Behavioral reproductive isolation inDrosophila silvestris,D. heteroneura,and their F1 hybrids (Diptera: Drosophilidae)

We investigated the role that courtship and aggressive interactions may have for the maintenance of reproductive isolation betweenDrosophila silvestris andD. heteroneura. We examined the behavioral bases of reproductive isolation between the parental species and we examined the courtship success of each sex of both reciprocal F₁ hybrids when paired with the parental species. We found reduced copulation success among heterotypic parental pairs compared to homotypic pairs, which was primarily due to the lack of courtship initiation betweenD. silvestris males andD. heteroneura females. When hybrid males from both reciprocal crosses were paired with parental females their copulation successes were not significantly different from that of parental males. In contrast, hybrid females from both crosses had reduced copulation success withD. silvestris males, which in turn was primarily due to a reduced success of reaching later stages of courtship. The time spent in copulation by hybrid males was intermediate between the two parental males. We studied aggression by observing the interactions of males of heterotypic pairs, both between the parental species and between the hybrids and parental males. A lack of aggressive interactions betweenD. silvestris males andD. heteroneura males in addition to the lack of courtship suggests thatD. silvestris males do not respond toD. heteroneura individuals of either sex. Hybrid males were equally successful in winning fights with bothD. silvestris andD. heteroneura males. These results indicate that the behavioral isolation betweenD. silvestris andD. heteroneura may be largely a consequence of the earliest stages of interactions. The two species may differ either in activity levels or in morphological or chemical traits that are important for species and mate recognition. The relatively high copulation and aggressive success of hybrids indicates that sexual selection against hybrids alone is unlikely to be a sufficient force to reduce gene flow and maintain species distinctions.     

INTERSPECIFIC HYBRIDS OF DROSOPHILA HETERONEURA AND D. SILVESTRIS I. COURTSHIP SUCCESS

Drosophila heteroneura and D. silvestris are well-defined, sympatric species of the planitibia subgroup of Hawaiian Drosophila. D. silvestris can be subdivided into two allopatric morphotypes that differ in the number of bristle rows on the front tibia (two rows versus three rows). We measured courtship success of intraspecific and interspecific hybrids as the proportion of females inseminated during a two-week period with a single sib male. Proportions were arcsin-transformed so that the values were asymptotically normal in distribution, and tests of homogeneity and of mean differences were performed. Of key importance is the discovery of genetic variation for the proportion of inseminated females within both D. heteroneura and D. silvestris. The interspecific crosses and the D. silvestris intraspecific crosses also provide evidence for a coadapted gene complex with some dominance or heterosis. This coadapted gene complex correlates with the morphotypes of these flies, rather than with the D. heteroneura/D. silvestris contrasts per se. This observation stresses the importance of recognizing both behavioral and morphological components of the mate-recognition system. The incompatible coadaptation that separates the two-row from the three-row forms also supports recent molecular studies which indicate that the three-row form split from the two-row form prior to the split between D. heteroneura and two-row D. silvestris. The observations of intraspecific variability and coadaptation support the predictions of a genetic-transilience model which explains the origin of a new mate-recognition system in terms of sexual selection in the context of a founder-flush event.     

Sexual Selection, Epistasis and Species Boundaries in Sympatric Hawaiian Picture-winged Drosophila

The Hawaiian picture-winged flies in the genus Drosophila are a spectacular example of rapid evolutionary diversification in which sexual selection is considered an important mechanism for reproductive isolation and speciation. We investigated the behavioral reproductive isolation of two closely related and sympatric Hawaiian picture-winged Drosophila species, D. silvestris and D. heteroneura, which are known to hybridize in nature and produce viable and fertile hybrids. We compared the mating success of parental, F1 and backcross males in pairings with D. heteroneura females. The F1 males were produced by mating D. heteroneura males with D. silvestris females, and the backcross males were produced by mating F1 females with D. heteroneura males. The mating success of backcross males paired with D. heteroneura females were significantly reduced relative to that of parental and F1 males. This reduced mating success occurred primarily at a late stage of courtship where female choice of mate may be important. Two- and three-gene models demonstrate that epistasis involving a few genes could account for the observed variation in male mating success. These results are consistent with negative epistasis in the backcross generation and support the importance of sexual selection and negative epistasis in the evolution and maintenance of these species.     

ON THE ORIGIN OF INCIPIENT REPRODUCTIVE ISOLATION: THE CASE OF DROSOPHILA ALBOMICANS AND D. NASUTA

The nasuta subgroup of Drosophila consists of 12 known species classified within the immigrans group. D. nasuta and D. albomicans are two sibling species widely distributed throughout the Indo-Pacific tropics, which, although morphologically indistinguishable, have different meta-phase-chromosome configurations: chromosomes X and 3 are attached in D. albomicans, so that about 60% of its genes are sex-linked. Our experiments show that, at least in the laboratory, there is no sexual, mechanical, or gametic isolation between the two species. There is, however, hybrid “breakdown” expressed in three ways: 1) reduction in the number of F₂ hybrids produced per culture; 2) reduction in the fertility of F₂ (males) and F₃ (males and females) hybrid progenies; and 3) abnormal sex ratios in the progenies of crosses between strains of certain localities. In experimental populations, the karyotypes of both species are still present in substantial frequencies after 20 generations, although the frequencies of the two karyotypes vary depending on the geographic origin of the strains. Our results support the hypothesis that, in allopatry, the evolution of postzygotic isolation precedes that of prezygotic isolation. The mtDNA is polymorphic in both D. nasuta and D. albomicans and fairly similar between them. Assuming typical rates of mtDNA evolution, the two species would have diverged from each other about 500,000 years ago, whereas the African and Indian populations of D. nasuta (considered to be different subspecies by some authors) might have diverged some 350,000 years ago.     

Hybridization and sequential components of reproductive isolation between parapatric walnut‐infesting sister species Rhagoletis completa and Rhagoletis zoqui

Hybridization can provide a window into how populations diverge to form new species. Here, we confirm hybridization between Rhagoletis completa Cresson, 1929 and Rhagoletis zoqui Bush, 1966, two species of walnut husk‐infesting flies that geographically overlap in a narrow area of parapatry in Northeastern Mexico. Rhagoletis completa and R. zoqui are members of a species group (Rhagoletis suavis) that has been hypothesized to speciate in allopatry. Sexual selection has been argued to be a potentially important factor for generating pre‐mating isolation among walnut husk flies, because of the differences in wing morphology and body coloration, and the existence of sexual dimorphism within species. However, there was no evidence for pre‐mating isolation between R. completa and R. zoqui, based on choice and no‐choice mating experiments conducted on adults of fly populations outside the contact zone. There was also no support for reduced fertility of hybrid matings or for F₁ inviability; however, F₁ hybrids appeared to have lower fertility and F₂ offspring have reduced survivorship. Postzygotic isolation in later generation hybrids of mixed ancestry therefore appears to be the first intrinsic barrier to gene flow evolving between R. completa and R. zoqui. We discuss the implications of our results for allopatric speciation in walnut flies and the potential evolutionary fate of R. zoqui and R. completa if they were to come into broad geographic contact in the future. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Evolutionary distances in hawaiian drosophila measured by DNA reassociation

Summary Comparisons of the sequence divergence of three species of Hawaiian Drosophila have been made by hybridization of single-copy tracer DNA of each of the species with driver DNA from each species, and measurement of the average melting temperature (Tma) in a chaotropic solvent (2.4 M tetraethylammonium chloride) which minimizes differences due to base composition. Correction was made for the length of hybrid duplex regions to obtain the reduction in thermal stability due to divergence.An accuracy of ± 0.2°C was achieved and the mean reduction in Tm for hybridization betweenD. heteroneura andD. silvestris (found only on the island of Hawaii) was 0.55°C and betweenD. picticornis, found only on the island of Kauai, and the other two species was 2.13°C. The rate of DNA change is estimated to be between 0.2 and 0.4%/My by assuming that theD. heteroneura-D. silvestris divergence occurred 0.8 My ago and the divergence between these species andD. picticornis between 4 and 6 My ago.The general single copy DNA sequence divergence appears to be very much greater than the minimal coding region sequence divergence previously estimated from allozyme studies.     

THE GENETIC BASIS OF SEXUAL ISOLATION BETWEEN DROSOPHILA MELANOGASTER AND D. SIMULANS

The genetic analysis of sexual isolation between the closely-related species Drosophila melanogaster and Drosophila simulans involved two experiments with no-choice tests. The efficiency of sexual isolation was measured by the frequency of courtship initiation and interspecific mating. We first surveyed the variation in sexual isolation between D. melanogaster strains and D. simulans strains of different geographic origin. Then, to investigate variation in sexual isolation within strains, we made F₁ diallel sets of reciprocal crosses within strains of D. melanogaster and D. simulans. The F₁ diallel progeny of one sex were paired with the opposite sex of the other species. The first experiment showed significant differences in the frequency of interspecific mating between geographic strains. There were more matings between D. simulans females and D. melanogaster males than between D. melanogaster females and D. simulans males. The second experiment uncovered that the male genotypes in the D. melanogaster diallel significantly differed in interspecific mating frequency, but not in courtship initiation frequency. The female genotypes in the D. simulans diallel were not significantly different in courtship initiation and interspecific mating frequency. Genetic analysis reveals that in D. melanogaster males sexual isolation was not affected by either maternal cytoplasmic effects, sex-linked effects, or epistatic interaction. The main genetic components were directional dominance and overdominance. The F₁ males achieved more matings with D. simulans females than the inbred males. The genetic architecture of sexual isolation in D. melanogaster males argues for a history of weak or no selection for lower interspecific mating propensity. The behavioral causes of variation in sexual isolation between the two species are discussed.     

QUANTITATIVE GENETIC ANALYSIS OF DIVERGENCE IN MALE SECONDARY SEXUAL TRAITS BETWEEN DROSOPHILA SIMULANS AND DROSOPHILA MAURITIANA

The sibling species Drosophila simulans and D. mauritiana differ significantly in a number of male secondary sexual traits, providing an ideal system for genetic analysis of interspecific morphological divergence. In the experiment reported here, F₁ hybrids from a cross of two inbred lines were backcrossed in both directions and about 200 flies from each backcross were scored for several traits (bristle numbers and cuticle areas), as well as 18 markers distributed throughout the genome. Each trait was analyzed by composite interval mapping to identify quantitative trait loci (QTL) and estimate their effects. For each trait, from one to eight loci were detected, with more divergent traits showing evidence for greater numbers of QTL. Estimates of additive effects varied widely, with a range of 0.4 to 4.1 environmental standard deviation units and an average of 2.2 units. There was substantial evidence for nonadditive effects, since the magnitude of estimates often differed significantly between the two backcrosses. The sign of the estimated effect differed among QTL for bristle traits, but not for cuticle area traits, suggesting that these two types of trait may have undergone different types of selection. Finally, several similarities were found between different traits in the estimated positions of QTL, suggesting that pleiotropy and/or linkage of QTL may have been important in the evolution of these traits.     

Evolution of the transposable element Uhu in five species of Hawaiian Drosophila

The complete DNA sequence of three independent isolates of Uhu, a member of the Tc1-like class of transposable elements from D. heteroneura (Uhu-1, Uhu-3, and Uhu-4), has been determined. These isolates have between 95 and 96.4% nucleotide sequence identity indicating that Uhu is well conserved within this species. A comparison of the DNA sequences of Uhu and the D. melanogaster Hb1 transposable element shows that the nucleotide substitution rate for Uhu is comparable to the synonymous rate for the Adh gene in these species. Uhu has been identified in four other species of endemic Hawaiian Drosophila, D. silvestris, D. differens, D. planitibia and D. picticornis, and nine Uhu elements were isolated from genomic libraries of these four species. A 444 base pair region from within the coding region of the Uhu element, with well conserved ends, was amplified by the polymerase chain reaction and used for sequence comparison of elements from different species. The analysis of the sequence similarities between the elements within and between the species shows a grouping of the two pairs of most closely related species (D. heteroneura and D. silvestris, and D. differens and D. planitibia), but shows a much larger variation within the most recently diverged species (D. heteroneura and D. silvestris) than expected. There are extensive nucleotide substitutions and deletions in the Uhu elements from D. picticornis showing that they are degenerating and being lost in this species. These observations indicate that the Uhu element has been transmitted vertically and that transposition may have been activated at the time of formation of each species as it colonized the newly formed islands of the Hawaiian archipelago.     

Genetic Variation in Hawaiian Drosophila. IV. Allozymic Similarity between D. SILVESTRIS and D. HETERONEURA from the Island of Hawaii

The species are endemic to the newest island in the archipelago and are broadly sympatric. They are easily distinguished morphologically in both sexes. Using standard electrophoretic procedures, we have examined 25 loci encoding for structural proteins from 539 silvestris and 325 heteroneura collected at three widely-separated localities where the two species are sympatric. Pairwise comparisons within and between the species show the following coefficients of genetic identity (Nei''s I): within silvestris, 0.961 ± 0.01; within heteroneura, 0.949 ± 0.02; between silvestris and heteroneura, 0.939 ± 0.01. Neither the differences within nor between the species are significant. There are no fixed allelic differences either within or between the species. At the three areas of sympatry, the species show gene frequency differences (P < 0.05) at 9, 11 and 13 loci respectively. This is not much different from the variation within either one of the species across the three localities. The two species have similar heterozygosity ( H) levels (silvestris, 0.083; heteroneura, 0.089) and percent of polymorphic loci (both 0.37). It is suggested that despite their morphological divergence, these species are much more newly formed than classical sibling species. Significant allozymic differences may not have had time to accumulate.     

CYTOLOGICAL CHARACTERIZATION OF PREMEIOTIC VERSUS POSTMEIOTIC DEFECTS PRODUCING HYBRID MALE STERILITY AMONG SIBLING SPECIES OF THE DROSOPHILA MELANOGASTER COMPLEX

In accordance with Haldane's rule, hybridizations between species of the Drosophila simulans clade produce fertile females but sterile males. In this study, a comprehensive characterization was undertaken on the six types of F₁ males that were the result of the crosses between D. simulans, D. sechellia, and D. mauritiana. With the use of light and electron microscopy, it was shown that while each particular hybrid genotype exhibited a specific sterility phenotype, these phenotypes fell into two distinct classes. The two hybrid genotypes that possessed D. mauritiana X-chromosomes contained spermatogenic defects that caused arrests in premeiotic spermatogenic stages. The other four F₁ hybrids possessed postmeiotic spermatogenic defects. Nonsynchronous cell divisions, underdeveloped mitochondrial derivative-axonemal associations, and microtubule abnormalities were common to all of these hybrids. Each particular postmeiotically defective hybrid genotype demonstrated characteristically distinct profiles in sperm bundle number in addition to characteristic spermiogenic arrests in the furthest developed spermatids. These results in species hybrids contrast with the absence of significant differences in spermatogenic characters between species of this clade. In addition, by utilizing an attached-X cross, we investigated the influence of maternal effects and cytoplasmic factors on the sterility of D. simulans F₁ hybrids and found none. However, we discovered a strain of D. simulans (2119) that caused a large shift in sterility from postmeiotic to premeiotic when crossed to D. sechellia. This suggests that D. simulans is polymorphic for genes involving premeiotic and postmeiotic sterility and that the two types of sterilities between species may have a simple genetic basis.     

Geographic patterns of postzygotic isolation between two closely related widespread dung fly species (Sepsis cynipsea and Sepsis neocynipsea; Diptera: Sepsidae)

Identifying the contribution of pre‐ and postzygotic barriers to gene flow is a key goal of speciation research. The widespread dung fly species Sepsis cynipsea and Sepsis neocynipsea offer great potential for studying the speciation process over a range of opportunities for gene exchange within and across sister species (cross‐continental allopatry, continental parapatry and sympatry). We examined the role of postcopulatory isolating barriers by comparing female fecundity and egg‐to‐adult viability of F₁ and F₂ hybrids, as well as backcrosses of F₁ hybrids with the parental species, via replicated crosses of sym‐, para‐ and allopatric populations. Egg‐to‐adult viability was strongly but not totally suppressed in hybrids, and offspring production approached nil in the F₂ generation (hybrid breakdown), indicating yet unspecified intrinsic incompatibilities. Viable F₁ hybrid offspring showed almost absolute male (the heterogametic sex) sterility while females remained largely fertile, in accordance with Haldane's rule. Hybridization between the two species in European areas of sympatry (Swiss Alps) indicated only minor reinforcement based on fecundity traits. Crossing geographically isolated European and North American S. neocynipsea showed similar albeit weaker isolating barriers that are most easily explained by random genetic drift. We conclude that in this system with a biogeographic continuum of reproductive barriers, speciation is mediated primarily by genetic drift following dispersal of flies over a wide (allopatric) geographic range, with some role of natural or sexual selection in incidental or direct reinforcement of incompatibility mechanisms in areas of European sympatry. S(ubs)pecies status of continental S. neocynipsea appears warranted.     

SEXUAL SELECTION AGAINST HYBRIDS BETWEEN SYMPATRIC STICKLEBACK SPECIES: EVIDENCE FROM A FIELD EXPERIMENT

Sexual selection against viable, fertile hybrids may contribute to reproductive isolation between recently diverged species. If so, then sexual selection may be implicated in the speciation process. Laboratory measures of the mating success of hybrids may underestimate the amount of sexual selection against them if selection pressures are habitat specific. Male F₁ hybrids between sympatric benthic and limnetic sticklebacks (Gasterosteus aculeatus complex) do not suffer a mating disadvantage when tested in the laboratory. However, in the wild males choose different microhabitats and parental females tend to be found in the same habitats as conspecific males. This sets up the opportunity for sexual selection against male hybrids because they must compete with parental males for access to parental females. To test for sexual selection against adult F₁ hybrid males, we examined their mating success in enclosures in their preferred habitat (open, unvegetated substrate) where limnetic males and females also predominate. We found significantly reduced mating success in F₁ hybrid males compared with limnetic males. Thus, sexual selection, like other mechanisms of postzygotic isolation between young sister species, may be stronger in a wild setting than in the laboratory because of habitat-specific selection pressures. Our results are consistent with, but do not confirm, a role for sexual selection in stickleback speciation.     

Interspecific hybrids of Antheraea roylei and A. pernyi — a cytogenetic reassessment

Summary A rare case of interspecific hybridization between the Indian oak feeding silkworm Antheraea roylei (n=31) and Chinese oak feeding silkworm A. pernyi (n=49) yielding fertile and vigorous offspring is reported. The F₁ and the backcross (A. roylei X A. pernyi X A. pernyi male individuals of the above cross and the F₂₃ and F₃₂ male offspring derived from an earlier cross between another race of A. roylei (n=30) and A. pernyi (n=49) were cytogenetically analysed in order to study their chromosome dynamics. The F₁ hybrids showed 18 trivalents and 13 bivalents in the first meiotic prophase and metaphase. The backcross individuals possessed either 9 trivalents and 31 bivalents or 49 bivalents, in Metaphase I cells. The F₂₃ and F₃₂ individuals were karyotypically alike and exhibited 49 bivalents. The following conclusions were drawn from the above observations: (a) in spite of allopatry and karyotypic divergence in number, a high degree of homology exists between the chromosomal complements of the two species; (b) A. pernyi possibly evolved from A. roylei, during the course of which 18 chromosomes of the latter underwent fission to give rise to the 36 chromosomes of the former. This is demonstrated by trivalent formation and pairing affinities in F₁ hybrids; (c) selection has favoured the elimination of large A. roylei chromosomes which participated in trivalent formation in successive generations of inbred hybrids to establish a stable Karyotype like that of A. pernyi.     

Sexual isolation between two sibling species with overlapping ranges: Drosophila santomea and Drosophila yakuba

Abstract.— .Drosophila yakuba is widespread in Africa, whereas D. santomea, its newly discovered sister species, is endemic to the volcanic island of São Tomé in the Gulf of Guinea. Drosophila santomea probably formed after colonization of the island by a D. yakuba‐like ancestor. The species presently have overlapping ranges on the mountain Pico do São Tome, with some hybridization occurring in this region. Sexual isolation between the species is uniformly high regardless of the source of the populations, and, as in many pairs of Drosophila species, is asymmetrical, so that hybridizations occur much more readily in one direction than the other. Despite the fact that these species meet many of the conditions required for the evolution of reinforcement (the elevation of sexual isolation by natural selection to avoid maladaptive interspecific hybridization), there is no evidence that sexual isolation between the species is highest in the zone of overlap. Sexual isolation is due to evolutionary changes in both female preference for heterospecific males and in the vigor with which males court heterospecific females. Heterospecific matings are also slower to take place than are homospecific matings, constituting another possible form of reproductive isolation. Genetic studies show that, when tested with females of either species, male hybrids having a D. santomea X chromosome mate much less frequently with females of either species than do males having a D. yakuba X chromosome, suggesting that the interaction between the D. santomea X chromosome and the D. yakuba genome causes behavioral sterility. Hybrid F₁ females mate readily with males of either species, so that sexual isolation in this sex is completely recessive, a phenomenon seen in other Drosophila species. There has also been significant evolutionary change in the duration of copulation between these species; this difference involves genetic changes in both sexes, with at least two genes responsible in males and at least one in females.     

Clarification of the genetic component of hybrids between Phyllodoce caerulea and Phyllodoce aleutica (Ericaceae) in Hokkaido,northern Japan

Natural hybridization provides great opportunities to understand the interaction of genetics and ecology in determining species boundaries. We examined the genetic relationships of Phyllodoce taxa and revealed that most hybrids were fertile F₁s and an extremely small number of backcross and no F₂ plants were established in natural conditions. Because this trend was irrespective of regions, we conclude that negative endogenous selection may act after the germination of F₁ seeds and prevent the establishment of later‐generation hybrids. Based on these results, we discuss the ecological and evolutionary significance of natural hybridization in Phyllodoce taxa.     

GENETIC VARIATION IN A COURTSHIP-RELATED MALE CHARACTER IN DROSOPHILA SILVESTRIS FROM A SINGLE HAWAIIAN LOCALITY

The large Drosophila species of Hawaii display sexual dimorphism and elaborate species-specific courtship patterns. Male characters related to courtship attract particular attention since they frequently constitute the most conspicuous taxonomic differences between Hawaiian species. The present study concerns intraspecific genetic variation in a courtship-related male character. D. silvestris, endemic to the geologically new volcanoes of Hawaii island, displays a brush of large dorsal cilia on the tibia of the foreleg of the male fly. This is used to stimulate the female during courtship. Genetic variation between local populations has previously been shown to exist for both sexual behavior and cilia number. The present paper reports the results of a study of cilia number variation in males collected at a single site over the period 1976-1980. Male progeny of females captured in nature were also studied. The mean number of cilia in the natural population was stable except for 1979 when it rose significantly, falling back again in 1980. The data indicate the existence of ample genetic variance for this character, existing as a balanced polymorphism in the natural population. The character appears to be under stabilizing selection. It is hypothesized that sexual selection is a contributing factor.     

GENETICS OF SEXUAL ISOLATION AND COURTSHIP DYSFUNCTION IN MALE HYBRIDS OF DROSOPHILA PSEUDOOBSCURA AND DROSOPHILA PERSIMILIS

Despite the importance of sexual isolation to speciation, few studies have analyzed the genetic basis of interspecific mating discrimination, particularly using hybrid males. In this study, I investigated the genetic basis of sexual isolation using male hybrids of Drosophila pseudoobscura and D. persimilis. Hybrid male mating success was caused by interactions between the X-chromosome and autosomes (or Y-chromosome), and different arms of the X-chromosome contributed to mating success with females of each species. Further, although there was an X-chromosome component to mating success, its magnitude was not disproportionately large when compared with the proportion of the genome contained on this chromosome. Some hybrid males courted with an anomalously low intensity, so I simultaneously mapped the genetic basis of this “courtship dysfunction.” The courtship dysfunction was caused by an interaction between the left arm of the X-chromosome in D. persimilis with the autosomes or Y-chromosome from D. pseudoobscura. Anomalous courtship behavior in interspecific hybrids can obscure the conclusions of studies of the genetics of sexual isolation, so courtship intensity should be evaluated in all such investigations.     

Impact of Asymmetrical Hybridization Followed By Sterile F1 Hybrids on Species Replacement in Pseudorasbora

Pseudorasbora pumila, one of the endangered freshwater minnows in eastern Japan, has been largely replaced by the accidentally introduced species, P. parva, which originated from western Japan. In the contact zone, P. pumila and P. parva have hybridized intensively, producing sterile F₁ hybrids. The present study determined the maternal parent of F₁ hybrids using mtDNA haplotypes to investigate the mating system between P. parva and P. pumila in the hybrid zone. We also pursued the successive changes in the genetic structures of hybridizing populations over a 5-year period using allozymes. A total of 100 natural F₁ hybrids collected from six different populations had P. pumila mtDNA without exception, suggesting that sterile F₁ hybrids resulted from mating only between P. pumila females and P. parva males. Such asymmetrical hybridization implies that P. pumila females waste considerably greater reproductive efforts compared with P. parva males. The data suggest that the rapid replacement of P. pumila by P. parva has been promoted by asymmetrical hybridization, resulting in sterile F₁ hybrids.     

A study of wing morphology and fluctuating asymmetry in interspecific hybrids between Drosophila buzzatii and D. koepferae

In this work we investigate the effect of interspecific hybridization on wing morphology using geometric morphometrics in the cactophilic sibling species D. buzzatii and D. koepferae. Wing morphology in F₁ hybrids exhibited an important degree of phenotypic plasticity and differs significantly from both parental species. However, the pattern of morphological variation between hybrids and the parental strains varied between wing size and wing shape, across rearing media, sexes, and crosses, suggesting a complex genetic architecture underlying divergence in wing morphology. Even though there was significant fluctuating asymmetry for both, wing size and shape in F₁ hybrids and both parental species, there was no evidence of an increased degree of fluctuating asymmetry in hybrids as compared to parental species. These results are interpreted in terms of developmental stability as a function of a balance between levels of heterozygosity and the disruption of coadaptation as an indirect consequence of genomic divergence.