Berit Gehrke

Estimates of the number of vascular plant species currently under threat of extinction are shockingly high, with the highest extinction rates reported for narrow-range, woody plants, especially in biodiversity hotspots with Mediterranean and tropical climates. The large genus Erica is a prime example, as a large proportion of its 851 species, all shrubs or small trees, are endemic to the Cape Floristic Region (CFR) of South Africa. Almost two hundred are known to be threatened and a further hundred are ‘Data Deficient’. We need to target conservation efforts and research to fill the most problematic knowledge gaps. This can be especially challenging in large genera, such as Erica, with numerous threatened species that are closely related. One approach involves combining knowledge of phylogenetic diversity with that of IUCN threat status to identify the most Evolutionarily Distinct and Globally Endangered (EDGE) species. We present an expanded and improved phylogenetic hypothesis for Erica (representing 65% of described species diversity) and combine this with available threat and distribution data to identify species and geographic areas that could be targeted for conservation effort to maximise preservation of phylogenetic diversity (PD). The resulting 39 EDGE taxa include 35 from the CFR. A further 32 high PD, data deficient taxa are mostly from outside the CFR, reflecting the low proportion of assessed taxa outside South Africa. The most taxon-rich areas are found in the south-western CFR. They are not the most phylogenetically diverse, but do include the most threatened PD. These results can be cross-referenced to existing living and seed-banked ex situ collections and used to target new and updated threat assessments and conservation action.

Estimates of the number of vascular plant species currently under threat of extinction are shockingly high, with the highest extinction rates reported for narrow-range, woody plants, especially in biodiversity hotspots with Mediterranean and tropical climates. The large genus Erica is a prime example, as a large proportion of its 851 species, all shrubs or small trees, are endemic to the Cape Floristic Region (CFR) of South Africa. Almost two hundred are known to be threatened and a further hundred are ‘Data Deficient’. We need to target conservation efforts and research to fill the most problematic knowledge gaps. This can be especially challenging in large genera, such as Erica, with numerous threatened species that are more or less closely related. One approach involves combining knowledge of phylogenetic diversity with that of IUCN threat status to identify the most evolutionarily distinct and globally endangered (EDGE) species. We present an expanded and improved phylogenetic hypothesis for Erica (representing 65% of described species diversity) and combine this with available threat and distribution data to identify species and geographic areas that could be targeted for conservation effort to maximise preservation of phylogenetic diversity (PD). The resulting 39 EDGE taxa include 35 from the CFR. A further 32 high PD, data deficient taxa are mostly from outside the CFR, reflecting the low proportion of assessed taxa outside South Africa. The most taxon rich areas are found in the south-western CFR. They are not the most phylogenetically diverse, but do include the most threatened PD. These results can be cross-referenced to existing living and seed banked ex situ collections and used to target new and updated threat assessments and conservation action.

Summary: Poales are one of the most species‐rich, ecologically and economically important orders of plants and often characterise open habitats, enabled by unique suites of traits. We test six hypotheses regarding the evolution and assembly of Poales in open and closed habitats throughout the world, and examine whether diversification patterns demonstrate parallel evolution. We sampled 42% of Poales species and obtained taxonomic and biogeographic data from the World Checklist of Vascular Plants database, which was combined with open/closed habitat data scored by taxonomic experts. A dated supertree of Poales was constructed. We integrated spatial phylogenetics with regionalisation analyses, historical biogeography and ancestral state estimations. Diversification in Poales and assembly of open and closed habitats result from dynamic evolutionary processes that vary across lineages, time and space, most prominently in tropical and southern latitudes. Our results reveal parallel and recurrent patterns of habitat and trait transitions in the species‐rich families Poaceae and Cyperaceae. Smaller families display unique and often divergent evolutionary trajectories. The Poales have achieved global dominance via parallel evolution in open habitats, with notable, spatially and phylogenetically restricted divergences into strictly closed habitats.

Significance: Resilience is required to withstand or mitigate the effect of human-induced climate change. Today whole ecosystems are affected by climate change, but our understanding of their evolution and natural response is limited, often restricted to individual populations or species. The enigmatic flora on the tops of the African sky islands is isolated and unique, showing striking adaptations to the harsh tropical alpine conditions. Here we analyze genome data from a large fraction of afroalpine plants and show that this remarkable flora has a dynamic history with frequent colonizations and extinctions, most likely caused by previous natural climate changes during the ice-age cycles. The flora will be particularly vulnerable to human-induced climate warming, reducing alpine habitat into successively smaller areas.

The monumental work of Olov Hedberg provided deep insights into the spectacular and fragmented tropical alpine flora of the African sky islands. Here we review recent molecular and niche modelling studies and re-examine Hedberg’s hypotheses and conclusions. Colonisation started when mountain uplift established the harsh diurnal climate with nightly frosts, accelerated throughout the last 5 Myr (Plio-Pleistocene), and resulted in a flora rich in local endemics. Recruitment was dominated by long-distance dispersals (LDDs) from seasonally cold, remote areas, mainly in Eurasia. Colonisation was only rarely followed by substantial diversification. Instead, most of the larger genera and even species colonised the afroalpine habitat multiple times independently. Conspicuous parallel evolution occurred among mountains, e.g., of gigantism inLobeliaandDendrosenecioand dwarf shrubs inAlchemilla. Although the alpine habitat was ~ 8 times larger and the treeline was ~ 1000 m lower than today during the Last Glacial Maximum, genetic data suggest that the flora was shaped by strong intermountain isolation interrupted by rare LDDs rather than ecological connectivity. The new evidence points to a much younger and more dynamic island scenario than envisioned by Hedberg: the afroalpine flora is unsaturated and fragile, it was repeatedly disrupted by the Pleistocene climate oscillations, and it harbours taxonomic and genetic diversity that is unique but severely depauperated by frequent bottlenecks and cycles of colonisation, extinction, and recolonisation. The level of intrapopulation genetic variation is alarmingly low, and many afroalpine species may be vulnerable to extinction because of climate warming and increasing human impact.

Background: The coincidence of long distance dispersal (LDD) and biome shift is assumed to be the result of a multifaceted interplay between geographical distance and ecological suitability of source and sink areas. Here, we test the influence of these factors on the dispersal history of the flowering plant genus Erica (Ericaceae) across the Afrotemperate. We quantify similarity of Erica climate niches per biogeographic area using direct observations of species, and test various colonisation scenarios while estimating ancestral areas for the Erica clade using parametric biogeographic model testing. Results: We infer that the overall dispersal history of Erica across the Afrotemperate is the result of infrequent colonisation limited by geographic proximity and niche similarity. However, the Drakensberg Mountains represent a colonisation sink, rather than acting as a “stepping stone” between more distant and ecologically dissimilar Cape and Tropical African regions. Strikingly, the most dramatic examples of species radiations in Erica were the result of single unique dispersals over longer distances between ecologically dissimilar areas, contradicting the rule of phylogenetic biome conservatism. Conclusions: These results highlight the roles of geographical and ecological distance in limiting LDD, but also the importance of rare biome shifts, in which a unique dispersal event fuels evolutionary radiation.

Plant species tend to retain their ancestral ecology, responding to temporal, geographic and climatic changes by tracking suitable habitats rather than adapting to novel conditions. Nevertheless, transitions into different environments or biomes still seem to be common. Especially intriguing are the tropical alpine-like areas found on only the highest mountainous regions surrounded by tropical environments. Tropical mountains are hotspots of biodiversity, often with striking degrees of endemism at higher elevations. On these mountains, steep environmental gradients and high habitat heterogeneity within small spaces coincide with astounding species diversity of great conservation value. The analysis presented here shows that the importance ofin situspeciation in tropical alpine-like areas has been underestimated. Additionally and contrary to widely held opinion, the impact of dispersal from other regions with alpine-like environments is relatively minor compared to that of immigration from other biomes with a temperate (but not alpine-like) climate. This suggests that establishment in tropical alpine-like regions is favoured by preadaptation to a temperate, especially aseasonal, freezing regime such as the cool temperate climate regions in the Tropics. Furthermore, emigration out of an alpine-like environment is generally rare, suggesting that alpine-like environments – at least tropical ones – are species sinks.

The coincidence of long distance dispersal and biome shift is assumed to be the result of a multifaceted interplay between geographical distance and ecological suitability of source and sink areas. Here, we test the influence of these factors on the dispersal history of the flowering plant genus Erica (Ericaceae) across the Afrotemperate. We quantify similarity of Erica climate niches per biogeographic area using direct observations of species, and test various colonisation scenarios while estimating ancestral areas for the Erica clade using parametric biogeographic model testing. We infer that the overall dispersal history of Erica across the Afrotemperate is the result of infrequent colonisation limited by geographic proximity and niche similarity. However, the Drakensberg Mountains represent a colonisation sink, rather than acting as a “stepping stone” between more distant and ecologically dissimilar Cape and Tropical African regions. Strikingly, the most dramatic examples of species radiations in Erica were the result of single unique dispersals over longer distances between ecologically dissimilar areas, contradicting the rule of phylogenetic biome conservatism. These results highlight the importance of rare biome shifts, in which a unique dispersal event fuels evolutionary radiation.

As one of the largest genera of flowering plants, the richness of species in Erica (Ericaceae) is all the more remarkable because > 80% of the > 800 species are endemic to the smallest floral kingdom, the Cape Floristic Region (CFR) of South Africa. In the CFR, pockets of narrowly endemic taxa appear in close juxtaposition with their widespread and variable relatives. The taxonomic challenges of Cape Erica are epitomized by the complex Cape ‘abietina/viscaria clade’, currently comprising at least 25 species. We reassess species boundaries and patterns of regional endemism in this clade using a phylogenetic tree inferred from multiple nuclear ribosomal and plastid DNA sequences. We show that the seven currently recognized subspecies of E. abietina represent at least three independent, morphologically distinct lineages with non-overlapping geographical distributions. We resurrect the name E. grandiflora to include E. abietina subsp. aurantiaca and subsp. perfoliosa and we provide a new name for E. abietina subsp. petraea (E. situshiemalis). This means that E. abietina is now an additional endemic species for the Cape Peninsula, including the natural World Heritage Site, Table Mountain National Park.