Why is the Cape Peninsula so rich in plant species? An analysis of the independent diversity components

With 2285 species of higher plants crammed into 471 km², the flora of South Africa's Cape Peninsula is exceptionally rich. Similar sized areas in other Mediterranean-climate region biodiversity hot-spots support between 4.7 and 2.7 times fewer species. The high plant species richness of the Cape Peninsula is due to the exceptionally high turnover between moderately species-rich sites in different habitats (beta diversity) and between sites in similar habitats along geographical gradients (gamma diversity). Highest beta diversity, encompassing almost complete turnover, was recorded along soil fertility gradients. Although similar patterns for these independent components explain the richness of other regions in the Cape Floristic Region, it is the very long and steep habitat gradients of the Cape Peninsula that makes this region exceptionally rich. Furthermore, the flora is characterized by a high degree of rarity, a phenomenon that undoubtedly influences the turnover. Future research should focus on developing a biological and ecological understanding of the different forms of rarity and integrating this into management plans for the maintenance of biodiversity.     

The Cape Peninsula, South Africa: physiographical, biological and historical background to an extraordinary hot-spot of biodiversity

The Cape Peninsula, a 470 km² area of rugged scenery and varied climate, is located at the southwestern tip of the Cape Floristic Region, South Africa. The Peninsula is home to 2285 plant species and is a globally important hot-spot of biodiversity for higher plants and invertebrates. This paper provides a broad overview of the physiography, biological attributes and history of human occupation of the Peninsula. The Peninsula is characterized physiographically by extremely high topographical heterogeneity, very long and steep gradients in annual rainfall, and a great diversity of nutrient-poor soils. Thus, the Peninsula supports a high number of habitats and ecological communities. The predominant vegetation is fynbos, a fire-prone shrubland, and 12 broadly characterized fynbos types have been described on the Peninsula. Animal community structure, especially with regard to invertebrates, is poorly known. Vertebrate community structure is probably strongly influenced by nutrient poverty and recurrent fire. Generally, most vertebrates are small and typically occur in low numbers. Some invertebrates play keystone roles in facilitating ecological processes. Human occupation of the Peninsula was limited, until relatively recently, by nutrient poverty. After Dutch colonization in 1652, direct and indirect impacts on the natural ecosystems of the Peninsula escalated dramatically, and by 1994, some 65% of original natural habitat was either transformed by urbanization and agriculture, or invaded by alien plants. Nonetheless, there is still excellent potential to conserve the Cape Peninsula's remaining biodiversity.     

Aquatic invertebrates in riverine landscapes

1. Riverine systems consist of a mosaic of patches and habitats linked by diverse processes and supporting highly complex communities. Invertebrates show a high taxonomic and functional diversity in riverine systems and are in several ways important components of these systems. Their distribution patterns, movements and effects on ecological flows, testify to their importance in various landscape ecological processes. This paper reviews the invertebrate literature with respect to patterns and processes in the riverine landscape. 2. The distribution of invertebrates in riverine habitats is governed by a number of factors that typically act at different scales. Hence, the local community structure can be seen as the result of a continuous sorting process through environmental filters ranging from regional or catchment‐wide processes, involving speciation, geological history and climate, to the small‐scale characteristics of individual patches, such as local predation risk, substratum porosity and current velocity. 3. Dispersal is an important process driving invertebrate distribution, linking different ecological systems across boundaries. Dispersal occurs within the aquatic habitat as well as into the terrestrial surrounding, and also over land to other waterbodies. New genetic techniques have contributed significantly to the understanding of aquatic invertebrate dispersal and revealed the importance of factors such as physical barriers, synchrony of emergence and taxonomic affiliation. 4. Invertebrates affect the cycling of nutrients and carbon by being a crucial intermediate link between primary producers, detritus pools or primary consumers, and predators higher up in the trophic hierarchy. Suspension feeders increase the retention of carbon. The subsidies of aquatic invertebrates to the terrestrial ecosystem have been shown to be important, as are reciprocal processes such as the supply of terrestrial invertebrates that fall into the water. 5. Future studies are needed both to advance theoretical aspects of landscape ecology pertaining to the invertebrates in riverine systems and to intensify the experimental testing of hypotheses, for example with respect to the scaling of processes and to linkages between the terrestrial and aquatic systems. Another promising avenue is to take advantage of naturally steep environmental gradients, and of systems disturbed by humans, such as regulated rivers. By comparison with unimpaired reference sites, the mechanisms involved might be identified. The use of `natural' experiments, especially where environmental gradients are steep, is another technique with great potential.     

An overview of the Cape geophytes

The Cape Region (here treated as the winter rainfall region of southern Africa, thus including fynbos, renosterveld and succulent karoo vegetation) is the world's foremost centre of geophyte diversity. Some 2100 species in 20 families have been recorded from this area, 84% of them endemic. The most important families, with more than a hundred geophyte species each, are Iridaceae, Oxalidaceae, Hyacinthaceae, Orchidaceae, Amaryllidaceae and Asphodelaceae. Although southern Africa does not appear to have been the main diversification centre for the plant orders with highest geophyte representation (Asparagales and Liliales), it represents an active centre of transition to geophytism, such transitions having occurred independently in numerous plant groups, often followed by rapid speciation. Several Cape geophyte groups have consequently expanded across Africa to the Mediterranean Basin, and possibly to other winter rainfall regions. Remarkably high local species diversity in renosterveld vegetation, even in relatively homogeneous environments, suggests that pollinator specificity and phenology play an important role in niche partitioning. However, character diversity is also high in storage organs and leaves, and this could account for the high species diversity values recorded at larger spatial scales, especially across environmental gradients. Long-term climatic stability, combined with topoclimatic and edaphic diversity and regular fire occurrence, is likely to be responsible for the remarkable geophyte diversity of the Cape, as compared to other mediterranean-climate regions.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 27–43.     

Exceptionally poor land snail fauna of central Yakutia (NE Russia): climatic and habitat determinants of species richness

There is an ongoing debate on the causes of the latitudinal diversity gradient, but diversity decline towards high latitudes is poorly documented for many invertebrate taxa. Therefore, we sampled land snail assemblages at 79 sites and in various habitat types in central Yakutia, a region with extremely continental, cool and dry climate. We tested whether habitats lacking suitable shelters for winter survival harbour less species than those with vegetation cover that softens climatic extremes. Both local species diversity and regional species diversity were extremely low: 13 species were recorded in total with an average of 1.4 species per site. While the majority of grassland sites were without snails (26 of 34 sites), forest sites supported at least one snail species in most cases (38 of 45 sites). Within grasslands, snail occurrences were associated with a higher herb-layer biomass. Numbers of snail species correlated with the amount of available calcium only in forests, in which species accumulation towards more favourable habitats was possible due to softening of climate harshness. As minute snails are known to be effective passive dispersers and the study area was not glaciated during the last glacial stage, there was certainly enough time for colonization of all favourable habitats. Our results suggest climatically driven limitations of both local and regional land snail diversity in central Yakutia. We conclude that the hypothesis of climate harshness remains the most probable explanation of a sharp drop in land snail diversity in high-latitude areas with cold climate.     

The contribution of edaphic heterogeneity to the evolution and diversity of Burseraceae trees in the western Amazon

Environmental heterogeneity in the tropics is thought to lead to specialization in plants and thereby contribute to the diversity of the tropical flora. We examine this idea with data on the habitat specificity of 35 western Amazonian species from the genera Protium, Crepidospermum, and Tetragastris in the monophyletic tribe Protieae (Burseraceae) mapped on a molecular-based phylogeny. We surveyed three edaphic habitats that occur throughout terra firme Amazonia: white-sand, clay, and terrace soils in eight forests across more than 2000 km in the western Amazon. Twenty-six of the 35 species were found to be associated with only one of three soil types, and no species was associated with all three habitats; this pattern of edaphic specialization was consistent across the entire region. Habitat association mapped onto the phylogenetic tree shows association with terrace soils to be the probable ancestral state in the group, with subsequent speciation events onto clay and white-sand soils. The repeated gain of clay association within the clade likely coincides with the emergence of large areas of clay soils in the Miocene deposited during the Andean uplift. Character optimizations revealed that soil association was not phylogenetically clustered for white-sand and clay specialists, suggesting repeated independent evolution of soil specificity is common within the Protieae. This phylogenetic analysis also showed that multiple cases of putative sister taxa with parapatric distributions differ in their edaphic associations, suggesting that edaphic heterogeneity was an important driver of speciation in the Protieae in the Amazon basin.     

Explaining the uniqueness of the Cape flora: incorporating geomorphic evolution as a factor for explaining its diversification

The plant diversity of the Cape Floristic Region is regarded as being exceptional in an ecological and evolutionary context. The region supports about double the number of species predicted by models based on water-energy variables for regional floras globally. However, contemporary diversity patterns are profoundly influenced by evolutionary processes contingent upon idiosyncrasies of history and geography. The relatively recent appearance of dated molecular phylogenies, and their optimization in relation to habitat and geography, has provided hitherto unsurpassed opportunities to generate knowledge about the evolution of the Cape flora. Almost all studies invoke climatic deterioration during the Mio-Pliocene as the major trigger of radiations and subsequent speciation of Cape clades. While some do show the importance of edaphic heterogeneity for clade radiation, the evolution of this heterogeneity is not considered. Here, we review the literature on the late Cenozoic geomorphic evolution of the Cape in order to assess the extent to which the changing nature of scenery and soils could act as a stimulus for plant diversification. Despite dating uncertainties associated with both the phylogenetic and geomorphic data, it appears that moderate uplift in the early and late Miocene, which significantly increased the topo-edaphic heterogeneity of the Cape was an important driver of plant diversification. In particular, the massive increase in heterogeneity after the late Miocene event probably acted in synergy with rapid climatic deterioration, to produce the extraordinarily rapid diversification recorded for some Cape clades at that time. A comparison of the plant diversity and palaeoenvironmetal patterns of mediterranean-climate regions provide insights regarding the "remarkable environmental conditions" of the Cape that have generated the high diversification and low extinction rates necessary to produce such a rich flora. These conditions are a gradual increase in topo-edaphic heterogeneity and relative climatic stability during the late Cenozoic.     

THE CONTRIBUTION OF EDAPHIC HETEROGENEIYT TO THE EVOLUTION AND DIVERSITY OF BURSERACEAR TREES IN THE WESTERN AMAZON

Abstract —Environmental heterogeneity in the tropics is thought to lead to specialization in plants and thereby contribute to the diversity of the tropical flora. We examine this idea with data on the habitat specificity of 35 western Amazonian species from the genera Protium, Crepidospermum, and Tetragastris in the monophyletic tribe Protieae (Burseraceae) mapped on a molecular‐based phylogeny. We surveyed three edaphic habitats that occur throughout terra firme Amazonia: white‐sand, clay, and terrace soils in eight forests across more than 2000 km in the western Amazon. Twenty‐six of the 35 species were found to be associated with only one of three soil types, and no species was associated with all three habitats; this pattern of edaphic specialization was consistent across the entire region. Habitat association mapped onto the phylogenetic tree shows association with terrace soils to be the probable ancestral state in the group, with subsequent speciation events onto clay and white‐sand soils. The repeated gain of clay association within the clade likely coincides with the emergence of large areas of clay soils in the Miocene deposited during the Andean uplift. Character optimizations revealed that soil association was not phylogenetically clustered for white‐sand and clay specialists, suggesting repeated independent evolution of soil specificity is common within the Protieae. This phylogenetic analysis also showed that multiple cases of putative sister taxa with parapatric distributions differ in their edaphic associations, suggesting that edaphic heterogeneity was an important driver of speciation in the Protieae in the Amazon basin.     

Tree Diversity,Forest Structure and Productivity along Altitudinal and Topographical Gradients in a Species-Rich Ecuadorian Montane Rain Forest

We studied the spatial heterogeneity of tree diversity, and of forest structure and productivity in a highly diverse tropical mountain area in southern Ecuador with the aim of understanding the causes of the large variation in these parameters. Two major environmental gradients, elevation and topography, representing a broad range of climatic and edaphic site conditions, were analyzed. We found the highest species richness of trees in valleys <2100 m. Valleys showed highest values of basal area, leaf area index and tree basal area increment as well. Tree diversity also increased from ridges to valleys, while canopy openness decreased. Significant relationships existed between tree diversity and soil parameters (pH, total contents of Mg, K, Ca, N and P), and between diversity and the spatial variability of pH and Ca and Mg contents suggesting a dependence of tree diversity on both absolute levels and on the small-scale heterogeneity of soil nutrient availability. Tree diversity and basal area increment were positively correlated, partly because both are similarly affected by soil conditions. We conclude that the extraordinarily high tree species richness in the area is primarily caused by three factors: (1) the existence of steep altitudinal and topographic gradients in a rather limited area creating a small-scale mosaic of edaphically different habitats; (2) the intermingling of Amazonian lowland plant species, that reach their upper distribution limits, and of montane forest species; and (3) the geographical position of the study area between the humid eastern Andean slope and the dry interandean forests of South Ecuador.     

Distribution of the endemic Balkan flora on serpentine I. – obligate serpentine endemics

Serpentine (ophiolithic) substrate covers large areas in the Balkans, more so than in any other part of Europe. These areas extend from north to south mainly in the mountainous central regions and represent specialized habitats for basiphilous-calcifugal plants. Biodiversity in the area is high, with a great number of interesting local and regional endemics. The high number of endemics indicates the importance of serpentine habitats as centres for floristic differentiation and speciation. The number of Balkan endemics growing on serpentine is c. 335 taxa (species and subspecies) of which 123 are obligate. Their distribution is presented in 50 × 50 km UTM squares as adopted in the Atlas Florae Europaeae project coordinated at Helsinki. The richest (in number of taxa) squares are situated in NW Greece (Epirus), the island of Evvia, N Albania together with SW Serbia, and N Greece (Vourinos). They indicate important centres of plant diversity in the Balkans, areas to be noted for conservation strategy. Features responsible for the distribution and abundance of these obligate serpentine endemics include: 1) edaphic isolation in relation to type of bedrock (lime, dolomite, marble, schist, etc.), 2) mountain island isolation (Smolikas, Vourinos, Ostrovica, etc.), 3) island isolation (Evvia) and 4) continuous long-term isolation without interruption or disturbance of speciation.     

Potential impact of viticulture expansion on habitat types in the Cape Floristic Region, South Africa

The wine industry in the Western Cape, South Africa has expanded over the past decade, particularly since the lifting of trade sanctions in 1992. Wine grapes are cultivated on fertile soils upon which threatened biodiversity habitat units of the Cape Floristic Region occur naturally. There is a concern as to whether further expansion of the wine industry, which would benefit the economy through increased foreign exchange, would encroach on the little remaining vegetation in vineyard-producing areas. Predictive land use modeling using logistic regression techniques was applied to determine suitable areas for vineyard cultivation according to climatic, topographic, and soil/geology variables. Of the most threatened habitats, 14849 hectares are particularly suitable for vineyards. Breede fynbos/renosterveld mosaic was the habitat most likely to be converted, and was considered 89.3% irreplaceable to current conservation goals. Also vulnerable are Ashton inland renosterveld and Boland coast renosterveld, the latter being 100% irreplaceable. Although the high rate in vine replanting suggests that the need for untransformed land will not be great immediately, an economic analysis showed that protection of these areas against future ploughing will be vital if targets of adequately representing each habitat in the Cape Floristic Region are to be met. Land use change modeling, especially if done in a spatially explicit and integrated manner with expert input, was shown to be an important technique for the extrapolation of historical patterns to understand the forces that shape landscapes, allowing for the assessment of management alternatives, and testing our understanding of key processes in land use changes that effect conservation planning.     

The Role of Nitrogen Deposition in Widespread Plant Community Change Across Semi-natural Habitats

Experimental studies have shown that deposition of reactive nitrogen is an important driver of plant community change, however, most of these experiments are of short duration with unrealistic treatments, and conducted in regions with elevated ambient deposition. Studies of spatial gradients of pollution can complement experimental data and indicate whether the potential impacts demonstrated by experiments are actually occurring in the ‘real world’. However, targeted surveys exist for only a very few habitats and are not readily comparable. In a coordinated campaign, we determined the species richness and plant community composition of five widespread, semi-natural habitats across Great Britain in sites stratified along gradients of climate and pollution, and related these ecological parameters to major drivers of biodiversity, including climate, pollution deposition, and local edaphic factors. In every habitat, we found reduced species richness and changed species composition associated with higher nitrogen deposition, with remarkable consistency in relative species loss across ecosystem types. Whereas the diversity of mosses, lichens, forbs, and graminoids declines with N deposition in different habitats, the cover of graminoids generally increases. Considered alongside previous experimental studies and survey work, our results provide a compelling argument that nitrogen deposition is a widespread and pervasive threat to terrestrial ecosystems.     

Orchid biogeography and factors associated with rarity in a biodiversity hotspot,the Southwest Australian Floristic Region

Aim The causes of orchid diversification and intrinsic rarity are poorly resolved. The Orchidaceae of the Southwest Australian Floristic Region use a diversity of pollination strategies and sites of mycorrhizal infection, and occupy a diversity of habitats. We combined a biogeographic analysis with analysis of factors associated with rarity to establish: (1) the landscape features correlated with taxon turnover and speciation, and (2) the possible role in taxon rarity of geographic region, pollination strategy, edaphic habitat and site of mycorrhizal infection. Location Southwest Australian Floristic Region. Methods The distributions of 407 orchid taxa (species and subspecies) were mapped at the quarter‐degree scale using 13,267 collections in the Western Australian Herbarium. This database was used to map taxon richness, for a biogeographic analysis and to quantify rarity of taxa. Using herbarium records, rarity was expressed as mean abundance, mean distribution and incidence of rarity based on abundance and distribution for each genus. We tested for differences in rarity of species between pollination strategies, edaphic habitats and sites of mycorrhizal infection. Results Taxon richness was highest in the High Rainfall Province. Biogeographic provincial boundaries for orchids were aligned with rainfall, while district boundaries tended to follow geological formations. When rarity was defined as either low abundance or small distribution, the greatest number of rare taxa occurred in areas of high taxon richness and naturally fragmented edaphic environments. For both abundance and distributional extent, sexual deception had a significantly higher incidence of rarity than food‐rewarding taxa. There was no significant difference in rarity with site of mycorrhizal infection. Main conclusions While large‐scale edaphic and climatic variation are correlated with orchid taxon turnover and speciation in a similar fashion to the flora in general, the processes responsible for patterns of diversity may differ. Fragmented edaphic environments appear to be associated with a higher incidence of rare species due to limited dispersal/colonization opportunities or radiations of taxa in allopatry. The high incidence of rarity in sexually deceptive taxa could be due to either low fruit set or the risk of specializing on a single pollinator species.     

Multi-scale determinants of dung beetle assemblage structure across abiotic gradients of the Kalahari–Nama Karoo ecotone, South Africa

Aim Changing conditions across spatial gradients are primary determinants of biotic regions, local habitats, and distributional edges. We investigate how a climatic gradient and edaphic mosaic interact as multi‐scale drivers of spatial patterns in scarabaeine dung beetles. The patterns are tested for congruency with ecoregion and floral boundaries over the same gradient, as responses to physical factors often differ among higher taxa. Location Southern Africa and the Nama Karoo–Kalahari ecotone, Northern Cape, South Africa. Methods Data consisted of the climatic distributions of 104 species and their abundances at 223 sites in two ecoregions/floral biomes, four bioregions, and 13 vegetation units. Factor analyses determined the biogeographical composition of the species, and regional‐ to local‐scale patterns in species abundance structure. Hierarchical analysis of oblique factors determined the proportional contribution of spatial variance to patterns. One‐way anova was used to test for significant separation of patterns along factor axes. Stepwise multiple regression was used to determine correlations of five physical attributes with species richness, Shannon‐Wiener diversity, and factor loadings for the study sites. Results Four biogeographical influences overlap in the study region, although rank contribution declines from south‐west arid through north‐east savanna to widespread and south‐east highland taxa. Species abundance structure comprises five subregional patterns, two centred to the north‐east (Kalahari, Isolated Kalahari Dune) dominated by Kalahari influence, and three to the south‐west (Nama Karoo subdivisions: Bushmanland, ‘Upper’, ‘Stony Prieska’) dominated by south‐west arid influence. Kalahari deep sands are characterized especially by a warmer, moister climate, whereas the Nama Karoo mosaic of deep or stony soils is characterized especially by north‐west aridity (Bushmanland), south‐east cooler temperatures (‘Upper’), or excessively stony soils (‘Stony Prieska’). Four of the subregional patterns each comprised three localized patterns related primarily to relative stoniness, edge effects from geographical position, or incidence of rainfall. Species richness and diversity declined with decreasing rainfall and increasing stoniness. Main conclusions Climatic and edaphic factors are important multi‐scale determinants of spatial patterns in dung beetle assemblage structure, with edaphic factors becoming more important at local spatial scales. The patterns are roughly congruent with the Kalahari Savanna–Nama Karoo ecotone at the floral biome or ecoregion scale, but show limited coincidence with finer‐scale floral classification.     

Edaphic niches of metallophytes from southeastern Democratic Republic of Congo: Implications for post-mining restoration

In southeastern D. R. Congo, about 550 metallophytes grow on soils with high copper and cobalt concentrations, 57 of which are endemics to these metalliferous environments. About 70% of those endemics are considered threatened by destruction of habitats through mining activities. To provide guidelines for future restoration programs, the edaphic ecological niches of eight endemic metallophytes (i.e. copper endemics) were studied using a pragmatic sampling method adapted for urgent conservation needs. Niches were modelled using violin plot along Cu, Co and C:N gradients representing the two main independent edaphic gradients among nine edaphic variables (C, N, C:N, K, P, pH, Co, Cu, and Mn). Copper endemics presented distinct edaphic niches along the copper and cobalt gradients, but differentiation was lower along the C:N gradient. In addition, edaphic elements presented covariations among them and metalliferous soils had higher nutrient and element content compared to the non-metalliferous soils of the region dominated by the Miombo woodland. The complexity of the soil composition and the edaphic niches of copper endemics revealed an important challenge in the implementation of the species conservation and the habitat restoration strategies of post-mining sites.     

Endemism and speciation in a lowland flora from the Gape Floristic Region

Taxonomic, edaphic and biological aspects of endemism were studied in a phanerogamous flora from the Agulhas Plain, a coastal lowland area of the Cape Floristic Region. Of the 1751 species in the flora, 23.6% were regional endemics and 5.7% were local endemics. Families which were over-represented in terms of endemics included the Ericaceae, Rutaceae, Proteaceae and Polygalaceae. Under-represented families included the Poaceae, Cyperaceae, Scrophulariaceae and Orchidaceae. Highest levels of local endemism were recorded on limestone and colluvial acid sand. Sixty-nine percent of regional endemics and 85% of local endemics were confined to a single substratum. An analysis of the frequency of biological traits associated with species with different categories of endemism enabled the establishment of a biological profile of a local endemic: a dwarf to low, non-sprouting shrub with soil stored seeds which are ant-dispersed and/or form a symbiotic relationship with microbes. It is argued that lineages with these characteristics are vulnerable to severe population reduction or even local extinction. An effect of this would be the promotion of rapid, edaphic speciation as a result of catastrophic selection. Thus, certain traits (e.g. non-sprouting) prevail or even predominate in the flora not because of any adaptive advantage but because high speciation rates of lineages which possess them, overwhelm low survival rates.     

Are forest‐shrubland mosaics of the Cape Floristic Region an example of alternate stable states?

The idea of alternate stable states (ASS) has been used to explain the juxtaposition of distinct vegetation types within the same climate regime. ASS may explain the co‐existence of relatively inflammable closed‐canopy Afrotemperate forest patches (‘Forest’) within fire‐prone open‐canopy Fynbos in the Cape Floristic Region (CFR) on sandstone‐derived soils. We evaluated the hypothesis that although fire and local topography and hydrology likely determined the paleogeographic boundaries of Forest, present‐day boundaries are additionally imposed by emergent edaphic properties and disturbance histories. We studied vegetation and edaphic properties of Forest‐Transition‐Fynbos vegetation at two sites within the CFR on sandstone‐derived soils and tracked historical change using aerial photography. Whereas Forest and Fynbos have changed little in extent or density since 1945, transition vegetation increased into areas formerly occupied by Fynbos. Forest soils were ubiquitously more nutrient‐rich than Fynbos soils, with transition soils being intermediate. These edaphic differences are not due to geological differences, but instead appear to have emerged as a consequence of different nutrient cycling within the different ecosystems. Soil nutrients are now so different that a switch from Fynbos to Forest is unlikely, in the short term (i.e. decades). Floristically and nutritionally, transitional vegetation is more similar to Fynbos than Forest and may be less resilient to changes in exogenous drivers (e.g. fire). Our findings are consistent with the idea that geologically Forest and Fynbos are largely fire‐derived long‐term ASS, with the stability of each state reinforced by marked soil nutrient differences. In contrast, the intermediate transitional vegetation that might switch states is unlikely to be stable.     

Biodiversity and environmental stability

Although levels of biological diversity may seem to be equivalent in different areas, diversity is created and maintained by a range of different ]processes: overlap of habitat on gradients; a dynamic mosaic of communities; and accumulation and evolution of taxa in extremely stable areas. These different communities will respond in very different ways to disturbance. The most fragile are those whose component taxa are genetically adapted to the stability of a predictable environment. These areas are often under pressure from local rural populations and require intensive local conservation management actions. In other areas, where diversity is adapted to dynamism, communities are more resilient to disturbance and conservation can be best effected by policy instruments.     

Frequent and parallel habitat transitions as driver of unbounded radiations in the Cape flora

The enormous species richness in the Cape Floristic Region (CFR) of Southern Africa is the result of numerous radiations, but the temporal progression and possible mechanisms of these radiations are still poorly understood. Here, we explore the macroevolutionary dynamics of the Restionaceae, which include 340 species that are found in all vegetation types in the Cape flora and are ecologically dominant in fynbos. Using an almost complete (i.e., 98%) species‐level time calibrated phylogeny and models of diversification dynamics, we show that species diversification is constant through the Cenozoic, with no evidence of an acceleration with the onset of the modern winter‐wet climate, or a recent density‐dependent slowdown. Contrary to expectation, species inhabiting the oldest (montane) and most extensive (drylands) habitats did not undergo higher diversification rates than species in the younger (lowlands) and more restricted (wetland) habitats. We show that the rate of habitat transitions is more closely related to the speciation rate than to time, and that more than a quarter of all speciation events are associated with habitat transitions. This suggests that the unbounded Restionaceae diversification resulted from numerous, parallel, habitat shifts, rather than persistence in a habitat stimulating speciation. We speculate that this could be one of the mechanisms resulting in the hyperdiverse Cape flora.     

The radiation of the Cape flora, southern Africa

The flora of the south-western tip of southern Africa, the Cape flora, with some 9000 species in an area of 90,000 km2 is much more speciose than can be expected from its area or latitude, and is comparable to that expected from the most diverse equatorial areas. The endemism of almost 70%, on the other hand, is comparable to that found on islands. This high endemism is accounted for by the ecological and geographical isolation of the Cape Floristic Region, but explanations for the high species richness are not so easily found. The high species richness is accentuated when its taxonomic distribution is investigated: almost half of the total species richness of the area is accounted for by 33 'Cape floral clades'. These are clades which may have initially diversified in the region, and of which at least half the species are still found in the Cape Floristic Region. Such a high contribution by a very small number of clades is typical of island floras, not of mainland floras. The start of the radiation of these clades has been dated by molecular clock techniques to between 18 million years ago (Mya) (Pelargonium) and 8 Mya (Phylica), but only six radiations have been dated to date. The fossil evidence for the dating of the radiation is shown to be largely speculative. The Cenozoic environmental history of southern Africa is reviewed in search of possible triggers for the radiations, climatic changes emerge as the most likely candidate. Due to a very poor fossil record, the climatic history has to be inferred from larger scale patterns, these suggest large-scale fluctuations between summer wet (Palaeocene, Early Miocene) and summer dry climates (Oligocene, Middle Miocene to present). The massive speciation in the Cape flora might be accounted for by the diverse limitations to gene flow (dissected landscapes, pollinator specialisation, long flowering times allowing much phenological specialisation), as well as a richly complex environment providing a diversity of selective forces (geographically variable climate, much altitude variation, different soil types, rocky terrain providing many micro-niches, and regular fires providing both intermediate disturbances, as well as different ways of surviving the fires). However, much of this is based on correlation, and there is a great need for (a) experimental testing of the proposed speciation mechanisms, (b) more molecular clock estimates of the age and pattern of the radiations, and (c) more fossil evidence bearing on the past climates.