Scholar
  40. Helme NA, Trinder-Smith TH (2006) The endemic flora of the cape peninsula, South Africa. S Afr J Bot 72:205–210. Scholar
  41. Holt BG, Lessard J-P, Borregaard MK, Fritz SA, Araújo MB, Dimitrov D, Fabre P-H, Graham CH, Graves GR, Jønsson KA, Nogués-Bravo D, Wang Z, Whittaker RJ, Fjeldså J, Rahbek C (2013) An update of Wallace’s zoogeographic regions of the world. Science 339:74–78. Scholar
  42. Johnson SD (1992) Plant-animal relationships. In: Cowling RM (ed) The ecology of Fynbos: nutrients, fire and diversity. Oxford University Press, Cape Town, pp 175–205Google Scholar
  43. Johnson SD (1996) Pollination, adaptation and speciation models in the cape flora of South Africa. Taxon 45:59–66CrossRefGoogle Scholar
  44. Johnson SD (2010) The pollination niche and its role in the diversification and maintenance of the southern African flora. Philos Trans R Soc B 365(1539):499–516. Scholar
  45. Jürgens N (1991) A new approach to the Namib region .1. Phytogeographic subdivision. Vegetatio 97:21–38Google Scholar
  46. Jürgens N (1997) Floristic biodiversity and history of African arid regions. Biodivers Conserv 6:495–514CrossRefGoogle Scholar
  47. Kemp JE, Ellis AG (2017) Significant local-scale plant-insect species richness relationship independent of abiotic effects in the temperate cape floristic region biodiversity hotspot. PLoS One 12:e0168033. Scholar
  48. Levyns MR (1952) Clues to the past in the cape flora of today. S Afr J Sci 49:155–164Google Scholar
  49. Levyns MR (1964) Migrations and origin of the cape flora. Trans R Soc S Afr 37:85–107CrossRefGoogle Scholar
  50. Linder HP (1985) Gene flow, speciation, and species diversity patterns in a species-rich area: the cape flora. In: Vrba ES (ed) Species and speciation, Transvaal Museum Monograph No. 4. Transvaal Museum, Pretoria, pp 53–57Google Scholar
  51. Linder HP (1991) Environmental correlates of patterns of species richness in the south-western Cape-Province of South-Africa. J Biogeogr 18:509–518CrossRefGoogle Scholar
  52. Linder HP (2003) The radiation of the cape flora, southern Africa. Biol Rev 78:597–638CrossRefPubMedGoogle Scholar
  53. Linder HP (2005) Evolution of diversity: the cape flora. Trends Plant Sci 10:536–541CrossRefPubMedGoogle Scholar
  54. Linder HP, de Klerk HM, Born J, Burgess ND, Fjeldsa J, Rahbek C (2012) The partitioning of Africa: statistically defined biogeographical regions in sub-Saharan Africa. J Biogeogr 39:1189–1205. Scholar
  55. Manning JC, Goldblatt P (2012) Plants of the greater cape floristic region 1: the Core cape flora, Strelitzia 29. South African National Biodiversity Institute, PretoriaGoogle Scholar
  56. Marloth R (1908) Das Kapland, insonderheit das Reich der Kapflora, das Waldgebiet und die Karroo, pflanzengeographisch dargestelt. In: Chun C (ed) Wissenschaffliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898–1899. Gustav Fischer, Jena, pp 1–427Google Scholar
  57. Maurin O, Davies TJ, Burrows JE, Daru BH, Yessoufou K, Muasya AM, Van der Bank M, Bond WJ (2014) Savanna fire and the origins of the ‘underground forests’ of Africa. New Phytol 204:201–214CrossRefPubMedGoogle Scholar
  58. Mecenero S, Altwegg R, Colville JF, Beale CM (2015) Roles of spatial scale and rarity on the relationship between butterfly species richness and human density in South Africa. PLoS One 10:e0124327. Scholar
  59. Midgley GF, Hannah L, Roberts R, MacDonald DJ, Allsopp JC (2001) Have Pleistocene climatic cycles influenced species richness in the greater Cape Mediterranean region. J Med Ecol 2:137–144Google Scholar
  60. Midgley GF, Reeves G, Klak C (2005) Late tertiary and quaternary climate change and centres of endemism in the southern African flora. In: Purvis A, Gittleman JL, Brooks T (eds) Phylogeny and conservation. Cambridge Univeristy Press, Cambridge, pp 230–242CrossRefGoogle Scholar
  61. Mucina L, Rutherford MC (2006) The vegetation of South Africa, Lesotho and Swaziland. In: Strelitzia, vol 19. South African National Biodiversity Institute, PretoriaGoogle Scholar
  62. Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403:853–858. Scholar
  63. Nic Lughadha E, Baillie J, Barthlott W, Brummitt NA, Cheek MR, Farjon A, Govaerts R, Hardwick KA, Hilton-Taylor C, Meagher TR, Moat J, Mutke J, Paton AJ, Pleasants LJ, Savolainen V, Schatz GE, Smith P, Turner I, Wyse-Jackson P, Crane PR (2005) Measuring the fate of plant diversity: towards a foundation for future monitoring and opportunities for urgent action. Philos Trans R Soc B 360:359–372. Scholar
  64. Ojeda F (1998) Biogeography of seeder and resprouter Erica species in the cape floristic region - where are the resprouters? Biol J Linn Soc 63:331–347Google Scholar
  65. Oliver EGH, Linder HP, Rourke JP (1983) Geographical distribution of present-day cape taxa and their phytogeographical significance. Bothalia 14:427–440CrossRefGoogle Scholar
  66. Peron G, Altwegg R (2015) Departures from the energy-biodiversity relationship in south African passerines: are the legacies of past climates mediated by behavioral constraints on dispersal? PLoS One 10:e0133992. Scholar
  67. Picker MD, Samways MJ (1996) Faunal diversity and endemicity of the cape peninsula, South Africa - a first assessment. Biodivers Conserv 5:591–606CrossRefGoogle Scholar
  68. Pio DV, Engler R, Linder HP, Monadjem A, Cotterill FPD, Taylor PJ, Schoeman MC, Price BW, Villet MH, Eick G, Salamin N, Guisan A (2014) Climate change effects on animal and plant phylogenetic diversity in southern Africa. Glob Chang Biol 20:1538–1549. Scholar
  69. Procheş Ş, Cowling RM (2006) Insect diversity in cape fynbos and neighbouring south African vegetation. Glob Ecol Biogeogr 15:445–451CrossRefGoogle Scholar
  70. Procheş Ş, Cowling RM, du Preez DR (2005) Patterns of geophyte diversity and storage organ size in the winter-rainfall region of southern Africa. Divers Distrib 11:101–109CrossRefGoogle Scholar
  71. Procheş Ş, Wilson JRU, Cowling RM (2006) How much evolutionary history in a 10x10 m plot? Proc R Soc Lond B Biol Sci 273(1590):1143–1148CrossRefGoogle Scholar
  72. Procheş Ş, Forest F, Veldtman R, Chown SL, Cowling RM, Johnson SD, Richardson DM, Savolainen V (2009) Dissecting the plant-insect diversity relationship in the cape. Mol Phylogenet Evol 51:94–99CrossRefPubMedGoogle Scholar
  73. Rosauer D, Laffan SW, Crisp MD, Donnellan SC, Cook LG (2009) Phylogenetic endemism: a new approach for identifying geographical concentrations of evolutionary history. Mol Ecol 18:4061–4072CrossRefPubMedGoogle Scholar
  74. Sharratt NJ, Picker MD, Samways MJ (2000) The invertebrate fauna of the sandstone caves of the cape peninsula (South Africa): patterns of endemism and conservation priorities. Biodivers Conserv 9:107–143CrossRefGoogle Scholar
  75. Smid J, Tolley KA (2017) The use of phylogenetic diversity for inferring biogeographical patterns - the case of south African reptiles. African Herp News 66:50–51Google Scholar
  76. Snijman DA (2013) Plants of the greater cape floristic region 2: the extra cape flora, Strelitzia 30. South African National Biodiversity Institute, PretoriaGoogle Scholar
  77. Switala AK, Sole CL, Scholtz CH (2014) Phylogeny, historical biogeography and divergence time estimates of the genus Colophon Gray (Coleoptera : Lucanidae). Invertebr Syst 28:326–336CrossRefGoogle Scholar
  78. Takhtajan A (1986) Floristic regions of the world. University of California Press, BerkeleyGoogle Scholar
  79. Thuiller W, Midgley GF, Rouget M, Cowling RM (2006) Predicting patterns of plant species richness in megadiverse South Africa. Ecography 29:733–744. Scholar
  80. Tolley KA, Chase BM, Forest F (2008) Speciation and radiations track climate transitions since the Miocene climatic optimum: a case study of southern African chameleons. J Biogeogr 35:1402–1414CrossRefGoogle Scholar
  81. Treurnicht M, Colville JF, Joppa LN, Huyser O, Manning JC (2017) Counting complete? Finalising the plant inventory of a global biodiversity hotspot. PeerJ 5:e2984. Scholar
  82. Tucker CM, Cadotte MW, Carvalho SB, Davies TJ, Ferrier S, Fritz SA, Grenyer R, Helmus MR, Jin LS, Mooers AO, Pavoine S, Purschke O, Redding DW, Rosauer DF, Winter M, Mazel F (2017) A guide to phylogenetic metrics for conservation, community ecology and macroecology. Biol Rev 92:698–715. Scholar
  83. Vane-Wright RI, Humphries CJ, Williams PH (1991) What to protect –systematics and the agony of choice. Biol Conserv 55:235–254CrossRefGoogle Scholar
  84. Verboom GA, Archibald JK, Bakker FT, Bellstedt DU, Conrad F, Dreyer LL, Forest F, Galley C, Goldblatt P, Henning JF, Mummenhoff K, Linder HP, Muasya AM, Oberlander KC, Savolainen V, Snijman DA, van der Niet T, Nowell TL (2009) Origin and diversification of the greater cape flora: ancient species repository, hot-bed of recent radiation, or both? Mol Phylogenet Evol 51:44–53CrossRefPubMedGoogle Scholar
  85. Verboom GA, Linder HP, Forest F, Hoffmann V, Bergh NG, Cowling RM (2014) Cenozoic assembly of the greater cape flora. In: Allsopp N, Colville JF, Verboom GA (eds) Fynbos: ecology, evolution, and conservation of a megadiverse region. Oxford University Press, Oxford, pp 93–118CrossRefGoogle Scholar
  86. Verboom GA, Bergh NG, Haiden SA, Hoffman V, Britton MN (2015) Topography as a driver of diversification in the cape floristic region of South Africa. New Phytol 207:368–376. Scholar
  87. Webb CO, Ackerly DD, McPeek MA, Donoghue MJ (2002) Phylogenies and community ecology. Annu Rev Ecol Syst 33:475–505CrossRefGoogle Scholar
  88. Weimarck H (1941) Phytogeographical groups, centres, and intervals within the cape Flora. Acta Universitatis Lundensis 37:1–143Google Scholar
  89. Werger MJA (1978) Biogeography and ecology of southern Africa. Dr W. Junk bv Publishers, The HagueCrossRefGoogle Scholar
  90. White F (1976) The vegetation map of Africa – the history of a completed project. Boissiera 24:659–666Google Scholar

Copyright information

© Springer International Publishing AG, part of Springer Nature 2018

Authors and Affiliations

  • Félix Forest
    • 1
    Email author
  • Jonathan F. Colville
    • 2
    • 3
  • Richard M. Cowling
    • 4
  1. 1.Royal Botanic GardensKew, RichmondUK
  2. 2.Kirstenbosch Research Centre, South African National Biodiversity InstituteClaremontSouth Africa
  3. 3.Statistics in Ecology, Environment and Conservation, Department of Statistical SciencesUniversity of Cape TownRondeboschSouth Africa
  4. 4.African Centre for Coastal PaleoscienceNelson Mandela Metropolitan UniversityPort ElizabethSouth Africa

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