Coalescent analyses support multiple mainland-to-island dispersals in the evolution of Malagasy Triaenops bats (Chiroptera: Hipposideridae)

Aim We investigate the directionality of mainland‐to‐island dispersals, focusing on a case study of an African‐Malagasy bat genus, Triaenops (Hipposideridae). Taxa include T. persicus from east Africa and three Triaenops species from Madagascar (T. auritus, T. furculus, and T. rufus). The evolution of this bat family considerably post‐dated the tectonic division of Madagascar from Africa, excluding vicariance as a viable hypothesis. Therefore, we consider three biogeographical scenarios to explain these species' current ranges: (A) a single dispersal from Africa to Madagascar with subsequent speciation of the Malagasy species; (B) multiple, unidirectional dispersals from Africa to Madagascar resulting in multiple, independent Malagasy lineages; or (C) early dispersal of a proto‐species from Africa to Madagascar, with later back‐dispersal of a descendant Malagasy taxon to Africa. Location East Africa, Madagascar, and the Mozambique Channel. Methods We compare the utility of phylogenetic and coalescent methodologies to address the question of directionality in a mainland‐to‐island dispersal event for recently diverged taxa. We also emphasize the application of biologically explicit demographic systems, such as the non‐equilibrium isolation‐with‐migration model. Here, these methods are applied to a four‐species haploid genetic data set, with simulation analyses being applied to validate this approach. Results Coalescent simulations favour scenario B: multiple, unidirectional dispersals from Africa to Madagascar resulting in multiple, independent Malagasy bat lineages. From coalescent dating, we estimate that the genus Triaenops was still a single taxon approximately 2.25 Ma. The most recent Africa to Madagascar dispersal occurred much more recently (c. 660 ka), and led to the formation of the extant Malagasy species, T. rufus. Main conclusions Haploid genetic data from four species of Triaenops are statistically most consistent with multiple, unidirectional dispersals from mainland Africa to Madagascar during the late Pleistocene.     

Defining spatial and temporal patterns of phylogeographic structure in Madagascar's iguanid lizards (genus Oplurus)

Understanding the remarkably high species diversity and levels of endemism found among Madagascar’s flora and fauna has been the focus of many studies. One hypothesis that has received much attention proposes that Quaternary climate fluctuations spurred diversification. While spatial patterns of distribution and phylogenetic relationships can provide support for biogeographic predictions, temporal estimates of divergence are required to determine the fit of these geospatial patterns to climatic or biogeographic mechanisms. We use multilocus DNA sequence data to test whether divergence times among Malagasy iguanid lizards of the subfamily Oplurinae are compatible with a hypotheses of Pliocene–Pleistocene diversification. We estimate the oplurine species tree and associated divergence times under a relaxed‐clock model. In addition, we examine the phylogeographic structure and population divergence times within two sister species of Oplurus primarily distributed in the north‐west and south‐west of Madagascar (Oplurus cuvieri and Oplurus cyclurus, respectively). We find that divergence events among oplurine lineages occurred in the Oligocene and Miocene and are thus far older and incompatible with the hypothesis that recent climate fluctuations are related to current species diversity. However, the timing of intraspecific divergences and spatial patterns of population genetic structure within O. cuvieri and O. cyclurus suggest a role for both intrinsic barriers and recent climate fluctuations at population‐level divergences. Integrating information across spatial and temporal scales allows us to identify and better understand the mechanisms generating patterns diversity.     

Differential effects of historical migration,glaciations and human impact on the genetic structure and diversity of the mountain pasture weed Veratrum album L.

Aim Today’s genetic population structure and diversity of species can be understood as the result of range expansion from the area of origin, past climatic oscillations and contemporary processes. We examined the relative importance of these factors in Veratrum album L., a toxic weed of mountain grasslands. Location Continental Europe. Methods Forty populations from the Asian border (Urals and Caucasus) to Portugal were studied using amplified fragment length polymorphisms (AFLPs) combined with selected plant and population measures. The data were analysed with phylogenetic, population genetic and regression methods inferring both genetic structure and diversity from geographic and ecological factors. Results Fragment frequency clines together with genetic distance clustering and principal coordinates analysis indicated an east–west direction in the genetic structure of V. album, suggesting ancient migration into Europe from a proposed Asian origin. However, the strong geographic pattern in the genetic structure, pronounced isolation by distance (R² = 0.74) and moderate overall population differentiation (F_ST = 0.13) suggests high historical gene flow, possibly during glacials, and vicariance into mountainous regions during interglacials. Occurrence of V. album during the last glaciation in several areas along the periphery of the Alps and recolonization of this mountain range from both eastern and central–western areas was indicated. Genetic diversity was highest in central Europe, a pattern that did not agree with the expectations from east–west migration into Europe. Furthermore, managed habitats showed higher levels of genetic diversity compared to unmanaged habitats. Stepwise linear regression determined shoot density and soil phosphorus as the main predictors of within‐population genetic diversity (R² = 0.40). Main conclusions Our results showed that V. album retained genetic imprints of historical range expansion into Europe, although this was alleviated by the influence of climatic oscillations and contemporary processes. For example, genetic population structure was strongly affected by post‐glacial vicariance while patterns of genetic diversity seemed mainly to be influenced by human land use. Our findings highlight the importance of applying a synthetic approach, testing the influence of both historical and contemporary processes on genetic structure and diversity in order to understand complex phylogeographic patterns. This may especially apply to widespread species, such as weeds. Implications of our findings for biological control are briefly discussed.     

Genetic structure and phylogeography of a European flagship species,the white‐tailed sea eagle Haliaeetus albicilla

We analysed 120 white‐tailed sea eagles Haliaeetus albicilla from eastern (Poland and Estonia) and southeastern (Serbian Danube population) Europe for genetic variability and structuring at the mitochondrial control region and seven nuclear microsatellite loci. We combined this new dataset with sequence and genotype data from previous analyses covering Greenland and Eurasia (total sample sizes of 420 and 186 individuals for mtDNA and microsatellites, respectively) to address the following questions: 1) does the large eastern population in Europe add significantly to the species‘ overall genetic diversity? 2) Do the new sequence data match the clinal distribution pattern (west to east) of the two major mtDNA lineages? 3) Does the preliminary hypothesis of two nuclear genetic clusters recently found in this species hold for the whole of Europe, and do these clusters show a geographic pattern? Our results confirmed Europe as a stronghold of genetic diversity in white‐tailed sea eagles, and the east of the continent contributed disproportionately to this, the reason being the admixture of eagles with different genetic background. As hypothesised, both mitochondrial lineages were recovered also in eastern Europe, but the globally more eastern lineage was dominant. The presence of two microsatellite clusters was also confirmed, and these groups, too, show a non‐random geographic distribution, with, except for Poland, a high proportion of ‘eastern‐type’ eagles in the populations of east–central and eastern Europe.     

Vicariance patterns in the Mediterranean Sea: east–west cleavage and low dispersal in the endemic seagrass Posidonia oceanica

Aim The seagrass, Posidonia oceanica is a clonal angiosperm endemic to the Mediterranean Sea. Previous studies have suggested that clonal growth is far greater than sexual recruitment and thus leads to low clonal diversity within meadows. However, recently developed microsatellite markers indicate that there are many different genotypes, and therefore many distinct clones present. The low resolution of markers used in the past limited our ability to estimate clonality and assess the individual level. New high‐resolution dinucleotide microsatellites now allow genetically distinct individuals to be identified, enabling more reliable estimation of population genetic parameters across the Mediterranean Basin. We investigated the biogeography and dispersal of P. oceanica at various spatial scales in order to assess the influence of different evolutionary factors shaping the distribution of genetic diversity in this species. Location The Mediterranean. Methods We used seven hypervariable microsatellite markers, in addition to the five previously existing markers, to describe the spatial distribution of genetic variability in 34 meadows spread throughout the Mediterranean, on the basis of an average of 35.6 (± 6.3) ramets sampled. Results At the scale of the Mediterranean Sea as a whole, a strong east–west cleavage was detected (amova) . These results are in line with those obtained using previous markers. The new results showed the presence of a putative secondary contact zone at the Siculo‐Tunisian Strait, which exhibited high allelic richness and shared alleles absent from the eastern and western basins. F statistics (pairwise θ ranges between 0.09 and 0.71) revealed high genetic structure between meadows, both at a small scale (about 2 to 200 km) and at a medium scale within the eastern and western basins, independent of geographical distance. At the intrameadow scale, significant spatial autocorrelation in six out of 15 locations revealed that dispersal can be restricted to the scale of a few metres. Main conclusions A stochastic pattern of effective migration due to low population size, turnover and seed survival is the most likely explanation for this pattern of highly restricted gene flow, despite the importance of an a priori seed dispersal potential. The east–west cleavage probably represents the outline of vicariance caused by the last Pleistocene ice age and maintained to this day by low gene flow. These results emphasize the diversity of evolutionary processes shaping the genetic structure at different spatial scales.     

Postglacial recolonization shaped the genetic diversity of the winter moth (Operophtera brumata) in Europe

Changes in climate conditions, particularly during the Quaternary climatic oscillations, have long been recognized to be important for shaping patterns of species diversity. For species residing in the western Palearctic, two commonly observed genetic patterns resulting from these cycles are as follows: (1) that the numbers and distributions of genetic lineages correspond with the use of geographically distinct glacial refugia and (2) that southern populations are generally more diverse than northern populations (the “southern richness, northern purity” paradigm). To determine whether these patterns hold true for the widespread pest species the winter moth (Operophtera brumata), we genotyped 699 individual winter moths collected from 15 Eurasian countries with 24 polymorphic microsatellite loci. We find strong evidence for the presence of two major genetic clusters that diverged ~18 to ~22 ka, with evidence that secondary contact (i.e., hybridization) resumed ~ 5 ka along a well‐established hybrid zone in Central Europe. This pattern supports the hypothesis that contemporary populations descend from populations that resided in distinct glacial refugia. However, unlike many previous studies of postglacial recolonization, we found no evidence for the “southern richness, northern purity” paradigm. We also find evidence for ongoing gene flow between populations in adjacent Eurasian countries, suggesting that long‐distance dispersal plays an important part in shaping winter moth genetic diversity. In addition, we find that this gene flow is predominantly in a west‐to‐east direction, suggesting that recently debated reports of cyclical outbreaks of winter moth spreading from east to west across Europe are not the result of dispersal.     

The east‐west‐north colonization history of the Mediterranean and Europe by the coastal plant Carex extensa (Cyperaceae)

Coastal plants are ideal models for studying the colonization routes of species because of the simple linear distributions of these species. Carex extensa occurs mainly in salt marshes along the Mediterranean and European coasts. Variation in cpDNA sequences, amplified fragment length polymorphisms (AFLPs) and simple sequence repeats (SSRs) of 24 populations were analysed to reconstruct its colonization history. Phylogenetic relationships indicate that C. extensa together with the South American Carex vixdentata and the southern African Carex ecklonii form a monophyletic group of halophilic species. Analyses of divergence times suggest that early lineage diversification may have occurred between the late Miocene and the late Pliocene (Messinian crisis). Phylogenetic and network analyses of cpDNA variation revealed the monophyly of the species and an ancestral haplotype contained in populations of the eastern Mediterranean. The AFLP and SSR analyses support a pattern of variation compatible with these two lineages. These analyses also show higher levels of genetic diversity and differentiation in the eastern population group, which underwent an east‐to‐west Mediterranean colonization. Quaternary climatic oscillations appear to have been responsible for the split between these two lineages. Secondary contacts may have taken place in areas near the Ligurian Sea in agreement with the gene flow detected in Corsican populations. The AFLP and SSR data accord with the ‘tabula rasa’ hypothesis in which a recent and rapid colonization of northern Europe took place from the western Mediterranean after the Last Glacial Maximum. The unbalanced west‐east vs. west‐north colonization may be as a result of ‘high density blocking’ effect.     

Longevity in some Malagasy rainforest passerines

Data on longevity is a prerequisite to understand the life-history strategies of a species. For Malagasy songbirds no information has been previously published. Therefore, we studied their longevity in a capture–recapture effort in a rainforest in eastern Madagascar (Maromizaha, Andasibe) between 2003 and 2016. We present first data on the longevity of 23 species of Malagasy songbirds. A female Dark Newtonia Newtonia amphichroa (Vangidae) and a Grey-crowned Greenbul Bernieria cinereiceps (Bernieridae) attained an age of at least 12 years, followed by two male Madagascar Brush Warblers Nesillas typica (Acrocephalidae), female Madagascar Bulbul Hypsipetes madagascariensis (Pycnonotidae) and Madagascar Drongo Dicrurus forficatus (Dicruridae) that attained at least 10 years. There was much variation within some taxonomic groups, longevity did not increase with the mass of a bird species and most insectivorous birds lived longer than granivorous ones.     

The role of altitude and associated habitat stability in determining patterns of population genetic structure in two species of Atalophlebia (Ephemeroptera: Leptophlebiidae)

1. Previous studies have identified lowland areas as barriers to gene flow (dispersal) between distinct mountain ranges in montane species of aquatic insects. In this study, we investigated the population genetic structure of two closely related Atalophlebia (mayfly) species inhabiting lowland areas of south‐east Queensland, Australia, with the expectation of widespread gene flow throughout the low‐altitude environment and associated homogeneous genetic structure. 2. In particular, we asked whether species with lower‐altitude distributions demonstrate greater spatial distribution of mtDNA (COI) alleles than the upland species studied previously. This pattern would be expected if good dispersal ability is associated with population persistence in these drought‐prone habitats. 3. The two species demonstrated contrasting genetic population structure. Atalophlebia sp. AV13 D revealed strong population structure, with populations on each side of the low‐altitude area isolated from each other for a long time (c.350 kya), and the presence of an isolation‐by‐distance pattern over relatively small geographical distances (<40 km). In contrast, Atalophlebia sp. AV13 A was panmictic at the scale investigated (≤160 km), with no history of past population fragmentation. 4. Examination of sample distribution along the altitudinal gradient reveals that Atalophlebia sp. AV13 D may have a more upland distribution (associated with greater habitat stability) than previously supposed, while Atalophlebia sp. AV13 A inhabits more xeric lowland areas, where freshwater habitats are less stable. We consequently hypothesise that these contrasting genetic population structures result from differences in habitat stability along the altitudinal gradient, only species with good dispersal ability being able to persist in unstable habitats. These findings may be applicable to other regions of the globe where habitat instability is associated with altitudinal gradients.     

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.     

Integrating multiple data sources to assess the distribution and abundance of bottlenose dolphins Tursiops truncatus in Scottish waters

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Heterogeneity, speciation/extinction history and climate: explaining regional plant diversity patterns in the Cape Floristic Region

This paper investigates the role of heterogeneity and speciation/extinction history in explaining variation in regional scale (c. 0.1–3000 km²) plant diversity in the Cape Floristic Region of south‐western Africa, a species‐ and endemic‐rich biogeographical region. We used species‐area analysis and analysis of covariance to investigate geographical (east vs. west) and topographic (lowland vs. montane) patterns of diversity. We used community diversity as a surrogate for biological heterogeneity, and the diversity of naturally rare species in quarter degree squares as an indicator of differences in speciation/extinction histories across the study region. We then used standard statistical methods to analyse geographical and topographic patterns of these two measures. There was a clear geographical diversity pattern (richer in the west), while a topographic pattern (richer in mountains) was evident only in the west. The geographical boundary coincided with a transition from the reliable winter‐rainfall zone (west) to the less reliable non‐seasonal rainfall zone (east). Community diversity, or biological heterogeneity, showed no significant variation in relation to geography and topography. Diversity patterns of rare species mirrored the diversity pattern for all species. We hypothesize that regional diversity patterns are the product of different speciation and extinction histories, leading to different steady‐state diversities. Greater Pleistocene climatic stability in the west would have resulted in higher rates of speciation and lower rates of extinction than in the east, where for the most, Pleistocene climates would not have favoured Cape lineages. A more parsimonious hypothesis is that the more predictable seasonal rainfall of the west would have favoured non‐sprouting plants and that this, in turn, resulted in higher speciation and lower extinction rates. Both hypotheses are consistent with the higher incidence of rare species in the west, and higher levels of beta and gamma diversity there, associated with the turnover of species along environmental and geographical gradients, respectively. These rare species do not contribute to community patterns; hence, biological heterogeneity is uniform across the region. The weak topography pattern of diversity in the west arises from higher speciation rates and lower extinction rates in the topographically complex mountains, rather than from the influence of environmental heterogeneity on diversity.     

Low local but high beta diversity of tropical forest dung beetles in Madagascar

Aims We have compared local (alpha) and regional (beta) species diversities of dung beetles in wet forests in the main tropical regions including Madagascar. Madagascar is exceptional in lacking native large herbivorous mammals which produce the key resource for dung beetles elsewhere. Location Central and South America, mainland Africa, Madagascar and Southeast Asia. Methods Trapping data on dung beetles and data on mammalian faunas were obtained from published and unpublished studies. We used our original data for Madagascar. Results Species richness of dung beetles and that of large‐bodied (> 15 mm length) species in particular were highly significantly explained by the regional number of large‐bodied (> 10 kg) mammals (R² from 50 to 80%). For a given pairwise spatial distance between two communities, beta diversity was significantly higher in Madagascar than elsewhere, explaining the very high total species richness in Madagascar in spite of low local diversity. Main conclusion The presence and numbers of large herbivorous mammals greatly influence the species richness of dung beetles in tropical wet forests. The lack of native large herbivores rather than a limited species pool explains the low local diversity in Madagascar. Exceptionally high beta diversity in Madagascar suggests a pattern of old radiation involving extensive allopatric speciation.     

Contrasting genetic diversity and population structure among three sympatric Madagascan shorebirds: parallels with rarity,endemism, and dispersal

Understanding the relative contributions of intrinsic and extrinsic factors to population structure and genetic diversity is a central goal of conservation and evolutionary genetics. One way to achieve this is through comparative population genetic analysis of sympatric sister taxa, which allows evaluation of intrinsic factors such as population demography and life history while controlling for phylogenetic relatedness and geography. We used ten conserved microsatellites to explore the population structure and genetic diversity of three sympatric and closely related plover species in southwestern Madagascar: Kittlitz's plover (Charadrius pecuarius), white‐fronted plover (C. marginatus), and Madagascar plover (C. thoracicus). Bayesian clustering revealed strong population structure in the rare and endemic Madagascar plover, intermediate population structure in the white‐fronted plover, and no detectable population structure in the geographically widespread Kittlitz's plover. In contrast, allelic richness and heterozygosity were highest for the Kittlitz's plover, intermediate for the white‐fronted plover and lowest for the Madagascar plover. No evidence was found in support of the “watershed mechanism” proposed to facilitate vicariant divergence of Madagascan lemurs and reptiles, which we attribute to the vagility of birds. However, we found a significant pattern of genetic isolation by distance among populations of the Madagascar plover, but not for the other two species. These findings suggest that interspecific variation in rarity, endemism, and dispersal propensity may influence genetic structure and diversity, even in highly vagile species.     

Landscape, habitat characteristics and the genetic population structure of two caddisflies

1. In a region of south‐eastern England, we investigated the hierarchical genetic structure of populations of two stream‐dwelling caddisflies (Trichoptera: Polycentropodidae) with contrasting distributions: Plectrocnemia conspersa inhabits numerous small, patchily distributed seeps and streams, while the confamilial Polycentropus flavomaculatus is found in fewer but larger streams and rivers. We also contrasted the genetic structure of P. conspersa in the lowland south‐east with that in an upland region in the north west. 2. Microsatellite genotypes were obtained from samples of both species taken from a ‘core area’ and at sites 15, 40 and 100 km from this core (two regions for P. conspersa, totalling 45 sites and 1405 larvae; one region for P. flavomaculatus, totalling 10 sites and 269 larvae). 3. The genetic structure of P. conspersa differed in the two regions. In the upland north‐west, significant genetic differentiation was observed at a spatial scale of around 40 km from the core, while there was no structure in the lowland south‐east up to around 100 km. Areas of high altitude did not appear directly to reduce gene flow, whereas other potential landscape barriers, including particular geological formations, large urban areas and the sea had a pronounced effect. 4. Weak genetic differentiation in P. conspersa across large distances, particularly in the lowland south‐east, suggests that it disperses strongly, facilitating gene flow within and between catchments. Conversely, for P. flavomaculatus we found strong genetic differentiation between almost all sites, suggesting that dispersal is much more limited. 5. Greater dispersal in the patchily distributed P. conspersa than in P. flavomaculatus, which occupies larger and presumably more persistent habitats, could be a general feature of other similarly distributed aquatic insects. While higher relief is potentially a partial barrier to dispersal, P. conspersamust have effective gene flow through such apparently inhospitable terrain, perhaps attributable to dispersal between neighbouring small and ephemeral populations. Indeed, its exploitation of headwaters and seeps requires the ability to disperse between such sites. Apparently it cannot, however, overcome more continuous barriers, consisting of large tracts of landscape with few habitable larval sites. Such landscapes, including those created by humans, may have a stronger effect on population connectivity and colonization in the longer term.     

Identifying refugia from climate change using coupled ecological and genetic data in a transitional Mediterranean‐temperate tree species

Populations occurring in areas of overlap between the current and future distribution of a species are particularly important because they can represent “refugia from climate change”. We coupled ecological and range‐wide genetic variation data to detect such areas and to evaluate the impacts of habitat suitability changes on the genetic diversity of the transitional Mediterranean‐temperate tree Fraxinus angustifolia. We sampled and genotyped 38 natural populations comprising 1006 individuals from across Europe. We found the highest genetic diversity in western and northern Mediterranean populations, as well as a significant west to east decline in genetic diversity. Areas of potential refugia that correspond to approximately 70% of the suitable habitat may support the persistence of more than 90% of the total number of alleles in the future. Moreover, based on correlations between Bayesian genetic assignment and climate, climate change may favour the westward spread of the Black Sea gene pool in the long term. Overall, our results suggest that the northerly core areas of the current distribution contain the most important part of the genetic variation for this species and may serve as in situ macrorefugia from ongoing climate change. However, rear‐edge populations of the southern Mediterranean may be exposed to a potential loss of unique genetic diversity owing to habitat suitability changes unless populations can persist in microrefugia that have facilitated such persistence in the past.