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Lice (Insecta: Phthiraptera) have long been considered to compose a monophyletic group of insects on the basis of external morphological character- istics. However, a recent phylogenetic analysis of 18S rDNA sequences suggested that 'Phthiraptera' have arisen twice within the order Psocoptera (booklice and barklice). The external features of lice are highly specialized to a parasitic lifestyle, and convergence may be frequent for such characters. To provide a further test between traditional and recent molecular-based phylogenetic hypotheses, a phy- logenetic analysis of lice and relatives based on morphological characters that are independent from the selective pressures of a parasitic lifestyle is needed. Here, we examined the morphology of the male phallic organ in lice and relatives ('Psocoptera': suborders Troctomorpha and Psocomorpha) and detected some novel modifications that were stable within each group and useful for higher level phylogenetic reconstruction. Phylogenetic analysis based on these characters provided a concordant result with the 18S-based phylogeny. In particular, the apomorphic presence of articulations between the basal plate, mesomere and ventral plate (! sclerite on the permanently everted endophallus) is observed consistently throughout the psocid families Pachytroctidae and Liposcelididae and the louse suborder Amblycera, providing support for a clade composed of these three groups, although possible homoplasy was detected in some Ischnocera. This is the first study to provide morphological support for the polyphyly of lice.

The order Zoraptera (angel insects) is one of the least known insect groups, containing only 32 extant species. The phylogenetic position of Zoraptera is poorly understood, but it is generally thought to be closely related to either Paraneoptera (hemipteroid orders: booklice, lice, thrips, and bugs), Dictyoptera (blattoid orders: cockroaches, termites, and mantis), or Embioptera (web spinners). We inferred the phylogenetic position of Zoraptera by analyzing nuclear 18S rDNA sequences, which we aligned according to a secondary structure model. Maximum likelihood and Bayesian analyses both supported a close relationship between Zoraptera and Dictyoptera with relatively high posterior probability. The 18S sequences of Zoraptera exhibited several unusual properties: (1) a dramatically increased substitution rate, which resulted in very long branches; (2) long insertions at helix E23; and (3) modifications of secondary structures at helices 12 and 18.

SUMMARY Sensitivity kernels for fundamental mode surface waves at finite frequency for 2-D phase speed and 3-D shear wave speed are constructed based on the Born and Rytov approximations working with a potential representation for surface waves. The use of asymptotic Green’s functions for scalar wave equations provides an efficient way to calculate the Born or Rytov kernels. The 2-D sensitivity kernels enable us to incorporate the finite-frequency effects of surface waves, as well as off-great-circle propagation, in tomographic inversions for phasespeed structures. We derive examples of the 2-D sensitivity kernels both for a homogeneous background model (or a spherically symmetric model), and for a laterally heterogeneous model. The resulting distortions of the shape of the sensitivity kernels for a heterogeneous background model indicate the importance of the use of proper kernels to account of the heterogeneity in the real Earth. By combining a set of 2-D sensitivity kernels with 1-D vertical sensitivity kernels for a particular frequency range and taking the inverse Fourier transform, we can derive 3-D sensitivity kernels for surface waves in the time domain. Such 3-D kernels are useful for efficient forward modelling of surface waveforms incorporating finite-frequency effects, and will also enable us to perform direct inversion of surface waveforms into 3-D structure taking account of finite-frequency effects.

We derive estimates of temperature of the Australian continental mantle between 80 and 350 km depth from two published S-velocity models. Lithospheric temperatures range over about 1000 °C, with a large-scale correlation between temperature and tectonic age. In detail however, variations ranging from 200 to 700 °C occur within each tectonic province. At the current seismic resolution, strictly Proterozoic and Archean blocks do not have substantially different temperatures, nor does the Phanerozoic lithosphere east and west of the Tasman line. Temperatures close to an average (moist) MORB source mantle solidus characterize the eastern seaboard and its offshore. Differences between the temperatures derived from the two velocity models illustrate the importance of well-constrained absolute velocities and gradients for physical interpretation. The large range of lithospheric temperatures cannot be explained solely with documented variability in crustal heat production, but requires significant variations in mantle heat flow as well.

The genus Podopterocus Banks has been known only from male specimens and has been characterized by the thickened first antennal flagellomere and the expanded hind tibia. The genus Dinopsocus Banks has been characterized by the thickened first antennal flagellomere. Detailed morphological examinations and brief molecular-based identification (only for one species) suggest that both taxa are congeneric. Based on characters of female terminalia and fore wing venation, both taxa are furthermore considered to be closely related to the genus Sigmatoneura Enderlein. The discovery of two new species and observations on Sigmatoneura spp. reveal the limited significance of the diagnostic characters of Podopterocus and Dinopsocus. Therefore we consider both Podopterocus and Dinopsocus as junior synonyms of Sigmatoneura. Monophyly of the genus Sigmatoneura, including Podopterocus, Dinopsocus, and the subgenus Longifolia Li, is well supported by the unique fore wing venation of females and by sexually dimorphic fore wing venation and coloration. Two new species, Sigmatoneura kakisayap sp. n. and Sigmatoneura lemahsayap sp. n., which would be classified under Podopterocus or Dinopsocus by the previous generic definitions, are described. Sigmatoneura longicornis comb. n., the type species of Podopterocus, is redescribed and transferred to Sigmatoneura and the female of this species is described for the first time. Dinopsocus atratus, the type species of Dinopsocus, is synonymized with S. longicornis. Sigmatoneura semicolorata comb. n. is redescribed and transferred from Dinopsocus to Sismatoneura.

Phylogenetic relationships among species groups of Trichadenotecnum were inferred based on morphology and the partial sequences of five gene regions (mitochondrial 12S rDNA, 16S rDNA, cytochrome oxidase I, NADH dehydro- genase subunit 5 and nuclear 18S rDNA). All analyses supported the monophyly of Trichadenotecnum and all previously proposed species groups, except that T. circularoides was excluded from the spiniserrulum group. To examine the phylogenetic usefulness of morphological data, the morphological characters used in the construction of an earlier taxonomic system for Trichadenotecnum were mapped parsimoniously on the molecular tree. As a result: (1) commonly used forewing marking features (sparsely or extensively spotted) are considered to be very homoplastic and less informative of higher-level phylogenetic relation- ships; (2) a broadly expanded epiproct lobe is considered to be independently evolved at least two or three times, and a detailed morphological re-examination allows recognition of these convergent structures; (3) the short ventral valve of gonapophyses independently evolved at least three or four times, although this character was used initially to diagnose the spiniserrulum group.

Two new Trichadenotecnum species, T. corniculum and T. germinatum , were described from Japan. These species were considered to compose a monophyletic group newly defined here as the corniculum group. Judging from morphology of male terminalia, the corniculum group was considered to represent the basal most clade of the genus Trichadenotecnum .

SUMMARY 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.

SUMMARY A new technique for multimode dispersion measurement has been developed by employing fully non-linear waveform inversion for a path-specific 1-D profile using a neighbourhood algorithm (NA). One-dimensional models derived from waveform inversion are quite sensitive to the model parametrization and the reference model used to start the inversion. With different approaches to the parametrization of the shear wave speed profile, we can find models with significant differences in velocity variation with depth, which provide similar levels of fit to the observed waveforms. Although the models differ, the calculated phase dispersion for the first few modes of the surface waves are very close indeed. We therefore regard the 1-D models derived from the multimode waveform inversion as an implicit description of the path-specific dispersion for each of the modes. The inversion procedure using NA samples a substantial number of models and we select the 1-D wave speed model that achieves minimum misfit. Phase speeds for each mode branch are then calculated from the model parameters with the minimum misfit. Because we use the 1-D models as a summary of the average multimode dispersion along the path, we do not need to make assumptions concerning the nature of anisotropy. We can employ simple isotropic models and the method can be applied to both Rayleigh and Love waves independently. The new method is applied to sets of paths in the Australian region and provides stable measurements of multimode dispersion from a single seismogram. The multimode phase speeds measured using the NA inversion can be used to retrieve phase velocity maps as a function of frequency for higher-mode branches as well as the fundamental mode, which will be crucial constraints on 3-D Earth models.

SUMMARY A three-stage inversion scheme for surface wave tomography working with multimode phase dispersion as a function of frequency provides a means of combining a wide range of data in a common framework. The phase average approximation is applied directly to phase slowness and there is no need to invoke perturbation arguments for the interpretation of path-averaged velocity models derived from waveform inversion of surface waves. By treating such wave speed profiles as summaries of path specific dispersion behaviour it is possible not only to combine results from different style of inversion but also to provide maximum exploitation of Love and Rayleigh wave information. Inversions of all suitable waveforms can be undertaken in terms of isotropic models. Dispersion information from all paths is combined to form multimode phase speed distributions as a function of frequency in linearized inversion which takes account of path bending and finite frequency effects. The final inversion for 3-D wave speed structure is based on a cellular inversion of the multimode frequency dispersion including angular effects in terms of a local stratified model including anisotropy. The smoothing from inclusion of finite frequency effects and damping of the linearized inversion for the phase speed distributions will control the smoothness of the 3-D shear wave speed model.

Species of the bark louse genus Trichadenotecnum Enderlein (Insecta: Psocodea) from Peninsular Malaysia and Singapore are revised with illustrations and identification keys. Twenty species are here recognised, with four new species and ten recorded for the first time from this region, together with an unnamed species represented by a single female. The previously described species T. marginatum NewThornton is not included because its generic assignment is questionable. Females of T. cinnamonum EndangNew, T. imrum NewThornton and T. sibolangitense Endang, ThorntonNew, and the male of T. kerinciense EndangNew are described for the first time. A new species group is defined for T. krucilense Endang, ThorntonNew.

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 resolving power of polarization data compared with that of phase data is investigated by employing both synthetic and observed data sets, using the linear relationship between the phase velocity perturbation and the phase or polarization anomaly. In order to investigate the intrinsic differences between phase and polarization data, a synthetic test is first undertaken using a white noise model with sufficiently uniform coverage of ray paths. This test shows that polarization data can retrieve higher-order heterogeneities of degrees up to 20 almost completely, despite damping and smoothing effects, whilst phase data can only retrieve those of degrees lower than 8 with reasonable damping. Next, about 4000 phase and 2500 polarization records are collected for minor- and major-arc Rayleigh waves (R1 and R2) in the frequency range 4–12 mHz. To correct the Rayleigh wave polarization data, the misorientation of each station is estimated from the polarization of long-period P waves propagating mainly in the lower mantle. The phase and polarization data are then inverted for the phase velocity distribution in spherical harmonics with degrees up to 15. The phase velocity maps derived from the phase data are quite consistent with previous studies, whilst those from the polarization data show some discrepancies. For example, the correlation between the phase and polarization models is quite good for low even degrees such as 2, 4 and 6, but not for low odd degrees or degrees higher than 8. The gradients of ampli-tude spectra from the polarization data are smaller than those from the phase data, especially at degrees higher than 6, which suggests a slightly higher sensitivity of the polarization data to higher-order heterogeneities. Nevertheless, the overall spectral characteristics of both models are similar; that is, low-order heterogeneities are dominant whilst higher orders are clearly reduced. Further investigation using a synthetic test with the same uneven paths as the observed data shows the suppression of higher-order heterogeneities. Since the synthetic test with even paths retrieves higher-order heterogeneities sufficiently, this result strongly suggests the path averaging effect of uneven ray paths that is intrinsic in the ray theoretical approach used in this study as well as almost all the global inversions. Although inversions based on geometrical ray theory have some limited resolving power with the current status of global records, polarization data are indeed helpful in resolving higher-order lateral heterogeneities with the dense and uniform path coverage that is becoming available.

A new genus, Atrichadenotecnum, is described in the tribe Psocini of the family Psocidae (Psocoptera). Atrichadenotecnum ryukyuense sp. n., A. tayal sp. n., and A. quadripunctatum sp. n. are described and Atrichadenotecnum nudum (Thornton, 1961) comb. n. and Atrichadenotecnum quinquepunctatum (McLachlan, 1872) comb. n. are transferred from the genus Trichadenotecnum Enderlein. The new genus is regarded as the sister-group of the genus Psocus Latreille.

A new genus, Idatenopsocus, represented by Idatenopsocus orientalis (Vishnyakova, 1986), comb, nov., is described and the species is redescribed. Within the family Mesopsocidae, this new genus is considered as the sister group of a monophyletic group composed of the genera Mesopsocus, Microtrichipsocus, Metapsocus, Cyrtopsochus, Hexacyrtoma, and Rhinopsocus.

Extreme elongation of a part of the intromittent organ, the flagellum, has occurred several times in Criocerinae (Chrysomelidae). These leaf beetles have acquired a specialized pocket to store the flagellum in the abdominal cavity, at the same time allowing a quick control of movements of this structure during copulation. We investigated the morphogenesis of the intromittent organs of species with and without a flagellum to discuss the evolutionary background of parallel evolution of novel structures. We found that the specialized pocket is formed by the invagination of an epidermal layer and a resultant rotation of the primary gonopore. Invagination itself is a well-known phenomenon in morphogenetic processes, which leads us to hypothesize that the novelty is formed by co-opting a previously acquired genetic system. A large open-space is present within the intromittent organ during the entire morphogenesis in species without a flagellum, and the invagination in the species with a flagellum grows in the corresponding area. This means that there are no physical impediments for the growth of a large pocket. In addition the sites of muscular attachments in the species with a flagellum are also different from those without it. The differentiation of muscles is completed immediately before adult emergence, which means the muscles are adjustable during the entire morphogenesis in this group. Simple modifications probably based on a co-option of previously acquired genetic systems, the potential space for adding a new element, and an adjustable factor in morphogenesis of the intromittent organ facilitate the parallel evolution of the extreme elongation.

Acercaria display an unusually broad array of adhesive devices occurring on different parts of the legs. Attachment structures of all major subgroups are described and illustrated. Nineteen characters of the distal leg region were combined with a data matrix containing 99 additional morphological characters of different body parts. The results of the cladistic analysis are largely congruent with current hypotheses. Zoraptera are not retrieved as close relatives of Acercaria. The monophyly of the entire lineage and of the major subgroups Psocodea, Phthiraptera, and Hemiptera is confirmed. Our data also support the monophyly of Auchenorrhycha and a sister-group relationship between Thysanoptera and Hemiptera (Condylognatha). In contrast to other lineages of insects, the hairy type of adhesive device is present only in one group within the Acercaria (Heteroptera, Cimicomorpha). The arolium is present in the groundplan but missing in several groups (e.g. Psocodea, Cicadoidea, Aphidoidea). Pretarsal pulvilli evolved several times independently. Tarsal euplantulae and different specialized clasping devices have evolved within Phthiraptera, whereas pretarsal attachment devices are missing in this ectoparasitic group. The potential to modify pretarsal attachment devices in their structural details has probably contributed to the very successful diversification of the predominantly phytophagous Hemiptera.

The squirrel Callosciurus erythraeus (Pallas) (Rodentia: Sciuridae) was intentionally introduced to Japan in 1935 and has become established throughout much of the country. Although they live mainly in forests, Pallas squirrels come into gardens and are frequently fed by people or kept as pets, so their ectoparasites could be of potential medical as well as veterinary importance. During 2001-2003 we conducted the first ectoparasite survey of Pallas squirrels in Japan. From 105 C. erythraeus captured in Kamakura District of Kanagawa Prefecture on Honshu Island, three types of ectoparasite were found: 52 specimens of the sucking louse Neohaematopinus callosciuri Johnson (Anoplura: Haematopinidae), 26 fleas Ceratophyllus (Monopsyllus) anisus Rothschild (Siphonaptera: Ceratophyllidae) and four nymphs of the tick Haemaphysalis flava Neumann (Acari: Ixodidae) on 22, 13 and one squirrels, respectively. Evidently in Japan C. erythraeus carries relatively few ectoparasite species; this may be a contributory factor to their invasive success. Further investigations are needed to assess risks of zoonotic transmission of plague or murine typhus by C. anisus, of louse-borne typhus by N. callosciuri and of tularaemia and especially Japanese spotted fever (Rickettsia japonica) by H. flava.

A review of species of the genus Neopsocopsis Badonnel, 1936 is presented. Four species are redescribed, viz. N. hirticornis (Reuter, 1893), N. quinquedentata (Li & Yang, 1988), N. profunda (Li, 1995), and N. flavida (Li, 1989), as well as the description of one new species, N. convexa sp. n. Seven new synonymies are proposed as follows: Pentablaste obconica Li syn. n. and P. clavata Li syn. n. of N. hirticornis, P. tetraedrica Li syn. n. of N. longiptera, Neoblaste schizopetala Li syn. n. and Nb. flavae Li syn. n. of N. profunda, Blastopsocidus pini Li syn. n. and P. lanceolata Li syn. n. of N. flavida. N. hirticornis (Reuter, 1893) is recorded from Japan and China for the first time, and N. longiptera Vishnyakova, 1986 is newly recorded from China. Illustrated keys to adult males and females are presented.

For modern lineages of birds and mammals, few fossils have been found that predate the Cretaceous–Palaeogene (K–Pg) boundary. However, molecular studies using fossil calibrations have shown that many of these lineages existed at that time. Both birds and mammals are parasitized by obligate ectoparasitic lice (Insecta: Phthiraptera), which have shared a long coevolutionary history with their hosts. Evaluating whether many lineages of lice passed through the K–Pg boundary would provide insight into the radiation of their hosts. Using molecular dating techniques, we demonstrate that the major louse suborders began to radiate before the K–Pg boundary. These data lend support to a Cretaceous diversification of many modern bird and mammal lineages.

We investigated the utility of indel data for genealogical and population genetic analyses using the Tpi intron of the leaf mining moth Acrocercops transecta (Insecta: Lepidoptera). Genealogical analyses revealed that indel data were less homoplasious than DNA sequence data and that indel data contained a sufficient signal to provide a high resolution tree that was highly congruent with the tree estimated from DNA sequences. Although some conflicts were identified in the distributions of multi-residue indels, such conflicts were especially useful for the unambiguous detection of recombinations. For the first time, we adopted a Bayesian clustering method for indel characters to infer genetic structure of the moth. We concluded that indel characters have the potential to be a powerful tool in the analysis of population genetics and population structure as well as in the detection of gene flow.

Recently diverged populations often exhibit incomplete reproductive isolation, with a low level of gene flow continuing between populations. Previous studies have shown that, even under a low level of gene flow, genetic divergence between populations can proceed at the loci governing local adaptation and reproductive isolation but not at other neutral loci. A leaf-mining moth, Acrocercops transecta, consists of Juglans- and Lyonia-associated host races. The two host races differ in host preferences of ovipositing females and in larval adaptation to host plants but mate readily in the laboratory, producing fertile hybrids. The Juglans and Lyonia races are often sympatric in the wild, implying that gene introgression could occur in nature between the two host races. We tested this hypothesis by combining phylogenetic analyses with coalescent simulations, focusing on mitochondrial genes (COI and ND5) and the nuclear Tpi, Per and Ldh genes located on the Z-chromosome. The mitochondrial genes clearly distinguished the Lyonia race from the Juglnas race, whereas the Tpi, Per and Ldh genealogies did not reflect the two host races. Coalescent simulations indicated gene flow at the three Z-linked genes in both directions, whereas there was no introgression in the mitochondrial genes. The lack of introgression in mitochondrial genes suggests that female host preference is the primary force leading to the bifurcation of maternally inherited loci. Thus, the results show that a low level of gene flow coupled with the inflexible female host preference differentiates histories of divergence between maternally and biparentally inherited genes in this host race system.

[1] A series of steps in the lithospheric thickness of eastern Australia are revealed by the latest seismic surface wave tomographic model and calculations of the horizontal gradient of shear wave speed. The new images incorporate data from the recent Tasmal experiment, improving resolution in continental Australia. Through comparisons with surface geology and geochemical studies, it is possible to infer that the steps in lithospheric thickness are related to boundaries between blocks of different age. The westernmost boundary marks the edge of the Archaean to Early-Proterozoic core of the continent. A second lithospheric boundary is observed in the central part of east Australia. To the west of this line, geochemical evidence suggests that there is Proterozoic lithospheric mantle, and this boundary may therefore represent the change from Proterozoic to Phanerozoic basement. The structure on the eastern margin of the continent is dominated by slow velocities, suggesting that in this area the continental lithosphere is very thin. There is a strong correlation between the slow wave speeds and the location of both the highest topography and recent volcanic activity. Inland of the continental margin, a zone of strong gradients in the seismic wave speed is observed, indicating a distinct step in lithospheric structure. If the step in lithospheric thickness was in place prior to volcanism, it may have acted as a boundary, with volcanism mainly occurring beneath the thinner lithosphere to the east.

We revised the homology of wing base structure in Ephemeroptera (Insecta: Pterygota) proposed by Willkommen and Hörnschemeyer in a recent issue of Arthropod Structure and Development. The first free sclerite (s1) in Ephemeroptera should be homologized with a part of the first axillary sclerite (1Ax) of Neoptera, together with the second free sclerite, whereas the authors recognized s1 as a detached part of the anterior notal wing process. The fifth free sclerite of Ephemeroptera should be homologized with the median notal wing process of Neoptera, rather than it being homologous with a part of 1Ax in Neoptera, as the authors postulated. Hypothesized secondary fusion of the axillary sclerites in Ephemeroptera and Odonata proposed by the authors is premature, because the basal phylogeny of Pterygota is still poorly understood, and an alternative interpretation of morphological evolution (i.e., that undifferentiated axillary sclerites represent the ground plan of Pterygota) can also be drawn from the Ephemeroptera + Neoptera hypothesis.

Background Maternal care (egg-nymph guarding behavior) has been recorded in some genera of Acanthosomatidae. However, the origin of the maternal care in the family has remained unclear due to the lack of phylogenetic hypotheses. Another reproductive mode is found in non-caring species whose females smear their eggs before leaving them. They possess pairs of complex organs on the abdominal venter called Pendergrast’s organ (PO) and spread the secretion of this organ onto each egg with their hind legs, which is supposed to provide a protective function against enemies. Some authors claim that the absence of PO may be associated with the presence of maternal care. No study, however, has tested this hypothesis of a correlated evolution between the two traits. Results We reconstructed the molecular phylogeny of the subfamily Acanthosomatinae using five genetic markers sequenced from 44 species and one subspecies with and without maternal care. Eight additional species from the other two acanthosomatid subfamilies were included as outgroups. Our results indicated that maternal care has evolved independently at least three times within Acanthosomatinae and once in the outgroup species. Statistical tests for correlated evolution showed that the presence of maternal care is significantly correlated with the secondary loss or reduction of PO. Ancestral state reconstruction for the node of Acanthosoma denticaudum (a non-caring species in which egg smearing with developed POs occurs) and A. firmatum (a caring species with reduced POs) suggested egg smearing was still present in their most recent common ancestor and that maternal care in A. firmatum has evolved relatively recently. Conclusions We showed that maternal care is an apomorphic trait that has arisen multiple times from the presence of PO within the subfamily Acanthosomatinae. The acquisition of maternal care is correlated with the reduction or loss of PO, which suggests an evolutionary trade-off between the two traits resulting from physiological costs. This prediction also implies that presence of maternal care can be highly expected for those groups lacking behavioral data, which invariably also lack the organ. No secondary loss of maternal care was detected in the present tree. We suggest that the loss of maternal care may be suppressed due to the vulnerability of the PO-free condition, which thus maintains maternal care.

[1] We obtained three-dimensional (3-D) shear wave speed structure beneath the Philippine Sea and the surrounding region from seismograms recorded by land-based and long-term broadband ocean bottom seismographic stations. The ocean bottom data gave us a better station coverage to obtain a higher spatial resolution (about 300–400 km) in the Philippine Sea than in previous studies. We employed a new technique of surface wave tomography, in which multimode phase speeds are measured and inverted for a 3-D shear wave speed structure by incorporating the effects of finite frequency and ray bending. There is a sharp speed contrast along the Izu-Bonin-Mariana trench, across which the Philippine Sea side has a significantly slower upper mantle than the Pacific Ocean side. In the upper 120 km, the shear wave speed structure is well correlated with the age of the provinces. At depths greater than 160 km, the pattern is dominated by fast anomalies of the subducted slabs of the Pacific plate and two slow anomalies to the south of the Daito ridge and in the southernmost part of the Philippine Sea.

[1] We determined three-dimensional shear wave speed structure beneath the South Pacific superswell down to a depth of 200 km by analyzing Rayleigh wave records from broadband ocean bottom seismograph stations and island stations in the Pacific Ocean. The ocean bottom stations were deployed from 2003 to 2005 on the seafloor in the French Polynesian region, which enabled us to study the upper mantle structure beneath the Superswell with unprecedentedly high resolution. We measured the dispersions of fundamental mode Rayleigh waves at periods between 40 and 140 seconds by the two-station method. We found pronounced slow anomalies near the hot spots and in the Lau Basin. The slow anomalies beneath the hot spots have deeper-rooted than those associated with the Lau basin. The slow anomalies near the Society, Macdonald, Marquesas, and Pitcairn hot spots continue down to at least 200 km depth.

Insect herbivores such as gall formers and leaf miners are often highly specialized and adapted to their respective natal host plants. Due to the specialization and adaptation, it is presumed that host shifts readily occur among closely related plant species. Leaf-mining moths, the Acrocercops leucophaea complex, consist of three species, A. leucophaea, A. defigurata, and A. transecta. Larvae of all the species of the complex feed on Juglandaceae plants, but A. leucophaea and A. transecta are also associated with an Ericaceae plant, which is quite distantly related to Juglandaceae. Such a host utilization as in this species complex is very rare among phytophagous insects. In the present study, we estimate the history of host shifts by reconstructing the phylogeny of the A. leucophaea complex using molecular data (partial sequence of mitochondrial COI, 12S rDNA, and ND5). Parsimony and maximum likelihood analyses indicated that the common ancestor of the A. leucophaea complex used Juglandaceae only, and that the association with Ericaceae has evolved in A. leucophaea and A. transecta independently. Parametric bootstrap analysis also supported multiple origins of the association with Ericaceae in this complex. These results imply that there are ecological and biochemical factors that promote host shifting between Juglandaceae and Ericaceae despite the two families being not closely related.

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.

Phthiraptera (chewing and sucking lice) and Psocoptera (booklice and barklice) are closely related to each other and compose the monophyletic taxon Psocodea. However, there are two hypotheses regarding their phylogenetic relationship: (1) monophyletic Psocoptera is the sister group of Phthiraptera or (2) Psocoptera is paraphyletic, and Liposcelididae of Psocoptera is the sister group of Phthiraptera. Each hypothesis is supported morphologically and/or embryologically, and this problem has not yet been resolved. In the present study, the phylogenetic position of Phthiraptera was examined using mitochondrial 12S and 16S rDNA sequences, with three methods of phylogenetic analysis. Results of all analyses strongly supported the close relationship between Phthiraptera and Liposcelididae. Results of the present analyses also provided some insight into the elevated rate of evolution in mitochondrial DNA (mtDNA) in Phthiraptera. An elevated substitution rate of mtDNA appears to originate in the common ancestor of Phthiraptera and Liposcelididae, and directly corresponds to an increased G + C content. Therefore, the elevated substitution rate of mtDNA in Phthiraptera and Liposcelididae appears to be directional. A high diversity of 12S rDNA secondary structure was also observed in wide range of Phthiraptera and Liposcelididae, but these structures seem to have evolved independently in different clades.

Phylogenetic relationships among all 24 families of suborder Psocomorpha (Insecta: Psocodea: ‘Psocoptera’) are inferred based on adult morphology. Monophyly of Psocomorpha is strongly supported by six autapomorphies. The presently accepted four infraorders − Psocetae, Homilopsocidea, Epipsocetae and Caeciliusetae − are regarded as monophyletic, but Archipsocidae and Hemipsocidae, previously assigned to Homilopsocidea and Psocetae, respectively, are regarded as the basalmost clades of the suborder. Based on the results of the cladistic analysis, a higher classification for Psocomorpha is proposed. Six infraorders (two new − Archipsocetae, Hemipsocetae − and the four aforementioned) are recognized. Four new superfamilies are recognized within Homilopsocidea: Elipsocoidea, Lachesilloidea, Pseudocaecilioidea and Peripsocoidea . Two superfamilies are recognized within Caeciliusetae: Asiopsocoidea and Caeciliusoidea. Descriptions of taxa above family level are provided. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society , 2002, 136 , 371 − 400

Three new species of the uncommonly encountered insect order Zoraptera are described and figured from Peninsular Malaysia— Zorotypus magnicaudelli sp. n., Zorotypus cervicornis sp. n., and Zorotypus impolitus sp. n. Another species from the region, identified as Zorotypus caudelli Karny, 1927, was also collected and is reevaluated herein based on new material. A brief discussion of characters used in zorapteran systematics is provided, and a key to the species of Peninsular Malaysia provided. This is the first report for the order Zoraptera from Peninsular Malaysia.

Atrichadenotecnum multispinosus sp. n. is described from southwestern China. Psocomesites and Clematostigma from China are discussed, with five species transferred to Atrichadenotecnum and four species placed as new synonyms. Keys to adult males and females of Atrichadenotecnum species are presented.

Background The 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. Results Minicircular 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). Conclusions Minicircular 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.

The systematic status of Kaindipsocinae (formerly Kaindipsocini) is revised based on morphology of the male terminalia and on molecular data. The genera Clematostigma, Lasiopsocus and Tanystigma are newly assigned to this subfamily. The ‘Blaste’ lunulata species-group is also placed within Kaindipsocinae and is probably closest to Kaindipsocus. Both morphological and molecular data provide strong support for monophyly of Kaindipsocinae and molecular data support a sister relationship between this subfamily and the rest of Psocidae.

Over the past century, the geosciences have developed an impressive capability to understand and anticipate events occurring within the Earth system.The past decade has seen an explosion in quantitative geoscience. Progress is continuing to accelerate as advances in observational systems and computational tools are allowing simulations and predictions of geophysical processes with a temporal and spatial resolution unprecedented in human history. With these advances comes the necessity to reflect on the current state of geoscientific research, to reassess the direction in which research should be progressing, and to redefine the goals that can be realistically achieved.

The genus Trichadenotecnum Enderlein, 1909 is redefined as a monophyletic group. The following 22 Japanese species are treated and classified into five species-groups except one species not placed in a group: sexpunctatum-group – T. album, sp. nov., T. incognitum Roesler, 1939; medium-group – T. takahashii, sp. nov., T. mixtum, sp. nov., T. magnomixtum, sp. nov., T. amamiense, sp. nov., T. okinawense, sp. nov., T. yonaguniense, sp. nov., T. kumejimense, sp. nov., T. latebrachium, sp. nov., T. pseudomedium, sp. nov., T. yaeyamense, sp. nov.; alexanderae-group – T. alexanderae Sommerman, 1948, T. castum Betz, 1983, T. sexpunctellum (Enderlein, 1907); majus-group – T. yamatomajus, sp. nov., T. nothoapertum, sp. nov.; spiniserrulum-group – T. falx, sp. nov., T. furcalingum, sp. nov., T. pardidum Thornton, 1961, T. circularoides Badonnel, 1955; incertae sedis – T. fuscipennis, sp. nov. The monophyly and phylogenetic relationships of these species-groups are discussed. A key to Japanese species of Trichadenotecnum is given.

We present some singular characteristics of the 2011 off the Pacific coast of Tohoku Earthquake in comparison with other megathrust earthquakes, such as the 1960 Chilean and the 2004 Sumatra-Andaman earthquakes. In addition to the conventional along-strike segmentation, along-dip segmentation of the fault area or subduction zone is an important feature for the Tohoku subduction zone, as indicated by the difference in background seismicity: virtually no seismicity in shallow segments but active with large events repeating in deep segments. The interaction between along-dip segments (deep and shallow segments) led to the great 2011 Tohoku earthquake. The along-dip segmentation results in plane or two-dimensional rupture propagation on a coseismic fault. Significant along-strike variability is also important for the 2011 Tohoku earthquake, with segments of both weak (e.g., slow or tsunami earthquakes) and strong plate couplings located adjacent to each other. In contrast, every segment appears to be with strong plate coupling for other megathrust earthquakes. One exception is the 1964 Alaska earthquake that shares a certain degree of common characteristics with the 2011 Tohoku earthquake: two distinct seismogenic zones along the dip direction of the trench, that is, the along-dip segmentation is noticeable. Significant along-strike variability also characterizes the activities in and around the subduction zone of the 1964 Alaska earthquake, including a creeping segment and a tsunami-earthquake segment.