Variation in female mate preference across a grasshopper hybrid zone

A method for studying variation in female preference among populations free from the confounding effects of variation in male characters is described. This method is applied to sixteen populations in a transect across a hybrid zone in the grasshopper, Chorthippus parallelus. Significant variation in preference is revealed among populations. The pattern of this variation is analysed in the context of the reinforcement model of speciation. While the data do indicate an increased homogamic preference on one side of the zone relative to more distant populations, this pattern cannot be distinguished statistically from an abrupt transition in preference over a distance of less than 1 km. We argue that either of these patterns implies that selection operates on female preferences in the hybrid zone.     

A hybrid zone between two subspecies of Chorthippus parallelus. X-chromosome variation through a contact zone

Two grasshopper subspecies, Chorthippus parallelus parallelus (Cpp) and Chorthippus parallelus erythropus (Cpe), meet along the Pyrenees where they hybridize and produce a hybrid zone. A contact zone located in the Col de Portalet has been analyzed for the distribution of chromosome markers on the sex (X) chromosome in pure and hybrid populations. C-banding allowed us to distinguish both pure subspecific forms and recombinant forms, and to demonstrate their different frequencies through the contact zone. Interestingly, the distal C-band (P) on the X chromosome that characterizes Cpp individuals occurs at very high frequency through the zone and then drops sharply, close to where pure Cpe populations are found. A novel interstitial C-band (H) has been found, probably derived from that characterizing Cpe individuals (E). This marker band is only present in hybrid populations. These data are discussed in the light of the dynamics of the hybrid zone.     

CAN GENE FLOW PREVENT REINFORCEMENT?

A model of reinforcement in a hybrid zone is developed in which an allele causing reinforcement may only be favored in the center of the hybrid zone and is selected against elsewhere. When reinforcement is favored only in the hybrid zone, the swamping effect of gene flow severely impedes the evolution of reinforcement: reinforcement can only occur when β < sγ²/4 (s is selection against hybrids, γ is the strength of reinforcement, and β is the strength of selection against reinforcement). Although the region in which reinforcement can occur is narrower when selection against hybrids is stronger, strong selection is still more likely to be reinforced despite the effects of gene flow. Another modifier is considered which reduces postmating isolation. The conditions for increase in frequency of this modifier reduce to the same conditions as those for the reinforcing allele. Both kinds of modification reduce the strength of selection and, therefore, cause the cline to widen. Reinforcement causes an increase in premating isolation between races, while modification of selection reduces postmating isolation.     

Divergence in mate finding behavior between two subspecies of the meadow grasshopperChorthippus parallelus (Orthoptera: Acrididae)

Two parapatric subspecies of the grasshopper Chorthippus parallelusform a hybrid zone in the Pyrenees. Partial assortative mating between subspecies must result from differences in elements of the sequence of behaviors that leads to mating. It was already known that the subspecies differ in the structure of the acoustic mating signal produced by males and in the amount of song produced under laboratory conditions. Here we investigate whether any differences exist in male mating strategies in the field, concentrating on the prediction that male C. p. erythropussing less and compensate for this by active searching (males divide their time between singing and moving in search of females). Results of field observations show that the two subspecies do in fact allocate their time differently. C. p. erythropusmales spend almost twice as much time moving around the habitat as C. p. parallelusmales. Furthermore, C. p. parallelusmales divide their time into long bouts of continuous singing alternating with periods of movement. C. p. erythropusmales, on the other hand, sing in short bursts interspersed with movement. The possible reasons for this divergence in male mating behavior and its consequences are discussed.     

Seeking signatures of reinforcement at the genetic level: a hitchhiking mapping and candidate gene approach in the house mouse

Reinforcement is the process by which prezygotic isolation is strengthened as a response to selection against hybridization. Most empirical support for reinforcement comes from the observation of its possible phenotypic signature: an accentuated degree of prezygotic isolation in the hybrid zone as compared to allopatry. Here, we implemented a novel approach to this question by seeking for the signature of reinforcement at the genetic level. In the house mouse, selection against hybrids and enhanced olfactory‐based assortative mate preferences are observed in a hybrid zone between the two European subspecies Mus musculus musculus and M. m. domesticus, suggesting a possible recent reinforcement event. To test for the genetic signature of reinforcing selection and identify genes involved in sexual isolation, we adopted a hitchhiking mapping approach targeting genomic regions containing candidate genes for assortative mating in mice. We densely scanned these genomic regions in hybrid zone and allopatric samples using a large number of fast evolving microsatellite loci that allow the detection of recent selection events. We found a handful of loci showing the expected pattern of significant reduction in variability in populations close to the hybrid zone, showing assortative odour preference in mate choice experiments as compared to populations further away and displaying no such preference. These loci lie close to genes that we pinpoint as testable candidates for further investigation.     

CLINES FOR HYBRID DYSFUNCTION IN A GRASSHOPPER HYBRID ZONE

Two subspecies of the grasshopper Chorthippus parallelus meet in the Pyrenees forming a hybrid zone several kilometers wide. Crosses between the two pure taxa result in sterile male offspring and normal females (i.e., Haldane's rule applies). However, no such dysfunction has been detected in hybrid males collected through the center of the hybrid zone. By assessing the level of dysfunction in the offspring of reciprocal crosses, it was possible to map clines for the genes responsible for dysfunction through the zone. This analysis shows that there is no abrupt transition between incompatible genomes in the field. Crosses were also made between females collected from a transect spanning the hybrid zone and pure males of both subspecies. This reveals noncoincident clines for dysfunction near the center of the hybrid zone such that the dysfunction expressed in the offspring of these crosses is less than expected from simple models. More complex models involving interaction among genes must be invoked. Also, the possibility exists that since the postglacial contact of these two grasshopper taxa, hybrid dysfunction has become ameliorated by the evolution of modifiers. This hybrid zone is thought to be a tension zone, maintained by a balance between selection against hybrid genotypes and dispersal into the zone center. The lessening of hybrid disadvantage over time through the breakdown of epistatic interactions by recombination or through modification could account for the general lack of dysfunction in field collected hybrids today.     

REINFORCING SELECTION IS EFFECTIVE UNDER A RELATIVELY BROAD SET OF CONDITIONS IN A MOSAIC HYBRID ZONE

Many hybrid zones have a mosaic structure, yet we know of no theoretical work that examines the impact of mosaicism on the outcome of evolution. We developed a computer simulation model designed to test whether the outcome of reinforcing selection differs in a mosaic and a clinal hybrid zone. Our model was a one-dimensional stepping-stone model. The mosaic and clinal hybrid zones that we modeled were, respectively, a mosaic maintained by differential fitness of the interacting taxa in patchy habitats and a tension zone. We modeled changes in gene frequency at two biallelic loci, A and B. Hybrids at the A locus were selected against. An allele at the B locus caused assortative mating at the A locus, which promoted reinforcement; there was a selective cost to this allele. In a mosaic hybrid zone, spatial variation in the fitness of A-locus homozygotes in different patches caused gene and genotype frequencies at the A and B loci to differ greatly from those in a tension zone. Compared to a tension zone, a mosaic hybrid zone had a broader region in which hybrids could be formed and, thus, a broader region in which the assortative-mating allele provided a net selective advantage (via decreased production of the less fit A-locus hybrids). This caused the assortative-mating allele to be favored under a broader set of conditions in a mosaic hybrid zone than in a tension zone. In mosaic and tension hybrid zones, both low and high levels of migration could prevent the establishment of the allele that promoted reinforcement, but the allele could establish under a wider range of migration rates in a mosaic than in a tension zone. In a tension zone, both low and high levels of selection against A-locus hybrids could prevent the establishment of the assortative-mating allele. In a mosaic hybrid zone, the assortative-mating allele established under lower levels of selection against hybrids than in a tension zone, and high levels of selection did not impede the establishment of this allele. Overall, our work illustrates how the structure of a hybrid zone can alter the outcome of an important evolutionary process, in this case, reinforcement.     

Wolbachia effects in natural populations of Chorthippus parallelus from the Pyrenean hybrid zone

We evaluate for the first time the effect of Wolbachia infection, involving two different supergroups, on the structure and dynamics of the hybrid zone between two subspecies of Chorthippus parallelus (Orthoptera) in the Pyrenees. Wolbachia infection showed no effects on female fecundity or a slight increment in females infected by F supergroup, although in the last case it has to be well established. Cytoplasmic incompatibility (CI) is confirmed in crosses carried out in the field between individuals from a natural hybrid population. This CI, registered as the relative reduction in embryo production (s_h), was of s_h = 0.355 and s_h = 0.286 in unidirectional crosses involving B and F supergroups, respectively. CI also occurred in bidirectional crosses (s_h = 0.147) but with a weaker intensity. The transmission rates of the two Wolbachia strains (B and F) were estimated by the optimization of a theoretical model to reach the infection frequencies observed in certain population. To fit this scenario, both supergroups should present transmission rates close to 1. Further, we have simulated the infection dynamics, and hence, the capacity of Wolbachia to structure the population of the host insects and to affect to reproduction and genetic introgression in the hybrid zone. This represents a first example of the influence of Wolbachia in an insect natural hybrid zone.     

Bilateral Song Convergence in a Passerine Hybrid Zone: Genetics Contribute in One Species Only

Hybridization can drive the convergence of territorial and sexual signals. However, non-genetic processes such as competition, environment matching, or cultural transmission, also generate this pattern. We investigated the effect of hybridization on song convergence between two interspecifically territorial warblers in a moving hybrid zone. We confirmed song convergence in each species. Using an AFLP-based genetic index, we detected an effect of genetics on song convergence in Hippolais polyglotta, the expanding species. Evidence was weaker for H. icterina, the receding species. In moving zones, introgression is expected to be larger in the expanding species than in the receding. Thus, the asymmetric contribution of the genetic index to convergence was consistent with expectations for genetically determined traits in moving hybrid zones, and the observed introgression pattern of AFLP markers. However, the geographical location of individuals had an effect on song variation too when genetics was accounted for, suggesting that convergence also has non-genetic explanations. We examine the possible role of alternative processes to that of hybridization and discuss their conflicting effects on reinforcement and hybrid zone dynamics.     

PATTERNS OF HYBRIDIZATION IN THE PIRIQUETA CAROLINIANA COMPLEX IN CENTRAL FLORIDA: EVIDENCE FOR AN EXPANDING HYBRID ZONE

We have identified a broad zone of hybridization between two morphologically and ecologically distinct herbaceous perennial taxa (morphotypes) within the Piriqueta caroliniana complex, which extends more than 300 km across the central Florida peninsula. Phylogeographic analyses indicate that the caroliniana morphotype has been present in north and central Florida since the early Pleistocene and that the viridis morphotype has immigrated into southern Florida much more recently. We examine the distribution of diagnostic morphological characters and nuclear genetic markers to assess the extent and patterns of introgression in this system. The results are consistent with the hypothesis that this hybrid zone has expanded north in recent history as viridis alleles have introgressed into regions that were previously occupied by populations of caroliniana. Genetic markers diagnostic for caroliniana have consistently high frequencies across the hybrid zone, whereas markers for viridis are extremely variable among populations with frequency reversals in adjacent populations. The latter pattern is probably the result of the combined stochastic effects of dispersal and drift on viridis alleles as they introgressed northward. Additional evidence for the recent expansion of this hybrid zone comes from patterns of variation for morphological and genetic markers. As expected for an expanding hybrid zone, within-population morphological variation was greatest toward the advancing front of introgression and levels of genetic variation for neutral diagnostic markers were greatest in the region of initial contact and lower in areas of recent expansion. The observed patterns of variation suggest that at least some hybrid genotypes have high fitnesses, which has led to the expansion of the hybrid zone via the displacement of parental genotypes in central Florida.     

A Narrow Heliconius cydno (Nymphalidae; Heliconiini) Hybrid Zone With Differences in Morph Sex Ratios1

In the western cordillera of the Cauca valley, Colombia, there is a narrow hybrid zone between two races of Heliconius cydno, one of which is polymorphic. Three large broods show that most of the phenotypic variation observed can be explained by four loci of major effect, named Sb, Yb, L, and K. Two further loci, G and Wo, were identified that control more minor phenotypic variation. Sb, Yb, and Wo are linked and the latter is differentially expressed between the sexes. The transition between H. c. cydnides in the north and H. c. weymeri to the south occurs over approximately 15 km. Collections from a single site near the center of the hybrid zone show that gene frequencies have been stable over the 10 yr from 1991 to 2001 and that color‐pattern genes are in Hardy–Weinberg equilibrium, with little evidence for linkage disequilibrium. This suggests that mating is random between color‐pattern forms. Nonetheless, there is evidence for variation in the sex ratio, with parental phenotypes and the most melanic hybrid phenotypes showing a fairly even sex ratio compared to a strong male bias observed in the remaining hybrids. We hypothesize that this might be explained by differential selection between sexes and phenotypes, phenomena that could have important implications for hybrid zone analysis and the evolution of Müllerian mimicry.     

Postglacial expansion and genome subdivision in the European grasshopper Chorthippus parallelus

A noncoding nuclear DNA marker sequence (Cpnl-1) was used to investigate subdivision in the grasshopper Chorthippus parallelus and deduce postglacial expansion patterns across its species range in Europe. Investigation of the spatial distribution of 71 Cpnl-1 haplotypes and estimation of levels of genetic differentiation (K_ST values) between populations and geographic regions provided evidence for subdivision of C. parallelus into at least five major geographic regions and indicated that the French form of C. parallelus originated after range expansion from a Balkan refugium, Further evidence for subdivision of C. parallelus between Italy and northern Europe suggests that the Alps may have formed a significant barrier to gene flow in this grasshopper.     

Detection of Spiroplasma and Wolbachia in the Bacterial Gonad Community of Chorthippus parallelus

We have recently detected the endosymbiont Wolbachia in multiple individuals and populations of the grasshopper Chorthippus parallelus (Orthoptera: acrididae). This bacterium induces reproductive anomalies, including cytoplasmic incompatibility. Such incompatibilities may help explain the maintenance of two distinct subspecies of this grasshopper, C. parallelus parallelus and C. parallelus erythropus, which are involved in a Pyrenean hybrid zone that has been extensively studied for the past 20 years, becoming a model system for the study of genetic divergence and speciation. To evaluate whether Wolbachia is the sole bacterial infection that might induce reproductive anomalies, the gonadal bacterial community of individuals from 13 distinct populations of C. parallelus was determined by denaturing gradient gel electrophoresis analysis of bacterial 16S rRNA gene fragments and sequencing. The study revealed low bacterial diversity in the gonads: a persistent bacterial trio consistent with Spiroplasma sp. and the two previously described supergroups of Wolbachia (B and F) dominated the gonad microbiota. A further evaluation of the composition of the gonad bacterial communities was carried out by whole cell hybridization. Our results confirm previous studies of the cytological distribution of Wolbachia in C. parallelus gonads and show a homogeneous infection by Spiroplasma. Spiroplasma and Wolbachia cooccurred in some individuals, but there was no significant association of Spiroplasma with a grasshopper’s sex or with Wolbachia infection, although subtle trends might be detected with a larger sample size. This information, together with previous experimental crosses of this grasshopper, suggests that Spiroplasma is unlikely to contribute to sex-specific reproductive anomalies; instead, they implicate Wolbachia as the agent of the observed anomalies in C. parallelus.     

A narrow hybrid zone between the grasshoppers Stenobothrus clavatus and Stenobothrus rubicundus (Orthoptera: Gomphocerinae): female preferences for courtship songs

Two grasshopper species, Stenobothrus rubicundus and Stenobothrus clavatus, were previously shown to hybridize in a narrow contact zone on Mount Tomaros in northern Greece. The species are characterized by complex and completely different courtship songs. In the present study, we investigated female preferences for the courtship songs of S. rubicundus, S. clavatus and hybrids in playback experiments. Playback of the courtship songs revealed assortative preferences in females of the parental species: they significantly more often preferred the songs of conspecific males. Hybrid females showed a lower selectivity than parental females, responding somewhat equally eager to playback of the songs of S. clavatus, S. rubicundus, and natural hybrid song, although less actively to the F₁ hybrid song. The results suggest that hybrid males may lose to males of parental species, whereas hybrid females would even have an advantage over parental females. Comparison of responses of females from allopatric populations and Mount Tomaros to different song types shows no evidence for reinforcement. Asymmetry found in female preferences may have implications for the structure of the hybrid zone. © 2013 The Linnean Society of London     

Hybridization effects and genetic diversity of the common and black‐tufted marmoset (Callithrix jacchus and Callithrix penicillata) mitochondrial control region

Hybridization is continually documented in primates, but effects of natural and anthropogenic hybridization on biodiversity are still unclear and differentiating between these contexts remains challenging in regards to primate evolution and conservation. Here, we examine hybridization effects on the mitochondrial DNA (mtDNA) control region of Callithrix marmosets, which provide a unique glimpse into interspecific mating under distinct anthropogenic and natural conditions. DNA was sampled from 40 marmosets along a 50‐km transect from a previously uncharacterized hybrid zone in NE Brazil between the ranges of Callithrix jacchus and Callithrix penicillata. DNA was also collected from 46 marmosets along a 30‐km transect in a hybrid zone in Rio de Janeiro state, Brazil, where exotic marmosets appeared in the 1980s. Combining Callithrix DNA sampled inside and outside of these hybrid zones, phylogenetic and network analyses show C. jacchus and C. penicillata being parental species to sampled hybrids. We expand limited Callithrix population genetics work by describing mtDNA diversity and demographic history of these parental species. We show ancient population expansion in C. jacchus and historically constant population size in C. penicillata, with the latter being more genetically diverse than the former. The natural hybrid zone contained higher genetic diversity relative to the anthropogenic zone. While our data suggest hybrid swarm formation within the anthropogenic zone due to removed physical reproductive barriers, this pattern is not seen in the natural hybrid zone. These results suggest different genetic dynamics within natural and anthropogenic hybridization contexts that carry important implications for primate evolution and conservation. Am J Phys Anthropol 155:522–536, 2014. © 2014 Wiley Periodicals, Inc.     

Genetic variation and self-incompatibility within and outside a Rorippa hybrid zone (Brassicaceae)

Rorippa amphibia and R. sylvestris are both self-incompatible, perennial species. In northern Germany they show a geographic pattern of interspecific gene flow: R. amphibia and R. sylvestris form a hybrid zone at the river Elbe but do not hybridise elsewhere in northern Germany. In the present paper we analyse the relationship between molecular diversity, seed set and seed germination within and outside the hybrid zone. In both species we observed high genetic variation and high seed set in the hybrid zone, and low genetic variation and low seed set outside the hybrid zone. In R. amphibia, high genetic variation is correlated with high seed set, but with low germination rates of the seeds. The results of an isolation experiment show that self-incompatibility is maintained in the parental species and their hybrids. The impact of genetic self-incompatibility on hybrid zone formation and hybrid fitness is discussed.     

ELECTROPHORETIC CONFIRMATION OF INTERSPECIFIC HYBRIDIZATION IN AESCULUS (HIPPOCASTANACEAE) AND THE GENETIC STRUCTURE OF A BROAD HYBRID ZONE

Within a broad (>200 km wide) hybrid zone involving three parapatric species of Aesculus, we observed coincident clines in allele frequency for 6 of 14 electrophoretic loci. The cooccurrence of alleles characteristic of A. pavia, A. sylvatica, and A. flava was used to estimate genetic admixtures in 48 populations involving various hybrids between these taxa in the southeastern United States. High levels of allelic polymorphism (up to 40% greater than the parental taxa) were observed in hybrid populations and also in some populations bordering the hybrid zone. A detailed analysis of a portion of the hybrid zone involving A. pavia and A. sylvatica revealed a highly asymmetrical pattern of gene flow, predominantly from Coastal Plain populations of A. pavia into Piedmont populations of A. sylvatica. Computer simulations were used to generate expected genotypic arrays for parental, F₁; and backcross individuals, which were compared with natural populations using a character index scoring system. In these comparisons, hybrid individuals could be distinguished from either parent, but F₁ and backcross progeny could not be distinguished from each other. Most hybrid populations were found to include hybrids and one of the parental taxa, but never both parents. Three populations appeared to be predominantly hybrids with no identifiable parental individuals. Hybrids occurred commonly at least 150 km beyond the range of A. pavia, but usually not more than 25 km beyond the range of A. sylvatica. Introgression, suggested by genetically hybrid individuals and significant gene admixtures of two or more species in populations lacking morphological evidence of hybridization, may extend the hybrid zone further in both directions. The absence of one or both parental species from hybrid populations implies a selective disadvantage to parentals in the hybrid zone and/or that hybridization has occurred through long-distance gene flow via pollen, primarily from A. pavia into A. sylvatica. Long-distance pollen movement in plants may generate hybrid zones of qualitatively different structure than those observed in animals, where gene flow involves dispersal of individuals.     

Plant hybrid zones as centers of biodiversity: the herbivore community of two endemic Tasmanian eucalypts

We found the hybrid zone between Eucalyptus amygdalina and Eucalyptus risdonii to be a center of insect and fungal species richness and abundance. Of 40 taxa examined, 73% were significantly more abundant in the hybrid zone than in pure zones, 25% showed on significant differences, and 2% were most abundant on a pure host species. The average hybrid tree supported 53% more insect and fungal species, and relative abundances were, on average, 4 times greater on hybrids than on either eucalypt species growing in pure stands. Hybrids may act as refugia for rare species: 5 of 40 species were largely restricted to the hybrid zone. Also, 50% of the species coexisted only in the hybrid zone, making for mique species assemblages. Although hybrids support more species and greater abundances, all hybrids are not equal: 68% of the 40 taxa examined were significantly more abundant on one hybrid phenotype than another. While herbivore concentrations on F1 type intermediates were rare, concentrations were common on phenotypes resembling backcrosses either to E. amygdalina or E. risdonii. For specialist herbivores, the hybrid phenotype most heavily utilized appears to be determined by its phenotypic affinity to its host species. Generalists exhibit an overall greater abundance on hybrids, but are less likely to utilize one hybrid phenotype over another. Mechanistic explanations for these distributions are numerous and probably species specific, but are likely to include: increased genetic susceptibility of hybrids due to hybrid breakdown; increased stress in the hybrid zone resulting in greater plant susceptibility; and a greater diversity of resources in the hybrid zone which could support more species. Seed capsule production by hybrids and their parental species is negatively correlated with herbivory. However, it is difficult to determine whether herbivores cause this pattern as hybrids may have inherently lower sexual reproduction. Laws enacted to protect rare and endangered species do not include hybrids. We argue that a re-examination of our current hybrid policy is warranted. Plant hybrid zones are centers of plant evolution and speciation, sources of economically important plants and potential biocontrol agents, and, as our study suggests, also provide essential habitats for phytophagous communities.     

Spatiotemporally consistent genomic signatures of reproductive isolation in a moving hybrid zone

Studies of hybrid zone dynamics often investigate a single sampling period and draw conclusions from that temporal snapshot. Stochasticity can, however, result in loci with spurious outlier patterns, which is exacerbated by limited temporal or geographic sampling. Comparing admixed populations from different geographic regions is one way to detect repeatedly divergent genomic regions potentially involved in reproductive isolation. Temporal comparisons also allow us to control partially for the role of stochasticity, but the power of temporal sampling has not yet been adequately explored. In North America, black‐capped (Poecile atricapillus) and Carolina (P. carolinensis) chickadees hybridize in a contact zone extending from New Jersey to Kansas. The hybrid zone is likely maintained by strong intrinsic selection against hybrids, and it is moving north. We used a reduced representation genomic approach and temporally spaced sampling—two samples of ～80 individuals separated by a decade—to determine the pattern and consistency of selection and genomic introgression in the chickadee hybrid zone. We report consistently low introgression for highly divergent loci between P. atricapillus and P. carolinensis in this moving hybrid zone. This is strong evidence that these loci may be linked to genomic regions involved in reproductive isolation between chickadees.     

A MODEL OF A HYBRID ZONE BETWEEN TWO CHROMOSOMAL RACES OF THE COMMON SHREW (SOREX ARANEUS)

The common shrew (Sorex araneus) is subdivided into several karyotypic races in Britain. Two of these races meet near Oxford o form the “Oxford-Hermitage” hybrid zone. We present a model which describes this system a; a “tension zone,” i.e., a set of clines maintained by a balance between dispersal and selection against chromosomal heterozygotes. The Oxford and Hermitage races differ by Robertsonian fusions with monobrachial homology (kq, no versus ko), and so Fl hybrids between them would have low fertility. However, the acrocentric karyotype is found at high frequency within the hybrid zone, so that complex Robertsonian heterozygotes (kg no/q ko n) are replaced by more fertile combinations, such as (kg no/k q n o). This suggests that the hybrid zone has been modified so as to increase hybrid fitness. Mathematical analysis and simulation show that, if selection against complex heterozygotes is sufficiently strong relative to selection against simple heterozygotes, acrocentrics increase, and displace the clines for kg and no from the cline for ko. Superimposed on this separation is a tendency for the hybrid zone to move in favor of the Oxford (kg no) race. We compare the model with estimates of linkage disequilibrium and cline shape made from field data.