The genome of the blind bee louse fly reveals deep convergences with its social host and illuminates Drosophila origins.

Social insects' nests harbor intruders known as inquilines,1 which are usually related to their hosts.2,3 However, distant non-social inquilines may also show convergences with their hosts,4,5 although the underlying genomic changes remain unclear. We analyzed the genome of the wingless and blind bee louse fly Braula coeca, an inquiline kleptoparasite of the western honey bee, Apis mellifera.6,7 Using large phylogenomic data, we confirmed recent accounts that the bee louse fly is a drosophilid8,9 and showed that it had likely evolved from a sap-breeder ancestor associated with honeydew and scale insects' wax. Unlike many parasites, the bee louse fly genome did not show significant erosion or strict reliance on an endosymbiont, likely due to a relatively recent age of inquilinism. However, we observed a horizontal transfer of a transposon and a striking parallel evolution in a set of gene families between the honey bee and the bee louse fly. Convergences included genes potentially involved in metabolism and immunity and the loss of nearly all bitter-tasting gustatory receptors, in agreement with life in a protective nest and a diet of honey, pollen, and beeswax. Vision and odorant receptor genes also exhibited rapid losses. Only genes whose orthologs in the closely related Drosophila melanogaster respond to honey bee pheromone components or floral aroma were retained, whereas the losses included orthologous receptors responsive to the anti-ovarian honey bee queen pheromones. Hence, deep genomic convergences can underlie major phenotypic transitions during the evolution of inquilinism between non-social parasites and their social hosts.

Genetic characterization of the honeybee ectoparasitic mite Varroa destructor from Benin (West Africa) using mitochondrial and microsatellite markers.

Varroa destructor is one of the scourges of global beekeeping. It was detected for the first time in Benin in 2011 on the honeybee Apis mellifera adansonii. The aim of this study was to identify the strain of Varroa sp. found and study its genetic diversity. In total 183 Varroa mites were sampled in 21 municipalities in Benin. The COI intergenic region of each mite mtDNA was amplified by PCR. The SacI restriction enzyme was used to determine the strains of Varroa sp. Only the Korean (K) haplotype, identical to the most prevalent strain in Africa, was detected. Analysis of the genetic diversity of Varroa mites with eight microsatellite loci (Simple Sequence Repeats) indicated a very low diversity of genotypes. Thus, V. destructor populations from Benin appear to make up a single group. Their clonal wealth ranges from 0.00 to 0.47. This study is an important step forward in the monitoring of the infestation of V. destructor.

MtDNA COI-COII marker and drone congregation area: an efficient method to establish and monitor honeybee (Apis mellifera L.) conservation centres.

Honeybee subspecies have been affected by human activities in Europe over the past few decades. One such example is the importation of nonlocal subspecies of bees which has had an adverse impact on the geographical repartition and subsequently on the genetic diversity of the black honeybee Apis mellifera mellifera. To restore the original diversity of this local honeybee subspecies, different conservation centres were set up in Europe. In this study, we established a black honeybee conservation centre Conservatoire de l'Abeille Noire d'Ile de France (CANIF) in the region of Ile-de-France, France. CANIF's honeybee colonies were intensively studied over a 3-year period. This study included a drone congregation area (DCA) located in the conservation centre. MtDNA COI-COII marker was used to evaluate the genetic diversity of CANIF's honeybee populations and the drones found and collected from the DCA. The same marker (mtDNA) was used to estimate the interactions and the haplotype frequency between CANIF's honeybee populations and 10 surrounding honeybee apiaries located outside of the CANIF. Our results indicate that the colonies of the conservation centre and the drones of the DCA show similar stable profiles compared to the surrounding populations with lower level of introgression. The mtDNA marker used on both DCA and colonies of the conservation centre seems to be an efficient approach to monitor and maintain the genetic diversity of the protected honeybee populations.

A fifth major genetic group among honeybees revealed in Syria.

BACKGROUND: Apiculture has been practiced in North Africa and the Middle-East from antiquity. Several thousand years of selective breeding have left a mosaic of Apis mellifera subspecies in the Middle-East, many uniquely adapted and survived to local environmental conditions. In this study we explore the genetic diversity of A. mellifera from Syria (n = 1258), Lebanon (n = 169) and Iraq (n = 35) based on 14 short tandem repeat (STR) loci in the context of reference populations from throughout the Old World (n = 732). RESULTS: Our data suggest that the Syrian honeybee Apis mellifera syriaca occurs in both Syrian and Lebanese territories, with no significant genetic variability between respective populations from Syria and Lebanon. All studied populations clustered within a new fifth independent nuclear cluster, congruent with an mtDNA Z haplotype identified in a previous study. Syrian honeybee populations are not associated with Oriental lineage O, except for sporadic introgression into some populations close to the Turkish and Iraqi borders. Southern Syrian and Lebanese populations demonstrated high levels of genetic diversity compared to the northern populations. CONCLUSION: This study revealed the effects of foreign queen importations on Syrian bee populations, especially for the region of Tartus, where extensive introgression of A. m. anatolica and/or A. m. caucasica alleles were identified. The policy of creating genetic conservation centers for the Syrian subspecies should take into consideration the influence of the oriental lineage O from the northern Syrian border and the large population of genetically divergent indigenous honeybees located in southern Syria.

Thermal phenotypic plasticity of body size in Drosophila melanogaster: sexual dimorphism and genetic correlations.

Thirty isofemale lines collected in three different years from the same wild French population were grown at seven different temperatures (12-31 °C). Two linear measures, wing and thorax length, were taken on 10 females and 10 males of each line at each temperature, also enabling the calculation of the wing/thorax (W/T) ratio, a shape index related to wing loading. Genetic correlations were calculated using family means. The W-T correlation was independent of temperature and on average, 0.75. For each line, characteristic values of the temperature reaction norm were calculated, i.e. maximum value, temperature of maximum value and curvature. Significant negative correlations were found between curvature and maximum value or temperature of maximum value. Sexual dimorphism was analysed by considering either the correlation between sexes or the female/male ratio. Female-male correlation was on average 0.75 at the within line, within temperature level but increased up to 0.90 when all temperatures were averaged for each line. The female-male ratio was genetically variable among lines but without any temperature effect. For the female/male ratio, heritability (intraclass correlation) was about 0.20 and evolvability (genetic coefficient of variation) close to 1. Although significant, these values are much less than for the traits themselves. Phenotypic plasticity of sexual dimorphism revealed very similar reaction norms for wing and thorax length, i.e. a monotonically increasing sigmoid curve from about 1.11 up to 1.17. This shows that the males are more sensitive to a thermal increase than females. In contrast, the W/T ratio was almost identical in both sexes, with only a very slight temperature effect.

Sexual dimorphism of body size and sternopleural bristle number: a comparison of geographic populations of an invasive cosmopolitan drosophilid.

Zaprionus indianus is a cosmopolitan drosophilid, of Afrotropical origin, which has recently colonized South America. The sexual dimorphism (SD) of body size is low, males being almost as big as females. We investigated 10 natural populations, 5 from America and 5 from Africa, using the isofemale line technique. Three traits were measured on each fly: wing and thorax length and sternopleural bristle number. Two indices of SD were compared, and found to be highly correlated (r > 0.99). For the sake of simplicity, only the female/male (F/M) ratio was further considered. A significant genetic variability of SD was found in all cases, although with a low heritability (intra-class correlation of 0.13), about half the value found for the traits themselves. For size SD, we did not find any variation among continents or any latitudinal trend, and average values were 1.02 for wing length and 1.01 for thorax length. Bristle number SD was much greater (1.07). Among mass laboratory strains, SD was genetically much more variable than in recently collected populations, a likely consequence of laboratory drift. Altogether, SD, although genetically variable and prone to laboratory drift, is independent of size variations and presumably submitted to a stabilizing selection in nature.

Phenotypic plasticity of body size in a temperate population of Drosophila melanogaster: when the temperature-size rule does not apply.

A natural population of Drosophila melanogaster in southern France was sampled in three different years and 10 isofemale lines were investigated from each sample. Two size-related traits, wing and thorax length, were measured and the wing/thorax ratio was also calculated. Phenotypic plasticity was analysed after development at seven different constant temperatures, ranging from 12 degrees C to 31 degrees C. The three year samples exhibited similar reaction norms, suggesting a stable genetic architecture in the natural population. The whole sample (30 lines) was used to determine precisely the shape of each reaction norm, using a derivative analysis. The practical conclusion was that polynomial adjustments could be used in all cases, but with different degrees: linear for the wing/thorax ratio, quadratic for thorax length, and cubic for wing length. Both wing and thorax length exhibited concave reaction norms, with a maximum within the viable thermal range. The temperatures of the maxima were, however, quite different, around 15 degrees C for the wing and 19.5 degrees C for the thorax. Assuming that thorax length is a better estimate of body size, it is not possible to state that increasing the temperature results in monotonically decreasing size (the temperature-size rule), although this is often seen to be the case for genetic variations in latitudinal clines. The variability of the traits was investigated at two levels-within and between lines-and expressed as a coefficient of variation. The within-line (environmental) variability revealed a regular, quadratic convex reaction norm for the three traits, with a minimum around 21 degrees C. This temperature of minimum variability may be considered as a physiological optimum, while extreme temperatures are stressful. The between-line (genetic) variability could also be adjusted to quadratic polynomials, but the curvature parameters were not significant. Our results show that the mean values of the traits and their variance are both plastic, but react in different ways along a temperature gradient. Extreme low or high temperatures decrease the size but increase the variability. These effects may be considered as a functional response to environmental stress.

REML estimates of genetic parameters of sexual dimorphism for wing and thorax length in Drosophila melanogaster.

Restricted maximum likelihood was used to estimate genetic parameters of male and female wing and thorax length in isofemale lines of Drosophila melanogaster, and results compared to estimates obtained earlier with the classical analysis of variance approach. As parents within an isofemale line were unknown, a total of 500 parental pedigrees were simulated and mean estimates computed. Full and half sibs were distinguished, in contrast to usual isofemale studies in which animals were all treated as half sibs and hence heritability was overestimated. Heritability was thus estimated at 0.33, 0.38, 0.30 and 0.33 for male and female wing length and male and female thorax length, respectively, whereas corresponding estimates obtained using analysis of variance were 0.46, 0.54, 0.35 and 0.38. Genetic correlations between male and female traits were 0.85 and 0.62 for wing and thorax length, respectively. Sexual dimorphism and the ratio of female to male traits were moderately heritable (0.30 and 0.23 for wing length, 0.38 and 0.23 for thorax length). Both were moderately and positively correlated with female traits, and weakly and negatively correlated with male traits. Such heritabilities confirmed that sexual dimorphism might be a fast evolving trait in Drosophila.

Genetic variability of sexual size dimorphism in a natural population of Drosophila melanogaster: an isofemale-line approach.

Most animal species exhibit sexual size dimorphism (SSD). SSD is a trait difficult to quantify for genetical purposes since it must be simultaneously measured on two kinds of individuals, and it is generally expressed either as a difference or as a ratio between sexes. Here we ask two related questions: What is the best way to describe SSD, and is it possible to conveniently demonstrate its genetic variability in a natural population? We show that a simple experimental design, the isofemale-line technique (full-sib families), may provide an estimate of genetic variability, using the coefficient of intraclass correlation. We consider two SSD indices, the female-male difference and the female/male ratio. For two size-related traits, wing and thorax length, we found that both SSD indices were normally distributed. Within each family, the variability of SSD was estimated by considering individual values in one sex (the female) with respect to the mean value in the other sex (the male). In a homogeneous sample of 30 lines of Drosophila melanogaster, both indices provided similar intraclass correlations, on average 0.21, significantly greater than zero but lower than those for the traits themselves: 0.50 and 0.36 for wing and thorax length respectively. Wing and thorax length were strongly positively correlated within each sex. SSD indices of wing and thorax length were also positively correlated, but to a lesser degree than for the traits themselves. For comparative evolutionary studies, the ratio between sexes seems a better index of SSD since it avoids scaling effects among populations or species, permits comparisons between different traits, and has an unambiguous biological significance. In the case of D. melanogaster grown at 25 degrees C, the average female/male ratios are very similar for the wing (1.16) and the thorax (1.15), and indicate that, on average, these size traits are 15-16% longer in females.

Allometric and nonallometric components of Drosophila wing shape respond differently to developmental temperature.

Phenotypic plasticity of wing size and shape of Drosophila simulans was analyzed across the entire range of viable developmental temperatures with Procrustes geometric morphometric method. In agreement with previous studies, size clearly decreases when temperature increases. Wing shape variation was decomposed into its allometric (24%) and nonallometric (76%) components, and both were shown to involve landmarks located throughout the entire wing blade. The allometric component basically revealed a progressive, monotonous variation along the temperature. Surprisingly, nonallometric shape changes were highly similar at both extremes of the thermal range, suggesting that stress, rather than temperature per se, is the key developmental factor affecting wing shape.