Role of micromorphological leaf traits and molecular data in taxonomy of three sympatric white oak species and their hybrids (Quercus L.)

Hybridisation and introgression occur with high frequency in the genus Quercus and interspecific hybrid individuals show patterns of morphological traits that might be influenced in different ways. Micromorphological leaf traits appear to be positive and stable in Quercus species, and by combining genetic and micromorphological analyses, it is possible to compare the patterns of variation in micromorphological leaf traits of pure and hybrid individuals. Trichomes and stomatal traits were examined using scanning electron microscopy at 150–2000 × magnification in sympatric oak species collected in a natural deciduous wood. Q. frainetto, Q. petraea and Q. pubescens appear to have a relatively predictable complement of trichome types. Both the pattern and quantitative values of each micromorphological trait examined (stomata and trichomes) have an important role in identifying hybrids and pure species; putative hybrids show a pattern of trichomes that is a combination of the parental types. These results, combined with the fact that micromorphological traits generally exhibit higher consistency, indicate that this source of information can be an excellent clue to hybridisation and introgression and useful in taxonomical, systematic and evolutionary studies on the European white oaks.     

Utility of multilocus genotypes for taxon assignment in stands of closely related European white oaks from Switzerland

Background and Aims

European white oaks (Quercus petraea, Q. pubescens, Q. robur) have long puzzled plant biologists owing to disputed species differentiation. Extensive hybridization or shared ancestry have been proposed as alternative hypotheses to explain why genetic differentiation between these oak species is low. Species delimitation is usually weak and often shows gradual transitions in leaf morphology. Hence, individual identification may be difficult, but remains a critical step for both scientific work and practical management.

Methods

Multilocus genotype data (five nuclear microsatellites) were used from ten Swiss oak stands for taxon identification without a priori grouping of individuals or populations, using model-based Bayesian assignment tests.

Key Results

Three groups best structured the data, indicating that the taxonomical signal was stronger than the spatial signal. Most individuals showed high posterior probabilities for either of three genetic groups that were best circumscribed as taxonomical units. The assignment of a subset of trees, whose taxonomic status had been previously characterized in detail, supported this classification scheme.

Conclusions

Molecular-genetic assignment tests are useful in the identification of species status in critical taxon complexes such as the European white oaks. Such an approach is of practical importance for forest management, e.g. for stand certification or in seed trade to trace the origin of forest products.Key words: Assignment test, Bayesian inference, multilocus genotype, nuclear microsatellites, Quercus sp., species complex     

Molecular and morphological support for a Florida origin of the Cuban oak

Aim The origins of the Cuban biota are of long‐standing interest in biogeography, and the source of a small live oak (Quercus series Virentes) population on Cuba remains unresolved. Based on morphological evidence, previous authors have hypothesized a Florida origin from either Q. geminata or Q. virginiana or both; a Mexican origin from Q. oleoides; or a hybrid origin from both sources. We use molecular data and taxonomically informative leaf morphology to identify the source species and timing of colonization. Location Cuba, Central America, Mexico and the south‐eastern United States. Methods  We collected representative samples of Cuban oaks and each putative source species and genotyped each sample at 12 nuclear microsatellites and two chloroplast DNA sequences. We estimated population structure using a Bayesian clustering analysis and F‐statistics, pairwise migration rates among taxa, and divergence time using an isolation‐with‐migration model. We measured seven leaf traits and conducted an analysis of similarity (ANOSIM) to determine which putative source species was most similar to Cuban oaks. Results Cuban oak contains one chloroplast DNA haplotype, which is common in southern Florida. Bayesian clustering analysis of microsatellites revealed that the Cuban oak forms a distinct and pure population cluster, and F‐statistics showed that Cuban oaks are differentiated least from Q. virginiana and most from Q. geminata. Migration rates were highest out of Cuba to Q. oleoides. Molecular diversity, the ratio of allelic richness to allele size range, and effective population size of the Cuban oak were relatively low, suggesting a founder effect. Divergence time estimates fell entirely within the Pleistocene (628–6 ka), considering a range of mutation rates and generation times. Cuban oaks were morphologically most similar to Q. virginiana and least similar to Q. geminata. Main conclusions Molecular and morphological data support a Pleistocene dispersal of Q. virginiana from Florida to Cuba, followed by isolation and divergence, then limited dispersal and introgression from Cuba to Q. oleoides in Central America. Birds could have dispersed acorns to Cuba during a glacial period when sea levels were low. These results highlight the varied origin of the Cuban biota and the possible role of Pleistocene glaciations in the establishment of temperate taxa in the tropics.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

Combining geometric morphometrics with molecular genetics to investigate a putative hybrid complex: a case study with barbels Barbus spp. (Teleostei: Cyprinidae)

This integrative study examined the morphological and genetic affinities of three endemic barbel species from Italy (brook barbel Barbus caninus, Italian barbel Barbus plebejus and horse barbel Barbus tyberinus) and of putative hybrid specimens to their species of origin. Two of the species frequently occur together with the non‐native barbel Barbus barbus. DNA barcoding indicates that mitochondrial (mt) haplotypes often do not match the species expected from morphology. Linear distance measurements and meristics are not informative for discrimination of the species and putative hybrids, but a discriminant analysis of principal components (DAPC) of geometric landmark data produces reassignments largely in congruence with mt and nuclear genetic data. Cyto‐nuclear conflicts confirm the presence of hybridization in B. plebejus and B. tyberinus and identify additional introgressed specimens. A comparison between mixed genotypes and their morphology‐based assignment reveals no predictable pattern. The finding that most individuals of the morphologically similar B. plebejus and B. tyberinus have very high assignment probabilities to their respective species suggests that the presented approach may serve as a valuable tool to distinguish morphologically very similar taxa.     

Phylogeography and climatic niche evolution in live oaks (Quercus series Virentes) from the tropics to the temperate zone

Aim We investigated the phylogeography, geographical variation in leaf morphology, freezing tolerance and climatic niches of two widespread evergreen sister oak species (Quercus) in the series Virentes. Location South‐eastern USA, Mexico and Central America. Methods Nuclear microsatellites and non‐recombining nuclear and chloroplast DNA sequences were obtained from trees throughout the range of two sister lineages of live oaks, represented by Quercus virginiana in the temperate zone and Q. oleoides in the tropics. Divergence times were estimated for the two major geographical and genetic breaks. Differentiation in leaf morphology, analysed from field specimens, was compared with the molecular data. Freezing sensitivities of Q. virginiana and Q. oleoides populations were assessed in common garden experiments. Results The geographical break between Q. virginiana and Q. oleoides was associated with strong genetic differentiation of possible early Pleistocene origin and with differentiation in freezing sensitivity, climatic envelopes and leaf morphology. A second important geographical and genetic break within Q. oleoides between Costa Rica and the rest of Central America showed a mid‐Pleistocene divergence time and no differentiation in leaf morphology. Population genetic differentiation was greater but genetic diversity was lower within the temperate Q. virginiana than within the tropical Q. oleoides, and genetic breaks largely corresponded to breaks in leaf morphology. Main conclusions Two major breaks, one between Mexico and the USA at the boundary of the two species, and a more recent one within Q. oleoides between Honduras and Costa Rica, implicate climatic changes as potential causes. The latter break may be associated with the formation of the Cordillera de Guanacaste, which was followed by seasonal changes in precipitation. In the former case, an ‘out of the tropics’ scenario is hypothesized, in which the acquisition of freezing tolerance in Q. virginiana permitted colonization of and expansion in the temperate zone, while differences in climatic tolerances between the species limited secondary contact. More pronounced Pleistocene changes in climate and sea level in the south‐eastern USA relative to coastal Mexico and Central America may explain the greater population differentiation within temperate Q. virginiana and greater genetic diversity in tropical Q. oleoides. These patterns are predicted to hold for other taxa that span temperate and tropical zones of North and Central America.     

Effects of plant hybridization on the structure and composition of a highly rich community of cynipid gall wasps: the case of the oak hybrid complex <Emphasis Type="Italic">Quercus magnoliifolia</Emphasis> x <Emphasis Type="Italic">Quercus resinosa</Emphasis> in Mexico

The richness and composition of herbivore communities can be influenced by the genetic variation of host plants. Hybrid plant populations are ideal to test these effects because they usually harbor high genetic variation and display a mosaic of phenotypic characters. The goal of this study was to examine the effect of hybridization between two Mexican white oaks, Q. magnoliifolia and Q. resinosa, on the composition and diversity of the associated cynipid gall wasp community. We used eight nuclear microsatellite markers to genotype 150 oak individuals sampled at three different altitudes at the Tequila volcano and conducted monthly samplings of galls in each individual over the course of 2 years. A Bayesian assignment analysis indicated genetic admixture between the two oak species at the study site and allowed classifying individuals as Q. magnoliifolia, Q. resinosa or hybrids. Gall morphospecies richness was significantly higher in the hybrids, intermediate in Q. magnoliifolia and lower in Q. resinosa. Overall, 48 different gall morphospecies were found, with 21 of them being shared among the three groups of plants, 13 between two groups of plants, and 14 were unique to one group of plants, with eight of these being found in hybrids. Several of the shared galls showed differences in abundance among plant groups. Therefore, genetic structure in this oak complex significantly influences the diversity and composition of the associated gall wasp community, and hybrid individuals are probably acting as potential sinks and bridges for the colonization of plant hosts by these highly specialized insect species.     

Coincidence of small-scale spatial discontinuities in leaf morphology and nuclear microsatellite variation of Quercus petraea and Q. robur in a mixed forest

BACKGROUND AND AIMS: The taxon complex comprising Quercus petraea and Q. robur shows distinct morphologies and ecological preferences, but mostly low differentiation in various types of molecular markers at a broad spatial range. Local, spatially explicit analyses may reveal patterns induced by microevolutionary processes operating mainly over short distances. However, no attempts have been made to date to explore the potential of spatial analyses combining morphological and genetic data of these oaks. METHODS: A mixed oak stand was studied to elucidate the small-scale population genetic structure. All adult individuals were classified and putative hybrids were identified using multivariate discrimination analysis of leaf morphological characters. Likewise, all trees were genotyped with five nuclear microsatellites, and a Bayesian assignment method was applied based on maximum likelihood of multilocus genotypes for taxon and putative hybrid classification. KEY RESULTS: Multivariate analyses of leaf morphological data recognized two groups with few individuals as putative hybrids. These groups were significantly differentiated at the five microsatellites, and genetic taxon assignment coincided well with morphological classification. Furthermore, most putative hybrids were assigned to the taxon found in their spatial neighbourhood. When grouping trees into clusters according to their spatial positions, these clusters were clearly dominated by one taxon. Discontinuities in morphological and genetic distance matrices among these clusters showed high congruence. CONCLUSIONS: The spatial-genetic analyses and the available literature led to the assumption that reproductive barriers, assortative mating, limited seed dispersal and microsite-induced selection in favour of the locally adapted taxon at the juvenile stage may reinforce taxon-specific spatial aggregation that fosters species separation. Thus, the results tend to support the hypothesis that Q. petraea and Q. robur are distinct taxa which share a recent common ancestry. Occasional hybrids are rarely found in adults owing to selection during establishment of juveniles.     

Dermaphis coccidiformis sp. nov. (Hemiptera), an aphid species with asymmetrically sclerotized apterae and “winter alates”

Dermaphis coccidiformis sp. nov. (Hormaphidinae: Nipponaphidini) is described from Japan. Apterous adults of the species were found between winter buds (or between a winter bud and a leaf petiole) of the evergreen oaks Quercus glauca, Q. myrsinifolia and Q. salicina. Their morphology is peculiar in that their tergites are heavily sclerotized only in the part that seems to have been exposed to sunlight. The new species is also peculiar in that nymphs to be alates (sexuparae) were found on the upper surfaces of leaves of the host oak only during winter, from December to March or early April, before the bud break of the oak. Our molecular phylogenetic analysis indicated that the new species is closely related to Dermaphis spp., therefore it was placed in the genus. The analysis incidentally indicated that “Dinipponaphis” autumna, a monoecious species forming galls on Distylium racemosum, was included in the clade of the genus Dermaphis, and therefore it was transferred to this genus.     

Interspecific gene flow in a multispecies oak hybrid zone in the Sierra Tarahumara of Mexico

Background and Aims

Interspecific gene flow can occur in many combinations among species within the genus Quercus, but simultaneous hybridization among more than two species has been rarely analysed. The present study addresses the genetic structure and morphological variation in a triple hybrid zone formed by Q. hypoleucoides, Q. scytophylla and Q. sideroxyla in north-western Mexico.

Methods

A total of 247 trees from ten reference and 13 presumed intermediate populations were characterized using leaf shape variation and geometric morphometrics, and seven nuclear microsatellites as genetic markers. Discriminant function analysis was performed for leaf shape variation, and estimates of genetic diversity and structure, and individual Bayesian genetic assignments were obtained.

Key Results

Reference populations formed three completely distinct groups according to discriminant function analysis based on the morphological data, and showed low, but significant, genetic differentiation. Populations from the zone of contact contained individuals morphologically intermediate between pairs of species in different combinations, or even among the three species. The Bayesian admixture analysis found that three main genetic clusters best fitted the data, with good correspondence of reference populations of each species to one of the genetic clusters, but various degrees of admixture evidenced in populations from the contact area.

Conclusions

The three oak species have formed a complex hybrid zone that is geographically structured as a mosaic, and comprising a wide range of genotypes, including hybrids between different species pairs, backcrosses and probable triple hybrids.     

Parasitoid complexes of lepidopteran leaf miners on oaks (Quercus dentata andQuercus mongolica) in Hokkaido,Japan

Parasitoid complexes of four microlepidopteran leaf-mining genera,Phyllonorycter, Caloptilia (Gracillariidae),Tischeria (Tischeriidae) andStigmella (Nepticulidae), which make mines on two oaks,Quercus dentata andQuercus mongolica, were studied at the Ishikari Coast, Hokkaido, northern Japan. In total, twenty-five parasitoid species were recognized.Phyllonorycter showed the highest parasitoid-species richness among the four leaf-mining genera. The number of parasitoid species associated with leaf miners onQ. dentata was nearly equal to that associated with miners onQ. mongolica, but a considerable difference was observed in the faunal make-up of parasitoids between leaf miners on the two oaks. The dominant parasitoid species differed betweenPhyllonorycter leucocorona andPhyllonorycter similis even on the same oak,Q. dentata. Factors influencing the host ranges of parasitoids and the parasitoid richness of host species or genera are discussed on the basis of the koinoparasitism/idioparasitism categorization.     

Suitability of native and ornamental oak species in California for Agrilus auroguttatus

Goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is a new invasive species in southern California, USA. The extent of the host range of this insect is not known, but this knowledge will have a major impact on assessment of the risks that this pest poses to oaks [Quercus spp. (Fagaceae)]. We conducted laboratory tests to determine the potential suitability of native and ornamental oak species for larvae and adults of A. auroguttatus. We infested 179 cut logs (from 163 different trees) with eggs or larvae, measured neonate survival and, after 5 months, counted feeding galleries, and noted the proportion of galleries with late instars. Initial larval survival was generally high when larvae penetrated the phloem (range: 62–98% among oak species), and low by the time larvae began to feed at the phloem/xylem interface (range: 0–25% among oak species). The majority of larvae that established a visible feeding gallery survived to the fourth instar (total of 73% for all oak species). Larval galleries were established with greater frequency in red oaks (Section Lobatae) compared with other oaks (19 vs. 7 or 4%). All red oaks tested (Q. agrifolia Née, Q. kelloggii Newberry, and Q. wislizeni A. DC.) were likely suitable hosts for larvae. Larvae were apparently able to feed on some of the other oaks (Q. chrysolepis Leibmann, Q. suber L., Q. lobata Née, and Q. douglasii Hook & Arn), although it remains unclear whether these species would be preferred hosts under natural conditions. Adult longevity and fecundity varied little by species of oak foliage fed to adults. The host range of A. auroguttatus is likely limited by suitability of oak species for the larval rather than the adult life stage. Results support published field observations that red oaks are more suitable hosts than white oaks.     

Mediterranean pine and oak distribution in southern Spain: Is there a mismatch between regeneration and adult distribution?

Questions: What is the current distribution of pine and oak species along environmental gradients in southern Spain? Do pine and oak regeneration niches differ from the environmental niches of adults? Is oak species regeneration favoured under the canopy of pine forests? Location: Forest areas of Andalusia (～87 600 km², southern Spain). Methods: We compiled extensive forest inventory data to explore differences in abundance (basal area, m² ha^?1) patterns of adults (dbh >7.4 cm) and regeneration (dbh ≤7.4 cm) of five pine and five oak species. Canonical correspondence analysis (CCA) and generalized linear models were applied to explore species–environment relationships along climatic, edaphic, topographic and fire‐frequency gradients. Results: Both pines and oaks segregated along complex environmental gradients, with pines generally dominating in more severe (colder and drier) environments, while oaks dominated in milder, wetter winter areas. In 40‐55% of mature pine stands there was a lack of regeneration in the understorey, while in two oak species (Q. suber and Q. canariensis) 70% of stands did not show regeneration. Pine recruits were found at a higher frequency and abundance under the canopy of their congeners, whereas some oaks (Q. ilex) had greater regeneration under mixed pine–oak canopies. Conclusions: Climatic limitations and soil properties partly explained the regional distribution of pines and oaks. We found evidence for an upward shift of Q. ilex recruits towards areas with colder conditions in pine forests, which could be explained by a possible facilitative effect of the pine canopy on seedling establishment.     

Geometric morphometric analyses of leaf variation in four oak species of the subgenus Quercus (Fagaceae)

Abstract

This paper defines a protocol for the study of leaf morphology and morphometric relationships among Quercus frainetto, Quercus robur, Quercus petraea and Quercus pubescens. Leaf shapes were investigated in an attempt to characterize and discriminate among these four species. Landmark and outline data were compared in order to ascertain their relative efficacy for describing variation in leaf morphology. MANOVA was performed to assess intra‐specific differentiation and to compare the distribution of variance in individuals, populations and species. A strong correlation was found between landmark and outline datasets of the four species: both methods proved capable of discriminating among the four species. Finally, a Two‐Block Partial Least‐Squares analysis was performed for each species in order to assess the correlation between intra‐specific shape variation and environmental factors.     

Pure species in a continuum of genetic and morphological variation: sympatric oaks at the edge of their range

Background and Aims Studies on oaks (Quercus spp.) have often been hampered by taxonomic confusion, a situation further compounded by the occurrence of extensive interspecific hybridization. In the present study, a combination of genetic and morphological analyses was used to examine sympatric populations of Q. petraea and Q. robur at the north-western edge of their ranges in Northern Ireland, since it had previously been suggested that hybridization could facilitate the apparent rapid, long-distance dispersal of oaks following the glaciations.Methods Samples were collected from 24 sites across Northern Ireland that had been previously designated as ancient or semi-natural woodland. Genotypes were obtained from a total of 950 trees using 12 nuclear microsatellite loci, and admixture coefficients were calculated based on a Bayesian clustering approach. Individuals were also classified as Q. petraea, Q. robur or hybrids based on two objective morphometric characters shown previously to delineate pure individuals effectively. Genetically ‘pure’ individuals of both species, as defined by the Bayesian clustering, were also genotyped for five chloroplast microsatellites.Key Results Genetic and morphological analyses both indicated the presence of pure individuals of both species, as well as a continuum of intermediates. There was a good agreement between the molecular and morphological classification, with a generally clear separation between pure individuals.Conclusions Despite millennia of hybridization and introgression, genetically and morphologically pure individuals of both Q. petraea and Q. robur can be found at the edge of their range, where both species occur sympatrically. The high proportion of individuals exhibiting introgression compared with previous studies may reflect the historical role of hybridization in facilitating dispersal following the glaciations. This is further supported by the significantly higher chloroplast diversity in Q. robur compared with Q. petraea.     

Spread and potential host range of the invasive oak lace bug [Corythucha arcuata (Say, 1832) – Heteroptera: Tingidae] in Eurasia

1. The North American oak lace bug feeds on leaves of ‘white oaks” in its native range. In Europe, it was first discovered in northern Italy in 2000. In recent years, it has subsequently spread rapidly and population outbreaks have been observed in several European countries. In the present study, we summarize the steps of its expansion.
2. To predict its potential host range, we checked 48 oak species in 20 sentinel gardens in seven countries between 2013 and 2018.
3. In total, 27 oak species were recorded as suitable hosts; 13 of them are globally new ones, 23 out of the 29 in section Quercus (～ white oaks, an intrageneric taxonomic unit within genus Quercus), including Asian oaks, native to Japan, Korea and China, and four out of five in section Cerris (another intrageneric unit of the same genus), were accepted as hosts. None of the species in section Lobatae (red oaks) or in the Ilex group was accepted.
4. Host records were also collected in forest stands of 10 countries. We found 11 oak species that were infested. Outbreak populations were most commonly found on Quercus robur, Quercus frainetto, Quercus petraea and Quercus cerris, comprising widespread and outstandingly important oaks species in Europe.
5. Based on our findings, we conclude that suitable hosts for oak lace bug are present in most of Europe and Asia. This means that a lack of hosts will likely not restrict further range expansion.

     

Carbon and water relations of juvenile Quercus species in tall‐grass prairie

Abstract. In ecosystems where environments are extreme, such as deserts, adult plant species may facilitate the establishment and growth of seedlings and juveniles. Because high temperatures and evaporative demand characterize tall‐grass prairies of the central United States (relative to forests), we predicted that the grassland‐forest ecotone, by minimizing temperature extremes and moderating water stress, may function to facilitate the expansion of Quercus species into undisturbed tall‐grass prairie. We assessed the carbon and water relations of juvenile Quercus macrocarpa and Q. muhlenbergii, the dominant tree species in gallery forests of northeast Kansas, in ecotone and prairie sites. To evaluate the potentially competitive effects of neighboring herbaceous biomass on these oaks, juveniles (< 0.5 m tall) of both species also were subjected to either: (1) removal of surrounding above‐ground herbaceous biomass, or (2) control (prairie community intact) treatments. Herbaceous biomass removal had no significant effect on gas exchange or water relations in these oak species in either the prairie or the ecotone environment. Although the ecotone did alleviate some environmental extremes, photosynthetic rates and stomatal conductance were ca. 20 % higher (p < 0.05) in both oaks in prairie sites vs. the ecotone. Moreover, although leaf temperatures on average were higher in oaks in the prairie, high leaf temperatures in the ecotone had a greater negative effect on photosynthesis. These data suggest that the grassland‐forest ecotone did not facilitate the growth of Quercus juveniles expanding into this grassland. Moreover, the carbon and water relations of juvenile oaks in the prairie appeared to be unaffected by the presence of the dominant C₄ grasses.     

Hybridisation and colonisation dynamics in European oaks

Summary

Extensive hybridisation between the two sympatric species Quercus petraea and Q.robur is suggested by the near lack of genetic differentiation between the two species and supported by controlled crosses and mating system analysis in mixed stands. Further ecological and genetic evidence suggest that hybridisation does not impede the ecological specialisation of the two species, raising the issue of its evolutionary significance in oaks. Preferential unidirectional hybridization (pollen Q. petraea to ovule Q. robur) has been shown in various mixed stands and facilitates the introduction of sessile oak in existing pedunculate stands. If this unidirectional trend is reinforced in later backcrosses, then hybridisation leads to the dispersal of Q. petraea in existing stands of Q. robur. Hybridisation can therefore be seen as a ‘pollen-mediated’ dispersal mechanism, and has most likely contributed to the rapid migration of Q. petraea in Europe. Given the extant distribution of the species in Europe, migration through pollen swamping should be seen at the edges of the natural distribution of Q. petraea where the demographic imbalance of the two species will reinforce backcrosses.