Variability of Hybridization in Topminnows

Pairs of species that exhibit broadly overlapping distributions, and multiple geographically isolated contact zones, provide opportunities to investigate the mechanisms of reproductive isolation. Such naturally replicated systems have demonstrated that hybridization rates can vary substantially among populations, raising important questions about the genetic basis of reproductive isolation. The topminnows, Fundulus notatus and F. olivaceus, are reciprocally monophyletic, and co-occur in drainages throughout much of the central and southern United States. Hybridization rates vary substantially among populations in isolated drainage systems. We employed genome-wide sampling to investigate geographic variation in hybridization, and to assess the possible importance of chromosome fusions to reproductive isolation among nine separate contact zones. The species differ by chromosomal rearrangements resulting from Robertsonian (Rb) fusions, so we hypothesized that Rb fusion chromosomes would serve as reproductive barriers, exhibiting steeper genomic clines than the rest of the genome. We observed variation in hybridization dynamics among drainages that ranged from nearly random mating to complete absence of hybridization. Contrary to predictions, our use of genomic cline analyses on mapped species-diagnostic SNP markers did not indicate consistent patterns of variable introgression across linkage groups, or an association between Rb fusions and genomic clines that would be indicative of reproductive isolation. We did observe a relationship between hybridization rates and population phylogeography, with the lowest rates of hybridization tending to be found in populations inferred to have had the longest histories of drainage sympatry. Our results, combined with previous studies of contact zones between the species, support population history as an important factor in explaining variation in hybridization rates

Status of the Blackstripe (Fundulus notatus) and Blackspotted (F. olivaceus) Topminnows in the Ozark uplands of Central Missouri

The topminnow species Fundulus notatus and F. olivaceus have broadly overlapping geographic distributions that extend throughout much of the central and southern United States. In the northern portion of their respective ranges, in Missouri, the regional distributions of the two species coincide largely with recognized ecoregions. In the unglaciated southern half of Missouri, F. olivaceus is distributed throughout Ozark upland habitats while F. notatus is abundant in marginal large river and prairie habitats along the Ozark borders. An exception to this partitioning is the historical report of abundant F. notatus in the Bourbeuse and upper Meramec River drainages within the Ozark uplands ecoregion. We conducted an extensive survey of the Bourbeuse and Dry Fork Meramec Rivers to determine topminnow species composition in these systems. Our surveys found abundant F. olivaceus populations throughout these drainages and failed to uncover any F. notatus individuals. A review of museum accessions from the 1940s and 1960s confirms the historical presence of F. notatus in these river drainages, suggesting that a significant shift in topminnow species abundance has occurred in the past half century

Variable Rates Of Hybridization Among Contact Zones Between A Pair Of Topminnow Species, Fundulus Notatus And F. Olivaceus

Pairs of species that exhibit broadly overlapping distributions, and multiple geographically isolated contact zones, provide opportunities to investigate the mechanisms of reproductive isolation. Such naturally replicated systems have demonstrated that hybridization rates can vary substantially among populations, raising important questions about the genetic basis of reproductive isolation. The topminnows, Fundulus notatus and F. olivaceus, are reciprocally monophyletic, and co-occur in drainages throughout much of the central and southern United States. Hybridization rates vary substantially among populations in isolated drainage systems. We employed genome-wide sampling to investigate geographic variation in hybridization, and to assess the possible importance of chromosome fusions to reproductive isolation among nine separate contact zones. The species differ by chromosomal rearrangements resulting from Robertsonian (Rb) fusions, so we hypothesized that Rb fusion chromosomes would serve as reproductive barriers, exhibiting steeper genomic clines than the rest of the genome. We observed variation in hybridization dynamics among drainages that ranged from nearly random mating to complete absence of hybridization. Contrary to predictions, our use of genomic cline analyses on mapped species-diagnostic SNP markers did not indicate consistent patterns of variable introgression across linkage groups, or an association between Rb fusions and genomic clines that would be indicative of reproductive isolation. We did observe a relationship between hybridization rates and population phylogeography, with the lowest rates of hybridization tending to be found in populations inferred to have had the longest histories of drainage sympatry. Our results, combined with previous studies of contact zones between the species, support population history as an important factor in explaining variation in hybridization rates

Evolution of a Sexually Dimorphic Trait In a Broadly Distributed Topminnow (\u3ci\u3eFundulus olivaceus\u3c/i\u3e)

Understanding the interaction between sexual and natural selection within variable environments is crucial to our understanding of evolutionary processes. The handicap principle predicts females will prefer males with exaggerated traits provided those traits are indicators of male quality to ensure direct or indirect female benefits. Spatial variability in ecological factors is expected to alter the balance between sexual and natural selection that defines the evolution of such traits. Male and female blackspotted topminnows (Fundulidae: Fundulus olivaceus) display prominent black dorsolateral spots that are variable in number across its broad range. We investigated variability in spot phenotypes at 117 sites across 13 river systems and asked if the trait was sexually dimorphic and positively correlated with measures of fitness (condition and gonadosomatic index [GSI]). Laboratory and mesocosm experiments assessed femalemate choice and predation pressure on spot phenotypes. Environmental and community data collected at sampling locations were used to assess predictive models of spot density at the individual, site, and river system level. Greater number of spots was positively correlated with measures of fitness in males. Males with more spots were preferred by females and suffered greater mortality due to predation. Water clarity (turbidity) was the best predictor of spot density on the drainage scale, indicating that sexual and natural selection for the trait may be mediated by local light environments

Evolution of a Sexually Dimorphic Trait in a Broadly Distributed Topminnow (Fundulus Olivaceus)

Understanding the interaction between sexual and natural selection within variable environments is crucial to our understanding of evolutionary processes. The handicap principle predicts females will prefer males with exaggerated traits provided those traits are indicators of male quality to ensure direct or indirect female benefits. Spatial variability in ecological factors is expected to alter the balance between sexual and natural selection that defines the evolution of such traits. Male and female blackspotted topminnows (Fundulidae: Fundulus olivaceus) display prominent black dorsolateral spots that are variable in number across its broad range. We investigated variability in spot phenotypes at 117 sites across 13 river systems and asked if the trait was sexually dimorphic and positively correlated with measures of fitness (condition and gonadosomatic index [GSI]). Laboratory and mesocosm experiments assessed female mate choice and predation pressure on spot phenotypes. Environmental and community data collected at sampling locations were used to assess predictive models of spot density at the individual, site, and river system level. Greater number of spots was positively correlated with measures of fitness in males. Males with more spots were preferred by females and suffered greater mortality due to predation. Water clarity (turbidity) was the best predictor of spot density on the drainage scale, indicating that sexual and natural selection for the trait may be mediated by local light environments

Population Genetics of Death Valley Pupfishes (Cyprinodontidae:Cyprinodon Spp.) and the Identification of a New Retrotransposable Element Family

Study of the genetic relationships and evolutionary histories of pupfish populations (Cyprinodontidae: Cyprinodon spp.) from the remnant aquatic habitats of Death Valley was approached by exploring the genetic structure and divergence within and among populations using mitochondrial and nuclear DNA markers. The findings of these studies illustrate the influences of population size and isolation time in the divergence of small, fragmented populations largely via genetic drift. The information revealed in this study has implications for assessing priorities in the conservation of the unique evolutionary heritage among populations of the Death Valley pupfishes. A new retrotransposable element family was identified and characterized. This family of genetic elements was uncovered during a search of the pupfish genome for transposable elements to be used as molecular markers for population analyses. The description of this element family, named Swimmer 1 (SW1), provides new insights into the evolution of long interspersed nuclear elements (LINEs) in vertebrates. Therefore, a full characterization of the SW1 element family was undertaken in the Japanese medaka (Oryzias latipes) as well as in the pupfish genome. The Japanese medaka is a model organism widely used for genetic and developmental biology studies

Evolutionary Genetics of Death Valley Pupfish Populations: Mitochondrial DNA Sequence Variation and Population Structure

The pupfishes of Death Valley (genus: Cyprinodon) are a classic example of rapid (post-Pleistocene) allopatric divergence with several populations exhibiting striking morphological, physiological, and behavioural differences. However, genetic changes accompanying or causing this divergence have been poorly resolved and/or difficult to detect. In order to investigate the phylogeography of the system and infer historical effective population sizes, mtDNA sequence variation was assessed within and among populations. In total, mtDNA D-loop sequences revealed 11 haplotypes among 278 individuals from 16 populations. Genetic diversity within populations was generally low (1-3 haplotypes per population) suggesting relatively small effective population sizes. Most variation occurred among populations, resulting in extensive genetic structure. The genealogical relationships of mtDNA haplotypes were determined by a combined phylogenetic analysis of both D-loop and ND2 sequences. These relationships revealed that the present distribution of haplotypes in Death Valley has probably resulted from stochastic, and in some cases, incomplete sorting of ancestral variation. The presence of highly divergent haplotypes in some populations may indicate that ancestral population sizes were substantially larger, perhaps rivalling population sizes observed in contemporary coastal pupfish species. In contrast, other Death Valley populations of equal contemporary size appeared monomorphic suggesting historical genetic bottleneck events. The Death Valley pupfishes provide an important example of the limitations inherent in defining evolutionarily significant units (ESUs) for conservation purposes based on mtDNA sequence variation alone

Variation and Divergence of Death Valley Pupfish Populations at Retrotransposon-Defined Loci

A population survey of the Death Valley pupfishes (Cyprinodontidae: Cyprinodon sp.) for insertional variation associated with \u27Swimmer 1\u27 (SW1), a retrotransposon family, was conducted with Southern blot hybridization. Numerous polymorphic insertion sites were detected, providing compelling evidence that SW1 has been retrotranspositionally active in the recent history of the Death Valley pupfishes. This extensive variation revealed marked genetic divergence among some populations that were indistinguishably monomorphic by other molecular techniques. Large disparities were also detected among populations in the levels of genetic diversity exhibited at SW1defined loci. These differences may have resulted from either variability among populations in SW1 retrotranspositional activity (i.e., mutation rates) or variable rates of genetic drift mediated by differences in effective population size. The patterns of genetic variation suggest that most polymorphic sites derive from a common ancestor and that recent population divergence has occurred primarily through loss of variability via drift

Variation in Contact Zone Dynamics Between Two Species of Topminnows, \u3ci\u3eFundulus notatus\u3c/i\u3e and \u3ci\u3eF. olivaceus\u3c/i\u3e, Across Isolated Drainage Systems

Spatially variable selection pressure within heterogeneous environments can result in the evolution of specialist phenotypes that facilitate co-occurrence of closely related species and limit genetic exchange. If divergent selection pressures maintain reproductive isolation, hybridization is expected to correlate with the strength of underlying ecological gradients and the traits shaped by adaptive processes. We sampled ten replicate topminnow (Fundulus olivaceus and Fundulus notatus) hybrid zones in isolated drainages throughout central and southern North America. In all drainages, species were distributed in an upstream–downstream manner with contact zones localized at confluences featuring abrupt shifts from tributary to river habitat. In two drainages, the typical up and downstream positions of species were reversed. Phenotype differences between the species reflect predicted selection differences along stream gradients. Downstream populations (lower food availability and greater predator pressure) generally showed larger investment in reproduction (higher gonadal somatic index), smaller body size and lower somatic condition compared to upstream populations. Phenotypic differences between the species in the two reversed drainages were consistent with convergence of life history traits in the respective habitats. Phenotypes of individuals of hybrid origin (F1 hybrids or backcrosses) were not significantly different from the average of the two parental forms, though there were trends towards reduced fitness. The prevalence of hybridization among drainages ranged from no hybrids in two drainages to near random mating. The strongest correlates of hybridization rate among replicate hybrid zones were similarity in body shape and the homogeneity of habitat through tributary-river confluences. The two reversed orientation hybrid zones also exhibited high prevalence of hybrids suggesting that phenotypic convergence could lead to increased hybridization