Insects are the most speciose group of animals, but the phylogenetic relationships of many major lineages remain unresolved. We inferred the phylogeny of insects from 1478 protein-coding genes. Phylogenomic analyses of nucleotide and amino acid sequences, with site-specific nucleotide or domain-specific amino acid substitution models, produced statistically robust and congruent results resolving previously controversial phylogenetic relations hips. We dated the origin of insects to the Early Ordovician [~479 million years ago (Ma)], of insect flight to the Early Devonian (~406 Ma), of major extant lineages to the Mississippian (~345 Ma), and the major diversification of holometabolous insects to the Early Cretaceous. Our phylogenomic study provides a comprehensive reliable scaffold for future comparative analyses of evolutionary innovations among insects.

Hemipteroid insects (Paraneoptera), with over 10% of all known insect diversity, are a major component of terrestrial and aquatic ecosystems. Previous phylogenetic analyses have not consistently resolved the relationships among major hemipteroid lineages. We provide maximum likelihood-based phylogenomic analyses of a taxonomically comprehensive dataset comprising sequences of 2,395 single-copy, protein-coding genes for 193 samples of hemipteroid insects and outgroups. These analyses yield a well-supported phylogeny for hemipteroid insects. Monophyly of each of the three hemipteroid orders (Psocodea, Thysanoptera, and Hemiptera) is strongly supported, as are most relationships among suborders and families. Thysanoptera (thrips) is strongly supported as sister to Hemiptera. However, as in a recent large-scale analysis sampling all insect orders, trees from our data matrices support Psocodea (bark lice and parasitic lice) as the sister group to the holometabolous insects (those with complete metamorphosis). In contrast, four-cluster likelihood mapping of these data does not support this result. A molecular dating analysis using 23 fossil calibration points suggests hemipteroid insects began diversifying before the Carboniferous, over 365 million years ago. We also explore implications for understanding the timing of diversification, the evolution of morphological traits, and the evolution of mitochondrial genome organization. These results provide a phylogenetic framework for future studies of the group.

A major fraction of the diversity of insects is parasitic, as herbivores, parasitoids or vertebrate ectopara sites. Understanding this diversity requires information on the origin of parasitism in various insect groups. Parasitic lice (Phthiraptera) are the only major group of insects in which all members are permanent parasites of birds or mammals. Lice are classified into a single order but are thought to be closely related to, or derived from, book lice and bark lice (Psocoptera). Here, we use sequences of the nuclear 18S rDNA gene to investigate the relationships among Phthiraptera and Psocoptera and to identify the origins of parasitism in this group (termed Psocodea). Maximum-likelihood (ML), Bayesian ML and parsimony analyses of these data indicate that lice are embedded within the psocopteran infraorder Nanopsocetae, making the order Psocoptera paraphyletic (i.e. does not contain all descendants of a single common ancestor). Furthermore, one family of Psocoptera, Liposcelididae, is identified as the sister taxon to the louse suborder Amblycera, making parasitic lice (Phthiraptera) a polyphyletic order (i.e. descended from two separate ancestors). We infer from these results that parasitism of vertebrates arose twice independently within Psocodea, once in the common ancestor of Amblycera and once in the common ancestor of all other parasitic lice.

An approximate description of the zone of influence around the propagation path for a surface wave is provided by investigating the Fresnel zones for the frequency range of interest. The influence zone about surface wave paths, over which surface waves are coherent in phase, is identified as approximately one-third of the width of the first Fresnel zone. A technique called Fresnel-area ray tracing (FRT) for surface waves has been used to estimate this region around the ray path for each frequency. The FRT technique is developed by combining two standard ray tracing methods, i.e. kinematic ray tracing (KRT) and dynamic ray tracing (DRT). To obtain the exact Fresnel area in a laterally heterogeneous structure would require the solution of a large number of KRT equations. In contrast, the FRT approach requires just a few ray tracing calculations. In the first step, the trajectory of the surface wave is computed by solving the KRT system for the phase-velocity distribution at the required frequency. In the next step, the behaviour of rays in the zone surrounding the KRT path is calculated by solving the DRT system twice; once from the source to the receiver and once more from the receiver to the source along the same trajectory. Finally, combining the solutions of these ray tracing systems using paraxial ray theory, the Fresnel area around a central ray can be estimated. Using FRT, stationary-phase fields can be constructed around a central ray path in a laterally heterogeneous structure. The influence zone around the ray path is then estimated from the stationary-phase function with simple assumptions concerning the perturbed wavefield. The estimate of the influence zone can be efficiently calculated in laterally heterogeneous structure by using the FRT technique, and allows an extension of current methods of surface wave analysis, which have commonly been based on geometrical ray theory and on the approximation of great-circle propagation. This approach allows the treatment of finite-width rays as well as deviations in propagation from the great circle induced by moderate lateral heterogeneity as revealed by recent tomography models. Such finite-width rays should be of major benefit in enhancing ray-based surface wave tomography

[1] A three-stage inversion technique for surface wave tomography is applied to the Australian region. The inversion procedure consists of three independent processes. In the first stage, path-specific one-dimensional (1-D) shear velocity profiles are derived from multimode waveform inversion to provide dispersion information. The information from all paths is then combined to produce multimode phase speed maps as a function of frequency. The first version of these phase speed maps is derived from linearized inversion based on the assumption of surface wave propagation along great circle paths. Subsequently, the 2-D phase speed maps are updated by including ray tracing and finite frequency effects through the influence zone around the surface wave paths over which the phase is coherent. Finally, in the third stage the 3-D shear wave speed distribution is reconstructed from the set of updated multimode phase speed maps. This three-stage inversion of surface waves has significant benefits because it is possible to incorporate multimode dispersion, off-great circle propagation, and finite frequency effects for surface waves in a common framework. The final 3-D model, which includes the effects of ray bending and finite frequency, shows improvement in the definition of the model in regions with high gradients in shear velocity, such as near tectonic boundaries, especially in eastern Australia. Despite the natural smoothing imposed by considering the influence zone around the surface wave paths, the final models still require rapid change in shear wave properties in the neighborhood of the edge of the craton.

Summary Phylogenetic relationships among three paraneopteran clades (Psocodea, Hemiptera and Thysanoptera) were analysed based on the morphology of forewing base structure. Monophyly of Paraneoptera was supported by nine autapomorphies, monophyly of Condylognatha (= Thysanoptera + Hemiptera) by two autapo‐ morphies, monophyly of Thysanoptera by five autapomorphies and monophyly of Hemiptera by one autapomorphy. Thus, (Psocodea + (Thysanoptera + Hemiptera)) were proposed to be the phylogenetic relationships within Paraneoptera. A homoplastic similarity of the third axillary sclerite was observed between Thysanoptera and Heteroptera, and a possible evolutionary factor providing this homoplasy was discussed. The present analysis also suggested a monophyletic Auchenorrhyncha, and reduction of the proximal median plate was considered as an autapomorphy of this clade.

The mantle component of the Australian Seismological Reference Model (AuSREM) has been constructed from Australian-specific sources, primarily exploiting the wealth of seismic sources at regional distances around Australia recorded at portable and permanent stations on the continent. AuSREM is designed to bring together the existing information on Australia, from both body wave and surface wave studies and provide a synthesis in the form of a 3-D model that can provide the basis for future refinement. The model is grid based with a 0.5 • sampling in latitude and longitude, and is designed to be fully interpolable, so that properties can be extracted at any point.For the upper mantle the primary source of information comes from seismic surface wave tomography, supplemented by analysis of body wave arrivals and regional tomography which provide useful constraints on the relation between P-and S-wave speeds in the mantle lithosphere. A representative model has been developed to capture the features of mantle structure drawing on a range of studies. The mantle structure is represented by grid values at 25 km intervals in depth from 75 to 300 km. Shallower structure is linked to the AuSREM crust through the recent Moho depth model of Kennett et al., which exploits all available sources of seismological information. Below 300 km depth and in the surrounding area AuSREM is linked to the S40RTS model of Ritsema et al.

BackgroundThe gene composition, gene order and structure of the mitochondrial genome are remarkably stable across bilaterian animals. Lice (Insecta: Phthiraptera) are a major exception to this genomic stability in that the canonical single chromosome with 37 genes found in almost all other bilaterians has been lost in multiple lineages in favour of multiple, minicircular chromosomes with less than 37 genes on each chromosome.ResultsMinicircular mt genomes are found in six of the ten louse species examined to date and three types of minicircles were identified: heteroplasmic minicircles which coexist with full sized mt genomes (type 1); multigene chromosomes with short, simple control regions, we infer that the genome consists of several such chromosomes (type 2); and multiple, single to three gene chromosomes with large, complex control regions (type 3). Mapping minicircle types onto a phylogenetic tree of lice fails to show a pattern of their occurrence consistent with an evolutionary series of minicircle types. Analysis of the nuclear-encoded, mitochondrially-targetted genes inferred from the body louse, Pediculus, suggests that the loss of mitochondrial single-stranded binding protein (mtSSB) may be responsible for the presence of minicircles in at least species with the most derived type 3 minicircles (Pediculus, Damalinia).ConclusionsMinicircular mt genomes are common in lice and appear to have arisen multiple times within the group. Life history adaptive explanations which attribute minicircular mt genomes in lice to the adoption of blood-feeding in the Anoplura are not supported by this expanded data set as minicircles are found in multiple non-blood feeding louse groups but are not found in the blood-feeding genus Heterodoxus. In contrast, a mechanist explanation based on the loss of mtSSB suggests that minicircles may be selectively favoured due to the incapacity of the mt replisome to synthesize long replicative products without mtSSB and thus the loss of this gene lead to the formation of minicircles in lice.