Altitudinal and seasonal variation in microsatellite allele frequencies of Drosophila buzzatii

Variation in climate, particularly temperature, is known to affect the genetic composition of populations. Although there have been many studies of latitudinal variation, comparisons of populations across altitudes or seasons, particularly for animal species, are less common. Here, we study genetic variation (microsatellite markers) in populations of Drosophila buzzatii collected along altitudinal gradients and in different seasons. We found no differences in genetic variation between 2 years or between seasons within years. However, there were numerous cases of significant associations between allele frequencies or expected heterozygosities and altitude, with more than half showing nonlinear relationships. While these associations indicate possible selection and local altitudinal adaptation, direct tests gave strong evidence for selection affecting two loci and weaker evidence for five other loci. Two loci that are located within an inversion (including the one with strongest evidence for selection) show a linear increase in genetic diversity with altitude, likely due to thermal selection. Parallel associations with altitude here and with latitude in Australian populations indicate that selection is operating on chromosomal regions marked by some of the loci.     

A microsatellite linkage map for Drosophila montana shows large variation in recombination rates,and a courtship song trait maps to an area of low recombination

Current advances in genetic analysis are opening up our knowledge of the genetics of species differences, but challenges remain, particularly for out‐bred natural populations. We constructed a microsatellite‐based linkage map for two out‐bred lines of Drosophila montana derived from divergent populations by taking advantage of the Drosophila virilis genome and available cytological maps of both species. Although the placement of markers was quite consistent with cytological predictions, the map indicated large heterogeneity in recombination rates along chromosomes. We also performed a quantitative trait locus (QTL) analysis on a courtship song character (carrier frequency), which differs between populations and is subject to strong sexual selection. Linkage mapping yielded two significant QTLs, which explained 3% and 14% of the variation in carrier frequency, respectively. Interestingly, as in other recent studies of traits which can influence speciation, the strongest QTL mapped to a genomic region partly covered by an inversion polymorphism.     

The adaptive significance of colour pattern polymorphism in the Australian scincid lizard Lampropholis delicata

Females of Lampropholis delicata are dimorphic for colour pattern, the difference between morphs being the presence or absence of a distinct white mid-lateral stripe. A less distinct striped morph occurs also in males. We evaluated alternative hypotheses for the maintenance of this polymorphism by examining temporal and spatial variation in morph frequency, testing for differential selection among morphs using data on body size and reproductive traits from preserved specimens, and experimentally manipulating colour pattern in free-ranging lizards of both sexes, to assess the influence of the lateral stripe on survival rates. We found that the relative frequency of striped individuals varied among populations and decreased from north to south in both sexes, coincident with an increasing incidence of regenerated tails. Morph frequencies did not change through time within a population. Striped gravid females appeared to survive better and produced larger clutches than did non-striped females. In our experimental study, the relationship between survival and colour morph differed between the two sexes; males painted with a white lateral stripe had lower survival than control (brown stripe) males, but survival did not differ between striped and control females. The different response in the two sexes may be due partly to differences in temperature and microhabitat selection. We propose that the white lateral stripe decreases susceptibility to predators in gravid females but increases risk of predation in males, especially in combination with low temperatures. The polymorphism might be maintained by: (1) opposing fitness consequences of the stripe in males and females; (2) sex-specific habitat selection; and (3) gene flow in combination with spatial variation in relative fitness of the two morphs.     

Epistasis, phenotypic disequilibrium and contrasting associations with climate in the land snail Theba pisana

Hotter conditions favour effectively unbanded (EUB) shells in the snail Theba pisana. T. pisana is also polymorphic for colour of the shell's apex, determined by a pair of alleles at a locus linked to the banding locus. Apex colour is epistatic to shell banding, such that banded snails with a dark apex have darker bands. Annual censuses over 22 years across an ecotone between a sheltered Acacia thicket and open dune vegetation showed a persistent association of both EUB shells and pale apex with the Open habitat. The parallel variation was due partly to strong phenotypic disequilibrium, as the combination of EUB with dark apex was rare. Nevertheless, in fully banded shells the frequency of pale apex was also higher in the Open habitat, confirming independent, parallel associations of the two contributors to paleness. Within the Acacia habitat, temporal variation of the frequencies of banding morphs was much greater than for apex colour, and EUB shells were associated with hotter summers. Consistent with its primary effect only on the very small snails, apex colour did not vary with summer conditions, but instead, higher frequencies of pale apices were associated with sunnier winters. The intensity of selection was lower on apex colour than shell banding, due partly to the constraint of phenotypic disequilibrium. The shell traits in T. pisana are an example of complex responses to climatic variation, in which phenotypic disequilibrium constrains evolution of apex colour, but separate mechanisms of selection are evident.     

The effect of color polymorphism on mortality in the aphidMacrosiphoniella yomogicola

We examined body color polymorphism in the aphidMacrosiphoniella yomogicola from July to September 1993. We classified body color into eight types: green 1, green 2, red 1, red 2, white, orange, yellow and mist. The frequencies of body color varied with time and among patches of the host plant, yomogi (Artemisia spp.). Color diversity within a shoot was calculated using the Shannon diversity index. Of five usable data sets, three showed negative relationships between color diversity and mortality. The regression coefficients for two of these relationships were significant. No significant relationship between mortality and the number of aphids was found. The color diversity was not significantly related to a particular body color found on a yomogi shoot. Color polymorphism may be maintained because selection may favor a high color diversity on the host plant shoot.     

Studies of colour polymorphism in some marginal populations of the aposematic jersey tiger moth Callimorpha quadripunctaria

Callimorpha quadripunctaria (Lepidoptera, Arctiidae) is both genetically polymorphic for hindwing colour and well protected from predators. Polymorphic populations are, however, only found in the extreme northwest of its extensive distribution in Europe and Asia Minor. Use of a colour analyser confirmed the existence of three reasonably discrete colour classes: bright red, orange and yellow. These could each be subdivided into two further groupings of phenotypes, one of moths of pure colour and the other with a secondary colour. The species is polymorphic in SW England, where it became well established in villages and towns around the Exe estuary in the late 19th century. It now extends about 30 km east and west along the south coast of Devon and up to a similar distance inland along the river valleys. The rate of spread is consistent with the substantial dispersal of up to 500 m detected in a capture-recapture experiment performed at Exeter over the whole flight period in August 1984. This experiment also found no differences in timing of emergence, survivorship or within-habitat movement between the sexes or the colour classes. Males may show a higher rate of emigration than females. The species is polymorphic throughout S Devon. Its stronghold to the west of the Exe basin is characterized by a comparatively uniform morph frequency with 5045% red, 20–40% orange and 10–20% yellow. Yellow moths have their highest frequency in this area, red moths in the east and west and orange to the north. Adult activity is concentrated in the 2–3 h after dusk. Air temperatures of higher than 15°C are probably critical to pairing which involves sex pheromones released by females. This may be an important factor limiting the species' range. Our results suggest that climatic selection pressures involving non-visual differences in fitness account for the geographical variation. The lack of daytime activity of moths in comparison to that found by other workers on Rhodes and which is likely in other warmer climates suggests the hypothesis that expression of the polymorphism is favoured in the marginal populations of NW Europe by a relaxation of the stabilizing influence on the warning coloration exerted by visual predators.     

Role of cytogenetics for breeding management of squirrel monkey colonies

Breeding colonies of squirrel monkeys in zoos are often a mix of Saimiri taxa, which leads to the possibility of hybridization, loss of classification features, and possible reduction of fertility among hybrids. Two zoos, Potawatomi and Brookfield, were used to assess chromosomal composition of colonies. Chromosomes were analyzed from lymphocyte cultures and animals were classified into cytogenetic types. At both zoos there was a predominance of Peruvian types, but hybrids (Peruvian/Colombian) at Potawatomi and (Peruvian/Colombian and Peruvian/Guyanan) at Brookfield were also identified. Analysis of the Brookfield lineage (three generations) revealed karyotypic Peruvians derived from hybrid parents. The Gothic versus Roman arch of periocular white hair shows intermediate or quite variable expression in hybrids and offspring derived from hybrids. Given the relative length of both pericentric inversions and the segments distal to each, a detectable reduction in fertility is predicted when compared with similar-sized inversions in humans. Somatic segregation of inversion heterozygotes, which would reconstitute homozygosity and elevate fertility, was not detected.     

Multiple quantitative trait loci influence intra-specific variation in genital morphology between phylogenetically distinct lines of Drosophila montana

The evolution of animal genitalia has gained renewed interest because of their potential roles during sexual selection and early stages of species formation. Although central to understanding the evolutionary process, knowledge of the genetic basis of natural variation in genital morphology is limited to a very few species. Using an outbred cross between phylogenetically distinct lines of Drosophila montana, we characterized quantitative trait loci (QTLs) affecting the size and shape of the distiphallus, a prominent part of the male intromittent organ. Our microsatellite-based linkage analysis shows that intra-specific variation in the distiphallus involves several QTLs of largely additive effect and that a highly significant QTL co-localizes with the same inversion where we have earlier localized a large QTL for a sexually selected courtship song trait. The latter indicates that inversions can play an important role in shaping the evolution of rapidly evolving traits with a potential influence on speciation.     

Frequency dependent selection: homage to E. B. Poulton

Nearly a hundred years ago, E. B. Poulton (1884) set out clearly the argument for frequency-dependent selection by predators acting on non-mimetic polymorphic prey (i.e. for apostatic selection), anticipating many of the points made by later workers. We draw attention to his remarkable insight.     

Ecological genetic studies in the chromosomal form Mopti ofAnopheles gambiae s.str. in Mali,West Africa

Among the sibling species of the AfrotropicalAnopheles gambiae complex, the nominal taxon (An. gambiae s.str.) is the major malaria vector. Its bionomics suggest a man-dependent speciation process which involves, in West Africa, various incipient species chromosomally recognized by different combinations of 2R paracentric inversions. One of the most recent evolutionary steps of such a speciation process appears to be the chromosomal form Mopti, which is associated with dry season irrigation in arid zones, and is characterized by a remarkable ecological flexibility related to three 2R alternative arrangements, namelybc, u and +, whose expected karyotypes are found in Hardy-Weinberg equilibrium. The study of this chromosomal polymorphism in samples from a 16-locality transect in Mali shows wide variations and highly significant correlation with both temporal and spatial climatic differences. Mosquitoes homokaryotypic for 2Rbc are the actual dry season and arid areas breeders. The regular rise of 2Rbc frequency, up to fixation, during each dry season, corresponds to the South-North clinal increase of the same arrangement along the transect, from about 30% in the humid savanna to near fixation in the South-Saharan zone. This coherent ecological genetics case provides full support to the hypothesis of the adaptive nature of paracentric inversions. Moreover, the very peculiar system of combinations of contiguous 2R inversions, utilized by Mopti as well as by other chromosomal forms ofAn. gambiae, suggests a process of polygenic reorganization based on linkage disequilibria and involving the inversions as driving selection units.