Did Polyommatus icarus (Lepidoptera: Lycaenidae) have distinct glacial refugia in southern Europe? Evidence from population genetics

Pleistocene climatic oscillations strongly influenced the genetic composition of many species which are often divided into several genetic lineages. In this context, we studied the allozymes of a common and widely distributed butterfly, the common blue Polyommatus icarus, over a large part of Europe. The species had a rather high genetic diversity within populations with a strikingly high mean number of alleles per locus (2.98). In contrast, differentiation between populations was very low ( F _ST: 0.0187). Only a marginal trend of decline in genetic diversity from the south to the north was observed. Isolation-by-distance existed on a European scale ( r =  0.826), but not at a regional level. Regional differentiation between populations in western Germany was extremely low ( F _ST: 0.0041). It is probable that P. icarus was widely distributed in the Mediterranean region during the last ice age and expanded into central Europe in the postglacial period without major genetic erosion. Moderate present and past gene flow in an intact metapopulation structure may have occurred on local, regional and perhaps even continental scales.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 529–538.     

Regeneration dynamics of Tsuga canadensis in the southern Appalachian Mountains, USA

Tsuga canadensis is often considered a foundation species because of its influence on the biological and ecological processes of upland forest systems. With hemlock woolly adelgid (Adelges tsugae; HWA)-induced mortality of T. canadensis in southern Appalachian upland forests, there is a crucial need to understand the regeneration dynamics of this tree species. This research was conducted within mature T. canadensis forests of Great Smoky Mountains National Park (GSMNP) prior to widespread HWA infestation. Tsuga canadensis regeneration abundance is lower than has been reported in more northern locales, but varies widely among sites in the southern Appalachian Mountains. Ordination results suggest landscape-scale patterns of regeneration abundance that are influenced by gradients in understory Rhododendron maximum cover, soil sand and nutrient content. Regression models reinforce the observed gradients in regeneration abundance. Sites with abundant T. canadensis regeneration have less R. maximum cover, soil sand and litter depth, but more soil nutrient availability. Tsuga canadensis seedlings and saplings tend to be spatially aggregated at the 14 × 21 m plot scale. Sites with spatially aggregated seedlings and saplings have greater canopy openness and more species with gap affinity. This research elucidates associations between successful T. canadensis regeneration and site factors such as soil nutrient availability and canopy dynamics. More forest surveys are required to understand the conditions necessary for successful T. canadensis regeneration. Land managers involved in forest restoration efforts may have to utilize appropriate silvicultural techniques conjoined with R. maximum removal operations to regenerate T. canadensis in southern Appalachian upland forests.     

Evidence of a phylogeographic break in the Romanian brown bear (Ursus arctos) population from the Carpathians

We analysed 33 brown bears from the Romanian Carpathians and the Italian Apennines at sequences of the mitochondrial control region and nine polymorphic microsatellite loci with regard to genetic variability and haplotype distribution. The Italian brown bears were monomorphic for mtDNA sequences. The Romanian bears yielded the highest variability found so far in this species. Haplotypes of both previously identified mtDNA lineages (western and eastern) were found in Romania. In the eastern part of the Carpathians western and eastern haplotypes occurred sympatrically, the bears from the western part of the mountain range only exhibited western-type sequences. This pattern provides evidence of a mitochondrial phylogeographic break in the distribution of the eastern lineage within the Romanian Carpathians. Conservation implications of this finding are discussed.     

A new apameine genus and species from the southern Appalachian Mountains,USA (Lepidoptera,Noctuidae, Noctuinae)

Cherokeeagen. n. is proposed for a rarely collected apameine moth species from the southern Appalachian Mountains, C. attakullakullasp. n. It is recorded from foothills and lower montane habitats of North Carolina and Georgia where hill cane, Arundinaria appalachiana Triplett, Weakley & L.G. Clark is found. Adults and their genitalia are figured and a mtDNA barcode sequence is given.     

Phylogeny of the millipede genus Sphaeriodesmus Peters, 1864 (Polydesmida: Sphaeriodesmidae) based on morphological characters

In order to understand the evolutionary relationships among the species encompassed within the genus Sphaeriodesmus Peters, 1864, a cladistic analysis including 63 species was conducted. Ninety-five morphological characters were used for the phylogenetic reconstruction. The results suggested that the current composition of the genus Sphaeriodesmus does not circumscribe a monophyletic group; instead, the genera Eusphaeriodesmus, Colobodesmus, and Proeilodesmus are here synonymized under Sphaeriodesmus. Although raw morphological data had suggested the genus Lophocyclus as the sister taxon of Sphaeriodesmus, the phylogenetic analysis under implied weight identified the genus Cyphodesmus as the taxon most closely related to Sphaeriodesmus. Sphaeriodesmus isolatus Chamberlin, 1940 is a subjective synonym of Sphaeriodesmus conformans Chamberlin, 1925. The putative subdivisions previously proposed within Sphaeriodesmus do not hold as monophyletic either. Low stability was observed concerning the higher-level phylogenetic relationships of Sphaeriodesmus. Sphaeriodesmus crucis (Loomis, 1974), S. mecistonyx (Hoffman, 1990), and S. triramus (Kraus, 1954) are new combinations.     

Phylogeographic evidence for the postglacial colonization of the North and Baltic Sea coasts from inland glacial refugia by Triglochin maritima L.

We investigated the geographical distribution of genetic variation in 67 individuals of Triglochin maritima from 38 localities across Europe using AFLP markers. Analysis of genetic variation resulted in the recognition of two major genetic groups. Apart from few geographical outliers, these are distributed (1) along the Atlantic coasts of Portugal, Spain and France and (2) in the North Sea area, the Baltic Sea area, at central European inland localities, the northern Adriatic Sea coast and the Mediterranean coast of southwest France. Considering possible range shifts of T. maritima in reaction to Quaternary climatic changes as deduced from the present-day northern temperature limit of the species, Quaternary changes of coastline in the North Sea area and the very recent origin of the Baltic Sea, we conclude that the coastal populations of T. maritima in the North Sea and Baltic Sea areas originated from inland populations.     

Stand dynamics of a yellow-poplar (Liriodendron tulipifera L.) forest in the Appalachian Mountains, Virginia, USA

Yellow-poplar (Liriodendron tulipifera L.) is a large, rapidly growing, shade-intolerant tree species common after disturbances on moist sites in the Appalachian Mountains. The species is typically scattered throughout old-growth mesophytic forests, where periodic gap formation creates conditions favorable for yellow-poplar establishment and growth. On abandoned agricultural fields, however, it is common for nearly monospecific forests of yellow-poplar to develop.This study examines stand dynamics of a yellow-poplar forest in western Virginia, USA that was established on agricultural fields abandoned in the late 1940s. Increment cores were collected from yellow-poplar trees growing on exposed ridgetops and in a more sheltered hollow. Tree-ring data show that the forest is even-aged. Tree establishment began about 5 years earlier on the ridgetops than in the hollow. Major ice storms disturbed the forest in 1978 and 1994, with two separate events in 1994. Ice storms disturb forests by depositing heavy loads of freezing rain on trees, breaking or uprooting them. The dendroecological data collected for this study provide little support for the hypothesis that ice storm disturbance promotes the establishment of new yellow-poplar cohorts. However, the data show that radial growth of some trees increased after ice storm disturbance, a pattern that reflects the increased availability of light following disturbance. Radial growth declined in some other trees as a consequence of severe injury during the storms.Radial growth responses following the 1978 ice storm were stronger on the ridgetops than in the hollow, suggesting that tree damage was more severe on the higher, more exposed sites. Growth responses were relatively mild following the storms of 1994, and did not exhibit pronounced topographic variations.     

Biotic and abiotic factors affecting the genetic structure and diversity of butternut in the southern Appalachian Mountains,USA

The abundance of butternut (Juglans cinerea L.) trees has severely declined rangewide over the past 50 years. An important factor in the decline is butternut canker, a disease caused by the fungus Ophiognomonia clavigigenti-juglandacearum, which has left the remaining butternuts isolated and sparsely distributed. To manage the remaining populations effectively, information regarding how butternut’s population genetic structure is affected by environmental and historical factors is needed. In this study, we assessed genetic structure and diversity of 161 butternut trees from 19 adjacent watersheds in the southern portion of butternut’s range using 12 microsatellite markers. We assessed the genetic diversity and genetic differentiation among trees grouped at various spatial scales. Our goal was to use historical abundance and land use data for these watersheds, which are now all a part of the Great Smoky Mountains National Park (GSMNP), to understand the ecological and evolutionary forces that challenge the conservation and management of butternut. In general, butternuts within the 19 neighboring watersheds were all part of one continuous population, with gene flow throughout. Significant genetic differentiation was detected between some groups of trees, but the differentiation was quite small and may not represent an ecologically significant distinction. The mean heterozygosity in all watersheds remained high, despite extensive mortality. Overall, genetic diversity and rare alleles were evenly distributed across all watersheds, with some variability in subpopulations containing butternut-Japanese walnut hybrids (Juglans x bixbyi or buarts). These results indicate that management of this species should focus on protection from future hybridization with Japanese walnut, promotion of regeneration, and persistence of all remaining butternut trees, which still retain high levels of genetic diversity.     

Taxonomy and species boundaries in the coral genus Favia Milne Edwards and Haime, 1857 (Cnidaria: Scleractinia) from Thailand revealed by morphological and genetic data

While Faviidae is a widely and uniformly distributed coral family throughout the Indo-Pacific, the extensive phenotypic plasticity of colony surface and corallite features often confounds the use of macromorphological characters in species identification, and contributes to conflict between traditional classification and molecular analyses of the group. Recent advances in morphological and molecular techniques now provide a suite of methods to re-address coral taxonomy in complex groups, such as that represented by the Faviidae. This study combines morphologic measurements including “3D coordinates landmarks” data with phylogenetic assessments of nuclear (ITS) and mitochondrial (COI-trnM) DNA to assess species boundaries in nine species of Faviidae with para-septothecal walls from Thailand. Strong concordance was found between morphological features and a priori groupings based on both morphospecies and genetically defined groups (ITS and COI-trnM). Favia truncatus was the most well-defined species based on morphological analyses, and it was also shown to be monophyletic using phylogenetic analyses. Besides F. truncatus, the only other species that was found to be monophyletic in analyses of both genes was F. cf. helianthoides, but its skeletal morphology overlapped with the F. favus species complex (comprised of F. favus, F. speciosa, F. matthaii and F. rotumana). Although not genetically monophyletic, the F. favus species complex and F. pallida were fairly well delineated morphologically. Morphospecies within the F. favus species complex are therefore possibly a result of genetic drift and/or stable polymorphisms driven by divergent selection. These results represent a first step toward a taxonomic revision of the Indo-Pacific Favia, which will integrate morphological methods with the study of type material, genetic information, reproductive data, and tests of phenotypic plasticity—given that multiple lines of evidence are needed to resolve ambiguous species and assign species names.     

Molecular genetic data reveal hybridization between Typha angustifolia and Typha latifolia across a broad spatial scale in eastern North America

A recent increase in the abundance of cattails (Typha spp.) in North American wetlands has been anecdotally linked with hybridization between Typha latifolia and Typha angustifolia. In this study, we used molecular genetic markers (microsatellites) to investigate whether the hybrid lineage (Typha × glauca) is restricted to The Great Lakes region, or exists across a much broader spatial scale. We also investigated the possibility of backcrossing and genetic introgression in natural populations. Parental species could be distinguished from one another based on the distribution of alleles at six microsatellite loci. Species identification based on genetic data corresponded well with species identifications based on leaf width, a key morphological trait that can distinguish the two parental species. We found that hybrids occur in Ontario, Quebec, New Brunswick, and Nova Scotia, but we did not detect hybrids in Maine. F₁s are more abundant than backcrossed or intercrossed hybrids, although we also found evidence of backcrossing, particularly in Ontario. This indicates that hybrids are fertile, and are therefore potential conduits of gene flow between the parental species. Further work is needed to determine whether T. × glauca is particularly successful in the Great Lakes region relative to other areas in which the two parental species co-exist, and to assess whether introgression may lead to increased invasiveness in the species complex.     

Gradual decline in genetic diversity in Swiss stone pine populations (Pinus cembra) across Switzerland suggests postglacial re-colonization into the Alps from a common eastern glacial refugium

Molecular genetic markers may reveal informative patterns of population processes such as historical migration, which may substantiate inference on postglacial re-colonization inferred, e.g., from fossil records. Palynological records of Swiss stone pine (Pinus cembra) suggest that the species has re-colonized the central Alps from a southeastern Alpine refugium after the last glacial maximum. Such a migration pathway likely resulted in a gradual decrease in genetic diversity with increasing distance to the glacial refugium, owing to founder events at the leading range edge. The present distribution of P. cembra in Switzerland consists of two rather distinct ranges, namely the inner-alpine parts of the Grisons and Valais, respectively, and additional disjunct occurrences in the northern and southern periphery of the Alps as well as between the two main ranges. We screened chloroplast microsatellite loci on 39 Swiss P. cembra populations and show that the genetic structure detected was congruent with a common ancestry from a single glacial refugium, likely located at the (south-)eastern periphery of the Alps. In contrast, our data rejected the alternative hypothesis of a distinct genetic separation of the two main ranges of Swiss stone pine in Switzerland. We further show that low genetic diversity within and high differentiation among peripheral populations in the northern Alps as well as the genetic differentiation between core and peripheral populations reflect genetic drift as a consequence of colonization history and limited gene flow by pollen and seed.

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Zusammenfassung  Molekulargenetische Marker enthalten wertvolle Information über Populationsprozesse wie historische Wanderungen, wodurch Annahmen zur postglazialen Wiederbesiedelung, beispielsweise abgeleitet von Fossilfunden, unterstützt werden k?nnen. Palynologische Funde von Arve (Pinus cembra) lassen vermuten, dass diese Art nach dem letzten glazialen Maximum von einem Refugium in den süd?stlichen Alpen wieder in die zentralen Alpen eingewandert ist. Ein solcher Rückwanderungsweg dürfte aufgrund von Gründereffekten an der Wanderungsfront eine graduelle Verringerung der genetischen Variation mit zunehmender Distanz zum Glazialrefugium bewirkt haben. Die heutige Verbreitung von P. cembra in der Schweiz weist zwei deutlich getrennte Gebiete auf, n?mlich inneralpine Bereiche der Kantone Graubünden und Wallis, mit jeweils isolierten Vorkommen in den n?rdlichen und südlichen Randalpen. Wir untersuchten Chloroplasten-Mikrosatelliten in 39 Schweizer Populationen von P. cembra und zeigen, dass die gefundene genetische Struktur übereinstimmt mit der Annahme einer gemeinsamen Abstammung aus einem einzigen Glazialrefugium, welches vermutlich am (süd-)?stlichen Rand der Alpen lag. Im Gegensatz dazu widerlegen unsere Daten die alternative Hypothese einer deutlichen genetischen Trennung der zwei Hauptvorkommen der Arve in der Schweiz. Im Weiteren zeigen unsere Resultate eine geringe genetische Variation innerhalb und einen hohen Differenzierungsgrad zwischen n?rdlichen Randalpenvorkommen, sowie eine genetische Differenzierung zwischen zentralen und peripheren Populationen. Dies weist auf genetische Drift hin, welche die Besiedlungsgeschichte und einen beschr?nkten Genfluss durch Pollen und Samen widerspiegelt.

     

Late Pleistocene freshwater fish (Cottidae) trackways from New England (USA) glacial lakes and a reinterpretation of the ichnogenus Broomichnium Kuhn

New trace fossil material from Late Pleistocene glaciolacustrine varves of the Connecticut River Valley, New England, USA represent the first evidence of freshwater sculpin in glacial Lake Hitchcock. Paleobiogeographic data constrain the timing of the last reinhabitation of freshwater cottids from a Wisconsinan glacial refugium. Freshwater sculpin were present in the area of study by ~ 13.7 ky BP, moving approximately 400 km in 5000 yr, and following the ice margin at distances as close as 35 km. The trace fossils warrant erection of a new ichnospecies, Broomichnium flirii isp. nov. Comparison of this new ichnospecies to Broomichnium permianum reveals distinct similarities, and it is possible that the Permian examples of B. permianum also were made by fish, which would reconcile a long-running controversy. Many groups of fish are demersal and make ventral body contact with the substrate and could potentially leave similar traces. Identifying new forms of trace fossils made by fish that use alternative modes of locomotion will prove useful in paleoenvironmental interpretations.     

Diversification across an altitudinal gradient in the Tiny Greenbul (Phyllastrephus debilis) from the Eastern Arc Mountains of Africa

Background  

The Eastern Arc Mountains of Africa have become one of the focal systems with which to explore the patterns and mechanisms of diversification among montane species and populations. One unresolved question is the extent to which populations inhabiting montane forest interact with those of adjacent lowland forest abutting the coast of eastern Africa. The Tiny Greenbul (Phyllastephus debilis) represents the only described bird species within the Eastern Arc/coastal forest mosaic, which is polytypic across an altitudinal gradient: the subspecies albigula (green head) is distributed in the montane Usambara and Nguru Mountains whereas the subspecies rabai (grey head) is found in Tanzanian lowland and foothill forest. Using a combination of morphological and genetic data, we aim to establish if the pattern of morphological differentiation in the Tiny Greenbul (Phyllastrephus debilis) is the result of disruptive selection along an altitudinal gradient or a consequence of secondary contact following population expansion of two differentiated lineages.     

Patchy population structure in a short-distance migrant: evidence from genetic and demographic data

Species often occur in subdivided populations as a consequence of spatial heterogeneity of the habitat. To describe the spatial organization of subpopulations, existing theory proposes three main population models: patchy population, metapopulation and isolated populations. These models differ in their predicted levels of connectivity among subpopulations, and in the risk that a subpopulation will go extinct. However, spatially discrete subpopulations are commonly considered to be organized as metapopulations, even though explicit tests of metapopulation assumptions are rare. Here, we test predictions of the three models on the basis of demographic and genetic data, a combined approach so far surprisingly little used in mobile organisms. From 2002 to 2005, we studied nine subpopulations of the wetland-restricted reed bunting ( Emberiza schoeniclus ) in the southeastern part of the Canton Zurich (Switzerland), from which local declines of this species have been reported. Here, wetlands are as small as 2.7 ha and separated through intensively used agricultural landscapes. Demographic data consisted of dispersal of colour-banded individuals among subpopulations, immigration rates and extinction-/recolonization dynamics. Genetic data were based on the distribution of genetic variability and gene flow among subpopulations derived from the analysis of nine microsatellite loci. Both demographic and genetic data revealed that the patchy population model best described the spatial organization of reed bunting subpopulations. High levels of dispersal among subpopulations, high immigration into the patchy population, and genetic admixture suggested little risk of extinction of both subpopulations and the entire patchy population. This study exemplifies the idea that spatially discrete subpopulations may be organized in ways other than a metapopulation, and hence has implications for the conservation of subpopulations and species.     

Phylogeography of the burnet moth Zygaena transalpina complex: molecular and morphometric differentiation suggests glacial refugia in Southern France,Western France and micro‐refugia within the Alps

Patterns of common recolonization routes from glacial refugia to Central Europe during the Pleistocene are generalized to paradigms of postglacial recolonization in Europe. Recent studies indicate, however, that the actual phylogeographic history of many species might be more complex and cannot be simplified to generalized patterns. Burnet moths of the Zygaena transalpina complex represent a group of closely related taxa, which are considered as a typical example for these generalized patterns. At present, three groups are recognized that are assumed to have spread from three classical refugia in Western Europe, Italy and the Balkans to Central Europe. Here, we re‐investigate their phylogeography using a combined molecular and morphometric approach. Phylogenetic and nested clade phylogeographic analyses of 476 samples from 55 localities taken from Southern and Central Europe reveal that the Zygaena transalpina complex consists of three distinct haplotype clusters, which geographically roughly correspond to possible refugia in Western Europe, Italy and the Balkans. A synthesis of the data with a geometric morphometry dataset of 425 specimens from 46 localities corroborates this molecular result but differs in several aspects. Important new aspects are multiple refugia of the western ‘hippocrepidis’ branch and micro‐habitats within the Alps of the central ‘transalpina’ branch. Further, our results display a more complex phylogeographic pattern for this species complex, which is not tractable with a rigid, generalized pattern.     

Morphologic variability of the coccolithophorid Calcidiscus leptoporus in the plankton, surface sediments and from the Early Pleistocene

On a global scale, morphological variability of the extant coccolithophorid Calcidiscus leptoporus (Murray and Blackman, 1898) Loeblich and Tappan was investigated in surface sediments and plankton samples and from an Early Pleistocene time-slice (1.8 Ma to 1.6 Ma). In the bivariate space coccolith diameter versus number of rays in the distal shield, Holocene samples follow a single, unimodal morphocline. Sample means of coccolith size and number of elements group in three clusters, I, II and III, which are of biogeographic significance. Clusters II and III coccoliths (mean coccolith size of 5.0 μm and 20.9 elements, and 6.6 μm and 25.6 elements, respectively) are found in a tropical belt extending from 11 °N to 17 °S with an annual minimum sea-surface temperature above 23.5 °C. Cluster I coccoliths (5.8 μm, 20.7 elements) are found in samples outside that belt. The distribution of coccoliths in the surface sediments is tentatively interpreted to be a result of mixing to a varying degree of at least three different morphotypes (‘small’, ‘intermediate’ and ‘large’), which were identified in the living plankton, and which are separated from each other at 5 μm and 8 μm mean coccolith diameter, respectively. A comparison of the surface sediments with the Early Pleistocene assemblages revealed that between 1.6 Ma and 1.8 Ma two morphoclines A and B existed, the first of which persisted until the Holocene in the form of C. leptoporus, while the second comprises only extinct morphotypes including Calcidiscus macintyrei as one end-member. During the Early Pleistocene morphocline A was more homogeneous and no clusters were evident.Morphocline B shows a clear bimodality with a separation of morphotypes at 9.5 μm. Our observations suggest that morphoclines are subsets within the total stratigraphical range of a single species, and represent the global variability of that species in a particular time interval. Morphotypes, which belong to a morphocline, represent the infra-specific variability of that species within the biogeographic and stratigraphic limits of that species.