An Evaluation of Genetic Distances for Use with Microsatellite Loci

Mutations of alleles at microsatellite loci tend to result in alleles with repeat scores similar to those of the alleles from which they were derived. Therefore the difference in repeat score between alleles carries information about the amount of time that has passed since they shared a common ancestral allele. This information is ignored by genetic distances based on the infinite alleles model. Here we develop a genetic distance based on the stepwise mutation model that includes allelic repeat score. We adapt earlier treatments of the stepwise mutation model to show analytically that the expectation of this distance is a linear function of time. We then use computer simulations to evaluate the overall reliability of this distance and to compare it with allele sharing and Nei's distance. We find that no distance is uniformly superior for all purposes, but that for phylogenetic reconstruction of taxa that are sufficiently diverged, our new distance is preferable.     

Molecular phylogeny of butterflies Parnassius glacialis and P. stubbendorfii at various localities in East Asia.

The phylogeny of butterflies, Parnassius stubbendorfii and P. glacialis, collected at various localities in the Japan archipelago and the eastern part of the Asian continent was analyzed using mitochondrial DNA sequences coding for NADH dehydrogenase subunit 5 (805 bp). The molecular phylogenetic trees revealed that P. glacialis and P. stubbendorfii diverged from a common ancestor, and then the populations inhabiting the Japan archipelago and the Asian continent diverged in each species. The reliability of these divergences was supported by high bootstrap values. The divergences within the Japan archipelago and within the Asian continent in each species were unclear because of low bootstrap values. The genetic distance and a rough time-estimation in the UPGMA tree suggest that the both populations of P. glacialis and P. stubbendorfii may have been isolated in the Japan archipelago at the early time (about 1.7-2.0 Mya) of the glacial period in the Pleistocene. The genetic distance between the Japanese and the continental subspecies may be large enough that they can be classified as different species, in comparison with the genetic distances among some other parnassian species.     

Phylogenetics beyond biology

Evolutionary processes have been described not only in biology but also for a wide range of human cultural activities including languages and law. In contrast to the evolution of DNA or protein sequences, the detailed mechanisms giving rise to the observed evolution-like processes are not or only partially known. The absence of a mechanistic model of evolution implies that it remains unknown how the distances between different taxa have to be quantified. Considering distortions of metric distances, we first show that poor choices of the distance measure can lead to incorrect phylogenetic trees. Based on the well-known fact that phylogenetic inference requires additive metrics, we then show that the correct phylogeny can be computed from a distance matrix \({\mathbf {D}}\) if there is a monotonic, subadditive function \(\zeta\) such that \(\zeta ^{-1}({\mathbf {D}})\) is additive. The required metric-preserving transformation \(\zeta\) can be computed as the solution of an optimization problem. This result shows that the problem of phylogeny reconstruction is well defined even if a detailed mechanistic model of the evolutionary process remains elusive.     

Flies on thistles: support for synchronous speciation?

Synchronous speciation of hosts and herbivorous insects predicts a congruent topology of host and insect phylogenies and similar evolutionary ages of host and insect taxa. To test these predictions for the specialized herbivorous fly genus Urophora (Diptera: Tephritidae), we used three different approaches. (i) We generated a phylogenetic tree of 11 European Urophora species from allozyme data and constructed a phylogeny of their hosts from published sources. Superimposing the Urophora tree on the host-plant tree we found no evidence for general congruence. (ii) We correlated genetic distances (Nei distances) of the host plants vs. the genetic distances of associated Urophora species. Overall, the relationship was not positive. Nevertheless, for some pairs of Urophora species and host plants genetic distances were in the same order of magnitude. (iii) We collected allozyme data for pairs of thistle taxa and pairs of herbivores on thistles together with independent time estimates. With these data we calibrated a molecular clock. There was a non-linear relationship between phylogenetic age and genetic distance, rendering the dating of deep events in thistle–insect evolution difficult. Nevertheless the derived molecular clock showed that the split of insect taxa lagged behind the split of hosts.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 775–783.     

PATTERNS OF SPECIATION IN DROSOPHILA

To investigate the time course of speciation, we gathered literature data on 119 pairs of closely related Drosophila species with known genetic distances, mating discrimination, strength of hybrid sterility and inviability, and geographic ranges. Because genetic distance is correlated with divergence time, these data provide a cross-section of taxa at different stages of speciation. Mating discrimination and the sterility or inviability of hybrids increase gradually with time. Hybrid sterility and inviability evolve at similar rates. Among allopatric species, mating discrimination and postzygotic isolation evolve at comparable rates, but among sympatric species strong mating discrimination appears well before severe sterility or inviability. This suggests that prezygotic reproductive isolation may be reinforced when allopatric taxa become sympatric. Analysis of the evolution of postzygotic isolation shows that recently diverged taxa usually produce sterile or inviable male but not female hybrids. Moreover, there is a large temporal gap between the evolution of male-limited and female hybrid sterility or inviability. This gap, which is predicted by recent theories about the genetics of speciation, explains the overwhelming preponderance of hybridizations yielding male-limited hybrid sterility or inviability (Haldane's rule).     

The ecology of differences: assessing community assembly with trait and evolutionary distances

Species enter and persist in local communities because of their ecological fit to local conditions, and recently, ecologists have moved from measuring diversity as species richness and evenness, to using measures that reflect species ecological differences. There are two principal approaches for quantifying species ecological differences: functional (trait‐based) and phylogenetic pairwise distances between species. Both approaches have produced new ecological insights, yet at the same time methodological issues and assumptions limit them. Traits and phylogeny may provide different, and perhaps complementary, information about species' differences. To adequately test assembly hypotheses, a framework integrating the information provided by traits and phylogenies is required. We propose an intuitive measure for combining functional and phylogenetic pairwise distances, which provides a useful way to assess how functional and phylogenetic distances contribute to understanding patterns of community assembly. Here, we show that both traits and phylogeny inform community assembly patterns in alpine plant communities across an elevation gradient, because they represent complementary information. Differences in historical selection pressures have produced variation in the strength of the trait‐phylogeny correlation, and as such, integrating traits and phylogeny can enhance the ability to detect assembly patterns across habitats or environmental gradients.     

A model of immunological distances in systematics

While immunological distances among taxa have had wide use in systematics, there has been some doubt about their utility because of the observed non-metricity of such distance matrices. A model is presented here relating observed immunological distance to the actual number of antigenic site differences between taxa. This model accounts for the observed departures of these distances from the metric condition of reciprocity and triangle inequality. Based upon the model, two procedures are suggested for the transformation of immunological distances to metric distances appropriate for phylogenetic analysis. The model implies that the usual scaling adjustments applied to the immunological distance matrix are inappropriate; however, the same transformation applied instead to an initial similarity matrix will solve a scaling problem. Non-reciprocity of the distances is shown to remain a problem independent of this initial scaling problem. It is suggested that further transformation of these re-scaled distances may be obtained through an extension of the ADCLUS procedure developed in psychology. This approach suggests a general strategy for a transformation to metric distances, given a particular model of non-metricity for the data.     

Distances that perfectly mislead

Given a collection of discrete characters (e.g., aligned DNA sites, gene adjacencies), a common measure of distance between taxa is the proportion of characters for which taxa have different character states. Tree reconstruction based on these (uncorrected) distances can be statistically inconsistent and can lead to trees different from those obtained using character-based methods such as maximum likelihood or maximum parsimony. However, in these cases the distance data often reveal their unreliability by some deviation from additivity, as indicated by conflicting support for more than one tree. We describe two results that show how uncorrected (and miscorrected) distance data can be simultaneously perfectly additive and misleading. First, multistate character data can be perfectly compatible and define one tree, and yet the uncorrected distances derived from these characters are perfectly treelike (and obey a molecular clock), only for a completely different tree. Second, under a Markov model of character evolution a similar phenomenon can occur; not only is there statistical inconsistency using uncorrected distances, but there is no evidence of this inconsistency because the distances look perfectly treelike (this does not occur in the classic two-parameter Felsenstein zone). We characterize precisely when uncorrected distances are additive on the true (and on a false) tree for four taxa. We also extend this result to a more general setting that applies to distances corrected according to an incorrect model.     

中国沙塘鳢属鱼类线粒体12S rRNA基因序列分析

沙塘鳢属鱼类为东亚特有的小型淡水经济鱼类,中国产沙塘鳢属鱼类分类问题长期存在争议。本文测定了中国产沙塘鳢属鱼类全部种类的线粒体12S rRNA基因部分序列,结合GenBank中下载的2种日本沙塘鳢属鱼类和塘鳢科鱼类同源序列,探讨中国产4种沙塘鳢属鱼类的物种有效性,分析沙塘鳢属鱼类的系统发育关系。作者所使用的同源序列长度为690bp,其中变异位点258个,简约信息位点201个,包括插入/缺失位点34个,转换/颠换平均值为3.0,表明12S rRNA基因是研究沙塘鳢属鱼类系统发育关系的合适分子标记。基于p-distance模型的6种沙塘鳢属鱼类种内遗传距离为0.000—0.024,种间遗传距离为0.058—0.064,支持暗色沙塘鳢和中华沙塘鳢为不同种,中国产沙塘鳢属鱼类包括中华沙塘鳢、河川沙塘鳢、海丰沙塘鳢、鸭绿江沙塘鳢4个种的观点;至于中国还有没有新的沙塘鳢属鱼类,尚有待进一步研究。系统发育分析表明海丰海塘鳢是河川沙塘鳢的姐妹群,暗色沙塘鳢与O.hikimius的亲缘关系最为密切,而同属其余类群之间的系统发育关系则由于自展数据支持率较低而尚不明确。中国产沙塘鳢和日本产沙塘鳢并未单独分群,推测沙塘鳢属鱼类的共同原始祖先可能广泛分布于中国、朝鲜和日本等东亚地区,约在4.9—6.5百万年前的上新世开始分化,系统发育过程比较适合离散假说。     

Fast computation of distance estimators

Background  

Some distance methods are among the most commonly used methods for reconstructing phylogenetic trees from sequence data. The input to a distance method is a distance matrix, containing estimated pairwise distances between all pairs of taxa. Distance methods themselves are often fast, e.g., the famous and popular Neighbor Joining (NJ) algorithm reconstructs a phylogeny of n taxa in time O(n ³). Unfortunately, the fastest practical algorithms known for Computing the distance matrix, from n sequences of length l, takes time proportional to l·n ². Since the sequence length typically is much larger than the number of taxa, the distance estimation is the bottleneck in phylogeny reconstruction. This bottleneck is especially apparent in reconstruction of large phylogenies or in applications where many trees have to be reconstructed, e.g., bootstrapping and genome wide applications.     

Prediction of heterosis in crosses between inbred lines of Drosophila melanogaster

Summary The aim of the experiment was to determine if the estimated genetic distance between two populations could be used to predict the amount of heterosis that would occur when they were crossed. Eight lines of known relatedness to each other were produced by eight generations of sib mating and sub-lining. This produced lines that varied in coefficient of coancestry from zero to 0.78. Fourteen reciprocal crosses of these lines were used to measure heterosis for larval viability and adult fecundity. Gene frequencies at six polymorphic enzyme loci were used to estimate the genetic distances between lines, which were then compared with the known degrees of coancestry. The estimated genetic differences were poorly correlated with the known coancestry coefficients (r=0.4), possibly due to the small number of loci typed. Also genetic distances were only about 1/3 of what was expected. Selection acting on blocks of genes linked to the enzyme loci probably prevented the expected increase in homozygosity. Coancestry coefficient was correlated with heterosis (r=0.44–0.71). This level of correlation implied differences in heterosis among parent lines with the same level of coancestry. This variability is expected if a small number of loci explain most of the heterosis. The average level of heterosis was less than expected after eight generations of sib mating. This is most likely due to selection opposing the increase in homozygosity caused by inbreeding. The combination of these two imperfect correlations resulted in no significant correlation between genetic distance estimated from markers and heterosis.     

phylin: an r package for phylogeographic interpolation

phylin is a package for the r programming environment which offers different methods to spatially interpolate genetic information from phylogeographic data. These interpolations can be used to predict the spatial occurrence of different lineages within a phylogeny using a modified method of kriging, which allows the usage of a genetic distance matrix to derive a model of spatial dependence. phylin improves the available methods to generate interpolated surfaces from a phylogenetic trees by assessing the autocorrelation structure of the genetic information, interpolating the genetic data based on a statistical model, estimating the uncertainty of the predictions and identifying lineage occurrence and contact zones probability without projection of pairwise genetic distances into mid‐points between sample locations. The package also includes methods to plot interpolation surfaces and provide summary tables from the generated data and models. We provide an example of the usefulness of this tool by inferring the spatial occurrence of distinct historical evolutionary lineages of the Lataste's viper (Vipera latastei Boscá, 1878) in the Iberian Peninsula and identifying potential contact areas. The maps of phylogenetic patterns obtained with these methods provide a spatial context to test hypotheses related to processes underlying the geographic distribution of genetic diversity and to inform conservation planning.     

Hierarchical components of genetic variation at a species boundary: population structure in two sympatric varieties of Lupinus microcarpus (Leguminosae)

Lupinus microcarpus is a self-compatible annual plant that forms a species complex of morphologically variable but indeterminate varieties. In order to examine the hypothesis that varieties of L. microcarpus comprise genetically differentiated and reproductively isolated species, populations of L. microcarpus var. horizontalis and var. densiflorus were sampled from an area of sympatry in central California and genotyped using six microsatellite loci. Bayesian clustering divided the total sample into two groups corresponding to the named varieties with extremely low levels of inferred coancestry. Similarly, maximum likelihood and distance methods for genetic assignment placed individuals in two nonoverlapping groups. Evidence for isolation by distance (IBD) within each variety was found at shorter distance classes, but varieties remained differentiated in sympatry. Furthermore, coalescent estimates of divergence time indicate separation within the past 950-5050 generations, with minimal gene flow after divergence. A four-level hierarchical analysis of molecular variance (amova) found significant levels of genetic differentiation among varieties (theta(P) = 0.292), populations within varieties (theta(S) = 0.449), subpopulations within populations (theta(SS) = 0.623), and individuals within subpopulations (f = 0.421); but the greatest degree of differentiation was at the subpopulation level. Although it is sometimes assumed that the magnitude of genetic differences (e.g. F(ST)) should be greater between species than among populations or subpopulations of the same species, shared ancestral polymorphism may lead to relatively low levels of differentiation at the species level, even as the stochastic effects of genetic drift generate higher levels of differentiation at lower hierarchical levels. These results suggest that L. microcarpus var. horizontalis and var. densiflorus are recently diverged yet reproductively isolated species, with high levels of inbreeding resulting from the combined effects of limited gene flow, demographic bottlenecks, and partial selfing in finite, geographically structured populations.     

Phylogenetic uncertainty revisited: Implications for ecological analyses

Ecologists and biogeographers usually rely on a single phylogenetic tree to study evolutionary processes that affect macroecological patterns. This approach ignores the fact that each phylogenetic tree is a hypothesis about the evolutionary history of a clade, and cannot be directly observed in nature. Also, trees often leave out many extant species, or include missing species as polytomies because of a lack of information on the relationship among taxa. Still, researchers usually do not quantify the effects of phylogenetic uncertainty in ecological analyses. We propose here a novel analytical strategy to maximize the use of incomplete phylogenetic information, while simultaneously accounting for several sources of phylogenetic uncertainty that may distort statistical inferences about evolutionary processes. We illustrate the approach using a clade‐wide analysis of the hummingbirds, evaluating how different sources of uncertainty affect several phylogenetic comparative analyses of trait evolution and biogeographic patterns. Although no statistical approximation can fully substitute for a complete and robust phylogeny, the method we describe and illustrate enables researchers to broaden the number of clades for which studies informed by evolutionary relationships are possible, while allowing the estimation and control of statistical error that arises from phylogenetic uncertainty. Software tools to carry out the necessary computations are offered.     

Age rank/clade rank metrics--sampling,taxonomy, and the meaning of "stratigraphic consistency"

Paleontologists frequently contrast clade rank (i.e., nodal or patristic distance from the base of a cladogram) with age rank (i.e., relative first known appearances of the analyzed taxa) to measure the degree of congruence between the estimated phylogeny and the fossil record. Although some potential biases of these methods have been examined (e.g., the effect of tree imbalance), other properties of age rank/clade rank (ARCR) comparisons have not been studied in detail. A basic premise of ARCR metrics is that outgroup taxa diverged earlier than ingroups and thus should first appear in older strata. For example, given phylogeny (A,(B,C)), then taxon A should be sampled before either taxon B or taxon C. We examine this premise in the context of (1) phylogenetic theory, (2) taxonomic practice, (3) sampling intensity (R), and (4) factors other than sampling intensity (including cladogram accuracy). Simulations combining clade evolution and sampling over time indicate a poor relationship between ARCR metrics and R when all taxa are apomorphy-based monophyletic groups. However, a good relationship exists when taxa are either stem-based monophyletic groups or if workers include taxa without a priori decisions about monophyly or paraphyly. These results are not surprising because cladograms predict the order in which lineages diverged (which applies to stem-based monophyletic taxa) and the order in which morphologic grades appeared (which applies to paraphyletic taxa relative to derived monophyletic groups). Other factors that increase ARCR metrics when the average R stays the same include high temporal variation in R, budding instead of bifurcating speciation patterns, low extinction rates, cladogram inaccuracy, and (to a much lesser extent) large clade size. These results suggest several plausible explanations for patterned differences in ARCR metrics among clades, thereby compromising their validity as measures of the quality of the fossil record.     

Phylogenetic relationships within the eared seals (Otariidae: Carnivora): implications for the historical biogeography of the family.

Phylogenetic relationships within the family Otariidae were investigated using two regions of the mitochondrial genome. A 360-bp region of the cytochrome b gene was employed for the primary phylogenetic analysis, while a 356-bp segment of the control region was used to enhance resolution of the terminal nodes. Traditional classification of the family into the subfamilies Arctocephalinae (fur seals) and Otariinae (sea lions) is not supported, with the fur seal Callorhinus ursinus having a basal relationship relative to the rest of the family. This is consistent with the fossil record which suggests that this genus diverged from the line leading to the remaining fur seals and sea lions about 6 million years ago (mya). There is also little evidence to support or refute the monophyly of sea lions. Four sea lion clades and five fur seal clades were observed, but relationships among these clades are unclear. Similar genetic divergences between the sea lion clades (D(a) = 0.054-0.078), as well as between the major Arctocephalus fur seal clades (D(a) = 0.040-0.069) suggest that these groups underwent periods of rapid radiation at about the time they diverged from each other. Rapid radiations of this type make the resolution of relationships between the resulting species difficult and indicate the requirement for additional molecular data from both nuclear and mitochondrial genes. The phylogenetic relationships within the family and the genetic distances among some taxa highlight inconsistencies in the current taxonomic classification of the family.     

基于18S rDNA遗传距离与GC含量对游走类纤毛虫的分子系统学研究

为了揭示游走类纤毛虫的系统发生,对寄生于淡水鱼类的车轮虫科中的6种车轮虫进行了18S rDNA的测序并获得了9个序列。采用了最大似然法(ML)与贝叶斯法(BI)对GenBank中所有游走类纤毛虫的18S rDNA序列进行了系统树的构建,并首次将SPSS与18S rDNA遗传距离结合分析了游走类纤毛虫的系统发生。研究结果进一步证实了车轮虫属(Trichodina)的非单系发生与小车轮虫属 (Trichodinella) 的有效性。此外,研究结合18S rDNA 的GC含量与遗传距离分析提出了游走类纤毛虫科属及种间新的鉴定依据: 18S rDNA 的GC含量可用于游走类纤毛虫的科属区分,且与游走类纤毛虫的分化密切相关; 18S rDNA的遗传距离在游走类纤毛虫的不同阶元中具有一定的阈值范围,即通常种内遗传距离阈值范围为0.000-0.005,属种间阈值范围为0.005-0.150,当遗传距离大于0.150时,则达到了科间水平。     

Phylogenetic signal and its decay in mitochondrial SSU and LSU rRNA gene fragments of Anisoptera

The phylogeny of Anisoptera, dragonflies in the strict sense, has proven to be notoriously difficult to resolve. Based on morphological characters, several recent publications dealing with the phylogeny of dragonflies proposed contradicting inter- and intrafamily relationships. We explored phylogenetic information content of mitochondrial large-subunit (LSU) and small-subunit (SSU) ribosomal gene fragments for these systematic problems. Starting at published universal primers, we developed primer sets suitable for amplifying large parts of the LSU and SSU rRNA genes within dragonflies. These fragments turned out to harbor sufficient phylogenetic information to satisfyingly resolve intrafamily relationships, but they contain insufficient phylogenetic structure to permit reliable conclusions about several interfamily relationships. We demonstrate that decay of phylogenetic signal progresses from intrafamily to interfamily to outgroup relationships and is correlated with an increase of genetic distances. As expected, signal decay is most pronounced in fast-changing sites. Additionally, base composition among fast-changing sites significantly deviates from the expected homogeneity. Homogeneity of base composition among all included taxa was restored only after removing fast-changing sites from the data set. The molecular data tentatively support interfamily relationships proposed by the most recent publication based on morphological characters of fossil and extant dragonflies.     

Optimum contribution selection in large general tree breeding populations with an application to Scots pine

Development of selection methods that optimises selection differential subject to a constraint on the increase of inbreeding (or coancestry) in a population is an important part of breeding programmes. One such method that has received much attention in animal breeding is the optimum contribution (OC) dynamic selection method. We implemented the OC algorithm and applied it to a diallel progeny trial of Pinus sylvestris L. (Scots pine) focussing on two traits (total tree height and stem diameter). The OC method resulted in a higher increase in genetic gain (8–30%) compared to the genetic gain achieved using standard restricted selection method at the same level of coancestry constraint. Genetic merit obtained at two different levels of restriction on coancestry showed that the benefit of OC was highest when restriction was strict. At the same level of genetic merit, OC decreased coancestry with 56 and 39% for diameter and height, respectively, compared to the level of coancestry obtained using unrestricted truncation selection. Inclusion of a dominance term in the statistical model resulted in changes in contribution rank of trees with 7 and 13% for diameter and height, respectively, compared to results achieved by using a pure additive model. However, the genetic gain was higher for the pure additive model than for the model including dominance for both traits.     

RNA SEQUENCE DATA IN PHYLOGENETIC RECONSTRUCTION: TESTING THE LIMITS OF ITS RESOLUTION

Abstract— 18S ribosomal RNA sequences from 11 echinoderms are analysed using parsimony to investigate phylogenetic relationships. Their estimated divergence limes range from less than 20 Ma to more than 550 Ma before present. Phylogenies based on 18S rRNA sequence data are compared with well-established morphological phylogenies to discover at what evolutionary distance the two approaches start to produce incongruent results. Three regions of the 18S rRNA molecule are analysed separately and together, and paired and unpaired sites are also treated separately and combined.
Results show that a parsimony analysis of sequence data produces reliable results only when taxa have diverged more recently than about 100 Ma. At greater evolutionary distances (up to 250 Ma), paired nucleotides produce more reliable results than unpaired, while paired and unpaired data combined produce intermediate results. All trees within about 1% of the most parsimonious solution ought to be accepted. Transversions give results almost as reliable as paired regions though there were relatively few informative sites. The relationships of echinoderm classes, which diverged 450–550 Ma ago, are unresolved by 18S rRNA data.