Genetic variation in Rhipicephalus appendiculatus (Acari: Ixodidae) from Zambia: correlating genetic and ecological variation with Rhipicephalus appendiculatus from eastern and southern Africa.

Based on their ecology, Rhipicephalus appendiculatus ticks from eastern and southern Africa have been divided into three groups. We investigated how two geographic genetically differentiated stocks of R. appendiculatus from the southern and the eastern provinces of Zambia, representing two ecological groups, i.e., southern African and transition groups, respectively, genetically compare to stocks from east Africa (Rwanda) and southern Africa (Zimbabwe and South Africa). The ITS2 and two mitochondrial genes segments, 12s rDNA and COI, were used in the investigations. The ITS2 tree did not show support for differentiation into any groups, while the two mitochondrial genes trees (12s rDNA and COI) showed two genetically differentiated groups: an east African genetic group which included specimens from Rwanda and the plateau area of the eastern province of Zambia, and a southern African genetic group represented by specimens from South Africa, Zimbabwe and specimens collected on the fringes of the eastern province plateau in the Nyimba district of Zambia. This suggests that the two geographically differentiated stocks of the southern and eastern provinces of Zambia might be part of two wider geographic genetically differentiated R. appendiculatus groups that extend beyond Zambia. Stocks of "transition" ecology (eastern province) belong to the east African genetic group and the differences in ecology within this genetic grouping may be due to genetic polymorphism, phenotypic plasticity, and other local factors.     

Regional genetic structuring and evolutionary history of the impala Aepyceros melampus

Samples of 162 impala antelope (Aepyceros melampus) from throughout its distribution range in sub-Saharan Africa were surveyed using eight polymorphic microsatellite loci. Furthermore, 155 previously published mitochondrial DNA (mtDNA) sequences from the same localities were reanalyzed. Two subspecies of impala are presently recognized--the isolated black-faced impala (Aepyceros melampus petersi) in southwest Africa and the common impala (Aepyceros melampus melampus) abundant in southern and east Africa. All tests performed indicated significant genetic differentiation at the subspecific level. Furthermore, individual-based analyses split the common impala subspecies into two distinct genetic groups, conforming with regional geographic affiliation to southern or east Africa. This was supported by assignment tests, genetic distance measures, pairwise theta values, and analysis of molecular variance. We suggest that the presence of such previously unknown regional structuring within the subspecies reflects a pattern of colonization from a formerly large panmictic population in southern Africa toward east Africa. This scenario was supported by a progressive decline in population diversity indices toward east Africa and a significant increase in the quantity theta/(1 - theta). Both microsatellite and mtDNA data indicated a genetic distinctiveness of the Samburu population in Kenya.     

A previously unrecognized radiation of ranid frogs in Southern Africa revealed by nuclear and mitochondrial DNA sequences

In sub-Saharan Africa, amphibians are represented by a large number of endemic frog genera and species of incompletely clarified phylogenetic relationships. This applies especially to African frogs of the family Ranidae. We provide a molecular phylogenetic hypothesis for ranids, including 11 of the 12 African endemic genera. Analysis of nuclear (rag-1, rag-2, and rhodopsin genes) and mitochondrial markers (12S and 16S ribosomal RNA genes) provide evidence for an endemic clade of African genera of high morphological and ecological diversity thus far assigned to up to five different subfamilies: Afrana, Cacosternum, Natalobatrachus, Pyxicephalus, Strongylopus, and Tomopterna. This clade has its highest species diversity in southern Africa, suggesting a possible biogeographic connection with the Cape Floral Region. Bayesian estimates of divergence times place the initial diversification of the southern African ranid clade at approximately 62-85 million years ago, concurrent with the onset of the radiation of Afrotherian mammals. These and other African ranids (Conraua, Petropedetes, Phrynobatrachus, and Ptychadena) are placed basally within the Ranoidae with respect to the Eurasian groups, which suggests an African origin for this whole epifamily.     

Re-evaluation of species limits in the genus Atlapetes based on mtDNA sequence data

Brush-finches of the genus Atlapetes present a complicated pattern of differentiation along the Andes. Forms with different colours may replace each other sharply at different altitudes on the same slopes, in similar humid habitats on different slopes or on adjacent humid and semi-humid slopes. In the current classification, grey and yellow-and-green species are placed in different groups, and similar-coloured populations replacing each other in corresponding ecological zones on different slopes are ranked as subspecies of the same species. However, Remsen and Graves have suggested that different-coloured forms inhabiting adjacent mountains could be representatives of the same species with different pigment saturations, rather than competing species. The relationships between and within Atlapetes species are assessed based on mtDNA sequence data. Three main branches are demonstrated, comprising southern (Bolivian, northwest Argentinean), central (south and central Peruvian] and northern/western species. Clearly, the traditional groupings based on plumage colours did not demonstrate evolutionary trajectories in the genus. The data also show that many forms, currently ranked as subspecies, are genetically more divergent than sympatric species. Species rank is suggested for several forms hitherto treated as subspecies of geographically disjunct species.     

Differentiation and phylogeny of the olivaceous warbler Hippolais pallida species complex

The pattern of phenotypic and molecular variation within the polytypic olivaceous warbler H. pallida was examined. This species is distributed in the southern parts of the western Palaearctic, central Asia and in the arid parts of northern Africa, and also in parts of the sub-Saharan Sahel zone. Based on morphology, five subspecies, at times assigned to three groups, have been identified. By comparing morphological, behavioural, vocal and molecular variation, we investigated the phenotypic and phylogenetic relationships within the Hippolais pallida group. The morphological and genetic data of the present study support the view of splitting the olivaceous warbler into a western (former subspecies opaca) and an eastern form (former subspecies elaeica, pallida, reiseri and laeneni). Opaca is consistently and significantly larger than the other taxa in all size measurements, even if it does share morphological properties with elaeica in characters associated with migration. The song of the polytypic pallida differs clearly from that of opaca, and is remarkably consistent within its wide range which comprises four subspecies. The molecular analysis, and the resulting phylogenetic pattern, clearly separated opaca on a single branch distant from the other subspecies. The eastern form consists of two non-overlapping haplotype groups: elaeica with relatively diverse mtDNA variation, and the three African subspecies pallida, reiseri and laeneni, which all share the same or closely related mtDNA haplotypes. It remains open whether the African taxa should be regarded as three valid subspecies of the Eastern olivaceous warbler, or if they are better treated as a single African subspecies.     

Contrasting patterns of radiation in African and Australian Restionaceae

The floras of the Mediterranean-climate areas of southern Africa and southwestern Australia are remarkably species rich. Because the two areas are at similar latitudes and in similar positions on their respective continents, they have probably had similar Cenozoic climatic histories. Here we test the prediction that the evolution of the species richness in the two areas followed a similar temporal progression by comparing the rates of lineage accumulation for African and Australian Restionaceae. Restionaceae (Poales) are typical and often dominant elements in the fynbos vegetation of the Cape Floristic Region of southern Africa and the kwongan vegetation of the Southwestern Floristic Province of Western Australia. The phylogeny of the family was estimated from combined datasets for rbcL and trnL-F sequences and a large morphological dataset; these datasets are largely congruent. The monophyly of Restionaceae is supported and a basal division into an African clade (approximately 350 species) and an Australian clade (146 species) corroborated. There is also support for a futher subdivision of these two large sister-clades, but the terminal resolution within the African clade is very weak. Fossil pollen records provided a minimum age of the common ancestor of Australian and African Restionaceae as 64-71 million years ago, and this date was used to calibrate a molecular clock. A molecular clock was rejected by a likelihood ratio test; therefore, rate changes between the lineages were smoothed using nonparametric rate smoothing. The rate-corrected ages were used to construct a plot of lineages through time. During the Palaeogene the Australian lineage diversity increased consistent with the predictions of the constant birthrate model, while the African lineage diversity showed a dramatic increase in diversification rate in the Miocene. Incomplete sampling obscures the patterns in the Neogene, but extending the trends to the modern extant diversity suggests that this acceleration in the speciation rate continued in the African clade, whereas the Australian clade retained a constant diversification rate. The substantial morphological and anatomical similarity between the African and Australian Restionaceae appear to preclude morphological innovations as possible explanations for the intercontinental differences. Most likely these differences are due to the greater geographical extent and ecological variation in temperate Australia than temperate Africa, which might have provided refugia for basal Restionaceae lineages, whereas the more mountainous terrain of southern Africa might have provided the selective regimes for a more rapid, recent speciation.     

Where are you from,stranger? The enigmatic biogeography of North African pond turtles (Emys orbicularis)

The European pond turtle (Emys orbicularis) is a Nearctic element in the African fauna and thought to have invaded North Africa from the Iberian Peninsula. All North African populations are currently identified with the subspecies E. o. occidentalis. However, a nearly range-wide sampling in North Africa used for analyses of mitochondrial and microsatellite DNA provides evidence that only Moroccan populations belong to this taxon, while eastern Algerian and Tunisian pond turtles represent an undescribed distinct subspecies. These two taxa are most closely related to E. o. galloitalica with a native distribution along the Mediterranean coast of northern Spain through southern France to western and southern Italy. This group is sister to a clade comprising several mitochondrial lineages and subspecies of E. orbicularis from Central and Eastern Europe plus Asia, and the successive sisters are E. o. hellenica and E. trinacris. Our results suggest that E. orbicularis has been present in North Africa longer than on the Iberian Peninsula and that after an initial invasion of North Africa by pond turtles from an unknown European source region, there was a phase of diversification in North Africa, followed by a later re-invasion of Europe by one of the African lineages. The differentiation of pond turtles in North Africa parallels a general phylogeographic paradigm in amphibians and reptiles, with deeply divergent lineages in the western and eastern Maghreb. Acknowledging their genetic similarity, we propose to synonymize the previously recognized Iberian subspecies E. o. fritzjuergenobsti with E. o. occidentalis sensu stricto. The seriously imperiled Moroccan populations of E. o. occidentalis represent two Management Units different in mitochondrial haplotypes and microsatellite markers. The conservation status of eastern Algerian pond turtles is unclear, while Tunisian populations are endangered. Considering that Algerian and Tunisian pond turtles represent an endemic taxon, their situation throughout the historical range should be surveyed to establish a basis for conservation measures.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

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.     

Regional variation in the cheetah (Acinonyx jubatus) revisited: Morphology of wild and captive populations

The cheetah (Acinonyx jubatus) is listed as a vulnerable species by the International union for the conservation of nature (IUCN), including two critically endangered subspecies, the Saharan cheetah, and the Iranian cheetah, so it is imperative that we understand variation in cheetah morphology to make good decisions regarding the conservation of this species. Here, we aim to determine whether northeastern African cheetahs have smaller body sizes than southern African cheetahs. This study also adds to our knowledge of cheetah morphology from two cheetah populations that do not yet have comprehensive published data: Kenya, and northeastern Africa, including captive individuals. We calculated means and standard deviations on cranial and body measurements of live or in few cases, freshly dead, cheetahs from the aforementioned populations, plus previously published data on Namibian and Botswanan cheetahs and compared them to one another using multivariate analysis of variance. Results show that northeastern African cheetahs have smaller body sizes than southern and eastern African populations. We also found that captive cheetahs retain the morphological characteristics of their ancestral population- captive cheetahs from southern Africa have similar body sizes to wild southern African cheetahs and larger body sizes than captives from northeastern Africa. Other analyses regarding cheetah growth agree with previous studies on Namibian and Botswanan cheetah populations rates. As such, this study can serve as a baseline for the care of captive cheetah populations to maintain healthy weights and body proportions.     

Patterns of stable isotope signatures in willow warbler Phylloscopus trochilus feathers collected in Africa

We conducted stable isotope analyses of nitrogen and carbon on feathers obtained from willow warblers in Africa to find an explanation for a previously observed pattern of different δ¹⁵N and δ¹³C values across a migratory divide in central Scandinavia. A new data set confirms that north Scandinavian birds of the subspecies P. t. acredula have higher δ¹⁵N values than south Scandinavian birds of the subspecies P. t. trochilus . In Africa, we found significant differences for both δ¹⁵N and δ¹³C values among feathers collected from major geographical regions as well as between countries within regions. Isotope signatures of δ¹⁵N and δ¹³C in Scandinavian acredula matched well with those of willow warblers sampled in southern parts of Africa, but differed from samples obtained in East and West Africa. Isotope signatures in Scandinavian trochilus did not agree with the pattern in any of the three African regions (West, East or South). However, a more detailed analysis of the isotopic data in feathers from countries within West Africa, which is the wintering region of Swedish trochilus based on ringing recoveries, revealed a correspondence with samples from Liberia, the Ivory Coast and Nigeria.     

The pollination niche and its role in the diversification and maintenance of the southern African flora

The flora of southern Africa has exceptional species richness and endemism, making it an ideal system for studying the patterns and processes of evolutionary diversification. Using a wealth of recent case studies, I examine the evidence for pollinator-driven diversification in this flora. Pollination systems, which represent available niches for ecological diversification, are characterized in southern Africa by a high level of ecological and evolutionary specialization on the part of plants, and, in some cases, by pollinators as well. These systems are asymmetric, with entire plant guilds commonly specialized for a particular pollinator species or functional type, resulting in obvious convergent floral evolution among guild members. Identified modes of plant lineage diversification involving adaptation to pollinators in these guilds include (i) shifts between pollination systems, (ii) divergent use of the same pollinator, (iii) coevolution, (iv) trait tracking, and (v) floral mimicry of different model species. Microevolutionary studies confirm that pollinator shifts can be precipitated when a plant species encounters a novel pollinator fauna on its range margin, and macroevolutionary studies confirm frequent pollinator shifts associated with lineage diversification. As Darwin first noted, evolutionary specialization for particular pollinators, when resulting in ecological dependency, may increase the risk of plant extinction. I thus also consider the evidence that disturbance provokes pollination failure in some southern African plants with specialized pollination systems.     

Evolutionary history of the Persian Jird,Meriones persicus,based on genetics,species distribution modelling and morphometric data

The Persian Jird, Meriones persicus, is distributed from Eastern Anatolia to Afghanistan and western Pakistan. Six subspecies were described based on skull features and coat colours, but the validity of these subspecies is uncertain, and no molecular work has ever been conducted on this species. Iran appears to be a key geographical region in which to revise the systematic and evolutionary history of this species, because five of the six subspecies are present in this country. To evaluate the phylogeographical history and taxonomy of this species in Iran, we used a combination of genetic (cytochrome b gene sequences of 70 specimens) and geometric morphometric (2D landmarks on the ventral side of skull of 258 specimens) analyses. We also used ecological niche modelling to make inferences about the evolutionary history of these lineages. Our molecular data highlight the existence of four genetic lineages, but they only partly correspond to the previously described subspecies. Our molecular and morphometric data confirm the validity of M. p. rossicus and show that it has a wider geographical range than previously thought. M. p. gurganensis and M. p. baptistae are genetically very close. The skull of M. p. gurganensis is morphologically distinguishable from other subspecies. The subspecies M. p. persicus and M. p. baptistae are genetically distinct, but morphologically close. Meriones p. ambrosius is genetically close to M. p. persicus, and additional analyses with more specimens are needed to validate its subspecific status. The genetic structure observed in Iran seems to fit the topography and biogeography of the country and emphasize the role of the Abarkooh, Central and Lut deserts as barriers to gene flow. All intraspecific divergent events within the Persian Jird occurred during the last 1.4 My, suggesting that climatic changes probably trigger diversification within this species. Our genetic and species niche modelling results suggest that potential refugial areas persisted during glacial periods for this species in north‐western Zagros Mountains, north‐eastern Alborz Mountains and Kohrud Mountains.     

Song differences between populations of seven subspecies of the African forest weaver Ploceus bicolor Vieillot (Aves: Passeriformes)

We present an analysis of song type differences between populations of seven forest weaver (Ploceus bicolor) subspecies. Phonology and syntax of the melodious parts of song types differ between these populations to the same extent as they differ between population-specific dialects within a subspecies. Phonological and syntactical song features are socially transmitted; they can be transmitted even between representatives of taxonomically different subspecies. The Harsh Call, a complex element embedded in the melody, is of similar structure in populations of four South African subspecies, but shows markedly different characteristics in populations of the subspecies from Kenya and Cameroon. Song differences between populations are suggested to result from cultural drift and geographic isolation. A correspondence between genetically determined morphological structures on the one hand and socially transmitted song structure on the other seems to be a coincidence due to geographic separation.     

Nuclear and mtDNA-based phylogeny of southern African sand lizards, Pedioplanis (Sauria: Lacertidae)

The diversity of lacertid lizards in Africa is highest in the southern African subcontinent, where over two-thirds of the species are endemic. With eleven currently recognized species, Pedioplanis is the most diverse among the southern African genera. In this study we use 2200 nucleotide positions derived from two mitochondrial markers (ND2 and 16S rRNA) and one nuclear gene (RAG-1) to (i) assess the phylogeny of Pedioplanis and (ii) estimate divergence time among lineages using the relaxed molecular clock method. Individual analyses of each gene separately supported different nodes in the phylogeny and the combined analysis yielded more well supported relationships. We present the first, well-resolved gene tree for the genus Pedioplanis and this is largely congruent with a phylogeny derived from morphology. Contrary to previous suggestions Heliobolus/Nucras are sister to Pedioplanis. The genus Pedioplanis is monophyletic, with P. burchelli/P. laticeps forming a clade that is sister to all the remaining congeners. Two distinct geographic lineages can be identified within the widespread P. namaquensis; one occurs in Namibia, while the other occurs in South Africa. The P. undata species complex is monophyletic, but one of its constituent species, P. inornata, is paraphyletic. Relationships among the subspecies of P. lineoocellata are much more complex than previously documented. An isolated population previously assigned to P. l. pulchella is paraphyletic and sister to the three named subspecies. The phylogeny identifies two biogeographical clades that probably diverged during the mid-Miocene, after the development of the Benguella Current. This probably led to habitat changes associated with climate and, in conjunction with physical barriers (Great Escarpment), contributed towards speciation within the genus Pedioplanis.     

Genetic diversification without obvious genitalic morphological divergence in harvestmen (Opiliones, Laniatores, Sclerobunus robustus) from montane sky islands of western North America

The southern Rocky Mountains and adjacent Intermontane Plateau Highlands region of western North America is a geographically diverse area with an active geologic history. Given the topological complexity and extensive geologic activity, organisms inhabiting this region are expected to show some degree of morphological and genetic divergence, especially populations found on the southern montane 'sky islands' of this region. Here we examine the phylogeographic history and diversification of a montane forest inhabiting harvestmen, Sclerobunus robustus, using a combination of genetic and morphological data. Divergence time estimates indicate that much of the diversification within and between major groups S. robustus predate the Pleistocene glacial cycles. The most widespread subspecies, Sclerobunus robustus robustus, is recovered as six genetically distinct, geographically cohesive mitochondrial phylogroups. Gene tree data for a single nuclear gene reveals congruent, albeit slightly more conservative, patterns of genetic divergence. Despite high levels of genetic divergence throughout their distribution, phylogroups show extreme conservation in somatic and reproductive morphology. This uncoupling of morphological and genetic differentiation may be due to morphological conservatism associated with a conserved microhabitat preference. Based on these data, it is obvious that S. robustus has undergone some level of cryptic diversification.     

Taxonomy of the African giant pouched rats (Nesomyidae: Cricetomys): molecular and craniometric evidence support an unexpected high species diversity

Our study combined a mitochondrial cytochrome b phylogeny with cranial measurements from giant pouched rats collected across sub‐Saharan Africa. The mitochondrial phylogeny resolves two West African clades and a clade with east and central Africa representatives. This last clade can be further divided into four subclades. Altogether they represent six species (Cricetomys gambianus, Cricetomys ansorgei, Cricetomys emini, and three undescribed taxa) that can be distinguished on the basis of their mitochondrial DNA sequences and craniometry. In the absence of adequate craniometric data the existence of Cricetomys kivuensis cannot be confirmed by our data. Our combined molecular and craniometric data allowed us to broadly delineate the distribution ranges of the detected species. Cricetomys gambianus occurs in the savannah and forest clearings of West Africa. Cricetomys ansorgei is distributed in the savannah of East and southern Africa. Cricetomys emini, as currently recognized across the Guineo‐Congolian forest of Africa, is shown to be diphyletic. Cricetomys sp. 1, a separate operational taxonomic unit closely resembling C. emini, occurs in the forest zone of West Africa. An undescribed sister‐species of C. ansorgei, Cricetomys sp. 2, occurs in the forest of Central Africa along the left bank of the Congo River. Cricetomys sp. 3 occurs on the right bank of the Congo River from Cameroon to the Republic of Congo, whereas the true C. emini also occurs on the right bank of the Congo River but appears to be restricted to the Democratic Republic of Congo. Cranial phenotype within the genus tends to conform to ecological zonation (either forest or savannah) rather than to phylogenetic affiliation of the species concerned, suggesting that diversifying selection across environmental gradients could be responsible for biological diversification within the genus.     

Molecular probes of phylogeny and biogeography in toads of the widespread genus Bufo

Genetic relationships among 25 species of Central and South American Bufo and among representative North, Central, and South American, Asian, and African Bufo were probed, using the quantitative immunological technique of microcomplement fixation (MC'F) which indicated a clear separation of North, Central, and South American lineages of Bufo. The South American lineage likely diverged from the Central and North American lineages in the Eocene; the latter two lineages diverged later, probably in the mid-Oligocene. Some species groups of South American toads, defined on the basis of traditional morphological studies, are genetically quite similar within groups, whereas others are genetically divergent. The amount of albumin evolution does not appear to parallel the amount of karyotypic, morphological, ecological, or behavioral evolution documented. Comparisons suggest that the African lineages separated from the American and Asian lineages in the late Cretaceous, corresponding to the time of the final separation of Gondwanaland, the southern supercontinent including the modern continents of South America, Africa, Australia, Antarctica, and India. The Asian lineages diverged from the lineage giving rise to all of the American species in the early Paleocene.      

Tectonics,climate and the diversification of the tropical African terrestrial flora and fauna

Tropical Africa is home to an astonishing biodiversity occurring in a variety of ecosystems. Past climatic change and geological events have impacted the evolution and diversification of this biodiversity. During the last two decades, around 90 dated molecular phylogenies of different clades across animals and plants have been published leading to an increased understanding of the diversification and speciation processes generating tropical African biodiversity. In parallel, extended geological and palaeoclimatic records together with detailed numerical simulations have refined our understanding of past geological and climatic changes in Africa. To date, these important advances have not been reviewed within a common framework. Here, we critically review and synthesize African climate, tectonics and terrestrial biodiversity evolution throughout the Cenozoic to the mid-Pleistocene, drawing on recent advances in Earth and life sciences. We first review six major geo-climatic periods defining tropical African biodiversity diversification by synthesizing 89 dated molecular phylogeny studies. Two major geo-climatic factors impacting the diversification of the sub-Saharan biota are highlighted. First, Africa underwent numerous climatic fluctuations at ancient and more recent timescales, with tectonic, greenhouse gas, and orbital forcing stimulating diversification. Second, increased aridification since the Late Eocene led to important extinction events, but also provided unique diversification opportunities shaping the current tropical African biodiversity landscape. We then review diversification studies of tropical terrestrial animal and plant clades and discuss three major models of speciation: (i) geographic speciation via vicariance (allopatry); (ii) ecological speciation impacted by climate and geological changes, and (iii) genomic speciation via genome duplication. Geographic speciation has been the most widely documented to date and is a common speciation model across tropical Africa. We conclude with four important challenges faced by tropical African biodiversity research: (i) to increase knowledge by gathering basic and fundamental biodiversity information; (ii) to improve modelling of African geophysical evolution throughout the Cenozoic via better constraints and downscaling approaches; (iii) to increase the precision of phylogenetic reconstruction and molecular dating of tropical African clades by using next generation sequencing approaches together with better fossil calibrations; (iv) finally, as done here, to integrate data better from Earth and life sciences by focusing on the interdisciplinary study of the evolution of tropical African biodiversity in a wider geodiversity context.     

Out of Africa: biogeographic history of the open‐habitat chats (Aves,Muscicapidae: Saxicolinae) across arid areas of the old world

Arid and semi‐arid areas constitute a prominent feature of the earth today, especially in Asia and Africa. Their formation started in the middle Miocene with increased stepwise aridification since the Pliocene. This aridification had strong ecological and evolutionary consequences and not only led to fragmentation of moist‐adapted biota, but also fostered the evolution of arid‐adapted taxa from mesic ancestors and triggered speciation within arid areas. The open‐habitat chats, a clade within Saxicolinae (Aves, Muscicapidae), constitute one of the most significant arid‐adapted passerine groups of Africa and Eurasia. Here, we present a temporal and spatial framework for the diversification of open‐habitat chats, using probabilistic approaches for the reconstruction of their biogeographic history based on a time‐calibrated multilocus molecular phylogenetic hypothesis. The diversification of open‐habitat chats was initiated in the late Miocene at around 7.4 Ma, most likely in sub‐Saharan Africa. Southern Africa and the Horn of Africa acted as centres of diversification and biogeographic expansion. From the latter area, the Arabo‐Sindic region and subsequently further parts of Eurasia and North Africa were colonized. The colonization history out of sub‐Saharan Africa contrasts with that of several other songbird clades, where a biogeographic expansion from Eurasia or northern Africa to southern Africa was prevalent. Habitat fragmentation through forest expansions during intermittent wetter periods in Africa influenced diversification in several clades. However, phases of increased aridity, with hyperarid regions acting as drivers of vicariance, seem to have also been important in radiations of the Arabo‐Sindic region and the Horn of Africa during the Pleistocene. Different processes such as colonization of new areas followed by vicariance or speciation across ecotones might have played a role throughout the radiation of open‐habitat chats.