Clinal genetic variation and the 'rare allele phenomenon' in random mating populations of <Emphasis Type="Italic">Urophora cardui</Emphasis> (Diptera: Tephritidae)

In the present study we investigate a contact zone between two population groups of the tephritid fly Urophora cardui. We investigate scenarios that may have produced the genetic differentiation of the two groups, and we describe the 'rare allele phenomenon' from the contact zone. The rare allele phenomenon refers to alleles that are found at high frequency in contact zones but are rare or lacking outside the contact zone. The phenomenon is often observed in hybrid zones between subspecies of limited reproductive compatibility, but seldom in populations with random mating. Clinal genetic variation was observed at three loci in the contact zone. Three alleles at the locus Aat showed steep clines, between 20--70 km wide. A rare Aat-A allele occurred at high frequency in the centre of the contact zone. Two further loci, Hk and Pgd, showed less steep clinal genetic variation, the transition being in and slightly south of the centre of the Aat cline. Populations showed Hardy--Weinberg proportions and there was no evidence for linkage dis-equlibrium. These findings suggest random mating and gradual introgression between the population systems, which may originate from at least two range expansions. Aat's steep clines and rare allele may indicate selection on Aat alleles, although we presently can not quantify any agents. Because U. cardui experiences random mating in the contact zone with no apparent 'hybrid' incompatibility, mating experiments offer the possibility for future enquiries about the genetic basis of the rare allele phenomenon.     

Different cell size and cell number contribution in two newly established and one ancient body size cline of Drosophila subobscura

Latitudinal genetic clines in body size occur in many ectotherms including Drosophila species. In the wing of D. melanogaster, these clines are generally based on latitudinal variation in cell number. In contrast, differences in wing area that evolve by thermal selection in the laboratory are in general based on cell size. To investigate possible reasons for the different cellular bases of these two types of evolutionary response, we compared the newly established North and South American wing size clines of Drosophila subobscura. The new clines are based on latitudinal variation in cell area in North America and cell number in South America. The ancestral European cline is also based on latitudinal variation in cell number. The difference in the cellular basis of wing size variation in the American clines, which are roughly the same age, together with the similar cellular basis of the new South American cline and the ancient European one, suggest that the antiquity of a cline does not explain its cellular basis. Furthermore, the results indicate that wing size as a whole, rather than its cellular basis, is under selection. The different cellular bases of different size clines are most likely explained either entirely by chance or by different patterns of genetic variance--or its expression--in founding populations.     

A morphological cline in Eucalyptus: a genetic perspective

The putative hybrid zone between Eucalyptus populnea and E. brownii is examined using morphological and molecular techniques. This species complex displays continuous morphological variation across the study area, which has been previously interpreted as the product of hybridization between allopatric species. A microsatellite analysis indicates that there was little genetic structuring across the morphological cline and only low levels of population differentiation. The nested clade analysis of the JLA+ region of the chloroplast DNA (cpDNA) indicates that the geographical distribution of cpDNA haplotypes is unlikely to be the result of historical hybridization events, and that restricted seed-mediated gene flow with isolation by distance is responsible for the phylogeographical distribution. A more plausible explanation for the origin and persistence of the morphological cline is that the process of continuous morphological diversification has been promoted by a directional selection gradient. This study addresses species status within Eucalyptus and the belief that hybridization is widespread and is an important process in the group's evolution.     

Latitudinal clines in<Emphasis Type="Italic">Drosophila melanogaster</Emphasis>: Body size,allozyme frequencies,inversion frequencies,and the insulin-signalling pathway

Many latitudinal clines exist inDrosophila melanogaster: in adult body size, in allele frequency at allozyme loci, and in frequencies of common cosmopolitan inversions. The question is raised whether these latitudinal clines are causally related. This review aims to connect data from two very different fields of study, evolutionary biology and cell biology, in explaining such natural genetic variation inD. melanogaster body size and development time. It is argued that adult body size clines, inversion frequency clines, and clines in allele frequency at loci involved in glycolysis and glycogen storage are part of the same adaptive strategy. Selection pressure is expected to differ at opposite ends of the clines. At high latitudes, selection onD. melanogaster would favour high larval growth rate at low temperatures, and resource storage in adults to survive winter. At low latitudes selection would favour lower larval critical size to survive crowding, and increased male activity leading to high male reproductive success. Studies of the insulin-signalling pathway inD. melanogaster point to the involvement of this pathway in metabolism and adult body size. The genes involved in the insulin-signalling pathway are associated with common cosmopolitan inversions that show latitudinal clines. Each chromosome region connected with a large common cosmopolitan inversion possesses a gene of the insulin transmembrane complex, a gene of the intermediate pathway and a gene of the TOR branch. The hypothesis is presented that temperateD. melanogaster populations have a higher frequency of a ’thrifty’ genotype corresponding to high insulin level or high signal level, while tropical populations possess a more ’spendthrift’ genotype corresponding to low insulin or low signal level.     

Postglacial migration and the loss of allozyme variation in northern populations of Asclepias exaltata (Asclepiadaceae)

The recent Wisconsin glaciation has provided opportunities for examining the effects of postglacial recolonization on the population genetics of plant and animal communities. In this study allozyme Variation was examined in 19 populations of the herbaceous perennial Asclepias exaltata occurring in previously glaciated regions of North America. These northern populations of A. exaltata possess significantly fewer polymorphic loci (46.31 ± 2.7; mean ± 1 SD), alleles per polymorphic locus (1.84 ± 0.24), and expected heterozygosity (0.133 ± 0.031) than populations found in the Pleistocene refugium in the southern Appalachians. Population-level allozyme diversity decreased linearly from south to north and from east to west. Nineteen uncommon alleles previously observed in southern Appalachian populations were undetected in the northern region. Seven common alleles exhibited a clinal change in allele frequency. Of these, only Pgd-1a and Mnr-1c were at low-frequency in the southern Appalachians and increased significantly with increasing latitude and longitude, respectively. Despite this loss of allozyme diversity following postglacial migration, northern populations of A. exaltata have higher allozyme diversity and lower population differentiation (G" = 0.1 17) than mean values for other long-lived herbaceous perennials. Increased habitat fragmentation in northern regions and potential habitat loss in the southern Appalachians are likely to reduce the historically rich gene pool that has provided the genetic stock for postglacial recoveries.     

Origin and decay of the P element-associated latitudinal cline in Australian Drosophila melanogaster

The latitudinal cline in P transposable element-associated characteristics in eastern Australian populations of Drosophila melanogaster has changed between 1986 and 1991–1994. New collections were made in 1991–1994 from localities along the eastern coast of Australia. P element-associated properties of 256 isofemale lines from 43 localities were evaluated using gonadal dysgenesis and/or singed-weak hypermutability assays. The overall results indicate that both P activity and P susceptibility have declined, with all populations showing a tendency towards a state with little P activity potential but with P repressor function (neutral or ‘Q’). P repressor function is strong in all populations except some of the most southerly. P activity potential peaks at about 27° SLat, and drops off to the south (as in 1983–1986 collections) and to the north (in contrast to 1983–1986 collections); thus the cline is no longer a simple P-to-Q-to-M pattern from north to south, but is now Q-P-Q-M. A mtDNA RFLP that putatively distinguishes North American and European populations varies in frequency among the populations but the frequency does not vary clinally with latitude, ruling out massive introductions from North America and Europe as causing the cline. This revised version was published online in July 2006 with corrections to the Cover Date.     

EVOLUTION OF RESISTANCE IN CULEX PIPIENS: ALLELE REPLACEMENT AND CHANGING ENVIRONMENT

Fixation of adaptive mutations in populations is often constrained by pleiotropic fitness costs. The evolutionary pathways that compensate such fitness disadvantages are either the occurrence of modifier genes or replacement of the adaptive allele by less costly ones. In this context, 23 years of evolution of insecticide resistance genes in the mosquito Culex pipiens from southern France are analyzed. The aim of this study is to answer the following points. Is there a fitness cost associated with these resistance genes in natural populations? Does evolution proceed through allele replacement or through selection of modifiers? And finally, how do environmental changes affect the evolution of resistance genes? Samples from the same transect, crossing the boundary between an insecticide-treated and a nontreated area, are analyzed. Clinal analyses indicate a variable fitness cost among the resistance genes and show that allele replacement has been the primary mechanism of resistance evolution in this area. It is also shown that replacement was probably due to environmental changes corresponding to modification in pesticide-treatment intensity.     

Variation in frequency of hybrids and spatial structure among Ipomopsis (Polemoniaceae) contact sites

Rates of hybridization vary among angiosperm taxa. Among-taxon variation in hybridization rate has been used to compare the importance of pre- and post-zygotic reproductive isolating mechanisms. Variation in rates of hybridization within a single-species pair would suggest that local conditions also affect reproductive isolation within a single taxonomic context. In this study, contact sites of Ipomopsis aggregata-Ipomopsis tenuituba were surveyed for variation in frequency of hybrids, and spatial structure. Floral morphology was used to identify parent species and hybrids in seven contact sites in the western Rocky Mountains, USA. Contact sites varied widely in elevational range, the degree to which morphological variation was clinal rather than mosaic and the frequency of hybrids. Two sites provided a strong contrast between a clinal, unimodal site and a mosaic, bimodal site. This natural variation among contact sites of the same species pair provides an opportunity to assess the effect of local ecological conditions and spatial structure of parent populations on reproductive isolation, while controlling for between-taxon variation.     

Clinal variation in a brown lemur (Eulemur spp.) hybrid zone: Combining morphological,genetic and climatic data to examine stability

Studies of hybrid zones can inform our understanding of reproductive isolation and speciation. Two species of brown lemur (Eulemur rufifrons and E. cinereiceps) form an apparently stable hybrid zone in the Andringitra region of south‐eastern Madagascar. The aim of this study was to identify factors that contribute to this stability. We sampled animals at 11 sites along a 90‐km transect through the hybrid zone and examined variation in 26 microsatellites, the D‐loop region of mitochondrial DNA, six pelage and nine morphological traits; we also included samples collected in more distant allopatric sites. Clines in these traits were noncoincident, and there was no increase in either inbreeding coefficients or linkage disequilibrium at the centre of the zone. These results could suggest that the hybrid zone is maintained by weak selection against hybrids, conforming to either the tension zone or geographical selection‐gradient model. However, a closer examination of clines in pelage and microsatellites indicates that these clines are not sigmoid or stepped in shape but instead plateau at their centre. Sites within the hybrid zone also occur in a distinct habitat, characterized by greater seasonality in precipitation and lower seasonality in temperature. Together, these findings suggest that the hybrid zone may follow the bounded superiority model, with exogenous selection favouring hybrids within the transitional zone. These findings are noteworthy, as examples supporting the bounded superiority model are rare and may indicate a process of ecologically driven speciation without geographical isolation.     

Decline in lizard species diversity,abundance and ectoparasite load across an elevational gradient in the Australian alps

The rapid changes in altitude, and associated habitat, of mountain ecosystems make them ideal natural laboratories for testing the effect of environmental heterogeneity on species assemblage. Our understanding of the sensitivity of Australian reptiles to elevational clines is limited. We examined lizard distribution across three elevation zones (montane, subalpine and alpine), spanning from 900 to 1840 m above sea level, in the Australian alps. We aimed to examine how elevation influences species diversity and abundance, and ectoparasite load, and whether species alter their habitat use amongst different elevational zones. Active searches were conducted across the elevation zones to identify lizard community structure (at least 16 species) across elevational zones, along with skink habitat preferences and the ectoparasite load. Skink diversity and abundance were negatively correlated with increased elevation. The alpine zone had significantly lower diversity and abundance of skinks. Habitat use differed amongst both elevations and species. Ectoparasite prevalence was also significantly diminished in the alpine zone. Ectoparasites only infected a subset of the skink community, with ectoparasite load increasing as the active season progressed. This study provides evidence of the complex interplay between elevation and species diversity, as well as the differences in ectoparasite pressure along elevational gradients in the Australian alps.