Forest refugia and riverine barriers promote diversification in the West African pygmy shrew (Crocidura obscurior complex,Soricomorpha)

The Crocidura obscurior or West African pygmy shrew complex is endemic to West African forests from south‐eastern Guinea, eastern Liberia, southern Côte d'Ivoire and south‐western Ghana. We explore the genetic and morphometric diversity of 239 individuals of the C. obscurior complex from 17 localities across its geographical range. Using genetic data from three mitochondrial (16S, cytochrome b and COI) and four nuclear markers (BRCA1, STAT5A, HDAC2 and RIOK3) and skull geometric morphometrics, we show that this complex is composed of two cryptic and sympatric species, C. obscurior and C. eburnea. We then test several hypotheses to infer their evolutionary history. The observed phylogeographical pattern based on cytochrome b and COI sequences fits the forest refuge theory: during arid phases of the Plio‐Pleistocene, around 3.5, 2.1, 1 and 0.5 Mya, a small number of populations survived in isolated forest patches and diverged allopatrically. During wetter climatic periods, forests expanded, leading to secondary contacts between previously isolated populations. Our results also suggest the possible contribution of episodes of isolation in subrefuges. Historical variation of the West African hydrographic network could also have contributed to the observed patterns of genetic differentiation. Rivers such as the Volta and Sassandra may act as past and/or current barriers to gene flow. Although these two species have sympatric distributions, their phylogeographical histories are somewhat dissimilar due to small differences in their dispersal abilities and ecological requirements.     

Molecular and morphometric variation in two sibling species of the genus Praomys (Rodentia: Muridae): implications for biogeography

The rodent genus Praomys is widely distributed in the African tropics. The species are cryptic, rendering the species taxonomy unclear. There are differences of opinion concerning the specific status of Praomys misonnei and Praomys tullbergi, and their geographical distribution. We sequenced the cytochrome b and/or the 16S gene of 221 specimens from 12 countries in order to evaluate the genetic variability within these two species, and to precisely determine their geographical distribution. Morphological and morphometrical analyses on the sequenced specimens were also performed to find criteria useful for the identification of museum specimens. Our results confirm that P. misonnei and P. tullbergi are two valid species that can be separated by molecular data. However, no single discrete morphological character or simple metric measurement can be used to discriminate them. The percentage of misclassified individuals in multivariate discriminant analysis is relatively high (10%). The two species have allopatric distributions: P. tullbergi occurs in West Africa, from eastern Guinea to western Ghana, and P. misonnei is widely distributed from eastern Ghana to western Kenya. Within P. misonnei we identified three or four major geographical clades: a West Central African clade, an East African clade, a Nigerian clade, and a possible West African clade. Within P. misonnei, high geographical morphometrical variability was also identified. The role of both rivers and Pleistocene forest refugia in promoting speciation within the genus Praomys is discussed. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 397–419.     

Guinean biodiversity at the edge: Rodents in forest patches of southern Mali

Southern Mali mainly belongs to the Sudanian savanna bioclimatic zone, but forest patches showing botanical affinities with Guinean humid forest remain as gallery forests or ravine forests. To characterize the rodent diversity of this area and check for the presence of some species of Guinean affinities in this group, rodent assemblages were sampled in four regions of southern Mali, using trapping and observational data in forest and surrounding habitats. Twenty-four species were recorded, comprising a representative sample of the expected overall diversity in this group according to rarefaction curves. Praomys rostratus was the dominant species in the most humid, closed lowland forest. Praomys daltoni was also present in this habitat type, being all the more abundant as habitat degradation was apparent. It became the dominant species in ravine forest on rocky substrate where P. rostratus was completely absent. In Sudanian savanna habitats and in herbaceous and cultivated areas, Mastomys erythroleucus dominated a diverse rodent community. A few species were found that testified for Guinean affinities of the most humid forest patches, especially in the extreme southeast of the country (region of Sikasso). Rodent assemblages of the Bafing and Mts Mandingues areas, in the western part of the study area, showed the highest similarity, in relation with environmental characteristics of this region representing an extension of the Fouta Djallon plateau in Guinea. The results obtained highlight the high biodiversity value of this forest-savanna mosaic, and provide new arguments in favour of the preservation of West African forest patches and their surrounding habitats.     

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, ??, ??–??.     

The roles of rivers and Pleistocene refugia in shaping genetic diversity in Praomys misonnei in tropical Africa

Aim This study aims to elucidate the phylogeography of the murid rodent Praomys misonnei and to document whether or not rain forest refugia and rivers structure patterns of diversity within this species. Location Tropical Africa, from Ghana to Kenya. Methods Patterns of genetic structure and signatures of population history (cytochrome b gene) were assessed in a survey of 229 individuals from 54 localities. Using maximum likelihood, Bayesian, network and genetic structure analyses, we inferred intra‐specific relationships and tested hypotheses for historical patterns of gene flow within P. misonnei. Results Our phylogenetic analyses reveal a strong phylogeographical structure. We identified four major geographical clades within P. misonnei: one clade in Ghana and Benin, a Nigerian clade, a West Central African clade and a Central and East African clade. Several subclades were identified within these four major clades. A signal of population expansion was detected in most clades or subclades. Coalescence within all of the major clades of P. misonnei occurred during the Middle Pleistocene and/or the beginning of Late Pleistocene. Main conclusions Our results suggest a role for both Pleistocene refugia and rivers in structuring genetic diversity in P. misonnei. This forest‐dwelling rodent may have been isolated in a number of forest fragments during arid periods and expanded its range during wetter periods. Potential forest refugia may have been localized in Benin–Ghana, south‐western Cameroon, southern Gabon, northern Gabon and eastern Democratic Republic of Congo–western Uganda. The Niger and/or the Cross Rivers, the Oubangui‐Congo, Sanaga, Ogooue and/or Ivindo Rivers probably stopped the re‐expansion of the species from relict areas.     

Size and shape variability in relation to species differences and climatic gradients in the African rodent Oenomys

Aim The modern African murine rodent Oenomys is a semiarboreal genus characteristic of the rain forest zone. The size and shape geographical differentiation is studied in order to discuss the possible evolutionary patterns and constraints leading to this distribution. Location Two species of Oenomys have been previously recognized, corresponding to the West African forest block on the one hand and to the main forest block on the other hand. Both species have been sampled, and forty localities permit the study of the whole geographical range of Oenomys, from Guinea to Kenya. Methods A Fourier analysis applied to the outlines of the first upper and lower molars allows a quantification of the size and shape variations across the geographical range of Oenomys. Results This morphometrical analysis defines three morphological groups of Oenomys, corresponding to the West, Central, and East African forest zone. This result suggests the existence of an additional cryptic species. The Western group is characterized by broad asymmetrical molars and a small size. The Central African group has more elongated and larger molars while the Eastern African group displays extremely constricted molars and a small size. Main conclusion The shape can be considered as charactheristic of each group and its variations appear to be mostly related to phylogeny while size exhibits variations within each group related to climatic gradients. This discrepancy could be related to different genetic determinants for both characters, the shape being strongly genetically constrained while size can vary according to a wide range of environmental factors.     

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.     

Plio-Pleistocene history of West African Sudanian savanna and the phylogeography of the Praomys daltoni complex (Rodentia): the environment/geography/genetic interplay

Rodents of the Praomys daltoni complex are typical inhabitants of the Sudanian savanna ecosystem in western Africa and represent a suitable model for testing the effects of Quaternary climatic oscillations on extant genetic variation patterns. Phylogeographical analyses of mitochondrial DNA sequences (cytochrome b) across the distribution range of the complex revealed several well‐defined clades that do not support the division of the clade into the two species currently recognized on the basis of morphology, i.e. P. daltoni (Thomas, 1892) and Praomys derooi ( Van der Straeten & Verheyen 1978 ). The observed genetic structure fits the refuge hypothesis, suggesting that only a small number of populations repeatedly survived in distinct forest‐savanna mosaic blocks during the arid phases of the Pleistocene, and then expanded again during moister periods. West African rivers may also have contributed to genetic differentiation, especially by forming barriers after secondary contact of expanding populations. The combination of three types of genetic markers (mtDNA sequences, microsatellite loci, cytogenetic data) provides evidence for the presence of up to three lineages, which most probably represent distinct biological species. Furthermore, incongruence between nuclear and mtDNA markers in some individuals unambiguously points towards a past introgression event. Our results highlight the importance of combining different molecular markers for an accurate interpretation of genetic data.     

Congruent phylogeographical patterns of eight tree species in Atlantic Central Africa provide insights into the past dynamics of forest cover

Cycles of Quaternary climatic change are assumed to be major drivers of African rainforest dynamics and evolution. However, most hypotheses on past vegetation dynamics relied on palaeobotanical records, an approach lacking spatial resolution, and on current patterns of species diversity and endemism, an approach confounding history and environmental determinism. In this context, a comparative phylogeographical study of rainforest species represents a complementary approach because Pleistocene climatic fluctuations may have left interpretable signatures in the patterns of genetic diversity within species. Using 1274 plastid DNA sequences from eight tree species (Afrostyrax kamerunensis, A. lepidophyllus, Erythrophleum suaveolens, Greenwayodendron suaveolens, Milicia excelsa, Santiria trimera, Scorodophloeus zenkeri and Symphonia globulifera) sampled in 50 populations of Atlantic Central Africa (ACA), we averaged divergence across species to produce the first map of the region synthesizing genetic distinctiveness and standardized divergence within and among localities. Significant congruence in divergence was detected mostly among five of the eight species and was stronger in the northern ACA. This pattern is compatible with a scenario of past forest fragmentation and recolonization whereby forests from eastern Cameroon and northeastern Gabon would have been more affected by past climatic change than those of western Cameroon (where one or more refugia would have occurred). By contrast, southern ACA (Gabon) displayed low congruence among species that may reflect less drastic past forest fragmentation or a more complex history of vegetation changes. Finally, we also highlight the potential impact of current environmental barriers on spatial genetic structures.     

Effects of past climate on Passiflora actinia (Passifloraceae) populations and insights into future species management in the Brazilian Atlantic forest

The Brazilian Atlantic Rainforest is one of the most diverse and threatened ecoregions on the planet and displays high levels of endemism. Despite several population analyses and phylogeographical studies, the origins of its species richness and the evolutionary processes that gave rise to this diversification remain poorly understood, especially at the southern edge of the Atlantic Forest. Passiflora actinia is an indigenous species from the southern Atlantic Forest and, as such, was influenced by climatic changes during the Pleistocene. In this study, we investigated the effects of past climate changes on the genetic diversity of P. actinia, using nuclear and plastid markers. We subsequently suggest strategies for the preservation of this species in particular and the whole ecoregion in general. We employed phylogeographical methods and combined these results with past, present and future ensemble niche models. Genetic variability in P. actinia was similar to that of other species with similar geographical distributions, and geographical structuring was not observed based on either type of genetic marker. Diversification in P. actinia was dated to the Pleistocene, suggesting that climate changes could have influenced the distribution of genetic diversity in this species. Our results predicted that suitable P. actinia habitat will persist in the highlands but will be reduced in the lowlands, especially with higher greenhouse gas concentrations. Conservation efforts should focus on populations with unique genetic units and populations from areas with greater climatic instability. Habitat loss due to deforestation in the Atlantic Forest constitutes a major risk to this species, especially to small populations or those with low diversity indices.     

A tale of the nearly tail‐less: the effects of Plio‐Pleistocene climate change on the diversification of the African avian genus Sylvietta

Sylvietta is a broadly distributed group of African species inhabiting a wide range of habitats and presents an interesting opportunity to investigate the historic mechanisms that have impacted the biogeography of African avian species. We collected sequence data from 50 individuals and used model‐based phylogenetic methods, molecular divergence estimates and ancestral area estimates to construct a time‐calibrated phylogeny and estimation of biogeographic history. We estimate a southern African origin for Sylvietta, with an initial divergence splitting the genus into two clades. The first consists of arid‐adapted species, with a southern African origin and subsequent diversification north into Ethiopia–Somalia. The second clade is estimated as having a Congolian forest origin with an eastward pattern of colonization and diversification as a result of Plio‐Pleistocene forest dynamics. Additionally, two members of the genus Sylvietta display interesting patterns of intraspecific diversification. Sylvietta rufescens is an arid‐adapted species inhabiting southern Africa, and we recover two subclades with a divergence dating to the Pleistocene, a unique pattern for avian species which may be explained via isolation in arid habitat fragments in the early Pleistocene. Second, Sylvietta virens, a species endemic to Afro‐tropical forests, is recovered with geographically structured genetic diversification across its broad range, an interesting result given that recent investigations of several avian forest species have found similar and substantial geographically structured genetic diversity relating to Plio‐Pleistocene forest fragmentation. Overall, Plio‐Pleistocene habitat cycling played a significant role in driving diversification in Sylvietta, and this investigation highlights the substantial impact of climate‐driven habitat dynamics on the history of sub‐Saharan species.     

Phylogeography of the Guinea multimammate mouse (Mastomys erythroleucus): a case study for Sahelian species in West Africa

Aim To investigate the phylogeographical structure of the Guinea multimammate mouse, Mastomys erythroleucus (Temminck, 1853), a widespread murid rodent in sub‐Saharan (Sahel and Sudan) savannas, for a better understanding of the impacts of geographical and historical factors on the evolutionary history of this species, in the context of the growing database of phylogeographical studies of African savanna mammal species. Location Sahel and Sudan savannas, Africa. Methods We sequenced the whole cytochrome b gene in 211 individuals from 59 localities distributed from Senegal to Ethiopia. Sequence data were analysed using both phylogenetic (several rooted tree‐construction methods, median‐joining networks) and population genetic methods (spatial analyses of molecular variance, mismatch distributions). Results Haplotypes were distributed into four major monophyletic groups corresponding to distinct geographical regions across a west–east axis. Diversification events were estimated to have occurred between 1.16 and 0.18 Ma. Main conclusions Vicariance events related to the fragmentation of savanna habitats during the Pleistocene era may explain the phylogeographical patterns observed. Genetic structure was consistent with a role of major Sahelian rivers as significant barriers to west–east dispersal. Recent demographic expansions probably occurred during arid phases of the Holocene with the southward expansion of savannas.     

Population genetic structure and phylogeographical pattern of rice grasshopper, <Emphasis Type="Italic">Oxya hyla intricata</Emphasis>, across Southeast Asia

The rice grasshopper, Oxya hyla intricata, is a rice pest in Southeast Asia. In this study, population genetic diversity and structure of this Oxya species was examined using both DNA sequences and AFLP technology. The samples of 12 populations were collected from four Southeast Asian countries, among which 175 individuals were analysed using mitochondrial DNA cytochrome c oxidase subunit I (COI) sequences, and 232 individuals were examined using amplified fragment length polymorphisms (AFLP) to test whether the phylogeographical pattern and population genetics of this species are related to past geological events and/or climatic oscillations. No obvious trend of genetic diversity was found along a latitude/longitude gradient among different geographical groups. Phylogenetic analysis indicated three deep monophyletic clades that approximately correspond to three geographical regions separated by high mountains and a deep strait, and TCS analysis also revealed three disconnected networks, suggesting that spatial and temporal separations by vicariance, which were also supported by AMOVA as a source of the molecular variance presented among groups. Gene flow analysis showed that there had been frequent historical gene flow among local populations in different regions, but the networks exhibited no shared haplotype among populations. In conclusion, the past geological events and climatic fluctuations are the most important factor on the phylogeographical structure and genetic patterns of O. hyla intricata in Southeast Asia. Habitat, vegetation, and anthropogenic effect may also contribute to gene flow and introgression of this species. Moreover, temperature, abundant rainfall and a diversity of graminaceous species are beneficial for the migration of O. hyla intricata. High haplotype diversity, deep phylogenetic division, negative Fu’s F _s values and unimodal and multimodal distribution shapes all suggest a complicated demographic expansion pattern of these O. hyla intricata populations, which might have been caused by climatic oscillations during glacial periods in the Quaternary.     

DNA barcode discovers two cryptic species and two geographical radiations in the invasive drosophilid Zaprionus indianus

Comparing introduced to ancestral populations within a phylogeographical context is crucial in any study aiming to understand the ecological genetics of an invasive species. Zaprionus indianus is a cosmopolitan drosophilid that has recently succeeded to expand its geographical range upon three continents (Africa, Asia and the Americas). We studied the distribution of mitochondrial DNA (mtDNA) haplotypes for two genes (CO‐I and CO‐II) among 23 geographical populations. mtDNA revealed the presence of two well‐supported phylogenetic lineages (phylads), with bootstrap value of 100%. Phylad I included three African populations, reinforcing the African‐origin hypothesis of the species. Within phylad II, a distinct phylogeographical pattern was discovered: Atlantic populations (from the Americas and Madeira) were closer to the ancestral African populations than to Eastern ones (from Madagascar, Middle East and India). This means that during its passage from endemism to cosmopolitanism, Z. indianus exhibited two independent radiations, the older (the Eastern) to the East, and the younger (the Atlantic) to the West. Discriminant function analysis using 13 morphometrical characters was also able to discriminate between the two molecular phylads (93.34 ± 1.67%), although detailed morphological analysis of male genitalia using scanning electron microscopy showed no significant differences. Finally, crossing experiments revealed the presence of reproductive barrier between populations from the two phylads, and further between populations within phylad I. Hence, a bona species status was assigned to two new, cryptic species: Zaprionus africanus and Zaprionus gabonicus, and both were encompassed along with Z. indianus and Zaprionus megalorchis into the indianus complex. The ecology of these two species reveals that they are forest dwellers, which explains their restricted endemic distribution, in contrast to their relative cosmopolitan Z. indianus, known to be a human‐commensal. Our results reconfirm the great utility of mtDNA at both inter‐ and intraspecific analyses within the frame of an integrated taxonomical project.     

Genetic structure of Anogeissus dhofarica (Combretaceae) populations endemic to the monsoonal fog oases of the southern Arabian Peninsula

Anogeissus dhofarica (Combretaceae) is an endemic tree of the monsoon affected coastal mountains of the southern Arabian Peninsula, being the character species of the Hybantho durae–Anogeissetum dhofaricae association, a drought deciduous, monsoon forest community found only in the Dhofar region of southern Oman and the eastern Al‐Mahra region of south‐east Yemen. Due to the steep precipitation gradient from the centre to the edges in this monsoon affected area, A. dhofarica is found in two different habitat types: in continuous woodland belts of the Hawf and Dhofar mountains, and in isolated, scattered woodland patches, as found especially in the Fartak Mts (south‐east Yemen). Fifteen populations (212 individuals) from across the whole distribution area of the species were analysed using amplified fragment length polymorphism fingerprinting to: (1) evaluate the consequences of population fragmentation on the genetic diversity harboured in isolated patches versus cohering stands of the species and (2) to reconstruct the phylogeographical pattern of A. dhofarica as a consequence of oscillations in the monsoon activity during the Pleistocene and Holocene. The analysis of among‐population genetic differentiation and within‐population genetic diversity in A. dhofarica populations resulted in a lack of genetic pauperization and genetic differentiation of populations of the distinctly isolated patches of the Fartak Mts compared to the more luxurious forests of the Hawf and Dhofar regions. This is considered to be due to the high buffer capacity against the loss of genetic diversity caused by the long‐lived life‐form of the species combined with the capability to propagate clonally and the relatively recent fragmentation of Anogeissus forests into the described patches rather than due to high values of gene flow among remnant populations caused by bee pollination and anemochorical and hydrochorical diaspore dispersal. The phylogeographical pattern of the species argues for a quite recent fragmentation of a once continuous forest belt of A. dhofarica that is rather connected with climate changes in the Holocene than triggered by aridity–humidity oscillations reported for the Pleistocene. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 97 , 40–51.     

Phylogeographic structure and regional history of Lemniscomys striatus (Rodentia: Muridae) in tropical Africa

Aim This study aims to elucidate the phylogeography of the murid rodent Lemniscomys striatus and to evaluate the relative roles of ecological change, habitat patchiness, rivers and geological barriers in structuring patterns of diversity. Location Sub‐Saharan Africa. Methods The extent of phylogeographic patterns and molecular genetic diversity (cytochrome b gene) were addressed in a survey of 128 individuals of L. striatus from 42 localities. Using maximum parsimony, maximum likelihood, Bayesian, network and genetic structure analyses, we inferred intraspecific relationships and tested hypotheses for historical patterns of gene flow within L. striatus. Results Our results identified four major geographical clades within L. striatus: a West African clade, a Benin‐Nigeria clade, a Central African clade, and an East African clade. Several subclades were identified within these four major clades. Restricted gene flow with isolation by distance was recorded, which is congruent with the low dispersal ability of such a small murid rodent. No clear signal of population expansion was detected within clades or subclades. Main conclusions The western rift system and the Volta and Niger rivers may have acted as long‐term extrinsic barriers to gene flow, resulting in the emergence of the four main clades of L. striatus. The observed pattern of mitochondrial variation observed within each clade probably results from late Pleistocene climatic and vegetation changes: during adverse conditions (forest expansion), L. striatus may have survived only in refugia, and then experienced range expansion under favourable conditions (savanna expansion).     

Distribution and population genetic structure of the Mediterranean pine shoot beetle Tomicus destruens in the Iberian Peninsula and Southern France

1 The Mediterranean pine shoot beetle Tomicus destruens has long been indistinguishable from its congeneric Tomicus piniperda. Both species attack pines, and can be found in sympatry. The geographical distribution of T. destruens is still unclear in most of the Mediterranean Basin. 2 We aimed to describe the geographical distribution and zones of sympatry of both species in the Iberian Peninsula and France, and to study the molecular phylogeographical pattern of T. destruens. 3 Tomicus spp. adults were sampled in Portugal, Spain and France, and a portion of the mitochondrial genes COI and COII was sequenced for 84 individuals. Sequences were aligned to a data set previously obtained from French localities. 4 Tomicus destruens was found in all populations, except for one locality in Portugal and in the Landes (France). It was in sympatry with T. piniperda in two locations on Pinus pinaster and one location on Pinus radiata. 5 Within‐population genetic diversity was high, but we found a significant pattern of spatial distribution of genetic variation, as well as a significant effect of the host tree. 6 The data suggest the existence of two glacial refugia, from which T. destruens recolonized its current range. One refugium was located in Portugal where the beetle probably evolved on P. pinaster. The corresponding haplotypes show a West–East frequency gradient. The other refugium was probably in the eastern range, where the beetles evolved on Pinus halepensis and P. pinea. The corresponding haplotypes show an East–West frequency gradient.     

Phylogeography of the Eurasian green woodpecker (Picus viridis)

Aim In this paper we investigate the evolutionary history of the Eurasian green woodpecker (Picus viridis) using molecular markers. We specifically focus on the respective roles of Pleistocene climatic oscillations and geographical barriers in shaping the current population genetics within this species. In addition, we discuss the validity of current species and subspecies limits. Location Western Palaearctic: Europe to western Russia, and Africa north of the Sahara. Methods We sequenced two mitochondrial genes and five nuclear introns for 17 Eurasian green woodpeckers. Multilocus phylogenetic analyses were conducted using maximum likelihood and Bayesian algorithms. In addition, we sequenced a fragment of the cytochrome b gene (cyt b, 427 bp) and of the Z‐linked BRM intron 15 for 113 and 85 individuals, respectively. The latter data set was analysed using population genetic methods. Results Our phylogenetic results support the monophyly of Picus viridis and suggest that this taxon comprises three allopatric/parapatric lineages distributed in North Africa, the Iberian Peninsula and Europe, respectively. The North African lineage split from the Iberian/European clade during the early Pleistocene (1.6–2.2 Ma). The divergence event between the Iberian and the European lineages occurred during the mid‐Pleistocene (0.7–1.2 Ma). Our results also support a post‐glacial range expansion of these two lineages from distinct refugia located in the Iberian Peninsula and possibly in eastern Europe or Anatolia, which led to the establishment of a secondary contact zone in southern France. Main conclusions Our results emphasize the crucial role of both Pleistocene climatic oscillations and geographical barriers (Strait of Gibraltar, Pyrenees chain) in shaping the current genetic structure of the Eurasian green woodpecker. Our molecular data, in combination with diagnosable plumage characters, suggest that the North African green woodpecker (Levaillant’s woodpecker) merits species rank as Picus vaillantii (Malherbe, 1847). The two European lineages could be distinguished by molecular and phenotypic characters over most of their respective geographical ranges, but they locally exchange genes in southern France. Consequently, we prefer to treat them as subspecies (P. viridis viridis, P. viridis sharpei) pending further studies.     

Evidence of history in explaining diversity patterns in tropical riverine fish

Aim Documentation of the ongoing effect of rain forest refuges at the last glacial maximum (LGM) on patterns of tropical freshwater fish diversity. Location Tropical South and Central America, and West Africa. Methods LGM rain forest regions and species richness by drainage were compiled from published data. GIS mapping was applied to compile drainage area and contemporary primary productivity. We used multiple regression analyses, applied separately for Tropical South America, Central America and West Africa, to assess differences in species richness between drainages that were connected and disconnected to rain forest refuge zones during the LGM. Spatial autocorrelation of the residuals was tested using Moran's I statistic. We added an intercontinental comparison to our analyses to see if a historical signal would persist even when a regional historical effect (climate at the LGM) had already been accounted for. Results Both area and history (contact with LGM rain forest refuge) explained the greatest proportions of variance in the geographical pattern of riverine species richness. In the three examined regions, we found highest richness in drainages that were connected to the rain forest refuges. No significant residual spatial autocorrelation was detected after considering area, primary productivity and LGM rain forest refuges. These results show that past climatic events still affect West African and Latin American regional and continental freshwater fish richness. At the continental scale, we found South American rivers more species‐rich than expected on the basis of their area, productivity and connectedness to rain forest refuge. Conversely, Central American rivers were less species‐diverse than expected by the grouped model. African rivers were intermediate. Therefore, a historical signal persists even when a regional historical effect (climate at the LGM) had already been accounted for. Main conclusions It has been hypothesized that past climatic events have limited impact on species richness because species have tracked environmental changes through range shifts. However, when considering organisms with physically constrained dispersal (such as freshwater fish), past events leave a perceptible imprint on present species diversity. Furthermore, we considered regions that have comparable contemporary climatic and environmental characteristics, explaining the absence of a productivity effect. From the LGM to the present day (a time scale of 18,000 years), extinction processes should have played a predominant role in shaping the current diversity pattern. By contrast, the continental effects could reflect historical contingencies explained by differences in speciation and extinction rates between continents at higher time scales (millions of years).     

Phylogeography of the humbug damselfish,Dascyllus aruanus (Linnaeus, 1758): evidence of Indo‐Pacific vicariance and genetic differentiation of peripheral populations

The phylogeographical structure of coral‐associated reef fishes may have been severely affected, more than species from deeper habitats, by habitat loss during periods of low sea level. The humbug damselfish, Dascyllus aruanus, is widely distributed across the Indo‐West Pacific, and exclusively inhabits branching corals. We used mitochondrial cytochrome b sequence and seven microsatellite loci on D. aruanus samples (260 individuals) from 13 locations across the Indo‐West Pacific to investigate its phylogeographical structure distribution‐wide. A major genetic partition was found between the Indian and Pacific Ocean populations, which we interpret as the result of geographical isolation on either side of the Indo‐Pacific barrier during glacial periods. The peripheral populations of the Red Sea and the Society Islands exhibited lower genetic diversity, and substantial genetic differences with the other populations, suggesting relative isolation. Thus, vicariance on either side of the Indo‐Pacific barrier and peripheral differentiation are considered to be the main drivers that have shaped the phylogeographical patterns presently observed in D. aruanus. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 931–942.