@article{11334,
  abstract     = {Hybridization is a common evolutionary process with multiple possible outcomes. In vertebrates, interspecific hybridization has repeatedly generated parthenogenetic hybrid species. However, it is unknown whether the generation of parthenogenetic hybrids is a rare outcome of frequent hybridization between sexual species within a genus or the typical outcome of rare hybridization events. Darevskia is a genus of rock lizards with both hybrid parthenogenetic and sexual species. Using capture sequencing, we estimate phylogenetic relationships and gene flow among the sexual species, to determine how introgressive hybridization relates to the origins of parthenogenetic hybrids. We find evidence for widespread hybridization with gene flow, both between recently diverged species and deep branches. Surprisingly, we find no signal of gene flow between parental species of the parthenogenetic hybrids, suggesting that the parental pairs were either reproductively or geographically isolated early in their divergence. The generation of parthenogenetic hybrids in Darevskia is, then, a rare outcome of the total occurrence of hybridization within the genus, but the typical outcome when specific species pairs hybridize. Our results question the conventional view that parthenogenetic lineages are generated by hybridization in a window of divergence. Instead, they suggest that some lineages possess specific properties that underpin successful parthenogenetic reproduction.},
  author       = {Freitas, Susana and Westram, Anja M and Schwander, Tanja and Arakelyan, Marine and Ilgaz, Çetin and Kumlutas, Yusuf and Harris, David James and Carretero, Miguel A. and Butlin, Roger K.},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {5},
  pages        = {899--914},
  publisher    = {Wiley},
  title        = {{Parthenogenesis in Darevskia lizards: A rare outcome of common hybridization, not a common outcome of rare hybridization}},
  doi          = {10.1111/evo.14462},
  volume       = {76},
  year         = {2022},
}

@article{12234,
  abstract     = {Hybrid speciation—the origin of new species resulting from the hybridization of genetically divergent lineages—was once considered rare, but genomic data suggest that it may occur more often than once thought. In this study, Noguerales and Ortego found genomic evidence supporting the hybrid origin of a grasshopper that is able to exploit a broader range of host plants than either of its putative parents.},
  author       = {Stankowski, Sean},
  issn         = {1558-5646},
  journal      = {Evolution},
  keywords     = {General Agricultural and Biological Sciences, Genetics, Ecology, Evolution, Behavior and Systematics},
  number       = {11},
  pages        = {2784--2785},
  publisher    = {Wiley},
  title        = {{Digest: On the origin of a possible hybrid species}},
  doi          = {10.1111/evo.14632},
  volume       = {76},
  year         = {2022},
}

@article{12247,
  abstract     = {Chromosomal inversions have been shown to play a major role in a local adaptation by suppressing recombination between alternative arrangements and maintaining beneficial allele combinations. However, so far, their importance relative to the remaining genome remains largely unknown. Understanding the genetic architecture of adaptation requires better estimates of how loci of different effect sizes contribute to phenotypic variation. Here, we used three Swedish islands where the marine snail Littorina saxatilis has repeatedly evolved into two distinct ecotypes along a habitat transition. We estimated the contribution of inversion polymorphisms to phenotypic divergence while controlling for polygenic effects in the remaining genome using a quantitative genetics framework. We confirmed the importance of inversions but showed that contributions of loci outside inversions are of similar magnitude, with variable proportions dependent on the trait and the population. Some inversions showed consistent effects across all sites, whereas others exhibited site-specific effects, indicating that the genomic basis for replicated phenotypic divergence is only partly shared. The contributions of sexual dimorphism as well as environmental factors to phenotypic variation were significant but minor compared to inversions and polygenic background. Overall, this integrated approach provides insight into the multiple mechanisms contributing to parallel phenotypic divergence.},
  author       = {Koch, Eva L. and Ravinet, Mark and Westram, Anja M and Johannesson, Kerstin and Butlin, Roger K.},
  issn         = {1558-5646},
  journal      = {Evolution},
  keywords     = {General Agricultural and Biological Sciences, Genetics, Ecology, Evolution, Behavior and Systematics},
  number       = {10},
  pages        = {2332--2346},
  publisher    = {Wiley},
  title        = {{Genetic architecture of repeated phenotypic divergence in Littorina saxatilis evolution}},
  doi          = {10.1111/evo.14602},
  volume       = {76},
  year         = {2022},
}

@article{8743,
  abstract     = {Montane cloud forests are areas of high endemism, and are one of the more vulnerable terrestrial ecosystems to climate change. Thus, understanding how they both contribute to the generation of biodiversity, and will respond to ongoing climate change, are important and related challenges. The widely accepted model for montane cloud forest dynamics involves upslope forcing of their range limits with global climate warming. However, limited climate data provides some support for an alternative model, where range limits are forced downslope with climate warming. Testing between these two models is challenging, due to the inherent limitations of climate and pollen records. We overcome this with an alternative source of historical information, testing between competing model predictions using genomic data and demographic analyses for a species of beetle tightly associated to an oceanic island cloud forest. Results unequivocally support the alternative model: populations that were isolated at higher elevation peaks during the Last Glacial Maximum are now in contact and hybridizing at lower elevations. Our results suggest that genomic data are a rich source of information to further understand how montane cloud forest biodiversity originates, and how it is likely to be impacted by ongoing climate change.},
  author       = {Salces-Castellano, Antonia and Stankowski, Sean and Arribas, Paula and Patino, Jairo and Karger, Dirk N.  and Butlin, Roger and Emerson, Brent C.},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {2},
  pages        = {231--244},
  publisher    = {Wiley},
  title        = {{Long-term cloud forest response to climate warming revealed by insect speciation history}},
  doi          = {10.1111/evo.14111},
  volume       = {75},
  year         = {2021},
}

@article{9252,
  abstract     = {This paper analyses the conditions for local adaptation in a metapopulation with infinitely many islands under a model of hard selection, where population size depends on local fitness. Each island belongs to one of two distinct ecological niches or habitats. Fitness is influenced by an additive trait which is under habitat‐dependent directional selection. Our analysis is based on the diffusion approximation and accounts for both genetic drift and demographic stochasticity. By neglecting linkage disequilibria, it yields the joint distribution of allele frequencies and population size on each island. We find that under hard selection, the conditions for local adaptation in a rare habitat are more restrictive for more polygenic traits: even moderate migration load per locus at very many loci is sufficient for population sizes to decline. This further reduces the efficacy of selection at individual loci due to increased drift and because smaller populations are more prone to swamping due to migration, causing a positive feedback between increasing maladaptation and declining population sizes. Our analysis also highlights the importance of demographic stochasticity, which exacerbates the decline in numbers of maladapted populations, leading to population collapse in the rare habitat at significantly lower migration than predicted by deterministic arguments.},
  author       = {Szep, Eniko and Sachdeva, Himani and Barton, Nicholas H},
  issn         = {1558-5646},
  journal      = {Evolution},
  keywords     = {Genetics, Ecology, Evolution, Behavior and Systematics, General Agricultural and Biological Sciences},
  number       = {5},
  pages        = {1030--1045},
  publisher    = {Wiley},
  title        = {{Polygenic local adaptation in metapopulations: A stochastic eco‐evolutionary model}},
  doi          = {10.1111/evo.14210},
  volume       = {75},
  year         = {2021},
}

@article{9374,
  abstract     = {If there are no constraints on the process of speciation, then the number of species might be expected to match the number of available niches and this number might be indefinitely large. One possible constraint is the opportunity for allopatric divergence. In 1981, Felsenstein used a simple and elegant model to ask if there might also be genetic constraints. He showed that progress towards speciation could be described by the build‐up of linkage disequilibrium among divergently selected loci and between these loci and those contributing to other forms of reproductive isolation. Therefore, speciation is opposed by recombination, because it tends to break down linkage disequilibria. Felsenstein then introduced a crucial distinction between “two‐allele” models, which are subject to this effect, and “one‐allele” models, which are free from the recombination constraint. These fundamentally important insights have been the foundation for both empirical and theoretical studies of speciation ever since.},
  author       = {Butlin, Roger K. and Servedio, Maria R. and Smadja, Carole M. and Bank, Claudia and Barton, Nicholas H and Flaxman, Samuel M. and Giraud, Tatiana and Hopkins, Robin and Larson, Erica L. and Maan, Martine E. and Meier, Joana and Merrill, Richard and Noor, Mohamed A. F. and Ortiz‐Barrientos, Daniel and Qvarnström, Anna},
  issn         = {1558-5646},
  journal      = {Evolution},
  keywords     = {Genetics, Ecology, Evolution, Behavior and Systematics, General Agricultural and Biological Sciences},
  number       = {5},
  pages        = {978--988},
  publisher    = {Wiley},
  title        = {{Homage to Felsenstein 1981, or why are there so few/many species?}},
  doi          = {10.1111/evo.14235},
  volume       = {75},
  year         = {2021},
}

@article{9383,
  abstract     = {A primary roadblock to our understanding of speciation is that it usually occurs over a timeframe that is too long to study from start to finish. The idea of a speciation continuum provides something of a solution to this problem; rather than observing the entire process, we can simply reconstruct it from the multitude of speciation events that surround us. But what do we really mean when we talk about the speciation continuum, and can it really help us understand speciation? We explored these questions using a literature review and online survey of speciation researchers. Although most researchers were familiar with the concept and thought it was useful, our survey revealed extensive disagreement about what the speciation continuum actually tells us. This is due partly to the lack of a clear definition. Here, we provide an explicit definition that is compatible with the Biological Species Concept. That is, the speciation continuum is a continuum of reproductive isolation. After outlining the logic of the definition in light of alternatives, we explain why attempts to reconstruct the speciation process from present‐day populations will ultimately fail. We then outline how we think the speciation continuum concept can continue to act as a foundation for understanding the continuum of reproductive isolation that surrounds us.},
  author       = {Stankowski, Sean and Ravinet, Mark},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {6},
  pages        = {1256--1273},
  publisher    = {Oxford University Press},
  title        = {{Defining the speciation continuum}},
  doi          = {10.1111/evo.14215},
  volume       = {75},
  year         = {2021},
}

@article{6680,
  abstract     = {This paper analyzes how partial selfing in a large source population influences its ability to colonize a new habitat via the introduction of a few founder individuals. Founders experience inbreeding depression due to partially recessive deleterious alleles as well as maladaptation to the new environment due to selection on a large number of additive loci. I first introduce a simplified version of the Inbreeding History Model (Kelly, 2007) in order to characterize mutation‐selection balance in a large, partially selfing source population under selection involving multiple non‐identical loci. I then use individual‐based simulations to study the eco‐evolutionary dynamics of founders establishing in the new habitat under a model of hard selection. The study explores how selfing rate shapes establishment probabilities of founders via effects on both inbreeding depression and adaptability to the new environment, and also distinguishes the effects of selfing on the initial fitness of founders from its effects on the long‐term adaptive response of the populations they found. A high rate of (but not complete) selfing is found to aid establishment over a wide range of parameters, even in the absence of mate limitation. The sensitivity of the results to assumptions about the nature of polygenic selection are discussed.},
  author       = {Sachdeva, Himani},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {9},
  pages        = {1729--1745},
  publisher    = {Wiley},
  title        = {{Effect of partial selfing and polygenic selection on establishment in a new habitat}},
  doi          = {10.1111/evo.13812},
  volume       = {73},
  year         = {2019},
}

@article{1851,
  abstract     = {We consider mating strategies for females who search for males sequentially during a season of limited length. We show that the best strategy rejects a given male type if encountered before a time-threshold but accepts him after. For frequency-independent benefits, we obtain the optimal time-thresholds explicitly for both discrete and continuous distributions of males, and allow for mistakes being made in assessing the correct male type. When the benefits are indirect (genes for the offspring) and the population is under frequency-dependent ecological selection, the benefits depend on the mating strategy of other females as well. This case is particularly relevant to speciation models that seek to explore the stability of reproductive isolation by assortative mating under frequency-dependent ecological selection. We show that the indirect benefits are to be quantified by the reproductive values of couples, and describe how the evolutionarily stable time-thresholds can be found. We conclude with an example based on the Levene model, in which we analyze the evolutionarily stable assortative mating strategies and the strength of reproductive isolation provided by them.},
  author       = {Priklopil, Tadeas and Kisdi, Eva and Gyllenberg, Mats},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {4},
  pages        = {1015 -- 1026},
  publisher    = {Wiley},
  title        = {{Evolutionarily stable mating decisions for sequentially searching females and the stability of reproductive isolation by assortative mating}},
  doi          = {10.1111/evo.12618},
  volume       = {69},
  year         = {2015},
}

@article{9931,
  abstract     = {Gene duplication is important in evolution, because it provides new raw material for evolutionary adaptations. Several existing hypotheses about the causes of duplicate retention and diversification differ in their emphasis on gene dosage, subfunctionalization, and neofunctionalization. Little experimental data exist on the relative importance of gene expression changes and changes in coding regions for the evolution of duplicate genes. Furthermore, we do not know how strongly the environment could affect this importance. To address these questions, we performed evolution experiments with the TEM-1 beta lactamase gene in Escherichia coli to study the initial stages of duplicate gene evolution in the laboratory. We mimicked tandem duplication by inserting two copies of the TEM-1 gene on the same plasmid. We then subjected these copies to repeated cycles of mutagenesis and selection in various environments that contained antibiotics in different combinations and concentrations. Our experiments showed that gene dosage is the most important factor in the initial stages of duplicate gene evolution, and overshadows the importance of point mutations in the coding region.},
  author       = {Dhar, Riddhiman and Bergmiller, Tobias and Wagner, Andreas},
  issn         = {1558-5646},
  journal      = {Evolution},
  number       = {6},
  pages        = {1775--1791},
  publisher    = {Wiley},
  title        = {{Increased gene dosage plays a predominant role in the initial stages of evolution of duplicate TEM-1 beta lactamase genes}},
  doi          = {10.1111/evo.12373},
  volume       = {68},
  year         = {2014},
}

@article{7781,
  abstract     = {Males are predicted to compete for reproductive opportunities, with sexual selection driving the evolution of large body size and weaponry through the advantage they confer for access to females. Few studies have explored potential trade‐offs of investment in secondary sexual traits between different components of fitness or tested for sexually antagonistic selection pressures. These factors may provide explanations for observed polymorphisms in both form and quality of secondary sexual traits. We report here an analysis of selection on horn phenotype in a feral population of Soay sheep (Ovis aries) on the island of Hirta, St. Kilda, Scotland. Soay sheep display a phenotypic polymorphism for horn type with males growing either normal or reduced (scurred) horns, and females growing either normal, scurred, or no (polled) horns; further variation in size exists within horn morphs. We show that horn phenotype and the size of the trait displayed is subject to different selection pressures in males and females, generating sexually antagonistic selection. Furthermore, there was evidence of a trade‐off between breeding success and longevity in normal‐horned males, with both the normal horn type and larger horn size being associated with greater annual breeding success but reduced longevity. Therefore, selection through lifetime breeding success was not found to act upon horn phenotype in males. In females, a negative association of annual breeding success within the normal‐horned phenotype did not result in a significant difference in lifetime fitness when compared to scurred individuals, as no significant difference in longevity was found. However, increased horn size within this group was negatively associated with breeding success and longevity. Females without horns (polled) suffered reduced longevity and thus reduced lifetime breeding success relative the other horn morphs. Our results therefore suggest that trade‐offs between different components of fitness and antagonistic selection between the sexes may maintain genetic variation for secondary sexual traits within a population.},
  author       = {Robinson, Matthew Richard and Pilkington, Jill G. and Clutton-Brock, Tim H. and Pemberton, Josephine M. and Kruuk, Loeske E.B.},
  issn         = {0014-3820},
  journal      = {Evolution},
  number       = {10},
  pages        = {2168--2181},
  publisher    = {Wiley},
  title        = {{Live fast, die young: Trade-offs between fitness components and sexually antagonistic selection on weaponry in soay sheep}},
  doi          = {10.1111/j.0014-3820.2006.tb01854.x},
  volume       = {60},
  year         = {2007},
}

@article{4254,
  abstract     = {Chromosomal rearrangements can promote reproductive isolation by reducing recombination along a large section of the genome. We model the effects of the genetic barrier to gene flow caused by a chromosomal rearrangement on the rate of accumulation of postzygotic isolation genes in parapatry. We find that, if reproductive isolation is produced by the accumulation in parapatry of sets of alleles compatible within but incompatible across species, chromosomal rearrangements are far more likely to favor it than classical genetic barriers without chromosomal changes. New evidence of the role of chromosomal rearrangements in parapatric speciation suggests that postzygotic isolation is often due to the accumulation of such incompatibilities. The model makes testable qualitative predictions about the genetic signature of speciation.},
  author       = {Navarro, Arcadio and Barton, Nicholas H},
  issn         = {0014-3820},
  journal      = {Evolution; International Journal of Organic Evolution},
  number       = {3},
  pages        = {447 -- 459},
  publisher    = {Wiley-Blackwell},
  title        = {{Accumulating postzygotic isolation genes in parapatry: a new twist on chromosomal speciation}},
  doi          = {10.1111/j.0014-3820.2003.tb01537.x},
  volume       = {57},
  year         = {2003},
}

@article{4338,
  abstract     = {Mosaic hybrid zones arise when ecologically differentiated taxa hybridize across a network of habitat patches. Frequent interbreeding across a small-scale patchwork can erode species differences that might have been preserved in a clinal hybrid zone. In particular, the rapid breakdown of neutral divergence sets an upper limit to the time for which differences at marker loci can persist. We present here a case study of a mosaic hybrid zone between the fire-bellied toads Bombina bombina and B. variegata (Anura: Discoglossidae) near Apahida in Romania. In our 20 × 20 km study area, we detected no evidence of a clinal transition but found a strong association between aquatic habitat and mean allele frequencies at four molecular markers. In particular, pure populations of B. bombina in ponds appear to cause massive introgression into the surrounding B. variegata gene pool found in temporary aquatic sites. Nevertheless, the genetic structure of these hybrid populations was remarkably similar to those of a previously studied transect near Pescenica (Croatia), which had both clinal and mosaic features: estimates of heterozygote deficit and linkage disequilibrium in each country are similar. In Apahida, the observed strong linkage disequilibria should stem from an imperfect habitat preference that guides most (but not all) adults into the habitats to which they are adapted. In the absence of a clinal structure, the inferred migration rate between habitats implies that associations between selected loci and neutral markers should break down rapidly. Although plausible selection strengths can maintain differentiation at those loci adapting the toads to either permanent or temporary breeding sites, the divergence at neutral markers must be transient. The hybrid zone may be approaching a state in which the gene pools are homogenized at all but the selected loci, not dissimilar from an early stage of sympatric divergence.},
  author       = {Vines, Timothy and Kohler, S C and Thiel, M and Ghira, Ioan and Sands, T R and Maccallum, Catriona and Barton, Nicholas H and Nürnberger, Beate},
  issn         = {0014-3820},
  journal      = {Evolution},
  number       = {8},
  pages        = {1876 -- 1888},
  publisher    = {Wiley-Blackwell},
  title        = {{On the maintenance of reproductive isolation in a mosaic hybrid zone between the toads Bombina bombina and B. variegata}},
  doi          = {10.1111/j.0014-3820.2003.tb00595.x},
  volume       = {57},
  year         = {2003},
}

@article{3622,
  abstract     = {The extent of genetic variation in fitness and its components and genetic variation's dependence on environmental conditions remain key issues in evolutionary biology. We present measurements of genetic variation in preadult viability in a laboratory-adapted population of Drosophila melanogaster, made at four different densities. By crossing flies heterozygous for a wild-type chromosome and one of two different balancers (TM1, TM2), we measure both heterozygous (TM1/+, TM2/+) and homozygous (+/+) viability relative to a standard genotype (TM1/TM2). Forty wild-type chromosomes were tested, of which 10 were chosen to be homozygous viable. The mean numbers produced varied significantly between chromosome lines, with an estimated between-line variance in loge numbers of 0.013. Relative viabilities also varied significantly across chromosome lines, with a variance in loge homozygous viability of 1.76 and of loge heterozygous viability of 0.165. The between-line variance for numbers emerging increased with density, from 0.009 at lowest density to 0.079 at highest. The genetic variance in relative viability increases with density, but not significantly. Overall, the effects of different chromosomes on relative viability were remarkably consistent across densities and across the two heterozygous genotypes (TM1, TM2). The 10 lines that carried homozygous viable wild-type chromosomes produced significantly more adults than the 30 lethal lines at low density and significantly fewer adults at the highest density. Similarly, there was a positive correlation between heterozygous viability and mean numbers at low density, but a negative correlation at high density.},
  author       = {Gardner, Michael and Fowler, Kevin and Patridge, Linda and Barton, Nicholas H},
  issn         = {0014-3820},
  journal      = {Evolution},
  number       = {8},
  pages        = {1609 -- 1620},
  publisher    = {Wiley-Blackwell},
  title        = {{Genetic variation for preadult viability in Drosophila melanogaster}},
  doi          = {10.1111/j.0014-3820.2001.tb00680.x},
  volume       = {55},
  year         = {2001},
}

@article{4265,
  abstract     = {The reasons that sex and recombination are so widespread remain elusive. One popular hypothesis is that sex and recombination promote adaptation to a changing environment. The strongest evidence that increased recombination may evolve because recombination promotes adaptation comes from artificially selected populations. Recombination rates have been found to increase as a correlated response to selection on traits unrelated to recombination in several artificial selection experiments and in a comparison of domesticated and nondomesticated mammals. There are, however, several alternative explanations for the increase in recombination in such populations, including two different evolutionary explanations. The first is that the form of selection is epistatic, generating linkage disequilibria among selected loci, which can indirectly favor modifier alleles that increase recombination. The second is that random genetic drift in selected populations tends to generate disequilibria such that beneficial alleles are often found in different individuals; modifier alleles that increase recombination can bring together such favorable alleles and thus may be found in individuals with greater fitness. In this paper, we compare the evolutionary forces acting on recombination in finite populations subject to strong selection, To our surprise, we found that drift accounted for the majority of selection for increased recombination observed in simulations of small to moderately large populations, suggesting that, unless selected populations are large, epistasis plays a secondary role in the evolution of recombination.},
  author       = {Otto, Sarah and Barton, Nicholas H},
  issn         = {0014-3820},
  journal      = {Evolution; International Journal of Organic Evolution},
  number       = {10},
  pages        = {1921 -- 1931},
  publisher    = {Wiley-Blackwell},
  title        = {{Selection for recombination in small populations}},
  doi          = {10.1111/j.0014-3820.2001.tb01310.x},
  volume       = {55},
  year         = {2001},
}

@article{4269,
  author       = {Coyne, Jerry and Barton, Nicholas H and Turelli, Michael},
  issn         = {0014-3820},
  journal      = {Evolution; International Journal of Organic Evolution},
  number       = {1},
  pages        = {306 -- 317},
  publisher    = {Wiley-Blackwell},
  title        = {{Is Wright’s shifting balance process important in evolution?}},
  doi          = {10.1554/0014-3820(2000)054[0306:IWSSBP]2.0.CO;2},
  volume       = {54},
  year         = {2000},
}

@article{4277,
  abstract     = {Reproductive isolation between two taxa may be due to endogenous selection, which is generated by incompatibilities between the respective genomes, to exogenous selection, which is generated by differential adaptations to alternative environments, or to both. The continuing debate over the relative importance of either mode of selection has highlighted the need for unambiguous data on the fitness of hybrid genotypes. The hybrid zone between the fire-bellied toad (Bombina bombina) and the yellow-bellied toad (B. variegata) in central Europe involves adaptation to different environments, but evidence of hybrid dysfunction is equivocal. In this study, we followed the development under laboratory conditions of naturally laid eggs collected from a transect across the Bombina hybrid zone in Croatia. Fitness was significantly reduced in hybrid populations: Egg batches from the center of the hybrid zone showed significantly higher embryonic and larval mortality and higher frequencies of morphological abnormalities relative to either parental type. Overall mortality from day of egg collection to three weeks after hatching reached 20% in central hybrid populations, compared to 2% in pure populations. There was no significant difference in fitness between two parental types. Within hybrid populations, there was considerable variation in fitness, with some genotypes showing no evidence of reduced viability. We discuss the implications of these findings for our understanding of barriers to gene flow between species.},
  author       = {Kruuk, Loeske and Gilchrist, Jason and Barton, Nicholas H},
  issn         = {0014-3820},
  journal      = {Evolution; International Journal of Organic Evolution},
  number       = {5},
  pages        = {1611 -- 1616},
  publisher    = {Wiley-Blackwell},
  title        = {{Hybrid dysfunction in fire-bellied toads (Bombina)}},
  doi          = {10.2307/2640907},
  volume       = {53},
  year         = {1999},
}

@article{3629,
  abstract     = {This paper demonstrates the effect of habitat heterogeneity and a habitat preference on the genetic structure of a hybrid zone between the toads Bombina bombina and B. variegata (Anura: Discoglossidae); 1613 toads from 85 sites across a transect near Pešćenica, Croatia, were scored for five unlinked diagnostic allozyme markers. These were found to be largely concordant. Aside from minor systematic deviations, there was little variance in allele frequency among loci within sites. Yet the allele frequencies did not follow a smooth cline, but formed a mosaic in the center, such that neighboring sites could differ markedly in their enzyme score. A detailed ecological survey revealed a correlation between this pattern and habitat. In keeping with the typical breeding sites of the parental taxa, B. bombina-like hybrids were found more often in ponds, whereas B. variegata-like hybrids were more common in puddles. In addition, there was significant heterozygote deficit (FIS) and strong linkage disequilibrium (R), both of which were stronger on the B. bombina side of the transect, and stronger in puddles than ponds. Mark-recapture data showed: (1) that the animals disperse beyond the scale of the habitat pattern; (2) frequent turn-over of individuals within sites; and (3) nonrandom movement between two sites of different habitat type. We conclude that an active habitat preference must contribute to the observed association between marker alleles and habitat. As a consequence, there is incomplete mixing of the two gene pools, which could explain the high level of FIS and R. The asymmetry in these parameters may reflect asymmetry in the preference or in the distribution of habitats across the zone. We discuss the implications of habitat preference for the dynamics of hybrid zones.},
  author       = {Maccallum, Catriona and Nürnberger, Beate and Barton, Nicholas H and Szymura, Jacek},
  issn         = {0014-3820},
  journal      = {Evolution},
  number       = {1},
  pages        = {227 -- 239},
  publisher    = {Wiley-Blackwell},
  title        = {{Habitat preference in the Bombina hybrid zone in Croatia}},
  doi          = {10.1111/j.1558-5646.1998.tb05156.x},
  volume       = {52},
  year         = {1998},
}

@article{4287,
  abstract     = {We evaluate Sewall Wright's three-phase "shifting balance" theory of evolution, examining both the theoretical issues and the relevant data from nature and the laboratory. We conclude that while phases I and II of Wright's theory (the movement of populations from one "adaptive peak" to another via drift and selection) can occur under some conditions, genetic drift is often unnecessary for movement between peaks. Phase III of the shifting balance, in which adaptations spread from particular populations to the entire species, faces two major theoretical obstacles: (1) unlike adaptations favored by simple directional selection, adaptations whose fixation requires some genetic drift are often prevented from spreading by barriers to gene flow; and (2) it is difficult to assemble complex adaptations whose constituent parts arise via peak shifts in different demes. Our review of the data from nature shows that although there is some evidence for individual phases of the shifting balance process, there are few empirical observations explained better by Wright's three-phase mechanism than by simple mass selection. Similarly, artificial selection experiments fail to show that selection in subdivided populations produces greater response than does mass selection in large populations. The complexity of the shifting balance process and the difficulty of establishing that adaptive valleys have been crossed by genetic drift make it impossible to test Wright's claim that adaptations commonly originate by this process. In view of these problems, it seems unreasonable to consider the shifting balance process as an important explanation for the evolution of adaptations. },
  author       = {Coyne, Jerry and Barton, Nicholas H and Turelli, Michael},
  issn         = {0014-3820},
  journal      = {Evolution; International Journal of Organic Evolution},
  number       = {3},
  pages        = {643 -- 671},
  publisher    = {Wiley-Blackwell},
  title        = {{Perspective: A critique of Sewall Wright's shifting balance theory of evolutionight's shifting balance theory of evolution}},
  doi          = {10.1111/j.1558-5646.1997.tb03650.x},
  volume       = {51},
  year         = {1997},
}

@article{3636,
  abstract     = {Observations on the means, variances, and covariances of quantitative traits across hybrid zones can give information similar to that from Mendelian markers. In addition, they can identify particular traits through which the cline is maintained. We describe a survey of six traits across the hybrid zone between Bombina bombina and Bombina variegata (Amphibia: Discoglossidae) near Pescenica in Croatia. We obtained laboratory measuments of the belly pattern, skin thickness, mating call, skeletal form, egg size, and the developmental time of tadpoles. Although offspring from hybrid populations showed no evidence of reduced viability, a third of the F1 families failed completely, irrespective of the direction of the cross. All traits differed significantly between the taxa. Clines in belly pattern, skin thickness, mating call, and skeletal form were closely concordant with clines in four diagnostic enzyme loci. However, the cline in developmental time was displaced towards bombina, and the cline in egg size was displaced towards variegata. This discordance could be because the traits are not inherited additively or because they are subject to different selection pressures. We favor the latter explanation in the case of developmental time. We show that moderate selection acting directly on a trait suffices to shift its position; rather stronger selection is needed to change its width appreciably. Within hybrid populations, there are significant associations among quantitative traits, and between traits and enzymes. Phenotypic variances also increase in hybrid populations. These observations can be explained by linkage disequilibria among the underlying loci. However, the average magnitude of the covariance between traits is about half that expected from the linkage disequilibria between enzyme loci. The discrepancy is not readily explained by nonadditive gene action. This puzzle is now unresolved and calls for further investigation.},
  author       = {Nürnberger, Beate and Barton, Nicholas H and Maccallum, Catriona and Gilchrist, Jason and Appleby, Michael},
  issn         = {0014-3820},
  journal      = {Evolution},
  number       = {6},
  pages        = {1224 -- 1238},
  publisher    = {Wiley-Blackwell},
  title        = {{Natural selection on quantitative traits in the Bombina hybrid zone}},
  doi          = {10.1111/j.1558-5646.1995.tb04449.x},
  volume       = {49},
  year         = {1995},
}

