DEVELOPMENTAL STABILITY,FITNESS, AND TRAIT SIZE IN LABORATORY HYBRIDS BETWEEN EUROPEAN SUBSPECIES OF THE HOUSE MOUSE

The effects of hybridization on developmental stability and size of tooth characters were investigated in intersubspecific crosses between random-bred wild strains of the house mouse (Mus musculus domesticus and M. m. musculus). Fluctuating asymmetry (FA) and trait size were compared within and between parental, F₁, backcross, and F₂ hybrid groups. The relationship between FA and reproductive fitness within the F₁ hybrids was also studied. The results indicated that both FA and character size levels differed significantly between the two subspecies. The F₁ hybrids and the recombined groups (backcrosses and F₂ hybrids) showed heterosis for both parameters. No significant differences in the FA of fertile and sterile F₁ hybrid individuals were found. Comparison of the FA levels obtained in this study with those found in wild populations from the hybrid zone in Denmark showed that the levels of FA were lower in laboratory-bred samples than in the wild populations. This study provides further evidence that, in hybrids, the developmental processes underlying most of the morphological traits we studied benefit from a heterotic effect, despite the genomic incompatibilities between the two European house mice revealed by previous genetical and parasitological studies.     

MORPHOLOGICAL VARIABILITY IN GENETICALLY DEFINED CATEGORIES OF ANURAN HYBRIDS

Hybridization phenomena in anurans have traditionally been studied through morphological comparisons, under the assumption that various hybrids (e.g., F₁‘s, backcrosses) are predictably intermediate to parental species. We critically evaluate this assumption by examining morphology in genetically categorized hybrids between the treefrogs Hyla cinerea and H. gratiosa. A total of 202 frogs from a hybridizing population in Alabama were assayed for allozyme and mitochondrial DNA genotype and for a large suite of osteological characters. Discriminant analyses demonstrated distinct morphological separation between the genetically “pure” parental species. Morphometric analyses of genetically identified hybrids showed: 1) an extreme range of phenotypic expression within F₁ and backcross classes, and 2) no apparent directional parental bias on the F₁ phenotype. Had morphology alone been used as a guide, over 40 percent of the individuals with known hybrid ancestry would have been misclassified as “pure” parental species, and about 25 percent of the backcross individuals would not have been distinguished from F₁‘s. These results exemplify the utility of joint comparisons of morphology and genotypic constitution in studies of natural hybridization, and they emphasize the limitations inherent in describing hybrid classes solely by morphological criteria.     

HETEROSIS AND OUTBREEDING DEPRESSION IN INTERPOPULATION CROSSES SPANNING A WIDE RANGE OF DIVERGENCE

The intertidal copepod Tigriopus californicus was used as a model organism to look at effects of crossing distance on fitness and to investigate the genetic mechanisms responsible. Crosses were conducted between 12 pairs of populations spanning a broad range of both geographic distance (5 m to 2007 km) and genetic distance (0.2% to 22.3% sequence divergence for a 606-bp segment of the mitochondrial COI gene). For each pair of populations, three fitness components (hatching number, survivorship number, and metamorphosis number) were measured in up to 16 cohorts including parentals, reciprocal F₁, F₂, F₃, and first-generation backcross hybrids. Comparisons of each set of cohorts allowed estimation of within- and between-locus gene interaction. Relative to parentals, F₁ hybrids showed a trend toward increased fitness, with no correspondence with population divergence, and a decrease in variance, which in some cases correlated with population divergence. In sharp contrast, F₂ hybrids had a decrease in fitness and an increase in variance that both corresponded to population divergence. Genetic interpretation of these patterns suggests that both the beneficial effects of dominance and the detrimental effects of breaking up coadaptation are magnified by increasing evolutionary distance between populations. Because there is no recombination in T. californicus females, effects of recombination can be assessed by comparing F₁ hybrid males and females backcrossed to parentals. Both recombinant and nonrecombinant backcross hybrids showed a decline in fitness correlated with population divergence, indicating that segregation among chromosomes contributes to the breakup of coadaptation. Although there was no difference in mean fitness between the two backcross types, recombinational backcrosses showed greater variance for fitness than nonrecombinational backcrosses, suggesting that the breakup of parental gene ombinations within chromosomes has both beneficial and detrimental effects.     

Characterization of a contemporaneous hybrid zone between two darter species (Etheostoma bison and E. caeruleum) in the Buffalo River System

Hybrid zones have long intrigued evolutionary biologists and provide a natural laboratory to explore the evolution of reproductive isolation (speciation). Molecular characterization of hybrid zone dynamics can provide insight into the strength of reproductive isolation as well as the underlying evolutionary processes shaping gene flow. Approximately one-third of darter species naturally hybridize making this species-rich North American freshwater teleost fish clade an ideal system to investigate the extent and direction of hybridization. The objective of this study was to use diagnostic microsatellite markers to calculate genetic hybrid index scores of two syntopic, but distantly related darter species, Etheostoma bison and Etheostoma caeruleum. A combination of hybrid index scores, assignment tests, and mitochondrial haplotype profiles uncovered mixed ancestry in approximately 6 % of sampled adult individuals, supporting contemporaneous hybridization that was previously undocumented in E. bison. Moreover, hybrids were not limited to the F₁ generation, but encompassed the entire suite of hybrid categories (F₁, F₂ and backcross hybrids). The low number of hybrids assigned to each hybrid category represents a bimodal hybrid zone, suggesting reproductive isolation is strong (but incomplete) and also advocates for the ability of hybrids to produce second-generation hybrids and backcross into both parental species, mediating introgression across species boundaries. To this end, cytonuclear profiles of the sampled parental species and hybrids were consistent with bidirectional gene flow, although there was an overall trend of asymmetric hybridization between E. caeruleum females and E. bison males. The spatiotemporal variation in hybridization rates and resulting cytonuclear patterns expanded on in this study provide a comparative genetic framework on which future studies can begin to elucidate the underlying processes that not only generate a mosaic hybrid zone, but maintain the distinctness of species in the face of gene flow.     

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.     

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.     

BIOCHEMICAL PHENOTYPIC AND GENETIC STUDIES OF TWO INTRODUCED FIRE ANTS AND THEIR HYBRID (HYMENOPTERA: FORMICIDAE)

Two introduced fire ants, Solenopsis invicta and S. richteri, and their hybrid were studied using phenotypic markers from gas chromatographic analysis of hydrocarbons and venom alkaloids, as well as genetic markers from enzyme electrophoresis. Both methods show that extensive gene introgression is occurring over a distance of at least 120 km at the contact zone between the two forms in eastern Mississippi. Genetic analyses suggest that the hybrid population does not depart dramatically from panmixia. Also, recombinant genotypes predominate in the hybrid zone, indicating that F₁ hybrids are viable. Allele frequency clines through the hybrid zone are apparent for four polymorphic loci. Data sets generated by the chromatographic and electrophoretic methods are highly concordant in that they differentiate completely between the two forms and agree in designating colonies from the contact zone as hybrid or parental in a high proportion (90%) of cases. The two methods can serve as complementary tools for studying closely related but genetically distinct populations in this, and perhaps other, groups of insects.     

Genetic introgression of hybrid Rhododendron x intermedium Tausch is habitat mediated: Evidences from south-eastern Alps (Italy)

Hybridization between Rhododendron ferrugineum L. and R. hirsutum L. in south-eastern Alps was examined in order to (i) evaluate the breeding direction and the extent of backcrossing between hybrids and the parental species, (ii) define which processes facilitate speciation and maintain species identities and (iii) clarify the role of rock geochemistry in hybridization events. Individuals of three hybrid populations were analysed by morphological and molecular markers. The internal transcribed spacer and trnH–psbA distinguished the parental species and F₁ hybrids while only the simple sequence repeat markers recognized genotype classes: F, H, F₁, F₂, BxF (backcross to R. ferrugineum) and BxH (backcross to R. hirsutum). Combining morphological and molecular data, we found that the tested populations had complex genetic structure: the F₁ individuals produce F₂ hybrids and backcross to parental species. Due to R. hirsutum phenology, most backcrossing events were with this parental species (asymmetric hybridization). Geochemical analyses indicate that alkaline soil conditions linked to calcareous dolomitic rocks promoted the genetic assimilation of R. hirsutum. In addition, R. x intermedium shows a higher edaphic adaptation than R. hirsutum as it can be found on a wide range of calcareous-dolomitic rocks as well as on weakly acidic soils of natural or anthropogenic origin.     

RIBOSOMAL-DNA,MITOCHONDRIAL-DNA,CHROMOSOMAL, AND ALLOZYMIC STUDIES ON A CONTACT ZONE IN THE POCKET GOPHER,GEOMYS

We studied 75 individuals of the Plains pocket gopher, Geomys bursarius, from eastern New Mexico, where the subspecies major and knoxjonesi hybridize. Each individual was examined for chromosome number, ribosomal DNA, mitochondrial DNA, and three protein systems for which reference parental populations were fixed for alternative alleles. Twenty individuals were indistinguishable from parental major, 14 individuals were indistinguishable from parental knoxjonesi, and 41 individuals had genotypes composed of combinations of character states that distinguish the two parental types. The parental types appear to represent discrete genetic entities that have restricted introgression across a narrow hybrid zone (width approximately 3 km, using the 20/80 criterion). Parental types overlap in geographic distribution near the center of the zone, and changes in mitochondrial DNA and the five nuclear markers are concordant across the zone. It is probable that there is premating isolation between knoxjonesi males and major females. The frequencies of individuals with certain genotypic combinations within our sample imply differential reproductive success of certain genotypes. We propose that F₁'s and highly heterozygous males are sterile and that hybrid females are less fertile than parental females. These postmating factors, along with premating isolation for one of the reciprocal crosses, probably account for the restriction of gene flow across the contact zone. The structure of the zone can be explained by the “dynamic equilibrium” model.     

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.     

Morphological variation of genetically confirmed Alouatta Pigra × A. palliata hybrids from a natural hybrid zone in Tabasco,Mexico

While hybridization has been reported for a large number of primate taxa, there is a general lack of data on hybrid morphology for wild individuals with known genetic ancestry. A confirmed hybrid zone for the closely related Neotropical primates Alouatta palliata and A. pigra has provided a unique opportunity to study primate hybrid morphological variation. Here we used molecular evidence based on mitochondrial, Y‐chromosome, and autosomal data to assess hybrid ancestry. We conducted univariate and multivariate statistical comparisons of morphometric data collected from individuals both outside and within the hybrid zone in Tabasco, Mexico. Our results show that of all the hybrids detected (N = 128), only 12% of them were approximately genetically intermediate, and none of them were first generation hybrids. Univariate pairwise comparisons among parental individuals, multigenerational backcrossed hybrids, and intermediate hybrids showed that overall, multigenerational backcrossed hybrids resemble the parental species with which they share most of their alleles. Conversely, intermediates were highly variable. Similarly, principal component analysis depicts an overlap between the parental species and their backcrosses when considering overall morphological differences. Finally, discriminant function analysis of the morphological variables was overall unreliable for classifying individuals into their assigned genotypic classes. Taken together, our results suggest that primate natural hybridization studies should incorporate molecular methods for determining ancestry, because morphology may not always be a reliable indicator of hybrid status. Hybrid zones could comprise a large number of multigenerational backcrossed hybrids that are indistinguishable from the parental species. The implications for studying hybridization in the primate fossil record are discussed. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

Discordant patterns of introgression across a narrow hybrid zone between two cryptic lineages of an Iberian endemic newt

The study of natural hybrid zones can illuminate aspects of lineage divergence and speciation in morphologically cryptic taxa. We studied a hybrid zone between two highly divergent but morphologically similar lineages (south‐western and south‐eastern) of the Iberian endemic Bosca's newt (Lissotriton boscai) in SW Iberia with a multilocus dataset (microsatellites, nuclear and mitochondrial genes). STRUCTURE and NEWHYBRIDS analyses retrieved few admixed individuals, which classified as backcrosses involving parental individuals of the south‐western lineage. Our results show asymmetric introgression of mtDNA beyond the contact from this lineage into the south‐eastern lineage. Analysis of nongeographic introgression patterns revealed asymmetries in the direction of introgression, but except for mtDNA, we did not find evidence for nonconcordant introgression patterns across nuclear loci. Analysis of a 150‐km transect across the hybrid zone showed broadly coincident cline widths (ca. 3.2–27.9 km), and concordant cline centres across all markers, except for mtDNA that is displaced ca. 60 km northward. Results from ecological niche modelling show that the hybrid zone is in a climatically homogenous area with suitable habitat for the species, suggesting that contact between the two lineages is unlikely to occur further south as their distributions are currently separated by an extensive area of unfavourable habitat. Taken together, our findings suggest the genetic structure of this hybrid zone results from the interplay of historical (biogeographic) and population‐level processes. The narrowness and coincidence of genetic clines can be explained by weak selection against hybrids and reflect a degree of reproductive isolation that is consistent with cryptic speciation.     

Developmental stability is not disrupted by extensive hybridization and introgression among populations of the marine bivalve molluscs Mytilus edulis (L.) and M. galloprovincialis (Lmk.) from south-west England

The effects of hybridization and introzgression were assessed among two naturally hybridizing bivalve molluscs (the mussels Mytilus edulis and M. galloprovincialis) from western Europe to estimate how disruptive these processes are to developmental stability (measured in terms of morphological variability). Ten shell traits were measured for 392 mussels from four allopatric populations (two each of At. edulis and At. galloprovincialis) and two hybrid populations. An index of variability (defined as Z_i= |y_i–y_i where y_; is the population mean of the length-standardized trait, and y_i is the individualcar;s length-standardized trait value) was constructed for each trait, and for the sum of the traits. The hybrid populations did not exhibit greater mean variability than the allopatric populations for any of the indices. Upon pooling, the hybrid populations had significantly lower variability than the pooled M. edulis populations and the pooled M. galloprovincialis populations in two analyses, and had similar means in the remaining nine analyses. Where significant differences existed, the pooled M. galloprovincialis had lower levels of mean trait variability than the pooled M. edulis. Among the two hybrid populations, the putative Fl hybrids and backcross individuals exhibited means of trait variability which were similar to those of the parental types. Thus, extensive hybridization and introgression do not adversely affect developmental stability among these mussel populations. There was a strong significant correlation between the ranking of indices (based on the amount of variability) across all six populations, indicating that a large genetic component determines the measured morphological variability. It is concluded that the genes or gene complexes which control morphological development in M. edulis and M. galloprovincialis arc very similar, providing further evidence of the close evolutionary relatedness of these mussel taxa.     

Fitness of natural willow hybrids in a pioneer mosaic hybrid zone

Hybrid fitness is an important parameter to predict the evolutionary consequences of a hybridization event and to characterize hybrid zones. We studied fitness parameters of F₁ and later‐generation hybrids between the lowland species Salix purpurea and the alpine S. helvetica that have recently emerged during colonization of an alpine glacier forefield. Fruit production (number of capsules per catkin and fruit set) did not differ between hybrids and parents, but the number of seeds per capsule of F₁ hybrids was slightly lower than that of later‐generation hybrids and of the parents. Germination rates and seedling growth were tested on three substrates (pH 4.5, 7.0, and 8.0). Germination rates of seeds collected from F₁ hybrids were lower on acid and neutral substrates, but equal at pH 8.0 compared to all other groups, while the seeds from later‐generation hybrids performed as well as the parents on all three substrates. In seedling growth, the colonizer S. purpurea performed better than all other taxa on all three substrates, while hybrids resembled the subalpine species S. helvetica. Results suggest that endogenous selection acts against F₁ hybrids, but favors fitter genotypes in later‐generation hybrids. Exogenous selection via soil pH appears to be weak during seedling establishment. The pioneer vegetation on the glacier forefield may offer sufficient niche space for hybrid seedlings. Owing to the relatively high fitness of the hybrids and the scattered distribution of hybrids and parental individuals on the glacier forefield, this hybrid zone can be assigned to a mosaic model, probably facilitating gene flow and introgression between the parental species. As establishment of the hybrid zone appears to be linked to a colonization process, we propose to call it a pioneer mosaic hybrid zone.     

ESTIMATING THE MATING BEHAVIOR OF A PAIR OF HYBRIDIZING HELICONIUS SPECIES IN THE WILD

Premating isolation between incipient species is rarely studied in nature, even though mating tests in captivity may give an inaccurate picture of natural hybridization. We studied premating barriers between the warningly colored butterflies Heliconius erato and H. himera (Lepidoptera) in a narrow contact zone in Ecuador, where hybrids are found at low frequency. Eggs obtained from wild-mated females, supplemented with eggs and young larvae collected from the wild, were reared to adulthood. Adult color patterns of these progeny were then used to infer how their parents must have mated. Likelihood was used to estimate both the frequencies of potential parental genotypes from adult phenotypes collected in the wild, and the degree of assortative mating from the inferred parents. The frequencies of parental genotypes varied across the hybrid zone, but our statistical method allowed estimates of hybrid deficit and assortative mating to be integrated across all sites sampled. The best estimate of the frequency of F₁ and backcross hybrid adults in the center of the hybrid zone was 10%, with support limits (7.1%, 13.0%; support limits are asymptotically equivalent to 95% confidence limits). Mating was highly assortative: in the center of the hybrid zone the cross-mating probability between H. erato and H. himera was only 5% (0.3%, 21.4%). Wild hybrids themselves mated with both pure forms, and the probabilities that they mated in any direction were not significantly lower than those among conspecifics. These results are consistent with earlier laboratory studies on mate choice, and suggest that selection against hybrids must be strong to prevent formation of a hybrid swarm. Unfortunately, the wide support limits on mating behavior precluded a measure of the strength of selection from these data alone. Our statistical approach provides a useful general method for estimating mate choice in the wild.     

Genetic structure of a hybrid zone between two violets,Viola rossii Hemsl. and V. bissetii Maxim.: dominance of F1 individuals in a narrow contact range

The genetic composition of a hybrid zone can provide insight into the evolution of diversification in plants. We carried out morphological and amplified fragment length polymorphism analyses to investigate the genetic composition of a hybrid zone between two violets, Viola bissetii Hemsl. and Viola rossii Maxim. Our aim was to clarify the formation and maintenance of hybrids between these Viola species. We found that most hybrid individuals (V. bissetii × V. rossii) were of the F₁ generation, with a few of the F₂ generation. We found no backcrosses. The scarcity of post‐F₁ hybrids indicates that a species barrier is established between the parental species. The F₁‐dominated hybrid zone occupied only a narrow, intermediate ecotone between the parental habitats, suggesting that selection by environmental factors against hybrids may help to maintain the current conditions in this hybrid zone.     

POPULATION VARIATION AND HYBRIDIZATION IN SEA-ROCKETS (CAKILE,CRUCIFERAE): SEED GLUCOSINOLATE CHARACTERS

In a study of intra- and interpopulation variation in seed glucosinolates, remarkable uniformity was found among individuals within the two chemically distinct subspecies of East Coast Cakile edentula. Glucosinolates were used as taxonomic markers in a study of hybridization between ssp. edentula and ssp. harperi where they form mixed populations on the Outer Banks of North Carolina. Thirteen plants with hybrid chemical profiles were detected in a sample of 89 individuals from four populations; the zone of hybridization is narrow, and has persisted for at least eight years. Artificial hybrids of the two subspecies are additive for kinds and intermediate for proportions of seed glucosinolates, and families of F₂'s show a range of recombinant chemical profiles. Natural hybridization was confirmed by recovering hybrid and parental types in 47 progeny of three presumptive wild hybrids. Glucosinolate profile characteristics segregated independently of the segregation patterns for fruit length and seed weight. Selection against particular glucosinolate phenotypes is one possible mechanism maintaining this narrow hybrid zone on the Outer Banks.     

Patterns of color and nectar variation across an Ipomopsis (Polemoniaceae) hybrid zone

Hybridization may uncouple adaptive trait combinations that are present in parental species. I studied variation in flower color and reward quality across a hybrid zone of Ipomopsis aggregata and I. tenuituba. Individuals from hybrid populations showed considerable variation in flower color using corolla reflectance measurements. Flower spectra of such individuals were either intermediate or else resembled those flowers from the parental species. Ipomopsis aggregata populations had consistently higher nectar production rates and higher nectar standing crops than either I. tenuituba or hybrids. Ipomopsis aggregata flowers also produced more dilute nectar than those of hybrids and I. tenuituba, but the actual concentration values were quite variable among populations of the same type. Overall, the nectar quality of hybrid flowers most resembled that of I. tenuituba flowers. Based on the observed interpopulation patterns of color-reward associations in this hybrid zone, pollinators should be able to discriminate against I. tenuituba and hybrid populations and against most individuals within hybrid populations. However, they may visit those hybrids that resemble the most rewarding flower type (I. aggregata). The results emphasize the need for studies that address how hybridization affects subsequent plant fitness and the evolutionary dynamics of the species involved.     

HYBRIDIZATION STUDIES ON THE HOST RACES OF EUROSTA SOLIDAGINIS: IMPLICATIONS FOR SYMPATRIC SPECIATION

We studied the inheritance of survival ability in host-associated populations of the tephritid fly, Eurosta solidaginis, to test predictions of sympatric speciation models. Eurosta solidaginis induces galls on two species of goldenrod, Solidago altissima and S. gigantea. The host-associated populations have been hypothesized to be host races that originated in sympatry (Craig et al. 1993). We found evidence for disruptive selection for host use, which is a critical assumption of sympatric speciation models. Each host race had higher survival rates on their host plant than on the alternative host. F₁ and backcross hybrids also had lower survival rates than the pure host-race flies on their host plant. Since assortative mating occurs due to host-plant preference (Craig et al. 1993) this would select for divergence in host preference. Low hybrid survival could have been due to strong genetic incompatibilities of the populations or due to host adaptation by each population. Strong genetic incompatibilities would result in poor survival on all host plants, while host adaptation could result in low overall survival with high hybrid survival on some host plants with particularly “benign” environments. High survival of F₁, F₂, and backcross hybrids on some plant genotypes in some years supported the host adaptation hypothesis. F₁ flies mated and oviposited normally and produced viable F₂ and backcross hybrids indicating gene flow is possible between the host races. A few flies developed and emerged on the alternative host plant. This demonstrates that genes necessary to utilize the alternative host exist in both host races. This could have facilitated the origin of one of the populations via a host shift from the ancestral host. The inheritance of survival ability appears to be an autosomal trait. We did not find evidence that survival ability was maternally influenced or sex linked.