Spatiotemporal allozyme variation in the damselfly, <Emphasis Type="BoldItalic">Lestes viridis</Emphasis> (Odonata: Zygoptera): gene flow among permanent and temporary ponds

Several insect species seem to persist not only in permanent but also in temporary ponds where they face particularly harsh conditions and frequent extinctions. Under such conditions, gene flow may prevent local adaptation to temporary ponds and may promote phenotypic plasticity, or maintain apparent population persistence. The few empirical studies on insects suggest the latter mechanism, but no studies so far quantified gene flow including both pond types. We investigated the effects of pond type and temporal variation on population genetic differentiation and gene flow in the damselfly Lestes viridis in northern Belgium. We report a survey of two allozyme loci (Gpi, Pgm) with polyacrylamide gel electrophoresis in 14 populations from permanent and temporary ponds, and compared these results with similar data from the same permanent populations one year before. The data suggested that neither pond-drying regime, nor temporal variation have a substantial effect on population genetic structuring and did not provide evidence for stable population differentiation in L. viridis in northern Belgium. Gene flow estimates were high within permanent and temporary ponds, and between pond types. Our data are consistent with a source-sink metapopulation system where temporary ponds act as sinks in dry years, and are quickly recolonized after local population extinction. This may create a pattern of apparent population persistence of this species in permanent and temporary ponds without clear local adaptation.     

Emergence of the damselfly Pyrrhosoma nymphula (Sulzer) (Zygoptera: Coenagrionidae) from two adjacent ponds in northern England

Emergence of the damselfly Pyrrhosoma nymphula was followed closely at two similar and adjacent ponds in northern England. Males emerged earlier than females at both ponds (significantly so at one). The sex ratio was significantly male biased at both ponds. Size of emerging adults declined through the emergence period at both ponds. Adults emerging from one pond did so significantly earlier and were significantly larger than those from the other pond. The density of larvae was approximately twice as high in the pond from which larvae emerged early; this pond was also slightly deeper and usually achieved higher maximum daily water temperatures. These findings are discussed in the light of the conventional view of seasonal regulation in a spring-emerging damselfly.     

Spatiotemporal analysis of gene flow in Chesapeake Bay Diamondback Terrapins (Malaclemys terrapin)

There is widespread concern regarding the impacts of anthropogenic activities on connectivity among populations of plants and animals, and understanding how contemporary and historical processes shape metapopulation dynamics is crucial for setting appropriate conservation targets. We used genetic data to identify population clusters and quantify gene flow over historical and contemporary time frames in the Diamondback Terrapin (Malaclemys terrapin). This species has a long and complicated history with humans, including commercial overharvesting and subsequent translocation events during the early twentieth century. Today, terrapins face threats from habitat loss and mortality in fisheries bycatch. To evaluate population structure and gene flow among Diamondback Terrapin populations in the Chesapeake Bay region, we sampled 617 individuals from 15 localities and screened individuals at 12 polymorphic microsatellite loci. Our goals were to demarcate metapopulation structure, quantify genetic diversity, estimate effective population sizes, and document temporal changes in gene flow. We found that terrapins in the Chesapeake Bay region harbour high levels of genetic diversity and form four populations. Effective population sizes were variable. Among most population comparisons, estimates of historical and contemporary terrapin gene flow were generally low (m ≈ 0.01). However, we detected a substantial increase in contemporary gene flow into Chesapeake Bay from populations outside the bay, as well as between two populations within Chesapeake Bay, possibly as a consequence of translocations during the early twentieth century. Our study shows that inferences across multiple time scales are needed to evaluate population connectivity, especially as recent changes may identify threats to population persistence.     

Allozyme and mitochondrial DNA variability within the New Zealand damselfly genera Xanthocnemis,Austrolestes, and Ischnura (Odonata)

Abstract

We collected larval damselflies from 17 sites in the North, South and Chatham Islands, and tested the hypotheses that: (1) genetic markers (e.g., allozymes, mtDNA) would successfully discriminate taxa; and (2) the dispersal capabilities of adult damselflies would limit differentiation among locations. Four species from three genera were identified based on available taxonomic keys. Using 11 allozyme loci and the mitochondrial cytochrome c‐oxidase subunit I (COI) gene, we confirmed that all taxa were clearly discernible. We found evidence for low to moderate differentiation among locations based on allozyme (meani F _ST = 0.09) and sequence (COI) divergence (<0.034). No obvious patterns with respect to geographic location were detected, although slight differences were found between New Zealand's main islands (North Island, South Island) and the Chatham Islands for A. colensonis (sequence divergence 0.030–0.034). We also found limited intraspecific genetic variability based on allozyme data (H_exp < 0.06 in all cases). We conclude that levels of gene flow/dispersal on the main islands may have been sufficient to maintain the observed homogeneous population structure, and that genetic techniques, particularly the COI gene locus, will be a useful aid in future identifications.     

Population genetics and gene variation in screwworms (Diptera: Calliphoridae) from Brazil

Allozyme variation in New World screwworm,Cochliomyia hominivorax (Coquerel), populations from Brazil was examined. Variability was observed in 8 of 13 enzyme loci and the frequency of the most common allele was <0.95 for seven loci. Observed and expected heterozygosities were 0.159 and 0.165, respectively. Comparisons of the Brazilian populations with previously studied populations from Costa Rica resulted in Nei's genetic distances of between 0.000 and 0.006, with the greatest distance being between populations within Brazil. Comparisons with Mexican populations using only three loci resulted in genetic distances ≤0.031. Goodness-of-fit statistics for Hardy-Weinberg equilibrium and Wright'sF statistics indicated small deviations from expected equilibrium genotype frequencies and low levels of differentiation between populations within Brazil. Differentiation among screwworm populations from Brazil, Costa Rica, and Mexico was minimal.     

Population genetic structure, gene flow and sex-biased dispersal in frillneck lizards (Chlamydosaurus kingii)

By using both mitochondrial and nuclear multiloci markers, we explored population genetic structure, gene flow and sex-specific dispersal of frillneck lizards ( Chlamydosaurus kingii ) sampled at three locations, separated by 10 to 50 km, in a homogenous savannah woodland in tropical Australia. Apart from a recombinant lizard, the mitochondrial analyses revealed two nonoverlapping haplotypes/populations, while the nuclear markers showed that the frillneck lizards represented three separate clusters/populations. Due to the small population size of the mtDNA, fixation may occur via founder effects and/or drift. We therefore suggest that either of these two processes, or a combination of the two, are the most likely causes of the discordant results obtained from the mitochondrial and the nuclear markers. In contrast to the nonoverlapping mitochondrial haplotypes, in 12 out of 74 lizards, mixed nuclear genotypes were observed, hence revealing a limited nuclear gene flow. Although gene flow should ultimately result in a blending of the populations, we propose that the distinct nuclear population structure is maintained by frequent fires resulting in local bottlenecks, and concomitant spatial separation of the frillneck lizard populations. Limited mark–recapture data and the difference in distribution of the mitochondrial and nuclear markers suggest that the mixed nuclear genotypes were caused by juvenile male-biased dispersal.     

Leaching from vegetation of soluble polyphenolic compounds,and their abundance in temporary ponds in the Doñana National Park (SW Spain)

The origin of allochthonous soluble polyphenols in the water of the temporary ponds of Doñana National Park during flood and heavy rainfall was examined. Soil and litter samples were found to leach large quantities of polyphenolic compounds. The concentration of these compounds showed a significant correlation with the organic content of the soil samples, but indicated no correlation with the litter samples. Rain simulation tests proved that fresh vegetation leached soluble polyphenolic compounds. Mediterranean scrub (Cystaceae) from dry poor-nutrient soils yielded the highest concentration (being furthest away from the ponds), whereas wetland (Ericaceae) and littoral vegetation (Juncaceae and Cyperaceae) rated second and third, respectively. The potential of the surrounding vegetation to leach high concentrations of polyphenolic compounds suggests that this is the main source of the rapid polyphenolic enrichment of these ponds during periods of heavy rain.     

Allozyme variation in stable flies (Diptera: Muscidae)

Polyacrylamide gel electrophoresis was used to resolve allozymes in the cosmopolitan blood-feeding stable fly,Stomoxys calcitrans (L.). Nineteen of 38 loci were polymorphic (53%). Mean heterozygosities among all loci and among only polymorphic loci were 0.096 and 0.182, respectively. These gene diversity measures are about half those among other muscid Diptera. Variation in gene frequencies was examined in 10 natural stable fly populations from Iowa and Minnesota. Gene frequencies were homogeneous at five of eight loci among six populations in 1990 and eight of eight loci among four populations in 1992. Wright'sF statistics showed no significant departure from random mating among stable flies. It was concluded that gene flow compensates for any local differentiation in stable fly populations.     

Microgeographic edaphic differentiation in allozyme polymorphisms of wild barley (Hordeum spontaneum,Poaceae)

Allozymic variation in proteins encoded by 22 loci was analyzed electrophoretically in 278 individual plants of wild barley,Hordeum spontaneum, the progenitor of cultivated barley, in four 100 meter transects, in Israel, each equally subdivided into basalt and terra rossa soil types. Significant differentiation according to soil was found in 9 alleles. Our results suggest that allozyme polymorphisms in wild barley are at least partly adaptive, and differentiate by edaphic natural selection rather than by stochastic processes, and/or neutrality of allozymic variants.     

Gene variation and gene flow inOrchis morio (Orchidaceae) from Italy

Data are presented on genetic variation at 27 enzyme loci of the Green-Winged orchid,Orchis morio, in 18 population samples from Italy. The existence in Italy of two subspecies, i.e. subspp.morio andpicta, is not supported by allozyme data. No genetic heterogeneity was found betweenmorio-like andpicta-like samples and specimens. Moreover, morphological transition between the two forms was observed in different Italian populations. The parameters of genetic variability estimated forO. morio populations are consistent with those found among monocotyledon plants, and among those outcrossing, animal-pollinated and with wind-dispersed seeds. Genetic diversity of ItalianO. morio is mostly within populations. Correspondingly, low values of interpopulational genetic distance were found. This appears to be due to high levels of gene flow, which were estimated with different methods. The lack ofO. longicornu from Italian samples, as well as of any hybrid withO. morio (F₁, backcrossed or recombinant individuals) is demonstrated on the basis of genetic data. It is concluded that recurrent reports ofO. longicornu from Italy are due to confusion withO. morio or with otherOrchis species.     

Estimates of gene flow, genetic substructure and population heterogeneity in bracken (Pteridium aquilinum)

Bracken [ Pteridium aquilinum (L.) Kuhn] is a cosmopolitan species and is a noxious weed in many areas. Because of its abundance, particularly in Britain, bracken affords an ideal system for investigating various aspects of population genetics and evolution. High mobility of dispersal units (spores) suggests that rates of gene flow among distant populations should be high. Gene flow is a major evolutionary force that influences the genetic structure of populations. To examine the effects of gene flow on population heterogeneity and population substructuring in bracken, starch gel electrophoresis of enzymes was used to provide the necessary genetic database. Allele frequency data at 21 loci were obtained for seven British populations, one Majorcan and one from the eastern United States. A model was employed to estimate the amount of gene flow ( Nm ) at several levels. Gene flow among British populations was extremely high ( Nm = 36.51), one of the highest estimates reported for plants. Among eight European populations gene flow was lower (but still considered high) at Nm = 2.47. Trans-Atlantic gene flow was low ( Nm = 0.0926).
F -statistics were used to assess population heterogeneity and substructuring. The data indicate that, compared with other species, there is very little genetic differentiation among British populations of bracken. Indeed, it appears that the whole island is behaving as a single randommating population. This result is consistent with high levels of gene flow. Only one population (on the Isle of Arran) showed statistically significant genetic substructuring. Habitat heterogeneity on the island and age structure are hypothesized as possible causes of this result.
The data reported here support previous studies demonstrating that bracken is genetically polymorphic and is an outcrossing species.     

Australian lungfish (Neoceratodus forsteri: Dipnoi) have low genetic variation at allozyme and mitochondrial DNA loci: a conservation alert?

Genetic variation at allozyme and mitochondrialDNA loci was investigated in the Australianlungfish, Neoceratodus forsteri Krefft1870. Tissue samples for genetic analysis weretaken non-lethally from 278 individualsrepresenting two spatially distinct endemicpopulations (Mary and Burnett rivers), as wellas one population thought to be derived from ananthropogenic translocation in the 1890's(Brisbane river). Two of 24 allozyme lociresolved from muscle tissue were polymorphic.Mitochondrial DNA nucleotide sequence diversityestimated across 2,235 base pairs in each of 40individuals ranged between 0.000423 and0.001470 per river. Low genetic variation atallozyme and mitochondrial loci could beattributed to population bottlenecks, possiblyinduced by Pleistocene aridity. Limited geneticdifferentiation was detected among rivers usingnuclear and mitochondrial markers suggestingthat admixture may have occurred between theendemic Mary and Burnett populations duringperiods of low sea level when the drainages mayhave converged before reaching the ocean.Genetic data was consistent with theexplanation that lungfish were introduced tothe Brisbane river from the Mary river. Furtherresearch using more variable genetic loci isneeded before the conservation status ofpopulations can be determined, particularly asanthropogenic demands on lungfish habitat areincreasing. In the interim we recommend amanagement strategy aimed at conservingexisting genetic variation within and betweenrivers.     

Gene flow among San Joaquin kit fox populations in a severely changed ecosystem

The San Joaquin kit fox (Vulpes macrotis mutica) was once ubiquitous throughout Californias San Joaquin Valley and its surrounds. However, most of its habitat has been lost to irrigated agriculture, urban development, and oil fields. The remaining foxes are concentrated in six areas, although there are several small pockets of foxes throughout the Valley. To help conserve kit foxes, we sought an ecological understanding of the level of genetic variation remaining in these locations and the extent of gene flow among them. We collected tissue from 317 kit foxes from 8 sites and estimated genetic variability in and gene flow among sites using data from 8 polymorphic, microsatellite markers. We found no differences in both observed and expected heterozygosity between locations using Bonferonni corrected paired t-tests. We found differences in mean number of alleles per locus, even after we used Monte Carlo simulations to adjust for sample size differences. Population subdivision was low among sites (F_st=0.043), yet a matrix of pairwise F_st values was correlated with a matrix of pairwise geographic distances. An assignment test classified only 45% of the individuals to the site where they were captured. Overall, these data suggest that kit fox dispersal between locations may still maintain genetic variation throughout most of the areas we sampled.     

Evolutionary history, population genetics, and gene flow in the common rosefinch (Carpodacus erythrinus)

We analyzed sequences of two mitochondrial DNA gene regions (control region and ND2) from 186 specimens obtained from 17 Eurasian localities covering most of the distribution of the common rosefinch to assess phylogeographic structure. Populations possessed a high level of nucleotide diversity relative to many other Palearctic species, suggesting that rosefinch populations are relatively old and possess high effective sizes. Mismatch distributions suggested that many localities experienced past population expansions, which are older than those expected for post-Pleistocene climate warming and reforestation of Eurasia. Our Phi(st) analysis revealed that 12.4% of total genetic variation was distributed among localities owing in part to the existence of three incompletely isolated groups: southwestern (subspecies C. e. kubanensis), northeastern (subspecies C. e. grebnitskii), and northwestern (subspecies C. e. erythrinus and C. e. ferghanensis). The three groups are not reciprocally monophyletic which suggests that they were formed relatively recently. Gene flow among groups is restricted. Coalescence analysis indicated dispersal asymmetry.     

Biochemical genetic variation among populations of the greenshell mussel, Perna canaliculus, from New Zealand: preliminary findings

Seven polymorphic and four monomorphic allozyme loci were assayed from nine wild populations and one cultured population of the endemic New Zealand greenshell mussel, Perna canaliculus. Genetic variation was examined to determine the extent of intra-population subdivision and inter-population variability. Five of seven polymorphic loci exhibited significant heterozygote deficiencies compared to Hardy-Weinberg equilibrium. Population F-statistics indicated that significant genetic heterogeneity exists among populations, indicating that there is insufficient gene flow between geographically isolated populations to create panmixia. The observed genetic heterogeneity among populations is best explained by an isolation-by-distance model of gene flow, which is modified by localized hydrographic conditions. These preliminary findings are discussed in the context of the one previous survey of population genetic variation in P. canaliculus and how this information relates to the gene flow of the greenshell mussel in New Zealand, which is often mediated by human transport from the main collection site of Kaitaia to aquaculture sites throughout the country.     

Genetic variation and putative hybridization inSalix alba andS. fragilis (Salicaceae): Evidence from allozyme data

Genetic variability was estimated by enzyme electrophoresis in 239 Belgian clones from theSalix alba-S. fragilis complex. This morphological complex suggested a high frequency of hybrids. To test this hypothesis, the clones were pooled as a single co-adapted species complex and secondly as belonging to either species, i.e. beingS. alba-like orS. fragilis-like. The standard genetic variability measures showed higher values for the complex than for the separate species. However, the observed mean heterozygosity was lower in the putative hybrid complex than for each of the species separately. The fixation indices were more variable at the species level and indicated that mostS. fragilis locations appeared fixed forlap-1, whereas no fixation occured inS. alba locations. Averaged at the regional (i.e. catchment) level, this difference between the two species remained and values ranged from 0.457 to 0.617 inS. alba and were much higher, fixed homozygous or monomorph inS. fragilis. Hierarchical F-statistics revealed that most of the differentiation occured at the lower levels of localities and tributaries and that there was no further differentiation between catchments. Tributaries which are 10–25km in length were proposed as the most likely entities for further examination of putative hybridization and events of allelic fixations. By considering the two abovementioned approaches of data input, it could be suggested that most of the allozyme differentiation was between the species and less between the regions.     

Temporal recruitment patterns and gene flow in kelp rockfish (Sebastes atrovirens)

Pelagic dispersal of marine organisms provides abundant opportunity for gene flow and presumably inhibits population genetic divergence. However, ephemeral, fine-scale, temporal and spatial genetic heterogeneity is frequently observed in settled propagules of marine species that otherwise exhibit broad-scale genetic homogeneity. A large variance in reproductive success is one explanation for this phenomenon. Here, genetic analyses of 16 microsatellite loci are used to examine temporal patterns of variation in young-of-year kelp rockfish (Sebastes atrovirens) recruiting to nearshore habitat in Monterey Bay, California, USA. Population structure of adults from central California is also evaluated to determine if spatial structure exists and might potentially contribute to recruitment patterns. Genetic homogeneity was found among 414 young-of-year sampled throughout the entire 1998 recruitment season. No substantial adult population structure was found among seven populations spanning 800 km of coastline that includes the Point Conception marine biogeographic boundary. Comparison of young-of-year and adult samples revealed no genetic differentiation and no measurable reduction in genetic variation of offspring, indicating little variance in reproductive success and no reduction in effective population size for this year class. Simulation analyses determined that the data set was sufficiently powerful to detect both slight population structure among adults and a small reduction in effective number of breeders contributing to this year class. The findings of high gene flow and low genetic drift have important implications for fisheries management and conservation efforts.     

Genetic diversity within Anatolian brown hares (Lepus europaeus Pallas, 1778) and differentiation among Anatolian and European populations

Genetic variability of Anatolian hares and relationships between Anatolian and European populations were assessed by a multilocus allozyme approach to infer evolutionary relationships between hares from Asia Minor and Europe. Of the 48 loci assayed, 19 (39.6%) were polymorphic with two to four alleles in the Anatolian hares. Among all Anatolian alleles, 14 were so far not found in the compared 717 brown hares from Europe. Overall, genetic diversity was highest in Anatolian hares, intermediate in brown hares from the southern and southeastern Balkans and lowest in central European populations. The rich genetic diversity in Anatolian hares might be a consequence of Anatolias biogeographic position with the chance of multiple gene flow from neighbouring regions, and the likelihood of long-term presence of hares during the last ice age, when large parts of more northern latitudes did not provide suitable habitats.However, among 28 loci used for the comparison between European and Anatolian populations, most common alleles of European brown hares were also common in Anatolian populations and no alternately fixed alleles were found for Anatolian and European populations. This together with only little or moderately varying allele frequencies produced low genetic divergence between Anatolian and European populations. Genetic differentiation among Anatolian populations was also low. Even between the two forms with different coat colour (brownish and yellowish) in Anatolian hares, there was little genetic differentiation. Altogether, all Anatolian hares studied presently are closely related to European brown hare populations, and only some distantly spaced population pairs revealed increased genetic divergence.

Zusammenfassung

Genetische Diversität anatolischer Feldhasen (Lepus europaeus Pallas, 1778) und Differenzierung zwischen anatolischen und europäischen PopulationenZur Beurteilung der phylogenetischen Beziehungen zwischen anatolischen Hasen und europäischen Feldhasenpopulationen wurde die allelische Variabilität anatolischer Hasen mittels horizontaler Stärkegelelektrophorese erfaßt und gemeinsam mit unmittelbar vergleichbaren Daten griechischer, bulgarischer und österreichischer Populationen aus früheren Studien populationsgenetischen Analysen unterzogen. Neunzehn der 48 untersuchten Loci der anatolischen Hasen zeigten allelische Variabilität. Unter den anatolischen Allelen kamen 14 bisher in den europäischen Polulationen nicht vor. Insgesamt zeigten anatolische Hasen die höchste und österreichische Populationen die niedrigste genetische Diversität; die jeweiligen Werte der griechischen und bulgarischen Populationen lagen dazwischen. Dies entspricht unserer Hypothese hoher genetischer Diversität in Anatolien, auf Grund der biogeografischen Position und der klimatischen bzw. Lebensraumbedingungen während des Pleistozäns, die, im Gegensatz zu Mitteleuropa, kontinuierliche Hasenpopulation in Anatolien wahrscheinlich erscheinen lassen. Kontinuierliche Populationen und Genflüsse aus verschiedenen Nachbarregionen könnten bei langfristig relative ungestörten Populationen zur Anreicherung genetischer Varianten in Anatolien geführt haben, während mitteleuropäische Feldhasenpopulationen im Zuge ihrer postglazialen Einwanderung aus Refugial-gebieten an genetischer Vielfalt eingebüßt haben. Allerdings waren die häufigen Allele der anatolischen Hasen ebenfalls häufig bei den europäischen Feldhasen vertreten; somit ergab sich insgesamt nur eine geringe genetische Differenzierung zwischen anatolischen und europäischen Feldhasen. Die zwei in Anatolien gefundenen Fellfärbungstypen (brauner vs. gelber Grundton) zeigten ebenfalls keine besondere genetische Differenzierung.