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When introduced or cultivated plants or animals hybridize with their native relatives, the spread of invasive genes into native populations might have biological, aesthetic, and legal implications. Models suggest that the rate of displacement of native by invasive alleles can be rapid and inevitable if they are favored by natural selection. We document the spread of a few introduced genes 90 km into a threatened native species (the California Tiger Salamander) in 60 years. Meanwhile, a majority of genetic markers (65 of 68) show little evidence of spread beyond the region where introductions occurred. Using computer simulations, we found that such a pattern is unlikely to emerge by chance among selectively neutral markers. Therefore, our results imply that natural selection has favored both the movement and fixation of these exceptional invasive alleles. The legal status of introgressed populations (native populations that are slightly genetically modified) is unresolved by the US Endangered Species Act. Our results illustrate that genetic and ecological factors need to be carefully weighed when considering different criteria for protection, because different rules could result in dramatically different geographic areas and numbers of individuals being protected.

Although the ecological consequences of species invasions are well studied, the ecological impacts of genetic introgression through hybridization are less understood. This is particularly true of the impacts of hybridization on \"third party\" community members not genetically involved in hybridization. We also know little about how direct interactions between hybrid and parental individuals influence fitness. Here, we examined the ecological effects of hybridization between the native, threatened California Tiger Salamander (Ambystoma californiense) and the introduced Barred Tiger Salamander (Ambystoma tigrinum mavortium). Native x introduced hybrids are widespread in California, where they are top predators in seasonal ponds. We examined the impacts of early generation hybrids (first 2 generations of parental crosses) and contemporary hybrids derived from ponds where hybrids have been under selection in the wild for 20 generations. We found that most classes of hybrid tiger salamander larvae dramatically reduced survival of 2 native community members, the Pacific Chorus Frog (Pseudacris regilla) and the California Newt (Taricha torosa). We also found that native A. californiense larvae were negatively impacted by the presence of hybrid larvae: Native survival and size at metamorphosis were reduced and time to metamorphosis was extended. We also observed a large influence of Mendelian dominance on size, metamorphic timing and predation rate of hybrid tiger salamanders. These results suggest that both genetic and ecological factors are likely to influence the dynamics of admixture, and that tiger salamander hybridization might constitute a threat to additional pond-breeding species of concern in the region.

Recently metamorphosed amphibians transport substantial biomass and nutrients from wetlands to terrestrial ecosystems. Previous estimates (except 1) were limited to either a subset of the community or a single year. Our goal was to examine temporal variability in biomass export of all amphibians within breeding ponds and the composition of that export. We completely encircled ponds with drift fences to capture, count, and weigh emerging recently metamorphosed individuals in Maine (four wetlands, six years) and Missouri (eight wetlands, 2–4 years). We estimated total amphibian biomass export, export scaled by pond surface area, species diversity, and percentage of biomass from anurans. Biomass export and export composition varied greatly among ponds and years. Our estimates were of similar magnitude to previous studies. Amphibian biomass export was higher when species diversity was low and the proportion of anurans was higher. Biomass estimates tended to be highest for juvenile cohorts dominated by a single ranid species: green frogs (Missouri) or wood frogs (Maine). Ranid frogs made up a substantial proportion of amphibian biomass export, suggesting that terrestrial impacts will likely occur in the leaf litter of forests. Future studies should examine the impacts of ranid juveniles on terrestrial ecosystem dynamics.

Background Hybrid zones represent valuable opportunities to observe evolution in systems that are unusually dynamic and where the potential for the origin of novelty and rapid adaptation co-occur with the potential for dysfunction. Recently initiated hybrid zones are particularly exciting evolutionary experiments because ongoing natural selection on novel genetic combinations can be studied in ecological time. Moreover, when hybrid zones involve native and introduced species, complex genetic patterns present important challenges for conservation policy. To assess variation of admixture dynamics, we scored a large panel of markers in five wild hybrid populations formed when Barred Tiger Salamanders were introduced into the range of California Tiger Salamanders. Results At three of 64 markers, introduced alleles have largely displaced native alleles within the hybrid populations. Another marker ( GNAT1 ) showed consistent heterozygote deficits in the wild, and this marker was associated with embryonic mortality in laboratory F2's. Other deviations from equilibrium expectations were idiosyncratic among breeding ponds, consistent with highly stochastic demographic effects. Conclusion While most markers retain native and introduced alleles in expected proportions, strong selection appears to be eliminating native alleles at a smaller set of loci. Such rapid fixation of alleles is detectable only in recently formed hybrid zones, though it might be representative of dynamics that frequently occur in nature. These results underscore the variable and mosaic nature of hybrid genomes and illustrate the potency of recombination and selection in promoting variable, and often unpredictable genetic outcomes. Introgression of a few, strongly selected introduced alleles should not necessarily affect the conservation status of California Tiger Salamanders, but suggests that genetically pure populations of this endangered species will be difficult to maintain.

Background Introductions of non-native tiger salamanders into the range of California tiger salamanders have provided a rare opportunity to study the early stages of secondary contact and hybridization. We produced first- and second-generation hybrid salamanders in the lab and measured viability among these early-generation hybrid crosses to determine the strength of the initial barrier to gene exchange. We also created contemporary-generation hybrids in the lab and evaluated the extent to which selection has affected fitness over approximately 20 generations of admixture. Additionally, we examined the inheritance of quantitative phenotypic variation to better understand how evolution has progressed since secondary contact. Results We found significant variation in the fitness of hybrids, with non-native backcrosses experiencing the highest survival and F2 hybrids the lowest. Contemporary-generation hybrids had similar survival to that of F2 families, contrary to our expectation that 20 generations of selection in the wild would eliminate unfit genotypes and increase survival. Hybrid survival clearly exhibited effects of epistasis, whereas size and growth showed mostly additive genetic variance, and time to metamorphosis showed substantial dominance. Conclusions Based on first- and second- generation cross types, our results suggest that the initial barrier to gene flow between these two species was relatively weak, and subsequent evolution has been generally slow. The persistence of low-viability recombinant hybrid genotypes in some contemporary populations illustrates that while hybridization can provide a potent source of genetic variation upon which natural selection can act, the sorting of fit from unfit gene combinations might be inefficient in highly admixed populations. Spatio-temporal fluctuation in selection or complex genetics has perhaps stalled adaptive evolution in this system despite selection for admixed genotypes within generations.

When introduced species hybridize with native relatives, spread of advantageous invasive genes into native populations (introgression) is a conservation concern. Genome-scale SNP (single nucleotide polymorphism) analysis can be a powerful approach to detect hybridization and identify candidate loci experiencing selection in hybrid zones. However, followup studies are critical to verify and interpret potentially impactful patterns of introgression. In an earlier publication we identified three outlier loci (out of 68 unlinked SNPs) where non-native alleles appeared to have introgressed 90 km into the range of a threatened native salamander, while the other 65 markers showed no evidence of spread further than 12 km. This was consistent with strong selection favoring a few invasive traits, but our inferences necessarily depended on limited reference samples of the native species. Here, we further tested our initial interpretation by interrogating the outlier markers in greater detail. First, we isolated DNA from two museum specimens of native salamanders collected several decades before the introduction. Both had the putatively invasive SNPs, indicating that the SNP alleles were present before the introduction and therefore not diagnostic for nonnative ancestry. Second, we developed a novel genealogical analysis of DNA sequences (rather than SNPs) to infer allelic ancestry, since genealogical analysis of haplotypes minimizes the ancestry assignment errors that can occur with SNPs. When applied to the original loci, this analysis confirmed that the genotypes formerly interpreted as ‘superinvasive’ are native variants, and non-native alleles remain limited to areas near the original introduction sites.

Leuctra Stephens, 1836 is the fourth most speciose genus of Plecoptera (Leuctridae) east of the Rocky Mountains with 31 recognised species, trailing only Isoperla Banks, 1906 (58), Allocapnia Claassen, 1928 (47), and Perlesta Banks, 1906 (34). Although Leuctra females are described in taxonomic literature, they are difficult to morphologically distinguish among regional congeners, and identifications are often made through inference only (i.e., presence of males). This is particularly problematic in the Appalachian Mountains of the United States of America, which host numerous Leuctra species. We sampled the stonefly fauna of Mount Mitchell, western North Carolina, United States of America, from April to October 2019. The mitochondrial cytochrome C oxidase subunit 1 gene was sequenced for 39 adult males and 239 adult females of Leuctra. This allowed us to confidently place species names on all of the latter individuals. Phylogenetic tree- and genetic distance–based methods consistently grouped females with males for nine recognised species. Two separate L. ferruginea (Walker, 1852) operational taxonomic units were recognised, albeit with low divergence values, and an additional undetermined Leuctra was identified based solely on females. Digital stereomicroscope images were taken from females of each species unit to identify variation among and between species. This approach allowed for a more robust assessment of regional biogeographic patterns.

It is important to identify and understand the critical habitat components of organisms inhabiting landscapes that are increasingly altered by human activities to adequately predict the effects of habitat alteration on natural populations. Our study identifies terrestrial habitats that are important to the gray treefrog (Hyla versicolor) during the nonbreeding season using radiotracking and mark-recapture/release of individuals captured in artificial arboreal refugia. High humidity and rainfall were associated with decreased captures in artificial refugia, while high ambient air temperatures were correlated with increases in the number of treefrog captures. Refugia placed in small trees recorded more small individuals than those in larger trees, and refugia in white oaks (Quercus alba) contained females more frequently than expected at random. We found that space-use estimates encompassing foraging and overwintering locations of gray treefrogs were relatively small and typically included only a few adjacent trees during the nonbreeding season. We discuss the physical and biological aspects of habitat patches that may be important in determining the persistence of gray treefrog populations.

Phenology is a key driver of population and community dynamics. Phenological metrics (e.g., first date that an event occurred) often simplify information from the full phenological distribution, which may undermine efforts to determine the importance of life history events. Data regarding full phenological distributions are especially needed as many species are shifting phenology with climatic change which can alter life-history patterns and species dynamics. We tested whether skewness, kurtosis or maximum duration of breeding phenology affected juvenile emigration phenology and survival in natural populations of ringed (Ambystoma annulatum) and spotted salamanders (A. maculatum) spanning a 7-year period at two study locations. We evaluated the relative importance of different phenological metrics in breeding phenology and larval density dependence on emigration phenology and survival. We found that variability in emigration phenology differed by species, with ringed salamanders having a shorter duration and distributions that were more often right-skewed and leptokurtic compared to spotted salamanders. Emigration phenology was not linked to any measure of variability in breeding phenology, indicating phenological variability operates independently across life stages and may be subject to stage-specific influences. Emigration duration and skewness were partially explained by larval density, which demonstrates how phenological distributions may change with species interactions. Further tests that use the full phenological distribution to link variability in timing of life history events to demographic traits such as survival are needed to determine if and how phenological shifts will impact species persistence.

Concern over amphibian population declines and loss of terrestrial and aquatic habitat have emphasized the need to define habitat requirements for each stage in a species' life history. The realization that pond-breeding amphibians spend most of their lives in the terrestrial environment suggests the need to protect terrestrial as well as aquatic habitat. Many studies on amphibian populations have focused on emigration from breeding sites to define habitat use; however these studies do not typically elucidate terrestrial activities of adults within the breeding season. We measured colonization rates of artificial pools by gray treefrogs (Hyla versicolor) at multiple distances from natural breeding ponds. We found a non-random distribution of egg deposition among distances, with 95% of eggs deposited within 15 m of the breeding pond. Additionally, we found that the time to first colonization of artificial pools increased with respect to distance. Our results indicate that adult gray treefrogs may travel up to 200 m within a breeding season, and that multiple breeding ponds may be considered part of a single population. We suggest that a minimum core terrestrial habitat of 60 m surrounding breeding sites is appropriate for protection of local populations of gray treefrogs.