Fossil pollen resolves origin of the South African Proteaceae as transcontinental not transoceanic.

BACKGROUND AND AIMS: The prevailing view from the areocladogenesis of molecular phylogenies is that the iconic South African Cape Proteaceae (subfamily Proteoideae) arrived from Australia across the Indian Ocean in the Upper Cretaceous (100‒65 million years ago, Ma). Since fossil pollen indicates that the family probably arose in North-West Africa during the early Cretaceous, an alternative view is that it migrated to the Cape from North-Central Africa. The plan therefore was to collate fossil pollen records throughout Africa to determine if they are consistent with an African (para-autochthonous) origin for the Cape Proteaceae, and to seek further support from other paleo-disciplines. METHODS: Palynology (identity, date and location of records), molecular phylogeny and chronogram preparation, biogeography of plate tectonics, and paleo-atmospheric and ocean circulation models. KEY RESULTS: Our collation of the rich assemblage of Proteaceae palynomorphs stretching back to 107 Ma (Triorites africaensis) in North-West Africa showed its progressive overland migration to the Cape by 75‒65 Ma. No key palynomorphs recorded in Australia-Antarctica have morphological affinities with African fossils but specific clade assignment of the preMiocene records is not currently possible. The Cape Proteaceae encompass three molecular-based clades (tribes) whose most-recent apparent ancestors are sisters to those in Australia. However, our chronogram shows that the major Adenanthos/Leucadendron-related clade, originating 54‒34 Ma, would have 'arrived' too late as species with Proteaceae affinities were already present ~20 My earlier. The Franklandia/Protea-related clade arose 118‒81 Ma so its distinctive pollen should have been the foundation for the scores of palynomorphs recorded at 100‒80 Ma but it was not. Also, the prevailing winds and ocean currents trended away from South Africa rather than towards, as the 'out-of-Australia' hypothesis requires. Based on the evidence assembled here, we list three points favouring an Australian origin and nine against; four points favouring an Antarctic origin and seven against; and nine points favouring a North-Central African origin and three against. CONCLUSIONS: We conclude that a gradual migration of the Proteaceae from North-Central Africa southeast→south→southwest to the Cape and surrounds occurred via adaptation and speciation during the period 90‒70 Ma. We caution that incorrect conclusions may be drawn from literal interpretations of molecular phylogenies that neglect the fossil record and do not recognize the possible confounding effects of selection under matched environments leading to parallel evolution and extinction of bona fide sister clades.

Plant richness, turnover, and evolutionary diversity track gradients of stability and ecological opportunity in a megadiversity center.

Research on global patterns of diversity has been dominated by studies seeking explanations for the equator-to-poles decline in richness of most groups of organisms, namely the latitudinal diversity gradient. A problem with this gradient is that it conflates two key explanations, namely biome stability (age and area) and productivity (ecological opportunity). Investigating longitudinal gradients in diversity can overcome this problem. Here we investigate a longitudinal gradient in plant diversity in the megadiverse Cape Floristic Region (CFR). We test predictions of the age and area and ecological opportunity hypotheses using metrics for both taxonomic and phylogenetic diversity and turnover. Our plant dataset includes modeled occurrences for 4,813 species and dated molecular phylogenies for 21 clades endemic to the CFR. Climate and biome stability were quantified over the past 140,000 y for testing the age and area hypothesis, and measures of topographic diversity, rainfall seasonality, and productivity were used to test the ecological opportunity hypothesis. Results from our spatial regression models showed biome stability, rainfall seasonality, and topographic heterogeneity were the strongest predictors of taxonomic diversity. Biome stability alone was the strongest predictor of all diversity metrics, and productivity played only a marginal role. We argue that age and area in conjunction with non-productivity-based measures of ecological opportunity explain the CFR's longitudinal diversity gradient. We suggest that this model may possibly be a general explanation for global diversity patterns, unconstrained as it is by the collinearities underpinning the latitudinal diversity gradient.

Fire-mediated germination syndromes in Leucadendron (Proteaceae) and their functional correlates.

A mechanistic understanding of fire-driven seedling recruitment is essential for effective conservation management of fire-prone vegetation, such as South African fynbos, especially with rare and threatened taxa. The genus Leucadendron (Proteaceae) is an ideal candidate for comparative germination studies, comprising 85 species with a mixture of contrasting life-history traits (killed by fire vs able to resprout; serotinous vs geosporous) and seed morphologies (nutlets vs winged achenes). Individual and combined effects of heat and smoke on seed germination of 40 species were quantified in the laboratory, and Bayesian inference applied to distinguish biologically meaningful treatment effects from non-zero, but biologically trivial, effects. Three germination syndromes were identified based on whether germination was dependent on, enhanced by, or independent of direct fire cues (heat and smoke). Seed storage location was the most reliable predictor of germination syndromes, with soil-stored seeds c. 80% more likely to respond to direct fire cues (primarily smoke) than canopy-stored seeds. Notable exceptions were L. linifolium, with an absolute requirement for smoke to germinate (the third serotinous species so reported), and two other serotinous species with smoke-enhanced germination. Nutlet-bearing species, whether serotinous or geosporous, were c. 70% more likely to respond to fire cues than winged seeds, but there was no evidence for an effect of phylogeny or persistence strategy on germination. This comprehensive account of seed germination characteristics and identification of germination syndromes and their predictors, supports propagation, conservation and restoration initiatives in this iconic fynbos genus and other fire-prone shrubs with canopy or soil-stored seeds.

Foraging potential of underground storage organ plants in the southern Cape, South Africa.

Underground storage organs (USOs) serve as a staple source of carbohydrates for many hunter-gatherer societies and they feature prominently in discussions of diets of early modern humans. While the way of life of hunter-gatherers in South Africa's Cape no longer exists, there is extensive ethnographic, historical, and archaeological evidence of hunter-gatherers' use of USOs. This is to be expected, given that the Cape supports the largest concentration of plant species with USOs globally. The southern Cape is the location of several Middle Stone Age sites that are highly significant to research on the origins of behaviourally modern humans, and this provided the context for our research. Here, we evaluate the foraging potential of USOs by identifying how abundant edible biomass is in the southern Cape, how easily it is gathered, and how nutritious it is. One hundred 5 × 5 m plots were assessed in terms of USO species and abundance. Nearly all of the sites sampled (83%) contained edible USOs and some had high concentrations of edible biomass. Extrapolating from these sites suggests that the edible USO biomass falls within the range of biomass observed in areas supporting extant hunter-gatherer communities. The nutritional content for six USO species was assessed; these contained between 40 and 228 calories/100 g. Furthermore, foraging events were staged to provide an indication of the potential return rates for the same six USOs. The target species grow near the soil surface, mostly in sandy soils, and were gathered with minimal effort. Some 50% of the foraging events conducted yielded enough calories to meet the daily requirements of a hunter-gatherer within two hours. In conclusion, we demonstrate that USOs are a readily available source of carbohydrates in the southern Cape landscape and, therefore, there is a strong possibility that USOs played a critical role in providing food for early humans.

Return rates from intertidal foraging from Blombos Cave to Pinnacle Point: Understanding early human economies.

The south coast of South Africa provides the earliest evidence for Middle Stone Age (MSA) coastal resource exploitation by early Homo sapiens. In coastal archaeology worldwide, there has been a debate over the general productivity of intertidal foraging, leading to studies that directly measure productivity in some regions, but there have been no such studies in South Africa. Here we present energetic return rate estimates for intertidal foraging along the southern coast of South Africa from Blombos Cave to Pinnacle Point. Foraging experiments were conducted with Khoi-San descendants of the region, and hourly caloric return rates for experienced foragers were measured on 41 days near low tide and through three seasons over two study years. On-site return rates varied as a function of sex, tidal level, marine habitat type and weather conditions. The overall energetic return rate from the entire sample (1492 kcal h(-1)) equals or exceeds intertidal returns reported from other hunter-gatherer studies, as well as measured return rates for activities as diverse as hunting mammals and plant collecting. Returns are projected to be exceptionally high (∼ 3400 kcal h(-1) for men, ∼ 1900 kcal h(-1) for women) under the best combination of conditions. However, because of the monthly tidal cycle, high return foraging is only possible for about 10 days per month and for only 2-3 h on those days. These experiments suggest that while intertidal resources are attractive, women and children could not have subsisted independently, nor met all their protein-lipid needs from marine resources alone, and would have required substantial additional energy and nutrients from plant gathering and/or from males contributing game.

Using counterfactuals to evaluate the cost-effectiveness of controlling biological invasions.

Prioritizing limited conservation funds for controlling biological invasions requires accurate estimates of the effectiveness of interventions to remove invasive species and their cost-effectiveness (cost per unit area or individual). Despite billions of dollars spent controlling biological invasions worldwide, it is unclear whether those efforts are effective, and cost-effective. The paucity of evidence results from the difficulty in measuring the effect of invasive species removal: a researcher must estimate the difference in outcomes (e.g. invasive species cover) between where the removal program intervened and what might have been observed if the program had not intervened. In the program evaluation literature, this is called a counterfactual analysis, which formally compares what actually happened and what would have happened in the absence of an intervention. When program implementation is not randomized, estimating counterfactual outcomes is especially difficult. We show how a thorough understanding of program implementation, combined with a matching empirical design can improve the way counterfactual outcomes are estimated in nonexperimental contexts. As a practical demonstration, we estimated the cost-effectiveness of South Africa's Working for Water program, arguably the world's most ambitious invasive species control program, in removing invasive alien trees from different land use types, across a large area in the Cape Floristic Region. We estimated that the proportion of the treatment area covered by invasive trees would have been 49% higher (5.5% instead of 2.7% of the grid cells occupied) had the program not intervened. Our estimates of cost per hectare to remove invasive species, however, are three to five times higher than the predictions made when the program was initiated. Had there been no control (counter-factual), invasive trees would have spread on untransformed land, but not on land parcels containing plantations or land transformed by agriculture or human settlements. This implies that the program might have prevented a larger area from being invaded if it had focused all of its clearing effort on untransformed land. Our results show that, with appropriate empirical designs, it is possible to better evaluate the impacts of invasive species removal and therefore to learn from past experiences.

Fossil evidence for a hyperdiverse sclerophyll flora under a non-Mediterranean-type climate.

The spectacular diversity of sclerophyll plants in the Cape Floristic Region in South Africa and Australia's Southwest Floristic Region has been attributed to either explosive radiation on infertile soils under fire-prone, summer-dry climates or sustained accretion of species under inferred stable climate regimes. However, the very poor fossil record of these regions has made these ideas difficult to test. Here, we reconstruct ecological-scale plant species richness from an exceptionally well-preserved fossil flora. We show that a hyperdiverse sclerophyll flora existed under high-rainfall, summer-wet climates in the Early Pleistocene in southeastern Australia. The sclerophyll flora of this region must, therefore, have suffered subsequent extinctions to result in its current relatively low diversity. This regional loss of sclerophyll diversity occurred at the same time as a loss of rainforest diversity and cannot be explained by ecological substitution of species of one ecological type by another type. We show that sclerophyll hyperdiversity has developed in distinctly non-Mediterranean climates, and this diversity is, therefore, more likely a response to long-term climate stability. Climate stability may have both reduced the intensity of extinctions associated with the Pleistocene climate cycles and promoted the accumulation of species richness by encouraging genetic divergence between populations and discouraging plant dispersal.

Expert-derived monitoring thresholds for impacts of megaherbivores on vegetation cover in a protected area.

Monitoring is meant to inform conservation authorities, yet managers often don't know when to respond to monitoring results. One of the reasons is that management often lacks consensus on monitoring thresholds for intervention. This results in aimless monitoring without a clear directive on when monitoring indicates a trajectory towards an unacceptable state or impending change, which possibly necessitates intervention. Although experts rarely provide simple, measureable and quantifiable monitoring thresholds as required by management, they are often more comfortable expressing opinions on whether a specific area is desirable or not. This allows thresholds to be reverse engineered: by getting experts to identify sites as desirable and undesirable, field variables can subsequently be measured to derive the boundary between subjectively identified desirable and undesirable states. Such a boundary provides a defendable point for management to assess and consider intervention. Here we describe the identification of monitoring thresholds by defining the limits of desirable canopy cover, derived from expert stakeholder preferences, in the Sundays Spekboom Thicket vegetation of the Addo Elephant National Park, South Africa. The park has experienced variable utilization intensity by large herbivores, especially elephant. For years managers have grappled with the question of what percentage shrub canopy cover is desirable as a management target, but science has failed to provide this. Using experts to assess pre-selected sites as desirable or undesirable across a range of canopy covers, we showed that a canopy cover of ∼65% (±15%) would be desirable for expert stakeholders. We then used satellite imagery to map canopy cover, providing managers for the first time with a large-scale map of canopy cover, indicating desirability status. This approach was useful for facilitating joint-decision making between conservation agencies and stakeholders on tangible indicators of achieving goals, and may be useful in fostering relationships, trust, mutual understanding and transparency, characteristics critical for managing complex socio-ecological systems.

A new research strategy for integrating studies of paleoclimate, paleoenvironment, and paleoanthropology.

Paleoanthropologists (scientists studying human origins) universally recognize the evolutionary significance of ancient climates and environments for understanding human origins. Even those scientists working in recent phases of human evolution, when modern humans evolved, agree that hunter-gatherer adaptations are tied to the way that climate and environment shape the food and technological resource base. The result is a long tradition of paleoanthropologists engaging with climate and environmental scientists in an effort to understand if and how hominin bio-behavioral evolution responded to climate and environmental change. Despite this unusual consonance, the anticipated rewards of this synergy are unrealized and, in our opinion, will not reach potential until there are some fundamental changes in the way the research model is constructed. Discovering the relation between climate and environmental change to human origins must be grounded in a theoretical framework and a causal understanding of the connection between climate, environment, resource patterning, behavior, and morphology, then move beyond the strict correlative research that continues to dominate the field.

Extinction risk and diversification are linked in a plant biodiversity hotspot.

It is widely recognized that we are entering an extinction event on a scale approaching the mass extinctions seen in the fossil record. Present-day rates of extinction are estimated to be several orders of magnitude greater than background rates and are projected to increase further if current trends continue. In vertebrates, species traits, such as body size, fecundity, and geographic range, are important predictors of vulnerability. Although plants are the basis for life on Earth, our knowledge of plant extinctions and vulnerabilities is lagging. Here, we disentangle the underlying drivers of extinction risk in plants, focusing on the Cape of South Africa, a global biodiversity hotspot. By comparing Red List data for the British and South African floras, we demonstrate that the taxonomic distribution of extinction risk differs significantly between regions, inconsistent with a simple, trait-based model of extinction. Using a comprehensive phylogenetic tree for the Cape, we reveal a phylogenetic signal in the distribution of plant extinction risks but show that the most threatened species cluster within short branches at the tips of the phylogeny--opposite to trends in mammals. From analyzing the distribution of threatened species across 11 exemplar clades, we suggest that mode of speciation best explains the unusual phylogenetic structure of extinction risks in plants of the Cape. Our results demonstrate that explanations for elevated extinction risk in plants of the Cape flora differ dramatically from those recognized for vertebrates. In the Cape, extinction risk is higher for young and fast-evolving plant lineages and cannot be explained by correlations with simple biological traits. Critically, we find that the most vulnerable plant species are nonetheless marching towards extinction at a more rapid pace but, surprisingly, independently from anthropogenic effects. Our results have important implications for conservation priorities and cast doubts on the utility of current Red List criteria for plants in regions such as the Cape, where speciation has been rapid, if our aim is to maximize the preservation of the tree-of-life.

Preserving the evolutionary potential of floras in biodiversity hotspots.

One of the biggest challenges for conservation biology is to provide conservation planners with ways to prioritize effort. Much attention has been focused on biodiversity hotspots. However, the conservation of evolutionary process is now also acknowledged as a priority in the face of global change. Phylogenetic diversity (PD) is a biodiversity index that measures the length of evolutionary pathways that connect a given set of taxa. PD therefore identifies sets of taxa that maximize the accumulation of 'feature diversity'. Recent studies, however, concluded that taxon richness is a good surrogate for PD. Here we show taxon richness to be decoupled from PD, using a biome-wide phylogenetic analysis of the flora of an undisputed biodiversity hotspot--the Cape of South Africa. We demonstrate that this decoupling has real-world importance for conservation planning. Finally, using a database of medicinal and economic plant use, we demonstrate that PD protection is the best strategy for preserving feature diversity in the Cape. We should be able to use PD to identify those key regions that maximize future options, both for the continuing evolution of life on Earth and for the benefit of society.

On the accuracy of conservation managers' beliefs and if they learn from evidence-based knowledge: a preliminary investigation.

Despite the significant impetus placed on the need for conservation managers to base their decisions on evidence-based findings, few studies have compared the accuracy of "evidence" versus experience-based knowledge. Furthermore we are not aware of any study that has tested the willingness of managers to change their beliefs after being exposed to evidence-based findings. Here, we tested nine managers' beliefs before-and-after being shown findings from an evidence-based study. The questions centered on the effectiveness of 'Working for Water' (WfW) in reducing invasive alien plant cover in two large catchment projects over a seven year period, as well as the managers' forecasts of WfW's effectiveness of reducing invasive alien plant cover, and the factors that underpin its effectiveness. We also assessed the financial cost of implementing the evidence-based assessment. We found that in comparison to the evidence-based findings, the managers underestimated the ineffectiveness of operations in reducing invasive alien plant cover in the one catchment and overestimated the ineffectiveness of the other catchment. All the managers whose estimates differed from the evidence-based findings were willing to change their beliefs. Surprisingly, however, when it came to forecasting WfW's effectiveness in the catchments, all the managers, with the exception of one project manager, were unwilling to reduce their optimistic estimates of the time required to control invasive alien plants from the two catchments. With regard to the drivers of effectiveness, the managers ranked their performance as the most important criterion whereas the data model emphasized variables related to site suitability for alien plant growth. Finally, we showed that it would only cost between 0.33% and 1.67% of the two projects' annual budgets to assess all sites, depending on the frequency of the monitoring. This preliminary investigation highlights how evidence-based findings alone, even if presented and explained to managers, might not result in managers learning and updating their beliefs.

The vegetation of Holocene coastal dunes of the Cape south coast, South Africa.

The vegetation of calcareous coastal dunes of Holocene age along the south coast of South Africa's Cape Floristic Region is poorly described. This vegetation comprises a mosaic of communities associated with two biomes, Fynbos and Subtropical Thicket. Previously, expert knowledge rather than quantitative floristic analysis has been used to identify and delimit vegetation units. In many areas, mapped units conflate vegetation on Holocene sand with that on unconsolidated sediments of late Pleistocene age, despite pronounced species turnover across this edaphic boundary. Despite dominance by Cape lineages and fynbos vegetation, dune vegetation in the eastern part of the region has been included in the Subtropical Thicket Biome rather than the Fynbos Biome. The high levels of local plant endemism associated with this dune vegetation and the small and fragmented configuration of these habitats, makes it an urgent conservation priority especially when placed in the context of rising sea levels, increasing development pressures and numerous other threats. Here we provide a quantitative analysis of 253 plots of the 620 km2 of Holocene dune vegetation of the study area using phytosociological and multivariate methods. We identified six fynbos and two thicket communities based on the occurrences of 500 species. Following a long tradition in Cape vegetation typology, we used the Strandveld (beach vegetation) concept as our first-order vegetation entity and identified six units based on the fynbos floras. These were, from east to west, Southeastern Strandveld, St Francis Strandveld, Goukamma Strandveld, Southwestern Strandveld and Grootbos Strandveld. Each unit was differentiated by a suite of differential species, most being Holocene dune endemics. The two thicket communities-Mesic and Xeric Dune Thicket-showed limited variation across the study area and were subsumed into the Strandveld units. We discussed our findings in terms of vegetation-sediment relationships, emphasizing the need for a greater geographical coverage of sediment ages to facilitate a better understanding of deposition history on vegetation composition. We also discussed the role of soil moisture and fire regime on structuring the relative abundance of fynbos and thicket across the Holocene dune landscape. Finally, we address the conservation implications of our study, arguing that all remaining Holocene dune habitat should be afforded the highest conservation priority in regional land-use planning processes.

Canopy plant composition and structure of Cape subtropical dune thicket are predicted by the levels of fire exposure.

Background: The subtropical dune thicket (hereafter "dune thicket") of the Cape Floristic Region experiences a wide range of fire exposure throughout the landscape, unlike other dry rainforest formations that rarely experience fire. We sought to determine how fire exposure influences species composition and the architectural composition of dune thicket. Methods: We used multivariate analysis and diversity indices based on cover abundance of species to describe the species composition, architectural guild composition and structure of dune thicket sites subject to different levels of fire exposure, namely low (fire return interval of >100 years), moderate (fire return interval of 50-100 years), and high (fire return interval of 10-50 years). Results: The diversity, cover abundance and architectural guild cover abundance of dune thicket canopy species were strongly influenced by the level of fire exposure such that each level was associated with a well-circumscribed vegetation unit. Dune thickets subject to low fire exposure comprises a floristically distinct, low forest characterized by shrubs with one-to-few upright stems (ca. 4-8 m tall) and a relatively small canopy spread (vertical growers). Of the 25 species in this unit, 40% were restricted to it. Dune thickets subject to moderate fire exposure had the highest abundance of lateral spreaders, which are multi-stemmed (ca. 3-6 m tall) species with a large canopy spread and lower stature than vertical growers. None of the 17 species found in this unit was restricted to it. Dune thickets subject to high fire exposure had the highest abundance of hedge-forming shrubs, these being low shrubs (ca. 0.6-1.4 m tall), with numerous shoots arising from an extensive system of below-ground stems. Of the 20 species in this unit, 40% were restricted to it. Multivariate analysis identified three floristic units corresponding to the three fire exposure regimes. Compositional structure, in terms of species and architectural guilds, was most distinctive for dune thickets subject to high and low fire exposure, while the dune thicket subject to moderate fire exposure showed greatest compositional overlap with the other units. Conclusion: Fire exposure profoundly influenced the composition and structure of dune thicket canopy species in the Cape Floristic Region. In the prolonged absence of fire, the thicket is invaded by vertical-growing species that overtop and outcompete the multi-stemmed, laterally-spreading shrubs that dominate this community. Regular exposure to fire selects for traits that enable thicket species to rapidly compete for canopy cover post-fire via the prolific production of resprouts from basal buds below- and above-ground. The trade-off is that plant height is constrained, as proportionately more resources are allocated to below-ground biomass.

The response of geophytes to continuous human foraging on the Cape south coast, South Africa and its implications for early hunter-gatherer mobility patterns.

Current ecological understanding of plants with underground storage organs (USOs) suggests they have, in general, low rates of recruitment and thus as a resource it should be rapidly exhausted, which likely had implications for hunter-gatherer mobility patterns. We focus on the resilience (defined here as the ability of species to persist after being harvested) of USOs to human foraging. Human foragers harvested all visible USO material from 19 plots spread across six Cape south coast (South Africa) vegetation types for three consecutive years (2015-2017) during the period of peak USO apparency (September-October). We expected the plots to be depleted after the first year of harvesting since the entire storage organ of the USO is removed during foraging, i.e. immediate and substantial declines from the first to the second harvest. However, over 50% of the total weight harvested in 2015 was harvested in 2016 and 2017; only after two consecutive years of harvesting, was there evidence of significantly lower yield (p = 0.034) than the first (2015) harvest. Novel emergence of new species and new individuals in year two and three buffered the decline of harvested USOs. We use our findings to make predictions on hunter-gatherer mobility patterns in this region compared to the Hadza in East Africa and the Alyawara in North Australia.

An operational model for mainstreaming ecosystem services for implementation.

Research on ecosystem services has grown markedly in recent years. However, few studies are embedded in a social process designed to ensure effective management of ecosystem services. Most research has focused only on biophysical and valuation assessments of putative services. As a mission-oriented discipline, ecosystem service research should be user-inspired and user-useful, which will require that researchers respond to stakeholder needs from the outset and collaborate with them in strategy development and implementation. Here we provide a pragmatic operational model for achieving the safeguarding of ecosystem services. The model comprises three phases: assessment, planning, and management. Outcomes of social, biophysical, and valuation assessments are used to identify opportunities and constraints for implementation. The latter then are transformed into user-friendly products to identify, with stakeholders, strategic objectives for implementation (the planning phase). The management phase undertakes and coordinates actions that achieve the protection of ecosystem services and ensure the flow of these services to beneficiaries. This outcome is achieved via mainstreaming, or incorporating the safeguarding of ecosystem services into the policies and practices of sectors that deal with land- and water-use planning. Management needs to be adaptive and should be institutionalized in a suite of learning organizations that are representative of the sectors that are concerned with decision-making and planning. By following the phases of our operational model, projects for safeguarding ecosystem services are likely to empower stakeholders to implement effective on-the-ground management that will achieve resilience of the corresponding social-ecological systems.

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.

A biome-wide experiment to assess the effects of propagule size and treatment on the survival of Portulacaria afra (spekboom) truncheons planted to restore degraded subtropical thicket of South Africa.

Insights from biome-wide experiments can improve efficacy of landscape-scale ecological restoration projects. Such insights enable implementers to set temporal and geographical benchmarks and to identify key drivers of success during the often decades-long restoration trajectory. Here we report on a biome-wide experiment aimed at informing the ecological restoration of thousands of hectares of degraded subtropical thicket dominated by the succulent shrub, Portulacaria afra (spekboom). Restoration using spekboom truncheons has the potential to sequester, for a semi-arid region, large amounts of ecosystem carbon, while regenerating a host of associated ecosystem services. This study evaluates, after about three years post-propagation, the effects of spekboom truncheon size and treatment on survivorship in 40 fence-enclosed (0.25 ha) plots located in target habitat across the entire spekboom thicket biome. In each plot, locally harvested spekboom truncheons, comprising eight size/treatment combinations, were planted in replicated rows of between 24 and 49 individuals, depending on treatment. The experiment assessed the role of truncheon size, spacing, application of rooting hormone and watering at planting on survivorship percentage as an indicator of restoration success. All eight combinations recorded extreme minimum survivorship values of zero, while the range of extreme maximum values was 70-100%. Larger truncheons (>22.5 mm diameter) had almost double the survivorship (ca. 45%) than smaller truncheons (< 15 mm) (ca. 25%). Planting large, untreated truncheons at 1 m intervals-as opposed to 2 m intervals recommended in the current restoration protocol-resulted in no significant change in survivorship. The application of rooting hormone and water at planting had no significant effect on restoration success for both large and small truncheons. While our results do not provide an evidence base for changing the current spekboom planting protocol, we recommend research on the financial and economic costs and benefits of different propagation strategies in real-world contexts.

Herbivory and misidentification of target habitat constrain region-wide restoration success of spekboom (Portulacaria afra) in South African subtropical succulent thicket.

Restoration of degraded subtropical succulent thicket, via the planting of Portulacaria afra (spekboom) truncheons, is the focus of a public works programme funded by the South African government. The goals of the programme, which started in 2004, are to create jobs, sequester carbon, restore biodiversity, reduce erosion, improve soil water holding capacity and catalyse private sector investment for upscaling of restoration. Here we report on a region-wide experiment to identify factors that can improve project success. Measures of success were survivorship and annual aboveground biomass carbon sequestration (ABCsr) of spekboom truncheons some 33-57 months after planting-starting in March 2008-into 173 fenced plots (0.25 ha) located throughout the global extent of spekboom thicket vegetation. We also collected data for 18 explanatory variables under the control of managers, and an additional 39 variables reflecting soil physical and chemical characteristics and rainfall patterns post restoration, all beyond the influence of managers. Since the latter covariates were available for only 83 plots, we analysed the two data sets separately. We used a prediction rule ensemble to determine the most important predictors of restoration success. There was great variation in percentage survivorship (median = 24, range = 0-100%) and ABCsr (median = 0.009, range = 0-0.38 t C ha-1 yr-1). The model using management variables explained less variance (53%) in survivorship than the model incorporating additional soil and rainfall covariates (62%). ABCsr models were better fits (78 and 88% variance explained, respectively). All model configurations identified browse intensity as a highly influential predictor of restoration success. Predicted success was highest for plots located in target habitat; however, only 45% were thus located, suggesting the need for expert input and habitat modelling for improving target habitat identification. Frost exposure was another important predictor influencing all models but was likely a consequence of locating sites off target habitat. Sites planted on equatorward slopes during the warm season showed reduced carbon sequestration, possibly due to elevated soil moisture stress associated with high radiation loads. Physiographic factors associated with improved restoration success were location on sloping ground (reduced frost exposure), increasing longitude (more warm-season rainfall) and increasing latitude (less frost coastwards). Few trends were evident among post-restoration climatic factors beyond the control of managers. Higher rainfall during the year post restoration had a negative impact on carbon sequestration while higher rain during the early months post restoration had a positive effect on both carbon sequestration and survivorship. Soil factors showed little importance for the survivorship model, whereas silt content, % K and Mg CEC emerged as predictors of carbon sequestration. Our results have direct relevance for improving the success of landscape-scale restoration projects envisioned for the ca. 8,930 km2 of degraded spekboom thicket.

Evolutionary stability, landscape heterogeneity, and human land-usage shape population genetic connectivity in the Cape Floristic Region biodiversity hotspot.

As human-induced change eliminates natural habitats, it impacts genetic diversity and population connectivity for local biodiversity. The South African Cape Floristic Region (CFR) is the most diverse extratropical area for plant biodiversity, and much of its habitat is protected as a UNESCO World Heritage site. There has long been great interest in explaining the underlying factors driving this unique diversity, especially as much of the CFR is endangered by urbanization and other anthropogenic activity. Here, we use a population and landscape genetic analysis of SNP data from the CFR endemic plant Leucadendron salignum or "common sunshine conebush" as a model to address the evolutionary and environmental factors shaping the vast CFR diversity. We found that high population structure, along with relatively deeper and older genealogies, is characteristic of the southwestern CFR, whereas low population structure and more recent lineage coalescence depict the eastern CFR. Population network analyses show genetic connectivity is facilitated in areas of lower elevation and higher seasonal precipitation. These population genetic signatures corroborate CFR species-level patterns consistent with high Pleistocene biome stability and landscape heterogeneity in the southwest, but with coincident instability in the east. Finally, we also find evidence of human land-usage as a significant gene flow barrier, especially in severely threatened lowlands where genetic connectivity has been historically the highest. These results help identify areas where conservation plans can prioritize protecting high genetic diversity threatened by contemporary human activities within this unique cultural UNESCO site.