Phylogenomic analysis of yellowjackets and hornets (Hymenoptera: Vespidae, Vespinae).

The phylogenetic relationships among genera of the subfamily Vespinae (yellowjackets and hornets) remain unclear. Yellowjackets and hornets constitute one of the only two lineages of highly eusocial wasps, and the distribution of key behavioral traits correlates closely with the current classification of the group. The potential of the Vespinae to elucidate the evolution of social life, however, remains limited due to ambiguous genus-level relationships. Here, we address the relationships among genera within the Vespinae using transcriptomic (RNA-seq) data. We sequenced the transcriptomes of six vespid wasps, including three of the four genera recognized in the Vespinae, combined our data with publicly available transcriptomes, and assembled two matrices comprising 1,507 and 3,356 putative single-copy genes. The results of our phylogenomic analyses recover Dolichovespula as more closely related to Vespa than to Vespula, therefore challenging the prevailing hypothesis of yellowjacket (Vespula+Dolichovespula) monophyly. This suggests that traits such as large colony size and high paternity arose in the genus Vespula following its early divergence from the remaining vespine genera.

Phylogenetic tests reject Emery's rule in the evolution of social parasitism in yellowjackets and hornets (Hymenoptera: Vespidae, Vespinae).

Social parasites exploit the brood-care behaviour and social structure of one or more host species. Within the social Hymenoptera there are different types of social parasitism. In its extreme form, species of obligate social parasites, or inquilines, do not have the worker caste and depend entirely on the workers of a host species to raise their reproductive offspring. The strict form of Emery's rule states that social parasites share immediate common ancestry with their hosts. Moreover, this rule has been linked with a sympatric origin of inquilines from their hosts. Here, we conduct phylogenetic analyses of yellowjackets and hornets based on 12 gene fragments and evaluate competing evolutionary scenarios to test Emery's rule. We find that inquilines, as well as facultative social parasites, are not the closest relatives of their hosts. Therefore, Emery's rule in its strict sense is rejected, suggesting that social parasites have not evolved sympatrically from their hosts in yellowjackets and hornets. However, the relaxed version of the rule is supported, as inquilines and their hosts belong to the same Dolichovespula clade. Furthermore, inquilinism has evolved only once in Dolichovespula.

Phylogeny and historical biogeography of the paper wasp genus Polistes (Hymenoptera: Vespidae): implications for the overwintering hypothesis of social evolution.

The phylogeny of the paper wasp genus Polistes is investigated using morphological and behavioural characters, as well as molecular data from six genes (COI, 12S, 16S, 28S, H3, and EF1-α). The results are used to investigate the following evolutionary hypotheses about the genus: (i) that Polistes first evolved in Southeast Asia, (ii) that dispersal to the New World occurred only once, and (iii) that long-term monogyny evolved as an adaptation to overwintering in a temperate climate. Optimization of distribution records on the recovered tree does not allow unambiguous reconstruction of the ancestral area of Polistes. While the results indicate that Polistes dispersed into the New World from Asia, South America is recovered as the ancestral area for all New World Polistes: Nearctic species groups evolved multiple times from this South American stock. The final tree topology suggests strongly that the genus first arose in a tropical environment, refuting the idea of monogyny as an overwintering adaptation.

Phylogenetic relationships of yellowjackets inferred from nine loci (Hymenoptera: Vespidae, Vespinae, Vespula and Dolichovespula).

Eusociality has arisen repeatedly and independently in the history of insects, often leading to evolutionary success and ecological dominance. Eusocial wasps of the genera Vespula and Dolichovespula, or yellowjackets, have developed advanced social traits in a relatively small number of species. The origin of traits such as effective paternity and colony size has been interpreted with reference to an established phylogenetic hypothesis that is based on phenotypic data, while the application of molecular evidence to phylogenetic analysis within yellowjackets has been limited. Here, we investigate the evolutionary history of yellowjackets on the basis of mitochondrial and nuclear markers (nuclear: 28S, EF1α, Pol II, and wg; mitochondrial: 12S, 16S, COI, COII, and Cytb). We use these data to test the monophyly of yellowjackets and species groups, and resolve species-level relationships within each genus using parsimony and Bayesian inference. Our results indicate that a yellowjacket clade is either weakly supported (parsimony) or rejected (Bayesian inference). However, the monophyly of each yellowjacket genus as well as species groups are strongly supported and concordant between methods. Our results agree with previous studies regarding the monophyly of the Vespula vulgaris group and the sister relationship between the V. rufa and V. squamosa groups. This suggests convergence of large colony size and high effective paternity in the vulgaris group and V. squamosa, or a single origin of both traits in the most recent common ancestor of all Vespula species and their evolutionary reversal in the rufa group.

Preservation of field samples for enzymatic and proteomic characterization: analysis of proteins from the trophallactic fluid of hornets and yellowjackets.

Proteomics is fast becoming one of the most interdisciplinary fields, bridging many chemical and biological disciplines. Major challenges, however, can limit the reach of proteomics to studies of model organisms. Challenges include the adequate preservation of field samples and the reliance of in-depth proteomics on sequenced genomes. Seeking to better establish the evolutionary relationships of hornets and yellowjackets comprising the subfamily Vespinae, we are combining classical morphological and genomic information with a functional genomics trait using proteomics. Vespine species form highly social colonies and exhibit division of labor in almost all aspects of colony life. An extreme digestive division of labor has been reported in Vespa orientalis, in which larvae but not adult workers exhibit the capacity to digest proteins fully. This makes the colony dependent upon the amino acid-rich trophallactic fluid released to adults by larvae and implies that the V. orientalis superorganism possesses larval-specific proteases. Identifying the proteases and the species exhibiting such extreme partitioning of digestive labor will allow for tracing the phylogenetic origins and elaboration of that digestive partitioning in the Vespinae. Herein we describe methods, generally applicable to field samples, showing the preservation of proteins and proteolytic activity from adult and larval vespine trophallactic fluid.

The conflation of ignorance and knowledge in the inference of clade posteriors.

The objective Bayesian approach relies on the construction of prior distributions that reflect ignorance. When topologies are considered equally probable a priori, clades cannot be. Shifting justifications have been offered for the use of uniform topological priors in Bayesian inference. These include: (i) topological priors do not inappropriately influence Bayesian inference when they are uniform; (ii) although clade priors are not uniform, their undesirable influence is negated by the likelihood function, even when data sets are small; and (iii) the influence of nonuniform clade priors is an appropriate reflection of knowledge. The first two justifications have been addressed previously: the first is false, and the second was found to be questionable. The third and most recent justification is inconsistent with the first two, and with the objective Bayesian philosophy itself. Thus, there has been no coherent justification for the use of nonflat clade priors in Bayesian phylogenetics. We discuss several solutions: (i) Bayesian inference can be abandoned in favour of other methods of phylogenetic inference; (ii) the objective Bayesian philosophy can be abandoned in favour of a subjective interpretation; (iii) the topology with the greatest posterior probability, which is also the tree of greatest marginal likelihood, can be accepted as optimal, with clade support estimated using other means; or (iv) a Bayes factor, which accounts for differences in priors among competing hypotheses, can be used to assess the weight of evidence in support of clades. ©The Willi Hennig Society 2009.

Topology-Bayes versus Clade-Bayes in phylogenetic analysis.

Several features of currently used Bayesian methods in phylogenetic analysis are discussed. The distinction between Clade-Bayes and Topology-Bayes is presented and illustrated with an empirical example. Three problems with Bayesian phylogenetic methods--exaggerated clade support, inconsistently biased priors, and the impossibility of hypothesis testing of cladograms--are shown to be the result of using a Clade-based Bayesian approach. Topology-based Bayesian methods do not share these shortcomings.

The new and improved PhyloCode, now with types, ranks, and even polyphyly: a conference report from the First International Phylogenetic Nomenclature Meeting.

A report from the first International Phylogenetic Nomenclature Meeting is presented. The meeting revealed that the PhyloCode, once implemented, will itself not require adherence to the three major tenets of philosophy that proponents have claimed required its creation. These include the abandonment of (1) non-monophyletic taxa, (2) ranks, and (3) types.

Parsimony overcomes statistical inconsistency with the addition of more data from the same gene.

Many authors have demonstrated that the parsimony method of phylogenetic analysis can fail to estimate phylogeny accurately under certain conditions when data follow a model that stipulates homogeneity of the evolutionary process. These demonstrations further show that no matter how much data are added, parsimony will forever exhibit this statistical inconsistency if the additional data have the same distributional properties as the original data. This final component-that the additional data must follow the same distribution as the original data-is crucial to the demonstration. Recent simulations show, however, that if data evolve heterogeneously, parsimony can perform consistently. Here we show, using natural data, that parsimony can overcome inconsistency if new data from the same gene are added to an analysis already exhibiting a condition indistinguishable from inconsistency.

Strange bayes indeed: uniform topological priors imply non-uniform clade priors.

While Bayesian analysis has become common in phylogenetics, the effects of topological prior probabilities on tree inference have not been investigated. In Bayesian analyses, the prior probability of topologies is almost always considered equal for all possible trees, and clade support is calculated from the majority rule consensus of the approximated posterior distribution of topologies. These uniform priors on tree topologies imply non-uniform prior probabilities of clades, which are dependent on the number of taxa in a clade as well as the number of taxa in the analysis. As such, uniform topological priors do not model ignorance with respect to clades. Here, we demonstrate that Bayesian clade support, bootstrap support, and jackknife support from 17 empirical studies are significantly and positively correlated with non-uniform clade priors resulting from uniform topological priors. Further, we demonstrate that this effect disappears for bootstrap and jackknife when data sets are free from character conflict, but remains pronounced for Bayesian clade supports, regardless of tree shape. Finally, we propose the use of a Bayes factor to account for the fact that uniform topological priors do not model ignorance with respect to clade probability.

How meaningful are Bayesian support values?

In this study, we used an empirical example based on 100 mitochondrial genomes from higher teleost fishes to compare the accuracy of parsimony-based jackknife values with Bayesian support values. Phylogenetic analyses of 366 partitions, using differential taxon and character sampling from the entire data matrix of 100 taxa and 7,990 characters, were performed for both phylogenetic methods. The tree topology and branch-support values from each partition were compared with the tree inferred from all taxa and characters. Using this approach, we quantified the accuracy of the branch-support values assigned by the jackknife and Bayesian methods, with respect to each of 15 basal clades. In comparing the jackknife and Bayesian methods, we found that (1) both measures of support differ significantly from an ideal support index; (2) the jackknife underestimated support values; (3) the Bayesian method consistently overestimated support; (4) the magnitude by which Bayesian values overestimate support exceeds the magnitude by which the jackknife underestimates support; and (5) both methods performed poorly when taxon sampling was increased and character sampling was not increases. These results indicate that (1) the higher Bayesian support values are inappropriate (in magnitude), and (2) Bayesian support values should not be interpreted as probabilities that clades are correctly resolved. We advocate the continued use of the relatively conservative bootstrap and jackknife approaches to estimating branch support rather than the more extreme overestimates provided by the Markov Chain Monte Carlo-based Bayesian methods.