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.     

Phylogeography of Mus (Nannomys) minutoides (Rodentia,Muridae) in West Central African savannahs: singular vicariance in neighbouring populations

We studied the phylogeography of the strict savannah pygmy mice Mus (Nannomys) minutoides in West Central Africa. A total of 846 base pairs of the cytochrome b sequence were obtained for 66 individuals collected in Gabon, Cameroon, Republic of Congo and Central African Republic. These sequences were compared to those of M. minutoides from other African countries and to eight other species of the genus Mus. We performed maximum likelihood, Bayesian and nested clade analyses, as well as neutrality tests and time estimates. We show that M. minutoides is a well‐differentiated monophyletic species that separated from other pygmy mice 1.17 Myr ago. A distinct West Central African M. minutoides clade diverged early from the other African populations of the species, with a more recent common ancestor dating 0.14 Myr. West Central African populations are globally homogeneous, despite the present fragmentation of savannahs by the rain forest. However, our analyses show an unexpected vicariance between geographically close savannahs, embedded in the rain forest in Central Gabon. One of these populations is genetically more similar to very distant peripheral populations than to three closely neighbouring populations situated on both sides of the Ogooué River. A non‐river geographical barrier probably persisted in this area, durably isolating these local populations. This hypothesis about the history of the savannah landscape should be testable through the biogeographical analysis of other strict savannah small mammal species.     

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.     

Dispersal and speciation of East African mountain bushcrickets of the genus Phlesirtes (Orthoptera: Tettigoniidae: Conocephalinae,Conocephalini) related to climate fluctuations

A phylogeny of the genus Phlesirtes Bolivar is presented, based on new sequence data of three genes (16S rDNA, COI, H3). Species of the genus Phlesirtes (subtribe Karniellina of the Tribe Conocephalini) occupy habitats of montane to afroalpine grasslands in East Africa. Phlesirtes is the most species‐rich genus of the subtribe Karniellina, a group of small flightless Ensifera restricted to eastern Africa. Taken together, the biogeographical patterns seen in Phlesirtes and its molecular phylogeny suggest a migration scenario: the mountain ranges acting as stepping stones, enabling a spread of Phlesirtes ancestors during periods of favourable climatic conditions in the past. The Pleistocene inland volcanoes, such as Mt Kenya or Mt Kilimanjaro, allow us to date speciation processes within the genus Phlesirtes. It is suggested that cooler humid periods of the past 3 Ma boosted speciation of Phlesirtes in East Africa.     

Colonization and diversification in the African 'sky islands' by Eurasian Lychnis L. (Caryophyllaceae)

Aim Many plants occurring on the isolated mountain peaks of Africa have their closest relatives in very remote areas, even in temperate Europe and Asia. Their biogeographical history is poorly understood. The Afro‐montane element of the primarily Eurasian genus Lychnis is a typical example of such a disjunction. Here, we aim to reconstruct the phylogeny of the Afro‐montane endemics of Lychnis and to infer the history of immigration into Africa and of the subsequent dispersal and speciation. Location The Afro‐montane representatives of Lychnis occur in Ethiopia, Kenya, Tanzania, Uganda, Rwanda, Burundi, the Democratic Republic of Congo, Cameroon, and Nigeria. Methods Silica‐dried plant materials collected in Ethiopia in 2004 as well as herbarium material were used for DNA extractions. We used parsimony analysis of nucleotide data from the rps16 intron and psbE‐petL region in plastids, and the internal transcribed spacer (ITS) and a region spanning exon 18–24 in the second largest subunit of RNA polymerase II (RPB2) from the nucleus. DAPI‐flow cytometry was used to determine relative genome size and to infer polyploidization events. Results Monophyly of the Afro‐montane Lychnis with the Eurasian diploid L. flos‐cuculi resolved as sister was strongly supported by the phylogenetic analyses. The Afro‐montane group was further divided into two strongly supported groups correlated with DNA ploidy levels. The relative genome size was species‐specific except for L. abyssinica, which had two genome‐size variants. Main conclusions Our results demonstrate that the biogeographical history of the Afro‐montane Lychnis has been highly dynamic, including polyploidization and both old and recent long‐distance dispersal events, even between Eastern and West Africa. The ancestor is inferred to have immigrated once from Eurasia via the Arabian Peninsula to the Ethiopian highlands. Divergence in the lineage containing diploid taxa was followed by several dispersals from Ethiopia to the Eastern and Western Rift Mountains, and recently to West Africa. Divergence of the lineage including the tetraploid L. crassifolia may have taken place in Ethiopia, from where it dispersed to the Eastern and Western Rift Mountains; alternatively, it may have taken place in the Eastern Rift Mountains, followed by dispersal from there to the Western Rift Mountains and then to Ethiopia.     

The distribution of forest passerine birds and Quaternary climatic change in tropical Africa

A cluster analysis of the distribution patterns of forest passerine birds in tropical Africa showed that they could be divided into three main distribution types: those of restricted distribution (144 species), those of disjunct distributions across West, Central and rarely to East Africa (65 species) and those of very wide distribution (81 species). Centres of species richness, endemism and disjunction coincide spatially, and are identified as forest refugia, where forest persisted throughout Quaternary climatic vicissitudes. These distribution patterns agree with modern interpretations of Quaternary palaeoclimatic changes, which show that glacial periods were arid and interglacials humid. Glacial periods were therefore too dry for montane forest to have spread into areas at present occupied by lowland forest, as advocated by Moreau and other supporters of ihe "pluvial" theory; montane forest is probably as extensive now as at any time during the Quaternary and there is no evidence of past connections between currently isolated montane forests. Bird distribution can be explained largely by slow dispersal outwards from refugia as climatic conditions allowed forest to spread, combined with a relaxing of the distinction between "montane" and "lowland" species under conditions of reduced interspecific competition.     

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.     

Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland forests in East Africa

Burgess, N.D. & Mlingwa, C.O.F. 2000. Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland forests in East Africa. Ostrich 71 (1 & 2): 184–190.

In this paper we assess the evidence for altitudinal movements of forest birds from the montane forests of the Eastern Arc mountains of East Africa to nearby lowland forest patches. For 34 montane species, including all the Eastern Arc endemics except Banded Green Sunbird Anthreptes rubritorques there is no evidence that they undertake seasonal movements to lower altitudes. An additional 26 montane species, of somewhat wider distribution, have been recorded at low (<500 m) altitudes during the cold/dry season (June to September). Most records of these montane birds at lower altitudes are from sites adjoining montane forest areas, although a few records are from lowland coastal forests at 100–240 km distance from montane areas. Only five of the 26 species (White-chested Alethe Alethe fulleborni, White-starred Forest Robin Pogonocichla stellata, Orange Ground Thrush Zoothera gurneyi, Evergreen Forest Warbler Bradypterus mariae and Barred Long-tailed Cuckoo Cercococcyx montanus) are regularly and commonly reported in the lowlands. They are also found in the lowlands in small numbers during the warm/wet season (October to February), when they may breed. The abundance of at least four, and probably more, of the forest birds with a more widespread distribution in the lowland and montane forests of East Africa declines greatly at high altitudes from the onset of the cold/wet season (February) and only increases again at the start of the warm/wet season (September). It is not known how far these species move as they cannot be easily separated from resident populations in lowland forests, and there are no ringing recoveries in different forests. Altitudinal migration of a proportion of the Eastern Arc avifauna is the most likely explanation for available data, although source-sink metapopulation theories may be helpful to explain the distributions of some species. As the movement of forest birds from the Eastern Arc to the lowland forests does not involve the rare endemics, they are of lower conservation concern, but the presence of montane and lowland forest may be important for the long-term survival of some more widely distributed forest species.     

Climatic fluctuations and orogenesis as motors for speciation in East Africa: case study on Parepistaurus Karsch, 1896 (Orthoptera)

Mechanisms of speciation of flightless grasshoppers in mountainous and coastal East Africa are inferred considering (i) phylogenies estimated with a combination of molecular markers (16S rRNA locus, COI and H3), (ii) ecological data and (iii) the geographic distribution of Parepistaurus species. The study suggests that coastal taxa of Parepistaurus belong to ancestral lineages from which evolved the high diversity of species found in the Eastern Arc Mountains of Tanzania and Kenya, which are geologically ancient mountain formations. Network analyses and a molecular clock approach, calibrated with the geological age of the volcanoes, suggested that speciation was boosted by climatic fluctuations affecting large areas of East Africa. With the aridification beginning 2.8 Ma, forest taxa were isolated due to forest fragmentation and populations were separated by extended grasslands, which are avoided by Parepistaurus species. However, a humid period between 2.7 and 2.5 Ma triggered a spread of coastal taxa along the Eastern Arc Mountains. Forests expanded again and riparian vegetation along rivers draining into the Indian Ocean probably served as corridors for the dispersal of coastal taxa to the hinterland. The inland volcanoes such as Mount Kilimanjaro are therefore good time markers because their geological age is known, limiting the available time for speciation processes of mountainous Parepistaurus in the area to a maximum of about 1–2 Ma. A third humid but cold period between 1.1 and 0.9 Ma probably further boosted the spread of several flightless and montane‐adapted Orthoptera taxa.     

Distribution of Asellia tridens (Chiroptera: Hipposideridae) lineages including representatives from Saudi Arabia

The genus Asellia was recently revised, giving an insight into the genetic and morphological characteristics of a group whose distribution spans from North Africa into Asia. We studied the genus further by considering additional mitochondrial markers and sampling localities. The deepest previously identified split in A. tridens is supported (Middle East/North Africa), along with the association of North and West African lineages. Central Saudi Arabia is found to represent the easternmost extent of a North African haplogroup when considering concatenated fragments of Cytochrome-b, NADH Dehydrogenase 2, and Cytochrome Oxidase 1 genes. New distribution data further emphasises the diversity found in the Middle East and suggests some mixing of haplotypes over long distances.     

Forest eternal? Endemic butterflies of the Bamenda highlands,Cameroon, avoid close‐canopy forest

The Gulf of Guinea Highlands, a centre of endemism and high conservation importance, represent the only large mountain system in West and Central Africa. We studied habitat use of three common endemic butterflies Colias electo manengoubensis, Bicyclus anisops and Mylothris jacksoni knutsoni, using time‐standardized surveys in four distinct habitats: close‐canopy forest, scrub and forest edges, bracken and grasslands. All three species avoided close‐canopy forests and bracken; the Colias preferring grassland, whereas Bicyclus and Mylothris scrub and forest edges. Ordination analyses of surrounding habitats indicated that all three taxa required heterogeneous landscape mosaics. We argue that the life history traits of taxa with limited geographic distribution may reflect past habitat conditions within their ranges, and that these habitat preferences can indicate the continuous existence of mosaic of forest and nonforest habitats in the West African mountains. Such a landscape was probably maintained by climatic fluctuation and large herbivores, further modified by human impact. This conclusion is consistent with the palaeoenvironmental record and with the requirements of Afromontane endemics from other groups. Recent conservation activities focus on patches of continuous forests, but the mosaic landscapes are no less threatened by intensive agriculture, and should be included to protected areas.     

Sky Islands of the Cameroon Volcanic Line: a diversification hot spot for puddle frogs (Phrynobatrachidae: Phrynobatrachus)

The continental highlands of the Cameroon Volcanic Line (CVL) represent biological ‘sky islands’ with high levels of species richness and endemism, providing the ideal opportunity to understand how orogenesis and historical climate change influenced species diversity and distribution in these isolated African highlands. Relationships of puddle frogs (Phrynobatrachus) endemic to the CVL are reconstructed to examine the patterns and timing of puddle frog diversification. Historical distributions were reconstructed using both elevation and geography data. Puddle frogs diversified in the CVL via several dispersal and vicariance events, with most of the locally endemic species distributed across the northern part of the montane forest area in the Bamenda‐Banso Highlands (Bamboutos Mts., Mt. Lefo, Mt. Mbam, Mt. Oku and medium elevation areas connecting these mountains). Two new species, P. jimzimkusi sp. n. and P. njiomock sp. n., are also described based on molecular analyses and morphological examination. We find that these new species are most closely related to one another and P. steindachneri with the ranges of all three species overlapping at Mt. Oku. Phrynobatrachus jimzimkusi sp. n. is distributed in the southern portion of the continental CVL, P. njiomock sp. n. is endemic to Mt. Oku, and P. steindachneri is present in the northeastern part of the montane forest area. Both new species can be distinguished from all other puddle frogs by a combination of morphological characters, including their large size, ventral coloration and secondary sexual characteristics present in males. These results highlight the Bamenda‐Banso Highlands, and specifically emphasize Mt. Oku, as a centre of diversification for puddle frogs, supporting the conservation importance of this region. Our results also provide new insights into the evolutionary processes shaping the CVL ‘sky islands’, demonstrating that lineage diversification in these montane amphibians is significantly older than expected with most species diverging from their closest relative in the Miocene. Whereas climatic changes during the Pliocene and Pleistocene shaped intraspecific diversification, most speciation events were significantly older and cannot be linked to Africa's aridification in response to Pleistocene climate fluctuations.     

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 Andean Hapalopsittaca parrots (Psittacidae,Aves): an example of montane‐tropical lowland vicariance

Quintero, E., Ribas, C. C. & Cracraft, J. (2012). The Andean Hapalopsittaca parrots (Psittacidae, Aves): an example of montane‐tropical lowland vicariance. —Zoologica Scripta, 42, 28–43. We describe a phylogenetic and biogeographical pattern connecting high montane biotas to those of the lowlands, as exemplified by the exclusively montane parrot genus Hapalopsittaca and its lowland sister genus Pyrilia, both nested within Tavares et al.’s “amazons and allies” clade. As Hapalopsittaca is the only genus within the “amazons and allies” clade that is exclusively distributed in the Andes, the optimization leads to the inference that the ancestral distribution of the branch leading to Pyrilia + Hapalopsittaca was lowland. Museum specimens were examined to determine basal diagnosably distinct taxonomic units. Based on this analysis, mitochondrial sequences (cyt b and ND2 genes) from 17 individuals, mostly from toe pads, and representing all basal taxa within Hapalopsittaca, were obtained. A divergence‐dating analysis using both nuclear (RAG‐1) and mitochondrial (cyt b) genes was conducted to explore whether the uplift of the Andes coincides temporarily with the origin of this montane group, and thus might be causally linked to its origin. Molecular dating estimates the split between Hapalopsittaca and Pyrilia to have occurred between 6.6 and 8.0 Myr; thus, the timing of this highland/lowland disjunction is consistent with that of the final uplift of the Central Andes, supporting a hypothesis of vicariance due to Andean uplift. These results suggest that the taxonomic assembly of montane biotas may be, at least in part, explained by events of Earth history rather than by long‐distance dispersal and colonization. Diversification within Hapalopsittaca and the origin of current species are more recent in time and probably are related to Pleistocene climatic oscillations, as has been shown in other montane groups.     

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).     

A new genus of African Karniellina (Orthoptera,Tettigoniidae, Conocephalinae,Conocephalini): integrating morphological,molecular and bioacoustical data

Melanoscirtes gen.n. is established within Karniellina. The members of this subtribe are small conocephaline bush crickets, confined to Africa. Melanoscirtes is erected on Phlesirtes kibonotensis, a species restricted to forest clearings and forest edge in the submontane and montane zones of Mt. Kilimanjaro. A subspecies, M. kibonotensis uguenoensis, is described from the North Pare mountains, a mountain range of the Eastern Arc adjacent to Mt. Kilimanjaro. Further species of Melanoscirtes occur on other mountain ranges of the northern branch of the Eastern Arc mountains of northern Tanzania and southern Kenya. The South Pare mountains harbour M. shengenae; the West Usambaras, M. usambarensis, and the Taita Hills, M. taitensis. All species and subspecies of Melanoscirtes exhibit a similar morphology and occupy analogous habitats on the respective mountains. The song patterns for all species found within this genus are very similar, and this, together with evidence from molecular data, suggests that allopatric speciation is the reason for the biogeographic pattern found in this genus. A key for the subspecies and species of Melanoscirtes is provided.     

Biogeography of Nicotiana section Suaveolentes (Solanaceae) reveals geographical tracks in arid Australia

Aim Nicotiana section Suaveolentes is largely endemic to Australia but includes one species endemic to Africa, one to New Caledonia and Tongatupa, and one to the Marquesas Islands in the Pacific. Other sections of Nicotiana are found in the New World. In Australia, Suaveolentes is widespread across the continent, with many taxa adapted to the Eremean zone. We aim to analyse the biogeography of the Australian clade, both to shed light on the evolution of the group and to determine general area relationships that provide insight into the history of the arid‐zone biota. Location Mesic and arid regions of continental Australia, the Central–South Pacific and Namibia, Africa. Methods A phylogeny of Suaveolentes, based on morphology and molecular data, was used to analyse the relationships of areas in which the taxa occur. The section is monophyletic, and all but three taxa were included (25). The method of paralogy‐free subtree analysis was employed, with the basal taxon Nicotiana africana used as the outgroup. Results Paralogy‐free subtree analysis found five area subtrees that, when combined, resulted in a minimal area cladogram with six resolved nodes. Pacific and mesic eastern Australia (including Lord Howe Island) are at the base of the area cladogram, followed by the differentiation of North West Australia and later South East Australia. Arid regions of Australia are related, revealing three biogeographical tracks: a northern track including the Great Sandy Desert and Tanami, which are related to the Pilbara; a central track relating the Western Desert, Central Ranges, Eastern Desert and North East Interzone; and a southern track relating the South West Interzone, Nullarbor, Adelaide/Eyre and the South East Interzone. Plesiomorphic taxa with chromosome number n = 24–23 occur on the periphery of the continent, and derived taxa with n = 21, 20, 18, 16–15 identify the tracks across arid Australia. Main conclusions The patterns of distribution and differentiation of Suaveolentes in Australia show that the age of the clade is at least Early Miocene, dating to before the onset of aridification in Australia about 15 Ma. The patterns are also interpreted as evidence that it was vicariance that largely shaped speciation in the Eremean zone, with range expansion of some widespread taxa probably occurring in the most recent cycles of severe drying and mobilization of desert dune sands.     

Systematics and diversification of Praomys species (Rodentia: Muridae) endemic to the Cameroon Volcanic Line (West Central Africa)

Missoup, A.D., Nicolas, V., Wendelen, W., Keming, E., Bilong Bilong, C.F., Couloux, A., Atanga, E., Hutterer, R. & Denys, C. (2012). Systematics and diversification of Praomys species (Rodentia: Muridae) endemic to the Cameroon Volcanic Line (West Central Africa). —Zoologica Scripta, 41, 327–345. Our integrative approach combines two mitochondrial genes (16S and cyt b gene), two nuclear genes (exon 10 GHR and exon 1 IRBP) and craniometrical data to test the status and to infer phylogenetic relationships of the three Praomys Cameroon Volcanic Line endemics (P. hartwigi, P. morio and P. obscurus). The taxonomic rank of the principal genus group is assessed and the mode of diversification of species of the P. tullbergi complex in Afrotropical forests is discussed based on estimates of times to the most recent common ancestors and on tree topologies. This study documents for the first time the molecular and morphometrical distinctiveness of P. hartwigi and P. morio within the P. tullbergi species complex. Further studies including specimens of P. hartwigi from all its distribution range are needed to conclude on the status of P. obscurus. The monophyly of the genus Praomys is refuted. Times to the most recent common ancestors of major clades within the P. tullbergi species complex are estimated for the last 2.5 Mya and during the last 1 or 2 Mya for different species or forms. The lowland forest refuge hypothesis might well explain the diversification of P. misonnei, P. rostratus and P. tullbergi in the guineo‐congolese forest block. The isolation of montane forests could have facilitated the divergence between the two montane forest forms P. hartwigi and P. obscurus and between populations of P. morio from the continent and those from the island of Bioko. Praomys populations (species) that inhabit the Cameroon Volcanic Line Praomys probably originated as lowland forms subsequently specialized to highland conditions.     

A 37,500-year pollen record from Lake Malaŵi and implications for the biogeography of afromontane forests

Abstract. Two 10-metre cores were analysed and a 37,500 year pollen stratigraphy was established for Lake Mala?i. A lowstand from 37,500 to 35,900 BP indicates extremely dry conditions. From 35,900 to 34,000 BP montane forests were widespread indicating cold, moist climate. Between 34,000 and 26,400 BP warm and dry conditions prevailed and forest area decreased. During the Last Glacial Maximum montane forest was widespread in the catchment, although woodlands apparently persisted at low altitudes: the expansion of the montane forest indicates that the aridity that affected equatorial Africa during this time period did not affect the Lake Mala?i Catchment. The cold and relatively moist conditions during the Late Pleistocene probably allowed biotic interchange between the highlands of East and West Africa via the highlands along the Zaire—Zambezi watershed and among now-isolated islands of Afromontane vegetation in the Lake Mala?i catchment. The Holocene is characterized by climate and vegetation quite similar to today, with indications of slightly wetter conditions between 6150 and 3000 bp and slightly drier conditions between 8000 and 6150 bp . The low percentages of montane forest pollen throughout the Holocene support the hypothesis that the montane grasslands of Mala?i are not recently anthropogenic.