Nicola G. Bergh

Cape diversification and repeated out-of-southern-Africa dispersal in paper daisies (Asteraceae-Gnaphalieae).

The large daisy tribe Gnaphalieae occurs in extra-tropical habitats worldwide, but is most diverse in southern Africa and in Australia. We explore the age and evolutionary history of the tribe by means of a phylogenetic hypothesis based on Bayesian analysis of plastid and nuclear DNA sequences, maximum likelihood reconstruction of ancestral areas, and relaxed Bayesian dating. Early diversification occurred in southern Africa in the Eocene-Oligocene, resulting in a grade of mostly Cape-centred lineages which subsequently began speciating in the Miocene, consistent with diversification times for many Cape groups. Gnaphalieae from other geographic regions are embedded within a southern African paraphylum, indicating multiple dispersals out of southern Africa since the Oligocene to Miocene which established the tribe in the rest of the world. Colonisation of Australia via direct long-distance trans-oceanic dispersal in the Miocene resulted in the radiation which produced the Australasian gnaphalioid flora. The similarly diverse regional gnaphalioid floras of Australasia and southern Africa thus exhibit very different temporal species accumulation histories. An examination of the timing and direction of trans-Indian Ocean dispersal events in other angiosperms suggests a role for the West Wind Drift in long-distance dispersal eastwards from southern Africa.

Topography as a driver of diversification in the Cape Floristic Region of South Africa

The rugged topography of the Cape Floristic Region (CFR), South Africa, is frequently invoked to explain the spectacular radiation of the Cape flora, but the mechanisms involved remain unclear. Where recent authors emphasize the importance of elevation gradients as stimuli for ecological speciation, earlier workers stressed the role of topography as an isolating mechanism, particularly in montane lineages. Using six Cape plant lineages, we tested whether elevation niches are phylogenetically conserved. We then assessed whether high-elevation species are more consistently range-restricted than low-elevation species, and whether high-elevation sisters show stronger range exclusivity (allopatry) and weaker ecological and phenotypic differentiation, suggestive of nonecological speciation. Elevation niches tend to be phylogenetically conserved. Also, high-elevation species are more consistently range-restricted than low-elevation species, potentially explaining the generally stronger range exclusivity of high-elevation sisters. While the high-elevation zone is less homogeneous ecologically, more data are required to demonstrate that high-elevation sister species show generally weaker ecological and phenotypic differentiation. Topographic complexity promotes geographical isolation at high elevations, thereby providing opportunities for nonecological, vicariant speciation. While recognizing the need for additional data, we suggest that the upland and lowland floras of the CFR may differ with regard to predominant speciation mode.

Erosive processes after tectonic uplift stimulate vicariant and adaptive speciation: evolution in an Afrotemperate-endemic paper daisy genus

BackgroundThe role of tectonic uplift in stimulating speciation in South Africa’s only alpine zone, the Drakensberg, has not been explicitly examined. Tectonic processes may influence speciation both through the creation of novel habitats and by physically isolating plant populations. We use the Afrotemperate endemic daisy genus Macowania to explore the timing and mode (geographic versus adaptive) of speciation in this region. Between sister species pairs we expect high morphological divergence where speciation has happened in sympatry (adaptive) while with geographic (vicariant) speciation we may expect to find less morphological divergence and a greater degree of allopatry. A dated molecular phylogenetic hypothesis for Macowania elucidates species’ relationships and is used to address the potential impact of uplift on diversification. Morphological divergence of a small sample of reproductive and vegetative characters, used as a proxy for adaptive divergence, is measured against species’ range distributions to estimate mode of speciation across two subclades in the genus.ResultsThe Macowania crown age is consistent with the hypothesis of post-uplift diversification, and we find evidence for both vicariant and adaptive speciation between the two subclades within Macowania. Both subclades exhibit strong signals of range allopatry, suggesting that geographic isolation was important in speciation. One subclade, associated with dry, rocky environments at high altitudes, shows very little morphological and ecological differentiation but high range allopatry. The other subclade occupies a greater variety of habitats and exhibits far greater morphological differentiation, but contains species with overlapping distribution ranges.ConclusionsSpecies in Macowania are likely to have diversified in response to tectonic uplift, and we invoke uplift and uplift-mediated erosion as the main drivers of speciation. The greater relative morphological divergence in sympatric species of Macowania indicates that speciation in the non-sympatric taxa may not have required obvious adaptive differences, implying that simple geographic isolation was the driving force for speciation (‘neutral speciation’).

Anomalous capitulum structure and monoecy may confer flexibility in sex allocation and life history evolution in the Ifloga lineage of paper daisies (Compositae: Gnaphalieae).

PREMISE OF THE STUDY Evolutionary significance of the Compositae capitulum and variation in its structure is poorly understood, although it may permit flexibility in sexual expression. Optimal sex ratio differs with life-history and reproductive strategy. We explore how the genus Ifloga and related members of southern African Gnaphalieae achieved different sex ratios, and the associations of these ratios with annual and perennial life history. METHODS Sex allocation was measured using the male to female ratio (M/F), a novel approximator of the pollen to ovule ratio (P/O). Life-history (annuality/perenniality), capitulum structure, capitular sexual system, and M/F were reconstructed on time-proportional phylogenies. Trait associations were examined using phylogenetically independent contrasts (PICs). KEY RESULTS Annual taxa have strongly female-biased capitula, as measured by M/F, and either gynomonoecious or monoecious sexual systems, while perennials have equal or male-biased capitula that are hermaphroditic or monoecious. These results are largely supported by PIC analysis. Different sexual systems afford differing flexibility in sex allocation, with hermaphrodites having the least, and monoecious taxa the greatest, range in M/F. Within Ifloga, the anomalous capitulum evolved in an annual, gynomonoecious ancestor, followed by two independent gains of monoecy. Two subsequent gains of perenniality occurred within a monoecious sublineage. CONCLUSIONS Different life histories have divergent sex allocation optima and are strongly associated with different sexual systems in gnaphalioid daisies. An anomalous capitulum structure in Ifloga may have facilitated the evolution of monoecy, which in turn may be linked to the evolution of life-history diversity in the genus.

Dominant pollinators drive non-random community assembly and shared flower colour patterns in daisy communities

Background and Aims As most plants rely on pollination for persistence in communities, pollination interactions should be important determinants of plant community assembly. Here, trait and phylogenetic null modelling approaches were combined with pollinator interaction networks to elucidate the processes structuring flower colour assembly patterns in Asteraceae communities in Namaqualand, South Africa. Methods Plant species were assigned to flower colour pattern categories (CPCs) that incorporate the complexity of the bulls-eye colour pattern, using pollinator vision models. Null models were used to assess whether daisy communities exhibit clustering (driven by filtering, facilitation or convergence) or overdispersion (driven by competitive exclusion or character displacement) of CPCs. Next, flower visitor networks were constructed for communities with non-random CPC assembly to confirm the functional role of pollinators in determining floral trait assembly. Key Results Plant species are unevenly distributed across CPCs, the majority of which are not phylogenetically conserved, suggesting that certain CPCs have a selective advantage. Clustering of CPCs in communities is more frequent than overdispersion, and this does not reflect non-random phylogenetic assembly. In most communities at least one CPC is overrepresented relative to null assemblages. Interaction networks show that each community has a single dominant pollinator that strongly interacts with the overrepresented CPC, suggesting a role for pollinator preferences in driving clustered assembly of CPCs within daisy communities. Conclusion This novel approach, which demonstrates non-random assembly of complex flower colour patterns and corroborates their functional association with particular pollinators, provides strong evidence that pollinators influence plant community assembly. Results suggest that in some community contexts the benefits of pollinator sharing outweigh the costs of heterospecific pollen transfer, generating clustered assembly. They also challenge the perception of generalized pollination in daisies, suggesting instead that complex daisy colour patterns represent a pollination syndrome trait linked to specific fly pollinators.

Species-level phylogenetic analysis in the Relhania clade of "everlastings" and a new generic treatment of species previously assigned to Macowania and Arrowsmithia (Asteraceae: Gnaphalieae)

The “Relhania clade” is a relatively species-poor lineage that is sister to the remaining members of the paper daisy tribe Gnaphalieae. Although the monophyly of the Relhania clade is well established, previous molecular studies are based on fairly sparse sampling. Here, we explore clade membership and generic circumscription using plastid and nuclear DNA sequence data, and near-complete species-level sampling of all putative member genera. Two subclades are recovered within the Relhania clade: (i) clade I housing the genera Alatoseta, Athrixia, Pentatrichia and Phagnalon, all monophyletic as currently circumscribed; and (ii) clade II, containing the genera Antithirixia, Comborhiza, Oedera, Oreoleyesera, Relhania and Rosenia; the annual or otherwise “short-lived” genera Leysera, Nestlera and Rhynchopsidium; as well as a subclade in which the monospecific Arrowsmithia is embedded within Macowania. A new taxonomic treatment involving 12 new combinations is proposed for the two latter genera. Only demonstrably monophyletic and diagnosable genera are recognised, involving the erection of a novel monospecific genus, Fluminaria, to house Macowania pinifolia, and the transfer of the remaining members of Macowania to Arrowsmithia. Updated descriptions and keys are provided.