Environmental factors, microgeographic patterns of endemism and hybrid zones in Monobella grassei (Insecta: Collembola: Neanuridae)

A complex pattern of narrow parapatric distributions with hybrid zones is described among subspecies of the Collembola species Monobella grassei in the eastern Pyrenees. Though hybrid zones are usually narrow, they vary from simple contact zones without recorded hybrids to relatively wide hybrid swarms, suggesting that large differences exist locally in the level of genetic divergence between subspecies. The course and structure of contact zones are analysed on morphological grounds, in relation to environmental (rainfall, vegetation, substrate) and topographical (altitude, hydrography) parameters. The results show that, unlike most cases documented so far, the courses of the contact zones in M. grassei are not connected to major ecological boundaries nor to ecological ecotones, on either the regional or the habitat scale. It is suggested that this is due to the great ecological similarity between the different parapatric subspecies of M. grassei , and their broad ecological requirements. There is no conclusive evidence that the distribution observed is linked to recent palaeoclimatic events such as glaciation. Endogenous selection could therefore be prevalent in this model to explain parapatry and the hybrid zone pattern.     

Species distribution models contribute to determine the effect of climate and interspecific interactions in moving hybrid zones

Climate is a major factor delimiting species’ distributions. However, biotic interactions may also be prominent in shaping geographical ranges, especially for parapatric species forming hybrid zones. Determining the relative effect of each factor and their interaction of the contact zone location has been difficult due to the lack of broad scale environmental data. Recent developments in species distribution modelling (SDM) now allow disentangling the relative contributions of climate and species’ interactions in hybrid zones and their responses to future climate change. We investigated the moving hybrid zone between the breeding ranges of two parapatric passerines in Europe. We conducted SDMs representing the climatic conditions during the breeding season. Our results show a large mismatch between the realized and potential distributions of the two species, suggesting that interspecific interactions, not climate, account for the present location of the contact zone. The SDM scenarios show that the southerly distributed species, Hippolais polyglotta, might lose large parts of its southern distribution under climate change, but a similar gain of novel habitat along the hybrid zone seems unlikely, because interactions with the other species (H. icterina) constrain its range expansion. Thus, whenever biotic interactions limit range expansion, species may become ‘trapped’ if range loss due to climate change is faster than the movement of the contact zone. An increasing number of moving hybrid zones are being reported, but the proximate causes of movement often remain unclear. In a global context of climate change, we call for more interest in their interactions with climate change.     

Hybridization of Bombina bombina and B. variegata (Anura, Discoglossidae) at a sharp ecotone in western Ukraine: comparisons across transects and over time

Abstract Bombina bombina and B. variegata are two anciently diverged toad taxa that have adapted to different breeding habitats yet hybridize freely in zones of overlap where their parapatric distributions meet. Here, we report on a joint genetic and ecological analysis of a hybrid zone in the vicinity of Stryi in western Ukraine. We used five unlinked allozyme loci, two nuclear single nucleotide polymorphisms and a mitochondrial DNA haplotype as genetic markers. Parallel allele frequency clines with a sharp central step occur across a sharp ecotone, where transitions in aquatic habitat, elevation, and terrestrial vegetation coincide. The width of the hybrid zone, estimated as the inverse of the maximum gradient in allele frequency, is 2.3 km. This is the smallest of four estimates derived from different clinal transects across Europe. We argue that the narrow cline near Stryi is mainly due to a combination of habitat distribution and habitat preference. Adult toads show a preference for either ponds ( B. bombina ) or puddles ( B. variegata ), which is known to affect the distribution of genotypes within the hybrid zones. At Stryi, it should cause a reduction of the dispersal rate across the ecotone and thus narrow the cline. A detailed comparison of all five intensively studied Bombina transects lends support to the hypothesis that habitat distribution plus habitat preference can jointly affect the structure of hybrid zones and, ultimately, the resulting barriers to gene flow between differentiated gene pools. This study also represents a resampling of an area that was last studied more than 70 years ago. Our allele-frequency clines largely coincide with those that were described then on the basis of morphological variation. However, we found asymmetrical introgression of B. variegata genes into B. bombina territory along the bank of a river.     

Temporal habitat variability and the maintenance of sex in host populations of the pea aphid

The evolutionary maintenance of sex, despite competition from asexual reproduction, has long intrigued the evolutionary biologists owing to its numerous apparent short-term costs. In aphids, winter climate is expected to determine the maintenance of sexual lineages in the high latitude zones owing to their exclusive ability to produce frost-resistant eggs. However, diverse reproductive modes may coexist at a local scale where climatic influence is counteracted by microgeographical factors. In this study, we tested the influence of local habitat characteristics on regional coexistence of reproductive modes in the pea aphid, Acyrthosiphon pisum. In the laboratory, the induction of sexual morph production of many pea aphid genotypes from the local fields of annual (pea and faba bean) and perennial (alfalfa and red clover) crops in Western France indicated that A. pisum lineages from annual crops had a significantly higher investment in sexual reproduction than A. pisum lineages from the perennial hosts. We propose that temporal habitat variability exerts a selective pressure to maintain the sexual reproduction in A. pisum. The ecological and evolutionary consequences of the association between the mode of reproduction and the host population on gene flow restriction and on ecological specialization are discussed.     

Intrinsic reproductive isolating mechanisms in the maintenance of a hybrid zone between ecologically divergent subspecies

Understanding factors involved in maintaining stable hybrid zones is important for predicting the ultimate fate of the interacting taxa, but the relative importance of mechanisms such as ecological selection and intrinsic reproductive isolation remains unclear. Most studies of reproductive isolation in hybrid zones have focused either on zones with strongly bimodal patterns in genotype or phenotype frequencies, with relatively strong isolation, or unimodal zones with relatively weak isolation, whereas less is known about more intermediate classes of hybrid zone. Here, we utilize a hybrid zone of this intermediate type occurring between northern and southern subspecies of Atlantic killifish, Fundulus heteroclitus, to identify isolating mechanisms playing a role in maintaining this type of zone. The two subspecies differ in environmental tolerance, and we found some evidence of microhabitat preference between subspecies within a small tidal creek at the centre of the hybrid zone. There was also an association between sex, mitochondrial genotype and habitat within this creek. Fertilization success did not differ between consubspecific and heterosubspecific crosses, but hatching success was significantly lower for crosses involving southern males and northern females, and crosses between southern females and northern males had altered developmental rates. Southern females and northern males showed patterns consistent with positive assortative mating. Together, these results indicate a role for a combination of factors including assortative mating and/or early hybrid inviability in the maintenance of this hybrid zone and suggest that hybrid zones with intermediate levels of reproductive isolation are likely to be maintained by multiple interacting isolating mechanisms.     

A spatially explicit individual-based model of reinforcement in hybrid zones

Abstract An individual-based model consisting of two dioecious populations in a two-dimensional environmental grid was constructed. Each population began with, and never exceeded, 1000 individuals; extinction was allowed. Genomes consisting of 30 biallelic loci for male sexual advertisement call, female mate preference, and population origin were constructed, and lineages of each individual in the starting populations were followed for 2000 generations. Type and level of hybrid disadvantage, initial population distribution, patchiness of environmental resources, and level of mate choice were varied. Persistence of bimodal hybrid zones was nonexistent at low levels of hybrid disadvantage and universal at high levels of hybrid disadvantage, with a narrow threshold in which persistence was unpredictable. Persistence occurred at lower levels of hybrid disadvantage when populations were initially parapatric rather than sympatric, and environments were patchy rather than homogeneous. Increased divergence in mating systems occurred when hybrid disadvantage was high, hybrids were infertile, populations were initially parapatric, and increased female choice was allowed. Mating system divergence was much higher in interacting populations compared with noninteracting populations, indicating that reinforcement caused most of the observed divergence. When hybrids were infertile, reinforcement contributed to speciation, because under hybrid infertility the probability of persistence at low levels of hybrid disadvantage was positively related to mate choice. The results agree with previous one-dimensional spatial models in finding that population persistence is more likely in parapatric and patchy population distributions. In addition, the results show that hybrid infertility may facilitate the process of reinforcement and speciation.     

When invaders meet locally adapted types: rapid moulding of hybrid zones between sculpins (Cottus, Pisces) in the Rhine system

Hybrid zones are commonly studied to dissect the processes that drive divergence among lineages, which have incomplete barriers of reproduction. Most hybrid zones have existed for an extended time making inferences on the initial mode of formation difficult. It is a priori unclear how fast a nascent hybrid zone would form as a response to endogenous and exogenous factors. We have studied several hybrid zones between two lineages of sculpins (Cottus spp.), which emerged due to a recent range expansion of one of the lineages along the river Rhine in the early 1990s. Applying a dense sampling across two contact areas and using a highly informative set of 45 microsatellite markers we found pronounced genetic structure. Steep genetic clines suggest that strong selective forces have shaped the respective hybrid zones from the beginning. We find that the zones are coupled to ecological transitions from small streams to larger rivers. The width of these zones is much smaller than estimates of annual individual dispersal distances, as estimated outside of the hybrid zones. The pattern is apparently not strongly affected by pre- or postzygotic reproductive isolation because numerous backcross hybrids occur within the zones. This suggests that strong natural selection acts against immigrant genotypes. The study exemplifies how local adaptation can play a key role in preventing admixture in dependence of the ecological context.     

Ecology and genetics of hybrid zones in the southern African Pycnonotus bulbul species complex

Lloyd, P., Craig, A.J.F.K., Hulley, P.E., Essop, M.F., Bloomer, P. & Crowe, T.M. 1997. Ecology and genetics of hybrid zones in the southern African Pycnonotus bulbul species complex. Ostrich 68 (2–4): 90–96.

The closely related Blackeyed Bulbul Pycnonotus barbatus, Cape Bulbul P. capensis and Redeyed Bulbul P. nigricans have parapatric to locally sympatric distributions within southern Africa. Extensive hybridization along narrow transition zones between each of the three species pairs is described in a region of the Eastern Cape province, South Africa. The transition zones coincide with ecotones between different vegetation types, which in turn follow escarpments or mountain ranges. The lack of population density depressions within the hybrid zones, together with the variability of the hybrids, suggests the hybrids are viable. Sharp step clines in various phenotypic characters are described across the P. barbatus/P. nigricans hybrid zone. A mtDNA analysis found evidence of possible introgression between P. barbatus and P. capensis. All eight P. barbatus x P. nigricans hybrids analysed possessed P. barbatus mtDNA, suggesting the existence of either positive assortative mating or strong directional selection, but our data are unable to distinguish which. Our results do not support the dynamic-equilibrium model, but are compatible with the bounded-hybrid-superiority model. We conclude that the maintenance of the parapatric distributions of the different taxa is due mainly to differences in environmentally-associated fitness between parental phenotypes or among parental and hybrid phenotypes along an ecotone, with the narrowness of the hybrid zones maintained by the steepness of the environmental gradients crossing them.     

Evidence for natural selection in a fern hybrid zone

Morphological characters, chloroplast DNA, and allozymes were used to analyze the distribution of individuals within a hybrid population of the ferns Polystichum munitum and P. imbricans in northwestern California. Microsites within the population were characterized according to soil moisture and light levels reaching the plants. In sites with low soil moisture and high light levels, all of the ferns were genetically and morphologically like P. imbricans. In contrast, ferns with the genetic and morphological identity of P. munitum predominated in moist shady sites. Intermediate sites supported very few P.munitum, a wide variety of hybrid recombinants, and a majority of ferns with P. imbricans characteristics. The pattern of variation within the population is noteworthy because of the close proximity of the habitat extremes and the long-range dispersal of fern spores. We conclude that natural selection along environmental gradients must be a major factor in determining the ecological and genetic associations within the hybrid zone. The results of this study are evaluated in the context of the fern life cycle and compared to the assumptions of models explaining the establishment and maintenance of hybrid zones, which vary in the role attributed to environmentally mediated natural selection.     

Do ecological niches differ between sexual and asexual lineages of an aphid species?

According to environmental-based theories on the maintenance of sexual reproduction, sexual and asexual populations may coexist if they occupy different ecological niches. The aphid Rhopalosiphum padi offers a good opportunity to test this hypothesis since sexual and asexual lineages show local coexistence during a large part of their respective life-cycles. Because these two reproductive variants are morphologically identical but genetically distinct, we first characterized them using genetic markers in populations of R. padi in areas where sexual and asexual lineages may occur in sympatry. We then inferred the natal host plant of sexual and asexual genotypes by analysing stable isotopic ratios and showed that sexual ones mostly originated from C₃ Poaceae while asexual ones originated from C₃ and C₄ plants, although the majority came from C₄ Poaceae. These findings indicate that ecological niches of sexual and asexual lineages of R. padi differ, offering a plausible explanation for the local coexistence of the two reproductive modes in this species through habitat specialisation.     

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

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

Reproductive interference, priority effects and the maintenance of parapatry in Skistodiaptomus copepods

Reproductive interactions among species, such as attempts to mate with heterospecifics, can have profound impacts on patterns of geographic range and co-occurrence. For example, several theoretical models demonstrate that reproductive interference-the negative influence of heterospecifics on the abilities of conspecifics to successfully mate-can lead to parapatry, even across homogeneous landscapes, when the parapatric species are initially allopatric. However, the potential consequences of reproductive interactions on patterns of range limits and co-occurrence have been largely ignored by ecologists. Here, I use a combination of laboratory mating experiments and a genetic survey for two parapatric species of Skistodiaptomus copepods to evaluate the potential importance of interspecific reproductive interactions on the maintenance of parapatry. The genetic survey demonstrates phylogenetic exclusivity of these species, suggesting that gene flow has not occurred between them. Moreover, laboratory crosses between both species demonstrate that their parapatric boundary is not maintained as a hybrid zone because the two species cannot form viable hybrids. However, Skistodiaptomus oregonensis and S. pygmaeus males mate indiscriminately and interfere with the ability of heterospecific females to successfully reproduce. I suggest that reproductive interference results in priority effects that strongly influence the maintenance of this parapatric boundary and stress that reproductive traits are important and often-overlooked in ecological models for species range limits.     

Phylogeography of a widespread lizard complex reflects patterns of both geographic and ecological isolation

A primary challenge for modern phylogeography is understanding how ecology and geography, both contemporary and historical, shape the spatial distribution and evolutionary histories of species. Phylogeographic patterns are the result of many factors, including geology, climate, habitat, colonization history and lineage‐specific constraints. Assessing the relative influences of these factors is difficult because few species, regions and environments are sampled in enough detail to compare competing hypotheses rigorously and because a particular phylogeographic pattern can potentially result from different evolutionary scenarios. The silky anoles (Anolis sericeus complex) of Central America and Mexico are abundant and found in all types of lowland terrestrial habitat, offering an excellent opportunity to test the relative influences of the factors affecting diversification. Here, we performed a range‐wide statistical phylogeographic analysis on restriction site‐associated DNA (RAD) markers from silky anoles and compared the phylogeographic patterns we recovered to historical and contemporary environmental and topographic data. We constructed niche models to compare niche overlap between sister lineages and conducted coalescent simulations to characterize how the major lineages of silky anoles have diverged. Our results revealed that the mode of divergence for major lineage diversification events was geographic isolation, resulting in ecological divergence between lineages, followed by secondary contact. Moreover, comparisons of parapatric sister lineages suggest that ecological niche divergence contributed to isolation by environment in this system, reflecting the natural history differences among populations in divergent environments.     

Phylogenetic evidence for hybrid origins of asexual lineages in an aphid species

Understanding the mode of origin of asexuality is central to ongoing debates concerning the evolution and maintenance of sexual reproduction in eukaryotes. This is because it has profound consequences for patterns of genetic diversity and ecological adaptability of asexual lineages, hence on the outcome of competition with sexual relatives both in short and longer terms. Among the possible routes to asexuality, hybridization is a very common mechanism in animals and plants. Aphids present frequent transitions from their ancestral reproductive mode (cyclical parthenogenesis) to permanent asexuality, but the mode of origin of asexual lineages is generally not known because it has never been thoroughly investigated with appropriate molecular tools. Rhopalosiphum padi is an aphid species with coexisting sexual (cyclically parthenogenetic) and asexual (obligately parthenogenetic) lineages that are genetically distinct. Previous studies have shown that asexual lineages of R. padi are heterozygous at most nuclear loci, suggesting either that they have undergone long-term asexuality (under which heterozygosity tends to increase) or that they have hybrid origins. To discriminate between these alternatives, we conducted an extensive molecular survey combining the sequence analysis of alleles of two nuclear DNA markers and mitochondrial DNA haplotypes in sexual and asexual lineages of R. padi. Both nuclear and cytoplasmic markers clearly showed that many asexual lineages have hybrid origins, the first such demonstration in aphids. Our results also indicated that asexuals result from multiple events of hybridization between R. padi and an unknown sibling species, and are of recent origin (contradicting previous estimates that asexual R. padi lineages were of moderate longevity). This study constitutes another example that putatively ancient asexual lineages are actually of much more recent origin than previously thought. It also presents a robust approach for testing whether hybrid origin of asexuality is also a common phenomenon in aphids.     

A hybrid zone dominated by fertile F1s: maintenance of species barriers in Rhododendron

Isolating barriers between interbreeding sympatric or parapatric interfertile species are maintained by processes that occur within their hybrid zones. Although the effects of intrinsic selection on hybrid fitness are well known, less is understood about extrinsic fitness variation. At Tiryal Dag, northeast Turkey, Rhododendron x sochadzeae (R. ponticum x caucasicum) forms large populations in which neither segregation nor backcrossing occur, in habitats intermediate between those of its parents. Using single-copy species-specific random amplified polymorphic DNA and inter simple sequence repeat markers, it was determined that most or all R. x sochadzeae plants are F1s, and that there are many separate genets present. Hand pollination and germination experiments showed that R. x sochadzeae plants can produce viable seed of F2s or backcrosses in either direction. Furthermore, adult backcrosses have been observed in habitats atypical for R. x sochadzeae. From this, all non-F1 hybrid derivatives appear to be eliminated in the hybrid zone at Tiryal Dag as a result of postgermination selection. This absence of post-F1 hybrid derivatives apparently prevents introgression. This type of hybrid population is here termed an F1-dominated hybrid zone (F1DZ), and also occurs in Encelia. The observed dominance of F1s within a narrow habitat range is best explained by habitat-mediated superiority of F1s over all other genotype classes. Therefore, habitat-mediated selection against the second hybrid generation might be preventing interspecific gene flow in R. x sochadzeae. F1DZ formation is postulated to require the formation of F1s in quantity, habitat-mediated superiority in F1s, and highly specific habitat conditions.     

Hybridization and introgression in two ecologically dissimilar Fundulus hybrid zones

Hybridization and introgression appear more common in rapidly evolving groups, suggesting an important role in the evolutionary process. Detailed studies of how extrinsic or intrinsic forces regulate hybridization and introgression have the potential for broadening our understanding of mechanisms generating diversity. Species in the Fundulus notatus species complex have broad overlapping ranges and occur in replicated hybrid zones along predictable stream gradients. Typical hybrid zone structure has Fundulus olivaceus in headwaters, F. notatus downstream, and hybrid zones near confluences or abrupt shifts in habitat. Rarely, the typical upstream‐downstream orientation is reversed raising questions as to how hybrid zones are formed and maintained. We used next‐generation sequencing data to study hybridization and introgression in hybrid zones in neighboring drainages that differ in orientation (typical and reversed). We predicted extrinsic forces linked to stream gradients would result in noticeable differences between the two. Contrary to predictions, the data indicate the hybrid zones are remarkably similar. We used individual‐based simulations to explore the potential role of intrinsic and extrinsic forces in generating and maintaining typical and reversed hybrid zones. Simulation results were consistent with reversed hybrid zones being formed from stochastic processes combined with strong intrinsic forces and weak extrinsic forces.     

Conservation Implications of Hybridization in African Cercopithecine Monkeys

Numerous field reports of hybrid monkeys and documented cases of persistent hybrid zones suggest that natural hybridization is common among African cercopithecines. Both theoretical considerations and a review of cases lead us to conclude that parapatric hybridization among closely related allotaxa is a widespread, usually natural process whose incidence may be modified by human influence. Sympatric hybridization, between species ecologically distinct enough to have overlapping ranges, is rarer, and in monkeys tends to occur in settings where natural or anthropogenic habitat edges restrict migration and hence access to unrelated conspecific mates. Although sympatric hybridization occurs in the absence of human disturbance, and may even have been a creative force in cercopithecine evolution, anthropogenic habitat fragmentation may increase its incidence. Hybridization with a more abundant form may increase the level of threat faced by a species whose numbers and range have been severely restricted, either naturally or artificially.     

Ecological and genetic assessment of spatial structure among replicate contact zones between two topminnow species

The spatial structure of contact zones is often described as disjunct, diffuse or mosaic and presumed to be related to underlying ecological gradients. However, the ecology of contact zones, how they are structured, and if that structure is predictable based on the strength and nature of ecological gradients is unknown. Large spatial scales and the unreplicated nature of many of the best studied contact zones has made it difficult to codify broader ecological patterns. Freshwater stream fish contact zones have the advantage of being potentially replicated with well defined boundaries and predictable linear gradients (river continuum concept). We sampled four replicate topminnow (Fundulus olivaceus and F. notatus) contact zones in Gulf of Mexico drainages. In each, we quantified contact zone spatial structure and the strength of ecological gradients (habitat, physicochemical variables and fish community functional traits). All three types of contact zone structure were represented. Systems with weaker gradients had diffuse contact zones, low species richness and were numerically dominated by generalist species. Rates of hybridization were also variable among systems. There was no hybridization detected in the mosaic zone while hybrids were found at most of the co-occurrence sites in the diffuse and disjunct zones. Overall, local ecology clearly influences contact zone structure and the two species interact in fundamentally different ways in these four systems.     

Characterizing a hybrid zone between a cryptic species pair of freshwater snails

Characterizing hybrid zones and their dynamics is a central goal in evolutionary biology, but this is particularly challenging for morphologically cryptic species. The lack of conspicuous divergence between parental types means intermediate hybrid forms often go undetected. We aimed to detect and characterize a suspected hybrid zone between a pair of morphologically cryptic lineages of the freshwater snail, Radix. We sampled Radix from across a contact zone between two mitochondrial lineages (Radix balthica and an undescribed lineage termed ‘MOTU3’) and detected admixture between two nuclear genotype clusters, which were significantly but not categorically associated with the mitochondrial lineages. Using a model selection approach, we show that the admixture cline is best explained by an interaction between precipitation and temperature gradients over the area, rather than geographic distance. We thus hypothesize that the correlation with climatic gradients suggests environmental selection has played a role in maintaining the hybrid zone. In a 2050 climate change scenario, we furthermore predict an expansion of one of the nuclear clusters and a widening of the hybrid zone as the climate warms and dries.     

Hybrid swarm between divergent lineages of mule deer (Odocoileus hemionus)

Studies of hybrid zones have revealed an array of evolutionary outcomes, yet the underlying structure is typically characterized as one of three types: a hybrid zone, a hybrid swarm or a hybrid taxon. Our primary objective was to determine which of these three structures best characterizes a zone of hybridization between two divergent lineages of mule deer (Odocoileus hemionus), mule deer and black-tailed deer. These lineages are morphologically, ecologically and genetically distinct, yet hybridize readily along a zone of secondary contact between the east and west slopes of the Cascade Mountains (Washington and Oregon, USA). Using microsatellite and mitochondrial DNA, we found clear evidence for extensive hybridization and introgression between lineages, with varying degrees of admixture across the zone of contact. The pattern of hybridization in this region closely resembles a hybrid swarm; based on data from 10 microsatellite loci, we detected hybrids that extend well beyond the F1 generation, did not detect linkage disequilibrium at the centre of the zone and found that genotypes were associated randomly within the zone of contact. Introgression was characterized as bidirectional and symmetric, which is surprising given that the zone of contact occurs along a sharp ecotone and that lineages are characterized by large differences in body size (a key component of mating success). Regardless of the underlying mechanisms promoting hybrid swarm maintenance, it is clear that the persistence of a hybrid swarm presents unique challenges for management in this region.