Fitness dynamics within a poplar hybrid zone: I. Prezygotic and postzygotic barriers impacting a native poplar hybrid stand

Hybridization and introgression are pervasive evolutionary phenomena that provide insight into the selective forces that maintain species boundaries, permit gene flow, and control the direction of evolutionary change. Poplar trees (Populus L.) are well known for their ability to form viable hybrids and maintain their distinct species boundaries despite this interspecific gene flow. We sought to quantify the hybridization dynamics and postzygotic fitness within a hybrid stand of balsam poplar (Populus balsamifera L.), eastern cottonwood (P. deltoides Marsh.), and their natural hybrids to gain insight into the barriers maintaining this stable hybrid zone. We observed asymmetrical hybrid formation with P. deltoides acting as the seed parent, but with subsequent introgression biased toward P. balsamifera. Native hybrids expressed fitness traits intermediate to the parental species and were not universally unfit. That said, native hybrid seedlings were absent from the seedling population, which may indicate additional selective pressures controlling their recruitment. It is imperative that we understand the selective forces maintaining this native hybrid zone in order to quantify the impact of exotic poplar hybrids on this native system.     

Plant genetic identity of foundation tree species and their hybrids affects a litter-dwelling generalist predator

The effects of plant genetics on predators, especially those not living on the plant itself, are rarely studied and poorly understood. Therefore, we investigated the effect of plant hybridization and genotype on litter-dwelling spiders. Using an 18-year-old cottonwood common garden, we recorded agelenid sheet-web density associated with the litter layers of replicated genotypes of three tree cross types: Populus fremontii, Populus angustifolia, and their F₁ hybrids. We surveyed 118 trees for agelenid litter webs at two distances from the trees (0–100 and 100–200 cm from trunk) and measured litter depth as a potential mechanism of web density patterns. Five major results emerged: web density within a 1-m radius of P. angustifolia was approximately three times higher than within a 1-m radius of P. fremontii, with F₁ hybrids having intermediate densities; web density responded to P. angustifolia and F₁ hybrid genotypes as indicated by a significant genotype × distance interaction, with some genotypes exhibiting a strong decline in web density with distance, while others did not; P. angustifolia litter layers were deeper than those of P. fremontii at both distance classes, and litter depth among P. angustifolia genotypes differed up to 300 %; cross type and genotype influenced web density via their effects on litter depth, and these effects were influenced by distance; web density was more sensitive to the effects of tree cross type than genotype. By influencing generalist predators, plant hybridization and genotype may indirectly impact trophic interactions such as intraguild predation, possibly affecting trophic cascades and ecosystem processes.     

A morphological and chemical study of Populus acuminata Rydberg

Evidence from morphology, flavonoid chemistry, and field observations suggests thatPopulus acuminata is of hybrid origin. The putative parents areP. angustifolia, the narrow leaf cottonwood, and deltoid leaved plants that are assigned toP. sargentii (P. deltoides var.occidentalis), P. fremontii, orP. wislizenii (P. fremontii var.wislizenii). Populus angustifolia exhibits a series of flavonol glycosides (kaempferol, quercetin, and myricetin) in its leaves. By contrast, the major leaf flavonoids of the broad leaved plants are flavone glycosides (apigenin and luteolin).Populus acuminata is intermediate between the suspected parents in morphological features. Additionally, the leaves of mostP. acuminata plants contain the exact summation of the flavonoid compounds characteristic of the putative parents. A diploid chromosome number of 2n = 38 was obtained for six plants, which confirms the one previous report for the species. Meiosis was regular in all cases. Correlated data indicate that the majority of plants ofP. acuminata represent F₁ hybrids and that complex hybridization is not common. Evidence from morphological and chemical studies is presented to show that in certain instances backcrossing to both parents has occurred. Results gathered in this study show thatP. ×andrewsii is undoubtedly “typical”P. acuminata, but the type specimen is from a sucker shoot, and thus has been interpreted as a backcross toP. sargentii. Populus acuminata var.rehderi is not considered worthy of taxonomic recognition.     

Geographical barriers and climate influence demographic history in narrowleaf cottonwoods

Studies of genetic variation can clarify the role of geography and spatio-temporal variation of climate in shaping demography, particularly in temperate zone tree species with large latitudinal ranges. Here, we examined genetic variation in narrowleaf cottonwood, Populus angustifolia, a dominant riparian tree. Using multi-locus surveys of polymorphism in 363 individuals across the species'' 1800 km latitudinal range, we found that, first, P. angustifolia has stronger neutral genetic structure than many forest trees (simple sequence repeat (SSR) F_ST=0.21), with major genetic groups corresponding to large apparent geographical barriers to gene flow. Second, using SSRs and putatively neutral sequenced loci, coalescent simulations indicated that populations diverged before the last glacial maximum (LGM), suggesting the presence of population structure before the LGM. Third, the LGM and subsequent warming appear to have had different influences on each of these distinct populations, with effective population size reduction in the southern extent of the range but major expansion in the north. These results are consistent with the hypothesis that climate and geographic barriers have jointly affected the demographic history of P. angustifolia, and point the importance of both factors as being instrumental in shaping genetic variation and structure in widespread forest trees.     

Asymmetric introgression between Magnolia stellata and M. salicifolia at a site where the two species grow sympatrically

In order to understand the ongoing evolutionary relationships between species, it is important to elucidate patterns of natural hybridization. In the zone where two species are sympatrically distributed, we examined 274 individuals of Magnolia stellata, Magnolia salicifolia, and their putative hybrids by means of 16 nuclear and three chloroplast microsatellite markers. Hybrid classes of individuals were estimated by admixture analyses. Morphological traits were also investigated for 64 of the 274 individuals. Admixture analyses revealed that 66 of the 274 individuals were classified as hybrids, comprising 17 F₁ and 19 F₂ individuals, 27 backcrosses to M. salicifolia, and 3 individuals of unknown origin. Morphological data from the 64 individuals agreed well with their genetic admixture rates. Spatial locations of F₁ and F₂ hybrids at the study site were intermediate between the two purebred species, indicating that the site preferences of hybrids are intermediate. The occurrences of F₂ and backcross hybrids indicate that F₁ hybrids are fertile. The chloroplast DNA haplotypes of all F₁ hybrids corresponded to those detected in M. salicifolia, so that maternal parents of the F₁ hybrids were all M. salicifolia. Furthermore, no hybrid individuals derived from a backcross to M. stellata were detected. These results suggest that the direction of hybridization and the subsequent introgression have been quite asymmetric and that the introgression occurred from M. stellata into M. salicifolia.     

Arthropod communities on hybrid and parental cottonwoods are phylogenetically structured by tree type: Implications for conservation of biodiversity in plant hybrid zones

Although hybridization in plants has been recognized as an important pathway in plant speciation, it may also affect the ecology and evolution of associated communities. Cottonwood species (Populus angustifolia and P. fremontii) and their naturally occurring hybrids are known to support different plant, animal, and microbial communities, but no studies have examined community structure within the context of phylogenetic history. Using a community composed of 199 arthropod species, we tested for differences in arthropod phylogenetic patterns within and among hybrid and parental tree types in a common garden. Three major patterns emerged. (1) Phylogenetic diversity (PD) was significantly different between arthropod communities on hybrids and Fremont cottonwood when pooled by tree type. (2) Mean phylogenetic distance (MPD) and net relatedness index (NRI) indicated that communities on hybrid trees were significantly more phylogenetically overdispersed than communities on either parental tree type. (3) Community distance (D_pw) indicated that communities on hybrids were significantly different than parental species. Our results show that arthropod communities on parental and hybrid cottonwoods exhibit significantly different patterns of phylogenetic structure. This suggests that arthropod community assembly is driven, in part, by plant–arthropod interactions at the level of cottonwood tree type. We discuss potential hypotheses to explain the effect of plant genetic dissimilarity on arthropod phylogenetic community structure, including the role of competition and environmental filtering. Our findings suggest that cottonwood species and their hybrids function as evolutionarily significant units (ESUs) that affect the assembly and composition of associated arthropod communities and deserve high priority for conservation.     

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.     

Repeated unidirectional introgression towards Populus balsamifera in contact zones of exotic and native poplars

As the evolutionary significance of hybridization is largely dictated by its extent beyond the first generation, we broadly surveyed patterns of introgression across a sympatric zone of two native poplars ( Populus balsamifera , Populus deltoides ) in Quebec, Canada within which European exotic Populus nigra and its hybrids have been extensively planted since the 1800s. Single nucleotide polymorphisms (SNPs) that appeared fixed within each species were characterized by DNA-sequencing pools of pure individuals. Thirty-five of these diagnostic SNPs were employed in a high-throughput assay that genotyped 635 trees of different age classes, sampled from 15 sites with various degrees of anthropogenic disturbance. The degree of admixture within sampled trees was then assessed through Bayesian clustering of genotypes. Hybrids were present in seven of the populations, with 2.4% of all sampled trees showing spontaneous admixture. Sites with hybrids were significantly more disturbed than pure stands, while hybrids comprised both immature juveniles and trees of reproductive age. All three possible F1s were detected. Advanced-generation hybrids were consistently biased towards P. balsamifera regardless of whether hybridization had occurred with P. deltoides or P. nigra. Gene exchange between P. deltoides and P. nigra was not detected beyond the F1 generation; however, detection of a trihybrid demonstrates that even this apparent reproductive isolation does not necessarily result in an evolutionary dead end. Collectively, results demonstrate the natural fertility of hybrid poplars and suggest that introduced genes could potentially affect the genetic integrity of native trees, similar to that arising from introgression between natives.     

Geographically extensive hybridization between the forest trees American butternut and Japanese walnut

We investigate the question of naturally occurring interspecific hybrids between two forest trees: the native North American butternut (Juglans cinerea L.) and the introduced Japanese walnut (Juglans ailantifolia Carrière). Using nuclear and chloroplast DNA markers, we provide evidence for 29 F₁ and 22 advanced generation hybrids in seven locations across the eastern and southern range of the native species. Two locations show extensive admixture (95% J. ailantifolia and hybrids) while other locations show limited admixture. Hybridization appears to be asymmetrical with 90.9 per cent of hybrids having J. ailantifolia as the maternal parent. This is, to our knowledge, the first genetic data supporting natural hybridization between these species. The long-term outcome of introgression could include loss of native diversity, but could also include transfer of useful traits from the introduced species.     

Testing the ‘Hybrid Susceptibility’ and ‘Phenological Sink’ Hypotheses Using the P. balsamifera – P. deltoides Hybrid Zone and Septoria Leaf Spot [Septoria musiva]

Hybrid genotypes that arise between plant species frequently have increased susceptibility to arthropod pests and fungal pathogens. This pattern has been attributed to the breakdown of plant defenses (‘Hybrid susceptibility’ hypothesis) and (or) to extended periods of susceptibility attributed to plant phenologies in zones of species overlap and (or) hybridization (‘phenological sink’ hypothesis). We examined these hypotheses by assessing the susceptibility of parental and hybrid Populus host genotypes to a leaf spot disease caused by the fungal pathogen Septoria musiva. For this purpose, 214 genotypes were obtained from morphologically pure zones of P. balsamifera and P. deltoides, and from an intervening zone of overlap and hybridization on the drainage of the Red Deer River, Alberta, Canada. Genotypes were identified as P. balsamifera, P. deltoides, or hybrid using a suite of 27 species-specific SNP markers. Initially the genetic structure of the hybrid zone was characterized with 27.7% of trees classified as admixed individuals. To test the hybrid susceptibility hypothesis, a subset of 52 genotypes was inoculated with four isolates of S. musiva. Levels of susceptibility were P. balsamifera > F₁ hybrid > P. deltoides. A further 53 genotypes were grown in a common garden to assess the effect of genotype on variation in leaf phenology. Leaf phenology was more variable within the category of hybrid genotypes than within categories of either parental species. Leaf phenology was also more variable for the category of trees originating in the hybrid (P. balsamifera – P. deltoides [hybrid and parental genotypes combined]) zone than in adjacent pure zones of the parental species. The results from the inoculation experiment support the hybrid intermediacy hypothesis. The results from the common garden experiment support the ‘phenological sink’ hypothesis. These findings have greatly increased our understanding of the epidemiology and ecology of fungal pathogens in plant hybrid zones.     

Molecular genetics of growth and development in Populus. I. Triploidy in hybrid poplars

Summary While constructing a genetic linkage map of a hybrid poplar genome (Populus trichocarpa x P. deltoides), we identified several restriction fragment length polymorphismus (RFLPs) for which the parental trees are heterozygous. Although 8 of the 11 F₁ hybrid offspring inherited, as expected, single RFLP alleles from each parent, 3 F₁ trees in the mapping pedigree inherited both maternal alleles along with a single paternal allele at some loci. Aneuploidy or polyploidy in these 3 F₁ trees due to partial or complete nondisj unction during female gametogenesis is the simplest explanation for this finding. Of the 3 f₁ offspring with supernumerary RFLP alleles 2 have triploid nuclear DNA contents as measured by fluorescence flow cytometry; the 3rd F₁ with supernumerary alleles has a sub-triploid nuclear DNA content and is probably aneuploid. Among the tri/aneuploid hybrids, leaf quantitative traits either are skewed toward those values characteristic of the P. trichocarpa female parent (adaxial stomate density, petiole length: blade length ratio; abaxial color) or show transgressive variation (epidermal cell size). Abaxial leaf color was used to screen a large population of P. trichocarpa x P. deltoides hybrids for further evidence of tri/aneuploidy. In each case where a white abaxial leaf surface was observed and the nuclear DNA content measured, the hybrid proved to be tri/aneuploid. All sexually mature female triploids examined were sterile, although the inflorescences completed their development in the absence of embryo formation. The (probably) aneuploid F₁ hybrid is a fertile female. Of 15 female P. trichocarpa parents used in crosses to P. deltoides, 10 produced one or more tri/aneuploid hybrid offspring. In an intraspecific cross using a P. trichocarpa female that had produced triploid hybrids with five different P. deltoides males, no tri/aneuploid offpsring were found.     

Relative importance of genetic, ontogenetic, induction, and seasonal variation in producing a multivariate defense phenotype in a foundation tree species

Plant adaptations for defense against herbivory vary both among species and among genotypes. Moreover, numerous forms of within-plant variation in defense, including ontogeny, induction, and seasonal gradients, allow plants to avoid expending resources on defense when herbivores are absent. We used an 18-year-old cottonwood common garden composed of Populus fremontii, Populus angustifolia, and their naturally occurring F₁ hybrids (collectively referred to as ??cross types??) to quantify and compare the relative influences of three hierarchical levels of variation (between cross types, among genotypes, and within individual genotypes) on univariate and multivariate phytochemical defense traits. Within genotypes, we evaluated ontogeny, induction (following cottonwood leaf beetle herbivory), and seasonal variation. We compared the effect sizes of each of these sources of variation on the plant defense phenotype. Three major patterns emerged. First, we observed significant differences in concentrations of defense phytochemicals among cross types, and/or among genotypes within cross types. Second, we found significant genetic variation for within-plant differences in phytochemical defenses: (a) based on ontogeny, levels of constitutive phenolic glycosides were nearly three times greater in the mature zone than in the juvenile zone within one cottonwood cross type, but did not significantly differ within another cross type; (b) induced levels of condensed tannins increased up to 65?% following herbivore damage within one cottonwood cross type, but were not significantly altered in another cross type; and (c) concentrations of condensed tannins tended to increase across the season, but did not do so across all cross types. Third, our estimates of effect size demonstrate that the magnitude of within-plant variation in a phytochemical defense can rival the magnitude of differences in defense among genotypes and/or cross types. We conclude that, in cottonwood and likely other plant species, multiple forms of within-individual variation have the potential to substantially influence ecological and evolutionary processes.     

Effects of flooding on leaf development,transpiration, and photosynthesis in narrowleaf cottonwood,a willow-like poplar

The narrowleaf cottonwood, Populus angustifolia, occurs in occasionally flooded, low elevation zones along river valleys near the North American Rocky Mountains. This small poplar has narrow leaves and fine branching and thus resembles willows, which are commonly flood-tolerant. We investigated the flood response of narrowleaf cottonwoods and a related native hybrid, jackii cottonwood (P. × jackii = P. balsamifera × P. deltoides), by studying saplings of 24 clones in a greenhouse, with some pots being inundated to provide the flood treatment. Flooding slightly reduced leaf numbers (−10%), and leaf sizes were reduced by about 21% in female P. angustifolia versus a 50% reduction in the female hybrids. Flooding-reduced stomatal conductance and net photosynthetic rate, and reduced transpiration particularly in P. × jackii. The effects on foliar gas exchange declined over a 5-week interval, suggesting compensation. The moderate impact of flooding supports the hypothesis that narrowleaf cottonwoods are flood-tolerant, and we anticipate that these trees could provide traits to increase the flood tolerance of fast-growing hybrid poplars. The results further indicate that female cottonwoods may be more flood-tolerant than males, and females could be more successful in lower, flood-prone sites.     

Detecting hybridization in mixed populations ofRhinanthus minor andRhinanthus angustifolius

Rhinanthus minor andRhinanthus angustifolius are known to hybridize in mixed populations in nature. These hybridization events can have important evolutionary consequences. The development and use of species-specific RAPD and ISSR markers allowed the detection of hybrid individuals not always distinguishable with morphological characters. Two mixed populations of different ages were studied. In a young mixed 2-year-old population, both individuals of the two parental species and F₁ hybrids were found using genetic analysis, showing that hybridization occurred rapidly. Flower morphology of F₁ hybrids was too variable to distinguish all these hybrids from the parental species. This morphological variability of F₁ hybrids was also confirmed in artificial crosses in the greenhouse. In an old and no longer mixed 30-year-old population, onlyR. angustifolius plants and a few genetically introgressed individuals close toR. angustifolius were present. Genetic markers showed traces of past hybridization and introgression. Unidirectional introgression ofR. minor intoR. angustifolius with the complete disappearance ofR. minor from this population was observed.     

Maintenance of strong morphological differentiation despite ongoing natural hybridization between sympatric species of Lomatia (Proteaceae)

Background and Aims

When species cohesion is maintained despite ongoing natural hybridization, many questions are raised about the evolutionary processes operating in the species complex. This study examined the extensive natural hybridization between the Australian native shrubs Lomatia myricoides and L. silaifolia (Proteaceae). These species exhibit striking differences in morphology and ecological preferences, exceeding those found in most studies of hybridization to date.

Methods

Nuclear microsatellite markers (nSSRs), genotyping methods and morphometric analyses were used to uncover patterns of hybridization and the role of gene flow in morphological differentiation between sympatric species.

Key Results

The complexity of hybridization patterns differed markedly between sites, however, signals of introgression were present at all sites. One site provided evidence of a large hybrid swarm and the likely presence of multiple hybrid generations and backcrosses, another site a handful of early generational hybrids and a third site only traces of admixture from a past hybridization event. The presence of cryptic hybrids and a pattern of morphological bimodality amongst hybrids often disguised the extent of underlying genetic admixture.

Conclusions

Distinct parental habitats and phenotypes are expected to form barriers that contribute to the rapid reversion of hybrid populations to their parental character state, due to limited opportunities for hybrid/intermediate advantage. Furthermore, strong genomic filters may facilitate continued gene flow between species without the danger of assimilation. Stochastic fire events facilitate temporal phenological isolation between species and may partly explain the bi-directional and site-specific patterns of hybridization observed. Furthermore, the findings suggest that F₁ hybrids are rare, and backcrosses may occur rapidly following these initial hybridization events.     

Hybridization and introgression between the exotic Siberian elm, Ulmus pumila, and the native Field elm, U. minor, in Italy

In response to the first Dutch elm disease (DED) pandemic, Siberian elm, Ulmus pumila, was planted to replace the native elm, U. minor, in Italy. The potential for hybridization between these two species is high and repeated hybridization could result in the genetic swamping of the native species and facilitate the evolution of invasiveness in the introduced species. We used genetic markers to examine the extent of hybridization between these two species and to determine the pattern of introgression. We quantified and compared the level of genetic diversity between the hybrids and the two parental species. Hybrids between U. pumila and U. minor were common. The pattern of introgression was not as strongly biased towards U. pumila as was previously observed for hybrids between U. rubra and U. pumila in the United States. The levels of heterozygosity were similar between U. minor and the hybrids and both groups had higher levels of heterozygosity relative to U. pumila. The programs Structure and NewHybrids indicated the presence of first- (F₁) and second- generation (F₂) hybrids and of backcrosses in the hybrid population. The presence of healthy DED resistant U. minor individuals combined with the self-compatibility of U. minor could help explain the presence of F₂ individuals in Italy. The presence of F₂ individuals, where most of the variability present in the hybrids will be released, could facilitate rapid evolution and the potential evolution of invasiveness of U. pumila in Italy.     

Hybrid populations selectively filter gene introgression between species

Hybrids have long been recognized as a potential pathway for gene flow between species that can have important consequences for evolution and conservation biology. However, few studies have demonstrated that genes from one species can introgress or invade another species over a broad geographic area. Using 35 genetically mapped restriction fragment length polymorphism (RFLP) markers of two species of cottonwoods (Populus fremontii x P. angustifolia) and their hybrids (n = 550 trees), we showed that the majority of the genome is prohibited from introgressing from one species into the other. However, this barrier was not absolute; Fremont cpDNA and mtDNA were found throughout the geographic range of narrowleaf cottonwood, and 20% of the nuclear markers of Fremont cottonwood introgressed varying distances (some over 100 km) into the recipient species' range. Rates of nuclear introgression were variable, but two nuclear markers introgressed as fast as the haploid, cytoplasmically inherited chloroplast and mitochondrial markers. Our genome-wide analysis provides evidence for positive, negative, and neutral effects of introgression. For example, we predict that DNA fragments that introgress through several generations of backcrossing will be small, because small fragments are less likely to contain deleterious genes. These results argue that recombination will be important, that introgression can be very selective, and that evolutionary forces within the hybrid population to effectively "filter" gene flow between species. A strong filter may make introgression adaptive, prevent genetic assimilation, lead to relaxed isolating mechanisms, and contribute to the stability of hybrid zones. Thus, rather than hybridization being a negative factor as is commonly argued, natural hybridization between native species may provide important genetic variation that impacts both ecological and evolutionary processes. Finally, we propose two hypotheses that contrast the likelihood of contemporary versus ancient introgression in this system.     

Natural and Anthropogenic Hybridization in Two Species of Eastern Brazilian Marmosets (Callithrix jacchus and C. penicillata)

Animal hybridization is well documented, but evolutionary outcomes and conservation priorities often differ for natural and anthropogenic hybrids. Among primates, an order with many endangered species, the two contexts can be hard to disentangle from one another, which carries important conservation implications. Callithrix marmosets give us a unique glimpse of genetic hybridization effects under distinct natural and human-induced contexts. Here, we use a 44 autosomal microsatellite marker panel to examine genome-wide admixture levels and introgression at a natural C. jacchus and C. penicillata species border along the São Francisco River in NE Brazil and in an area of Rio de Janeiro state where humans introduced these species exotically. Additionally, we describe for the first time autosomal genetic diversity in wild C. penicillata and expand previous C. jacchus genetic data. We characterize admixture within the natural zone as bimodal where hybrid ancestry is biased toward one parental species or the other. We also show evidence that São Francisco River islands are gateways for bidirectional gene flow across the species border. In the anthropogenic zone, marmosets essentially form a hybrid swarm with intermediate levels of admixture, likely from the absence of strong physical barriers to interspecific breeding. Our data show that while hybridization can occur naturally, the presence of physical, even if leaky, barriers to hybridization is important for maintaining species genetic integrity. Thus, we suggest further study of hybridization under different contexts to set well informed conservation guidelines for hybrid populations that often fit somewhere between “natural” and “man-made.”     

Detection of hybridization between two loach species (<Emphasis Type="Italic">Paramisgurnus dabryanus</Emphasis> and <Emphasis Type="Italic">Misgurnus anguillicaudatus</Emphasis>) in wild populations

Artificial interspecific hybrids between large scale loach P. dabryanus and tetraploid pond loach M. anguillicaudatus (Cobitidae, Cypriniformes) are viable. To detect the occurrence of possible natural hybridization, genetic analyses by using microsatellite markers were performed for natural populations of large scale loach and pond loach, the reciprocal laboratory hybrids, and “supposed hybrids” with ambiguous morphology. The fertility of the artificial hybrids was also tested. At one diagnostic microsatellite (Mac50), one out of 20 “supposed hybrids” was identified to be F₁ hybrid between the two loach species because it had the same genotype as that of the laboratory hybrids. The triploid hybrids between the two species were confirmed to be female-sterile. The results show that rare hybridization has occurred between diploid large scale loach and tetraploid pond loach in nature although it may have little effect in genetic introgression. This study is helpful for fish conservation and encourages further investigation on natural hybridization and introgression of loaches.     

Genetic Analysis of an Interspecific Hybrid Swarm of Populus: Occurrence of Unidirectional Introgression

Restriction fragment length polymorphisms were used to distinguish genotypes of two species of Populus, P. fremontii ('Fremont') and P. angustifolia ('narrowleaf'). Both inter- and intraspecific polymorphisms were detected in these cottonwood trees. The interspecific variation was much greater than the intraspecific variation. This permitted identification of parental genotypes within individual trees of a hybrid swarm which exists in an overlap zone between the two species. Within this hybrid swarm, individual trees are either F1 hybrids or backcrosses with a pure 'narrowleaf' parent; no progeny were found that could be attributed to crossing between F1 hybrid trees, or to backcrossing between F1 hybrid trees and 'Fremont'.