Geographic variation in hybridization across a reinforcement contact zone of chorus frogs (Pseudacris)

Reinforcement contact zones, which are secondary contact zones where species are diverging in reproductive behaviors due to selection against hybridization, represent natural laboratories for studying speciation‐in‐action. Here, we examined replicate localities across the entire reinforcement contact zone between North American chorus frogs Pseudacris feriarum and P. nigrita to investigate geographic variation in hybridization frequencies and to assess whether reinforcement may have contributed to increased genetic divergence within species. Previous work indicated these species have undergone reproductive character displacement (RCD) in male acoustic signals and female preferences due to reinforcement. We also examined acoustic signal variation across the contact zone to assess whether signal characteristics reliably predict hybrid index and to elucidate whether the degree of RCD predicts hybridization rate. Using microsatellites, mitochondrial sequences, and acoustic signal information from >1,000 individuals across >50 localities and ten sympatric focal regions, we demonstrate: (1) hybridization occurs and (2) varies substantially across the geographic range of the contact zone, (3) hybridization is asymmetric and in the direction predicted from observed patterns of asymmetric RCD, (4) in one species, genetic distance is higher between conspecific localities where one or both have been reinforced than between nonreinforced localities, after controlling for geographic distance, (5) acoustic signal characters strongly predict hybrid index, and (6) the degree of RCD does not strongly predict admixture levels. By showing that hybridization occurs in all sympatric localities, this study provides the fifth and final line of evidence that reproductive character displacement is due to reinforcement in the chorus frog contact zone. Furthermore, this work suggests that the dual action of cascade reinforcement and partial geographic isolation is promoting genetic diversification within one of the reinforced species.     

Phylogeography of the land snail Albinaria hippolyti (Pulmonata: Clausiliidae) from Crete, inferred from ITS-1 sequences

The polytypic Cretan land snail Albinaria hippolyti has a range that is partly fragmented and partly subdivided by hybrid zones. For this reason, it has served as a model species for investigating speciation and radiation in Mediterranean Clausiliidae. The first internal transcribed spacer ( ITS-1 ) of the nuclear ribosomal DNA was sequenced in 20 populations of A. hippolyti and phylogenetically analysed using maximum parsimony. We employed a novel method involving logarithmic weighting of gaps and topological constraints based on bootstrap values. The resulting phylogeography suggests that the species has undergone a recent cycle of range expansion and range reduction. Speciation cannot be linked to major geological vicariance events in the Miocene and Pliocene, as has been suggested previously. The subspecies A. h. arthuriana appears unrelated to other A. hippolyti subspecies, which supports recent suggestions, based on morphology, to regard it as a separate species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 317–326.     

The 'rare allele phenomenon' in a ribosomal spacer

We describe the increased frequency of a particular length variant of the internal transcribed spacer 1 (ITS-1) of the ribosomal DNA in a hybrid zone of the land snail Albinaria hippolyti. The phenomenon that normally rare alleles or other markers can increase in frequency in the centre of hybrid zones is not new. Under the term 'hybrizyme' or 'rare allele' phenomenon it has been recorded in many organisms and different genetic markers. However, this is the first time that it has been found in a multicopy locus. On the one hand, the pattern fits well with the view that purifying selection in hybrid populations works on many loci across the genome and should thus have its effect on many independent molecular markers. On the other hand, the results are puzzling, given that the multiple copies of rDNA are not expected to respond in unison. We suggest two possible explanations for these conflicting observations.     

RANDOM MATING IN THE NORTHERN FLICKER HYBRID ZONE: IMPLICATIONS FOR THE EVOLUTION OF BRIGHT AND CONTRASTING PLUMAGE PATTERNS IN BIRDS

The Red-shafted and Yellow-shafted Flickers hybridize in a narrow zone on the western Great Plains of North America. The two subspecies are markedly different in six plumage traits. Plumage phenotypes were scored for the male and female of 125 mated pairs from the hybrid zone. Correlation analyses revealed no tendency towards pairing of similar phenotypes. A binomial test was performed to determine whether flickers that have parental phenotypes and are presumably immigrants rather than hybrids tend to choose mates similar to themselves. This test again indicated random mating in the hybrid zone. These data complement earlier reports in providing an additional test of alternative hybrid zone theories. Random mating along with evidence of high reproductive success in hybrids suggests that reinforcement of premating reproductive isolation is not occurring and that speciation will not result from the dynamics of the hybrid zone. At least some of the plumage traits seem to be important in the integrated courtship and territorial behaviors of flickers. Since these plumage differences have no significance in mate choice, it is inferred that they evolved in an intrasexual selection context rather than in intersexual or species-recognition contexts. Selection gradients appear to be important in restricting hybridization to a narrow zone. The gradients could result from ecological selection or from sexual selection.     

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.     

Historical hybrid zone movement: More pervasive than appreciated

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

Ecological speciation in an island snail: evidence for the parallel evolution of a novel ecotype and maintenance by ecologically dependent postzygotic isolation

Speciation is the process by which reproductive isolation evolves between populations. Two general models of speciation have been proposed: ecological speciation, where reproductive barriers evolve due to ecologically based divergent selection, and mutation‐order speciation, where populations fix different mutations as they adapt to similar selection pressures. I evaluate these alternative models and determine the progress of speciation in a diverse group of land snails, genus Rhagada, inhabiting Rosemary Island. A recently derived keeled‐flat morphotype occupies two isolated rocky hills, while globose‐shelled snails inhabit the surrounding plains. The study of one hill reveals that they are separated by a narrow hybrid zone. As predicted by ecological speciation theory, there are local and landscape level associations between shell shape and habitat, and the morphological transition coincides with a narrow ecotone between the two distinct environments. Microsatellite DNA revealed a cline of hybrid index scores much wider than the morphological cline, further supporting the ecological maintenance of the morphotypes. The hybrid zone does not run through an area of low population density, as is expected for mutation‐order hybrid zones, and there is a unimodal distribution of phenotypes at the centre, suggesting that there is little or no prezygotic isolation. Instead, these data suggest that the ecotypes are maintained by ecologically dependent postzygotic isolation (i.e. ecological selection against hybrids). Mitochondrial and Microsatellite DNA indicate that the keeled‐flat form evolved recently, and without major historical disruptions to gene flow. The data also suggest that the two keeled‐flat populations, inhabiting similar rocky hills, have evolved in parallel. These snails provide a complex example of ecological speciation in its early stages.     

Distortion of symmetrical introgression in a hybrid zone: evidence for locus-specific selection and uni-directional range expansion

The fate of species integrity upon natural hybridization depends on the interaction between selection and dispersal. The relative significance of these processes may be studied in the initial phase of contact before selection and gene flow reach equilibrium. Here we study a hybrid zone of two salamander species, Lyciasalamandra antalyana and Lyciasalamandra billae, at the initial phase of hybridization. We quantify the degree and mode of introgression using nuclear and mtDNA markers. The hybrid zone can be characterized as an abrupt transition zone, the central hybrid zone being only c. 400 m, but introgressed genes were traced up to 3 km. Introgression was traced in both sexes but gene flow may be slightly male-biased. Indirect evidence suggests that hybrid males are less viable than females. Introgression occurred at two levels: (1) locus-specific selection led to different allelic introgression patterns independent of species, while (2) asymmetrical species-level introgression occurred predominately from L. antalyana to L. billae due to range expansion of the former. This indicates that foreign genes can be incorporated into novel genomic environments, which in turn may contribute to the great diversity of morphological variants in Lyciasalamandra.     

ECOLOGICAL GENETICS OF A MOSAIC HYBRID ZONE: MITOCHONDRIAL,NUCLEAR, AND REPRODUCTIVE DIFFERENTIATION OF CRICKETS BY SOIL TYPE

We investigated the effects that habitat variation has on the structure and dynamics of a hybrid zone between two closely related crickets in Connecticut. A collecting protocol was developed in which crickets were sampled from characteristic habitats on either side of the hybrid zone and from two distinct habitat types within the zone. Presumptive pure Gryllus pennsylvanicus were sampled from fields in northwestern Connecticut and represent “inland” populations. “Pure” Gryllus firmus were sampled from beaches along the coast and represent the “coastal” populations. Crickets from within the hybrid zone were sampled from two different soil types: the “loam” populations from loamy soils and the “sand” populations from sandy soils. Moreover, an attempt was made to identify closely adjacent sand and loam localities to determine the scale of habitat variation and its possible effects on hybrid-zone structure. In general, there was little variation in morphological traits or in allozyme and mtDNA genotype frequencies among localities from within each of the four habitat types. Between each of the closely situated sand and loam localities within the hybrid zone, however, there were very significant differences in each of these sets of markers. In addition, crickets from hybrid-zone populations were tested for reproductive isolation. The asymmetric outcome of hybrid crosses that exists across the zone (Harrison, 1983) also exists on a finer ecological scale within the zone. Thus, this hybrid zone is a mosaic of strikingly differentiated populations. The dynamics of hybrid zones with mosaic structures are discussed in contrast to the traditional clinal models. The data are also discussed in light of the semipermeable nature of species boundaries. The extent to which a species boundary is permeable varies not only from one genetic marker to the next, but also with the ecological and geographic context of species interaction.     

A hybrid zone between Chorthippus parallelus parallelus and Chorthippus parallelus erythropus (Orthoptera: Acrididae): morphological and electrophoretic characters

Two subspecies of the grasshopper Chorthippus parullelus have distributions that abut in the Pyrenees. We have studied morphological and electrophoretic variation in II populations near Mont Louis, Pyrénées-Orientales, France. The two subspecies differ in several morphological characters and at one enzyme locus, esterase-2, and in this area they hybridize forming a cline in morphology less than 5 km wide. Examination of museum specimens suggests that this hybrid zone extends for most of the length of the Pyrenees, possibly breaking down in the extreme west. As the two subspecies are known to differ in mating behaviour this hybrid zone is well suited to the study of reinforcement.     

SNP genotyping for detecting the ‘rare allele phenomenon’ in hybrid zones

Hybrid zones are regions where genetically distinct populations meet, mate and produce offspring. In such zones, genetically less compatible gene combinations are usually generated, resulting in reduced fitness, and hybrid zones are often maintained because of continuous removal of unfit genotypes, balanced by gene flow into the zone from the parental populations (and are then referred to as ‘tension zones’). Tension zones often display unexpectedly high frequencies of gene variants that are rare outside the zone. Previous work has shown that this ‘rare allele phenomenon’ is not the result of intragenic recombination or increased mutation rates. Further understanding of the population genetics of the phenomenon requires an approach in which both the numbers of individuals and the numbers of loci is increased. Here, we report an approach using a combination of Illumina next‐generation sequencing and mass spectrophotometer genotyping to identify markers that may be used for genome‐wide investigations of the rare allele phenomenon. We test this approach on a hybrid zone in the land snail Albinaria hippolyti from Greece.     

Reed frog diversification in the Gulf of Guinea: Overseas dispersal,the progression rule,and in situ speciation

Oceanic islands accumulate endemic species when new colonists diverge from source populations or by in situ diversification of resident island endemics. The relative importance of dispersal versus in situ speciation in generating diversity on islands varies with a number of archipelago characteristics including island size, age, and remoteness. Here, we characterize interisland dispersal and in situ speciation in frogs endemic to the Gulf of Guinea islands. Using mitochondrial sequence and genome‐wide single‐nucleotide polymorphism data, we demonstrate that dispersal proceeded from the younger island (São Tomé) to the older island (Príncipe) indicating that for organisms that disperse overseas on rafts, dispersal between islands may be determined by ocean currents and not island age. We find that dispersal between the islands is not ongoing, resulting in genotypically distinct but phenotypically similar lineages on the two islands. Finally, we demonstrate that in situ diversification on São Tomé Island likely proceeded in allopatry due to the geographic separation of breeding sites, resulting in phenotypically distinct species. We find evidence of hybridization between the species where their ranges are sympatric and the hybrid zone coincides with a transition from agricultural land to primary forest, indicating that anthropogenic development may have facilitated secondary contact between previously allopatric species.     

A narrow hybrid zone between two Cottus species in Wills Creek, Potomac drainage

We describe a narrow hybrid zone between the mottled sculpin (Cottus b. bairdi) and the Blue Ridge sculpin (C. caeruleomentum). Seven characters (dorsal fin rays, pectoral fin rays, caudal base band condition, male spawning coloration, and one frequency and two fixed allozyme differences) distinguish the two taxa in the hybrid zone. C. caeruleomentum and C. b. bairdi diverged in these characters in allopatry as indicated by their distribution on opposite sides of the Atlantic-Ohio divide. However, a stream capture placed these two taxa in secondary contact in Wills Creek, Potomac drainage (Atlantic slope). Allozyme data indicate the presence of post-F(1) hybrids in the zone of secondary contact. Changes in allozymes, morphology, and spawning coloration along a transect in Wills Creek reveal the hybrid zone is less than 20 river kilometers in length. Estimates of root mean square dispersal and gene flow tentatively suggest that selection is operating in the Wills Creek hybrid zone. C. b. bairdi and C. caeruleomentum are maintaining their identity in seven distinguishing characters on opposite ends of the hybrid zone revealing these two taxa are independent evolutionary lineages.     

Asymmetries in male aggression across an avian hybrid zone

Recent studies suggest that competitive asymmetries are causingthe hybrid zones between hermit and Townsend's warblers to move,such that Townsend's warblers are replacing hermit warblers.Here we examine the contribution of male aggression to thiscompetitive asymmetry by measuring aggressive responses to mounts.We presented male mounts of the two parental species to Townsend'sand hermit warblers outside the zone and to hybrids within the zone.Outside the zone, Townsend's males are more aggressive to both conspecificand heterospecific mounts than are hermit males. This asymmetry shouldmove the zone in the direction inferred from previous studies.Hybrids fall between parentals in their aggressiveness, whichshould accelerate the movement of the zone. Remarkably, we foundno relationship between phenotype and aggression in individualmales at a locality within the hybrid zone. The forces of selectionand dispersal that maintain narrow hybrid zones should generatesuch a correlation if aggressive differences between the parental speciesare genetically controlled. We resolve this conflict by proposinga behavioral model of competitive sorting within localities.If birds are sorted across the hybrid zone according to competitiveabilities, and competitive interactions within neighborhoodsare more or less complete, then the correlation between phenotypeand aggression within any given neighborhood will be eliminated.We tested this model by examining the relationship between phenotypeand aggression across the zone. Warblers in hybrid neighborhoodson the Townsend's side of the zone are more aggressive thanwarblers in hybrid neighborhoods on the hermit side, indicatingthat competitive sorting is occurring.     

Little and large: body size and genetic clines in a New Zealand gecko (Woodworthia maculata) along a coastal transect

Clinal variation can result from primary differentiation or secondary contact and determining which of these two processes is responsible for the existence of a cline is not a trivial problem. Samples from a coastal transect of New Zealand geckos (Woodworthia maculata) identified for the first time a body size cline 7-10 km wide. The larger geckos are almost twice the mass of the small adult geckos. Clines in allele and haplotype frequency were found at two of the four genetic loci examined. Estimated width of the morphological cline was concordant with neither the narrower mtDNA cline (3-7 m) nor the wider nuclear cline (RAG-2; 34-42 km), and cline centers were not coincident. Although the body size cline is narrow compared to the entire range of the species, it is 2-3 orders of magnitude greater than estimates of dispersal distance per generation for these geckos. No evidence of assortative mating, nor of hybrid disadvantage was identified, thus there is little evidence to infer that endogenous selection is maintaining a hybrid zone. We cannot distinguish secondary contact from primary origin of this body size cline but conclude that secondary contact is likely due to the occurrence of mtDNA haplotypes from three distinct clades within the coastal transect and the presence of two frequency clines within this region.     

基于NDVI绿洲-荒漠过渡带宽度识别——以河西走廊中部荒漠绿洲为例

绿洲-荒漠过渡带是荒漠与绿洲之间的生态缓冲区,在维持绿洲能量流动、物质循环和景观稳定方面具有重要作用。过渡带宽度和属性直接影响到了其在整个绿洲系统中的功能发挥。以河西走廊中部张掖绿洲的Landsat(OLI)NDVI(归一化植被指数,Normalized Difference Vegetation Index)数据为基础,结合野外调查将过渡带类型划分为绿洲与石质裸山、砾质荒漠、沙质荒漠和人工固沙区4类。采用缓冲分析、分段线性趋势分析和尺度分别为30 m、90 m、210 m、330 m焦点分析等方法研究了不同类型过渡带宽度和尺度依赖特征。结果表明,在不同尺度上绿洲外缘NDVI变化存在二种线性回归趋势,趋势线交点至绿洲边界距离可确定为过渡带宽度。不同尺度分析表明,绿洲-石质裸山过渡带宽度为165—220 m,在其内NDVI线性变化趋势显著(P0.05)。绿洲-砾质荒漠过渡带宽度保持在330 m,在其内NDVI变化趋势极显著(P0.001)。绿洲-沙质荒漠过渡带宽度变化在230—290 m,NDVI变化趋势也为极显著(P0.001)。绿洲-人工固沙区过渡带宽度变化在570—580 m,与其它类型不同地是在过渡带内存在二种变化趋势,在210—240 m范围内变化趋势极显著(P0.001),超出此范围线性回归趋势不显著(P0.05)。     

Stepping out: a computer simulation of hominid dispersal from Africa

A succession of new discoveries and the recent application of new dating methods provide strong evidence that Eurasia was colonized soon after 2.0m.y.a. In light of this new evidence many scenarios have been proposed regarding the influence of glacial/interglacial cycles on hominid dispersal, the role of mountain chains and deserts as barriers, the significance of land bridges and the possibility of sea-crossings. Such factors have been proposed to explain the apparent early arrival of hominids in East Asia and relatively late arrival in Europe, although the evidence in both regions remains open to various interpretations. While it is relatively easy to propose environmental factors that may have influenced dispersal patterns, it is more difficult to evaluate such proposals and to establish what the combined impact of several factors might have been. Moreover, the role of historical contingency in creating the observed pattern of dispersal has yet to be considered. This paper describes a computer simulation model of hominid dispersal which seeks to provide a means to evaluate environmental and ecological factors in hominid dispersal. It creates probability distributions for arrival dates at six key localities and compares these with current estimates from the archaeological and fossil records. It uses these to support some of the current arguments about dispersal and to challenge others.     

A rare model limits the distribution of its more common mimic: a twist on frequency-dependent Batesian mimicry

Batesian mimics are predicted to lose their fitness advantage not only in the absence of an unpalatable model, but also when the mimic becomes relatively abundant. The phenotypic hybrid zone between mimetic and nonmimetic admiral butterflies, comprising the polytypic Limenitis arthemis species complex, offers an ideal opportunity to test these predictions because the position of the hybrid zone is hypothesized to be controlled by the geographic range of Battus philenor , the chemically defended model. We used 29 years of observational field data from a continental-scale butterfly monitoring program, the 4th of July Butterfly Counts, to show that (1) the advantage of mimicry does not extend beyond the range of the model, (2) in contrast to expectations, the mimicry complex is maintained even where the model is rare and (3) the sharp phenotypic transition between mimetic and nonmimetic admiral populations occurs over a very narrow spatial scale corresponding to the limit of the model's range. These results suggest that, even at very low densities, there is selection for Batesian mimicry and it maintains the geographic position of this hybrid zone. Our findings highlight the value of large-scale, long-term citizen science monitoring programs for answering basic ecological and evolutionary questions.     

Spatio-temporal changes in the genetic structure of the Passerina bunting hybrid zone

Although theoretical models predict that the structure of a hybrid zone can change under a variety of scenarios, only a few empirical studies of hybrid zones have unequivocally demonstrated zone movement. These studies are rare because few data sets exist that include repeated, temporally spaced, samples of the same hybrid zone. We analysed mitochondrial DNA haplotype data from samples separated by 40-45 years from across the Passerina amoena (Lazuli Bunting) and Passerina cyanea (Indigo Bunting) hybrid zone to investigate whether the genetic structure of this zone has changed during that interval. Both cline and generalized linear mixed modelling analyses uncovered a significant narrowing and a substantial westward shift of the Passerina bunting hybrid zone, clearly illustrating hybrid zone movement. The cause of the change may be due to a combination of ecological, demographic and behavioural factors. Our results predict that the width of the hybrid zone will continue to narrow over time, a finding consistent with reinforcement theory.     

Genetic, morphological, and ecological characterization of a hybrid zone that spans a migratory divide

This study characterizes a hybrid zone that spans a migratory divide between subspecies of the Swainson's thrush (Catharus ustulatus), a long distance migratory songbird, in the Coast Mountains of British Columbia. To assess the potential for a barrier to gene flow between the subspecies, I: (1) analyzed the shape and width of genetic and morphological clines relative to estimates of dispersal distance, (2) assessed the ratio of parental to hybrid genotypes across the hybrid zone, (3) estimated population density across the hybrid zone, and (4) compared the spatial relationship between the hybrid zone and an existing environmental gradient. The results indicate that the hybrid zone is characterized by mostly concordant character clines that are narrow relative to dispersal, the absence of a hybrid swarm, and low population density at the center of the zone. This hybrid zone and additional regions of contact between these subspecies are found on the border between coastal and interior climatic regions throughout the Pacific Northwest. An identified shift in the location, but not the width, of the mtDNA cline relative to the nuclear clines is consistent with asymmetrical hybridization. Neutral diffusion of populations following secondary contact and hybrid superiority within an ecotone are insufficient explanations for the observed patterns. The hypothesis that best fits the data is that the Swainson's thrush hybrid zone is a tension zone maintained by dispersal and ecologically mediated barriers to gene flow.