Deciphering Pteronia's evolution in the Cape Floristic Region: A comprehensive study disputes polyploid deficiency and affirms diploid radiation.

The Greater Cape Floristic Region (GCFR) is renowned for its exceptional biodiversity, accommodating over 11 000 plant species, notable degree of endemism, and substantial diversification within limited plant lineages, a phenomenon ascribed to historical radiation events. While both abiotic and biotic factors contribute to this diversification, comprehensive genomic alterations, recognized as pivotal in the diversification of angiosperms, are perceived as uncommon. This investigation focuses on the genus Pteronia, a prominent representative of the Asteraceae family in the GCFR. Employing NGS-based HybSeq and RADSeq methodologies, flow cytometry, karyology, and ecological modeling, we scrutinize the intricacies of its polyploid evolution. Phylogenetic reconstructions using 951 low-copy nuclear genes confirm Pteronia as a well-supported, distinct clade within the tribe Astereae. The ingroup displays a structure indicative of rapid radiation likely antedating polyploid establishment, with the two main groups demarcated by their presence or absence in the fynbos biome. Genome size analysis encompasses 1293 individuals across 347 populations, elucidating significant variation ranging from 6.1 to 34.2 pg (2C-value). Pteronia demonstrates substantially large genome sizes within Astereae and phanerophytes. Polyploidy is identified in 31% of the studied species, with four discerned ploidy levels (2x, 4x, 6x, 8x). Cytotypes exhibit marked distinctions in environmental traits, influencing their distribution across biomes and augmenting their niche differentiation. These revelations challenge the presumed scarcity of polyploidy in the Cape flora, underscoring the imperative need for detailed population studies. The intricate evolutionary history of Pteronia, characterized by recent polyploidy and genome size variation, contributes substantially to the comprehension of diversification patterns within the GCFR biodiversity hotspot.

Strong habitat and seasonal phenology effects on the evolution of self-compatibility, clonality and pollinator shifts in Lachenalia (Asparagaceae: Scilloideae).

Plants employ a diversity of reproductive safeguarding strategies to circumvent the challenge of pollen limitation. Focusing on southern African Lachenalia (Asparagaceae: Scilloideae), we test the hypothesis that the evolution of reproductive safeguarding traits (self-compatibility, autonomous selfing, bird pollination and clonal propagation) is favoured in species occupying conditions of low insect abundance imposed by critically infertile fynbos heathland vegetation and by flowering outside the austral spring insect abundance peak. We trace the evolution of these traits and selective regimes on a dated, multi-locus phylogeny of Lachenalia and assess their evolutionary associations using ordinary and phylogenetic regression. Ancestral state reconstructions identify an association with non-fynbos vegetation and spring flowering as ancestral in Lachenalia, the transition to fynbos vegetation and non-spring flowering taking place multiple times. They also show that self-compatibility, autofertility, bird pollination and production of multiple clonal offsets have evolved repeatedly. Regression models suggest that bird pollination and self-compatibility are selected for in fynbos and in non-spring flowering lineages, with autofertility being positively associated with non-spring flowering. These patterns support the interpretation of these traits as reproductive safeguarding adaptations under reduced insect pollinator abundance. We find no evidence to support the interpretation of clonal propagation as a reproductive safeguarding strategy.

Pollinator shifts despite hybridisation in the Cape's hyperdiverse heathers (Erica, Ericaceae).

Interrogating the ecological and geographic factors that influence population divergence dynamics can reveal why some groups of organisms diversify more prolifically than others. One such group is the heathers (Erica, Ericaceae), the largest plant genus in the Cape Floristic Region. We study Erica abietina, a highly variable species complex with four subspecies differing in geographic range, habitat and pollination syndrome. We test for population differentiation, hybridisation, introgression and pollinator-driven divergence using genotyping-by-sequencing on samples across the entire distribution. We find five variably distinct genetic groups, with one subspecies comprising two independent lineages that are geographically isolated and occur on different soil types. Phylogenetic analysis suggests two independent shifts between bird and insect pollination, with accompanying genetic divergence. However, for one pair of populations with different pollinators, we uncover several individuals of hybrid origin at a site of sympatry. These results suggest that floral differentiation driven by divergent selection acts in concert with geographic isolation to maintain reproductive isolation and promote speciation. Our investigations reveal a highly dynamic system whose diversity has been shaped by a variety of interacting forces. We suggest that such a system could be a model for much of the diversification of the Cape flora.

Genome size variation in Cape schoenoid sedges (Schoeneae) and its ecophysiological consequences.

PREMISE: Increases in genome size in plants-often associated with larger, low-density stomata and greater water-use efficiency (WUE)-could affect plant ecophysiological and hydraulic function. Variation in plant genome size is often due to polyploidy, having occurred repeatedly in the austral sedge genus Schoenus in the Cape Floristic Region (CFR), while species in the other major schoenoid genus in the region, Tetraria, have smaller genomes. Comparing these genera is useful as they co-occur at the landscape level, under broadly similar bioclimatic conditions. We hypothesized that CFR Schoenus have greater WUE, with lower maximum stomatal conductance (gwmax) imposed by larger, less-dense stomata. METHODS: We investigated relationships between genome size and stomatal parameters in a phylogenetic context, reconstructing a phylogeny of CFR-occurring Schoeneae (Cyperaceae). Species' stomatal and functional traits were measured from field-collected and herbarium specimens. Carbon stable isotopes were used as an index of WUE. Genome size was derived from flow-cytometric measurements of leafy shoots. RESULTS: Evolutionary regressions demonstrated that stomatal size and density covary with genome size, positively and negatively, respectively, with genome size explaining 72-75% of the variation in stomatal size. Larger-genomed species had lower gwmax and C:N ratios, particularly in culms. CONCLUSIONS: We interpret differences in vegetative physiology between the genera as evidence of more-conservative strategies in CFR Schoenus compared to the more-acquisitive Tetraria. Because Schoenus have smaller, reduced leaves, they likely rely more on culm photosynthesis than Tetraria. Across the CFR Schoeneae, ecophysiology correlates with genome size, but confounding sources of trait variation limit inferences about causal relationships between traits.

Complex patterns of ploidy in a holocentric plant clade (Schoenus, Cyperaceae) in the Cape biodiversity hotspot.

BACKGROUND AND AIMS: It is unclear how widespread polyploidy is throughout the largest holocentric plant family - the Cyperaceae. Because of the prevalence of chromosomal fusions and fissions, which affect chromosome number but not genome size, it can be impossible to distinguish if individual plants are polyploids in holocentric lineages based on chromosome count data alone. Furthermore, it is unclear how differences in genome size and ploidy levels relate to environmental correlates within holocentric lineages, such as the Cyperaceae. METHODS: We focus our analyses on tribe Schoeneae, and more specifically the southern African clade of Schoenus. We examine broad-scale patterns of genome size evolution in tribe Schoeneae and focus more intensely on determining the prevalence of polyploidy across the southern African Schoenus by inferring ploidy level with the program ChromEvol, as well as interpreting chromosome number and genome size data. We further investigate whether there are relationships between genome size/ploidy level and environmental variables across the nutrient-poor and summer-arid Cape biodiversity hotspot. KEY RESULTS: Our results show a large increase in genome size, but not chromosome number, within Schoenus compared to other species in tribe Schoeneae. Across Schoenus, there is a positive relationship between chromosome number and genome size, and our results suggest that polyploidy is a relatively common process throughout the southern African Schoenus. At the regional scale of the Cape, we show that polyploids are more often associated with drier locations that have more variation in precipitation between dry and wet months, but these results are sensitive to the classification of ploidy level. CONCLUSIONS: Polyploidy is relatively common in the southern African Schoenus, where a positive relationship is observed between chromosome number and genome size. Thus, there may be a high incidence of polyploidy in holocentric plants, whose cell division properties differ from monocentrics.

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.

Limited dispersal and local adaptation promote allopatric speciation in a biodiversity hotspot.

Recently diverged or diverging populations can offer unobstructed insights into early barriers to gene flow during the initial stages of speciation. The current study utilised a novel insect system (order Mantophasmatodea) to shed light on the early drivers of speciation. The members of this group have limited dispersal abilities, small allopatric distributions and strong habitat associations in the Cape Floristic Region biodiversity hotspot in South Africa. Sister taxa from the diverse family Austrophasmatidae were chosen as focal species (Karoophasma biedouwense, K. botterkloofense). Population genetics and Generalized Dissimilarity Modelling (GDM) were used to characterise spatial patterns of genetic variation and evaluate the contribution of environmental factors to population divergence and speciation. Extensive sampling confirmed the suspected allopatry of these taxa. However, hybrids were identified in a narrow region occurring between the species' distributions. Strong population structure was found over short geographic distances; particularly in K. biedouwense in which geographic distance accounted for 32% of genetic variation over a scale of 50 km (r = .56, p < .001). GDM explained 42%-78% of the deviance in observed genetic dissimilarities. Geographic distance was consistently indicated to be important for between species and within population differentiation, suggesting that limited dispersal ability may be an important neutral driver of divergence. Temperature, altitude, precipitation and vegetation were also indicated as important factors, suggesting the possible role of adaptation to local environmental conditions for species divergence. The discovery of the hybrid-zone, and the multiple allopatric species pairs in Austrophasmatidae support the idea that this could be a promising group to further our understanding of speciation modes.

Biomechanics of nectar feeding explain flower orientation in plants pollinated by long-proboscid flies.

The function of flower orientation is much debated, with adaptation to pollinator mouthparts being a particularly compelling explanation, but also one that has lacked empirical support from broad-scale comparative studies. The two families of long-proboscid fly pollinators show similar hovering behaviour while feeding on nectar but differ in the biomechanics of their proboscides which can be up to 80 mm in length: Tabanidae have a fixed forward-pointing proboscis while Nemestrinidae can swivel their proboscis downwards. We predicted that this difference has implications for the evolution of flower orientation. We established the flower angles of 156 South African plant species specialised for pollination by long-proboscid flies. Using a phylogenetically corrected analysis, we found that flowers pollinated by Tabanidae tend to be horizontally orientated, while those pollinated by Nemestrinidae tend to be more variable in orientation and more often vertically orientated. These results confirm the importance of pollinator biomechanics for the evolution of floral traits and highlight a potential mechanism of reproductive isolation between sympatric plant species pollinated by different long-proboscid fly families.

Niche specificity influences gene flow across fine-scale habitat mosaics in Succulent Karoo plants.

While the tempo of diversification in biodiversity hotspots has received much attention, the spatial scale of diversification has often been overlooked. Addressing this deficiency requires understanding the drivers of population divergence and the spatial scales at which they operate in species-rich clades and ecosystems. South Africa's Succulent Karoo (SK) hotspot provides an excellent system for such research, being both compact (ca. 110,000 km2 ) and home to spectacular in-situ radiations, such as the ruschioid Aizoaceae. Here we use GBS to document genetic structure in two co-occurring ruschioid species, at both coarse (>10 km) and fine (<500 m) spatial scales. Where Ruschia burtoniae shows strong between-population genetic differentiation and no gene flow, Conophytum calculus shows weak differentiation, with high levels of admixture suggesting recent or ongoing gene flow. Community analysis and transplant experiments reveal that R. burtoniae occupies a narrow, low-pH edaphic niche, and at scales of a few hundred metres, areas of elevated genetic turnover correspond to patches of edaphically unsuitable habitat. In contrast, C. calculus occupies a broader niche and exhibits isolation-by-distance without a habitat effect. We suggest that edaphic specialisation, coupled with highly restricted seed and pollen dispersal in heterogeneous landscapes, has played a major role in driving rapid diversification at small spatial scales in this system. However, the contrasting patterns in our study species show that these factors do not influence all organisms uniformly, being strongly modulated by lineage-specific traits that influence both the spatial scale of gene flow and habitat specificity.

Intraspecific trait variation influences physiological performance and fitness in the South Africa shrub genus Protea (Proteaceae).

BACKGROUND AND AIMS: Global plant trait datasets commonly identify trait relationships that are interpreted to reflect fundamental trade-offs associated with plant strategies, but often these trait relationships are not identified when evaluating them at smaller taxonomic and spatial scales. In this study we evaluate trait relationships measured on individual plants for five widespread Protea species in South Africa to determine whether broad-scale patterns of structural trait (e.g. leaf area) and physiological trait (e.g. photosynthetic rates) relationships can be detected within natural populations, and if these traits are themselves related to plant fitness. METHODS: We evaluated the variance structure (i.e. the proportional intraspecific trait variation relative to among-species variation) for nine structural traits and six physiological traits measured in wild populations. We used a multivariate path model to evaluate the relationships between structural traits and physiological traits, and the relationship between these traits and plant size and reproductive effort. KEY RESULTS: While intraspecific trait variation is relatively low for structural traits, it accounts for between 50 and 100 % of the variation in physiological traits. Furthermore, we identified few trait associations between any one structural trait and physiological trait, but multivariate regressions revealed clear associations between combinations of structural traits and physiological performance (R2 = 0.37-0.64), and almost all traits had detectable associations with plant fitness. CONCLUSIONS: Intraspecific variation in structural traits leads to predictable differences in individual-level physiological performance in a multivariate framework, even though the relationship of any particular structural trait to physiological performance may be weak or undetectable. Furthermore, intraspecific variation in both structural and physiological traits leads to differences in plant size and fitness. These results demonstrate the importance of considering measurements of multivariate phenotypes on individual plants when evaluating trait relationships and how trait variation influences predictions of ecological and evolutionary outcomes.

Unexpected diversity and evolutionary lability in root architectural ecomorphs in the rushes of the hyperdiverse Cape flora.

Plants use roots to access soil resources, so differences in root traits and their ecological consequences could be a mechanism of species coexistence and niche divergence. Current views of the evolution of root diversity are informed by large-scale evolutionary analyses based on taxonomically coarse sampling and led to the 'root trait phylogenetic conservatism hypothesis'. Here we test this hypothesised conservatism among closely related species, and whether root variation plays an ecological role. We collected root architectural traits for the species-rich Cape rushes (Restionaceae) in the field and from herbaria. We used machine learning to interpolate missing data. Using model-based clustering we classified root syndromes. We modelled the proportion of the syndromes along environmental gradients using assemblages and environmental data of 735 plots. We fitted trait evolutionary models to test for the conservatism hypothesis. We recognised five root syndromes. Responses to environmental gradients are syndrome specific and thus these represent ecomorphs. Trait evolutionary models reveal an evolutionary lability in these ecomorphs. This could present the mechanistic underpinning of the taxonomic radiation of this group which has been linked to repeated habitat shifts. Our results challenge the perspective of strong phylogenetic conservatism and root trait evolution may more generally drive diversification.

Altered ignition catchments threaten a hyperdiverse fire-dependent ecosystem.

Human activities affect fire in many ways, often unintentionally or with considerable time-lags before they manifest themselves. Anticipating these changes is critical, so that insidious impacts on ecosystems, their biodiversity and associated goods and services can be avoided, mitigated or managed. Here we explore the impact of anthropogenic land cover change on fire and biodiversity in adjacent ecosystems on the hyperdiverse Cape Peninsula, South Africa. We develop a conceptual framework based on the notion of an ignition catchment, or the spatial extent and temporal range where an ignition is likely to result in a site burning. We apply this concept using fire models to estimate spatial changes in burn probability between historical and current land cover. This change layer was used to predict the observed record of fires and forest encroachment into fire-dependent Fynbos ecosystems in Table Mountain National Park. Urban expansion has created anthropogenic fire shadows that are modifying fire return intervals, facilitating a state shift to low-diversity, non-flammable forest at the expense of hyperdiverse, flammable Fynbos ecosystems. Despite occurring in a conservation area, these ecosystems are undergoing a hidden collapse and desperately require management intervention. Anthropogenic fire shadows can be caused by many human activities and are likely to be a universal phenomenon, not only contributing to the observed global decline in fire activity but also causing extreme fires in ecosystems where there is no shift to a less flammable state and flammable fuels accumulate. The ignition catchment framework is highly flexible and allows detection or prediction of changes in the fire regime, the threat this poses for ecosystems or fire risk and areas where management interventions and/or monitoring are required. Identifying anthropogenic impacts on ignition catchments is key for both understanding global impacts of humans on fire and guiding management of human-altered landscapes for desirable outcomes.

Environmental pressures on stomatal size may drive plant genome size evolution: evidence from a natural experiment with Cape geophytes.

BACKGROUND AND AIMS: The idea that genome (size) evolution in eukaryotes could be driven by environmental factors is still vigorously debated. In extant plants, genome size correlates positively with stomatal size, leading to the idea that conditions enabling the existence of large stomata in fossil plants also supported growth of their genome size. We test this inductive assumption in drought-adapted, prostrate-leaved Cape (South Africa) geophytes where, compared with their upright-leaved geophytic ancestors, stomata develop in a favourably humid microclimate formed underneath their leaves. METHODS: Stomatal parameters (leaf cuticle imprints) and genome size (flow cytometry) were measured in 16 closely related geophytic species pairs from seven plant families. In each pair, representing a different genus, we contrasted a prostrate-leaved species with its upright-leaved phylogenetic relative, the latter whose stomata are exposed to the ambient arid climate. KEY RESULTS: Except for one, all prostrate-leaves species had larger stomata, and in 13 of 16 pairs they also had larger genomes than their upright-leaved relatives. Stomatal density and theoretical maximum conductance were less in prostrate-leaved species with small guard cells (<1 pL) but showed no systematic difference in species pairs with larger guard cells (>1 pL). Giant stomata were observed in the prostrate-leaved Satyrium bicorne (89-137 µm long), despite its relatively small genome (2C = 9 Gbp). CONCLUSIONS: Our results imply that climate, through selection on stomatal size, might be able to drive genome size evolution in plants. The data support the idea that plants from 'greenhouse' geological periods with large stomata might have generally had larger genome sizes when compared with extant plants, though this might not have been solely due to higher atmospheric CO2 in these periods but could also have been due to humid conditions prevailing at fossil deposit sites.

Post-pollination barriers enable coexistence of pollinator-sharing ornithophilous Erica species.

Some evolutionary radiations produce a number of closely-related species that continue to coexist. In such plant systems, when pre-pollination barriers are weak, relatively strong post-pollination reproductive barriers are required to maintain species boundaries. Even when post-pollination barriers are in place, however, reproductive interference and pollinator dependence may strengthen selection for pre-pollination barriers. We assessed whether coexistence of species from the unusually speciose Erica genus in the fynbos biome, South Africa, is enabled through pre-pollination or post-pollination barriers. We also tested for reproductive interference and pollinator dependence. We investigated this in natural populations of three bird-pollinated Erica species (Erica plukenetii, E. curviflora and E. coccinea), which form part of a large guild of congeneric species that co-flower and share a single pollinator species (Orange-breasted Sunbird Anthobaphes violacea). At least two of the three pre-pollination barriers assessed (distribution ranges, flowering phenology and flower morphology) were weak in each species pair. Hand-pollination experiments revealed that seed set from heterospecific pollination (average 8%) was significantly lower than seed set from outcross pollination (average 50%), supporting the hypothesis that species boundaries are maintained through post-pollination barriers. Reproductive interference, assessed in one population by applying outcross pollen three hours after applying heterospecific pollen, significantly reduced seed set compared to outcross pollen alone. This may drive selection for traits that enhance pre-pollination barriers, particularly given that two of the three species were self-sterile, and therefore pollinator dependent. This study suggests that post-pollination reproductive barriers could facilitate the coexistence of congeneric species, in a recent radiation with weak pre-pollination reproductive barriers.

Intensifying postfire weather and biological invasion drive species loss in a Mediterranean-type biodiversity hotspot.

Prolonged periods of extreme heat or drought in the first year after fire affect the resilience and diversity of fire-dependent ecosystems by inhibiting seed germination or increasing mortality of seedlings and resprouting individuals. This interaction between weather and fire is of growing concern as climate changes, particularly in systems subject to stand-replacing crown fires, such as most Mediterranean-type ecosystems. We examined the longest running set of permanent vegetation plots in the Fynbos of South Africa (44 y), finding a significant decline in the diversity of plots driven by increasingly severe postfire summer weather events (number of consecutive days with high temperatures and no rain) and legacy effects of historical woody alien plant densities 30 y after clearing. Species that resprout after fire and/or have graminoid or herb growth forms were particularly affected by postfire weather, whereas all species were sensitive to invasive plants. Observed differences in the response of functional types to extreme postfire weather could drive major shifts in ecosystem structure and function such as altered fire behavior, hydrology, and carbon storage. An estimated 0.5 °C increase in maximum temperature tolerance of the species sets unique to each survey further suggests selection for species adapted to hotter conditions. Taken together, our results show climate change impacts on biodiversity in the hyperdiverse Cape Floristic Region and demonstrate an important interaction between extreme weather and disturbance by fire that may make flammable ecosystems particularly sensitive to climate change.

Leaps and bounds: geographical and ecological distance constrained the colonisation of the Afrotemperate by Erica.

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.

Should I stay or should I go? Pollinator shifts rather than cospeciation dominate the evolutionary history of South African Rediviva bees and their Diascia host plants.

Plant-pollinator interactions are often highly specialised, which may be a consequence of co-evolution. Yet when plants and pollinators co-evolve, it is not clear if this will also result in frequent cospeciation. Here, we investigate the mutual evolutionary history of South African oil-collecting Rediviva bees and their Diascia host plants, in which the elongated forelegs of female Rediviva have been suggested to coevolve with the oil-producing spurs of their Diascia hosts. After controlling for phylogenetic nonindependence, we found Rediviva foreleg length to be significantly correlated with Diascia spur length, suggestive of co-evolution. However, as trait correlation could also be due to pollinator shifts, we tested if cospeciation or pollinator shifts have dominated the evolution of Rediviva-Diascia interactions by analysing phylogenies in a cophylogenetic framework. Distance-based cophylogenetic analyses (PARAFIT, PACo) indicated significant congruence of the two phylogenies under most conditions. Yet, we found that phylogenetic relatedness was correlated with ecological similarity (the spectrum of partners that each taxon interacted with) only for Diascia but not for Rediviva, suggesting that phylogenetic congruence might be due to phylogenetic tracking by Diascia of Rediviva rather than strict (reciprocal) co-evolution. Furthermore, event-based reconciliation using a parsimony approach (CORE-PA) on average revealed only 11-13 cospeciation events but 58-80 pollinator shifts. Probabilistic cophylogenetic analyses (COALA) supported this trend (8-29 cospeciations vs. 40 pollinator shifts). Our study suggests that diversification of Diascia has been largely driven by Rediviva (phylogenetic tracking, pollinator shifts) but not vice versa. Moreover, our data suggest that, even in co-evolving mutualisms, cospeciation events might occur only infrequently.

Crassula, insights into an old, arid-adapted group of southern African leaf-succulents.

The Crassulaceae is an important family in the Greater Cape Floristic Region of southern Africa and is the seventh largest family in the arid Succulent Karoo Biome. After the Aizoaceae it is the largest group of leaf-succulents in southern Africa. This is the first investigation of a broad selection (68%) of the ±170 species of Crassula. We used data from three chloroplast and two nuclear gene-regions, which yielded many informative characters and provided good resolution among the species. We show that only five of the 20 sections in Crassula are monophyletic. However, the clades recovered show close correlation with the two subgenera that were once recognized. Crassula contains more than 25 succulent annual species which are not closely related to each other but form early-diverging branches in each of the three major clades. One of these major clades contains far more perennial species than the others and is the greatest diversification within Crassula. This diversification mostly arose within the last 10 million years (my) and spread across much of southern Africa. Members of the smaller two major clades are often soft- and flat-leaved perennials (many with basic chromosome number x = 8, with high levels of polyploidy). Those in the largest diversification (where a basic chromosome number of x = 7 predominates) show other arid-adaptations (more highly succulent leaves with a dense covering of hairs or papillae or a smooth xeromorphic epidermis). Their flowers are also more variable in shape and bee-, moth- and butterfly-pollinated species are known among them. We establish that Crassula arose in the Greater Cape Floristic Region of southern Africa. While much of its diversity has evolved in the last 10 my, Crassula nevertheless contains species that are much older and itself arose ±46 my ago. Since all its species are succulent it is possible that they are part of an early arid-adapted flora that contributed to the Succulent Karoo Biome in the western part of southern Africa. Consequently this Biome may not be assembled only from 'young lineages' as is usually thought to be the case.

Microscale trait-environment associations in two closely-related South African shrubs.

PREMISE OF THE STUDY: Plant traits are often associated with the environments in which they occur, but these associations often differ across spatial and phylogenetic scales. Here we study the relationship between microenvironment, microgeographical location, and traits within populations using co-occurring populations of two closely related evergreen shrubs in the genus Protea. METHODS: We measured a suite of functional traits on 147 plants along a single steep mountainside where both species occur, and we used data-loggers and soil analyses to characterize the environment at 10 microsites spanning the elevational gradient. We used Bayesian path analyses to detect trait-environment relationships in the field for each species. We used complementary data from greenhouse grown seedlings derived from wild collected seed to determine whether associations detected in the field are the result of genetic differentiation. KEY RESULTS: Microenvironmental variables differed substantially across our study site. We found strong evidence for six trait-environment associations, although these differed between species. We were unable to detect similar associations in greenhouse-grown seedlings. CONCLUSIONS: Several leaf traits were associated with temperature and soil variation in the field, but the inability to detect these in the greenhouse suggests that differences in the field are not the result of genetic differentiation.

Wandering spiders recover more slowly than web-building spiders after fire.

Fire is a natural feature of many ecosystems, with some vegetation types highly adapted to fire. However, very little is known about the effect of fire on spiders, especially as fires have become more frequent owing to human activity. We determine whether different spider functional guilds (web builders vs. wanderers) respond differently to fires in the sclerophyllous fynbos. We determine also the effect of rockiness as refuge for these guilds and whether it influences their post-fire recovery. There were three site categories of time-since-last fire: 3 months, 1 year, and 7 years. We found that fire caused a decline in spider richness and abundance, with the 3-month category supporting the lowest. In sites that were burned within 1 year, abundance of wanderers was as high as in sites that had 7 years to recover, whereas species richness and abundance of web builders in sites that were burned 1 year ago were as low as in recently burned sites. However, assemblages of wanderers differed among categories, while no differences were observed for web builders, highlighting that wanderers took longer time to recover than web builders. Species richness and abundance of both guilds were not affected by different levels of rockiness. However, rockiness is important in shaping assemblages of wanderers. The results emphasize that the assemblages of greatest conservation concern with increased fire frequencies are wanderers and are candidate surrogates for monitoring post-fire recovery. These results highlight the need to allow fynbos vegetation to recover fully between fire intervals and draws attention to the dangers of frequent unplanned fires.