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opencc-zeroAug 2020View details →

Secondary contact zones of closely-related Erebia butterflies overlap with narrow phenotypic and parasitic clines

Zones of secondary contact between closely related taxa are a common legacy of the Quaternary ice ages. Despite their abundance, the factors that keep species apart and prevent hybridisation are often unknown. Here we study a very narrow contact zone between three closely related butterfly species of the Erebia tyndarus species complex. Using genomic data, we first determined if gene flow occurs and then assessed whether it might be hampered by differences in chromosome number between some species. We found interspecific gene flow between sibling species that differ in karyotype by one chromosome. Conversely, only F1 hybrids occurred between two species that have the same karyotype, forming a steep genomic cline. In a second step, we fitted clines to phenotypic, ecological and parasitic data to identify the factors associated with the genetic cline. We found clines for phenotypic data and the prevalence of the endosymbiont parasite Wolbachia to overlap with the genetic cline, suggesting that they might be drivers for separating the two species. Overall our results highlight that some gene flow is possible between closely-related species despite different chromosome numbers, but that other barriers restrict such gene flow.

opencc-zeroAug 2020View details →

Contrasting signatures of introgression in North American box turtle (Terrapene spp.) contact zones

Hybridization occurs differentially across the genome in a balancing act between selection and migration. With the unprecedented resolution of contemporary sequencing technologies, selection and migration can now be effectively quantified such that researchers can identify genetic elements involved in introgression. Furthermore, genomic patterns can now be associated with ecologically relevant phenotypes, given availability of annotated reference genomes. We do so in North American box turtles (Terrapene) by deciphering how selection affects hybrid zones at the interface of species-boundaries and identifying genetic regions potentially under selection that may relate to thermal adaptations. Such genes may impact physiological pathways involved in temperature-dependent sex determination, immune system functioning, and hypoxia tolerance. We contrasted these patterns across inter- and intra-specific hybrid zones that differ temporally and biogeographically. We demonstrate hybridization is broadly apparent in Terrapene, but with observed genomic cline patterns corresponding to species boundaries at loci potentially associated with thermal adaptation. These loci display signatures of directional introgression within intra-specific boundaries, despite a genome-wide selective trend against intergrades. In contrast, outlier loci for inter-specific comparisons exhibited evidence of being under selection against hybrids. Importantly, adaptations coinciding with species-boundaries in Terrapene overlap with climatic boundaries and highlight the vulnerability of these terrestrial ectotherms to anthropogenic pressures.

opencc-zeroSep 2020View details →

Contact zone of two different chloroplast lineages and genetic guidelines for seed transfer in Quercus serrata and Q. crispula

Within their natural distribution ranges, plant species exhibit genetic structure which has been created by global climate change and natural selection over long periods. To conserve local forests with different genetic structures, genetic guidelines for seed and seedling transfer in individual species are therefore necessary. Genetic guidelines have been published for 43 Japanese tree species using population genetic data; however, for practical use, more detailed genetic borders between important genetic lineages should be clarified to inform seed collection and planting. Thus, we investigated in detail the genetic borders between two important Japanese oak species, Quercus serrata and Q. crispula, in the Chubu region of Japan using chloroplast and nuclear DNA markers, and we discuss the factors that influenced border creation using the results of species distribution modelling (SDM). Two distinct cpDNA haplotypes were found for each species (northern and southern haplotype) within the Chubu region of Japan but the difference in nuclear DNA between northern and southern haplotype populations was very small both in Q. serrata and Q. crispula. The results of SDM showed that during the LGM Q. serrata was distributed mostly along the coastline but Q. crispula was distributed not only along the coast but also in mountainous areas further inland. The cpDNA genetic borders of these two oak species are complex and seem to have been influenced by topography and their distribution during the LGM. We propose and discuss genetic guidelines for these two oak species based on the results of this study.

opencc-zeroSep 2020View details →

Parasite turnover zone at secondary contact: a new pattern in host-parasite population genetics

We introduce a new pattern of population genetic structure in a host-parasite system that can arise after secondary contact of previously isolated populations. Due to different generation time and therefore different tempo of molecular evolution the host and parasite populations reach different degrees of genetic differentiation during their separation (e.g. in refugia). Consequently, during the secondary contact the host populations are able to re-establish a single panmictic population across the area of contact, while the parasite populations stop their dispersal at the secondary contact zone and create a narrow hybrid zone. From the host's perspective, the parasite's hybrid zone functions on a microevolutionary scale as a "parasite turnover zone": while the hosts are passing from area A to area B, their parasites turn genetically from the area A genotypes to the area B genotypes. We demonstrate this novel pattern on a model composed of Apodemus mice and Polyplax lice by comparing maternally inherited markers (complete mitochondrial genomes, and complete genomes of vertically transmitted symbiont Legionella polyplacis) with SNPs derived from the louse genomic data. We discuss circumstances that may lead to this pattern and possible reasons why it has been overlooked in the studies on host parasite population genetics.

Figure 5 in Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species

Figure 5. Cline analyses of mitochondrial DNA and microsatellite data. Transects (top) through the two different contact zones of grass snake lineages (helvetica/eastern lineages – left; yellow/red lineages – right) and associated Maximum Likelihood clines for microsatellites (centre) and mtDNA (bottom). Grey: fuzzy 95% credible cline region. Red points (top) indicate cline centres. Maps were created using ARCGIS 10.2 (http://www. esri.com/arcgis) and ADOBE ILLUSTRATOR CS6 (http://www.adobe.com/products/illustrator.html).

Figure 4. PCA axes 1–2 in Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species

Figure 4. PCA axes 1–2 for microsatellite data. Samples are coloured according to mitochondrial lineages (top) or STRUCTURE clusters (bottom). Admixed individuals were identified according to HYBRIDLAB results. PCAs for the yellow and red lineages correspond to the samples from Fig. 3c. Non-native samples were excluded. The oval outlines represent 95% confidential intervals. For helvetica and the eastern lineages (left) the x axis explains 16.6% and the y axis 4.5% of variation. For the eastern lineages (right) the x axis explains 3.8% and the y axis 2.9% of variation. Analyses along axes 1–3 produced nearly identical results (see Supplementary Fig. S4).

Figure 3 in Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species

Figure 3. Genotypic structuring of grass snakes. On the left, the mitochondrial lineage of each sample is shown above the STRUCTURE diagrams, with haplotypes of Natrix natrix helvetica indicated in blue and haplotypes of the eastern lineages in colours corresponding to Fig. 1 (yellow, red, lilac, grey, green; white = missing data). In (a), orange and dark blue corresponds to non-native snakes (Italian lineages). Samples in STRUCTURE diagrams are arranged within each country from west to east (a) or from north to south (b,c). In STRUCTURE diagrams, an individual sample is represented by a vertical bar reflecting its inferred ancestry. In (a), the blue cluster corresponds to N. n. helvetica and the light green cluster to all other lineages. The isolated red/light green block (first row) represents the allochthonous population from the Neander valley, Germany. In (b), samples with genetic impact of helvetica are excluded. The pink cluster corresponds to samples from the yellow and red lineages. Brown percentages indicate genetic impact of adjacent lineages (lilac, grey, green). In (c) only samples from the yellow and red lineages and their hybrids, without genetic signatures of other lineages, were processed. Country abbreviations: Ba – Balkans (Albania, Bosnia and Herzegovina, Montenegro, Serbia, Kosovo, Former Yugoslav Republic of Macedonia, Romania, Bulgaria, and Greece), CH – Switzerland, CRO – Croatia, CZ – Czech Republic, FI – Finland, H – Hungary, N – Norway, NL – Netherlands, PL – Poland, S – Sweden. Maps were created using ARCGIS 10.2 (http://www.esri.com/arcgis) and ADOBE ILLUSTRATOR CS6 (http://www.adobe. com/products/ illustrator.html).

Figure 1 in Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species

Figure 1. Distribution of mitochondrial lineages of 1,580 grass snakes used in this study. Total sample size of each clade shown in the legend. Eight allochthonous grass snakes with haplotypes of Italian lineages caught in southern Great Britain and Hesse, Germany, not shown. Map was created using ARCGIS 10.2 (http://www.esri. com/arcgis) and ADOBE ILLUSTRATOR CS6 (http://www.adobe.com/products/illustrator.html). Inset: Natrix natrix helvetica (Linz am Rhein, Germany); photo: Wolfgang Böhme.

Figure 2 in Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species

Figure 2. Parsimony networks of mtDNA sequences. Symbol sizes reflect haplotype frequencies. Small black circles are missing node haplotypes; each line connecting two haplotypes corresponds to one mutation step, if not otherwise indicated by numbers. Haplotype colours correspond to lineages, i.e. Natrix natrix helvetica (h) in blue; eastern lineages in yellow (y) and in red (r).

opencc-zeroDec 2014View details →

Data from: Contact zone dynamics during early stages of speciation in a chorus frog (Pseudacris crucifer)

Characterizing the genetic and behavioural consequences of contact between previously geographically isolated lineages provides insights into the mechanisms underlying diversification and ultimately speciation. The spring peeper (Pseudacris crucifer) is a widespread Nearctic chorus frog with six divergent mitochondrial DNA (mtDNA) lineages, many of which came into secondary contact during the Holocene. We examined genetics, morphology, advertisement calls and female preference for two lineages that began diverging in allopatry in the Pliocene and now overlap in southwestern Ontario, Canada. We found non-coincident clines in mtDNA and nuclear DNA, mirroring directionality of premating isolation barriers. We also found divergence in a range of traits between these two lineages, displacement in male call attributes and female preference for calls of their natal lineage in sympatry. Hybrids were morphologically distinct from both parental lineages, but hybrid male calls were acoustically intermediate. Female hybrids showed asymmetrical preference for Eastern male calls. These results considered together provide evidence of either unidirectional hybridization or selection against hybrids, potentially implying reproductive character displacement. Our work demonstrates the utility of integrated, multi-character approaches to understanding the processes of divergence and the nature of speciation.

opencc-zeroDec 2015View details →

Data from: Does competitive interaction drive species recognition in a house mouse secondary contact zone?

Miscommunication may induce a high risk of unnecessary escalated fights between competitors (populations to species), resulting in selection favoring signal divergence through agonistic character displacement (ACD). When signals allowing discrimination between competitors are also involved in mate recognition, ACD could explain reproductive character displacement (RCD). We tested interference competition between males as a potential driver of RCD (here, subspecies recognition) in a secondary contact zone between two mouse subspecies (Mus musculus musculus and Mus musculus domesticus) displaying asymmetric dominance. Since such asymmetry could create a conflict between subspecies (compatibility) and quality (dominance) recognition in the contact zone, we tested for geographic variation in female preference for dominant males in the subordinate subspecies, musculus. We assessed competition between males and tested ACD during dyadic encounters comparing behavior displayed during trials between heterosubspecifics originating from populations close to the secondary contact ("contact") and further away ("allopatric"). We also compared behavior of contact versus allopatric males during homosubspecific versus heterosubspecific trials to test whether subspecies discrimination evolved under competitive interference. Although domesticus dominated most heterosubspecific trials regardless of geographic origin, agonistic behavior was more marked (i.e., lower attack latencies) during contact than allopatric encounters, suggesting that ACD occurred. Comparing behavior during homosubspecific and heterosubspecific encounters, only allopatric musculus displayed differences, that is, higher attack latencies toward heterosubspecifics, indicating that discrimination between competitors did not evolve with ACD. Finally, although allopatric musculus females seemed to prefer dominant males, their contact counterparts did not, suggesting that "compatibility" may have outweighed "quality" under a risk of hybridization.

opencc-zeroDec 2014View details →

Data from: Ecological differentiation, lack of hybrids involving diploids, and asymmetric gene flow between polyploids in narrow contact zones of Senecio carniolicus (syn. Jacobaea carniolica, Asteraceae)

Areas of immediate contact of different cytotypes offer a unique opportunity to study evolutionary dynamics within heteroploid species and to assess isolation mechanisms governing coexistence of cytotypes of different ploidy. The degree of reproductive isolation of cytotypes, i.e., the frequency of heteroploid crosses and subsequent formation of viable and (partly) fertile hybrids, plays a crucial role for the long-term integrity of lineages in contact zones. Here, we assessed fine-scale distribution, spatial clustering and ecological niches as well as patterns of gene-flow in parental and hybrid cytotypes in zones of immediate contact of di-, tetra- and hexaploid Senecio carniolicus (Asteraceae) in the Eastern Alps. Cytotypes were spatially separated also at the investigated micro-scale; the strongest spatial separation was observed for the fully interfertile tetra- and hexaploids. The three main cytotypes showed highly significant niche differences, which were, however, weaker than across their entire distribution ranges in the Eastern Alps. Individuals with intermediate ploidy levels were found neither in the diploid/tetraploid nor in the diploid/hexaploid contact zones indicating strong reproductive barriers. In contrast, pentaploid individuals were frequent in the tetraploid/hexaploid contact zone, albeit limited to a narrow strip in the immediate contact zone of their parental cytotypes. AFLP fingerprinting data revealed introgressive gene flow mediated by pentaploid hybrids from tetra- to hexaploid individuals, but not vice versa. The ecological niche of pentaploids differed significantly from that of tetraploids but not from hexaploids.

opencc-zeroDec 2013View details →

Data from: Haldane's rule revisited: do hybrid females have a shorter lifespan? Survival of hybrids in a recent contact zone between two large gull species

Haldane's rule predicts that particularly high fitness reduction should affect the heterogametic sex of interspecific hybrids. Despite the fact that hybridization is widespread in birds, survival of hybrid individuals is rarely addressed in studies of avian hybrid zones, possibly because of methodological constraints. Here, having applied capture-mark-recapture models to an extensive, 19-year-long dataset on individually marked birds, we estimate annual survival rates of hybrid individuals in the hybrid zone between herring (Larus argentatus) and Caspian (L. cachinnans) gulls. In both parental species, males have a slightly higher survival rate than females (model-weighted mean ± SE: herring gull males 0.88 ± 0.01, females 0.87 ± 0.01, Caspian gull males 0.88 ± 0.01, females 0.87 ± 0.01). Hybrid males do not survive for a shorter time than non-hybrid ones (0.88 ± 0.01), whereas hybrid females have the lowest survival rate among all groups of individuals (0.83 ± 0.03). This translates to a shorter adult (reproductive) lifespan (on average by 1.7-1.8 years, i.e. ca 25%) compared to non-hybrid females. We conclude that, in line with Haldane's rule, the lower survival rate of female hybrids may contribute to selection against hybrids in this hybrid zone.

opencc-zeroDec 2017View details →

Data from: Gene flow in the European coal tit, Periparus ater (Aves: Passeriformes): low among Mediterranean populations but high in a continental contact zone

Extant phylogeographic patterns of Palearctic terrestrial vertebrates are generally believed to have originated from glacial range fragmentation. Post-Pleistocene range expansions have led to the formation of secondary contact zones among genetically distinct taxa. For coal tits (Periparus ater), such a contact zone has been localized in Germany. In this study, we quantified gene flow between Fennoscandian and southern European coal tits using a set of 13 microsatellite loci. STRUCTURE analysis revealed four genetic clusters two of these on Mediterranean islands. German populations were genetically admixed but introgression of southern alleles was evident for Fennoscandian populations. In the South, we found negligible introgression of northern alleles (and haplotypes) but slight admixture of two southern genetic clusters in the Pyrenees and on the Balkan Peninsulae and near complete sorting of these two allelic lineages on the islands of Corsica and Sardinia. Genetic distinctiveness of the Mediterranean island populations reflects general patterns of endemism in the Corso-Sardinian fauna and the Cypriot fauna. Wide-range gene flow in Central Europe suggests a broad zone of intergradation between subspecies of the coal tit rather than a narrow contact zone. This is in accordance with low morphological and bioacoustic differentiation of European coal tit populations.

opencc-zeroDec 2010View details →

Data from: Strong selection against hybrids maintains a narrow contact zone between morphologically cryptic lineages in a rainforest lizard

Phenotypically cryptic lineages comprise an important yet understudied part of biodiversity; in particular, we have much to learn about how these lineages are formed and maintained. To better understand the evolutionary significance of such lineages, we studied a hybrid zone between two morphologically-cryptic phylogeographic lineages in the rainforest lizard, Lampropholis coggeri. Analyzing a multilocus genetic dataset through cline inference, individual-based methods and population measures of disequilibrium and using simulations to explore our genetic results in context of theoretical expectations, we inferred the processes maintaining this hybrid zone. We find that these lineages meet in a hybrid zone that is narrow (~400 m) relative to inferred dispersal rate. Further, the hybrid zone exhibits substantial genetic disequilibrium and sharply coincident and largely concordant clines. Based on our knowledge about the region's biogeography, the species' natural history, and our simulation results, we suggest that strong selection against hybrids structures this system. As all clines show a relatively narrow range of introgression, we posit that this hybrid zone might not yet be in equilibrium. Nonetheless, our results clearly show that phylogeographic lineages can evolve substantial reproductive isolation without concomitant morphological diversification, suggesting that such lineages can constitute a significant component of evolutionary diversity.

opencc-zeroDec 2013View details →

Data from: Individual performance in relation to cytonuclear discordance in a northern contact zone between long-toed salamander (Ambystoma macrodactylum) lineages

Cytonuclear discordance in contact zones between related lineages is common, with mitochondrial clines often being displaced from clines in nuclear allele frequency. Proposed explanations for such a pattern include adaptive introgression of mtDNA or a neutral wake of mtDNA being left behind following hybrid zone movement. However, studies investigating these hypotheses are rare. Our previous survey of genetic variation in the long-toed salamander (Ambystoma macrodactylum) highlighted a potential case of cytonuclear discordance between two lineages in western Canada. Here, we use additional markers and samples to clarify the extent of this discordance. We simultaneously assess the feeding performance of individuals in a common environment to test for an association between mitotype and individual performance. The genetic results confirm a general pattern of cytonuclear discordance in the focal region. However, we also observed more limited introgression of a diagnostic nuclear marker. Intriguingly, although there were differences in individual performance associated with the transition between mitotypes, these differences were not fully explained by mitotype. Instead, the lowest performance was observed in individuals demonstrating the greatest mismatch between mtDNA and all nuclear markers, suggesting the potential for cytonuclear incompatibilities to be acting. These results highlight the complexity of understanding the causes and consequences of mtDNA introgression and cytonuclear discordance in contact zones.