Phylogeography of two cryptic species of African desert jerboas (Dipodidae: Jaculus)

The lesser Egyptian jerboa Jaculus jaculus is a desert dwelling rodent that inhabits a broad Arabian–Saharan arid zone. Recently, two distant sympatric lineages were described in North‐West Africa, based on morphometric and molecular data, which may correspond to two cryptic species. In the current study, phylogenetic relationships and phylogeographical structure among those lineages and geographical populations from North Africa and the Middle East were investigated. The phylogeographical patterns and genetic diversity of the cytochrome b gene (1110 bp) were addressed on 111 jerboas from 41 localities. We found that the variation in Africa is partitioned into two divergent mitochondrial clades (10.5% divergence relating to 1.65–4.92 Mya) that corresponds to the two cryptic species: J. jaculus and J. deserti. Diversifications within those cryptic species/clades were dated to 0.23–1.13 Mya, suggesting that the Middle Pleistocene climatic change and its environmental consequences affected the evolutionary history of African jerboas. The third distant clade detected, found in the Middle East region, most likely represents a distinct evolutionary unit, independent of the two African lineages. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Molecular and morphological evidence for a Pleistocene radiation of laminate‐toothed rats (Otomys: Rodentia) across a volcanic archipelago in equatorial Africa

West African Mountains of the Cameroon Volcanic Line harbour two montane‐endemic species of laminated‐toothed rats (Otomys), which represent the most westerly occurrence of the genus. We explore here through mtDNA sequencing and cranial morphometrics the taxonomic status and phylogenetic relationships of O. burtoni (Mt Cameroon) and O. occidentalis (Mts Oku and Gotel). We conclude that both species are valid and can be discriminated by molecular data, as well as quantitative and qualitative cranial characters. From molecular data, O. occidentalis and O. burtoni are closest neighbours (p‐distance = 7.5–8.5%) and weakly associated sister species (suggesting a single West African radiation) and both are sister clades to a well supported clade of central, East and northeast African members of the O. typus s.l. and O. tropicalis s.l. species complexes from mountain ranges comprising the East African ‘Montane Circle’ and Ethiopian Highlands. Re‐evaluation of the evolutionary origins of the allopatric Otomys populations in equatorial Africa is undertaken in light of fossil evidence of a southern African origin of the genus. We can conclude that Otomys reached the Cameroon Volcanic Line via corridors of temperate grasslands during the Late Pliocene. Our data support the hypothesis that, following major peripatric speciation events at around 2.3 to 2.03 Ma (from East Africa into West and North Africa respectively), further speciation occurred across neighbouring mountain ranges in West, Central‐East and North‐East Africa. Estimated molecular dates of speciation events in Otomys reveal close congruence with well‐constrained geochronological estimates, pertinently the uplift of the Albertine Rift in the Early Pleistocene. These regional analyses reveal how peripatric speciation events established narrow‐range endemics of Otomys on principal stratovolcanoes across the East African plateau and Cameroon. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 320–344.     

Paternal lineages in Libya inferred from Y‐chromosome haplogroups

Many studies based on genetic diversity of North African populations have contributed to elucidate the modelling of the genetic landscape in this region. North Africa is considered as a distinct spatial‐temporal entity on geographic, archaeological, and historical grounds, which has undergone the influence of different human migrations along its shaping. For instance, Libya, a North African country, was first inhabited by Berbers and then colonized by a variety of ethnic groups like Phoenicians, Greeks, Romans, Arabs and, in recent times, Italians. In this study, we contribute to clarify the genetic variation of Libya and consequently, of North African modern populations, by the study of Libyan male lineages. A total of 22 Y‐chromosome‐specific SNPs were genotyped in a sample of 175 Libyan males, allowing the characterization of 18 Y‐chromosomal haplogroups. The obtained data revealed a predominant Northwest African component represented by haplogroup E‐M81 (33.7%) followed by J(xJ1a,J2)‐M304 (27.4%), which is postulated to have a Middle Eastern origin. The comparative study with other populations (～5,400 individuals from North Africa, Middle East, Sub‐Saharan Africa, and Europe) revealed a general genetic homogeneity among North African populations (F_ST = 5.3 %; P‐value < 0.0001). Overall, the Y‐haplogroup diversity in Libya and in North Africa is characterized by two genetic components. The first signature is typical of Berber‐speaking people (E‐M81), the autochthonous inhabitants, whereas the second is (J(xJ1a,J2)‐M304), originating from Arabic populations. This is in agreement with the hypothesis of an Arabic expansion from the Middle East, shaping the North African genetic landscape. Am J Phys Anthropol 157:242–251, 2015. © 2015 Wiley Periodicals, Inc.     

Cercopithecidae from the Pliocene Chiwondo beds, Malawi-rift

There are a total of 16 fossil cercopithecid specimens, representing at least 10 individuals, from the Chiwondo Beds of northern Malawi. The majority of this material is derived from the Middle Pliocene Unit 3A, but one specimen is from the Early Pliocene Unit 2. This latter specimen is from a papionin of indeterminate genus similar in size to Parapapio ado and Pliopapio alemui. Among the specimens from Unit 3A, two species can be diagnosed: an indeterminate species of Theropithecus, and a species of Parapapio similar in dental size to P. broomi. Neither of the genera from Unit 3A are exclusive to either East Africa or South Africa. Their relative abundances, however, are more similar to Middle Pliocene South African sites where Parapapio is the most common primate genus, and Theropithecus is comparatively rare. This is in contrast to similarly-aged East African sites where Theropithecus is by far the most abundant genus.     

Garra tibanica ghorensis subsp.nov. (Pisces: Cyprinidae), an African element in the cyprinid fauna of the Levant

Garra tibanica ghorensis subsp. nov. is described from the southern Dead Sea Valley. In the Levant it represents the only cyprinid fish with African affinities known to date. It is suggested that G. t. ghorensis reached the area of its present distribution from the south and is not to be regarded as a relict of an earlier migration of the species to Africa via the Levant.Results of the travels of R. Kinzelbach to the countries of the Middle East, No. 18.Results of the travels of R. Kinzelbach to the countries of the Middle East, No. 18.     

New adapiform primate of Old World affinities from the Devil's Graveyard Formation of Texas

Most adapiform primates from North America are members of an endemic radiation of notharctines. North American notharctines flourished during the Early and early Middle Eocene, with only two genera persisting into the late Middle Eocene. Here we describe a new genus of adapiform primate from the Devil’s Graveyard Formation of Texas. Mescalerolemur horneri, gen. et sp. nov., is known only from the late Middle Eocene (Uintan) Purple Bench locality. Phylogenetic analyses reveal that Mescalerolemur is more closely related to Eurasian and African adapiforms than to North American notharctines. In this respect, M. horneri is similar to its sister taxon Mahgarita stevensi from the late Duchesnean of the Devil’s Graveyard Formation. The presence of both genera in the Big Bend region of Texas after notharctines had become locally extinct provides further evidence of faunal interchange between North America and East Asia during the middle Eocene. The fact that Mescalerolemur and Mahgarita are both unknown outside of Texas also supports prior hypotheses that low-latitude faunal assemblages in North America demonstrate increased endemism by the late middle Eocene.     

African origin and global distribution patterns: Evidence inferred from phylogenetic and biogeographical analyses of ectomycorrhizal fungal genus Strobilomyces

Aim

The ectomycorrhizal genus Strobilomyces is widely distributed throughout many parts of the world, but its origin, divergence and distribution patterns remain largely unresolved. In this study, we aim to explore the species diversity, distribution and evolutionary patterns of Strobilomyces on a global scale by establishing a general phylogenetic framework with extensive sampling.

Location

Africa, Australasia, East Asia, Europe, North America, Central America and Southeast Asia.

Methods

The genealogical concordance phylogenetic species recognition method was used to delimit phylogenetic species. Divergence times were estimated using a Bayesian uncorrelated lognormal relaxed molecular clock. The ancestral area and host of Strobilomyces were inferred via the programs rasp and mesquite . The change of diversification rate over time was estimated using Ape, Laser and Bammtools software packages.

Results

We recognize a novel African clade and 49 phylogenetic species with morphological evidence, including 18 new phylogenetic species and 23 previously described ones. Strobilomyces probably originated in Africa, in association with Detarioideae/Phyllanthaceae/Monotoideae during the early Eocene. The dispersal to Southeast Asia can be explained by Wolfe's “Boreotropical migration” hypothesis. East Asia, Australasia, Europe and North/Central America are primarily the recipients of immigrant taxa during the Oligocene or later. A rapid radiation implied by one diversification shift was inferred within Strobilomyces during the Miocene.

Main conclusions

An unexpected phylogenetic species diversity within Strobilomyces was uncovered. The highest diversity, resulting probably from a rapid radiation, was found in East Asia. Dispersal played an important role in the current distribution pattern of Strobilomyces. The Palaeotropical disjunction is explained by species dispersal from Africa to Southeast Asia through boreotropical forests during the early Eocene. Species from the Northern Hemisphere and Australasia are largely derived from immigrant ancestors from Southeast Asia.     

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.     

Origin and taxonomic status of the Palearctic population of the stem borer Sesamia nonagrioides (Lefèbvre) (Lepidoptera: Noctuidae)

The major pest of maize in Mediterranean Europe, the stem borer Sesamia nonagrioides (Lefèbvre) (Lepidoptera: Noctuidae), has a fragmented distribution, north and south of the Sahara. The present study aimed: (1) to clarify the uncertain taxonomic status of the Palearctic and sub‐Saharan populations which were first considered as different species and later on as subspecies (Sesamia nonagrioides nonagrioides and Sesamia nonagrioides botanephaga) and (2) to investigate the origin of the Palearctic population which extends from Spain to Iran, outside what is considered typical for this mainly tropical genus. We reconstructed the evolutionary history of both populations using one nuclear and two mitochondrial genes. The sub‐Saharan taxon was fragmented in two isolated populations (West and East) whose mitochondrial genes were distant by 2.3%. The Palearctic population was included in the East African clade and its genes were close or identical to those of a population from Central Ethiopia, where the species was discovered for the first time. Similarly, in Africa, the alleles of the nuclear gene were distributed mainly in two West and East clades, whereas some Palearctic alleles belonged to the West clade. The Palearctic population originated therefore from East and West Africa and is the progeny of the cross between these two African populations. The main species concepts were in agreement, leading to the conclusion that the three populations are still conspecific. In the surveyed regions, the species therefore does not include two subspecies but three isolated populations. The Palearctic population suffered from severe bottlenecks that resulted in the fixation of one East African mitochondrial genome and the large reduction in its genetic diversity compared to the African populations. The data suggest that natural colonization of the Palearctic region was more plausible than human introduction. The allelic distribution of the Palearctic population was similar to that of species that survived the last glaciation. It is concluded that the African populations expanded during the last interglacial, crossed the Sahara and mixed in North Africa where fixation of the East mitochondrial genome occurred. The species then colonized Europe westward through only one eastern entrance. The coalescent‐based estimate of the time to the ancestor of the Palearctic population was 108 000 years, which is consistent with this scenario. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 904–922.     

The fossil vertebrate locality Kossom Bougoudi,Djurab desert,Chad: A window in the distribution of the carnivoran faunas at the Mio–Pliocene boundary in Africa

The fossil vertebrate bearing locality Kossom Bougoudi (KB) is situated in the Djurab desert (Chad, Africa), 600 km north-east of N’djamena. It has yielded about 1250 specimens with many mammalian remains, principally artiodactyls. Its geologic age has been estimated to be about 5 Ma by biochronologic estimation and about 5.3 Ma by radiometric studies on cosmogenic nuclides of beryllium (authigenic ¹⁰B/⁹B). The carnivoran fauna contains few specimens which belong to five different families. All the taxa were unknown in central Africa. A large lutrine is close to Sivaonyx but different from known species of the genus. Another large lutrine is similar by its size to a species described from the Middle Pliocene of Uganda. An edentulous mandible of a small machairodont cat resembles a small species of Dinofelis, while a distal humerus indicates the presence of a larger member of the same genus. A hunting hyaenid is also much like the European species. An unidentified canid reaches the size of the recent Canis aureus and an isolated calcaneum matches that of the large extant viverrid. This small fauna allows a first look at the guild of the carnivorans at the Latest Miocene–Pliocene boundary in Central Africa and is a milestone between North African, East African and South African carnivore faunas.     

Intercontinental disjunctions in Cryptotaenia (Apiaceae, Oenantheae): an appraisal using molecular data

Aim The angiosperm genus Cryptotaenia (family Apiaceae, tribe Oenantheae) exhibits an anomalous distribution pattern, with five of its eight species being narrow endemics geographically isolated from their presumed relatives. We examined the monophyly of the genus and ascertained the phylogenetic placements of its constituent members in order to explain their distribution patterns. Location Eastern North America, eastern Asia, the Caucasus, southern Italy, Macaronesia and Africa. Methods In total, 173 accessions were examined for nuclear rDNA ITS sequence variation, representing nearly all major lineages of Apiaceae subfamily Apioideae and seven species of Cryptotaenia. Sampling of tribes Oenantheae, Scandiceae and Pimpinelleae was comprehensive. Phylogenetic analyses included Bayesian, maximum parsimony and neighbour‐joining methods; biogeographical scenarios were inferred using dispersal–vicariance analysis (diva ). Results Cryptotaenia is polyphyletic and includes three distant lineages. (1) Cryptotaenia sensu stricto (C. canadensis, C. japonica, C. flahaultii and C. thomasii) is maintained within tribe Oenantheae; C. canadensis and C. japonica, representing an eastern North American–eastern Asian disjunction pattern, are confirmed to be sister species. (2) Cryptotaenia elegans, endemic to the Canary Islands, is placed within Scandiceae subtribe Daucinae along with two woody endemics of Madeira, Monizia edulis and Melanoselinum decipiens. The phylogeny of these Canarian and Madeiran endemics is unresolved. Either they constitute a monophyletic sister group to a clade comprising some Mediterranean and African species of Daucus and their relatives, or they are paraphyletic to this clade. The herbaceous/woody genus Tornabenea from Cape Verde, once included in Melanoselinum, is not closely related to the other Macaronesian endemics but to Daucus carota. (3) The African members of Cryptotaenia (C. africana, C. calycina and possibly C. polygama) comprise a clade with some African and Madagascan umbellifers; this entire clade is sister group to Eurasian Pimpinella. Main conclusions Elucidating the phylogeny of the biogeographically anomalous Cryptotaenia sensu lato enabled hypotheses on the biogeography of its constituent lineages. Cryptotaenia sensu stricto exhibits a holarctic distribution pattern, with its members occurring in regions that were important glacial refugia. The genus probably originated in eastern Asia and from there dispersed to Europe and North America. For the Macaronesian endemic species –C. elegans, M. edulis and M. decipiens–diva reconstructs either a single dispersal event to Macaronesia from the Mediterranean/African region, or a single dispersal followed by a back‐dispersal to the mainland. The radiation of Tornabenea from Cape Verde followed a second dispersal of Daucinae to Macaronesia. Woodiness in Melanoselinum/Monizia and Tornabenea, therefore, is a derived and independently acquired trait. The African members of Cryptotaenia are derived from an ancestor arriving from the Middle East.     

Mitochondrial CR-1 Variation in Sardinian Hares and Its Relationships with Other Old World Hares (Genus Lepus)

Among the European fauna, the Sardinian hare (Lepus sp.) is peculiar in that it differs from all other hares inhabiting the continent. Here, we report on the variation of a 461 bp sequence of hypervariable domain 1 of the mitochondrial control region, examined in 42 hares collected throughout Sardinia and compared to the corresponding sequences of different Lepus taxa. Seventeen novel haplotypes were found in the Sardinian population, resulting in a haplotype diversity of 0.840 and a nucleotide diversity of 0.012. As a result of Bayesian and principal coordinates analyses, Sardinian hares were grouped with North African hares, constituting a monophyletic clade that diverges from all other Old World hares, including Cape hares from South Africa and East Asia. Hence, our data agree that populations inhabiting North Africa and Sardinia form a distinct taxon, which could possibly be included in the L. capensis superspecies. Moreover, two corresponding lineages can be found in Sardinia and Tunisia, providing evidence of a common origin of the two populations and thus supporting the hypothesis that North African hares were introduced into the island in historical times. Our data show that the two lineages differ in their geographic distribution throughout the island and that the wild Sardinian population also shows the signature of a postintroduction demographic expansion.     

Molecular evidence for species level divergence in African Nile Crocodiles Crocodylus niloticus (Laurenti, 1786)

Relationships of the newly discovered dwarf crocodiles from Mauritania were inferred from mitochondrial 12S sequences. Specimens from 13 different Crocodylus niloticus populations (from East Africa, West Africa and Madagascar) were compared. Additional representatives of the genus Crocodylus (one from Africa and one from Australia), the African genus Osteolaemus and the South American alligatorid Paleosuchus palpebrosus (as outgroup) were included in the analysis. Maximum-likelihood and Bayesian analyses yielded relationships that were strikingly different from currently prevailing phylogenetic hypotheses. Both analyses consistently revealed two groups, one consisting of the monophyletic West- and Central African populations and the other of a paraphyletic group containing the East African and Madagascan populations. High genetic divergence between those groups indicates separation on the species level. Furthermore ‘C’ cataphractus is clearly shown not to be a member of the genus Crocodylus. The resulting nomenclatural changes are discussed. To cite this article: A. Schmitz et al., C. R. Palevol 2 (2003).     

Cryptic diversity within the Moroccan endemic day geckos Quedenfeldtia (Squamata: Gekkonidae): a multidisciplinary approach using genetic,morphological and ecological data

Quedenfeldtia (Boettger, 1883) is a genus of diurnal geckos, endemic to the Atlas Mountains in Morocco, with two species being recognized: Quedenfeldtia moerens and Quedenfeldtia trachyblepharus. Quedenfeldtia moerens is found across a wide variety of habitats, from sea level to 3000 m a.s.l., whereas Q. trachyblepharus occupies exclusively high mountain regions reaching up to 4000 m a.s.l. This differentiation, offers an interesting model for study biogeographical patterns and evolutionary scenarios in a North African endemic. Analysis of two mitochondrial (12S rRNA and ND4) and four nuclear (ACM4, MC1R, PDC, and Rag1) DNA markers revealed high genetic variation, consistent with other recent phylogeographical studies, and with the two currently described species. However, within each species, a subdivision into two groups with geographical consistence was found. Multivariate morphological analyses confirmed the existence of two main phenotypes, whereas ecological niche modelling identified various environmental variables associated with the distribution of each species, and helped to predict occurrences outside the confirmed ranges. The results obtained in the present study indicate the possible existence of additional ‘cryptic’ species within this genus, a condition found in many North African reptiles, and particularly common in geckos. In general, North African montane fauna appears to reflect the occurrence of diverse palaeoendemics, as seen in Central Africa Mountain systems, rather than the pattern of recent postglacial recolonization observed in Europe. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Are cryptic species of the Lesser Egyptian Jerboa,Jaculus jaculus (Rodentia,Dipodidae), really cryptic? Re‐evaluation of their taxonomic status with new data from Israel and Sinai

Two clades of the lesser Egyptian jerboa Jaculus jaculus sensu lato were recently described in North Africa and considered as cryptic species. Members of both clades are also found in Israel, where they can be easily identified according to fur and tail colouration and morphology of the male external genitalia, but cannot be separated confidently using skull characters. Examination of type specimens demonstrated that the correct names for the two species are Jaculus jaculus (Linnaeus 1758) and Jaculus hirtipes (Lichtenstein, 1823). Comparisons of geographic and habitat differences of the two species revealed a high niche divergence between them, slightly higher in the sympatric North African populations than in the parapatric populations of Israel and Sinai. A low niche divergence was detected between North African and Middle Eastern populations of J. jaculus, and a low niche convergence between North African and Middle Eastern populations of J. hirtipes. The levels of niche differentiation coincide with those of genetic differences.     

East African pigs have a complex Indian,Far Eastern and Western ancestry

In this study, we have characterized the mitochondrial diversity of 81 swine from Uganda. Median‐joining network analysis of D‐loop sequences from these individuals and others characterized in previous studies allowed us to determine that Ugandan pigs cluster with populations from the West (Europe/North Africa), Far East and India. In addition, partial sequencing of the Y‐chromosome UTY locus in 18 Ugandan domestic pigs revealed the segregation of a single HY1 lineage that has a cosmopolitan distribution. A Western and Far Eastern ancestry for East African pigs had been already reported, but this is the first study demonstrating an additional contribution from the Indian porcine gene pool. This result is consistent with the high frequency of zebuine alleles in cattle from East Africa. The geographic coordinates of East Africa, at the crossroads of many trading routes that, through the ages, linked Europe, Africa and Asia, might explain the rich and complex genetic heritage of livestock native to this area.     

Domestication ofEleusine coracana

Eleusine coracana (L.) Gaertn. subsp.coracana (Finger millet or eleusine) is a cereal that is widely cultivated in Africa and India. Archaeological records of finger millet are few and unsatisfactory. But, distribution, linguistic and historical evidence seem to suggest an African rather than Indian origin of the crop. Data from morphology, supplemented with cytogenetical observations and distribution revealed thatE. coracana subsp.africana is wild finger millet. This subspecies is widely distributed along the highlands of East Africa. Consequently, it is concluded that finger millet originated in the East African Highlands and was subsequently introduced into India.     

Evolutionary history of spiny‐tailed lizards (Agamidae: Uromastyx) from the Saharo‐Arabian region

The subfamily Uromastycinae within the Agamidae is comprised of 18 species: three within the genus Saara and 15 within Uromastyx. Uromastyx is distributed in the desert areas of North Africa and across the Arabian Peninsula towards Iran. The systematics of this genus has been previously revised, although incomplete taxonomic sampling or weakly supported topologies resulted in inconclusive relationships. Biogeographic assessments of Uromastycinae mostly agree on the direction of dispersal from Asia to Africa, although the timeframe of the cladogenesis events has never been fully explored. In this study, we analysed 129 Uromastyx specimens from across the entire distribution range of the genus. We included all but one of the recognized taxa of the genus and sequenced them for three mitochondrial and three nuclear markers. This enabled us to obtain a comprehensive multilocus time‐calibrated phylogeny of the genus, using the concatenated data and species trees. We also applied coalescent‐based species delimitation methods, phylogenetic network analyses and model‐testing approaches to biogeographic inferences. Our results revealed Uromastyx as a monophyletic genus comprised of five groups and 14 independently evolving lineages, corresponding to the 14 currently recognized species sampled. The onset of Uromastyx diversification is estimated to have occurred in south‐west Asia during the Middle Miocene with a later radiation in North Africa. During its Saharo‐Arabian colonization, Uromastyx underwent multiple vicariance and dispersal events, hypothesized to be derived from tectonic movements and habitat fragmentation due to the active continental separation of Arabia from Africa and the expansion and contraction of arid areas in the region.     

Why Africa matters: evolution of Old World Salvia (Lamiaceae) in Africa

Background and Aims

Salvia is the largest genus in Lamiaceae and it has recently been found to be non-monophyletic. Molecular data on Old World Salvia are largely lacking. In this study, we present data concerning Salvia in Africa. The focus is on the colonization of the continent, character evolution and the switch of pollination systems in the genus.

Methods

Maximum likelihood and Bayesian inference were used for phylogenetic reconstruction. Analyses were based on two nuclear markers [internal transcribed spacer (ITS) and external transcribed spacer (ETS)] and one plastid marker (rpl32-trnL). Sequence data were generated for 41 of the 62 African taxa (66 %). Mesquite was used to reconstruct ancestral character states for distribution, life form, calyx shape, stamen type and pollination syndrome.

Key Results

Salvia in Africa is non-monophyletic. Each of the five major regions in Africa, except Madagascar, was colonized at least twice, and floristic links between North African, south-west Asian and European species are strongly supported. The large radiation in Sub-Saharan Africa (23 species) can be traced back to dispersal from North Africa via East Africa to the Cape Region. Adaptation to bird pollination in southern Africa and Madagascar reflects parallel evolution.

Conclusions

The phenotypic diversity in African Salvia is associated with repeated introductions to the continent. Many important evolutionary processes, such as colonization, adaptation, parallelism and character transformation, are reflected in this comparatively small group. The data presented in this study can help to understand the evolution of Salvia sensu lato and other large genera.