Viability selection on overwintering eggs in a field cricket mosaic hybrid zone

Habitat segregation allows recently diverged taxa to minimize competition and maximize fitness. Consequently, the overall distribution of a species in part will be determined by interactions with their close relatives. Two recently diverged but hybridizing field crickets, Gryllus firmus and G. pennsylvanicus , form a mosaic hybrid zone in Connecticut in which each species is associated with a specific soil type: G. firmus is found on sand soils and G. pennsylvanicus is found on loam soils. Both species produce diapausing eggs, which spend the winter in the ground. We investigate how viability selection on overwintering eggs in different soils influences the distributions of the two species, habitat partitioning between them, and the structure of the mosaic hybrid zone. Our results suggest that selection on eggs by soil types is not important in determining the success of crickets on different soils. However, winter climate has a strong effect on egg viability, and variation in egg survivorship over different winter climate regimes likely plays an important role in determining the distribution of these cricket species and the position of the hybrid zone.     

PATTERNS OF VARIATION AND LINKAGE DISEQUILIBRIUM IN A FIELD CRICKET HYBRID ZONE

The distribution of multilocus genotypes found within a natural hybrid zone is determined by the sample of genotypes present when the hybrid zone first formed, by subsequent patterns of genetic exchange between the hybridizing taxa, and by drift and selection within each of the hybrid zone populations. We have used anonymous nuclear DNA restriction fragment polymorphisms (RFLPs) to characterize the array of multilocus genotypes present within a well-studied hybrid zone between two eastern North American field crickets, Gryllus pennsylvanicus and Gryllus firmus. These crickets hybridize along a zone of contact that extends from New England to Virginia. Previous studies have shown that both premating and postmating barriers exist between the two cricket species, but the absence of diagnostic morphological and allozyme markers has made it difficult to assess the consequences of these barriers for genetic exchange. Analyses based on four diagnostic anonymous nuclear markers indicate that hybrid zone populations in Connecticut contain few F₁ hybrids, and that nonrandom associations persist among nuclear gene markers, between nuclear and cytoplasmic markers, and between molecular markers and morphology. Field cricket populations within the hybrid zone are not “hybrid swarms” but consist primarily of crickets that are very much like one or the other of the parental species. Despite ample opportunity for genetic exchange and evidence for introgression at some loci, the two species remain quite distinct. Such a pattern appears to be characteristic of many natural hybrid zones.     

BARRIERS TO GENE EXCHANGE BETWEEN CLOSELY RELATED CRICKET SPECIES. II. LIFE CYCLE VARIATION AND TEMPORAL ISOLATION

The closely related field crickets Gryllus firmus and Gryllus pennsylvanicus are known to hybridize in a zone of contact that extends over more than 800 km from the Blue Ridge in Virginia to southern Connecticut. Here I present evidence that the extent of temporal reproductive isolation of the two cricket species varies along the length of the zone. Adults of G. firmus and G. pennsylvanicus occur synchronously in Connecticut but G. firmus matures significantly later than G. pennsylvanicus along the Blue Ridge in Virginia. Variation in the extent of temporal isolation is a consequence of intrinsic differences in the egg-to-adult development time of G. firmus from the two localities. In laboratory rearing experiments, there is little variation in development time among crickets from G. pennsylvanicus populations ranging from Vermont to Virginia. However, G. firmus from Virginia takes significantly longer to mature than G. firmus from Connecticut. Comparisons between species from neighboring localities show that lowland Virginia G. firmus take much longer to develop and are larger as adults than montane Virginia G. pennsylvanicus. In Connecticut, crickets of the two species differ very little in development time and body size. Patterns of variation in development time and body size are compared with similar data for frogs along one of the same environmental gradients and for field crickets along similar gradients elsewhere. Although G. firmus and G. pennsylvanicus are temporally isolated in Virginia, adults of the two species do occur together. Examination of Esterase genotypes of field-inseminated females and their progeny from a mixed population on the Blue Ridge suggests that pre-mating barriers other than temporal isolation are not effective in preventing gene exchange.     

GENETIC VARIATION IN CRICKET CALLING SONG ACROSS A HYBRID ZONE BETWEEN TWO SIBLING SPECIES

The sibling ground crickets Allonemobius fasciatus and A. socius meet along a mosaic hybrid zone at ≈ 40°N latitude in eastern North America. In this paper we report the findings of a genetic analysis of calling-song variation within and among six cricket populations sampled along a transect through the hybrid zone in southern New Jersey. We compared aspects of the calling song of both wild-caught and laboratory-reared crickets to test the hypothesis that population differences in song observed in the wild were genetically based. We found significant, species-level differences in all aspects of the calling song, and these differences persisted even after a generation of common-garden rearing in the laboratory, supporting the hypothesis that interspecific variation observed in the wild largely reflects genetic differentiation between the two taxa. A discriminant function analysis indicated that individual crickets could be assigned to the proper taxon with less than 10% error, supporting the premise that calling song could be used by female crickets as a mechanism for species recognition. One population, collected from within the hybrid zone and containing significant numbers of hybrid individuals, was intermediate in its calling song, presumably reflecting this population's mixed genetic makeup. In this hybrid zone population, song phenotype was highly correlated to a hybrid index score generated using species-specific alleles at four diagnostic allozyme markers, suggesting a multigenic basis to calling-song variation in these crickets as well as linkage disequilibrium between markers and song. Based on an analysis of laboratory-reared full-sib families, broad-sense heritabilities for calling-song characteristics were generally significant in the two A. socius populations, whereas many components of song showed no significant family effects in the three A. fasciatus populations. The genotypically mixed, hybrid zone population showed very high heritabilities for most calling-song components, which likely reflect the influence of interspecific gene flow on genetic variation for quantitative traits.     

A fine-scale spatial analysis of the mosaic hybrid zone between Gryllus firmus and Gryllus pennsylvanicus

Abstract.— The pattern of character variation within a hybrid zone, the hybrid zone structure, has been used to infer the processes that maintain hybrid zones. Unfortunately it is difficult to infer process from structure alone because many different processes can produce the same pattern of character variation. Mosaic hybrid zones may be maintained by exogenous selection in a heterogeneous environment and/or endogenous selection against hybrid individuals; habitat preference, premating isolating barriers and/or fertility selection can also contribute. The spatial scale at which a hybrid zone is sampled affects its apparent structure; a hybrid zone may appear clinal at one scale and mosaic at another. Here, we sample the mosaic hybrid zone between two field crickets, Gryllus firmus and G. pennsylvanicus , at a scale that spans the boundaries between individual soil-habitat patches. From our analysis, we find that at fine scales, the mosaic hybrid zone resolves into a set of steep clines across patch boundaries. Both morphological and molecular traits exhibit sharp and generally concordant clines. However, clines for mitochondrial DNA and one anonymous nuclear marker are clearly displaced as a result of current hybridization or past introgression (the "ghost of hybridization past"). Thus, scale is important for the structure of this and probably other hybrid zones. The extremely sharp, concordant clines across patch boundaries indicate that the cricket hybrid zone is undoubtedly structured by selection. However, the detailed mechanisms responsible for the maintenance of the hybrid zone–whether endogenous selection against hybrids, exogenous selection by the environment, and/or behavioral preferences for mates or habitats– remain to be elucidated. Determining these mechanisms will depend on closer inspection of the organisms themselves and their interactions, as is the case for all hybrid zones.     

SURVIVAL OF HYBRIDS IN A MOSAIC HYBRID ZONE

The ground crickets Allonemobius fasciatus and A. socius meet in a mosaic hybrid zone that stretches from New Jersey at least as far west as Illinois. Within mixed populations from the contact zone, “pure” species individuals predominate. To determine whether hybrids are less viable than pure-species individuals, and to assess whether the high proportion of pure-species individuals in mixed populations results from hybrid inviability, we performed a cohort analysis. In this study, five mixed populations from the hybrid zone were each sampled three to five times from the fall of 1986 to the spring of 1988. Individuals were assigned to one of three categories based on their genotypes: A. socius (individuals harboring only alleles unique to A. socius), hybrid (individuals with alleles unique to both species), and A. fasciatus (individuals harboring only alleles unique to A. fasciatus). This sampling and measurement scheme permitted monitoring of the proportion of hybrid individuals in a population over time. The results were fairly consistent. The relative survival of A. socius was greater than the relative survival of members of the other two groups in four of the five populations. The relative viability of A. fasciatus was greater than that of hybrids in one population, approximately equal to that of hybrids in two populations, and less than that of hybrids in two populations. These results not only shed light on an important component of fitness, viability from hatching to adulthood, but they demonstrate that loss of hybrid individuals during the course of the field season will not explain deviations from random mating expectations present in mixed populations in late summer. The deviations must be due to assortative mating or to a reproductive barrier operating prior to egg hatch.     

SPATIAL STRUCTURE AND HABITAT VARIATION IN A GRASSHOPPER HYBRID ZONE

Abstract A hybrid zone between the grasshoppers Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) in northern Spain has been analyzed for variation in morphology and ecology. These species are readily distinguished by the number of stridulatory pegs on the hind femur. Both sexes are fully winged and inhabit disturbed habitats throughout the study area. We develop a maximum-likelihood approach to fitting a two-dimensional cline to geographical variation in quantitative traits and for estimating associations of population mean with local habitat. This method reveals a cline in peg number approximately 30 km south of the Picos de Europa Mountains that shows substantial deviations in population mean compared with the expectations of simple tension zone models. The inclusion of variation in local vegetation in the model explains a significant proportion of the residual variation in peg number, indicating that habitat-genotype associations contribute to the observed spatial pattern. However, this association is weak, and a number of populations continue to show strong deviations in mean even after habitat is included in the final model. These outliers may be the result of long-distance colonization of sites distant from the cline center or may be due to a patchy pattern of initial contact during postglacial expansion. As well as contrasting with the smooth hybrid zones described for Chorthippus parallelus , this situation also contrasts with the mosaic hybrid zones observed in Gryllus crickets and in parts of the hybrid zone between Bombina toad species, where habitat-genotype associations account for substantial amounts of among-site variation.     

Structure of a mosaic hybrid zone between the field crickets Gryllus firmus and G. pennsylvanicus

Hybrid zones provide insight into the nature of species boundaries and the evolution of barriers to gene exchange. Characterizing multiple regions within hybrid zones is essential for understanding both their history and current dynamics. Here, we describe a previously uncharacterized region of a well‐studied hybrid zone between two species of field crickets, Gryllus pennsylvanicus and G. firmus. We use a combination of mitochondrial DNA sequencing, morphological data, and modeling of environmental variables to identify the ecological factors structuring the hybrid zone and define patterns of hybridization and introgression. We find an association between species distribution and natural habitat; Gryllus pennsylvanicus occupies natural habitat along forest edges and natural clearings, whereas G. firmus occupies more disturbed areas in agricultural and suburban environments. Hybridization and introgression occur across patch boundaries; there is evidence of substantial admixture both in morphological characters and mtDNA, over a broad geographic area. Nonetheless, the distribution of morphological types is bimodal. Given that F₁ hybrids are viable and fertile in the lab, this suggests that strong pre‐zygotic barriers are operating in this portion of the hybrid zone.     

Mitochondrial DNA phylogeny of North American field crickets: perspectives on the evolution of life cycles,songs, and habitat associations

North American field crickets (genus Gryllus) exhibit a diversity of life cycles, habitat associations, and calling songs. However, patterns of evolution for these ecological and behavioral traits remain uncertain in the absence of a robust phylogenetic framework. Analyses of morphological variation have provided few clues about species relationships in the genus Gryllus. Here we use comparisons of mitochondrial DNA restriction site maps for 29 individuals representing 11 species (including potential outgroups) to examine relationships among eastern North American field crickets. Initially chosen as likely outgroup taxa, the two European species of Gryllus do not obviously fall outside of an exclusively North American clade and (based on amount of sequence divergence) appear to have diverged from North American lineages at about the same time that major North American lineages diverged from each other. The egg-overwintering crickets comprise a strongly supported monophyletic group, but relationships among these three closely related species cannot be resolved. The mtDNA data are consistent with a single origin of egg diapause and do not support a model of recent life cycle divergence and allochronic speciation for Gryllus pennsylvanicus and G. veletis. The two crickets are not sister species, despite remarkable similarity in morphology, habitat, and calling song. This conclusion is consistent with published data on allozyme variation in North American field crickets. The habitat associations of eastern North American field crickets have been labile, but calling songs sometimes have remained virtually unchanged across multiple speciation events.     

CALLING SONG DISPLACEMENT IN A ZONE OF OVERLAP AND HYBRIDIZATION

The ground crickets Allonemobius fasciatus and A. socius meet in a mosaic zone of overlap and hybridization stretching from the East Coast to at least Illinois. To test whether male calling song differences were enhanced in sympatry, we analyzed the songs of crickets from inside and outside the zone of overlap along two transects. No evidence of calling song displacement was found in A. socius males from populations within the zone of overlap. On the other hand, A. fasciatus displayed calling song displacement in three populations. Our results are consistent with the hypothesis that the selective pressure exerted by the challenge from a related species is frequency dependent. While not a conclusive demonstration, the observed shifts in calling song are strongly suggestive of reproductive character displacement.     

Stuck in the middle with you: close concordance between geographical clines in a cricket hybrid zone

Students of speciation have long recognized that hybridization between populations does not affect all parts of the genome in the same way (Key 1968, Bazykin 1969, Wu 2001, Nosil et al. 2009). For example, divergence is expected to be high at loci involved in Dobzhansky–Muller incompatibilities or at loci under divergent natural selection, while those that are effectively neutral should show only weak divergence. Studies that examine geographical clines at divergent loci in a hybrid zone can be particularly powerful, as here one can estimate how net selection is affecting each locus (Payseur 2010). An excellent example of this approach appears in this issue (Larson et al. 2014) for a hybrid zone between the crickets Gryllus firmus and Gryllus pennsylvanicus in the eastern United States.     

Ecological differentiation and habitat unsuitability maintaining a ground beetle hybrid zone

Exogenous selection via interactions between organisms and environments may influence the dynamics of hybrid zones between species in multiple ways. Two major models of a hybrid zone allowed us to hypothesize that environmental conditions influence hybrid zone dynamics in two ways. In the first model, an environmental gradient determines the mosaic distribution at the boundary between ecologically differentiated species (mosaic hybrid zone model). In the second model, a patch of unsuitable habitat traps a hybrid zone between species whose hybrids are unfit (tension zone model). To test these, we examined the environmental factors influencing the spatial structure of a hybrid zone between the ground beetles Carabus maiyasanus and C. iwawakianus using GIS‐based quantification of environmental factors and a statistical comparison of species distribution models (SDMs). We determined that both of the hypothetical processes can be important in the hybrid zone. We detected interspecific differences in the environmental factors in presence localities and their relative contribution in SDMs. SDMs were not identical between species even within contact areas, but tended to be similar within the range of each species. These results suggest an association between environments and species, and provide evidence that ecological differentiation between species plays a role in the maintenance of the hybrid zone. Contact areas were characterized by a relatively high temperature, low precipitation, and high topological wetness. Thus, the contact areas were regarded as being located in an unsuitable habitat with a drier climate, where those populations are likely to occur in patches with limited precipitation concentrated. A comparison of spatial scales suggests that exogenous selection via environmental factors may be weaker than endogenous selection via genitalic incompatibility.     

Do Wolbachia infections play a role in unidirectional incompatibilities in a field cricket hybrid zone?

Two closely related field crickets, Gryllus firmus and G. pennsylvanicus, hybridize along an extensive north-south zone in the eastern United States. Crosses between G. firmus males and G. pennsylvanicus females produce viable and fertile F1, but the reciprocal cross consistently fails to produce offspring. Wolbachia, a bacterial parasite of arthropods that causes unidirectional incompatibilities in a variety of insect species, has been suggested as the cause of the observed incompatibility between G. pennsylvanicus and G. firmus. We examine the presence/absence of Wolbachia strains, defined by sequencing the ftsZ gene, in four cricket populations from the north-eastern United States. Most G. firmus individuals are infected (100% in Guilford, Connecticut; 65% in Seaside Park, New Jersey) and > 95% of those infected harbour a single strain of Wolbachia. All individuals in G. pennsylvanicus populations (Ithaca, New York; Sharon, Connecticut) are infected; the majority of individuals carry a second strain of Wolbachia, but a significant fraction carry the same strain found commonly in G. firmus. The presence of an apparently identical Wolbachia strain in crickets of both species means that some crosses between G. pennsylvanicus males and G. firmus females should be compatible. We have no evidence of such compatibility. Furthermore, if Wolbachia infections are responsible for the observed incompatibility between species, then incompatibilities must also exist within G. pennsylvanicus, because this species harbours both Wolbachia strains. Although some single pair crosses within G. pennsylvanicus do fail to produce offspring, the proportion is lower than expected if Wolbachia were responsible. Therefore, Wolbachia is unlikely to be involved in reproductive isolation between the two cricket species.     

Sex,death, and genetic variation: natural and sexual selection on cricket song

Male field crickets, Gryllus integer, in Texas, USA, produce a trilled calling song that attracts female crickets, resulting in enhanced mating success. Gravid female parasitoid flies, Ormia ochracea, are also attracted to male cricket calling song, resulting in the death of the male within about seven days. Using playbacks of field-cricket calling song in the natural habitat, we show that both female crickets and female parasitoid flies prefer male calling song with average numbers of pulses per trill. Thus female crickets exert stabilizing sexual selection, whereas flies exert disruptive natural selection on male song. Disruptive natural selection will promote genetic variation and population divergence. Stabilizing sexual selection will reduce genetic variation and maintain population cohesiveness. These forces may balance and together maintain the observed high levels of genetic variation (ca. 40%) in male calling song.     

Genetic and morphological evidence of a geographically widespread hybrid zone between two crocodile species,Crocodylus acutus and Crocodylus moreletii

Hybrid zones represent natural laboratories to study gene flow, divergence and the nature of species boundaries between closely related taxa. We evaluated the level and extent of hybridization between Crocodylus moreletii and Crocodylus acutus using genetic and morphological data on 300 crocodiles from 65 localities. To our knowledge, this is the first genetic study that includes the entire historic range and sympatric zone of the two species. Contrary to expectations, Bayesian admixture proportions and maximum‐likelihood estimates of hybrid indexes revealed that most sampled crocodiles were admixed and that the hybrid zone is geographically extensive, extending well beyond their historical region of sympatry. We identified a few geographically isolated, nonadmixed populations of both parental species. Hybrids do not appear to be F₁s or recent backcrosses, but rather are more likely later‐generation hybrids, suggesting that hybridization has been going on for several to many generations and is mostly the result of natural processes. Crocodylus moreletii is not the sister species of C. acutus, suggesting that the hybrid zone formed from secondary contact rather than primary divergence. Nonadmixed individuals from the two species were distinguishable based on morphological characters, whereas hybrids had a complex mosaic of morphological characters that hinders identification in the wild. Very few nonadmixed C. acutus and C. moreletii populations exist in the wild. Consequently, the last nonadmixed C. moreletii populations have become critically endangered. Indeed, not only the parental species but also the naturally occurring hybrids should be considered for their potential conservation value.     

SPATIAL SCALE OF GENETIC STRUCTURE AND AN INDIRECT ESTIMATE OF GENE FLOW IN EELGRASS,ZOSTERA MARINA

In this study, the first investigation of population structure in an aquatic angiosperm, I show that populations of a marine angiosperm (eelgrass, Zostera marina) are genetically differentiated at a number of spatial scales. I find also that there is no correspondence between geographic and genetic distances separating subpopulations, an increasingly common result in spatially stratified studies of genetic structure in marine invertebrates. F-statistics, calculated for two years from electrophoretic variation at five polymorphic allozyme loci, indicate significant genetic differentiation among sampling quadrats within each of two bays (θ = 0.064-0.208), between tide zones within a bay (θ = 0.025-0.157) and between bays (θ = 0.079). Spatial autocorrelation analysis was used to explore genetic differentiation at smaller spatial scales; estimated patch sizes (within which genetic individuals are randomly associated) indicated no appeciable genetic structure at scales less than 20 m × 20 m. Calculated values of F-statistics were a function of the spatial scale from which samples were drawn: increasing the size of the “subpopulation” included in calculation of fixation indices for the same “total” sample resulted in an increase in the magnitude of f (e.g., from 0.092 to 0.181) and a decrease in θ (e.g., from 0.186 to 0.025). On the basis of the best estimate of the spatial scale of subpopulations, the effective number of migrants per generation (N_em) ranges from 1.1 to 2.8. Genetic consequences of the disturbance regime in the eelgrass habitat sampled were extreme variation between years in the allele richness and proportion of heterozygotes in a sample and a positive relationship between the extinction probability of patches and the genetic variance among them. The changes in F-statistics as a function of sampling scale and the observation that θ among sampled quadrats was positively associated with the probability of extinction among quadrats indicated that indirect estimates of gene flow (N_em) calculated from θ should be cautiously interpreted in populations that may not yet be in drift-migration equilibrium.     

Genotype‐environment associations support a mosaic hybrid zone between two tidal marsh birds

Local environmental features can shape hybrid zone dynamics when hybrids are bounded by ecotones or when patchily distributed habitat types lead to a corresponding mosaic of genotypes. We investigated the role of marsh‐level characteristics in shaping a hybrid zone between two recently diverged avian taxa – Saltmarsh (Ammodramus caudacutus) and Nelson's (A. nelsoni) sparrows. These species occupy different niches where allopatric, with caudacutus restricted to coastal marshes and nelsoni found in a broader array of wetland and grassland habitats and co‐occur in tidal marshes in sympatry. We determined the influence of habitat types on the distribution of pure and hybrid sparrows and assessed the degree of overlap in the ecological niche of each taxon. To do this, we sampled and genotyped 305 sparrows from 34 marshes across the hybrid zone and from adjacent regions. We used linear regression to test for associations between marsh characteristics and the distribution of pure and admixed sparrows. We found a positive correlation between genotype and environmental variables with a patchy distribution of genotypes and habitats across the hybrid zone. Ecological niche models suggest that the hybrid niche was more similar to that of A. nelsoni and habitat suitability was influenced strongly by distance from coastline. Our results support a mosaic model of hybrid zone maintenance, suggesting a role for local environmental features in shaping the distribution and frequency of pure species and hybrids across space.     

DIFFERENT SPATIAL SCALES OF ADAPTATION IN THE CLIMBING BEHAVIOR OF PEROMYSCUS MANICULATUS: GEOGRAPHIC VARIATION,NATURAL SELECTION,AND GENE FLOW

Patterns of geographic variation in tree-climbing ability of Peromyscus maniculatus were used to examine the influence of spatial variation in natural selection and gene flow on the genetic divergence of climbing behavior among populations. Offspring of adults of two subspecies sampled from 10 localities in montane conifer forest, conifer woodland, and desert scrub/grassland habitats were raised in the laboratory and tested to determine their tree-climbing ability (the maximum diameter artificial rod that a mouse could climb). Comparisons of mean rod-climbing scores revealed that individuals of P. m. rufinus sampled from montane conifer forest and conifer woodland in Arizona were better climbers than P. m. sonoriensis sampled from conifer woodland and desert habitats in Nevada. This result was consistent with the hypothesis that natural selection has produced large-scale adaptation in climbing behavior. However, the climbing ability of P. m. sonoriensis sampled from conifer woodland habitats on isolated mountaintops in Nevada has not evolved in response to natural selection to the degree expected. In addition, populations sampled from desert grassland habitat, adjacent to woodland P. m. rufinus in Arizona, have climbing abilities that are not significantly different from conifer woodland populations. These observations indicate that local adaptation was constrained. An estimate of the heritability of climbing ability (h² = 0.352 ± 0.077) indicates that lack of a response to selection was not due to the absence of additive genetic variation. In addition, regressions of interpopulation differences on the degree of geographic isolation between pairs of populations do not support the hypothesis that gene flow between habitats has constrained evolution. Instead, a combination of historical events and insufficient time to respond to selection appears to have influenced geographic variation and the spatial scale of adaptation in climbing ability.     

Morphological clines across a karyotypic zone of house mice in Central Italy

Multivariate morphometric differentiation between chromosomal races of the mouse Mus domesticus in Central Italy was investigated using a population of 2n = 22 “CB” karyotype, three populations of standard 2n = 40 karyotype, five populations of 2n = 22 “CD” karyotype and three populations from the hybrid zone between the latter two karyotypes. Whilst populations of different karyotype generally have significantly different morphometry, canonical analysis does not reveal that the populations ordinate into distinct aggregations based on karyotype, largely because the 2n = 22 “CD” populations are so diverse. Nevertheless, canonical analysis does reveal a significant cline in morphology across the contact zone between the 2n = 40 and 2n = 22 “CD” mice. The nature of this transition, i. e. a cline 1. within the 2n = 40 range, 2. within the hybrid range (but unrelated to chromosome number) and 3. within the 2n = 22 “CD” range, tends to indicate that the morphometric divergence is due to adaptation to the different ecological regimes across which these mice are distributed rather than the phylogenetic divergence of the karyotypic races.     

REGIONAL,LOCAL AND MICROGEOGRAPHIC ALLELE FREQUENCY VARIATION BETWEEN APPLE AND HAWTHORN POPULATIONS OF RHAGOLETIS POMONELLA IN WESTERN MICHIGAN

In the preceding study (Feder et al., 1990), we report that paired apple and hawthorn infesting populations of Rhagoletis pomonella are genetically differentiated for six allozymes. Here, we show that patterns of intra- and inter-host allele frequency variation seen for these six loci across the eastern United States are consistent on a more fine grained spatial scale in western Michigan. Malic enzyme, Aconitase-2, Mannose phosphate isomerase, and Hydroxyacid dehydrogenase all displayed significant linear relationships with latitude among five “regional” hawthorn populations sampled along a north-south transect between the cities of Cadillac and Portage, Michigan. Clines were not as evident among “regional” apple populations in western Michigan, although allele frequencies for Malic enzyme¹⁰⁰, Mannose phosphate isomerase¹⁰⁰ and Aconitase-2⁹⁵ varied with latitude among six “local” apple populations within a 60 km² area near the town of Grant. Significant allele frequency differences were observed between hawthorn and apple populations at all “regional” and “local” collecting sites analyzed in the study (a total of 20 different apple and hawthorn populations). As was the case in the geographic survey of the eastern United States, the magnitude and pattern of inter-host frequency differences at “regional” and “local” sites were a function of latitude. Host related genetic differentiation was consistent on a “microgeographic” scale as well. Allele frequencies for Malic enzyme¹⁰⁰ and Aconitase-2⁹⁵ were significantly higher over a four-year period (1984 to 1987) for flies sampled from individual hawthorn trees (N = 6) than apple trees (N = 7) within an old field (0.09-km² area) located near Grant. The fine level of genetic subdivision between hawthorn and apple populations of R. pomonella in western Michigan substantiates the existence of host associated polymorphism in the fly and supports a sympatric mode of divergence for the “apple race”.