Biome specificity of distinct genetic lineages within the four-striped mouse Rhabdomys pumilio (Rodentia: Muridae) from southern Africa with implications for taxonomy

Within southern Africa, a link between past climatic changes and faunal diversification has been hypothesized for a diversity of taxa. To test the hypothesis that evolutionary divergences may be correlated to vegetation changes (induced by changes in climate), we selected the widely distributed four-striped mouse, Rhabdomys, as a model. Two species are currently recognized, the mesic-adapted R. dilectus and arid-adapted R. pumilio. However, the morphology-based taxonomy and the distribution boundaries of previously described subspecies remain poorly defined. The current study, which spans seven biomes, focuses on the spatial genetic structure of the arid-adapted R. pumilio (521 specimens from 31 localities), but also includes limited sampling of the mesic-adapted R. dilectus (33 specimens from 10 localities) to act as a reference for interspecific variation within the genus. The mitochondrial COI gene and four nuclear introns (Eef1a1, MGF, SPTBN1, Bfib7) were used for the construction of gene trees. Mitochondrial DNA analyses indicate that Rhabdomys consists of four reciprocally monophyletic, geographically structured clades, with three distinct lineages present within the arid-adapted R. pumilio. These monophyletic lineages differ by at least 7.9% (±0.3) and these results are partly confirmed by a multilocus network of the combined nuclear intron dataset. Ecological niche modeling in MaxEnt supports a strong correlation between regional biomes and the distribution of distinct evolutionary lineages of Rhabdomys. A Bayesian relaxed molecular clock suggests that the geographic clades diverged between 3.09 and 4.30Ma, supporting the hypothesis that the radiation within the genus coincides with paleoclimatic changes (and the establishment of the biomes) characterizing the Miocene-Pliocene boundary. Marked genetic divergence at the mitochondrial DNA level, coupled with strong nuclear and mtDNA signals of non-monophyly of R. pumilio, support the notion that a taxonomic revision of the genus is needed.     

Molecular genetics of Rhabdomys pumilio subspecies boundaries: mtDNA phylogeography and karyotypic analysis by fluorescence in situ hybridization

The phylogeography of the African four-striped mouse, Rhabdomys pumilio, was investigated using complete sequences of the mtDNA cytochrome b gene (1140 bp) and a combination of fluorescence in situ hybridization (FISH) and conventional cytogenetic banding techniques (G- and C-banding). Two cytotypes (2n=46 and 2n=48) were identified by cytogenetic analysis. There is no evidence of diploid number variation within populations, difference in gross chromosome morphology or of subtle interchromosomal rearrangements at levels detected by ZOO-FISH. Analysis of the mtDNA cytochrome b resulted in two major lineages that correspond roughly to the xeric and mesic biotic zones of southern Africa. One mtDNA clade comprises specimens with 2n=48 and the other representatives of two cytotypes (2n=48 and 2n=46). The mean sequence divergence (12%, range 8.3-15.6%) separating the two mtDNA clades is comparable to among-species variation within murid genera suggesting their recognition as distinct species, the prior names for which would be R. dilectus and R. pumilio. Low sequence divergences and the diploid number dichotomy within the mesic lineage support the recognition of two subspecies corresponding to R. d. dilectus (2n=46) and R. d. chakae (2n=48). Our data do not support subspecific delimitation within the nominate, R. pumilio. Molecular dating places cladogenesis of the two putative species at less than five million years, a period characterised by extensive climatic oscillations which are thought to have resulted in habitat fragmentation throughout much of the species range.     

Phylogeographic patterns in populations of the black-chinned tilapia complex (Teleostei,Cichlidae) from coastal areas in West Africa: support for the refuge zone theory

Within the tilapiines, a major African cichlid lineage, quite limited genetic informations are available on phylogenetic interrelationships and phylogeographical patterns at both macro- and micro-evolutionary scales. The present study examines the genetic diversity of the black-chinned tilapia complex from coastal areas in West Africa (Senegal to Congo-Brazzaville) based on mtDNA control region sequences. Phylogenetic inferences provide support for the monophyly of both taxa involved. It is suggested that Sarotherodon melanotheron and Sarotherodon nigripinnis have diverged probably since the early Pleistocene. The occurrence of the main intraspecific lineages could be dated back to about 500,000-900,000 years. Our data suggest that West African Pleistocene refuge zones may have strongly influenced the degree and level of genetic differentiation among populations and thus the observed phylogeographic patterns. The spatial distribution of mtDNA lineages and the centres of intrapopulational genetic diversity clearly correspond to proposed lowland refuge zones and core areas of Central Africa. Moreover, this study genetically supports the existence of Sarotherodon melanotheron leonensis for the first time and further confirms the validity of the most recently introduced subspecies, e.g., Sarotherodon nigripinnis dolloi.     

Diversification across biomes in a continental lizard radiation

Ecological opportunity is a powerful driver of evolutionary diversification, and predicts rapid lineage and phenotypic diversification following colonization of competitor‐free habitats. Alternatively, topographic or environmental heterogeneity could be key to generating and sustaining diversity. We explore these hypotheses in a widespread lineage of Australian lizards: the Gehyra variegata group. This clade occurs across two biomes: the Australian monsoonal tropics (AMT), where it overlaps a separate, larger bodied clade of Gehyra and is largely restricted to rocks; and in the larger Australian arid zone (AAZ) where it has no congeners and occupies trees and rocks. New phylogenomic data and coalescent analyses of AAZ taxa resolve lineages and their relationships and reveal high diversity in the western AAZ (Pilbara region). The AMT and AAZ radiations represent separate radiations with no difference in speciation rates. Most taxa occur on rocks, with small geographic ranges relative to widespread generalist taxa across the vast central AAZ. Rock‐dwelling and generalist taxa differ morphologically, but only the lineage‐poor central AAZ taxa have accelerated evolution. This accords with increasing evidence that lineage and morphological diversity are poorly correlated, and suggests environmental heterogeneity and refugial dynamics have been more important than ecological release in elevating lineage diversity.     

Divergence of mate recognition in the African striped mouse (Rhabdomys)

Mate recognition systems (MRSs) are complex signal-receivertraits. The present study addressed the roles of phylogeny,ecology, and geography in shaping the MRS of the African stripedmouse (Rhabdomys), which has a wide distribution in southernAfrica. Two putative species are recognized, which have differentecologies: Rhabdomys pumilio (arid) and Rhabdomys dilectus (mesic).The latter may be further subdivided into 2 subspecies (R. dilectusdilectus and R. dilectus chakae). Using 2 discrete populationsper taxon, we investigated within- and between-taxon variationsin male odor quality and female perception using habituation-discriminationand habituation-generalization tests, and female preferencein 2-way choice tests. Our results indicate: 1) no within-taxonvariation in odor quality, perception, or preference; 2) the2 subspecies of R. dilectus carry signals of different qualitiesbut share a common odor characteristic distinct from that ofR. pumilio; 3) female R. pumilio did not show a preference whentheir own species and R. d. chakae odors were presented simultaneouslybut displayed assortative preference when the alternative wasR. d. dilectus; 4) females of the 2 subspecies showed dissimilarpreferences: R. d. chakae for the genetically more similar taxonand R. d. dilectus for the most different one. Although we couldnot rule out the influence of ecology, we concluded that phylogenyappeared a more parsimonious explanation for the pattern ofdivergence in Rhabdomys. Further, we discuss our results inlight of current models on signal-receiver coevolution.     

Phylogeny and cryptic diversity in geckos (Phyllopezus; Phyllodactylidae; Gekkota) from South America's open biomes

The gecko genus Phyllopezus occurs across South America's open biomes: Cerrado, Seasonally Dry Tropical Forests (SDTF, including Caatinga), and Chaco. We generated a multi-gene dataset and estimated phylogenetic relationships among described Phyllopezus taxa and related species. We included exemplars from both described Phyllopezus pollicaris subspecies, P. p. pollicaris and P. p.przewalskii. Phylogenies from the concatenated data as well as species trees constructed from individual gene trees were largely congruent. All phylogeny reconstruction methods showed Bogertia lutzae as the sister species of Phyllopezus maranjonensis, rendering Phyllopezus paraphyletic. We synonymized the monotypic genus Bogertia with Phyllopezus to maintain a taxonomy that is isomorphic with phylogenetic history. We recovered multiple, deeply divergent, cryptic lineages within P. pollicaris. These cryptic lineages possessed mtDNA distances equivalent to distances among other gekkotan sister taxa. Described P. pollicaris subspecies are not reciprocally monophyletic and current subspecific taxonomy does not accurately reflect evolutionary relationships among cryptic lineages. We highlight the conservation significance of these results in light of the ongoing habitat loss in South America's open biomes.     

Birth of a biome: insights into the assembly and maintenance of the Australian arid zone biota

The integration of phylogenetics, phylogeography and palaeoenvironmental studies is providing major insights into the historical forces that have shaped the Earth's biomes. Yet our present view is biased towards arctic and temperate/tropical forest regions, with very little focus on the extensive arid regions of the planet. The Australian arid zone is one of the largest desert landform systems in the world, with a unique, diverse and relatively well-studied biota. With foci on palaeoenvironmental and molecular data, we here review what is known about the assembly and maintenance of this biome in the context of its physical history, and in comparison with other mesic biomes. Aridification of Australia began in the Mid-Miocene, around 15 million years, but fully arid landforms in central Australia appeared much later, around 1-4 million years. Dated molecular phylogenies of diverse taxa show the deepest divergences of arid-adapted taxa from the Mid-Miocene, consistent with the onset of desiccation. There is evidence of arid-adapted taxa evolving from mesic-adapted ancestors, and also of speciation within the arid zone. There is no evidence for an increase in speciation rate during the Pleistocene, and most arid-zone species lineages date to the Pliocene or earlier. The last 0.8 million years have seen major fluctuations of the arid zone, with large areas covered by mobile sand dunes during glacial maxima. Some large, vagile taxa show patterns of recent expansion and migration throughout the arid zone, in parallel with the ice sheet-imposed range shifts in Northern Hemisphere taxa. Yet other taxa show high lineage diversity and strong phylogeographical structure, indicating persistence in multiple localised refugia over several glacial maxima. Similar to the Northern Hemisphere, Pleistocene range shifts have produced suture zones, creating the opportunity for diversification and speciation through hybridisation, polyploidy and parthenogenesis. This review highlights the opportunities that development of arid conditions provides for rapid and diverse evolutionary radiations, and re-enforces the emerging view that Pleistocene environmental change can have diverse impacts on genetic structure and diversity in different biomes. There is a clear need for more detailed and targeted phylogeographical studies of Australia's arid biota and we suggest a framework and a set of a priori hypotheses by which to proceed.     

Geographical patterns of old and young species in African forest biota: the significance of specific montane areas as evolutionary centres

A widely accepted paradigm for speciation in tropical forests, the refuge theory, requires periodic habitat fragmentation driven by global climatic fluctuations to provide conditions for allopatric speciation. This implies that comparative species richness in refugia is due to loss of diverse communities in areas affected by climatic cycles. In this study we compare distribution patterns of bird and plant taxa which we consider to be of either deep phylogenetic lineages or recent radiations. It is demonstrated that lowland areas which have been postulated as Pleistocene refugia are dominated by species which represent lineages of pre-Pleistocene age. Since variations in species richness within these forest tracts reflect currently apparent environmental variables which might be considered to determine carrying capacity, we do not need to postulate that richness is the result of changes in forest cover in the past. Recently diversified taxa of plants and birds are found mainly at the periphery of the main rain forest blocks and in habitat islands outside them. Here, peak concentrations of young restricted-range species are often congruent with clusters of old and biogeographically relictual species. It is suggested that this reflects special intrinsic environmental properties of these areas, in the form of long-term environmental stability caused mainly by persistent orographic rain or mist. In this case, richness is not necessarily due to extinction outside these areas. Stability not only enables survival of relictual taxa, but also promotes morphological differentiation of radiating taxa, leading to aggregates of taxa of restricted distribution.     

A phylogenetic review of the African leaf chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation

The phylogenetic associations among 13 currently recognized African leaf chameleon species were investigated by making use of mitochondrial and nuclear DNA sequence data (44 taxa and 4145 characters). The gene tree indicates two divergent clades within Rhampholeon; this finding is congruent with previous morphological suggestions. The first clade (I) comprises three taxa (R. kerstenii, R. brevicaudatus and R. brachyurus) and is widely distributed in lowland forest and or non-forest biomes. The second clade (II) comprises the remaining Rhampholeon species and can be subdivided into three subclades. By contrast, most taxa belonging to clade II are confined to relict montane forest biotopes. Based on geographical, morphological and molecular evidence, it is suggested that the taxonomy of Rhampholeon be revised to include two genera (Rieppeleon and Rhampholeon) and three subgenera (Rhampholeon, Bicuspis and Rhinodigitum). There is a close correlation between geographical distribution and phylogenetic relatedness among Rhampholeon taxa, indicating that vicariance and climate change were possibly the most influential factors driving speciation in the group. A relaxed Bayesian clock suggests that speciation times coincided both with the northern movement of Africa, which caused the constriction of the pan African forest, and to rifting in east Africa ca. 20 Myr ago. Subsequent speciation among taxa was probably the result of gradual desiccation of forests between 20 and 5 Myr ago.     

Macro-scale assessment of demographic and environmental variation within genetically derived evolutionary lineages of eastern hemlock (<Emphasis Type="Italic">Tsuga canadensis</Emphasis>), an imperiled conifer of the eastern United States

Eastern hemlock (Tsuga canadensis) occupies a large swath of eastern North America and has historically undergone range expansion and contraction resulting in several genetically separate lineages. This conifer is currently experiencing mortality across most of its range following infestation of a non-native insect. With the goal of better understanding the current and future conservation potential of the species, we evaluate ecological differences among populations within these genetically defined clusters, which were previously inferred using nuclear microsatellite molecular markers from 58 eastern hemlock populations. We sub-divide these clusters into four genetic zones to differentiate putative north-central, north-east and southeast (SE) and southwest evolutionary lineages in eastern hemlock. We use demographic data (relative abundance, mortality, and seedling regeneration) from the Forest Inventory Analysis program in conjunction with environmental data to model how these lineages respond to current and future climatic gradients. Ecologically meaningful relationships are explored in the intraspecific context of hemlock abundance distribution and then related to genetic variation. We also assess hemlock’s colonization likelihood via a long distance dispersal model and explore its future genetic and ecological conservation potential by combining the future suitable habitats with colonization likelihoods. Results show that future habitats under climate change will markedly decline for eastern hemlock. The remaining areas with higher habitat quality and colonization potential are confined to the SE, the genetic zone nearest the species’ putative glacial refugia, pointing to the need to focus our conservation efforts on this ecologically and genetically important region.     

Ecological niche modelling of an invasive alien plant and its potential biological control agents

Invasive alien plants are of concern in South Africa. Pompom weed (Campuloclinium macrocephalum) is currently invading the Grassland and Savannah biomes of South Africa and is likely to continue spreading in the southern African sub- region. Two possible biological control agents (Liothrips tractabilis and Cochylis campuloclinium) have been identified for control of pompom weed. We used ecological niche modelling to predict which areas in southern Africa are likely to be suitable for pompom weed and the two potential biological control agents. The overlap between areas predicted to be highly suitable for pompom weed and areas suitable for the biological control agents was assessed. Methods of reducing sampling bias in a data set used for calibrating models were also compared. Finally, the performance of models calibrated using only native range data, only invaded range data and both were also compared. Models indicate that pompom weed is likely to spread across a greater region of southern Africa than it currently occupies, with the Savannah and Grassland biomes being at greatest risk of invasion. Poor overlap was found between the areas predicted to be highly suitable for pompom weed and those areas predicted to be suitable for the biological control agents. However, models of the potential distribution of the biological control agents are interpreted with caution due to the very small sample size of the data set used to calibrate the models. Models calibrated using both native range and invaded range data were found to perform best whilst models calibrated using only native range data performed the worst. There was little difference found between models that were calibrated using spatially reduced (selecting only one record per 30 min grid cell) and randomly reduced (randomly selecting 50% of available records) biased data sets.     

Quantity versus quality: Endemism and protected areas in the temperate forest of South America

Abstract Identification of biodiversity hotspots is essential to conservation strategies aimed at minimizing the possibility of losing half of the world's species in the next 50 years. The aims of the present study were: (i) to locate and designate zones of endemism in the temperate forest of South America; and (ii) to compare the distribution of these areas with the distribution of existing protected areas in this habitat type. Endemism areas were determined by using parsimonious analysis of endemism, which identified zones of endemism on the basis of sets of endemic species that were restricted to two or more study areas. We used distribution information for five unrelated taxa (ferns, trees, reptiles, birds and mammals) to provide more reliable results and patterns than would work with only a single taxon or related taxa. The northern part of this region has high endemism for all of the taxa considered in this study. We demonstrate that although the temperate forest of South America has more than 30% of its area under some type of protection, correlation between protected areas and the areas of endemism is remarkably low. In fact, less than 10% of protected areas are situated in areas that have the greatest value for conservation (i.e. high endemism). Under the current strategy, biodiversity within South America's temperate forest is in danger despite the large amount of protected area for this forest type.     

Identifying contact zone hotspots of passerine birds in the Palaearctic region

Birds are known to be a group rich in pairs of closely related species that have parapatric or allopatric distributions with relatively narrow contact zones. Here we analyse the geographical distribution of these contact zones for parapatric species pairs of passerine birds of the Palaearctic region. Their contact zones are located mainly in southwestern, northern and central-southern Asia, and in northwestern Africa, with a hotspot in the Middle East. A mid-domain effect null model, where contact zone hotspots are a neutral correlate of continental geometry, had a low explanatory power of 3.8%; the observed distribution of contact zones was not sufficiently predicted. Hypotheses involving range contractions and secondary contact in areas of high topographic and habitat diversity may offer more convincing explanations and offer promising perspectives for future studies.     

Radiation of southern African daisies: biogeographic inferences for subtribe Arctotidinae (Asteraceae, Arctotideae)

The majority of the approximately 80-90 species in subtribe Arctotidinae occur in southern Africa with the centre of diversity in the winter-rainfall region. Three species are restricted to afromontane eastern Africa and three species are endemic to Australia. To investigate biogeographic and phylogenetic relationships within Arctotidinae, sequence data from four cpDNA regions (psbA-trnH, trnT-trnL and trnL-trnF spacers and trnL intron) and the ITS nrDNA region for 59 Arctotidinae species were analyzed with parsimony and Bayesian-inference approaches. Eight well-supported major lineages were resolved. The earliest-diverging extant lineages are afromontane or inhabit mesic habitats, whereas almost all sampled taxa from the winter-rainfall and semi-arid areas have diverged more recently. Molecular dating estimated that the major clades diverged during the Miocene and Pliocene, which is coincident with the trend of increasing rainfall seasonality, aridification and vegetation changes in southwestern Africa. Trans-oceanic dispersal to Australia was estimated to have occurred during the Pliocene.     

Habitat characteristics and species interference influence space use and nest‐site occupancy: implications for social variation in two sister species

Nest‐site selection is an important component of species socio‐ecology, being a crucial factor in establishment of group living. Consequently, nest‐site characteristics together with space‐use proxies may reveal the social organization of species, which is critical when direct observation of social interactions is hindered in nature. Importantly, nest‐site choice is expected to be under strong selective pressures and the object of intra‐ and interspecific competition. Although the bulk of research on sociality focuses on its ecological drivers, our study introduces interspecific competition as a potential factor that could influence social evolution. We investigated the influence of habitat and interspecific competition on the social organization of two sister species of the African four striped mouse, Rhabdomys dilectus dilectus and Rhabdomys bechuanae, in a similar macroenvironment. These species diverged in allopatry and occupy distinct environmental niches. We radiotracked 140 adults to identify their nest‐sites, determine nest characteristics and record groups that shared nest‐sites. Group cohesion was estimated from nest‐site fidelity, group association strength, and home range overlap within versus between group members. We compared the two species in sympatry versus parapatry to determine the impact of species interference on sociality. In parapatry, the two species selected distinct nest‐site types, interpreted as different anti‐predator strategies: R. bechuanae selected fewer, spaced, less concealed nest‐sites whereas R. d. dilectus selected clumped and less visible nest‐sites. Rhabdomys bechuanae also showed more cohesive and stable social groups than R. d. dilectus. In sympatry, compared to R. bechuanae, R. d. dilectus occupied similar nest‐sites, however slightly more exposed and clumped, and displayed similar nest‐site fidelity and group association strength. We conclude that although habitat selection may be an important driver of social divergence in Rhabdomys, species interference, by limiting R. d. dilectus movements and forcing nest‐site sharing may induce new ecological pressures that could influence its social evolution.     

The second Southern African Bird Atlas Project: Causes and consequences of geographical sampling bias

Using the Southern African Bird Atlas Project (SABAP2) as a case study, we examine the possible determinants of spatial bias in volunteer sampling effort and how well such biased data represent environmental gradients across the area covered by the atlas. For each province in South Africa, we used generalized linear mixed models to determine the combination of variables that explain spatial variation in sampling effort (number of visits per 5′ × 5′ grid cell, or “pentad”). The explanatory variables were distance to major road and exceptional birding locations or “sampling hubs,” percentage cover of protected, urban, and cultivated area, and the climate variables mean annual precipitation, winter temperatures, and summer temperatures. Further, we used the climate variables and plant biomes to define subsets of pentads representing environmental zones across South Africa, Lesotho, and Swaziland. For each environmental zone, we quantified sampling intensity, and we assessed sampling completeness with species accumulation curves fitted to the asymptotic Lomolino model. Sampling effort was highest close to sampling hubs, major roads, urban areas, and protected areas. Cultivated area and the climate variables were less important. Further, environmental zones were not evenly represented by current data and the zones varied in the amount of sampling required representing the species that are present. SABAP2 volunteers' preferences in birding locations cause spatial bias in the dataset that should be taken into account when analyzing these data. Large parts of South Africa remain underrepresented, which may restrict the kind of ecological questions that may be addressed. However, sampling bias may be improved by directing volunteers toward undersampled regions while taking into account volunteer preferences.     

Home away from home — objective mapping of high-risk source areas for plant introductions

Prevention is the best way to slow the escalation of problems associated with biological invasions. Screening of potential introductions is widely applied for assessing the risk of species becoming invasive. Despite advances in the understanding of the determinants of invasiveness, screening still relies heavily on assessments of the potential of species to ‘fit in’ to the broad environmental conditions of a target region. Most screening systems ask whether species are native to, or are known to be naturalized or invasive in, regions with ‘similar’ climatic/environmental conditions to the target region. The level of similarity required to make the species a high‐risk introduction is generally not specified. This paper describes a protocol for making such assessments more objective, using South Africa as a test case. Using nonparametric niche‐based modelling (generalized additive model; GAM) calibrated on the current distribution of each South African biome, we mapped regions of the world that are climatically similar to South African biomes. Lists were produced of countries with the largest areas climatically similar to South Africa overall, and to each biome separately. Validation of the usefulness of the approach was sought by evaluating whether the main invasive plant species in South African biomes occur naturally, or have adventive ranges, in regions mapped as analogous to South African biomes. A very large part of the world is climatically similar to South Africa, with eight countries having larger areas of land classified as climatically similar to South African biomes than the total area of South Africa. Almost all the most prominent invasive species in South African biomes occur naturally or are invasive outside their natural range in areas with similar climates to those that occur in parts of South Africa. This confirms the value of objective climate matching in screening protocols. We examined climatic conditions for a representative sample of major invasive plants from other parts of the world. The analysis identified several species that are already invasive in regions that have matched climates in South Africa but that are not yet introduced or, if already present, have not yet invaded large areas. For example, the following known invasive species should be considered high‐risk species in South African grasslands: Alliaria petiolata, Cytisus scoparius, Gleditsia triacanthos, Heracleum mantegazzianum, Hieracium pilosella, Juniperus communis, Pinus contorta, P. monticola, P. ponderosa, P. sylvestris, Prunus laurcerasus, and P. serotina. Objectively matched climatic regions are also useful as a first‐cut assessment when evaluating species with no invasive history.     

Space Use Variation in Co-Occurring Sister Species: Response to Environmental Variation or Competition?

Coexistence often involves niche differentiation either as the result of environmental divergence, or in response to competition. Disentangling the causes of such divergence requires that environmental variation across space is taken into account, which is rarely done in empirical studies. We address the role of environmental variation versus competition in coexistence between two rodent species: Rhabdomys bechuanae (bechuanae) and Rhabdomys dilectus dilectus (dilectus) comparing their habitat preference and home range (HR) size in areas with similar climates, where their distributions abut (allopatry) or overlap (sympatry). Using Outlying Mean Index analyses, we test whether habitat characteristics of the species deviate significantly from a random sample of available habitats. In allopatry, results suggest habitat selection: dilectus preferring grasslands with little bare soil while bechuanae occurring in open shrublands. In sympatry, shrubland type habitats dominate and differences are less marked, yet dilectus selects habitats with more cover than bechuanae. Interestingly, bechuanae shows larger HRs than dilectus, and both species display larger HRs in sympatry. Further, HR overlaps between species are lower than expected. We discuss our results in light of data on the phylogeography of the genus and propose that evolution in allopatry resulted in adaptation leading to different habitat preferences, even at their distribution margins, a divergence expected to facilitate coexistence. However, since sympatry occurs in sites where environmental characteristics do not allow complete species separation, competition may explain reduced inter-species overlap and character displacement in HR size. This study reveals that both environmental variation and competition may shape species coexistence.     

Origin and diversification of the Greater Cape flora: ancient species repository, hot-bed of recent radiation, or both?

Like island-endemic taxa, whose origins are expected to postdate the appearance of the islands on which they occur, biome-endemic taxa should be younger than the biomes to which they are endemic. Accordingly, the ages of biome-endemic lineages may offer insights into biome history. In this study, we used the ages of multiple lineages to explore the origin and diversification of two southern African biomes whose remarkable floristic richness and endemism has identified them as global biodiversity hotspots (succulent karoo and fynbos). We used parsimony optimization to identify succulent karoo- and fynbos-endemic lineages across 17 groups of plants, for which dated phylogenies had been inferred using a relaxed Bayesian (BEAST) approach. All succulent karoo-endemic lineages were less than 17.5 My old, the majority being younger than 10 My. This is largely consistent with suggestions that this biome is the product of recent radiation, probably triggered by climatic deterioration since the late Miocene. In contrast, fynbos-endemic lineages showed a broader age distribution, with some lineages originating in the Oligocene, but most being more recent. Also, in groups having both succulent karoo- and fynbos-endemic lineages, there was a tendency for the latter to be older. These patterns reflect the greater antiquity of fynbos, but also indicate considerable recent speciation, probably through a combination of climatically-induced refugium fragmentation and adaptive radiation.