Patterns of local and regional genetic structuring in the meadow grasshopper,Chorthippus parallelus (Orthoptera: Acrididae), in Central Germany revealed using microsatellite markers

The meadow grasshopper, Chorthippus parallelus (Zetterstedt), is common and widespread in Central Europe, with a low dispersal range per generation. A population study in Central Germany (Frankenwald and Thüringer Schiefergebirge) showed strong interpopulation differences in abundance and individual fitness. We examined genetic variability using microsatellite markers within and between 22 populations in a short‐ to long‐distance sampling (19 populations, Frankenwald, Schiefergebirge, as well as a southern transect), and in the Erzgebirge region (three populations), with the latter aiming to check for effects as a result of historical forest cover. Of the 671 C. parallelus captured, none was macropterous (functionally winged). All populations showed a high level of expected and observed heterozygosity (mean 0.80–0.90 and 0.60–0.75, respectively), whereas there was evidence of inbreeding (F_IS values all positive). Allelic richness for all locus–population combinations was high (mean 9.3–11.2), whereas alleles per locus ranged from 15–62. At a local level, genic and genotypic differences were significant. Pairwise F_ST values were in the range 0.00–0.04, indicating little interpopulation genetic differentiation. Similarly, the calculated gene flow was very high, based on the respective F_ST (19.5) and using private alleles (7.7). A Neighbour‐joining tree using Nei's D_A and principal coordinate analysis separated two populations that were collected in the Erzgebirge region. Populations from this region may have escaped the effects of the historical forest cover. The visualization of the spatial arrangement of genotypes revealed one geographical barrier to gene flow in the short‐distance sampling. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 875–890.     

Assortative mating across a hybrid zone in Chorthippus parallelus (Orthoptera: Acrididae)

Two subspecies of the grasshopper Chorthippus parallelus meet and form a hybrid zone in the Pyrenees. Both strong hybrid dysfunction and a behavioural difference occur, which would seem to make the zone a suitable candidate for speciation by reinforcement. One of the classic ways of looking for this is to test for increased levels of assortative mating between populations from close to the region of contact. Here we show that, with virgin insects, such assortment decreases as one approaches the centre of the zone in C. parallelus. The pattern is different upon remating, with non-virgins showing a pattern more like that predicted by reinforcement. Overall there is little evidence for reinforcement. We argue that other tests of the model may be more appropriate for stable hybrid zones.     

Being a generalist herbivore in a diverse world: how do diets from different grasslands influence food plant selection and fitness of the grasshopper Chorthippus parallelus?

1. Generalist insect herbivores occupy a variety of habitats that differ in food plant composition. Dietary mixing has been proposed as a possibility for generalists to overcome nutritional deficiencies of single plant species, but only a few studies have investigated herbivore feeding and fitness for diets that resemble natural scenarios. We studied feeding behaviour, survival, and reproduction of the generalist grasshopper Chorthippus parallelus raised on food plants of four typical habitats. 2. Grasshopper diet consisted of grasses (92.5%), legumes (6.7%) and, in small quantities, other forbs (0.8%). Diet selection differed between the four food plant mixtures, and depended on grasshopper sex and developmental stage. There was no correlation between the relative abundance of plant species in the field and the fraction of these species in the grasshopper diet. 3. Grasshoppers survived on average for 40.4 ± 1.0 days before maturity, grew 106.8 mg until maturity moult, and females laid 4.1 ± 0.4 egg pods, each of which contained 8.5 ± 0.4 eggs. However, despite the differences in feeding behaviour, grasshopper fitness was the same in all of the four food plant mixtures. While the digestibility of ingested food was similar in the four different treatments, indices indicated differences in the conversion efficiency to body mass. 4. Our results show that C. parallelus is a plastic feeder with no fixed preferences in diet composition. The results emphasise that generalist herbivores can counteract putative quality deficiencies of single food plants by selective dietary mixing.     

Ecological energetics of Central European grasshoppers (Orthoptera: Acrididae)

Summary Biomass and energy budgets and food utilization efficiencies of laboratory and wild populations of three Central European grasshopper species, Chorthippus parallelus (Zetterstedt), C. biguttulus (L.), and Gomphocerus rufus (L.), were studied between 1979 and 1984. Larval consumption is relatively low in C. biguttulus (C=4.3 kJ/ind.) compared with C. parallelus and G. rufus (4.9 kJ/ind.). In the adult phase (maturity) consumption rates of C. biguttulus and G. rufus (6.9 kJ/ind.) are similar, but higher in C. parallelus (7.3 kJ/ind.). The energy content decreases from the egg (23.2–24.3 J/mg dw) and body tissue (22.1–23.2 J/mg dw) to faeces (16.6–18.1 J/mg dw). The energy contents of faeces differ significantly between the species, indicating different rates of food conversion (on Dactylis glomerata). On average, the assimilation rates are about 30%, slightly lower in G. rufus. Approximate digestibility (A/C) ranges from 28.2 (G. rufus) to 35.7 (C. biguttulus) without great differences between larvae and adults. In contrast, the efficiency of conversion of ingested food (P/C) differs significantly between larval (10–11) and adult stages (3.4–3.6), and so to a much higher degree does the efficiency of conversion of digested food (P/A), from 30.3–33.5 in the larval to 9.5–14.9 in the adult period. Based on 5-year (C. parallelus) and 2-year studies (G. rufus) of the population dynamics and life tables, the energy budgets of wild populations were calculated and summarized into diagrams. Depending on the annually fluctuating densities, in both populations about 1%–2% of net primary production was consumed, and another 5%–9% was cut and dropped. The proportion of production of the grasshopper populations (body tissue) used for egg production differs in C. parallelus (28%) and G. rufus (44%). The energy cycling owing to energy storage in the living eggs amounts to 2%–3% of the total energy consumed by the population. The results are compared with the available data for grasshoppers in the literature.     

Chromosomal differentiation through an Alpine hybrid zone in the grasshopper Chorthippus parallelus

The grasshopper Chorthippus parallelus is genetically differentiated over its range into at least five major geographical subdivisions. Two of these subdivisions, designated as subspecies, meet and form a hybrid zone in the Pyrenees. These subspecies differ for a wide range of morphological, behavioural and chromosomal characters, which vary clinally across the zone. A further geographical subdivision exists within Italy. Here we present the first evidence for chromosomal divergence between populations of the grasshopper north and south of the Alps. Across two Alpine cols (Col de Larche, between France and Italy; Passo de Resia, between Austria and Italy), these populations differ in both structural and functional components of the X-chromosome. Northern Alpine individuals possess an active nucleolar organizing region (NOR) at the distal end of the X-chromosome and an associated region of heterochromatin (C-band). Both these features are absent from individuals from the south of the Alps. However, all individuals examined carry distally located rDNA on the X-chromosome. Clinal transition was examined in the distal C-band in transects through the two cols. The cline centres are roughly coincident with the tops of the cols. Both clines were of similar widths (Col de Larche, 21.88 km; Passo de Resia, 24.05 km), and therefore much wider than those for an X-linked distal C-band in the Pyrenean hybrid zone. This suggests that there are different selective pressures on the cytogenetic characters in the Alps. The results are discussed in the context of the historical population dynamics of the species in relation to the climatic changes associated with the Pleistocene ice ages.     

Spontaneous chromosome mutations in Truxaline grasshoppers

Three distinct mutant conditions are described in single male individuals from three species of short horn grasshopper. Of these, one is an entire germ line mutant of Myrmeleotettix maculatus, heterozygous for a centric fusion between single M₄ and M₅ telocentric chromosomes. In contrast, the remaining two mutants are present in mosaic form. One is heterozygous for an L₁-M₄ interchange in Omocestus viridulus, the other tetrasomic for the M₄ chromosome in Chorthippus parallelus which in addition is characterised by the inclusion of a supernumerary heterochromatic segment on one S₈ homologue. Centric fusion in Myrmehotettix maculatus has neither disturbed the chiasma potential of the elements constituting the fusion multiple nor, has it apparently influenced the production of balanced gametes. The pattern of chiasma formation in the L₁-M₄ interchange multiple lends support to the contention that the process of chiasma formation originates near the distal end of chromosome arms in Omocestus viridulus. There is no interaction between the two mutant conditions of tetrasomy and the presence of supernumerary segments in Chorthippus parallelus. Moreover, because of the precocious nature of two of the four M₄ homologues there is little tendency to form multivalents. The two M₄ bivalents share a similar mean chiasma frequency.     

Complementarity effects through dietary mixing enhance the performance of a generalist insect herbivore

The ontogenetic niche concept predicts that resource use depends on an organism’s developmental stage. This concept has been investigated primarily in animals that show differing resource use strategies as juveniles and as adults, such as amphibians. We studied resource use and performance in the grasshopper Chorthippus parallelus (Orthoptera, Acrididae) provided with food plant mixtures of either one, three or eight plant species throughout their development. C. parallelus survival and fecundity was highest in the food plant mixture with eight plant species and lowest in the treatments where only one single plant species was offered as food. C. parallelus’ consumption throughout its ontogeny depended on sex, and feeding on different plant species was dependent on a grasshopper’s developmental stage. To depict grasshopper foraging in food plant mixtures compared to foraging on single plant species, we introduce the term “relative forage total” (RFT) based on an approach used in biodiversity research by Loreau and Hector (Nature 413:548–274, 2001). RFT of grasshoppers in food plant mixtures was always higher than what would have been expected from foraging in monocultures. The increase in food consumption was due to an overall increase in feeding on plant species in mixtures compared to consumption of the same species offered as a single diet. Thus we argue that grasshopper foraging exhibits complementarity effects. Our results reinforce the necessity to consider development-related changes in insect herbivore feeding. Thorough information on the feeding ontogeny of insect herbivores could not only elucidate their nutritional ecology but also help to shed light on their functional role in plant communities. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

THE INFLUENCE OF INTRASPECIFIC VARIATION IN DISPERSAL STRATEGIES ON THE GENETIC STRUCTURE OF PLANTHOPPER POPULATIONS

The hypothesis that levels of gene flow among populations are correlated with dispersal ability has typically been tested by comparing gene flow among species that differ in dispersal abilities, an approach that potentially confounds dispersal ability with other species-specific differences. In this study, we take advantage of geographic variation in the dispersal strategies of two wing-dimorphic planthopper species, Prokelisia marginata and P. dolus, to examine for the first time whether levels of gene flow among populations are correlated with intraspecific variation in dispersal ability. We found that in both of these coastal salt marsh–inhabiting species, population-genetic subdivision, as assessed using allozyme electrophoresis, parallels geographic variation in the proportion of flight-capable adults (macropters) in a population; in regions where levels of macroptery are high, population genetic subdivision is less than in regions where levels of macroptery are low. We found no evidence that geographic variation in dispersal capability influences the degree to which gene flow declines with distance in either species. Thus, both species provided evidence that intraspecific variation in dispersal strategies influences the genetic structure of populations, and that this effect is manifested in population-genetic structure at the scale of large, coastal regions, rather than in genetic isolation by distance within a region. This conclusion was supported by interspecific comparisons revealing that: (1) population-genetic structure (G_ST) of the two Prokelisia species correlated negatively with the mean proportion of flight-capable adults within a region; and (2) there was no evidence that the degree of isolation by distance increased with decreasing dispersal capability. Populations of the relatively sedentary P. dolus clustered by geographic region (using Nei's distances), but this was not the case for the more mobile P. marginata. Furthermore, gene flow among the two major regions we surveyed (Atlantic and Gulf Coasts) has been substantial in P. marginata, but relatively less in P. dolus. The results for P. marginata suggest that differences in the dispersal strategies of Atlantic and Gulf Coast populations occur despite extensive gene flow. We argue that gene flow is biased from Atlantic to Gulf Coast populations, indicating that selection favoring a reduction in flight capability must be intense along the Gulf. Together, the results of this study provide the first rigorous evidence of a negative relationship within a species between dispersal ability and the genetic structure of populations. Furthermore, regional variation in dispersal ability is apparently maintained by selective differences that outweigh high levels of gene flow among regions.     

GENETIC STRUCTURE AND MALE-MEDIATED GENE FLOW IN THE GHOST BAT (MACRODERMA GIGAS)

The Australian ghost bat is a large, opportunistic carnivorous species that has undergone a marked range contraction toward more mesic, tropical sites over the past century. Comparison of mitochondrial DNA (mtDNA) control region sequences and six nuclear microsatellite loci in 217 ghost bats from nine populations across subtropical and tropical Australia revealed strong population subdivision (mtDNA φ_ST = 0.80; microsatellites UR_ST = 0.337). Low-latitude (tropical) populations had higher heterozygosity and less marked phylogeographic structure and lower subdivision among sites within regions (within Northern Territory [NT] and within North Queensland [NQ]) than did populations at higher latitudes (subtropical sites; central Queensland [CQ]), although sampling of geographically proximal breeding sites is unavoidably restricted for the latter. Gene flow among populations within each of the northern regions appears to be male biased in that the difference in population subdivision for mtDNA and microsatellites (NT φ_ST = 0.39, UR_ST = 0.02; NQ φ_ST = 0.60, UR_ST = ?0.03) is greater than expected from differences in the effective population size of haploid versus diploid loci. The high level of population subdivision across the range of the ghost bat contrasts with evidence for high gene flow in other chiropteran species and may be due to narrow physiological tolerances and consequent limited availability of roosts for ghost bats, particularly across the subtropical and relatively arid regions. This observation is consistent with the hypothesis that the contraction of the species' range is associated with late Holocene climate change. The extreme isolation among higher-latitude populations may predispose them to additional local extinctions if the processes responsible for the range contraction continue to operate.     

GENETIC AND MORPHOMETRIC DIVERGENCE IN ANCESTRAL EUROPEAN AND DESCENDENT NEW ZEALAND POPULATIONS OF CHAFFINCHES (FRINGILLA COELEBS)

Descendent populations of chaffinches (Fringilla coelebs) introduced to New Zealand about 120 years ago were compared with “ancestral” populations in northern Europe and with those in a broader region of Europe (including Iberia) using protein electrophoresis at 42 loci and 12 skeletal measurements. The New Zealand populations exhibit very small scale differentiation in genetics (F_st = 0.040) and morphometrics, and the haphazard pattern of among-population variation does not align with environmental variation nor is it predicted by the geographic proximity of populations. Thus random drift is implicated in the differentiation among the descendent populations. The New Zealand chaffinches have diverged only slightly in morphometrics from an extant population in southern England, and constant heritability rate tests suggest that random drift alone could account for this small shift. In sharp contrast, the European populations are subdivided genetically (F_st = 0.222) and morphometrically, and this subdivision coincides with the Pyrenees mountains between Iberia and northern Europe which act as a barrier to gene flow between these regions. Iberian populations have smaller skulls and longer wings on average than northern European populations and are characterized by high frequencies of alternative common alleles at Ada and Np. Within both the Iberian and northern European regions, however, populations are effectively panmictic in protein-encoding genes, indicating that homogenizing gene flow is apparently extensive enough to prevent among-population differentiation in allozymes by drift. Variation in body size as represented by PC I is related to environmental productivity across Europe, unlike in New Zealand. These observations jointly suggest that longer term adaptive differentiation via selection for optimal body size has evolved in Europe. Because multilocus evolution is expected to proceed slowly in populations subject to the opposing forces of selection and homogenizing gene flow, I argue that local adaptation within “ancestral” populations in northern Europe may still be evolving.     

Population genetic structure of the Daubenton’s bat (<Emphasis Type="Italic">Myotis daubentonii</Emphasis>) in western Europe and the associated occurrence of rabies

The Daubenton’s bat is widespread and common in the UK and countries bordering the English Channel and North Sea. European bat lyssavirus 2 (EBLV-2), a rabies virus, has been detected in Daubenton’s bats in the UK and continental Europe. Investigating the relatedness of colonies and gene flow between these regions would allow regional estimates of the movement of Daubenton’s bats and thus the potential for disease transmission. The genetic structure of the Daubenton’s bat in western Europe was investigated by analysing variability at eight microsatellite loci. Genetic diversity was found to be high at all sites (H _E = 0.73–0.84), with little differentiation between bats sampled in the UK and continental Europe. Mantel tests indicated a significant correlation between geographic distance and pair-wise F _ST (P = 0.000), between colonies sampled in Scotland and northern England. However, this was not continuous throughout the sampled range, with evidence of panmixia within the area sampled in continental Europe. Assignment tests show no evidence that the (potential) EBLV-2 sero-positive and virus positive bats were more likely to have originated from the continental rather than UK populations. There is no sufficient significant genetic differentiation amongst most UK and continental colonies to conclude that EBLV-2 is maintained in the UK by immigration. Results show that it is likely to be maintained at a low endemic level within the UK. The relative genetic uniformity of UK and continental populations implies that there is no migration barrier to EBLV-2, between these regions.     

Genetic characterisation of Cylindrospermopsis raciborskii (Nostocales, Cyanobacteria) isolates from Africa and Europe

The invasive cyanobacterium Cylindrospermopsis raciborskii is increasingly spreading in temperate freshwater habitats worldwide and is of major concern due to its ability to produce potent toxins. It is, therefore, important to understand the mechanisms behind the dispersal of this species. Different hypotheses have been proposed to explain the phylogeography and mechanisms underlying the recent expansion of C. raciborskii into temperate latitudes, but there is still no conclusive evidence whether the obvious ecological success of C. raciborskii is due to selection mechanisms, physiological tolerance, climatic change or radiation after the last ice age. In the present study, new isolates of C. raciborskii from Europe and Africa were genetically characterised by sequencing the ITS1, PC-IGS, nifH and rpoC1 genes and compared to corresponding sequences of C. raciborskii available in GenBank in order to test different phylogeographical hypotheses. The strains were also morphologically examined and screened for production of the hepatotoxic cylindrospermopsin (CYN). We clearly demonstrate a variation among the populations of C. raciborskii from different geographical regions. The phylogenetic analyses revealed a clustering of the strains due to geographic origin. The ITS1 and nifH genes separated into American, European and Australian–African groups, whereas the PC-IGS and rpoC1 separated into American and European/Australian/African groups. An analysis of concatenated data supported the division into American, European and African/Australian groups, and even indicated a subdivision into an African and an Australian group. Our findings do not strongly support any of the existing hypotheses on the phylogeography of C. raciborskii, and most likely a combination of these hypotheses is the best approach to understand the evolution and dispersal of this species.     

The geography of speciation in narrow‐range endemics of the ‘Haenydra’ lineage (Coleoptera,Hydraenidae, Hydraena)

Aim We test whether species of western Mediterranean aquatic Coleoptera of the ‘Haenydra’ lineage (Hydraenidae, Hydraena) originated through: (1) successive periods of dispersal and speciation, (2) range fragmentation by random vicariance, or (3) range fragmentation by geographic isolation owing to a general reduction of population density. Location Europe. Methods To discriminate between scenarios we use contrasting predictions of the relationship between phylogenetic and geographic distance. The phylogeny was based on 3 kb of four mitochondrial and two nuclear gene fragments of about half of the known species of ‘Haenydra’, including most western Mediterranean taxa. Divergences were estimated using a molecular clock. The relationship between phylogenetic and geographic distance was tested using bivariate plots, Mantel tests and comparison of the observed phylogeny with the one minimizing geographic distances between species, as measured using Euclidean minimum spanning trees (EMSTs). Results The monophyly of ‘Haenydra’ was strongly supported, although its phylogenetic placement was not resolved. ‘Haenydra’ was estimated to be of late Miocene age, with most species originating during the Pleistocene. In two clades (Hydraena tatii and Hydraena emarginata clades) there was a significant association between geographic and phylogenetic distance, and the reconstructed phylogeny was identical to that obtained through the EMST, demonstrating a strong non‐randomness of the geographic distribution of the species. In two other clades (Hydraena iberica and Hydraena bitruncata clades) there was no association between geographic and phylogenetic distance, and the observed phylogeny was not the one minimizing geographic distances. In one of the clades this seems to be due to a secondary, recent range expansion of one species (H. iberica), which erased the geographic signal of their distributions. Main conclusions We show that it is possible to obtain strong evidence of stasis of the geographic ranges of narrow‐range endemic species through the study of their phylogenetic relationships and current distributions. In at least two of the studied clades, current species seem to have originated through the fragmentation of a more widely distributed species, without further range movements. A process of range expansion and fragmentation may have occurred repeatedly within the ‘Haenydra’ lineage, contributing to the accumulation of narrow‐range endemics in Mediterranean Pleistocene refugia.     

Detection of Spiroplasma and Wolbachia in the Bacterial Gonad Community of Chorthippus parallelus

We have recently detected the endosymbiont Wolbachia in multiple individuals and populations of the grasshopper Chorthippus parallelus (Orthoptera: acrididae). This bacterium induces reproductive anomalies, including cytoplasmic incompatibility. Such incompatibilities may help explain the maintenance of two distinct subspecies of this grasshopper, C. parallelus parallelus and C. parallelus erythropus, which are involved in a Pyrenean hybrid zone that has been extensively studied for the past 20 years, becoming a model system for the study of genetic divergence and speciation. To evaluate whether Wolbachia is the sole bacterial infection that might induce reproductive anomalies, the gonadal bacterial community of individuals from 13 distinct populations of C. parallelus was determined by denaturing gradient gel electrophoresis analysis of bacterial 16S rRNA gene fragments and sequencing. The study revealed low bacterial diversity in the gonads: a persistent bacterial trio consistent with Spiroplasma sp. and the two previously described supergroups of Wolbachia (B and F) dominated the gonad microbiota. A further evaluation of the composition of the gonad bacterial communities was carried out by whole cell hybridization. Our results confirm previous studies of the cytological distribution of Wolbachia in C. parallelus gonads and show a homogeneous infection by Spiroplasma. Spiroplasma and Wolbachia cooccurred in some individuals, but there was no significant association of Spiroplasma with a grasshopper’s sex or with Wolbachia infection, although subtle trends might be detected with a larger sample size. This information, together with previous experimental crosses of this grasshopper, suggests that Spiroplasma is unlikely to contribute to sex-specific reproductive anomalies; instead, they implicate Wolbachia as the agent of the observed anomalies in C. parallelus.     

Is life history a barrier to dispersal? Contrasting patterns of genetic differentiation along an oceanographically complex coast

Extreme variation in early life‐history strategies is considered a moderately good predictor of genetic subdivision and hence dispersal for a range of marine species. In reality, however, a good deal of population differentiation must reflect historical effects, more subtle variation in life histories, and, particularly, the interaction of larvae with oceanographic processes. Using a combination of allozyme and microsatellite markers, we show that the large‐scale genetic structure of populations of three species (direct and planktonically developing cushion stars and a planktonic developing sea anemone that is also asexually viviparous) varies consistently, in line with the predicted capacity for dispersal within three geographic regions. We detected high levels of genetic subdivision for the direct developing cushion star (F_ST = 0.6), low levels for the planktonically developing cushion star (F_ST = 0.009), and intermediate levels for the sexual/asexual sea anmone (F_ST = 0.19). These patterns are exhibited despite the highly variable patterns of current movement and the presence of biogeographic barriers. Our results suggest that, although there is large scale genetic differentiation for two species, patterns of population connectivity are remarkably consistent within major regions and do not reflect variation in major oceanographic processes or genetic discontinuity coincident with biogeographic boundaries. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 106–116.     

Thermogeography predicts the potential global range of the invasive European green crab (Carcinus maenas)

Aim The highly adaptable estuarine crab (Carcinus maenas) has successfully invaded five temperate geographic regions outside of its native Europe. Here, we determine which environmental factors predict the current distribution of C. maenas and what the potential geographic range of this species might be. We also investigated whether the invasion potential of C. maenas differs with respect to the origin of a native subpopulation. Location Models were developed using global observation records of C. maenas. Methods Boosted regression trees were used to model observations from the (1) native, (2) invasive, (3) southern European, (4) northern European and (5) the combined native and invasive geographic ranges of C. maenas. Results Most established invasions were predicted mainly based on temperature. Interestingly, the environment encountered by established invasions failed to predict the majority of northern European populations; suggesting that invasion potential may differ between distinct native populations. Supporting this suggestion, a model of northern European populations, distinguished from southern European populations based on genetic structure, only predicted established invasions south of Nova Scotia. By contrast, a model of southern European populations predicted most established invasions. Main conclusions These results suggest that invasion potential depends on the European origin of an invasive population and that most invasions have arisen from southern Europe. Finally, a model based on combined native and invasive ranges of C. maenas identified potential geographic range extension along many currently invaded coastlines and the potential invasion of countries like Chile, China, Russia, Namibia and New Zealand.     

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.     

Nucleolus size varies with sex,ploidy and gene dosage in insects

The nucleolus constitutes a cytologically visible phenotype for ribosomal DNA (rDNA). Nucleolar size, as determined by silver staining, is a good indicator of cell proliferation rate and biosynthetic activity. Nevertheless, the relationship between rDNA content and sexual dimorphism for nucleolar size is not well documented. In the present study, the impact of sex and ploidy level on nucleolar size is investigated in three haplo/diploid and three diplo/diploid species of insect. Nucleolar sizes are found to be proportional to ploidy level in the haplo/diploid hymenopterans Trypoxylon albitarse and Nasonia vitripennis. Conversely, in the ant Messor barbarus, nucleolar sizes are larger in haploid males (winged) than diploid females (apterous). Among the diplo/diploid species, evidence for gene dosage compensation on nucleolar activity is suggested by the absence of sex differences in Drosophila simulans, a species in which rDNA is limited to the X chromosome. By contrast, in the grasshopper Stenobothrus festivus, another species with rRNA genes restricted to the X chromosome, the size of the nucleolus is significantly larger in females than in males. Additionally, in the grasshopper Chorthippus parallelus, where rDNA is distributed evenly on several autosomes of males and females, the females also show larger nucleoli than males. In both grasshopper species, the magnitude of the female/male ratio for nucleolus area is very similar to the body size ratio, suggesting that body size, as well as sex, ploidy, gene dosage and physiological activity, may be an important determinant of nucleolus area.     

Nuclear and chloroplast microsatellites show multiple introductions in the worldwide invasion history of common ragweed, Ambrosia artemisiifolia

Background

Ambrosia artemisiifolia is a North American native that has become one of the most problematic invasive plants in Europe and Asia. We studied its worldwide population genetic structure, using both nuclear and chloroplast microsatellite markers and an unprecedented large population sampling. Our goals were (i) to identify the sources of the invasive populations; (ii) to assess whether all invasive populations were founded by multiple introductions, as previously found in France; (iii) to examine how the introductions have affected the amount and structure of genetic variation in Europe; (iv) to document how the colonization of Europe proceeded; (v) to check whether populations exhibit significant heterozygote deficiencies, as previously observed.

Principal Findings

We found evidence for multiple introductions of A. artemisiifolia, within regions but also within populations in most parts of its invasive range, leading to high levels of diversity. In Europe, introductions probably stem from two different regions of the native area: populations established in Central Europe appear to have originated from eastern North America, and Eastern European populations from more western North America. This may result from differential commercial exchanges between these geographic regions. Our results indicate that the expansion in Europe mostly occurred through long-distance dispersal, explaining the absence of isolation by distance and the weak influence of geography on the genetic structure in this area in contrast to the native range. Last, we detected significant heterozygote deficiencies in most populations. This may be explained by partial selfing, biparental inbreeding and/or a Wahlund effect and further investigation is warranted.

Conclusions

This insight into the sources and pathways of common ragweed expansion may help to better understand its invasion success and provides baseline data for future studies on the evolutionary processes involved during range expansion in novel environments.     

Comparative population structure of three snook species (Teleostei: Centropomidae) from the eastern central Pacific

Three snook species, Centropomus viridis, Centropomus medius, and Centropomus robalito, from the eastern central Pacific, representing three of the four proposed phyletic lineages in the genus, were analyzed for genetic variability by means of allozyme and RAPD to evaluate the divergence between populations at different levels of dispersal ability and to evaluate the importance of barriers to dispersal in the population subdivision and genetic diversity. Levels of genetic diversity among species estimated by allozymes were similar and consistent with the observed levels of differentiation in marine fish species. Mean heterozygosity ranged from 0.089 for C. viridis to 0.10 for C. robalito. Genetic diversity for the snook species studied was slightly higher than the mean estimation reported in allozymes for 106 marine fish (0.055) and for anadromous fish species (0.043 to 0.057). Multilocus allele frequency homogeneity tests and population-subdivision estimates for both allozyme and RAPD markers revealed the existence of population structure in C. viridis and C. medius, in coincidence with geographic separation of samples, whereas no divergence was detected in C. robalito. This finding may be attributed to the greater population size of C. robalito, which originated by a recent population range expansion, and hence the potential for dispersal is mediated by larval drift. Fluctuations in population size and population range expansion are used to explain discrepancies between levels of genetic diversity and population structure in the studied species. Supplementary material to this paper is available in electronic format at