Moving Hybrid Zone Research Articles

Moving Hybrid Zones; When Two Species Meet, Mate, and Compete

When parents of two different species have babies together, those babies are called hybrids. In nature, hybrids are often born in the region where the ranges of their parent species meet. This region is called a hybrid zone. We know that species change their ranges all the time, and we also know that some species compete with each other for food or living space. This means that, if one of the two parent species manages to expand its range, the other species may be forced to retreat. If that were to happen, the hybrid zone between the two species’ ranges should move, right? Even though researchers used to think that hybrid zone movement was rare, recent studies suggest otherwise. In this article, we will tell you what hybrid zones are, how they form, why their position may shift over time, and what we can learn from this movement.

Hybridization in late stages of speciation: Strong but incomplete genome-wide reproductive isolation and 'large Z-effect' in a moving hybrid zone.

In organisms reproducing sexually, speciation occurs when increasing divergence results in pre- or post-zygotic reproductive isolation between lineages. Studies focusing on reproductive isolation origin in early stages of speciation are common and many rely on genomic scans to infer introgression providing limited information on the genomic architecture of reproductive isolation long-term maintenance. This study analyses a natural hybrid zone between two species in a late stage of speciation. We used ddRADseq genotyping in the contact between Podarcis bocagei and P. carbonelli to examine admixture extent, analyse hybrid zone stability and assess genome-wide variation in selection against introgression. We confirmed strong but incomplete reproductive isolation in a bimodal hybrid zone. New findings revealed population genetic structure within P. carbonelli in the contact zone; geographical and genomic clines analysis suggested strong selection against gene flow, but a relatively small proportion of the loci can introgress, mostly within the narrow contact zone. However, geographical clines revealed that a few introgressed loci show signs of potential positive selection, particularly into P. bocagei. Geographical clines also detected a signal of hybrid zone movement towards P. bocagei distribution. Genomic cline analysis revealed heterogeneous patterns of introgression among loci within the syntopy zone, but the majority maintain a strong association with the genomic background of origin. However, incongruences between both cline approaches were found, potentially driven by confounding effects on genomic clines. Last, an important role of the Z chromosome in reproductive isolation is suggested. Importantly, overall patterns of restricted introgression seem to result from numerous strong intrinsic barriers across the genome.

Geographic variability of hybridization between red-breasted and red-naped sapsuckers.

Hybrid zones reveal the strength of reproductive isolation between populations undergoing speciation and are a key tool in evolutionary biology research. Multiple replicate transects across the same hybrid zone offer insight into the dynamics of hybridization in different environments, clarifying the role of extrinsic forces on the speciation process. Red-breasted and red-naped sapsuckers (Sphyrapicus ruber and Sphyrapicus nuchalis) have a long zone of contact over approximately 1,600 km from central British Columbia, Canada to central California, USA. We used Genotyping-by-Sequencing data from three independent sapsucker hybrid zone transects to compare hybridization dynamics between these species under variable geoclimatic conditions. We generated geographic clines of the genomic data to compare hybrid zone widths and used random forests models and linear regression to assess the relationship between climate and sapsucker ancestry along each transect. Our results show variation in the directionality of backcrossing, often indicative of moving hybrid zones. We note variable cline widths among transects, indicating differences in selection maintaining hybrid zone dynamics. Furthermore, random forests models identified different variables in close association with sapsucker ancestry across each transect. These results indicate a lack of repeatability across replicate transects and a strong influence of the local environment on hybrid zone dynamics.

Genetic traces of hybrid zone movement across a fragmented habitat.

Theoretical and empirical studies suggest that the structure and position of hybrid zones can change over time. Evidence for moving hybrid zones has been directly inferred by repeated sampling over time, or indirectly through the detection of genetic footprints left by the receding species and the resulting asymmetric patterns of introgression across markers. We here investigate a hybrid zone formed by two subspecies of the Iberian golden-striped salamander, Chioglossa lusitanica, using a panel of 35 nuclear loci (31 SNPs and 4 allozymes) and one mitochondrial locus in a transect in central Portugal. We found concordant and coincident clines for most of the nuclear loci (n=22, 63%), defining a narrow hybrid zone of ca. 6km wide, with the centre positioned ca. 15km south of the Mondego River. Asymmetric introgression was observed at another 14loci. Their clines are displaced towards the north, with positions located either close to the Mondego River (n=6) or further northwards (n=8). We interpret these profiles as genetic traces of the southward displacement of C. lusitanica lusitanica by C. l. longipes over the wider Mondego River valley. We noted the absence of significant linkage disequilibrium, and we inferred low levels of effective selection per locus against hybrids, suggesting that introgression in the area of species replacement occurred under a neutral diffusion process. A species distribution model suggests that the C. lusitanica hybrid zone coincides with a narrow corridor of fragmented habitat. From the position of the displaced clines, we infer that patches of locally suitable habitat trapped some genetic variants that became disassociated from the southward moving hybrid zone. This study highlights the influence of habitat availability on hybrid zone movement.

Haemosporidian prevalence and community composition vary little across a chickadee hybrid zone

Abstract Within animal hybrid zones, parasites may determine competitive outcomes between host species and thus affect hybridization dynamics. We addressed this hypothesis by evaluating haemosporidian prevalence and community composition in a rapidly moving hybrid zone between Black-capped Chickadees (Poecile atricapillus) and Carolina Chickadees (P. carolinensis). Using molecular methods, we screened for haemosporidians in multiple chickadee populations across the hybrid zone and investigated whether parasite prevalence varied as a function of admixture among these birds. We identified 36 parasite lineages from 3 haemosporidian genera (Plasmodium, Haemoproteus, and Leucocytozoon) but found no genera or lineages more likely to infect any particular chickadee taxon. Instead, haemosporidian prevalence varied across sites and seasons: Leucocytozoon was more prevalent during chickadees’ breeding season, whereas Haemoproteus prevalence peaked during nonbreeding periods. Leucocytozoon infected proportionally fewer birds at the leading edge of the hybrid zone than near its center. However, haemosporidian communities were similar among chickadee populations, and evidence for parasite exchanges between chickadee taxa was lacking. These results underscore the complexity of bird–parasite relationships and suggest that haemosporidians are unlikely to play a major role in the ongoing movement of this hybrid zone.

A genomic footprint of a moving hybrid zone in marbled newts

We developed a panel of 44 nuclear genetic markers and applied this to two species of marbled newts in the north (Triturus marmoratus) and the south (T. pygmaeus) of the Iberian Peninsula, to document pattern and process of interspecific gene flow. The northernmost occurrence of T. pygmaeus genetic material was in a T. marmoratus population north of the Vouga river estuary. This suggested the past presence of a hybrid zone, possibly coinciding with a natural river outlet at ca. 1200 A.D. Since 1808, the species contact has moved back south to a by then completed, man-made Vouga channel. We also found evidence for a T. marmoratus genomic footprint in T. pygmaeus from the Serra de Sintra, near Lisbon. In combination with a previously reported southern, relic occurrence of T. marmoratus in between both areas, the data point to the superseding with hybridization of T. marmoratus by T. pygmaeus. We estimate that the species hybrid zone has moved over a distance of ca. 215 km.

The Amazon river is a suture zone for a polyphyletic group of co‐mimetic heliconiine butterflies

The Amazon basin contains few obvious geographic barriers, yet it is the most biodiverse region on Earth. One hypothesis to explain its diversity is that the very large rivers promote allopatric divergence. Consistent with this, maps of heliconiine butterflies made from museum specimens show high subspecies richness close to the Amazon river, suggesting that it may produce or maintain intra‐specific phenotypic variability. However, museum data are subject to strong spatial biases (the ‘Wallacean shortfall' of distribution data), raising the possibility that this pattern is a sampling artefact. To test this, we systematically collected along a ~900 km north–south transect running through central Amazonia. We found a significant association between phenotypic diversity and major rivers, with distance from the Amazon river explaining 61% of the variance in the mean polymorphism of 25 species. This association is partly because many species exhibit different phenotypes on either side of the river. Nonetheless, we also find sites with high polymorphism close to the river, indicating continual cross‐river dispersal. Our results strongly suggest the presence of a suture zone (a region where multiple species have hybrid zones) near the city of Manaus. However, the effect of the river on spatial patterns of intra‐specific phenotypic diversity depends on a species' mimetic phenotype. Rather than being absolute barriers, our results support the idea that rivers can act as partial barriers that trap moving hybrid zones, resulting in a suture zone. As such, the wide Amazonian rivers help generate and maintain colour pattern diversity, but to date there is no evidence that they lead to speciation in our study group.

Genomic regions underlying metabolic and neuronal signaling pathways are temporally consistent in a moving avian hybrid zone.

The study of hybrid zones can provide insight into the genetic basis of species differences that are relevant for the maintenance of reproductive isolation. Hybrid zones can also provide insight into climate change, species distributions, and evolution. The hybrid zone between black-capped chickadees (Poecile atricapillus) and Carolina chickadees (Poecile carolinensis) is shifting northward in response to increasing winter temperatures but is not increasing in width. This pattern indicates strong selection against chickadees with admixed genomes. Using high-resolution genomic data, we identified regions of the genomes that are outliers in both time points and do not introgress between the species; these regions may be involved in the maintenance of reproductive isolation. Genes involved in metabolic regulation processes were overrepresented in this dataset. Several gene ontology categories were also temporally consistent-including glutamate signaling, synaptic transmission, and catabolic processes-but the nucleotide variants leading to this pattern were not. Our results support recent findings that hybrids between black-capped and Carolina chickadees have higher basal metabolic rates than either parental species and suffer spatial memory and problem-solving deficits. Metabolic breakdown, as well as spatial memory and problem-solving, in hybrid chickadees may act as strong postzygotic isolation mechanisms in this moving hybrid zone.

An amphibian species pushed out of Britain by a moving hybrid zone.

Classical theory states that hybrid zones will be stable in troughs of low population density where dispersal is hampered. Yet, evidence for moving hybrid zones is mounting. One possible reason that moving zones have been underappreciated is that they may drive themselves into oblivion and with just the superseding species remaining, morphological and genetic signals of past species replacement may be difficult to appreciate. Using genetic data (32 diagnostic single nucleotide polymorphisms) from a clinal hybrid zone of the common toad (Bufo bufo) and the spined toad (Bufo spinosus) in France for comparison, alleles of the latter species were documented in common toads in the south of Great Britain, at frequencies in excess of 10%. Because long distance dispersal across the Channel is unlikely, the conclusion reached was that the continental toad hybrid zone which previously extended into Britain, moved southwards and extirpated B. spinosus. Species distribution models for the mid‐Holocene and the present support that climate has locally changed in favour of B. bufo. The system bears resemblance with the demise of Homo neanderthalensis and the rise of Homo sapiens and provides an example that some paleoanthropologists demanded in support of a hominin “leaky replacement” scenario. The toad example is informative just because surviving pure B. spinosus and an extant slowly moving interspecific hybrid zone are available for comparison.

Historical hybrid zone movement: More pervasive than appreciated

AbstractHybrid zones are established where two divergent populations meet and interbreed, but experience some reproductive isolation. If one population expands its range at the expense of the other, their hybrid zone moves. While hybrid zone movement is generally considered to be uncommon and insignificant, recent studies challenge this idea. The commonality of contemporary hybrid zone movement—with shifts in hybrid zones tracked over years to decennia—cannot be disputed, given the many examples available. Cases of historical hybrid zone movement—covering centuries or millennia of mobility—are accumulating, with movement having been inferred from five lines of evidence: (1) range shifts documented in the fossil/pollen record; (2) distribution dynamics derived from species distribution modelling; (3) enclaves of a displaced population persisting inside the range of an expanding one; (4) a peak of linkage disequilibrium at the leading edge of a moving hybrid zone; and (5) genome‐wide genetic traces of a displaced population, left behind in an expanding one. While most of these lines of evidence are not straightforward to interpret and/or broadly applicable, the latter—a genomic footprint of hybrid zone movement—promises to be particularly suitable to determine whether a hybrid zone has been on the move since its inception. I argue that historical hybrid zone movement is likely to be prevalent and deserves wider acknowledgement in historical biogeography.

Tracing the footprints of a moving hybrid zone under a demographic history of speciation with gene flow.

A lack of optimal gene combinations, as well as low levels of genetic diversity, is often associated with the formation of species range margins. Conservation efforts rely on predictive modelling using abiotic variables and assessments of genetic diversity to determine target species and populations for controlled breeding, germplasm conservation and assisted migration. Biotic factors such as interspecific competition and hybridization, however, are largely ignored, despite their prevalence across diverse taxa and their role as key evolutionary forces. Hybridization between species with well‐developed barriers to reproductive isolation often results in the production of offspring with lower fitness. Generation of novel allelic combinations through hybridization, however, can also generate positive fitness consequences. Despite this possibility, hybridization‐mediated introgression is often considered a threat to biodiversity as it can blur species boundaries. The contribution of hybridization towards increasing genetic diversity of populations at range margins has only recently gathered attention in conservation studies. We assessed the extent to which hybridization contributes towards range dynamics by tracking spatio‐temporal changes in the central location of a hybrid zone between two recently diverged species of pines: Pinus strobiformis and P. flexilis. By comparing geographic cline centre estimates for global admixture coefficient with morphological traits associated with reproductive output, we demonstrate a northward shift in the hybrid zone. Using a combination of spatially explicit, individual‐based simulations and linkage disequilibrium variance partitioning, we note a significant contribution of adaptive introgression towards this northward movement, despite the potential for differences in regional population size to aid hybrid zone movement. Overall, our study demonstrates that hybridization between recently diverged species can increase genetic diversity and generate novel allelic combinations. These novel combinations may allow range margin populations to track favourable climatic conditions or facilitate adaptive evolution to ongoing and future climate change.

Testing an hypothesis of hybrid zone movement for toads in France.

Hybrid zone movement may result in substantial unidirectional introgression of selectively neutral material from the local to the advancing species, leaving a genetic footprint. This genetic footprint is represented by a trail of asymmetric tails and displaced cline centres in the wake of the moving hybrid zone. A peak of admixture linkage disequilibrium is predicted to exist ahead of the centre of the moving hybrid zone. We test these predictions of the movement hypothesis in a hybrid zone between common (Bufo bufo) and spined toads (B.spinosus), using 31 nuclear and one mtDNA SNPs along a transect in the northwest of France. Average effective selection in Bufo hybrids is low and clines vary in shape and centre. A weak pattern of asymmetric introgression is inferred from cline discordance of seven nuclear markers. The dominant direction of gene flow is from B.spinosus to B.bufo and is in support of southward movement of the hybrid zone. Conversely, a peak of admixture linkage disequilibrium north of the hybrid zone suggests northward movement. These contrasting results can be explained by reproductive isolation of the B.spinosus and B.bufo gene pools at the southern (B.spinosus) side of the hybrid zone. The joint occurrence of asymmetric introgression and admixture linkage disequilibrium can also be explained by the combination of low dispersal and random genetic drift due to low effective population sizes.

Hybrid zone formation and contrasting outcomes of secondary contact over transects in common toads.

Much progress in speciation research stems from documenting patterns of morphological and genetic variation in hybrid zones. Contrasting patterns of marker introgression in different sections of the contact can provide valuable insights on the relative importance of various evolutionary mechanisms maintaining species differences in the face of hybridization and gene flow and on hybrid zone temporal and spatial dynamics. We studied species interactions in the common toads Bufo bufo and B.spinosus in France and northwestern Italy using morphological and molecular data from the mitochondrial and nuclear genomes in an extensive survey, including two independent transects west and east of the Alps. At both, we found sharp, coincident and concordant nuclear genetic transitions. However, morphological clines were wider or absent and mtDNA introgression was asymmetric. We discuss alternative, nonexclusive hypotheses about evolutionary processes generating these patterns, including drift, selection, long-distance dispersal and spatial shifts in hybrid zone location and structure. The distribution of intraspecific mtDNA lineages supports a scenario in which B.bufo held a local refugium during the last glacial maximum. Present-day genetic profiles are best explained by an advance of B.spinosus from a nearby Iberian refugium, largely superseding the local B.bufo population, followed by an advance of B.bufo from the Balkans, with prongs north and south of the Alps, driving B.spinosus southwards. A pendulum moving hybrid zone, first northwards and then southwards, explains the wide areas of introgression at either side of the current position of the contact zones.

A genomic footprint of hybrid zone movement in crested newts.

Speciation typically involves a stage in which species can still exchange genetic material. Interspecific gene flow is facilitated by the hybrid zones that such species establish upon secondary contact. If one member of a hybridizing species pair displaces the other, their hybrid zone would move across the landscape. Although theory predicts that moving hybrid zones quickly stagnate, hybrid zones tracked over one or a few decades do not always follow such a limitation. This suggests that hybrid zones have the potential to traverse considerable distances over extended periods of time. When hybrid zones move, introgression is predicted to result in biased gene flow of selectively neutral alleles, from the receding species into the advancing species. We test for such a genomic footprint of hybrid zone movement in a pair of crested newt species (genus Triturus) for which we have a priori support for westward hybrid zone movement. We perform a multilocus phylogeographical survey and conduct Bayesian clustering analysis, estimation of ancestry and heterozygosity, and geographical cline analysis. In a 600 km wide area east of the present day hybrid zone a genomic footprint constitutes empirical evidence consistent with westward hybrid zone movement. The crested newt case suggests that hybrid zone movement can occur over an extensive span of time and space. Inferring hybrid zone movement provides fundamental insight into historical biogeography and the speciation process, and we anticipate that hybrid zones will prove to be far more mobile than currently appreciated.

Evidence for concerted movement of nuclear and mitochondrial clines in a lizard hybrid zone.

Moving hybrid zones provide compelling examples of evolution in action, yet long-term studies that test the assumptions of hybrid zone stability are rare. Using replicated transect samples collected over a 10-year interval from 2002 to 2012, we find evidence for concerted movement of genetic clines in a plateau fence lizard hybrid zone (Sceloporus tristichus) in Arizona. Cline-fitting analyses of SNP and mtDNA data both provide evidence that the hybrid zone shifted northward by approximately 2km during the 10-year interval. For each sampling period, the mtDNA cline centre is displaced from the SNP cline centre and maintaining an introgression distance of approximately 3km. The northward expansion of juniper trees into the Little Colorado River Basin in the early 1900s provides a plausible mechanism for hybrid zone formation and movement, and a broadscale quantification of recent land cover change provides support for increased woody species encroachment at the southern end of the hybrid zone. However, population processes can also contribute to hybrid zone movement, and the current stability of the ecotone habitats in the centre of the hybrid zone suggests that movement could decelerate in the future.

Differences in aggressive responses do not contribute to shifts in a sapsucker hybrid zone

ABSTRACT Interspecific interactions, such as aggressive territorial behaviors, can be important in establishing range boundaries, especially in hybrid zones where 2 divergent taxa meet and interbreed. Further, differences in the aggressive responses among hybridizing taxa can be important in moving hybrid zones, especially when there are strong asymmetries in their aggressive interactions. To determine how between-taxon interactions may be contributing to hybrid zone movement between 2 species of sapsucker, we studied an aspect of territory maintenance: aggression toward a territory intruder between species. Using a series of field experiments involving playback and taxidermic mounts, we tested whether Red-naped (Sphyrapicus nuchalis) and Red-breasted (S. ruber) sapsucker differ in their levels of aggressive behavior, thereby influencing the dynamic nature of the hybrid zone between these species. We found that 2 of the 3 aggressive response variables we measured differed between the 2 species. Red-naped ...

Stabilization of a salamander moving hybrid zone.

When related species meet upon postglacial range expansion, hybrid zones are frequently formed. Theory predicts that such zones may move over the landscape until equilibrium conditions are reached. One hybrid zone observed to be moving in historical times (1950–1979) is that of the pond‐breeding salamanders Triturus cristatus and Triturus marmoratus in western France. We identified the ecological correlates of the species hybrid zone as elevation, forestation, and hedgerows favoring the more terrestrial T. marmoratus and pond density favoring the more aquatic T. cristatus. The past movement of the zone of ca. 30 km over three decades has probably been driven by the drastic postwar reduction of the “bocage” hedgerow landscape, favoring T. cristatus over T. marmoratus. No further hybrid zone movement was observed from 1979 to the present. To explain the changing dynamics of the hybrid zone, we propose that it stalled, either because an equilibrium was found at an altitude of ca. 140 m a.s.l. or due to pond loss and decreased population densities. While we cannot rule out the former explanation, we found support for the latter. Under agricultural intensification, ponds in the study area are lost at an unprecedented rate of 5.5% per year, so that remaining Triturus populations are increasingly isolated, hampering dispersal and further hybrid zone movement.

Tracing reinforcement through asymmetrical partner preference in the European common vole Microtus arvalis.

BackgroundThe mechanistic basis of speciation and in particular the contribution of behaviour to the completion of the speciation process is often contentious. Contact zones between related taxa provide a situation where selection against hybridization might reinforce separation by behavioural mechanisms, which could ultimately fully isolate the taxa. One of the most abundant European mammals, the common vole Microtus arvalis, forms multiple natural hybrid zones where rapidly diverging evolutionary lineages meet in secondary contact. Very narrow zones of hybridization spanning only a few kilometres and sex-specific gene flow patterns indicate reduced fitness of natural hybrids and incipient speciation between some of the evolutionary lineages. In this study, we examined the contribution of behavioural mechanisms to the speciation process in these rodents by fine-mapping allopatric and parapatric populations in the hybrid zone between the Western and Central lineages and experimental testing of the partner preferences of wild, pure-bred and hybrid female common voles.ResultsGenetic analysis based on microsatellite markers revealed the presence of multiple parapatric and largely non-admixed populations at distances of about 10 km at the edge of the area of natural hybridization between the Western and Central lineages. Wild females from Western parapatric populations and lab-born F1 hybrids preferred males from the Western lineage whereas wild females of Central parapatric origin showed no measurable preference. Furthermore, wild and lab-born females from allopatric populations of the Western or Central lineages showed no detectable preference for males from either lineage.ConclusionsThe detected partner preferences are consistent with asymmetrical reinforcement of pre-mating reproductive isolation mechanisms in the European common vole and with earlier results suggesting that hybridization is more detrimental to the Western lineage. As a consequence, these differences in behaviour might contribute to a further geographical stabilization of this moving hybrid zone. Such behavioural processes could also provide a mechanistic perspective for frequently-detected asymmetrical introgression patterns in the largely allopatrically diversifying Microtus genus and other rapidly speciating rodents.

An Integrated Framework for Hybrid Zone Models

Hybridization, where distinguishable populations meet and interbreed resulting in hybrid offspring, is useful for understanding the mechanisms involved in the evolution of reproductive isolation. Hybridization can range from interactions with near-complete reproductive isolation, to stable zones, to complete intergradation. Several models explain how stable hybrid zones are maintained, but each model emphasizes a special case and some zones fit more than one model. The models also do not emphasize moving hybrid zones or occasional hybridization events that add genetic variation to the parental populations. This essay unifies the current models along a continuum of selection pressures organized by geography. Current models differ in assumed hybrid fitness and whether selection is intrinsic or extrinsic. The type of selection is a less general category than the direction of selection (for or against hybrids); multiple types of selection can exist in each zone depending on trait. Thus, the diversity and combinations of selection types make selection direction the most general category for sorting hybrid zones, and variation in selection over geography shapes the spatial extent (in concert with dispersal), movement, and stability of a hybrid zone. This framework is useful for considering zone structure both at the whole organism level and at individual genes or traits. Unifying hybrid zones models by the geographic range of selection pressure organizes each zone into context along the continuum regardless of its fit within one of the classic models, helping us explicitly consider what assumptions are made about the zone and providing direction for further research.

Integrating phylogeography and paleoecology to investigate the origin and dynamics of hybrid zones: insights from two widespread North American firs.

Secondary contact between closely related taxa routinely occurs during postglacial migrations. After initial contact, the location of hybrid zones may shift geographically or remain spatially stable over time in response to various selective pressures or neutral processes. Studying the extent and direction of introgression using markers having contrasted levels of gene flow can help unravel the historical dynamics of hybrid zones. Thanks to their contrasted maternal and paternal inheritance, resulting in different levels of gene flow for mitochondrial and chloroplast DNA (mtDNA and cpDNA), the Pinaceae stand out as a relevant biological model for this purpose. The objective of the study was to assess whether the hybrid zone between Abies balsamea and Abies lasiocarpa (two largely distributed Pinaceae) has moved or remained stable over time by analysing the distribution of cytoplasmic DNA variation as well as published palaeobotanical data. Interspecific gene flow was higher for cpDNA than mtDNA markers; hence, the geographic distribution of mitotypes was more congruent with species distributions than chlorotypes. This genetic signature was contrary to expectations under a moving hybrid zone scenario, as well as empirical observations in other conifers. Genetic evidence for this rare instance of stable hybrid zone was corroborated by the colonization chronology derived from published fossil data, indicating that the two fir species initially came into contact in the area corresponding to the current sympatric zone 11 kyr ago. While an explanatory analysis suggested the putative influence of various environmental factors on the relative abundance of cytoplasmic genome combinations, further research appears necessary to assess the role of both demographic history and selective factors in driving the dynamics of hybrid zones.