Experimental biogeography: the role of environmental gradients in high geographic diversity in Cape Proteaceae

One of the fundamental dimensions of biodiversity is the rate of species turnover across geographic distance. The Cape Floristic Region of South Africa has exceptionally high geographic species turnover, much of which is associated with groups of closely related species with mostly or completely non-overlapping distributions. A basic unresolved question about biodiversity in this global hotspot is the relative importance of ecological gradients in generating and maintaining high geographic turnover in the region. We used reciprocal transplant experiments to test the extent to which abiotic environmental factors may limit the distributions of a group of closely related species in the genus Protea (Proteaceae), and thus elevate species turnover in this diverse, iconic family. We tested whether these species have a “home site advantage” in demographic rates (germination, growth, mortality), and also parameterized stage-structured demographic models for the species. Two of the three native species were predicted to have a demographic advantage at their home sites. The models also predicted, however, that species could maintain positive population growth rates at sites beyond their current distribution limits. Thus the experiment suggests that abiotic limitation under current environmental conditions does not fully explain the observed distribution limits or resulting biogeographic pattern. One potentially important mechanism is dispersal limitation, which is consistent with estimates based on genetic data and mechanistic dispersal models, though other mechanisms including competition may also play a role. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A densely sampled ITS phylogeny of the Cape flagship genus Erica L. suggests numerous shifts in floral macro-morphology

Erica L. is the largest of the ‘Cape’ clades that together comprise around half of the disproportionately high species richness of the Cape Floristic Region (CFR) of South Africa. Around 840 species of Erica are currently recognised, C.680 of which are found in the CFR, the rest distributed across the rest of Southern Africa, the highlands of Tropical Africa and Madagascar, and Europe. Erica is taxonomically well documented, but very little is known about species-level relationships. We present the first densely sampled phylogenetic analysis of Erica, using nuclear ribosomal DNA sequences (internal transcribed spacers; ITS) of c. 45% of the species from across the full geographic range of the genus, both Calluna and Daboecia (Ericeae; monotypic genera and putative sister groups of Erica), and further Ericoideae outgroups. Our results show both morphological and geographic coherence of some clades, but numerous shifts in floral macro-morphology as represented by variation in individual morphological characters and pollination syndromes. European Ericeae is a paraphyletic grade subtending a monophyletic African/Malagasy Erica. Given the limited resolution of this single gene tree, more data are needed for further conclusions. Clades are identified that will serve as an effective guide for targeted sampling from multiple linkage groups.     

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.     

Pioneers and Perches—Promising Restoration Methods for Degraded Renosterveld Habitats?

Areas of abandoned agricultural fields are globally increasing and are also common features in the Cape Lowlands of South Africa. Previous restoration attempts in degraded West Coast renosterveld, a Mediterranean‐climate shrubland, have attained limited success and therefore novel approaches are needed for this area. The study reports on two restoration experiments, designed to re‐introduce key plant functional types back into this critically endangered habitat. The first experiment concentrated on a common pioneer species in renosterveld vegetation, Otholobium hirtum. Although in vitro experiments showed a significantly elevated germination response after scarification, in vivo experiments failed to produce establishment in an abandoned field. The second restoration experiment focused on bush clumps, a sub‐type of renosterveld vegetation that is characterized by broad‐leaved shrubs with fleshy bird‐dispersed diaspores. The effect of artificial bird perches and their potential to enhance diaspore dispersal by frugivorous birds in two abandoned field communities was tested. Results showed a significant increase in seed dispersal at artificial perch sites. However, in the next fruiting season, and after perch removal, seed germination and establishment in abandoned fields was not successful. The experiments revealed that restoration using early‐succession species and natural dispersal vectors appear not to produce demonstrable benefits, despite their promising potential and pre‐testing of effectiveness. Before launching large‐scale restoration programs in abandoned fields of renosterveld, preliminary studies in‐field are strongly recommended.     

Are Cape floral clades the same age? Contemporaneous origins of two lineages in the genistoids s.l. (Fabaceae)

The hypothesis that the elements of the modern species-rich flora of the Cape Floristic Region (CFR), South Africa, originated more or less simultaneously at the Miocene/Pliocene boundary, in response to the development of a mediterranean climate, has been challenged by numerous molecular dating estimates of Cape floral clades. These studies reveal a more gradual emergence, with the oldest clades originating in the Eocene, but others appearing later, some as recently as the Pliocene. That there are factors which might affect the dates recovered, such as choice of calibration point, analysis method, sampling density and the delimitation of Cape floral clades, suggests a need for further critical evaluation of the age estimates presented to date. In this study, the dates of origin of two Cape floral clades (the legume Crotalarieae p.p. and Podalyrieae) are estimated, constrained by a shared calibration point in a single analysis using an rDNA ITS phylogeny in which 633 taxa are sampled. The results indicate that these two clades arose contemporaneously 44-46 mya, not at the Miocene/Pliocene boundary as had been previously supposed. The contemporaneous origin of these Cape floral clades suggests that additional more inclusive analyses are needed before rejecting the hypothesis that a single environmental trigger explains the establishment of Cape floral clades.     

Impact of a dam on benthic macroinvertebrates in a small river in a biodiversity hotspot: Cape Floristic Region, South Africa

Suitable reservoirs and monitoring methods are needed to manage scarce water supplies in dry countries. We assessed here the impact on aquatic macroinvertebrates of the only dam on the Eerste River, which runs through the heart of a biodiversity hotspot, the Cape Floristic Region, South Africa. The dam and associated activities, were the only forms of disturbance in this otherwise pristine area. We sampled over 20,000 macroinvertebrate individuals and illustrated some categorical effects of the impoundment and its effects on macroinvertebrate assemblages. Macroinvertebrate species diversity below the dam was only half of that in the pristine catchment area above the dam. Furthermore, Ephemeroptera, Plecoptera and Trichoptera diversity and abundance dropped to almost zero as a result of the impoundment. In contrast, the abundance of the Diptera family Chironomidae increased substantially below the dam. These changes in macroinvertebrate diversity mirrored those recorded in biologically less diverse areas, but are of major concern in this biodiversity hotspot with its rich endemic fauna. We conclude that such an impoundment, while important for human welfare, results in a high price being paid in terms of loss of local biodiversity.     

New species and combinations in the African Restionaceae

Eight new species of the African Restionaceae (Restionoideae) are described, viz.: Cannomois anfracta, Cannomois arenicola, Cannomois grandis, Nevillea vlokii, Thamnochortus kammanassiae, Willdenowia pilleata, Restio uniflorus and Restio mkambatiae. A key to the species of Cannomois is provided, as well as a table comparing the characters of the three species in Nevillea. For all new species, notes on the affinities of the species and their habitats are provided. Two new combinations, Cannomois primosii (Pillans) H.P. Linder and Cannomois robusta (Kunth) H.P. Linder, are made.     

Montane Refugia for Endemic and Red Listed Dragonflies in the Cape Floristic Region Biodiversity Hotspot

One of the features of many endemic organisms is that they are highly spatially restricted, and habitat specialists. The Kogelberg Biosphere Reserve (KBR) is a major centre of plant endemism within a global hotspot, the Cape Floristic Region (CFR). Dragonflies in this botanical hotspot have a range of habitat specialization from narrow-range specialists to widespread generalists, with an unusually strong bias towards the specialists. A huge 53% of dragonfly individuals and 26% of taxa recorded are national endemics, and three species are Red Listed. Thus, a group of predatory insects, which are largely not dependent on plant composition, mirrors the level of habitat specialization and restricted distributions of the plants at the spatial scale of the whole reserve. Although some studies caution the use of one taxon as a surrogate for another, the results here show that at the reserve scale in this global hotspot there can be remarkable concordance, suggesting further studies on other taxa should be carried out to determine the full extent of taxonomic concordance in this irreplaceable area.     

Biogeographic patterns and phylogeography of dwarf chameleons (Bradypodion) in an African biodiversity hotspot

The southern African landscape appears to have experienced frequent shifts in vegetation associated with climatic change through the mid-Miocene and Plio-Pleistocene. One group whose historical biogeography may have been affected by these fluctuations are the dwarf chameleons (Bradypodion), due to their associations with distinct vegetation types. Thus, this group provides an opportunity to investigate historical biogeography in light of climatic fluctuations. A total of 138 dwarf chameleons from the Cape Floristic Region of South Africa were sequenced for two mitochondrial genes (ND2 and 16S), and resulting phylogenetic analyses showed two well-supported clades that are distributed allopatrically. Within clades, diversity among some lineages was low, and haplotype networks showed patterns of reticulate evolution and incomplete lineage sorting, suggesting relatively recent origins for some of these lineages. A dispersal-vicariance analysis and a relaxed Bayesian clock suggest that vicariance between the two main clades occurred in the mid-Miocene, and that both dispersal and vicariance have played a role in shaping present-day distributions. These analyses also suggest that the most recent series of lineage diversification events probably occurred within the last 3-6 million years. This suggests that the origins of many present-day lineages were founded in the Plio-Pleistocene, a time period that corresponds to the reduction of forests in the region and the establishment of the fynbos biome.     

Patterns of endemism in the limestone flora of South African lowland fynbos

Taxonomie and biological aspects of endemism and Red Data Book status were studied amongst the limestone endemics of the lowland fynbos in the Cape Floristic Region, South Africa. Of the 110 limestone endemics, 1.8% are widely distributed in the Cape Floristic Region and 56.4% are regional endemics. Relative to flora of non-limestone lowland fynbos (n=538 species), the families which were overrepresented in terms of limestone endemics included the Ericaceae, Fabaceae, Polygalaceae, Rutaceae and Sterculiaceae. The Restionaceae was the only underrepresented family. The local limestone endemics were not significantly different from regional endemics in terms of their biological attributes. An analysis of the frequency of the biological traits associated with the limestone-endemic flora established a biological profile for a limestone endemic: a dwarf-to-low shrub with soil-stored seeds which are ant or wind dispersed. In terms of the species richness of limestone endemics, the De Hoop Nature Reserve was the hotspot within the region. Relative to the total species richness, the Hagelkraal and Stilbaai areas contained higher-than-predicted numbers of rare species. These areas require urgent attention if the unique floral diversity associated with limestone substrata within the Bredasdorp-Riversdale centre of endemism is to be conserved.     

High ploidy diversity and distinct patterns of cytotype distribution in a widespread species of Oxalis in the Greater Cape Floristic Region

Background and Aims

Genome duplication is widely acknowledged as a major force in the evolution of angiosperms, although the incidence of polyploidy in different floras may differ dramatically. The Greater Cape Floristic Region of southern Africa is one of the world''s biodiversity hotspots and is considered depauperate in polyploids. To test this assumption, ploidy variation was assessed in a widespread member of the largest geophytic genus in the Cape flora: Oxalis obtusa.

Methods

DNA flow cytometry complemented by confirmatory chromosome counts was used to determine ploidy levels in 355 populations of O. obtusa (1014 individuals) across its entire distribution range. Ecological differentiation among cytotypes was tested by comparing sets of vegetation and climatic variables extracted for each locality.

Key Results

Three majority (2x, 4x, 6x) and three minority (3x, 5x, 8x) cytotypes were detected in situ, in addition to a heptaploid individual originating from a botanical garden. While single-cytotype populations predominate, 12 mixed-ploidy populations were also found. The overall pattern of ploidy level distribution is quite complex, but some ecological segregation was observed. Hexaploids are the most common cytotype and prevail in the Fynbos biome. In contrast, tetraploids dominate in the Succulent Karoo biome. Precipitation parameters were identified as the most important climatic variables associated with cytotype distribution.

Conclusions

Although it would be premature to make generalizations regarding the role of genome duplication in the genesis of hyperdiversity of the Cape flora, the substantial and unexpected ploidy diversity in Oxalis obtusa is unparalleled in comparison with any other cytologically known native Cape plant species. The results suggest that ploidy variation in the Greater Cape Floristic Region may be much greater than currently assumed, which, given the documented role of polyploidy in speciation, has direct implications for radiation hypotheses in this biodiversity hotspot.     

Was it an explosion? Using population genetics to explore the dynamics of a recent radiation within Protea (Proteaceae L.)

Adaptive radiations likely underlie much of the world’s diversity, especially that of hyper‐diverse regions. They are usually characterized by a burst of speciation early in their evolutionary history, a pattern which can be detected using population genetic tools. The Cape Floristic Region (CFR) of southwestern South Africa is home to many spectacular plant radiations. Here, we investigate the white proteas (Protea section Exsertae), a typical CFR radiation, to determine if it demonstrates the burst of speciation associated with adaptive radiations in recent models. Inferences from individual assignment, tree‐based population relationships, and pairwise F‐statistics based on 10 microsatellite loci reveal that while the white proteas radiated recently they did not radiate explosively. In addition, we found evidence that there is little gene flow between sampled populations of most species. Taken together, these results demonstrate that within a small clade, the processes underlying the radiation are different from those envisioned by current models of adaptive radiation and suggest that geographical isolation could have played a role in the diversification of the group. Our study implicates both adaptive and non‐adaptive processes in the evolution of botanical diversity of the CFR.     

Bryobartramia schelpei T.A.Hedderson,a new species to accommodate the South African populations of the genus

Abstract

South African populations of Bryobartramia have been treated as B. novae-valesiae, a species otherwise restricted to Australia. However, material from the two regions differs in a number of traits, and the African populations are here described as a new species, B. schelpei. The Cape species differs most obviously from B. novae-valesiae in the markedly prorate-papillose, narrow, thick-walled cells of the calyptra. Nested analysis of variance reveals further morphometric differences, including the relatively narrower, more cylindrical, calyptra with a longer rostrum, and the smaller leaf cells and spores. Bryobartramia schelpii is known only from arid portions of the winter rainfall region in the Northern and Western Cape Provinces of South Africa.