Pleistocene colonization of afro-alpine 'sky islands' by the arctic-alpine Arabis alpina

The afro-alpine region comprises the high mountains of Ethiopia and tropical East Africa, which represent biological 'sky islands' with high level of endemism. However, some primarily arctic-alpine plants also occur in the afro-alpine mountains. It has been suggested that these plants are Tertiary relicts, but a recent worldwide study of Arabis alpina suggests that this species colonized the region twice during the Pleistocene. Here we investigate the detailed colonization history of A. alpina in the afro-alpine region based on chloroplast DNA sequences from 11 mountain systems. The results confirm the twice-into-Africa scenario. The Asian lineage is confined to the mountains closest to the Arabian Peninsula, on opposite sides of the Rift Valley (Simen Mts and Gara Muleta in Ethiopia), suggesting long-distance dispersal of this lineage. The African lineage is divided into two phylogeographic groups with distinct geographic distribution. The observed pattern is consistent with isolation of the African lineage in at least two interglacial refugia, located on separated highlands, followed by range expansion in cooler period(s), when the afro-alpine habitat extended further down the mountains. Several long-distance dispersal events, also across the Rift Valley, are suggested by single haplotypes observed outside the area occupied by the phylogeographic groups they belonged to.     

Browser selectivity alters post-fire competition between <Emphasis Type="Italic">Erica arborea</Emphasis> and <Emphasis Type="Italic">E. trimera</Emphasis> in the sub-alpine heathlands of Ethiopia

Mammalian herbivores have the potential to alter the competitive relations of woody species, if consumption is unevenly distributed between species. At elevations above 3,500 m in the southern Ethiopian highlands, vegetation is dominated by Erica arborea and E. trimera. Both species can potentially grow into short trees, but are burnt on a rotation of 6–10 years, and regenerate by re-sprouting from belowground lignotubers. The regenerating scrub is heavily browsed by cattle. We set up browsing exclosures at three burnt sites to quantify the impact of browsing over a 3-year period. When protected from browsing, E. trimera had similar or better height growth than E. arborea, but in browsed vegetation, E. arborea instead grew taller. Browsing was more intense on E. trimera in the first years after fire, indicating a difference in palatability between the species. We checked if browse quality differed, by analysing shoot contents of acid detergent fibre (ADF), protein, phenolics and tannins. Contrary to expectations, the preferred E. trimera contained more ADF, less protein and had a higher tannin activity than E. arborea. Although the vegetative growth of E. arborea is favoured relative to E. trimera under high browsing pressure, rapid change in abundance would not be expected, since short-interval fire will repeatedly eradicate any gains in vegetative growth. However, within the typical fire return interval of less than 10 years, E. trimera barely reach a reproductive state, whereas E. arborea flower profusely. Under the current regime of fire and browsing, this may in the long run be more important than differences in height growth, leading to a gradual increase in the proportion of E. arborea.     

Phylogeography and conservation genetics of a giant lobelia (Lobelia giberroa) in Ethiopian and Tropical East African mountains

Lobelia giberroa is a giant rosette plant growing in the afro-montane belt of the afro-alpine environment, a unique and little-studied ecosystem occupying the high mountains of eastern Africa. We analysed amplified fragment length polymorphisms (AFLPs) from 11 mountain systems in Ethiopia and Tropical East Africa to infer the phylogeographical history of the species. A total of 191 individuals were investigated from 25 populations. Principal coordinate analysis and population structure analyses revealed three major phylogeographical groups: the Ethiopian mountains and one group on each side of the Rift Valley in Tropical East Africa, respectively: Elgon-Cherangani and Kenya-Aberdare-Kilimanjaro-Meru. Analysis of Molecular Variance showed 55.7% variance among the three groups, suggesting an old divergence. Together with a clear geographical substructure within the main groups, this pattern indicates gradual expansion and supports the montane forest bridge hypothesis, stating that the area occupied by forest was larger and more continuous in previous interglacials and earlier in the present interglacial. Genetic diversity was lower in Ethiopia than in the other two main groups, possibly due to an ancient founder effect when Ethiopia was colonized from the south.     

Sweet vernal grasses (Anthoxanthum) colonized African mountains along two fronts in the Late Pliocene,followed by secondary contact,polyploidization and local extinction in the Pleistocene

High tropical mountains harbour remarkable and fragmented biodiversity thought to a large degree to have been shaped by multiple dispersals of cold‐adapted lineages from remote areas. Few dated phylogenetic/phylogeographic analyses are however available. Here, we address the hypotheses that the sub‐Saharan African sweet vernal grasses have a dual colonization history and that lineages of independent origins have established secondary contact. We carried out rangewide sampling across the eastern African high mountains, inferred dated phylogenies from nuclear ribosomal and plastid DNA using Bayesian methods, and performed flow cytometry and AFLP (amplified fragment length polymorphism) analyses. We inferred a single Late Pliocene western Eurasian origin of the eastern African taxa, whose high‐ploid populations in one mountain group formed a distinct phylogeographic group and carried plastids that diverged from those of the currently allopatric southern African lineage in the Mid‐ to Late Pleistocene. We show that Anthoxanthum has an intriguing history in sub‐Saharan Africa, including Late Pliocene colonization from southeast and north, followed by secondary contact, hybridization, allopolyploidization and local extinction during one of the last glacial cycles. Our results add to a growing body of evidence showing that isolated tropical high mountain habitats have a dynamic recent history involving niche conservatism and recruitment from remote sources, repeated dispersals, diversification, hybridization and local extinction.     

Quaternary range dynamics of ecologically divergent species (Edraianthus serpyllifolius and E. tenuifolius,Campanulaceae) within the Balkan refugium

Aim We investigated Quaternary range dynamics of two closely related but ecologically divergent species (cold‐tolerant Edraianthus serpyllifolius and thermophilic Edraianthus tenuifolius) with overlapping distribution ranges endemic to the western Balkan Peninsula, an important yet understudied Pleistocene refugium. Our aims were: to test predictions of the ‘refugia‐within‐refugia’ model of strong genetic subdivisions due to population isolation in separate refugia; to explore whether two ecologically divergent species reacted differently to Pleistocene climatic fluctuations; and to test predictions of the displacement refugia model of stronger differentiation among populations in the thermophilic E. tenuifolius compared with the cold‐tolerant E. serpyllifolius. Location The western Balkan Peninsula. Methods We gathered amplified fragment‐length polymorphism (AFLP) data and plastid DNA sequences from two to five individuals from 10 populations of E. serpyllifolius and 22 populations of E. tenuifolius, spanning their entire respective distribution areas. AFLP data were analysed using a Bayesian clustering approach and a distance‐based network approach. Plastid sequences were used to depict relationships among haplotypes in a statistical parsimony network, and to obtain age estimates in a Bayesian framework. Results In E. serpyllifolius, both AFLP and plastid sequence data showed clear geographic structure. Western populations showed high AFLP diversity and a high number of rare fragments. In E. tenuifolius, both markers congruently identified a major phylogeographic split along the lower Neretva valley in central Dalmatia. The most distinct and earliest diverging chloroplast DNA (cpDNA) haplotypes were found further south in the south‐easternmost populations. North‐western populations, identified as a separate cluster by Bayesian clustering, were characterized by low genetic diversity and a low number of rare AFLP markers. Main conclusions Clear evidence for multiple Pleistocene refugia is found not only in the high‐elevation E. serpyllifolius, but also in the lowland E. tenuifolius, despite the lack of obvious dispersal barriers, in line with the refugia‐within‐refugia model. Genealogical relationships and genetic diversity patterns support the hypothesis that cold‐adapted E. serpyllifolius responded to climatic oscillations mostly by elevational range shifts, whereas thermophilic E. tenuifolius did so mainly by latitudinal range shifts, with different phases (and probably extents) of range expansion. In contrast to the displacement refugia hypothesis, the two elevationally differentiated species do not differ in their genetic diversity.     

Genetic population structure of the swimming crab, Portunus trituberculatus in the East China Sea based on mtDNA 16S rRNA sequences

Portunus trituberculatus is a commercially important fishery species. In this study, intraspecific variation was examined by using mitochondrial DNA 16S rRNA gene in 213 individual crabs sampled from six localities along the coast of the East China Sea. Twenty-two polymorphic sites defined 25 distinct haplotypes, revealing a moderately high haplotype diversity and relatively low sequence divergence among the six localities. An excess of within population unique haplotypes at most sample locations were detected, which might influence genetic structure of the swimming crab populations. Neither neighbor-joining tree nor minimum spanning network (MSN) based on the haplotype data indicated distinct patterns of phylogeographic structure among the 25 haplotypes. Analyses of molecular variance (AMOVA) and F_ST statistics supported the hypothesis that population samples from the East China Sea were genetically nonhomogenous, indicating that gene flow might be restricted across those regions, despite the high potential of dispersal. In addition, tests of neutral evolution and analysis of mismatch distribution suggested that P. trituberculatus might have undergone a population expansion, possibly within the last 127,000 and 429,000 years. Our study unraveled the extant population genetic structure of the P. trituberculatus, and addressed the related fishery management issues including artificial breeding, fishery stock identification and conservation.     

A long‐standing Pleistocene refugium in southern Africa and a mosaic of refugia in East Africa: insights from mtDNA and the common eland antelope

Aim Previous genetic studies of African savanna ungulates have indicated Pleistocene refugial areas in East and southern Africa, and recent palynological, palaeovegetation and fossil studies have suggested the presence of a long‐standing refugium in the south and a mosaic of refugia in the east. Phylogeographic analysis of the common eland antelope, Taurotragus oryx (Bovidae), was used to assess these hypotheses and the existence of genetic signatures of Pleistocene climate change. Location The sub‐Saharan savanna biome of East and southern Africa. Methods Mitochondrial DNA control‐region fragments (414 bp) from 122 individuals of common eland were analysed to elucidate the phylogeography, genetic diversity, spatial population structuring, historical migration and demographic history of the species. The phylogeographic split among major genetic lineages was dated using Bayesian coalescent‐based methods and a calibrated fossil root of 1.6 Ma for the split between the common eland and the giant eland, Taurotragus derbianus. Results Two major phylogeographic lineages comprising East and southern African localities, respectively, were separated by a net nucleotide distance of 4.7%. A third intermediate lineage comprised only three haplotypes, from Zimbabwe in southern Africa. The estimated mutation rate of 0.097 Myr^?1 revealed a more recent common ancestor for the eastern lineage (0.21 Ma; 0.07–0.37) than for the southern lineage (0.35 Ma; 0.10–0.62). Compared with the latter, the eastern lineage showed pronounced geographic structuring, lower overall nucleotide diversity, higher population differentiation, and isolation‐by‐distance among populations. Main conclusions The data support the hypothesis of Pleistocene refugia occurring in East and southern Africa. In agreement with palynological, palaeovegetation and fossil studies, our data strongly support the presence of a longer‐standing population in the south and a mosaic of Pleistocene refugia in the east, verifying the efficacy of genetic tools in addressing such questions. The more recent origin of the common eland inhabiting East Africa could result from colonization following extinction from the region. Only two other dated African ungulate phylogenies have been published, applying different methods, and the similarity of dates obtained from the three distinct approaches indicates a significant event c. 200 ka, which left a strong genetic signature across a range of ungulate taxa.     

Three times out of Asia Minor: the phylogeography of Arabis alpina L. (Brassicaceae)

Arabis alpina is a characteristic plant in arctic-alpine habitats and serves as a classical example to demonstrate biology, ecology and biogeography of arctic-alpine disjuncts. It has a wider distribution than most other arctic-alpine plants, covering all European mountain systems, the Canary Islands, North Africa, the high mountains of East Africa and Ethiopia, the Arabian Peninsula and mountain ranges of Central Asia in Iran and Iraq. Additionally it is found in the northern amphi-Atlantic area including northeastern North America, Greenland, Iceland, Svalbard and northwestern Europe. We used markers from the nuclear (internal transcribed spacer of ribosomal DNA) and chloroplast genome (trnL-F region) to reconstruct its phylogeographic history. Both markers revealed clear phylogeographic structure. We suggest that A. alpina originated in Asia Minor less than 2 million years ago based on synonymous mutation rates of different genes (plastidic matK, nuclear adh and chs). From the Asian ancestral stock one group migrated via the Arabian Peninsula to the East African high mountains. A second group gave rise to all European and northern populations, and also served as source for the northwest African populations. A third group, which is still centred in Asia, migrated independently southwards and came into secondary contact with the East African lineage in Ethiopia, resulting in high genetic diversity in this area. In the Mediterranean regions, the genetic diversity was relatively high with numerous unique haplotypes, but almost without geographic structure. In contrast, the populations in the northern amphi-Atlantic area were extremely depauperate, suggesting very recent (postglacial) expansion into this vast area from the south.     

Crossing barriers in an extremely fragmented system: two case studies in the afro-alpine sky island flora

The flora on the afro-alpine sky islands is renowned for extreme fragmentation, representing a unique natural experiment in biogeography. Here we address the roles of isolation and gene flow, in particular across the narrow Rift Valley (the RV barrier) that cuts through the Ethiopian Highlands (EH), and across the vast low-lying landscape that separates EH from the East African mountains (the EH–EA barrier). We inferred the history of two species with different dispersal mechanisms, but with similar geographic ranges and habitats based on Amplified fragment length polymorphisms (AFLPs). Contrary to our predictions, we found that the populations from opposite sides of the RV barrier were less similar than those from opposite sides of the EH–EA barrier, and that only the supposedly short distance-dispersed species (Trifolium cryptopodium) showed a strong signal of secondary gene flow across the RV barrier. In the wind-dispersed Carduus schimperi, we rather found an evidence for the gene flow between differentiated populations inhabiting different EA mountains. Both species harbored little genetic diversity but considerable genetic rarity in several individual mountains, suggesting long-term isolation and bottlenecks during climatically unfavorable periods. Our genetic data corroborate a division of C. schimperi into three subspecies, but with new delimitation of their ranges, and of T. cryptopodium into two intraspecific taxa. Our findings support the idea that stochasticity may play a major role in shaping extremely fragmented ecosystems such as the afro-alpine. After initial colonization of different mountains, periods of isolation may alternate with unpredictable episodes of intermountain gene flow.     

Comparative phylogeography of African savannah ungulates

The savannah biome of sub-Saharan Africa harbours the highest diversity of ungulates (hoofed mammals) on Earth. In this review, we compile population genetic data from 19 codistributed ungulate taxa of the savannah biome and find striking concordance in the phylogeographic structuring of species. Data from across taxa reveal distinct regional lineages, which reflect the survival and divergence of populations in isolated savannah refugia during the climatic oscillations of the Pleistocene. Data from taxa across trophic levels suggest distinct savannah refugia were present in West, East, Southern and South-West Africa. Furthermore, differing Pleistocene evolutionary biogeographic scenarios are proposed for East and Southern Africa, supported by palaeoclimatic data and the fossil record. Environmental instability in East Africa facilitated several spatial and temporal refugia and is reflected in the high inter- and intraspecific diversity of the region. In contrast, phylogeographic data suggest a stable, long-standing savannah refuge in the south.     

Southern isolation and northern long‐distance dispersal shaped the phylogeography of the widespread,but highly disjunct,European high mountain plant Artemisia eriantha (Asteraceae)

We investigated the range dynamics of Artemisia eriantha, a widespread, but rare, mountain plant with a highly disjunct distribution in the European Alpine System. We focused on testing the roles of vicariance and long‐distance dispersal in shaping the current distribution of the species. To this end, we collected AFLP and plastid DNA sequence data for 17 populations covering the entire distributional range of the species. Strong phylogeographical structure was found in both datasets. AFLP data suggested that almost all populations were genetically strongly differentiated, with 58% of the overall genetic variation partitioned among populations. Bayesian clustering identified five groups of populations: Balkans, Pyrenees, Central Apennines, one southwestern Alpine population and a Widespread cluster (eastern Pyrenees, Alps, Carpathians). Major groups were supported by neighbor‐joining and NeighbourNet analyses. Fourteen plastid haplotypes were found constituting five strongly distinct lineages: Alps plus Pyrenees, Apennines, Balkans, southern Carpathians, and a Widespread group (eastern Pyrenees, northern Carpathians, Mt. Olympus). Plastid DNA data suggested that A. eriantha colonized the European Alpine System in a westward direction. Although, in southern Europe, vicariant differentiation among the Iberian, Italian and Balkan Peninsulas predominated, thus highlighting their importance as glacial refugia for alpine species, in temperate mountain ranges, long‐distance dispersal prevailed. This study emphasizes that currently highly disjunct distributions can be shaped by both vicariance and long‐distance dispersal, although their relative importance may be geographically structured along, for instance, latitude, as in A. eriantha. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 214–226.     

Extrinsic and intrinsic influences on the phylogeography of the Austrocochlea constricta species group

Although extrinsic factors, such as oceanic currents and isolation induced by sea level maxima during Plio-Pleistocene glacial cycles, are often suggested as principal determinants of marine phylogeography, they are not always complete explanations. The counterexamples to predominant phylogeographic patterns in southeastern Australia suggest for example, that intrinsic factors such a habitat preference or reproductive mode can have significant influence.We collected DNA sequences from mitochondrial cytochrome c oxidase I (COI) and two nuclear gene introns from the Austrocochlea constricta species group to determine whether its porcata and constricta phenotypes are genetically distinguishable, to examine the phylogeographic effects of the Bassian Isthmus landbridge formed between Tasmania and Victoria during glacial maxima, and to investigate the importance of intrinsic factors in structuring its genetic variability.No fixed genetic differences between the porcata and constricta phenotypes were identified in any gene so the number of species comprised by the group cannot yet be determined. The two major clades recovered in COI analyses were respectively found principally east or west of the Bassian Isthmus. A. constricta is the first lecithotrophic or estuarine invertebrate known to show this pattern. There were no fixed differences, for any of the three genes, between specimens from estuarine and marine populations within either the eastern or western COI clades. Other intrinsic factors such as breeding period (possibly) and larval type (probably) may play roles in modulating phylogeographic patterns.     

Phylogeographic patterns, genetic affinities and morphological differentiation between Epipactis helleborine and related lineages in a Mediterranean glacial refugium

Background and Aims

In the Mediterranean basin, the Italian peninsula has been suggested to be one of the most important glacial refugia for temperate tree species. The orchid genus Epipactis is widely represented in the Italian peninsula by widespread species and several endemic, localized taxa, including selfing and outcrossing taxa. Here the phylogenetic and phylogeographic relationships in a group of closely related taxa in Epipactis are investigated with the aim of understanding the role of this refugial area for cladogenesis and speciation in herbaceous species, such as terrestrial orchids.

Methods

Ribosomal DNA (rDNA) was employed to assess phylogenetic relationships, and plastid sequence variation in the rbcL–accD spacer was used to reveal phylogeographic patterns among plastid haplotypes using a parsimony network.

Key Results

Low genetic variation and shared ribotypes were detected in rDNA, whereas high levels of sequence variation and a strong phylogeographic structure were found in the examined plastid region. The parsimony plastid haplotype network identified two main haplotype groups, one including E. atrorubens/microphylla/muelleri/leptochila and the other including all accessions of E. helleborine and several localized and endemic taxa, with a combination of widespread and rare haplotypes detected across the Italian peninsula. A greater genetic divergence separated the Italian and other European accessions of E. helleborine.

Conclusions

Phylogenetic and phylogeographic patterns support a working hypothesis in which the Italian peninsula has only recently been colonized by Epipactis, probably during the most recent phase of the Quaternary age and, nevertheless, it acted as a remarkable centre of diversification for this orchid lineage. Changes in pollination strategy and recurrent shifts in mating system (from allogamy to autogamy) could have represented the mechanism promoting this rapid diversification and the observed high taxonomic complexity detected in the E. helleborine species complex.     

Comparative phylogeography of diadromous and freshwater daces of the genus <Emphasis Type="Italic">Tribolodon</Emphasis> (Cyprinidae)

Far Eastern daces, genus Tribolodon (Cyprinidae), are thought to have diversified and developed unique diadromous life histories under changing conditions in the Sea of Japan and the surrounding environment. To examine the relationships between life history traits, distribution, and genetic population structures, we conducted a comparative phylogeographic analysis using partial mtDNA sequence data from samples collected over almost the full ranges of all four Tribolodon species. Phylogenetic analyses revealed several intraspecific haplotype groups that differentiated in the early Pleistocene to the Pliocene with or without geographic overlaps. A time-calibrated phylogeny suggested that the relatively smaller geographic ranges of the strictly freshwater species, T. sachalinensis and T. nakamurai, were explained not by the recent origins of these species, but by their limited dispersal abilities and smaller historical population sizes. The wider-ranging diadromous species, T. brandtii and T. hakonensis, exhibited similar major phylogeographic structures in their distributions, but the chronological order and timing of formation of this structure largely differed between the two species. In addition to those differences, the overlapping patterns of the differentiated intraspecific lineages in these species suggest dynamic, but somewhat restricted dispersal during the Plio-Pleistocene. Tribolodon hakonensis, one of the most widespread species of East Asian freshwater fishes, included both common and unique phylogeographic patterns compared to other fish species; the unique patterns (i.e., its wide range across freshwater biogeographic boundaries like the sea and mountains) would reflect its ecological features as a remarkable generalist inhabiting lakes, upper and lower reaches of rivers, and even coastal areas.     

The colonization history of Juniperus brevifolia (Cupressaceae) in the Azores Islands

Background

A central aim of island biogeography is to understand the colonization history of insular species using current distributions, fossil records and genetic diversity. Here, we analyze five plastid DNA regions of the endangered Juniperus brevifolia, which is endemic to the Azores archipelago.

Methodology/Principal Findings

The phylogeny of the section Juniperus and the phylogeographic analyses of J. brevifolia based on the coalescence theory of allele (plastid) diversity suggest that: (1) a single introduction event likely occurred from Europe; (2) genetic diversification and inter-island dispersal postdated the emergence of the oldest island (Santa Maria, 8.12 Ma); (3) the genetic differentiation found in populations on the islands with higher age and smaller distance to the continent is significantly higher than that on the younger, more remote ones; (4) the high number of haplotypes observed (16), and the widespread distribution of the most frequent and ancestral ones across the archipelago, are indicating early diversification, demographic expansion, and recurrent dispersal. In contrast, restriction of six of the seven derived haplotypes to single islands is construed as reflecting significant isolation time prior to colonization.

Conclusions/Significance

Our phylogeographic reconstruction points to the sequence of island emergence as the key factor to explain the distribution of plastid DNA variation. The reproductive traits of this juniper species (anemophily, ornithochory, multi-seeded cones), together with its broad ecological range, appear to be largely responsible for recurrent inter-island colonization of ancestral haplotypes. In contrast, certain delay in colonization of new haplotypes may reflect intraspecific habitat competition on islands where this juniper was already present.     

Plio-Pleistocene history and phylogeography of Acacia senegal in dry woodlands and savannahs of sub-Saharan tropical Africa: evidence of early colonisation and recent range expansion

Drylands are extensive across sub-Saharan Africa, socio-economically and ecologically important yet highly sensitive to environmental changes. Evolutionary history, as revealed by contemporary intraspecific genetic variation, can provide valuable insight into how species have responded to past environmental and population changes and guide strategies to promote resilience to future changes. The gum arabic tree (Acacia senegal) is an arid-adapted, morphologically diverse species native to the sub-Saharan drylands. We used variation in nuclear sequences (internal transcribed spacer (ITS)) and two types of chloroplast DNA (cpDNA) markers (PCR-RFLP, cpSSR) to study the phylogeography of the species with 293 individuals from 66 populations sampled across its natural range. cpDNA data showed high regional and rangewide haplotypic diversity (h_T(cpSSR)=0.903–0.948) and population differentiation (G_ST(RFLP)=0.700–0.782) with a phylogeographic pattern that indicated extensive historical gene flow via seed dispersal. Haplotypes were not restricted to any of the four varieties, but showed significant geographic structure (G_ST(cpSSR)=0.392; R_ST=0.673; R_ST>R_ST (permuted)), with the major division separating East and Southern Africa populations from those in West and Central Africa. Phylogenetic analysis of ITS data indicated a more recent origin for the clade including West and Central African haplotypes, suggesting range expansion in this region, possibly during the Holocene humid period. In conjunction with paleobotanical evidence, our data suggest dispersal to West Africa, and across to the Arabian Peninsula and Indian subcontinent, from source populations located in the East African region during climate oscillations of the Plio-Pleistocene.     

Dental and phylogeographic patterns of variation in gorillas

Gorilla patterns of variation have great relevance for studies of human evolution. In this study, molar morphometrics were used to evaluate patterns of geographic variation in gorillas. Dental specimens of 323 adult individuals, drawn from the current distribution of gorillas in equatorial Africa were divided into 14 populations. Discriminant analyses and Mahalanobis distances were used to study population structure.Results reveal that: 1) the West and East African gorillas form distinct clusters, 2) the Cross River gorillas are well separated from the rest of the western populations, 3) gorillas from the Virunga mountains and the Bwindi Forest can be differentiated from the lowland gorillas of Utu and Mwenga-Fizi, 4) the Tshiaberimu gorillas are distinct from other eastern gorillas, and the Kahuzi-Biega gorillas are affiliated with them. These findings provide support for a species distinction between Gorilla gorilla and Gorilla beringei, with subspecies G. g. diehli, G. g. gorilla, G. b. graueri, G. b. beringei, and possibly, G. b. rex-pygmaeorum. Clear correspondence between dental and other patterns of taxonomic diversity demonstrates that dental data reveal underlying genetic patterns of differentiation.Dental distances increased predictably with altitude but not with geographic distances, indicating that altitudinal segregation explains gorilla patterns of population divergence better than isolation-by-distance. The phylogeographic pattern of gorilla dental metric variation supports the idea that Plio-Pleistocene climatic fluctuations and local mountain building activity in Africa affected gorilla phylogeography. I propose that West Africa comprised the historic center of gorilla distribution and experienced drift-gene flow equilibrium, whereas Nigeria and East Africa were at the periphery, where climatic instability and altitudinal variation promoted drift and genetic differentiation. This understanding of gorilla population structure has implications for gorilla conservation, and for understanding the distribution of sympatric chimpanzees and Plio-Pleistocene hominins.     

Spatial and temporal patterns of genetic variation in the widespread antitropical deep‐sea coral Paragorgia arborea

Numerous deep‐sea species have apparent widespread and discontinuous distributions. Many of these are important foundation species, structuring hard‐bottom benthic ecosystems. Theoretically, differences in the genetic composition of their populations vary geographically and with depth. Previous studies have examined the genetic diversity of some of these taxa in a regional context, suggesting that genetic differentiation does not occur at scales of discrete features such as seamounts or canyons, but at larger scales (e.g. ocean basins). However, to date, few studies have evaluated such diversity throughout the known distribution of a putative deep‐sea species. We utilized sequences from seven mitochondrial gene regions and nuclear genetic variants of the deep‐sea coral Paragorgia arborea in a phylogeographic context to examine the global patterns of genetic variation and their possible correlation with the spatial variables of geographic position and depth. We also examined the compatibility of this morphospecies with the genealogical‐phylospecies concept by examining specimens collected worldwide. We show that the morphospecies P. arborea can be defined as a genealogical‐phylospecies, in contrast to the hypothesis that P. arborea represents a cryptic species complex. Genetic variation is correlated with geographic location at the basin‐scale level, but not with depth. Additionally, we present a phylogeographic hypothesis in which P. arborea originates from the North Pacific, followed by colonization of the Southern Hemisphere prior to migration to the North Atlantic. This hypothesis is consistent with the latest ocean circulation model for the Miocene.     

The Influence of Interspecific Competition and Host Preference on the Phylogeography of Two African Ixodid Tick Species

A comparative phylogeographic study on two economically important African tick species, Amblyomma hebraeum and Hyalomma rufipes was performed to test the influence of host specificity and host movement on dispersion. Pairwise AMOVA analyses of 277 mtDNA COI sequences supported significant population differentiation among the majority of sampling sites. The geographic mitochondrial structure was not supported by nuclear ITS-2 sequencing, probably attributed to a recent divergence. The three-host generalist, A. hebraeum, showed less mtDNA geographic structure, and a lower level of genetic diversity, while the more host-specific H. rufipes displayed higher levels of population differentiation and two distinct mtDNA assemblages (one predominantly confined to South Africa/Namibia and the other to Mozambique and East Africa). A zone of overlap is present in southern Mozambique. A mechanistic climate model suggests that climate alone cannot be responsible for the disruption in female gene flow. Our findings furthermore suggest that female gene dispersal of ticks is more dependent on the presence of juvenile hosts in the environment than on the ability of adult hosts to disperse across the landscape. Documented interspecific competition between the juvenile stages of H. rufipes and H. truncatum is implicated as a contributing factor towards disrupting gene flow between the two southern African H. rufipes genetic assemblages.     

The colonization history of the Mediterranean dwarf palm (<Emphasis Type="Italic">Chamaerops humilis</Emphasis> L., Palmae)

Chamaerops humilis L. (Mediterranean dwarf palm) is an important floristic element of the western Mediterranean region because it is the only palm species naturally distributed in both Europe and Africa. The combination of a time-calibrated phylogeny, a haplotype network and genetic diversity analyses based on plastid sequences, together with previous nuclear DNA fingerprint results, helped reconstruct the colonization history of the dwarf palm. Based on a sample of 218 individuals taken from 29 geographical areas that cover the current distribution of Chamaerops, we detected four plastid DNA (petA-psbJ) haplotypes distributed in two haplotype groups (lineage 1: haplotypes A/B; lineage 2: haplotypes C/D). Haplotypes A, B, and C showed a widespread geographical distribution in both Africa and Europe, whereas haplotype D was restricted to two African localities. Paleobotanical data, species distribution modeling and divergence time estimates suggest that Chamaerops diverged from Trachycarpus in the Miocene (27.05–6.05 Ma), followed by a split of the two C. humilis lineages that remained isolated during the Miocene-Pliocene. Divergence estimates also support a derived split into two haplotypes (A/B) in the Pleistocene, when the Mediterranean Sea barrier was in existence. This, together with geographical distribution of haplotypes A and B, strongly suggests that the disjunct distribution of C. humilis haplotypes in Europe and Africa is the result of long-distance dispersal (LDD) events rather than vicariance. In agreement with recurrent gene flow (events of LDD colonization), AMOVA revealed that most of the genetic variance was found among populations (61.52%). Irrespective of predominant plant translocations by humans or seed dispersal by natural means, our results support that current populations are the result of relatively recent contacts between Africa and Europe in the Quaternary.