The composition, geography, biology and assembly of the coastal flora of the Cape Floristic Region

The Cape Floristic Region (CFR) is globally recognized as a hotspot of plant diversity and endemism. Much of this diversity stems from radiations associated with infertile acid sands derived from sandstones of the geologically ancient Cape Fold Belt. These ancient montane floras acted as the source for most subsequent radiations on the Cape lowlands during the Oligocene (on silcretes) and Mio–Pliocene (on shales). The geomorphic evolution of the CFR during the Plio–Pleistocene led to the first large-scale occurrence of calcareous substrata (coastal dunes and calcarenites) along the Cape coast, providing novel habitats for plant colonization and ensuing evolution of the Cape coastal flora—the most recent diversification event in the Cape. Few studies have investigated the CFR’s dune and calcarenite floras, and fewer still have done so in an evolutionary context. Here, we present a unified flora of these coastal calcareous habitats of the CFR and analyze the taxonomic, biological and geographical traits of its component species to gain insights into its assembly. The Cape coastal flora, comprising 1,365 species, is taxonomically dominated by the Asteraceae, Fabaceae and Iridaceae, with Erica, Aspalathus and Agathosma being the most speciose genera. In terms of growth-form mix, there is a roughly equal split between herbaceous and woody species, the former dominated by geophytes and forbs, the latter by dwarf and low shrubs. Species associated with the Fynbos biome constitute the bulk of the flora, while the Subtropical Thicket and Wetland biomes also house a substantial number of species. The Cape coastal flora is a distinctly southern African assemblage, with 61% of species belonging to southern African lineages (including 35% of species with Cape affinity) and 59% being endemic to the CFR. Unique among floras from the Cape and coastal Mediterranean-climate regions is the relatively high proportion of species associated with tropical lineages, several of which are restricted to calcareous substrata of the CFR. The endemic, calcicolous component of the flora, constituting 40% of species, represents 6% of the Cape’s regional plant diversity—high tallies compared to other biodiversity hotspots. Most coastal-flora endemics emerged during the Plio–Pleistocene as a product of ecological speciation upon the colonization of calcareous substrata, with the calcifugous fynbos floras of montane acid substrata being the most significant source of this diversification, especially on the typically shallow soils of calcarenite landscapes. On the other hand, renosterveld floras, associated with edaphically benign soils that are widespread on the CFR lowlands, have not been a major source of lineages to the coastal flora. Our findings suggest that, over and above the strong pH gradient that exists on calcareous substrata, soil depth and texture may act as important edaphic filters to incorporating lineages from floras on juxtaposed substrata in the CFR.