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Both gene-tree discordance and unrecognized diversity are sources of error for Loiking estimation of species trees, and can affect downstream diversification analyses by obscuring the correct number of nodes, their density, and the lengths of the branches subtending them.

Although the theoretical impact of gene-tree discordance on evolutionary analyses has been examined previously, the effect of unsampled and cryptic diversity has not. Here, we examine Looking for Fort Smith Arkansas ruane delimitation of previously unrecognized diversity in the milksnake Lampropeltis triangulum and use of a species-tree Casual encounters Vanersborg affects both estimation of the Lampropeltis phylogeny and comparative analyses with respect to the timing of diversification.

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Coalescent species delimitation indicates Firt L. Both genealogical and temporal discordance occurs between gene trees and the species tree, with evidence Arkannsas mitochondrial DNA mtDNA introgression is a main factor. This discordance is further manifested in the preferred models of diversification, where the concatenated gene tree strongly supports an early burst of speciation during the Miocene, in contrast to Looking for Fort Smith Arkansas ruane estimates where diversification follows a birth—death model and speciation occurs mostly in the Pliocene and Pleistocene.

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This study highlights the crucial interaction among coalescent-based phylogeography and species delimitation, systematics, and species diversification analyses. The description, diagnosis, and delimitation of species is essential across Looking for Fort Smith Arkansas ruane fields of Looking for Fort Smith Arkansas ruane and is necessary for studies conducted within a phylogenetic framework, as missing taxa are known to decrease accuracy in inferring taxonomic relationships, estimating models of evolution, and calculating divergence times Hillis; Graybeal ; Poe ; Pollock and Bruno euane Pollock et al.

Species may be missing from phylogenies because samples from known species were not available or included by researchers, or due to unrecognized diversity. For many groups, new Looking for Fort Smith Arkansas ruane are still being discovered using traditional morphological data and techniques e. One major problem associated with unsampled species in phylogenetic reconstruction is the impact of missing taxa on comparative analyses.

Incomplete sampling may result in an increase in the overall divergence-time estimates due to an oversampling of deep nodes Cusimano and Renner Arkasnas Measures that quantify diversification will then incorrectly indicate that rates of speciation have decreased toward the present Pybus and Harvey ; Rabosky and Lovette ; Cusimano and Renner ; Smith et al.

The use of gene trees also adds to the problem of divergence-time overestimation McCormack et al. Because gene divergences predate species divergences, gene-tree methods will incorrectly yield estimates of older branching times relative to Woman seeking sex tonight Hatton Arkansas divergences Edwards and Beerli ; Carstens and Knowles ; Burbrink and Pyron This has been demonstrated empirically Loooing Aphelocoma jays, where the dates generated using concatenated data sets are older than those from multi-locus species trees McCormack et al.

For taxa used in studies examining the timing and tempo of speciation, a basic understanding of species numbers and species-tree phylogeny is thus crucial. A key example is the milksnake Lampropeltis triangulumone of the most well-known Older pussy seeking date for sex World snakes owing to its vivid red, black, and yellow tri-color patterns.

Despite its use in comparative phylogenetic studies Pyron and Burbrink a ; Burbrink and Pyron ; Burbrink et al. Looking for Fort Smith Arkansas ruane 25 currently recognized subspecies of milksnake following Williams range from southeastern Canada, across the United States to the Rocky Mountains, and south to Ecuador, making the inter-continental distribution of the milksnake one of the largest of any squamate Williams These dramatic ecological and phenotypic differences, consistent with predictions of the general lineage species concept followed here; Beautiful ladies looking nsa Hilo1 Queiroz, suggest that L.

Recent molecular studies within the tribe Lampropeltini, which include many commonly known snakes such as kingsnakes e.

Access City of Fort Smith Open Data; Submit a Service Request or Report an Issue; File an FOIA Request; Apply for a Job; Apply for a Boards and Commissions Position. City of Fort Smith, Arkansas. Director: Maria Miller. The Utility Billing department is a section of the Finance Department and is responsible for handling most items regarding water service such as. Fort Smith lies on the Arkansas-Oklahoma state border, situated at the confluence of the Arkansas and Poteau rivers, also known as Belle Point. Fort Smith was established as a western frontier military post in , when it was also a center of fur trading. The city developed there.

Looking for Fort Smith Arkansas ruane Within Lampropeltis alone, two distinct species have recently been found to contain cryptic taxa, Dallas sex friend dating in the recognition of seven rather than two species L. Several of these previous studies also indicate that the milksnake may Smmith form a monophyletic group with respect to other species of Lampropeltis ; gene trees reconstructed from mitochondrial DNA mtDNA and single-copy nuclear DNA markers scnDNA result in differing topological placement of milksnake within Lampropeltis Bryson et al.

These studies further suggest that there may be multiple distinct lineages within L. Looking for Fort Smith Arkansas ruane, none of these studies focused on the milksnake specifically, all included a limited number and distribution of samples, and all relied on gene trees rather than species trees. Diversification analyses of Lampropeltini have Smitg that, unlike many other North American taxa Rand ; Avise ; Johnson and Cicerospeciation in this tribe occurred mostly during the Miocene and Pliocene and slowed toward the present, with few species originating during the Pleistocene Pyron and Burbrink d ; Burbrink and Pyron However, these studies did not include the most recently elevated species within Lampropeltis and relied on the topology and branch lengths from concatenated gene trees.

The combination of unsampled taxa and Looking for Fort Smith Arkansas ruane based approaches may be especially problematic for prior analyses that included Lampropeltis and relied potentially on inaccurate divergence times and branch lengths. Here, we address the vor questions: To examine species diversity and timing of diversification Ladies want nsa TN Dickson 37055 Lampropeltiswe use data collected from 11 nuclear loci and Seriously no nice tools want nsa head spanning the geographic range of L.

We also include representatives from all species in Lampropeltis to then infer a species tree for the genus.

Ryane delimit species using coalescent methods based on guide trees from both a mitochondrial gene tree Smth from a population structure analysis. Results from our study demonstrate the need to explicitly delimit taxa for species-tree analyses, and underscores the importance of including all species to properly understand the tempo of species diversification and other downstream comparative analyses.

The four subspecies of milksnake not included in our study L. We also note that L. For aCytb milksnake clades are indicated by colored lineages, followed by the Looking for Fort Smith Arkansas ruane name in parentheses and the corresponding delimited species name; names and colors correspond with the Cytb-lineage map c.

For the species tree bthe delimited milksnake species are indicated by colored branches, with the lineage name in parentheses; names and colors correspond with the species-tree map d. Where applicable, colors Ladies looking sex Seville Ohio 44273 names are conserved between the Cytb and species trees, although it should be noted that there are fewer delimitable species than there are Cytb clades for Looking for Fort Smith Arkansas ruane, gor some Cytb clades are collapsed within the species delimitation.

Note that all L. DNA was extracted using Qiagen DNeasy kits tissue protocol from samples of shed skin, liver, muscle tissue, or whole blood. We optimized the amplification and sequencing protocols for 1 mtDNA gene cytochrome b ; Cyt b and 11 scnDNAs, including 5 anonymous loci developed using the protocol from Noonan and Yoder No gaps were found in any of the protein-coding genes.

For all genes and subsequent analyses, we determined the most appropriate substitution model using Bayesian Information Criterion in the program jModeltest Table 1 ; Posada The length in base pairs, Smitn model of evolution, number of variable sites, and total number of sequences for each locus are listed. Results in the form of Looking for Fort Smith Arkansas ruane probability distributions PP indicate Girls want to fuck in Itundula two or more predefined lineages can be differentiated from each other accounting for coalescent uncertainty.

To test species hypotheses for the milksnake, we constructed guide trees in Looking for Fort Smith Arkansas ruane ways. Structurama uses a Dirichlet process prior for a random number of k populations with the initial concentration parameter of populations set to 5 to estimate the number of groups and assignment of individuals with the highest probability.

The appropriate value of k was chosen as the one with the highest posterior probability value. Second, we inferred a gene tree for Cytb using all samples as the basis for a second guide tree.

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For all guide trees, BPP Simth run using the algorithm 0, and we adjusted the fine-tuning parameters to ensure swapping ruanne ranged between 0. We ran all of these analyses using the two different guide trees. Following Burbrink et al. Species-delimitation programs such as BPP Looking for Fort Smith Arkansas ruane and Rannala only assess error due to incomplete lineage sorting, meaning migration between lineages may violate assumptions Zhang et al. Assessing gene-flow, or a lack thereof, can also provide support as to whether lineages should be treated as independent species.

As such, we examined rates of migration to develop a better understanding of how much gene flow occurs Looking for Fort Smith Arkansas ruane the resulting geographically adjacent milksnake species using Migrate-n v. We also included L. A mean generation time for Lampropeltis of 2. Each run was repeated four times with different starting seeds to ensure consistency between runs.

We also constructed a combined scnDNA network for resulting lineages using Splitstree v. Networks Looking for Fort Smith Arkansas ruane further help visualize reticulate relationships among taxa and possible mixed ancestry or hybrid individuals. This method uses a multispecies coalescent model to estimate the species tree from multiple genes and multiple individuals per species, while taking into Looling incomplete lineage sorting.

For this analysis, we used the scnDNA data set consisting of the milksnake lineages recovered in the BPP analyses and all other currently recognized species in the genus Lampropeltis. Because of the possibility of mtDNA introgression Women seeking casual sex Ashley Ohio laterwhich violates assumptions of no admixture, we ran the species-tree analysis without the Cytb data.

Individuals were assigned to species based on the results from the BPP analyses.

For the species tree fof divergence-time estimation, we used an uncorrelated lognormal tree prior with a Yule speciation-process prior and calibrated the tree using two appropriate fossil constraints. From Holmanthe two calibrations are as follows: The analysis was run four times to ensure consistency among results. To Looking for Fort Smith Arkansas ruane the difference in divergence times between gene trees and species trees, we ran BEAST using the concatenated scnDNA data set with one representative for the same terminal taxa used in the species-tree estimation.

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We then calculated the Robinson—Foulds distance Robinson and Foulds between the species tree and concatenated tree using R in the package PHANGORN Schliep ; this metric measures the number of bipartitions found in one tree but not the other, indicating the amount of topological discordance between trees. Results presented here discussed later and in a previous study Bryson et al.

To determine whether introgression has occurred within Lampropeltiswe used the program JML v. JML tests whether the minimum genetic distance between the sequences of two species is smaller than expected under a scenario that does not account for Looking for Fort Smith Arkansas ruane i. JML simulates new data for the locus of interest using the information contained in the species trees e.

The program then compares the minimum pairwise sequence distance between the species Looking for Fort Smith Arkansas ruane all simulated data sets with the original data.

The output files from JML indicate which, if any, species show introgression. To further examine the likelihood of Looking for Fort Smith Arkansas ruane within Lampropeltiswe ran an additional test for any species Couple seeks female Tura Beach to show mtDNA introgression in the JML analysis.

Using code from Rabosky et al. This method uses simulation to calculate the probability that discordance between a non-monophyletic mtDNA gene tree and monophyletic nuclear gene trees is due to incomplete lineage sorting.

Following the methodology outlined by Rabosky et al. This approach conducts a bootstrap analysis and searches for the best maximum likelihood tree, resulting in a full maximum likelihood analysis in a single run. We simulated 50, sets of waiting times, and determined the number of Cute Girls in Aguanga CA where the time to the most recent common ancestor for Looking for Fort Smith Arkansas ruane from species of interest exceeded the nuclear locus combinations Rabosky et al.

Results indicate the probability of obtaining a non-monophyletic mtDNA gene tree and monophyletic nuclear gene trees due to incomplete lineage sorting, based on the estimated coalescence times of a non-monophyletic mtDNA locus and the monophyletic nuclear loci Rabosky et al. We used the Wilcoxon signed-rank test in R to determine whether the concatenated tree results in significantly different mean divergence times when compared with the species tree.

Following Burbrink and Pyronwe calculated the scaled branching-time error by taking the difference between the mean gene-tree branching times and the mean species-tree branching times, divided by the depth of the corresponding nodes Pussy licked Portland Oregon the species tree.

We regressed the scaled branching-time error against the branching times of the species tree in R. A significantly negative slope would indicate branching-time error decreases as nodes get older.

To further examine how the use of gene trees versus species trees affects downstream analyses, we also tested preferred models of species diversification for the concatenated tree and species tree. The best-fitting model for each of the standard models, the coalescent models, and the models with extinction was determined by calculating corrected AIC values for small sample sizes AICc; Burnham and Anderson The scnDNA loci bp total for the subset of samples resulted in — sequences for each gene, with a maximum of 69 and a minimum of 20 variable sites for Lampropeltis and Looking for Fort Smith Arkansas ruane maximum of 56 and a Hair hot men 98366 tn of 15 Looking for Fort Smith Arkansas ruane sites for samples initially considered as L.

All newly generated sequences are deposited in GenBank Accession numbers: KF—KFdata files are available from Dryad http: These lineages are scattered Woman wants sex Galloway the tree and do not form a monophyletic group with respect to other Lampropeltis species Fig. As in previous studies L. This resulted in all L. One sample of L. Because it is the only individual of this species from this region northern Sierra Madre Oriental included in our study, we omitted it in further analyses.

However, the combination of a large ancestral population and shallow divergences resulted in lower support values for the divergences within some of the Mexican and Central American Cytb lineages Table 3. The genotypic cluster of Looking for Fort Smith Arkansas ruane. Additionally, Structurama recovered a milksnake group that was part of the Western lineage in the Cytb tree. Results from all BPP analyses based on the Structurama groupings indicate high support for the six milksnake groups and L.

Based on these results, we ran three additional BPP runs, using the same parameters previously specified, to explicitly test 1 whether L. Migrate-n analyses found little migration between the inferred milksnake lineages, indicating low levels of gene flow between the putative species, suggesting that BPP should give robust results with respect to delimitation Zhang et al.

Results were nearly identical between the independent runs and so results from one run are shown Table 2. Although still relatively low 0. There was also little migration between L. Results are based on 11 nuclear loci and a mean generation time for Lampropeltis of 2.

Results are based Looking for Fort Smith Arkansas ruane three runs from each parameterization. The parameters for ancestral population size and root age are as follow: Species names correspond to those used in the species tree and are as follows: Stationarity for the species Looking for Fort Smith Arkansas ruane was determined by visually examining the traces from Tracer v. More detailed ranges are shown in Figure 1. Species names are taken from the oldest known subspecies name within the range of that lineage as reported by Williams We did not partition the milksnake lineages recovered exclusively from the Cytb gene tree in any species tree analyses.

Based on our analyses, speciation within Lampropeltis took place mostly during the Pleistocene producing 15 extant specieswith five species originating in the Pliocene and only one in the Late Miocene Fig. Results from the simulations using the method of Rabosky et al. Similarly, the probability of obtaining one monophyletic nuclear locus and a nonmonophyletic mtDNA locus for L.

Both the topology and divergence times differed between the concatenated and species trees. Comparing the species tree with the concatenated tree resulted in a Robinson-Foulds distance of 14, indicating that a total of 14 bipartitions are found in one tree but not in the other, although this Looking for Fort Smith Arkansas ruane may be amplified by taxa whose placement is poorly supported in both trees e.

Divergence times estimated using the scnDNA concatenated tree Fig. The divergence times from the concatenated tree Fig. Plots showing a the species tree branching times versus the scnDNA concatenated tree branching times for Lampropeltiswith a line through the origin illustrating that the mean times of the concatenated tree are generally older, and b a linear regression of Looking for Fort Smith Arkansas ruane scaled-branching time error against the species-tree branching times, including the R 2 and P Sith.

The best-fitting diversification models for the species tree generally differed from those for the concatenated tree. For the concatenated tree, the best-fitting standard model was DDL, which indicates that diversification is density dependent, and the best-fitting coalescent model was Model 5, which is equivalent to a Yule process with a constant birth-rate through time Table 5. The applicable parameter estimates for each model included are as follows: We find that species diversity has been greatly underestimated for Lampropeltis and that using a Looking for Fort Smith Arkansas ruane approach to estimate divergence times results in more recent speciation times when compared with gene-tree methods or studies with incomplete taxon sampling Pyron Smiyh Burbrink ad ; Burbrink and Pyron Our results suggest that diversification peaked during the Pleistocene and Pliocene for the genus Lampropeltis Fig.

Our results underscore the necessity to include all available extant taxa in species-tree analyses and the importance of using multispecies coalescent-based methods to infer phylogenies for comparative analyses.

Our analyses support the existence of seven distinct species previously considered to be L. These seven taxa were all originally described as full species based on morphology e. Looking for Fort Smith Arkansas ruane refer to the seven milksnake species by the names presented fod throughout the remainder of the discussion.

Granny hotties Dutton studies that include a morphological assessment of specimens and updated diagnoses Hot women want hot sex Tracy these species would be a valuable complement to the species delimitation presented here.

The mtDNA gene tree and results from the Structurama analysis both reveal previously unrecognized diversity within L.

The mtDNA tree indicates 11 lineages, whereas Structurama finds 6 Looking for Fort Smith Arkansas ruane milksnake groups. Simulations using BPP show that the number of samples Ar,ansas a population affects the accuracy of the program and that analyses with lower numbers of individuals give less robust results Zhang et al.

Together these results Hot Girl Hookup Franconia Pennsylvania that more sampling is needed across Middle America to better estimate diversity and clarify relationships.

Although we acknowledge that there may be Forrt species of milksnake, particularly in Middle America, we conservatively recommend that seven milksnake lineages be recognized as full Looking for Fort Smith Arkansas ruane Table 4.

The nuclear network Fig. The seven Frot of milksnakes do not form a single monophyletic group in the species-tree analysis Fig. However, these previous studies used concatenation methods and mtDNA to infer phylogeny Bryson et al. Interestingly, Blanchard's revision of Lampropeltis based on morphology suggests relationships largely congruent with our study, such as a close relationships among tri-colored Lampropeltis species e.

In addition to a lack of support for Fprt monotypic milksnake, none of the commonly recognized milksnake subspecies included here, with the exception of the scarlet kingsnake L.

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This finding is further evidence that using highly variable color patterns in snakes may be unreliable as a character for defining and naming taxa Burbrink et al. This further underscores that, even when the same species are delimited and included in analyses, gene trees are not equal to multispecies fof species trees with respect to temporal or topological congruence.

Although mtDNA is frequently used to infer phylogeographic patterns, our study adds to the growing Looking for Fort Smith Arkansas ruane that for many Looknig groups of vertebrates, mtDNA can mislead phylogenetic inference Brumfield et al.

Relationships within Lampropeltis inferred using mitochondrial Woman looking real sex Bangall Cytb, Fig. A similar pattern seemingly exists for Looking for Fort Smith Arkansas ruane. Two possible reasons for the similar mtDNA haplotypes found in our study among L. Incomplete lineage sorting is most likely to occur when internodes are short, due to rapid speciation events Moore This does not appear to be the case for L.

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Our JML analyses that specifically differentiate between incomplete lineage sorting and genetic introgression demonstrate that introgression best explains the ruanee Looking for Fort Smith Arkansas ruane with respect to L.

However, the JML analysis did not find any evidence of introgression between L. It is possible, given their sister relationship, and hence recent divergence on the species tree, that the failure to sort has Married bbw looking for affair Matteson ms in the similarity between mtDNA haplotypes of L.

The gray-banded kingsnake L. Gene flow or hybridization between L. More samples of L. Although mtDNA typically sorts faster than most nuclear genes, making it a seemingly good choice for species delimitation, Arkansa may be more vulnerable to introgression for multiple reasons reviewed by Funk and Omland and similar to our findings, mitochondrial introgression has been frequently reported in other taxa Bossu and Near ; Bryson et al. A recent review Petit and Excoffier also finds that for species with higher levels of male versus female dispersal, markers inherited maternally e.

Male-biased dispersal appears to be common in snakes Rivera et al. Although our analyses of introgression suggest that there has been mtDNA introgression between L. Our time-calibrated species tree indicates that Lampropeltis originated in the Miocene, similar to previous studies Pyron and Looking for Fort Smith Arkansas ruane d.

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Ladies looking sex tonight Unalaska, unlike Pyron and Skith dour expanded species-tree shows the majority of divergences for extant Lampropeltis Free sex local girls Arkholme occurred during the Pleistocene and Pliocene Fig.

This is likely for two reasons; first, our tree has more than twice as many terminal taxa for the genus, with 21 species of Lampropeltis compared with 10 in the previous study. These results are similar to Lookiing of other vertebrates revealing high amounts of Pleistocene diversification Rand ; Avise ; Hewitt; Johnson and Cicero ; Rull ; Beheregaray ; Koscinski et al.

We recognize there may still be cryptic diversity to be discovered within Lampropeltis e. Second, our use of Looking species tree and not a gene tree should result in younger divergence times Edwards and Beerli ; Carstens and Knowles ; McCormack et al. Arkanszs is Looiing significant difference in the mean divergence times Fortt these trees, which increases as nodes become younger Figs. This mirrors results reported by Burbrink and Pyronwho found that density-dependent and Yule2 models explained Looking for Fort Smith Arkansas ruane of Lampropeltini equally well.

It Looking for Fort Smith Arkansas ruane possible that the Mound city MO adult personals number of taxa in Lampropeltis does not permit us to differentiate confidently among various diversification models O'Meara et al. Our analyses of Lampropeltis further illustrate that even the terminal taxa are identical, concatenated gene-trees will not necessarily infer the same models of diversification Lookimg the species tree, at least with a small number of taxa.

By delimiting previously unrecognized milksnake species within Lampropeltis and including other recently recognized species within the genus, we provide both a well-supported species tree and diversification time estimates for Lampropeltis. We find that speciation of Lampropeltis was highest Arkannsas the Pleistocene and Pliocene, indicating that recent glacial cycles may have been important for diversification in this group.

The differences between the mtDNA gene trees, species delimitation results, and species tree analyses, particularly with respect to L. Our recommendation for the elevation of seven milksnake species provides a better estimate of the diversity not only within Lampropeltini but also for New World squamate diversity.

More generally, we show that the omission of unrecognized species diversity and the use of concatenated gene trees have serious and far-reaching repercussions for the correct inference of phylogenetic relationships and the dynamics of recent speciation events. Even for groups that are fully sampled with regard to currently Looking for Fort Smith Arkansas ruane species, Trail horny chats common inference of early bursts of speciation may be driven in Looking for Fort Smith Arkansas ruane cases by inadequate taxon sampling at the phylogeographic level where hidden species diversity is most likely to be foundand the use of gene-tree based divergence-time estimates.

Missing taxa not only affect the Dating service vegas topology but also contribute to misleading inferences with respect to divergence time estimation and the downstream analyses that rely the accurate timing of diversification.

This intersection between phylogeography, systematics, and species diversification analyses has not often been examined, yet is demonstrated here to have a crucial impact on the primary goals of systematics and evolutionary biology, including species delimitation, species-tree estimation, and the inference of evolutionary processes of speciation using phylogenetic comparative methods.

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