The wing base of the palaeodictyopteran genus Dunbaria Tillyard: Where are we now?

The structure of insect wing articulation is considered as reliable source of high level characters for phylogenetic analyses. However, the correct identification of homologous structures among the main groups of Pterygota is a hotly debated issue. Therefore, the reconstruction of the wing bases in Paleozoic extinct relatives is of great interest, but at the same time it should be treated with extreme caution due to distortions caused by taphonomic effects. The present study is focused on the wing base in Dunbaria (Spilapteridae). The articulation in Dunbaria quinquefasciata is mainly formed by a prominent upright axillary plate while the humeral plate is markedly reduced. Due to unique preservation of surface relief of the axillary plate, its composition shows a detailed pattern of three fused axillary sclerites and presumable position of the sclerite 3Ax. The obtained structures were compared among Spilapteridae and to other palaeodictyopterans Ostrava nigra (Homoiopteridae) and Namuroningxia elegans (Namuroningxiidae). The comparative study uncovered two patterns of 3Ax in Dunbaria and Namuroningxia, which correspond to their different suprafamilial classification. In contrast to previous studies these new results reveal the homologous structural elements in the wing base between Paleozoic Palaeodictyoptera and their extant relatives of Ephemeroptera, Odonata and Neoptera.     

The morphology and terminology of the hindwing articulation and wing base of the Coleoptera, with specific reference to the Scarabaeoidea

Abstract. Characters of the hindwing articulation and wing base are important for contributing to the solution of phylogenetic and systematic problems in the Coleoptera. In the Scarabaeoidea morphological terms proposed by previous authors do not cover many structures in sufficient detail and additional terms are needed to describe and utilize all characters used in systematic considerations; these can be used for all Coleoptera.
In this paper we identify new structures, the first basal plate and the second basal plate (two subdivisions of the coleopteran wing base), name the various yokes, braces and reinforcements found on them and propose names for various projections, lobes, indentations and embayments on the axillary sclerites (first axillary, head, neck and tail; second axillary, arm and body; third axillary, prong; basalare).     

Homology of the wing base sclerites in Ephemeroptera (Insecta: Pterygota) - a reply to Willkommen and Hörnschemeyer

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.     

Origin and evolution of insect wings and their relation to metamorphosis,as documented by the fossil record

In contemporary entomology the morphological characters of insects are not always treated according to their phylogenetic rank. Fossil evidence often gives clues for different interpretations. All primitive Paleozoic pterygote nymphs are now known to have had articulated, freely movable wings reinforced by tubular veins. This suggests that the wings of early Pterygota were engaged in flapping movements, that the immobilized, fixed, veinless wing pads of Recent nymphs have resulted from a later adaptation affecting only juveniles, and that the paranotal theory of wing origin is not valid. The wings of Paleozoic nymphs were curved backwards in Paleoptera and were flexed backwards at will in Neoptera, in both to reduce resistance during forward movement. Therefore, the fixed oblique-backwards position of wing pads in all modern nymphs is secondary and is not homologous in Paleoptera and Neoptera. Primitive Paleozoic nymphs had articulated and movable prothoracic wings which became in some modern insects transformed into prothoracic lobes and shields. The nine pairs of abdominal gillplates of Paleozoic mayfly nymphs have a venation pattern, position, and development comparable to that in thoracic wings, to which they are serially homologous. Vestigial equivalents of wings and legs were present in the abdomen of all primitive Paleoptera and primitive Neoptera. The ontogenetic development of Paleozoic nymphs was confluent, with many nymphal and subimaginal instars, and the metamorphic instar was missing. The metamorphic instar originated by the merging together of several instars of old nymphs; it occurred in most orders only after the Paleozoic, separately and in parallel in all modern major lineages (at least twice in Paleoptera, in Ephemeroptera and Odonata; separately in hemipteroid, blattoid, orthopteroid, and plecopteroid lineages of exopterygote Neoptera; and once only in Endopterygota). Endopterygota evolved from ametabolous, not from hemimetabolous, exopterygote Neoptera. The full primitive wing venation consists of six symmetrical pairs of veins; in each pair, the first branch is always convex and the second always concave; therefore costa, subcosta, radius, media, cubitus, and anal are all primitively composed of two separate branches. Each pair arises from a single veinal base formed from a sclerotized blood sinus. In the most primitive wings the circulatory system was as follows: the costa did not encircle the wing, the axillary cord was missing, and the blood pulsed in and out of each of the six primary, convex-concave vein pair systems through the six basal blood sinuses. This type of circulation is found as an archaic feature in modern mayflies. Wing corrugation first appeared in preflight wings, and hence is considered primitive for early (paleopterous) Pterygota. Somewhat leveled corrugation of the central wing veins is primitive for Neoptera. Leveled corrugation in some modern Ephemeroptera, as well as accentuated corrugation in higher Neoptera, are both derived characters. The wing tracheation of Recent Ephemeroptera is not fully homologous to that of other insects and represents a more primitive, segmental stage of tracheal system. Morphology of an ancient articular region in Palaeodictyoptera shows that the primitive pterygote wing hinge in its simplest form was straight and composed of two separate but adjoining morphological units: the tergal, formed by the tegula and axillaries; and the alar, formed by six sclerotized blood sinuses, the basivenales. The tergal sclerites were derived from the tergum as follows: the lateral part of the tergum became incised into five lobes; the prealare, suralare, median lobe, postmedian lobe and posterior notal wing process. From the tips of these lobes, five slanted tergal sclerites separated along the deep paranotal sulcus: the tegula, first axillary, second axillary, median sclerite, and third axillary. Primitively, all pteralia were arranged in two parallel series on both sides of the hinge. In Paleoptera, the series stayed more or less straight; in Neoptera, the series became V-shaped. Pteralia in Paleoptera and Neoptera have been homologized on the basis of the fossil record. A differential diagnosis between Paleoptera and Neoptera is given. Fossil evidence indicates that the major steps in evolution, which led to the origin first of Pterygota, then of Neoptera and Endopterygota, were triggered by the origin and the diversification of flight apparatus. It is believed here that all above mentioned major events in pterygote evolution occurred first in the immature stages.     

Higher‐level molecular phylogeny of jumping plant lice (Hemiptera: Sternorrhyncha: Psylloidea)

Jumping plant lice (Hemiptera: Psylloidea) are known for a few deleterious pest species worldwide, yet the phylogeny of the group has been poorly understood until very recently. Here, we reconstruct the higher‐level phylogeny for the superfamily Psylloidea based on multilocus DNA sequences, three mitochondrial (COI‐tRNA^leu‐COII, 12S, 16S) and five nuclear (18S, 28S D2, 28S D3, 28S D6–7a, 28S D9–10) gene fragments, using maximum likelihood and Bayesian inference phylogenetic frameworks. Our results are largely congruent with the recent phylogenomic study and partly support prior classification of Psylloidea based mainly on morphology, with the following major exceptions: the family Calophyidae is revealed as polyphyletic, Aphalaridae as paraphyletic with respect to most other taxa of Psylloidea, and Liviidae as paraphyletic with respect to Calophyinae, Psyllidae and Triozidae. Our phylogenetic hypothesis identifies Phacopteronidae and the genus Cecidopsylla Kieffer as the very basal taxa within extant Psylloidea. Sister‐group relationships of Rhinocolinae with Togepsyllinae and of Pachypsyllinae with Homotomidae are also suggested. We present specific discussions for each group of interest recovered in our phylogenetic analysis. One nomenclatorial change is proposed: Spanioneura longicauda (Konovalova) comb.n. , from Psylla Geoffroy.     

Descriptions of two new endoparasitic cecidomyiids (Diptera: Cecidomyiidae) from Japan

Two endoparasitic species of Cecidomyiidae (Diptera) new to science are reported from Japan. Females of Endaphis psyllophaga sp. nov. lay their eggs on the wing of Calophya nigridorsalis (Hemiptera: Psylloidea: Calophyidae) on Rhus succedanea (Anacardiaceae), and newly hatched larvae bore into the adult body. The six nominal species of the genus Endaphis are endoparasitoids of aphids. The genus Endopsylla, which is morphologically similar to the genus Endaphis, consists of two species whose larvae attack psyllids or tingids. Females of Endaphis muraii sp. nov. lay their eggs near colonies of host aphids and newly hatched larvae bore into the body of aphids such as Macrosiphum euphorbiae and Aphis glycines (Hemiptera: Aphididae). The two new species are described, illustrated, and compared to known congeners, and information is given for the two species on their distribution, host range and ecological traits. Now, E. muraii is considered to be a potential biological control agent against aphids.     

Description of two new genera, Santelmoa and Bentartia and two new species of Zoarcidae (Teleostei, Perciformes) from the Southern Ocean

Two new genera of lycodine zoarcid fish, Santelmoa and Bentartia, and two new species, Santelmoa carmenae and Bentartia cinerea, are described from 13 specimens collected from the Gerlache Strait, Southern Ocean, at 1,056-m depth. Santelmoa can be distinguished from all other lycodine genera by the combination of the following characters: anterior portion of frontals fused; scapular foramen open; ceratohyal–epihyal articulation interdigitating; cranium narrowed; supratemporal commissure and occipital pores absent; intercalar reaching the prootic; parasphenoid wing well developed; palatal arch well developed; posterior hyomandibular ramus short; post-temporal ventral ramus well developed; six branchiostegal rays; vertebrae asymmetrical; pelvic fin rays ensheathed; scales, lateral line, pyloric caeca, palatine and vomerine teeth present. Bentartia differs from the remaining lycodine genera by the following combination of characters: basioccipital and exoccipitals fused; supraoccipital–exoccipital articulation broadly contacting; ceratohyal–epihyal articulation interdigitating; post-temporal ventral ramus weak; two posterior nasal pores; cranium narrowed; supratemporal commissure and occipital pores absent; intercalar set posteriorly; palatal arch well developed; posterior hyomandibular ramus not elongate; parasphenoid wing high; six branchiostegal rays; vertebrae asymmetrical; pelvic fin rays ensheathed; scales, lateral line, pyloric caeca, palatine and vomerine teeth present. The relationships of the two new genera are discussed.     

Contribution to the Knowledge of the Pre-imaginal Stages of Neotropical Hydrobiosidae (Trichoptera): Metachorema griseum Schmid

Mature larvae and pupae of Metachorema griseum Schmid are described for the first time. The known distribution of the genus includes the province of Neuquén in Argentina and central and southern Chile. The distinctive characters of the larvae of this genus are: (1) prothoracic sternite consisting of a wide subrectangular central sclerite and a pair of elongated lateral sclerites, (2) anterior femora with a narrow basodistal process, (3) anterior tibia and tarsus fused and (4) anal prolegs with a short lateral sclerite with basal spine and claws simple with ventral spine and two ventral setae.     

The clavus and jugum venation in the wings of beetles (Coleoptera) and its genesis

The homology and nomenclature, as well as hypothesized pathways of the historical development of the clavus and jugal lobe of the beetle hind wings are discussed. The reconstructed plan of the clavus venation is largely similar to the venation patterns observed in some representatives of Corydalidae (Megaloptera). Its main apotypic characters are the following: the first anal cell was reduced at the wing base and the second anal or cuneiform cell appeared. This venation pattern is supposed to result from consolidation of the area around the claval furrow base at the earlier stages of the beetle wing evolution. In particular, longitudinal compressing the bases of RA, M, CuP, and 1A resulted in the development of a complex sclerite composed of the distal median plate and the bases of CuP and 1A. After this new reinforced connection between the remigium and the clavus appeared, the proximal parts of CuP and 1A were partly or completely reduced since they were no longer needed to maintain the structural integrity of the beetle wing.     

Geometric morphometrics as a tool for interpreting evolutionary transitions in the black fly wing (Diptera: Simuliidae)

A geometric morphometric analysis was conducted on wing‐vein landmarks on exemplar species of the family Simuliidae of the following genera: Parasimulium, Gymnopais, Twinnia, Helodon, Prosimulium, Greniera, Stegopterna, Tlalocomyia, Cnephia, Ectemnia, Metacnephia, Austrosimulium, and Simulium. Generalized least squares superimposition was performed on landmarks, followed by a principal component analysis on resulting Procrustes distances. Patterns of shape change along the principal component axes were visualized using the thin‐plate spline. The analysis revealed wing shape diversity through (1) the insertion points of the subcosta and R₁, resulting in the terminus of the costa exhibiting a trend towards a more apical position on the wing, and (2) the insertion point of the humeral cross vein, resulting in the anterior branch of the media exhibiting a trend toward a more basal position on the wing. Canonical variates analysis of Procrustes distances successfully assigned all exemplar species into their a priori taxonomic groupings. The diversity in wing shape reveals a trend towards decreased length of basal radial cell and increased costalization of anterior wing veins in the evolutionary transition from plesiomorphic prosimuliines to more derived simuliines. The functional significance of these evolutionary transitions is discussed. © 2013 The Linnean Society of London     

Mechanisms of wing rotating regulation in Calliphora erythrocephala (Insecta,Diptera)

Summary The blowfly Calliphora erythrocephala rotates its wings, i.e. changes the geometrical angle of attack, generating forces and moments for flight steering. There are two possibile ways to regulate this angle. The mechanisms for these movements are described. (1) The leading edge and the anterior part of the wing — between the costal vein and radial vein 4 — are pronated automatically due to the interaction of the moving parts during the downstroke. They are supinated during the upstroke. This is basic automatic regulation. (2) The posterior part of the wing — behind the anterior cross vein — is pronated and supinated independently of wing-drive. This is wing-drive independent additional regulation.Abbreviations a.c anterior cross vein - a.n anal veins - a.t.l anterior tergal lever - a.w anterior part of the wing - b.z bending zone - co costal vein - cr crossing of the tendons of the posterior notal wing process - c.s cross section - cu cubital vein - f fit or turning point of ventral radial vein 1 - h.a horizontal axis of pterale III - h.c humeral cross vein - h.co head of costal vein - h.r head of radial vein - k Klöppel - l.a longitudinal axis - me median vein - mp middle plate - ms mesoscutum - p anterior process of the anal veins - p.c posterior cross vein - pl pleurum - p.n.w.p posterior notal wing process - p.n.w.p 1–4 muscles 1–4 of the posterior notal wing process - pt I–III pterale I–III - p.t.l posterior tergal lever - p.w. posterior part of the wing - p.w.j pleural wing joint - r 1–4 radial veins 1–4 - r.s. ring stiffenings - sc subcostal vein - s.p semicircular part of the middle plate - s.t subalar tendon - t.c tip cross vein - te tegula - t.st thin strips - t.v.r tooth of ventral radial vein - v.a. vertical axis of pterale III - w wing - III 1–4 muscles 1–4 of pterale III     

How Insect Flight Steering Muscles Work

Insights into how exactly a fly powers and controls flight have been hindered by the need to unpick the dynamic complexity of the muscles involved. The wingbeats of insects are driven by two antagonistic groups of power muscles and the force is funneled to the wing via a very complex hinge mechanism. The hinge consists of several hardened and articulated cuticle elements called sclerites. This articulation is controlled by a great number of small steering muscles, whose function has been studied by means of kinematics and muscle activity. The details and partly novel function of some of these steering muscles and their tendons have now been revealed in research published in this issue of PLOS Biology. The new study from Graham Taylor and colleagues applies time-resolved X-ray microtomography to obtain a three-dimensional view of the blowfly wingbeat. Asymmetric power output is achieved by differential wingbeat amplitude on the left and right wing, which is mediated by muscular control of the hinge elements to mechanically block the wing stroke and by absorption of work by steering muscles on one of the sides. This new approach permits visualization of the motion of the thorax, wing muscles, and the hinge mechanism. This very promising line of work will help to reveal the complete picture of the flight motor of a fly. It also holds great potential for novel bio-inspired designs of fly-like micro air vehicles.The ability for powered flight has evolved four times in the animal kingdom and, thanks to their ability to fly, insects have diversified and moved into new regions and habitats with enormous success [1]. Powered flight requires an integrated system consisting of wings to generate aerodynamic force, muscles to move the wings, and a control system to modulate power output from the muscles. Insects are bewilderingly diverse with respect to flight morphology and behaviors, which in turn provides a real challenge to researchers wishing to understand how insects fly. In particular, the impressive flight maneuvers in flies, such as blowflies and fruit flies, have inspired scientists for many years [2]. The ability of a fly to accelerate, make tight turns, rolls, and loops that allow the creature to land upside down on a ceiling is unparalleled in any other organisms, as well as any manmade aircraft. Everybody knows how difficult it is to swat a fly with bare hands—the fly''s capacity for rapid take-off and accurate movement away from a perceived approaching threat is exquisite [3].The flight muscles of many insects, including flies, bees, and mosquitoes, are divided into a few large power muscles that simply contract cyclically to generate sheer power output and a greater number of smaller steering muscles that control the force transmission from the power muscles to the wing [4]–[6]. The power muscles of a fly consist of two sets of antagonistic muscles attached to the inside of the thorax (exoskeleton) (Figure 1). In many insects, including flies, these muscles are asynchronous, which means their contractions are uncoupled to the firing rate of the associated motor neuron [6],[7], i.e., the muscles continue to contract as long as the nerve tickles them. Another characteristic feature of the power muscles is that they are stretch-activated and contract as a response to being lengthened. Both sets of power muscles deform the thorax when contracted such that when the dorso-ventral muscles contract, the thorax is squeezed together dorso-ventrally while expanding longitudinally, and vice versa when the dorsal-longitudinal muscles contract as a response to prior lengthening. The result is an alternate contraction and lengthening of these perpendicular muscle groups and a resonance of the entire thorax that drives the wingbeat. Typical wingbeat frequencies are in the range from 100 Hz and even up to 1,000 Hz in the smallest species [5],[8].Open in a separate windowFigure 1The thorax with and dorsal longitudinal (upper left) and dorso-ventral (upper right) power flight muscles of a fly.The cartoon (bottom) shows a transverse section through the thorax with dorso-ventral muscles (DVM) and dorsal longitudinal muscles (DLM) indicated. The two upper illustrations are redrawn from [6].The forces from the flight muscles are transmitted to the wing through an intricate hinge mechanism (Figure 2). The hardened plates of cuticle between the thorax and wing (sclerites) are mobile and their positions relative to the thoracic outgrowths and wing determine the extent of the wing motion, i.e., the angular amplitude of the wingbeat [6].Open in a separate windowFigure 2Cartoon illustration of a transverse section of the thorax of a fly in rear view, showing some elements of the complex wing hinge of a fly, consisting of ridges and protrusions on the thorax and a number of hardened plates of cuticle (sclerites) between the body (thorax) and the wing root.The basalare sclerite (not shown) is positioned anterior of the first axillary sclerite (Ax1). The indicated structures are dorso-ventral power muscle (DVM), pleural wing process (PWP), post-medial notal process (PMNP), parascutal shelf (PSS), axial wing sclerites (Ax1, Ax2, Ax3), and radial stop (RS). Redrawn and modified from [18].Flight maneuvers arise owing to asymmetric force generation between the left and right wing. Aerodynamic force is proportional to the angle of attack (the angle between the wing surface and the airflow) and the speed squared relative to the air [9],[10]. Except from the turning points of each half-stroke, when the wings rotate about their span wise axes, the angle of attack is usually quite constant during the translational phases of the wingbeat [10], while asymmetric forces are mainly created by changing the wingbeat amplitude in flies [11]–[14]. With wingbeat frequency kept constant, changed amplitude changes the speed and hence force generated.The control of the elements forming the hinge mechanism of the wing is achieved by the steering muscles, which are tiny in terms of mass (<3% of the power muscle mass), but mean everything when it comes to making flight maneuvers. In contrast to the power muscles the steering muscles are synchronous, i.e., there is a 1∶1 correspondence between neural spikes and muscle contraction. No less than some 22 pairs of steering muscles are involved in the force transmission; a few of these indirectly modulate the output by affecting the resonating properties of the thorax, while others are directly attached to the sclerite elements of the hinge mechanism [6],[15]. Three small muscles (b1–b3) are attached to the basalare plate that is directly involved in wing articulation (Figure 3). The actual wing sclerites (Figure 2) are also controlled by specific steering muscles, also with the function of moving the sclerites in relation to required wing motion. The main control function of the hinge mechanism appears to be of the downward movement of the wing, i.e., the angle at the turning point at end of downstroke. For a detailed review about the steering muscles and their function see Dickinson and Tu [6].Open in a separate windowFigure 3The position of the three steering flight muscles b1–b3 inserted to the nail-shaped basalare sclerite.Contraction by the b1 and b2 muscles move the basalare forward and their antagonist b3 moves it backwards when contracted. Redrawn from [6].To date, the function of the steering muscles has been revealed mainly by electrophysiological studies on tethered subjects. Tethering means that the animal is glued to the end of a thin rod, often with force sensors attached to it, and then stimulated to “fly.” In many insects this can be achieved by simply blowing at them or placing them in a wind tunnel. On the tether the insect can either be presented with a visual stimulus or be rotated, which flies can sense via their halteres (hind wings modified to sensory gyroscopic sensory organs) [16]. By inserting electrode wires into the steering muscles, the neural impulses are measured at the same time as the wingbeat kinematics is recorded [13],[17]. What we know about the function of the steering muscles comes from the meticulous studies of correlations between muscle activity and the associated wing movement, including how the hinge mechanism works [6],[18]. Needless to say, such experiments are extremely difficult to achieve in small insects like blowflies and fruit flies that flap their wings at high frequencies. Recent studies of the wing and hinge kinematics provide some support for the hypothesis that the hinge may have a gear function that affects stroke amplitude, as well [18]. However, there are still many open questions regarding the exact function of the steering muscles and how they help in generating laterally asymmetric forces during a fly''s flight maneuver [6].In an article published in this issue of PLOS Biology, Walker and colleagues take a new approach for studying how steering muscles regulate the power output from power muscles [19], using time-resolved x-ray microtomography [20]. By rotating tethered blowflies (Calliphora vicina) in the X-ray beam, a 3D-movie was captured that shows how the steering muscles move. This by itself is a grand achievement at a wingbeat frequency of 145 Hz. As the flies could sense being rotated the steering muscles acted accordingly to achieve an asymmetric power output as a response to a perceived turn. The movies that accompany the article show how several of the key steering muscles and their sclerites operate in concert during the course of a wingbeat, and the visual results are supported by advanced statistical analyses of muscle strain rates and their phase offset. For example, the b1 and b3 muscles (Figure 3) work antagonistically, as was known before, but on the low-amplitude wing the oscillations are delayed by about a quarter of a wingbeat. The strain amplitudes of b1 and b3 were different between the two wings, which were found to be due to dorso-ventral movement of the basalare sclerite on the high-amplitude side and rotation on the low-amplitude side. This shows even higher complexity of the wing hinge than was previously envisaged.The measurements of strain rate in the muscle confirmed the results of a previous study, which showed that asymmetric power output is partially achieved by negative work [21], i.e., absorption of work, by the b1 muscle on the low-amplitude wing. As with other muscles, the steering muscles insert on the skeletal parts and sclerites by tendons. The tendon of the muscle (I1) associated with the first axillary sclerite was observed to buckle when the wing was elevated above the wing hinge, indicative of compressive force acting on it near the top of the wing stroke. This buckling of the tendon forces a reinterpretation of the function of this muscle: it is involved in reducing stroke amplitude at the bottom of the downstroke rather than exerting stress near the opposite end of the stroke. Tendon buckling was seen in some other muscles as well, and although this is its first observation, it may be a more general mechanism involved in control of insect wingbeat kinematics.What are the wider implications of this new study? First, it demonstrates the utility of a new approach to examine the in vivo operation of several insect flight muscles. This alone signals a methodological breakthrough that promises more. So far the flies were tethered and studied during one behavioral treatment (rotation about the yaw axis). Real flight maneuvers, however, also involve angular rotation about pitch and roll axes, acceleration, and braking. Thus, it remains to be seen how the steering muscles operate to control more subtle changes in wing kinematics during the turning saccades and advanced flight maneuvers that take place during free flight. The method involved exposure to lethal X-ray doses, which of course limits how long the experiments can be. Second, tethering is the prevailing paradigm for studying insect flight, but because it interrupts the sensory feedback loop [22], it would be useful for future studies to compare tethered and free flight in some commonly studied species. Furthermore, a more complete understanding of the flight muscle-hinge mechanism may help bio-inspired design of wing articulation systems for fly-like micro air vehicles. Until then, we can enjoy the stunning videos of the oscillating thorax and flight muscle system of the blowfly [19]. See the video from the related research article here (http://youtu.be/P6lBkK3J9wg) or [19].     

The lateral thoracico-abdominal region in adults and fifth instar nymphs of an aquatic bug,Notonecta undulata say (Insecta,Heteroptera)

Skeletal differences in the lateral thoracico-abdominal regions of fifth instar and adult Notonecta appear to reflect respiratory differences in the two stages. Changes in the epidermis of this region during the last instar are described, and the possible relationships between the epidermis, nymphal cuticle, and imaginal cuticle are discussed.Explanation of Figures A Abdominal segment - AC Antecosta (anterior boundary of abdominal segment) - AP Abdominal projection of second abdominal segment - AT Anterior tracheolar branch - B Posteromedial boundary of posterior projection - C Metacoxa - CM Metacoxal membrane - CP Metacoxal process - CS Coxal-subalar muscle - DL Dorsal longitudinal muscle of abdomen - EL External lateral muscle of abdomen - EM Metathoracic epimeron - ES Metathoracic episternum - EV Posterior evagination of metanotum - F Fold of metacoxal membrane - FM Functional thoracico-abdominal membrane - FW Metathoracic wing or wingpad - H Horizontal ridge or sulcus on metathoracic postnotum - HW Metathoracic wing or wingpad - HWM Costal margin of hindwing - ILL Internal lateral muscle of abdomen, lateral portion - ILM Internal lateral muscle of abdomen, medial portion - IN Insertions of wingbase muscles on subalar epidermis - LE Lobe of metathoracic episternum - MM Posterior moulting muscle - MP Process for attachment of Muscle EL 1 - MSP Pleuron of mesothorax - MST Tergum of mesothorax - N Metathoracic notum - P Metathoracic pleural ridge or sulcus - PA Postalar portion of metathorax - PAB Metathoracic postalar bridge - PH Third phragma - PN Metathoracic postnotum - PP Posterior projection of metathoracic epimeron - PT Posterior tracheolar branch - PW Posterior end of metathoracic wingbase - S Abdominal spiracle - SA Subalar portion of metathorax - SAM Subalar membrane - SAS Subalar sclerite - SAV Widened ventrolateral portion of subalar air space - SL Scutellum of metathoracic notum - SM Spiracular membrane - ST Spiracular tracheole - SU Scutum of metathoracic notum - T Tendon of metacoxal remotor muscle - TA Thoracico-abdominal sclerite - TAA Anterior portion of thoracicoabdominal sclerite - TAC Posterior portion of thoraeicoabdominal sclerite - TAM Membranous portion of lateral thoracico-abdominal region - TS Metathoracic spiracle - VA Anterior end of ventral abdominal air channel - WB Junction between subalar wall and metathoracic wing or wingpad - WH Hairs on margin of mesothoracic wingpad     

The Drosophila gene zfh2 is required to establish proximal-distal domains in the wing disc

Three main events characterize the development of the proximal-distal axis of the Drosophila wing disc: first, generation of nested circular domains defined by different combinations of gene expression; second, activation of wingless (wg) gene expression in a ring of cells; and third, an increase of cell number in each domain in response to Wg. The mechanisms by which these domains of gene expression are established and maintained are unknown. We have analyzed the role of the gene zinc finger homeodomain 2 (zfh2). We report that in discs lacking zfh2 the limits of the expression domains of the genes tsh, nub, rn, dve and nab coincide, and expression of wg in the wing hinge, is lost. We show that zfh2 expression is delimited distally by Vg, Nub and Dpp signalling, and proximally by Tsh and Dpp. Distal repression of zfh2 permits activation of nab in the wing blade and wg in the wing hinge. We suggest that the proximal-most wing fate, the hinge, is specified first and that later repression of zfh2 permits specification of the distal-most fate, the wing blade. We propose that proximal-distal axis development is achieved by a combination of two strategies: on one hand a process involving proximal to distal specification, with the wing hinge specified first followed later by the distal wing blade; on the other hand, early specification of the proximal-distal domains by different combinations of gene expression. The results we present here indicate that Zfh2 plays a critical role in both processes.     

Description of a new genus and species of zoarcid fish, <Emphasis Type="Italic">Bellingshausenia olasoi</Emphasis>, from the Bellingshausen Sea (Southern Ocean)

A new genus and species of zoarcid fish, Bellingshausenia olasoi, is described on the basis of five specimens collected from the Bellingshausen Sea, Southern Ocean, at depths of 602–615 m. Bellingshausenia is a lycodine that can be distinguished from all other zoarcid genera by the following combination of characters: seven branchiostegal rays, scapular foramen open, cranium narrowed, smooth ceratohyal-epihyal articulation, palatal arch well developed, supratemporal commissure and occipital pores absent, intercalar reduced and displaced backward and parasphenoid wing high. The relationships of the new genus are discussed.     

海南省蚋属一新种(双翅目：蚋科)

记述采自海南省尖峰蚋属蚋亚属Simulium(Simulium)一新种,乐东蚋Simulium(Simulium)ledongense sp.nov.。新种隶属多叉蚋组Simulium multistriatum-group,与秦氏蚋Simulium qiniCao et al.,1993和鞍阳蚋Simulium dphippiodum Chen and Wen,1999近缘。对其各虫期形态进行描述并与其近缘种比较作分类讨论。模式标本保存于遗阳医学院生物学教研室。     

Respiratory significance of the thoracic and abdominal morphology of the three aquatic bugs Ambrysus,Notonecta and Hesperocorixa (Insecta,Heteroptera)

The spiracular chambers of the three anterior pairs of spiracles are described, and their relationships with the air stores on the body are discussed. Morphological variations in the spiracular chambers of Ambrysus, Notonecta, and Hesperocorixa appear to be correlated with the manner in which these insects obtain atmospheric and dissolved oxygen.

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Explanation of Figures AB Abdominal segment - AC Antecosta - AF Flap on second abdominal segment - AM Alar membrane - AP Abdominal projection of second abdominal segment - AT Air trough of metathoracic spiracular chamber - AX Humeral axillary sclerite - CL Clavus of forewing - CM Coxal membrane - CO Corium of forewing - CP Metacoxal process - CS Claval suture of forewing - CX Coxa - D Wing-anchoring depression on costal margin of forewing - DVM Dorsoventral muscle of first abdominal segment - E Embolium of forewing - EM Epimeron - ES Episternum - FE Femur - FL Posterior flap on costal margin of forewing - FW Forewing - G Gap between forewing and body - OR Groove on abdominal projection - H Horizontal ridge or sulcus - HA Hairs on precoxal bridge HD Head - HL Laterosternal hairs - HW Hindwing - L Laterosternite of abdomen - LH Lateral hairs of ventral abdominal air channel - LM Epirneral lobe - LMO Opening into epimeral lobe - LN Notal lobe - LS Episternal lobe - MI-II Membrane between prothorax and mesothorax - MB Membrane of forewing - MH Medial hairs of ventral abdominal air channel - MP Process for dorsoventral muscle of first abdominal segment - N Notum - P Pleural ridge or sulcus - PB Postalar bridge - PN Postnotum - PP Postalar projection of postalar bridge - R Ridge on metathoracic episternum - RS Rounded swelling on costal margin of forewing - S Spiracle - SA Subalar air store - SC Spiracular chamber - SO Scolopophorous sense organ - ST Sternum - TA Thoracico-abdominal sclerite - TAL Thoracico-abdominal lobe of thoracico-abdominal sclerite - TAM Functional (not morphological) thoracico-abdominal membrane - TAO Opening into thoracico-abdominal lobe - TR Trochanter - V Vertical ridge or sulcus - VP Vertical plate of thoracico-abdominal sclerite - WA Wing-anchoring process of mesothoracic epimeron - WF Wing-anchoring flap on costal margin of forewing - WH Hairs on emboliar margin of forewing - WM Costal margin of forewing - WR Ridge on inner surface of forewing - XC Axillary cord of forewing - 2 or 3 PH Second or third phragma - II–III Boundary between mesothorax and metathorax     

COMPARATIVE MORPHOLOGY OF THE CARPEL IN THE ROSACEAE VI. POMOIDEAE: ERIOBOTRYA,HETEROMELES, PHOTINIA,POURTHIAEA, RAPHIOLEPIS,STRANVAESIA

The pomoid genera, Eriobotrya, Photinia, Pourthiaea, Raphiolepis, Stranvaesia, and Heteromeles, have compound inflorescences and biovulate carpels which become papery at maturity. The carpels of all of these except Heteromeles are fused with one another. There are open sutures in the carpels of Heteromeles, Photinia, Pourthiaea, and Raphiolepis, and in these four genera the extent of fusion of the ovular bundle with the wing bundle is related directly to the state of tegumentary fusion and to the extent of fusion of the carpel with the floral cup. In those species of Eriobotrya and Stranvaesia with closed sutures the integuments tend to be fused, as do the ovular and wing bundles, and the carpels are adnate with the floral cup for a considerable distance; in species with open sutures the integuments tend to be free, the ovular and wing bundles tend to be separate, and the extent of fusion of carpel with floral cup tends to be shorter. In genera with connate carpels the wing bundles of adjoining carpels may also be fused. The greatest extent of fusion occurs in Eriobotrya and Raphiolepis, in which there may also be attenuation and disappearance of the wing bundles above the region of ovular insertion and even reduction and disappearance of the carpellary margin.