Narrow hybrid zone between two subspecies of big sagebrush (Artemisia tridentata: Asteraceae). II. Selection gradients and hybrid fitness

The dynamic equilibrium hypothesis proposes that hybrid zones are stabilized by a balance between dispersal and selection against hybrids. A key prediction of this hypothesis is that hybrids should have lower fitness than either parental taxon, regardless of habitat. Hybrid big sagebrush (Artemisia tridentata ssp. tridentata × ssp. vaseyana) in two narrow hybrid zones do show greatly decreased recruitment. Hybrids in one zone also show increased browsing by grasshoppers, while those in the other zone have increased numbers of aphids. Overall herbivore loads, however, are not greater on the hybrids than on the parental subspecies. Browsing by mule deer is greatest on ssp. vaseyana in both hybrid zones. Incidence of galls is also greatest on ssp. vaseyana in one hybrid zone. Moreover, browsing by Artemisia weevils is greatest on ssp. tridentata in one hybrid zone. Hybrids produce more flowers than either ssp. tridentata or ssp. vaseyana, while seed production rates of hybrids do not differ from those of the parental taxa. Finally, hybrid seeds germinate as well as those of ssp. tridentata and better than those of ssp. vaseyana. Thus, our data do not support the dynamic equilibrium hypothesis.     

DYNAMICS OF A HYBRID ZONE IN PHLOX: AN EXPERIMENTAL DEMOGRAPHIC INVESTIGATION

The dynamics of a narrow hybrid zone between Phlox drummondii ssp. drummondii and ssp. mcallisteri was investigated using an experimental demographic approach. Three populations of each subspecies as well as those from the hybrid zone were involved. Experimental plots containing seeds from each of the nine populations were established in each of the eight sites. Germination, survivorship to reproduction, and fecundity were monitored at 4–6 wk intervals. Finite rates of increase were estimated from these data. There were significant site effects for each demographic variable and rate of increase as well as significant population effects for germination and finite rate of increase. Although there were population effects, no significant differences were found between plant types in sites of the subspecies or the hybrid zone. We reject the hypothesis that the narrow width of the hybrid zone is the result of hybrid disadvantage. We propose that the narrow width is a result of restricted gene flow and discuss the supporting evidence.     

AUTOTETRAPLOIDY IN HAPLOPAPPUS SPINULOSUS HYBRIDS: EVIDENCE FROM NATURAL AND SYNTHETIC TETRAPLOIDS

Haplopappus spinulosus (Asteraceae) is a herbaceous, perennial weed common throughout the western Great Plains of North America and includes both diploid and tetraploid populations. A number of populations in southeastern Colorado were analyzed cytogenetically and morphologically for two reasons. First, initial observations during a routine population survey showed they were morphologically intermediate between the diploid subspecies glaberrimus and spinulosus, suggesting they might have arisen via hybridization. Second, cytological examination revealed that they were tetraploid. Because there was indication of hybrid origin, it was of interest to determine whether the populations were behaving as autopolyploids or segmental allopolyploids. The distinction between these two polyploid types is not easily made since both are likely to form multivalents at meiosis, but equations derived from a model proposed by Jackson and Hauber (1982) have made it possible to determine statistically whether a tetraploid individual is behaving meiotically as an autotetraploid. Meiotic configuration frequencies at diakinesis were determined for each tetraploid plant sampled, and observed frequencies were compared to those expected for an autotetraploid having the same maximum number of chiasmata per bivalent, chiasma frequency and chromosome number. In general, the meiotic behavior of the tetraploids was no different from that expected for autoploids. The initial hypothesis that the populations were derived from hybridization was tested by a detailed cytogenetic and morphological study of the presumed parental subspecies, F, hybrids, and natural putative hybrids. The evidence supports the hypothesis that the natural autotetraploids arose from the hybridization of ssp. glaberrimus and ssp. spinulosus.     

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.     

Environmental (in)dependence of a hybrid zone: Insights from molecular markers and ecological niche modeling in a hybrid zone of Origanum (Lamiaceae) on the island of Crete

The role of environment and the relative significance of endogenous versus exogenous selection in shaping hybrid zones have been crucial issues in the studies of hybridization. Recent advances in ecological niche modeling (ENM) offer new methodological tools, especially in combination with the genotyping of individuals in the hybrid zone. Here, we study the hybrid zone between the widely known spices Origanum onites and Origanum vulgare ssp. hirtum in Crete. We analyze the genetic structure of both parental taxa and their hybrid Origanum × intercendens using AFLP markers on 15 sympatric and 12 allopatric populations and employ ecological niche modeling and niche similarity tests to study their niche patterns. We complement these analyses with seed viability measurements. Our study revealed that the hybridizing taxa O. onites and O. vulgare ssp. hirtum and the resulting genotypic classes showed geographical and environmental niche similarities based on the predictions of ENMs and the subsequent similarity tests. The occurrence of the hybrid zone is not directly dependent on environmental factors which favor the fitness of the hybrid compared to the parental taxa, but rather on aspects such as historical factors and management practices, which may contribute to the localization and maintenance of the contact zone between parental species. Our results suggest that if a minimum required niche differentiation between genotypic classes is not achieved, environmental dependence might not have a prominent role on the outcome of the hybridization.     

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.     

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.     

Role of seed germination in adaptation and reproductive isolation in Arabidopsis lyrata

Seed germination is an important developmental and life history stage. Yet, the evolutionary impact of germination has mainly been studied in the context of dormancy, or for its role in reproductive isolation between species. Here, we aim to examine multiple consequences of genetic divergence on germination traits between two Arabidopsis lyrata subspecies: ssp. petraea (Eurasia) and ssp. lyrata (North America). Postdormancy germination time, a potentially adaptive trait, showed differentiation between the populations, and quantitative trait loci (QTL) mapping revealed that the trait variation is mainly controlled by two antagonistic loci. These QTL areas contain several candidate genes with known function in postdormancy germination in A. thaliana. The sequence variation of three genes was consistent with differential selection, and they also included fixed nonsynonymous substitutions with potential to account for the phenotypic differentiation. We further show that the divergence between the subspecies has led to a slight but significant reduction in hybrid germination proportions, indicating incipient reproductive isolation. Comparison of reciprocal F₁ and F₂ progenies suggests that Bateson–Dobzhansky–Muller incompatibilities likely act through uniparentally inherited factors. Examination of genomewide transmission ratio distortion further revealed that cytonuclear interactions cause substantial pregermination inviability in the hybrids. These results confirm that seed germination has adaptive potential beyond the dormancy stage and that hybrid seed inviability can be one of the first reproductive barriers to arise during divergence.     

Ants exhibit asymmetric hybridization in a mosaic hybrid zone

Research on hybridization between species provides unparalleled insights into the pre‐ and postzygotic isolating mechanisms that drive speciation. In social organisms, colony‐level incompatibilities may provide additional reproductive barriers not present in solitary species, and hybrid zones offer an opportunity to identify these barriers. Here, we use genotyping‐by‐sequencing to sequence hundreds of markers in a hybrid zone between two socially polymorphic ant species, Formica selysi and Formica cinerea. We characterize the zone, determine the frequency of hybrid workers, infer whether hybrid queens or males are produced and investigate whether hybridization is influenced by colony social organization. We also compare cuticular hydrocarbon profiles and aggression levels between the two species. The hybrid zone exhibits a mosaic structure. The asymmetric distribution of hybrids skewed towards F. cinerea suggests a pattern of unidirectional nuclear gene flow from F. selysi into F. cinerea. The occurrence of backcrossed individuals indicates that hybrid queens and/or males are fertile, and the presence of the F. cinerea mitochondrial haplotype in 97% of hybrids shows that successful F1 hybrids will generally have F. cinerea mothers and F. selysi fathers. We found no evidence that social organization contributes to speciation, because hybrids occur in both single‐queen and multiple‐queen colonies. Strongly differentiated cuticular hydrocarbon profiles and heightened interspecific aggression further reveal that species recognition cues are both present and perceived. The discovery of fertile hybrids and asymmetrical gene flow is unusual in ants, and this hybrid zone will therefore provide an ideal system with which to investigate speciation in social insects.     

Genetic divergence and introgressive hybridization betweenAlnus sinuata andA. crispa (Betulaceae)

The actinorhizal genusAlnus contains numerous taxa that have been morphologically classified into different subgenera, species and subspecies. The genetic divergence has been evaluated within subg.Alnobetula between the parapatric taxaAlnus sinuata andA. crispa, using diversity of allozyme markers at 15 structural loci among 20 populations. Evidence for introgressive hybridization at the overlap of their ranges was noted in three populations. However, the width of the hybrid zone appeared tenuous. The average genetic distance derived from the comparisons of conspecific populations was much smaller than the interspecific distance (D = 0.047). This allelic divergence was also paralleled with larger amounts of allelic and genotypic diversity within and among populations ofA. sinuata, which are occupying a more heterogenous ecological niche. It is proposed that the repeated advances and retreats of the ice sheet during the Pleistocene may have promoted the divergence and allopatric evolution of these subspecies, and that secondary contact may have occurred repeatedly during the interglacial periods. The dynamic-equilibrium model would predict in such cases that narrow hybrid zones, formed at the contact of parapatric ranges, would impede gene exchange between parental taxa by selection against hybrids. The results obtained in this study seemed concordant with this hypothesis, as they were also in agreement with the existent taxonomical treatment of these taxa based on morphology.     

Genetic differentiation and inferred dynamics of a hybrid zone between Northern Spotted Owls (Strix occidentalis caurina) and California Spotted Owls (S. o. occidentalis) in northern California

Genetic differentiation among Spotted Owl (Strix occidentalis) subspecies has been established in prior studies. These investigations also provided evidence for introgression and hybridization among taxa but were limited by a lack of samples from geographic regions where subspecies came into close contact. We analyzed new sets of samples from Northern Spotted Owls (NSO: S. o. caurina) and California Spotted Owls (CSO: S. o. occidentalis) in northern California using mitochondrial DNA sequences (mtDNA) and 10 nuclear microsatellite loci to obtain a clearer depiction of genetic differentiation and hybridization in the region. Our analyses revealed that a NSO population close to the northern edge of the CSO range in northern California (the NSO Contact Zone population) is highly differentiated relative to other NSO populations throughout the remainder of their range. Phylogenetic analyses identified a unique lineage of mtDNA in the NSO Contact Zone, and Bayesian clustering analyses of the microsatellite data identified the Contact Zone as a third distinct population that is differentiated from CSO and NSO found in the remainder of the subspecies' range. Hybridization between NSO and CSO was readily detected in the NSO Contact Zone, with over 50% of individuals showing evidence of hybrid ancestry. Hybridization was also identified among 14% of CSO samples, which were dispersed across the subspecies' range in the Sierra Nevada Mountains. The asymmetry of hybridization suggested that the hybrid zone may be dynamic and moving. Although evidence of hybridization existed, we identified no F1 generation hybrid individuals. We instead found evidence for F2 or backcrossed individuals among our samples. The absence of F1 hybrids may indicate that (1) our 10 microsatellites were unable to distinguish hybrid types, (2) primary interactions between subspecies are occurring elsewhere on the landscape, or (3) dispersal between the subspecies' ranges is reduced relative to historical levels, potentially as a consequence of recent regional fires.     

Conservation of genetic diversity in British populations of the diploid endemic Coincya monensis ssp monensis (Isle of Man Cabbage): the risk of hybridisation with the tetraploid alien, Coincya monensis ssp cheiranthos

Coincya monensis is represented in the British flora by two, cytologically distinct subspecies. Coincya monensis ssp monensis is an endemic diploid with a coastal sand dune distribution that includes a number of isolated populations. Coincya monensis ssp cheiranthos is a tetraploid alien, well established in South Wales in early successional habitats. Both subspecies share similar life form traits, flowering times and pollinators. Cluster analysis and phylogenetic reconstruction based on sequences of the mitochondrial nad4 gene confirmed the distinction between alien and endemic taxa. Tetraploid populations carry more polymorphic RAPDs loci and their genetic diversity is partitioned more within than among populations. In contrast, C. monensis ssp monensis has a distinct population genetic structure. Analysis of the multilocus genetic data confirmed a structure of genetically isolated, endemic population clusters in Scotland, Arran, the Isle of Man and South Wales. Experimental hybridisation showed the two subspecies are interfertile. Multivariate analysis of RAPDs data resolved hybrids between alien and endemic clusters and hybrids contained a proportion of alien-specific polymorphic loci. Hybrids of alien maternal parentage contained the mitochondrial nad4 sequence characteristic of the alien subspecies. Since the alien subspecies can invade mobile sand dune communities from urban sites and compete for pollinators, there is a risk that alien and endemic populations will mix and introgress. Conservation of endemic genetic diversity in Britain will require protection for all C. monensis ssp monensis populations. Currently, the most disjunct endemic population in South Wales is most at risk from introgression.     

Molecular patterns of introgression in a classic hybrid zone between the Australian tree frogs,Litoria ewingii and L. paraewingi: evidence of a tension zone

Hybrid zones provide a rare opportunity to explore the processes involved in reproductive isolation and speciation. The southern hybrid zone between the southeastern Australian tree frogs Litoria ewingii and L. paraewingi has been comprehensively studied over the last 40 years, primarily using reproductive compatibility experiments and male advertisement calls. We used mitochondrial DNA (mtDNA) and eight nuclear microsatellite markers to characterize this hybrid zone along a historically studied transect and to test various dispersal‐dependent and dispersal‐independent hybrid zone models. The species are genetically distinct and the level of hybridization within the contact zone is low, with the majority of admixed individuals representing later‐generation hybrids. Based on previous experimental genetic compatibility studies, we predicted that hybrids with L. paraewingi mtDNA would be more frequent than hybrids with L. ewingii mtDNA. Surprisingly, a greater proportion of the identified hybrids had L. ewingii mtDNA. Geographical cline analyses showed a sharp transition in allele frequencies across the transect, and both the mtDNA and microsatellite data showed concordant cline centres, but were best supported by a model that allowed width to vary. Overall, the L. ewingii–L. paraewingi hybrid zone is best characterized as a tension zone, due to the narrow cline width, concordant genetic clines and low levels of hybridization.     

Narrow hybrid zone between two subspecies of big sagebrush (Artemisia tridentata: Asteraceae)

Several species of gall-forming insects specialize on big sagebrush (Artemisia tridentata), a species that shows much clinal and subspecific variation throughout its geographic range. Two of those subspecies, basin big sagebrush (A. t. ssp. tridentata) and mountain big sagebrush (A. t. ssp. vaseyana), form a narrow hybrid zone at Salt Creek, Utah. Reciprocal transplant experiments have shown that the hybrid big sagebrush at Salt Creek are more fit than either parental subspecies, but only in the hybrid zone. Do genotype and environment influence the density and distribution of galls on big sagebrush? We counted galls on parental and hybrid big sagebrush in three reciprocal transplant gardens at Salt Creek. Gardens were in each of the two parental zones and in the hybrid zone. Transplanted seedlings came from five source populations: two parental and three hybrid populations. We identified seven kinds of gall-forming flies (Rhopalomyia midges and Eutreta fruitflies) that produced identifiable galls. Densities of galls varied among the three gardens and five source populations, and there was also a significant garden by source interaction in gall density. In general, variation in gall density among gardens (i.e., environments) was much greater than the variation among source populations (i.e., genotypes). Nevertheless, significant genotype-environment interactions were observed for five of the seven kinds of galls. Overall density of galls, mostly due to Rhopalomyia ampullaria, was greatest in the high-elevation (mountain) garden and least in the low-elevation (basin) garden. To the best of our knowledge, this is the first reciprocal transplant experiment addressing herbivore richness in a hybrid zone.     

The Genus Primula: Section Muscarioides

ABSTRACT

Background: Hybrid zones provide excellent opportunities for studying plant adaptation and speciation.

Aims: We tested whether two herbaceous species of Senecio, S. vernalis and S. glaucus, hybridise in the eastern Mediterranean region and form a hybrid zone across an aridity gradient in the Jordan Rift Valley.

Methods: Allozyme variation surveyed across both species was analysed by the programme STRUCTURE to assign individuals to genetic groups and determine levels of admixture. Populations in the Jordan Rift Valley were subsequently subjected to a cline analysis.

Results: STRUCTURE showed that interspecific hybrids were produced at low frequency along the Israeli coastal plain where S. glaucus is represented by ssp. glaucus. In contrast, hybrids were more commonly produced in central populations of the Jordan Rift Valley. Here, the two species form a hybrid zone with S. vernalis occurring in mesic sites to the north and S. glaucus (ssp. coronopifolius) in arid sites to the south. Cline analysis showed that the hybrid zone is centred towards the northern end of the Dead Sea, but the analysis failed to distinguish how it is maintained.

Conclusions: Future detailed genetic and ecological analysis of the Senecio hybrid zone should improve our understanding of plant adaptation and speciation across aridity gradients.     

RELATIVE SALT TOLERANCE OF CAKILE EDENTULA (BRASSICACEAE) FROM LACUSTRINE AND MARINE BEACHES

Seeds of Cakile edentula ssp. edentula var. edentula (hereafter edentula) were collected from the Pacific Ocean shore of Washington, and Cakile edentula ssp. edentula var. lacustris (hereafter lacustris) seeds were collected from the Lake Huron shore of Ontario. Germination rates of the two varieties showed equal tolerance of substrate salinities ranging from 0–10,000 ppm. Early root growth of edentula seedlings was significantly stimulated by 1,000 ppm salts, but otherwise both varieties showed equal tolerance to 10,000 ppm. Edentula plants exposed for 4 wk to salt spray intensities of 0, 20, and 90 mg dm^–2 d^–1, applied in a growth chamber environment, showed no significant changes in shoot or root biomass, plant morphology, or reproductive effort. Lacustris plants exhibited significant declines in stem length and reproductive effort with increasing levels of salt spray. Our results suggest that lacustris and edentula have diverged physiologically over the past 9,000 years, and field observations from the literature corroborate our conclusion that this divergence in salt tolerance has ecological significance.     

Chromosome evolution in the Gibasis linearis group (Commelinaceae)

Karyotypes, DNA amounts, and meiotic behaviour were examined in population samples of two closely related species, Gibasis venustula and G. heterophylla, and their F₁ hybrids. All samples were diploid (2n=12). DNA amount was similar in G. heterophylla, G. venustula ssp. robusta, and some populations of G. venustula ssp. venustula but in 13 samples of the latter, it showed as much as a 60% difference despite karyotypic uniformity. Although DNA levels were related to the altitude of the habitat, there was some variation within two morphological and ecological races of subspecies venustula. Analysis of hybrid meiosis and pollen fertility demonstrated the cytogenetical discreteness of populations, races, and subspecies. Populations and races were concluded still to be evolving following ecological isolation. The significance of quantitative and qualitative genome changes in evolution and their bearing on the classification of these species is discussed.     

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.     

Secondary intergradation and genome incompatibility in Phlox pilosa (polemoniaceae)

Phlox pilosa ssp.pilosa andP. pilosa ssp.fulgida form a zone of secondary intergradation extending from northern Illinois and Indiana along the prairie-forest border to southwestern Missouri. The structure of the zone was analyzed in Illinois and Indiana. Trichome morphology offered the only diagnostic characteristic. Character gradients in the zone are highly irregular and steeply sloping. Closely adjacent populations often have disparate phenotype distributions. Most populations are strongly skewed toward the parental ends of the morphological spectrum. The properties of the zone are attributed to genome incompatibility, which is seen in the crossability barriers between the two subspecies and between the subspecies and hybrids, a reduction in pollen fertility, anther size and fecundity in hybrids, and the formation of multiple nucleoli in the hybrids. Subspeciesfulgida appears to be the primary recipient of alien gene flow. However, the position of the zone ostensibly is stabilized by the presence of an ecological barrier.