Phylogeographic Scenario Research Articles

Phylogeography of the endemic Gymnocypris chilianensis (Cyprinidae): Sequential westward colonization followed by allopatric evolution in response to cyclical Pleistocene glaciations on the Tibetan Plateau

The schizothoracine Gymnocypris chilianensis is restricted to the Shiyang, Ruoshui and Shule Rivers, listed from east to west, along the northeast edge of the Tibetan Plateau. This distribution provides a valuable system to test hypotheses about postglacial colonization. We used mitochondrial DNA sequence data (a control region and the cytochrome b gene; 1894 bp) to assess the phylogeographic structure of this species based on 278 specimens sampled from throughout the species’ entire geographical range. We found three lineages corresponding geographically to the three rivers, suggesting three independent glacial differentiation centers within the northeast edge of the Tibetan Plateau. The phylogenetic analysis suggested that the Shiyang River population forms a lineage that separated from the other populations of G. chilianensis at the basal phylogenetic split within this species. The molecular data further demonstrated a clear pattern of decreasing genetic diversity from the eastern Shiyang River towards the central Ruoshui River and western Shule River lineages, a pattern consistent with sequential western colonization. We therefore propose a phylogeographic scenario for G. chilianensis of a gradual westerly expansion from the Shiyang River population along the northeast edge of the Tibetan Plateau, with subsequent allopatric evolution at approximately 0.37 and 0.05 million years ago (Ma), through at least two glacial maxima. Together with the genetic evidence reported in other species, our findings suggest that this common biogeographic pattern emphasizes the importance of the northeastern edge region of the Tibetan Plateau as a hotspot of genetic diversity for some taxa.

Recent evolutionary history of the Iberian endemic lizards Podarcis bocagei (Seoane, 1884) and Podarcis carbonelli Pérez-Mellado, 1981 (Squamata: Lacertidae) revealed by allozyme and microsatellite markers

Podarcis bocagei and Podarcis carbonelli are two species of wall lizards endemic to the western Iberian Peninsula. A detailed phylogeographical study based on mitochondrial DNA (mtDNA) variation has shown that they responded differently to the Quaternary climatic oscillations. These differences have been attributed to their distribution patterns: P. bocagei is distributed in the north of the Peninsula and in a continuous fashion, whereas P. carbonelli has a more southern and fragmented distribution. In this study, we assessed whether nuclear markers reveal similar evolutionary patterns to those inferred from mtDNA variation. We studied a battery of allozyme and microsatellite loci in a geographically representative set of individuals from both species. For each species we evaluated overall levels of differentiation, patterns of geographical variation in genetic diversity, genetic relationships amongst localities, and applied model-based individual multilocus genotype clustering approaches to detect hidden population structure. Our results for P. bocagei are highly concordant with the phylogeographical scenario inferred from mtDNA variation: we found very low levels of population differentiation, consistent with survival in a single glacial refugium, and detected signatures of a rapid demographic and geographical expansion. The analyses of nuclear markers furthermore helped to identify a probable refugial area, as well as expansion routes. Additionally, in concordance with observations based on mtDNA variation, a low level of population differentiation was observed in P. carbonelli, but this was significantly higher than in P. bocagei. However, the geographical basis for differentiation in P. carbonelli is highly inconsistent between mtDNA and nuclear markers, suggesting a complex, albeit recent, history of fragmentation. A recent reduction of this species’ distribution has probably erased the signatures of glacial isolation and post-glacial expansion that are normally found in other Iberian species, suggesting that the currently observed pattern of genetic differentiation in this species was shaped more by recent genetic drift than by the Pleistocene climatic oscillations.

Phylogeography of the Italian vairone (Telestes muticellus, Bonaparte 1837) inferred by microsatellite markers: evolutionary history of a freshwater fish species with a restricted and fragmented distribution

BackgroundOwing to its independence from the main Central European drainage systems, the Italian freshwater fauna is characterized by a high degree of endemicity. Three main ichthyogeographic districts have been proposed in Italy. Yet, the validity of these regions has not been confirmed by phylogenetic and population genetic analyses and a phylogeographic scenario for Italy's primary freshwater fish fauna is still lacking. Here, we investigate the phylogeography of the Italian vairone (Telestes muticellus).ResultsWe sampled 38 populations representing the species' entire distribution range and covering all relevant drainage systems, and genotyped 509 individuals at eight variable microsatellite loci. Applying various population genetic analyses, we identify five distinct groups of populations that are only partly in agreement with the proposed ichthyogeographic districts. Our group I, which is formed by specimens from Veneto and the Po River system draining into the Adriatic Sea, corresponds to the Padano-Venetian ichthyogeographic district (PV), except for two Middle Adriatic drainages, which we identify as a separate group (III). The Tuscano-Latium district (TL) is equivalent to our group V. A more complex picture emerges for the Ligurian drainages: populations from Central Liguria belong to group I, while populations from West (group II) and East Liguria (group IV) form their own groups, albeit with affinities to PV and TL, respectively.ConclusionsWe propose a phylogeographic scenario for T. muticellus in which an initial T. muticellus stock became isolated from the 'Alpine' clade and survived the various glaciation cycles in several refugia. These were situated in the Upper Adriatic (groups I and II), the Middle Adriatic (group III), (East) Liguria (group IV) and Tuscano-Latium (group V). The population structure in the vairone is, in principal, in agreement with the two main ichthyogeographic districts (PV and TL), except for the two populations in the Middle Adriatic, which we identify as additional major "district".

Not just vicariance: phylogeography of a Sonoran Desert euphorb indicates a major role of range expansion along the Baja peninsula

To examine the generality of population-level impacts of ancient vicariance identified for numerous arid-adapted animal taxa along the Baja peninsula, we tested phylogeographical hypotheses in a similarly distributed desert plant, Euphorbia lomelii (Euphorbiaceae). In light of fossil data indicating marked changes in the distributions of Baja floristic assemblages throughout the Holocene and earlier, we also examined evidence for range expansion over more recent temporal scales. Two classes of complementary analytical approaches - hypothesis-testing and hypothesis-generating - were used to exploit phylogeographical signal from chloroplast DNA sequence data and genotypic data from six codominant nuclear intron markers. Sequence data are consistent with a scenario of mid-peninsular vicariance originating c. 1 million years ago (Ma). Alternative vicariance scenarios representing earlier splitting events inferred for some animals (e.g. Isthmus of La Paz inundation, c. 3 Ma; Sea of Cortez formation, c. 5 Ma) were rejected. Nested clade phylogeographical analysis corroborated coalescent simulation-based inferences. Nuclear markers broadened the temporal spectrum over which phylogeographical scenarios could be addressed, and provided strong evidence for recent range expansions along the north-south axis of the Baja peninsula. In contrast to previous plant studies in this region, however, the expansions do not appear to have been in a strictly northward direction. These findings contribute to a growing appreciation of the complexity of organismal responses to past climatic and geological changes - even when taxa have evolved in the same landscape context.

The limits of mtDNA phylogeography: complex patterns of population history in a highly structured Iberian lizard are only revealed by the use of nuclear markers

Phylogeographic analyses based on the sole use of the mitochondrial DNA (mtDNA) molecule reveal only a small part of the evolutionary history of a species or a set of related species. In this study, we have combined the application of slow- and fast-evolving nuclear markers (proteins and microsatellites, respectively) together with the analysis of two-gene genealogies to further understand the history of the Iberian endemic Schreiber's green lizard, Lacerta schreiberi, a species for which a well established phylogeographical scenario is available. In sharp contrast with the observation of four divergent and almost allopatric mtDNA clades, our nuclear data revealed how two groups of populations diverged, persisted and began to admix along the mountains of the Iberian Central System. In addition, the combination of mtDNA and nuclear data showed how the core area of the species distribution responded to ice ages, first by relatively old processes of population expansion to the south followed by episodes of contraction that are at the origin of present-day isolates, and more recently by a postglacial expansion to the Iberian Northwest where new habitats were made available after climatic amelioration. Taken together with recently published results for a variety of other organisms, our results suggest that complex processes of fragmentation, expansion and admixture can only be properly addressed through the use of several and complementary types of molecular markers. Finally, we also suggest that southern European refugia are both hotspots and melting pots of genetic diversity.

Genetic structure and diversity in Ramonda myconi (Gesneriaceae): effects of historical climate change on a preglacial relict species

The importance of the Mediterranean Basin as a long-term reservoir of biological diversity has been widely recognized, although much less effort has been devoted to understanding processes that allow species to persist in this area. Ramonda myconi (Gesneriaceae) is a Tertiary relict plant species restricted to the NE Iberian Peninsula. We used RAPD and chloroplast markers to assess the patterns of genetic structure in eight mountain regions covering almost the full species range, to identify the main historical processes that have shaped its current distribution and to infer the number and location of putative glacial refugia. While no cpDNA variation was detected, the species had relatively high levels of RAPD variation. Maximum levels of diversity were found within populations (71%), but there was also a significant differentiation between geographical regions (20%) and among populations within regions (9%). A spatial AMOVA identified three main groups of populations, corresponding to previously recognized centers of endemism and species richness. In addition, we found a marked geographical pattern of decreasing genetic diversity and increasing population differentiation from west to east. Our results support a complex phylogeographic scenario in the Iberian Peninsula of "refugia-within-refugia" and suggest that the higher diversity observed in western regions might be associated with prolonged and more stable climatic conditions in this area during the Quaternary.

Species delimitation in the Acomys cahirinus-dimidiatus complex (Rodentia, Muridae) inferred from chromosomal and morphological analyses

Our earlier chromosome banding studies of Acomys cahirinus and Acomys dimidiatus (the latter long considered to be a subspecies of the former) revealed that, despite very close diploid numbers (36 vs. 38), these taxa possess sharply different karyotypes and undoubtedly belong to different species. In this context, the taxonomic status and the relationship between the two chromosomal forms in Sinai (2n = 36) and Israel (2n = 38), chromosomally homozygous across a vast range except for a very narrow hybrid zone, remain poorly documented. Neither of these forms have previously been studied by chromosome banding; thus, the exact nature of chromosomal differences as well as the species to which these forms should be assigned remain unknown. Here, we present the data on comparative G-banding analysis and morphometrics of Acomys from Israel, Sinai, and Saudi Arabia, and a hybrid obtained in laboratory crosses between latter two. The analysis revealed that karyotype of Acomys from Israel is identical to that of Acomys from Saudi Arabia and both are different from that of Acomys from Sinai by one Robertsonian fusion. Therefore, karyotypically, all three are very different from A. cahirinus. It follows from the study that Sinai and probably Arabian peninsula and Minor Asia must be excluded from geographical distribution of A. cahirinus, which is limited from West Sahara to Egypt along Nile river (except Sinai). Furthermore, the synthesis of chromosomal and recent molecular data suggests a phylogeographical scenario explaining the modern distribution of Acomys in the Sinai and Arabian peninsulas and permits the update of the taxonomic status of these populations.

Genealogy of the nuclear β-fibrinogen locus in a highly structured lizard species: comparison with mtDNA and evidence for intragenic recombination in the hybrid zone

The study of nuclear genealogies in natural populations of nonmodel organisms is expected to provide novel insights into the evolutionary history of populations, especially when developed in the framework of well-established mtDNA phylogeographical scenarios. In the Iberian Peninsula, the endemic Schreiber's green lizard Lacerta schreiberi exhibits two highly divergent and allopatric mtDNA lineages that started to split during the late Pliocene. In this work, we performed a fine-scale analysis of the putative mtDNA contact zone together with a global analysis of the patterns of variation observed at the nuclear beta-fibrinogen intron 7 (beta-fibint7). Using a combination of DNA sequencing with single-strand conformational polymorphism (SSCP) analysis, we show that the observed genealogy at the beta-fibint7 locus reveals extensive admixture between two formerly isolated lizard populations while the two mtDNA lineages remain essentially allopatric. In addition, a private beta-fibint7 haplotype detected in the single population where both mtDNA lineages were found in sympatry is probably the result of intragenic recombination between the two more common and divergent beta-fibint7 haplotypes. Our results suggest that the progressive incorporation of nuclear genealogies in investigating the ancient demography and admixture dynamics of divergent genomes will be necessary to obtain a more comprehensive picture of the evolutionary history of organisms.

MtDNA Haplogroup Analysis of Black Brazilian and Sub-Saharan Populations: Implications for the Atlantic Slave Trade

Seventy individuals from two African and four black Brazilian populations were studied for the first hypervariable segment of mtDNA. To delineate a more complete phylogeographic scenario of the African mtDNA haplogroups in Brazil and to provide additional information on the nature of the Atlantic slave trade, we analyzed our data together with previously published data. The results indicate different sources of African slaves for the four major Brazilian regions. In addition, the data revealed patterns that differ from those expected on the basis of historical registers, thus suggesting the role of ethnic sex differences in the slave trade.

When morphometry meets genetics: inferring the phylogeography ofCarabus solieriusing Fourier analyses of pronotum and male genitalia

Population differentiation is a crucial step in the speciation process and is therefore a central subject in studies of microevolution. Assessing divergence and inferring its dynamics in space and time generally require a wide array of markers. Until now however, most studies of population structure are based on molecular markers and those concerning morphological traits are more scarce. In the present work, we studied morphological differentiation among populations of the ground beetle Carabus solieri, and tested its congruence with genetic population structure. The shape of pronotum and aedeagus was assessed using Dual Axis Fourier Shape Analysis. manova on Fourier coefficients revealed highly significant morphological variation between populations and a similar geographical pattern of differentiation for both structures. On the whole, morphological and genetic patterns were also found to be congruent. Our analysis confirms the phylogeographical scenario proposing that two entities of C. solieri differentiated during the last glaciation events before recolonizing the actual range of the species. It also indicates a large introgression between the two differentiated entities in the centre of the range.

Phylogeny and karyotype evolution of the Iberian Leptynia attenuata species complex (Insecta Phasmatodea)

An in-depth analysis of the Leptynia attenuata species complex has been performed by cytochrome oxidase subunit 2 ( cox2) gene sequencing as well as karyotype and allozyme analysis. The whole set of data allows to largely resolve the taxonomy of the group and suggests an overall trend of chromosomal repatternings through a progressive reduction of the chromosome number. A previously suggested new species has been also confirmed on a genetic basis. Data are discussed in order to depict a phylogenetic and phylogeographic scenario fitting the observed genetic relationships between the different species of the group. Chromosome rearrangements are proposed as the major speciation driving force within the group and androgenetic reproduction is suggested as a shortcut to overcome the problem of fixing chromosomal rearrangements that are strongly underdominant in heterozygotes.

Phylogeography of the California Thrasher (Toxostoma Redivivum) Based on Nested-Clade Analysis of Mitochondrial-DNA Variation

AbstractDistribution of genealogical lineages within a species is likely the result of a complicated series of ecological and historical events. Nested-clade analysis is specifically designed as an objective phylogeographic approach for inferring evolutionary processes on a spatial and temporal scale for small subclades within a larger set of intraspecific relationships. Here, we use nested-clade analysis as well as other phylogeographic methods to investigate the evolutionary history of California Thrasher (Toxostoma redivivum) populations. Inferences resulting from nested clade analysis suggest a history that includes past fragmentation, range expansion, and isolation-by-distance. Along with root information, those inferences enable the construction of a biogeographic scenario for this species involving general southern ancestry, an early north–south division, northward range expansion, and a southward back-expansion into an already populated southern region. Isolation-by-distance is also identified, particularly in southern California, indicating that gene flow between localities does occur but is restricted. Many conclusions drawn from this study are concordant with geologic data as well as phylogeographic scenarios drawn for other codistributed California taxa.

Setting conservation priorities: the case study of Carabus solieri (Col. Carabidae)

C. solieri is an endangered species of Carabidae which is mostly associated with humid forests of the Alps, in Prance and in Liguria (Italy). We reanalysed morphological data published by Bonadona and compared the results to the molecular data set presented here. We used partial sequences of the mitochondrial cytochrome b gene and seven microsatellite loci to evaluate genetic diversity among and between populations and to propose a phylogeographic scenario. We recognised two basic entities, considered as subspecies, which probably colonised the present distribution range from two main refugia (in France and in Italy). There is no strict agreement between morphological and molecular data and we propose that hybridisation and introgres-sion between the two subspecies have led to the observed patterns of distribution of the characters. Our results show that sampled populations differ significantly in microsatellite allele frequencies even though some populations are < 15 km apart. Gene flow estimates between the 19 sampled populations indicated very restricted exchange. This is in agreement with the low vagility observed for most species of Carabus. Consequently, the Solier’s Carab is genetically structured on an extremely fine scale, and local population within a definite forest should be considered as management units. In the light of our data on C. solieri we discuss the usefulness, when hybridisation occurs, of the two main criteria usually quoted for identifying candidate populations for conservation management (uniqueness and diversity) and give some conservation recommendations for C. solieri.

Refuge areas and suture zones in the Pyrenean and Cantabrian regions: geographic variation of the female MPI sex‐linked alleles among oviparous populations of the lizard Lacerta (Zootoca) vivipara

The viviparous lizard Lacerta (Zootoca) vivipara exhibits several alleles of the mannose‐6‐phosphate isomerase (MPI) enzyme that are carried exclusively on the female W sex‐chromosome. Previous studies showed that both the oviparous and viviparous forms of L. (Zootoca) vivipara have these female sex‐linked alleles. We document the existence of geographic variation of these alleles among the oviparous populations of southwestern France and northwestern Spain. Two oviparous subgroups were identified: all females from the eastern and central Pyrenees and most females from Aquitaine and from the northern slope of the western Pyrenees exhibited the fast migrating alleles MPI110 or MPI120, whereas all females from the Cantabric mountains, Spanish Basque country, and from the south slope of the western Pyrenees exhibited the slow migrating allele MPI90. Populations with both fast and slow migrating alleles occurred at some stations in the upper Ossau valley (western Pyrenees) and also at a lowland station of south Aquitaine. The hypothesis that several oviparous forms could have retreated to different places of the Pyreneo‐Iberian refugia during the Quaternary glaciations could explain the conservation or the evolution of the polymorphism of the MPI alleles, and that is consistent with the phylogeographic scenario previously proposed to account for the reproductive and cytogenetical variation observed in this species.