Conservation of genetic uniqueness in remaining populations of red squirrels (Sciurus vulgaris L.) in the South of England

The Eurasian red squirrel (Sciurus vulgaris) is an emblematic species for conservation, and its decline in the British Isles exemplifies the impact that alien introductions can have on native ecosystems. Indeed, red squirrels in this region have declined dramatically over the last 60 years due to the spread of squirrelpox virus following the introduction of the gray squirrel (Sciurus carolinensis). Currently, red squirrel populations in Britain are fragmented and need to be closely monitored in order to assess their viability and the effectiveness of conservation efforts. The situation is even more dramatic in the South of England, where S. vulgaris survives only on islands (Brownsea Island, Furzey Island, and the Isle of Wight). Using the D‐loop, we investigated the genetic diversity and putative ancestry of the squirrels from Southern England and compared them to a European dataset composed of 1,016 samples from 54 populations. We found that our three populations were more closely related to other squirrels from the British Isles than squirrels from Europe, showed low genetic diversity, and also harbored several private haplotypes. Our study demonstrates how genetically unique the Southern English populations are in comparison with squirrels from the continental European range. We report the presence of four private haplotypes, suggesting that these populations may potentially harbor distinct genetic lineages. Our results emphasize the importance of preserving these isolated red squirrel populations for the conservation of the species.     

Infestation of a novel host plant by <Emphasis Type="Italic">Tephritis conura</Emphasis> (Diptera: Tephritidae) in northern Britain: host-range expansion or host shift?

The addition of a novel host plant to a phytophagous insect’s diet may result in subsequent host-plant specialisation, and is believed to be a key cause for speciation in this trophic group. In northern Britain, the tephritid fly Tephritis conura has experienced a unique host-plant expansion, from the melancholy thistle Cirsium heterophyllum to the marsh thistle C. palustre. Here, we examine whether the incorporation of C. palustre in the repertoire of British T. conura flies has caused genetic divergence between populations infesting the old host and the novel host, and how British populations differ from populations infesting C. heterophyllum in continental Europe where C. palustre is not infested. No evidence for restricted gene flow among British C. palustre and C. heterophyllum flies was found. Significant differentiation between British and continental T. conura was found at only one allozyme locus, hexokinase, and caused by a new allele, Hex_95. Hexokinase is related to host-race formation in continental European flies infesting C. heterophyllum and C. oleraceum, and might be linked to loci determining host choice. Based on morphological and phenological data from previous studies, we suggest that T. conura in Britain has adapted to the novel host but that host-race formation is impeded by similar plant phenologies.     

Population genetic structure of Bombus terrestris in Europe: Isolation and genetic differentiation of Irish and British populations

The genetic structure of the earth bumblebee (Bombus terrestris L.) was examined across 22 wild populations and two commercially reared populations using eight microsatellite loci and two mitochondrial genes. Our study included wild bumblebee samples from six populations in Ireland, one from the Isle of Man, four from Britain and 11 from mainland Europe. A further sample was acquired from New Zealand. Observed levels of genetic variability and heterozygosity were low in Ireland and the Isle of Man, but relatively high in continental Europe and among commercial populations. Estimates of F_st revealed significant genetic differentiation among populations. Bayesian cluster analysis indicated that Irish populations were highly differentiated from British and continental populations, the latter two showing higher levels of admixture. The data suggest that the Irish Sea and prevailing south westerly winds act as a considerable geographical barrier to gene flow between populations in Ireland and Britain; however, some immigration from the Isle of Man to Ireland was detected. The results are discussed in the context of the recent commercialization of bumblebees for the European horticultural industry.     

Origin of, and conservation units in, the Irish red squirrel (Sciurus vulgaris) population

Knowledge of genetic relationships among wildlife populations is fundamental to their conservation, particularly where translocations are concerned. This study involved a survey of mitochondrial DNA variation in the Irish red squirrel population. Our main aims were: (1) to determine whether the Irish red squirrel population is distinct from that found in Britain, given known translocations that took place from Britain in the 1800’s; and (2) whether inclusion of Irish data into a reanalysis of European red squirrel data could reveal patterns of postglacial spread in Ireland. We found evidence that the current Irish red squirrel population may be a mixture of native and translocated stock, and relationships between Irish and European haplotypes supported a number of colonisation events of the island. Although only one haplotype was common to both Ireland and Britain, it is probable that the most common haplotypes in Ireland are British introductions that have since become extinct in Britain. There was a significant regional genetic structure in Ireland (P < 0.001), as well as between all Irish and British regions. Although it is likely that the red squirrel will not be fundamental in tracing the colonisation of Ireland by mammals, the data demonstrated that individual regions within Ireland, as well as the Irish population as a whole, are distinct both from the British population and from each other and, therefore, these populations should be treated as separate Management Units (MU) in conservation strategies.     

Palatability to a generalist herbivore,defence and growth of invasive and native <Emphasis Type="Italic">Senecio</Emphasis> species: testing the evolution of increased competitive ability hypothesis

This paper tests the prediction that introduced plants may become successful invaders because they experience evolutionary changes in growth and defence in their new range [evolution of increased competitive ability hypothesis (EICA)]. Interspecific and intraspecific binary feeding choices were offered to the snail Helix aspersa. The choices were between: (1) plants of the invasive Senecio inaequidens and Senecio pterophorus derived from populations in the introduced range (Europe) and plants of three indigenous species (Senecio jacobea, Senecio vulgaris and Senecio malacitanus) from populations in Europe; (2) plants of the invasive S. inaequidens and S. pterophorus from populations in the introduced range (Europe) and from populations in the native range (South Africa). We did not find a clear pattern of preference for indigenous or alien species of Senecio. However, we found that European invasive populations of S. inaequidens and S. pterophorus were less palatable than South African native populations. Moreover, in contrast to the predictions of the EICA hypothesis, the invasive genotypes of both species also showed a higher total concentration of pyrrolizidine alkaloids, and in the case of S. inaequidens we also found higher growth than in native genotypes. Our results are discussed with respect to the refinement of the EICA hypothesis that takes into account the difference between specialist and generalist herbivores and between qualitative and quantitative defences. We conclude that invasive populations of S. inaequidens and S. pterophorus are less palatable than native populations, suggesting that genetic differentiation associated with founding may occur and contribute to the plants’ invasion success by selecting the best-defended genotypes in the introduced range.     

Genetic structure of Eurasian badgers Meles meles (Carnivora: Mustelidae) and the colonization history of Ireland

The present study examined the contemporary genetic composition of the Eurasian badger, Meles meles, in Ireland, Britain and Western Europe, using six nuclear microsatellite loci and a 215‐bp fragment of the mitochondrial DNA control region. Significant population structure was evident within Europe (global multilocus microsatellite F_ST = 0.205, P < 0.001; global mitochondrial control region Φ_ST = 0.399, P < 0.001). Microsatellite‐based cluster analyses detected one population in Ireland, whereas badgers from Britain could be subdivided into several populations. Excluding the island populations of Ireland and Britain, badgers from Western Europe showed further structuring, with evidence of discrete Scandinavian, Central European, and Spanish populations. Mitochondrial DNA cluster analysis grouped the Irish population with Scandinavia and Spain, whereas the majority of British haplotypes grouped with those from Central Europe. The findings of the present study suggest that British and Irish badger populations colonized from different refugial areas, or that there were different waves of colonization from the source population. There are indications for the presence of an Atlantic fringe element, which has been seen in other Irish species. We discuss the results in light of the controversy about natural versus human‐mediated introductions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Harmonia axyridis in Great Britain: analysis of the spread and distribution of a non-native coccinellid

Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe H. axyridis is considered to be an invasive alien species. Although not known to have been deliberately introduced to Great Britain, it was first recorded there in 2004, in south-east England. Harmonia axyridis arrived in Great Britain by various means, primarily from mainland Europe, but also from Canada. Extensive national and local media coverage, and a dedicated website (), facilitated public involvement in recording H. axyridis in Great Britain; in excess of 4,000 verified records of the species were received between 2004 and 2006. Through detailed mapping, the objective of our study was to quantify and analyse the spread of H. axyridis in its early stages of invasion in Great Britain. Our data shows that between 2004 and 2006, the species spread north through Great Britain at the rate of 58 km year^-1 and west at the rate of 144.5 km year^-1. In England H. axyridis spread north to Yorkshire and west to Cornwall by the end of 2006, and also reached Wales. Recorded occurrence (of one or more H. axyridis individuals at larval, pupal and/or adult stage) in 10 km squares in Great Britain was: 2004—51; 2005—149; 2006—447. Records of juvenile H. axyridis extend through most of the recorded British range of the species, and we present evidence of bi-voltinism in the population in 2006.     

Population Genetics and Identity of an Introduced Terrestrial Slug: Arion subfuscus s.l. in the North-east USA (Gastropoda, Pulmonata, Arionidae)

Several European species of the terrestrial slug genus Arion have been introduced into North America. A case in point is the species complex A. subfuscus s.l. which has become one of the most abundant slug taxa in North America. In Europe this complex consists of at least two cryptic species, viz. A. fuscus and A. subfuscus s.s., the latter of which is further subdivided in five strongly divergent mtDNA lineages (A. subfuscus S1–S5). In order to determine which of these A. subfsucus s.l. taxa are present in the NE USA and in order to assess their population genetic structure, we compared mtDNA, nDNA and allozyme variation between populations from the NE USA and Europe. Our results show that (1) at least A. subfuscus S1 has become successfully established in the NE USA, (2) founder effects are the most likely explanation for the loss of a large amount of molecular genetic variation in populations from the NE USA (i.e. a loss of 96% of the 16S rDNA haplotypes, 67% of the ITS1 alleles and 46% of the alleles at polymorphic allozyme loci), and (3) part of the remaining genetic variation in NE USA populations was probably due to multiple introductions from the British Isles and the European mainland, and the hybrid structure of most of these source populations. Apparently, the extreme loss of molecular genetic variation in this introduced species has not prevented it from successfully establishing and spreading in novel environments.     

Phylogeography and genetic diversity of the red squirrel (Sciurus vulgaris) in China: Implications for the species' postglacial expansion history

The geographical distribution of Sciurus vulgaris spans much of the Palearctic from western Europe and the UK eastward to the pacific coast of East Asia. S. vulgaris occurs in China only in the far northwest and the northeast. Understanding of the species’ postglacial expansion history in East Asia has been limited by the paucity of molecular data. In this study, we used partial D-loop and cytochrome b gene sequences to assess mitochondrial DNA variation in S. vulgaris in China. Our objectives were to (1) determine phylogeographical patterns of S. vulgaris in China; (2) understand the species’ postglacial expansion history in this region; and (3) quantify genetic diversity levels within S. vulgaris populations in China. We identified a supported phylogenetic group in S. vulgaris from China, and found no tendency for haplotypes to cluster by geographic region. Our analysis of S. vulgaris from China and other regions supports the hypothesis that the Calabria region of southern Italy is a glacial refugium for the species. We tentatively propose a postglacial expansion pattern for the squirrels: migrating from Calabria via Central and Eastern Europe to Russia and from there to China, and firstly to the northwest and then to northeast in China. We found high levels of genetic diversity in S. vulgaris populations across China as a whole, and discussed its influential factors.     

Chloroplast DNA and isozyme evidence on the evolution ofSenecio vulgaris (Asteraceae)

Chloroplast DNA (cpDNA) and isozyme variation were analyzed over a range of populations of two infraspecific taxa of the tetraploidSenecio vulgaris. The isozyme data were supportive of the hypothesis that the weedy and cosmopolitanS. vulgaris var.vulgaris is an evolutionary derivative ofS. vulgaris subsp.denticulatus from the coasts of W Europe and montane altitudes in S Spain and Sicily. The two taxa exhibited a very high genetic identity with subsp.denticulatus containing slightly more isozyme diversity than was found in var.vulgaris. — Three cpDNA haplotypes (A, B, C) already known from other Mediterranean diploid species ofSenecio were resolved in var.vulgaris, and an additional fourth haplotype (E) was found in subsp.denticulatus. Two alternative hypotheses were chosen to account for the origin and maintenance of the observed cpDNA composition ofS. vulgaris. It either reflects (1) the retention of an ancestral polymorphism which stems from the recurrent and polytopic formation of ancestral tetraploid lineages; or (2)S. vulgaris originally was characterized by haplotype E, and haplotypes A, B and C were acquired through repeated introgressive hybridization with related diploid species. The finding that very low levels of nuclear (isozyme) diversity were present in both taxa ofS. vulgaris examined supports the second of these two hypotheses; however, more detailed analysis of nuclear genetic diversity is required before a firm conclusion can be reached on this matter.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Phylogeographic past and invasive presence of Arion pest slugs in Europe

Arion vulgaris (syn. A. lusitanicus) is the most destructive pest slug in Europe. The species has been regarded a classic case of an ongoing biological invasion with negative economic and ecological impact in many European countries, but this status has recently been contested. In this study, we assessed mitochondrial and autosomal genetic diversity in populations of A. vulgaris across the entire distribution range in order to characterize its evolutionary history. Mitochondrial diversity in A. vulgaris was strongly reduced compared with the closely related and largely codistributed noninvasive species A. rufus and A. ater, indicating a very rapid spread of A. vulgaris through Europe. Autosomal diversity assessed in 632 individuals from 32 populations decreased towards eastern and northern Europe which is consistent with the reported expansion of the species towards these regions in the last decades. Demographic simulations supported very recent population founding events in most of the European range. The short periods between the first detection of A. vulgaris in different countries and only a very weak association of genetic structuring among populations with geographical distances suggest a human contribution in the ongoing expansion of the slug. We propose that this contribution may ultimately prevent the exact localization of the debated region of origin of A. vulgaris. However, the reclassification of A. vulgaris as noninvasive would be premature. Without counter measures, the Eastern and Northern European countries can expect to see this biological invasion continued in the future.     

Interactions between the parasitoid wasp Dinocampus coccinellae and two species of coccinellid from Japan and Britain

The establishment, spread and increase of the invasive coccinellid Harmonia axyridis Pallas in North America and Britain have coincided with declines in native ladybirds. In Britain, this pattern was predicted soon after H. axyridis was first recorded. However, predictions of the impact that H. axyridis may have on the parasites, parasitoids and pathogens of native coccinellids have been less certain, largely because of uncertainty over whether H. axyridis would become an alternate host for such agents. Dinocampus coccinellae (Schrank) is a braconid parasitoid of ladybirds of the sub-family Coccinellinae. In Japan, this wasp uses both Coccinella septempunctata brucki Mulsant and H. axyridis as hosts, but successfully parasitizes a higher proportion of the former species. Data are here presented that indicate the same is true in Britain, but to a greater extent. This study investigates the interactions of D. coccinellae with C. septempunctata L. and H. axyridis from Japan and Britain. We show that coccinellid activity affects encounter rates with the wasp, with H. axyridis being more active than C. septempunctata in the tests. Escalation rates from investigation to ovipositional attack were higher for C. septempunctata than H. axyridis for both Japanese and British wasps. Wasp emergence was higher, relative to ovipositional attack, from C. septempunctata than from H. axyridis. Wasps did not discriminate between Japanese and British ladybirds of the same species. British wasp eclosion rates were higher from Japanese than from British H. axyridis. We consider both the effect D. coccinellae may have on H. axyridis population demography, and the effect H. axyridis may have on D. coccinellae in Europe.     

Evolution of an invasive phenotype: shift to belowground dominance and enhanced competitive ability in the introduced range

In response to novel selection pressures in an introduced range, non-native species may evolve more competitive phenotypes unique from those of their native range. We examined the existence of an invasive phenotype in the herbaceous perennial Artemisia vulgaris, a frequent invader of the Northeast and Mid-Atlantic US. Populations from both the native (European) and the introduced (North American) ranges were grown in intra-specific competition (same population), inter-specific competition with the native perennial herb Solidago canadensis, and alone in a common garden to quantify shifts in resource allocation and neighbor effects on performance and competitive ability. Without competition, introduced A. vulgaris populations were much shorter than native populations, but germinated earlier, produced more ramets, more belowground and total biomass, and maintained higher root-to-shoot ratios. Under inter- and intra-specific competitions, introduced A. vulgaris populations were shorter, but produced more ramets, belowground, and total biomass than native populations. S. canadensis belowground and total biomass were more highly suppressed by introduced than native A. vulgaris. Our data suggest that since the introduction to North America, A. vulgaris has evolved a more competitive invasive phenotype characterized by many short ramets with more extensive root/rhizome networks. This rapid evolutionary shift likely benefits A. vulgaris in its introduced range by allowing establishment and subsequent dominance in dense stands of existing vegetation.     

Population-specific migratory strategies of Twite Linaria flavirostris in Western Europe

Migratory connectivity describes the extent to which different individuals from specific breeding populations use similar non-breeding sites, with important consequences for the demography and population structure of birds. We used mark-recapture data from the EUring data bank to describe the non-breeding range of Twite Linaria flavirostris within Western Europe. We quantified the strength of migratory connectivity across all European breeding populations, and for the British and continental breeding populations separately. Twite from different parts of the British breeding range used different non-breeding areas from one another, whereas Twite from different parts of the continental breeding range used similar non-breeding areas. There was essentially no overlap during the non-breeding season between British and continental Twite.     

Morphology,geographical variation and the subspecies of Marsh Tit Poecile palustris in Britain and central Europe

Capsule All British Marsh Tits belong to subspecies Poecile palustris dresseri, being smaller than nominate P. p. palustris of central Europe.

Aims Determining the subspecies of Marsh Tit in Britain to test whether ssp. P. p. palustris occurs in northern England and Scotland, by assessing regional variation in size compared with central European birds.

Methods 1147 wing length and 250 tail length measurements from 953 Marsh Tits were compared between eight British locations to test for regional variation. Biometrics were compared between birds from Britain and six locations within the continental European range of ssp. palustris.

Results There was no regional variation in wing or tail lengths among British Marsh Tits, indicating that all resident birds belong to ssp. dresseri. There was no evidence supporting the existence of ssp. palustris in northern England. British birds were significantly smaller than those from continental Europe, with proportionately shorter tails, consistent across all age and sex classes.

Conclusion All British Marsh Tits should be considered as ssp. dresseri, with ssp. palustris being limited to continental Europe. With no evidence of regional variation in size within Britain, reliable sexing methods based on biometrics could be applied in demographic studies throughout the country.     

Recent hybrid origin and invasion of the British Isles by a self-incompatible species, Oxford ragwort (Senecio squalidus L., Asteraceae)

Senecio squalidus is a diploid hybrid species which originated in the British Isles following the introduction of material collected from a hybrid zone on Mount Etna, Sicily, approximately 300 years ago. Introduced hybrid material was cultivated in the Oxford Botanic Garden and gave rise to the stabilized diploid hybrid species, which later spread throughout much of the UK and into some parts of Ireland. Unusually for an invasive species, S. squalidus has a strong system of sporophytic self-incompatibility (SSI) that may have become modified as a result of its recent hybrid origin and spread. First, S. squalidus contains relatively few S alleles (between 2 and 6 S alleles within individual UK populations) compared to other species with SSI (estimates average ~17 S alleles per population). This most probably reflects the population bottleneck experienced by introduced hybrid material. Second, dominance relationships among S. squalidus S alleles are more extensive than those reported in other species with SSI. Third, although pseudo-self-compatibility occurs sporadically in S. squalidus, it is not widespread, indicating that SSI is maintained in the species despite potential mate availability restrictions imposed by low numbers of S alleles. Surveys of other forms of genetic diversity in S. squalidus show that allozyme variation is reduced relative to that within the progenitor species, but Randomly Amplified Polymorphic DNA variation is relatively high. Both types of genetic variation show little or no pattern of isolation-by-distance between populations in keeping with the recent range expansion of the species. During its spread in the British Isles, S. squalidus has hybridized with the native self-compatible (SC) tetraploid species, S. vulgaris, which has led to the origin of three new SC hybrid taxa: a radiate form of S. vulgaris (var. hibernicus), a tetrapoid hybrid species (S. eboracensis) and an allohexaploid (S. cambrensis).     

Phylogenomic analysis of pseudocryptic diversity reveals the new genus Deltalsia (Rhodomelaceae,Rhodophyta)

Molecular analyses, in combination with morphological studies, provide invaluable tools for delineating red algal taxa. However, molecular datasets are incomplete and taxonomic revisions are often required once additional species or populations are sequenced. The small red alga Conferva parasitica was described from the British Isles in 1762 and then reported from other parts of Europe. Conferva parasitica was traditionally included in the genus Pterosiphonia (type species P. cloiophylla in Schmitz and Falkenberg 1897), based on its morphological characters, and later transferred to Symphyocladia and finally to Symphyocladiella using molecular data from an Iberian specimen. However, although morphological differences have been observed between specimens of Symphyocladiella parasitica from northern and southern Europe they have yet to be investigated in a phylogenetic context. In this study, we collected specimens from both regions, studied their morphology and analyzed rbcL and cox1 DNA sequences. We determined the phylogenetic position of a British specimen using a phylogenomic approach based on mitochondrial and plastid genomes. Northern and southern European populations attributed to S. parasitica represent different species. Symphyocladiella arecina sp. nov. is proposed for specimens from southern Europe, but British specimens were resolved as a distant sister lineage to the morphologically distinctive Amplisiphonia, so we propose the new genus Deltalsia for this species. Our study highlights the relevance of using materials collected close to the type localities for taxonomic reassessments, and showcases the utility of genome-based phylogenies for resolving classification issues in the red algae.     

Isozyme evidence and the origin ofSenecio vulgaris (Compositae)

An electrophoretic survey of isozyme variation was conducted to test the hypothesis thatSenecio vulgaris L. (2n = 40) is of autotetraploid origin fromS. vernalis Waldst. & Kit. (2n = 20). It was established thatS. vulgaris exhibited fixed heterozygosity at three loci examined, showed disomic inheritance at all polymorphic loci, and contained a gene (Est-1) and an allele (Aat-3b) which were not present in the single population ofS. vernalis surveyed. From this it is concluded thatS. vulgaris is not of autotetraploid origin. Instead, the genetic evidence is in keeping with an allopolyploid origin ofS. vulgaris with the possibility thatS. vernalis acted as one of its two parents.