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Concept: Phasmatodea


The Lord Howe Island stick insect, Dryococelus australis, was once common on the island but was driven to extinction after the arrival of ship rats in the early 20(th) century [1, 2]. It was thought to be extinct for decades, until a tiny population of similar-looking stick insects was discovered 20 km away, on the islet of Ball’s Pyramid, in 2001 [2]. Individuals from this population are currently being reared in Australia and elsewhere in the world, with the eventual goal of recolonizing Lord Howe Island [3]. Recent surveys of the wild population on Ball’s Pyramid suggest that it is among the world’s rarest species. However, there are significant morphological differences between Ball’s Pyramid and museum specimens, and there has never been a genetic confirmation of the rediscovered population’s species identity. Because Dryococelus is monotypic, there are also no known extant relatives for comparison. Using shotgun genomic data from the Ball’s Pyramid population, we assembled a draft genome and the complete mitochondrial genome. We found that the genome is massive, over 4 Gb in size, and is most likely hexaploid. We re-sequenced mitochondrial genomes from historic museum specimens collected on Lord Howe Island before the extinction event. Sequence divergence between the two populations is less than 1% and is within the range of intraspecific differences between the museum specimens, suggesting that they are conspecific and that D. australis has successfully evaded extinction so far. This work highlights the importance of museum collections for taxonomic validation in the context of ongoing conservation efforts.

Concepts: Biology, Human Genome Project, Genome, Genomics, Insect, Phasmatodea, Extinction, Lord Howe Island


Pressure-sensitive adhesives such as tapes become easily contaminated by dust particles. By contrast, animal adhesive pads are able to self-clean and can be reused millions of times over a lifetime with little reduction in adhesion. However, the detailed mechanisms underlying this ability are still unclear. Here we test in adhesive pads of stick insects (Carausius morosus) (1) whether self-cleaning is enhanced by the liquid pad secretion, and (2) whether alternating push-pull movements aid the removal of particles. We measured attachment forces of insect pads on glass after contamination with 10 µm polystyrene beads. While the amount of fluid present on the pad showed no effect on the pads' susceptibility to contamination, the recovery of adhesive forces after contamination was faster when higher fluid levels were present. However, this effect does not appear to be based on a faster rate of self-cleaning since the number of spheres deposited with each step did not increase with fluid level. Instead, the fluid may aid the recovery of adhesive forces by filling in the gaps between contaminating particles, similar to the fluid’s function on rough surfaces. Further, we found no evidence that an alternation of pushing and pulling movements, as found in natural steps, leads to a more efficient recovery of adhesion than repeated pulling slides.

Concepts: Insect, Liquid, Carausius morosus, Phasmatodea, Adhesion, Adhesive, Dust, A Lifetime


Optimization of sensory processing during development can be studied by using photoreceptors of hemimetabolous insects (with incomplete metamorphosis) as a research model. We have addressed this topic in the stick insect Carausius morosus, where retinal growth after hatching is accompanied by a diurnal-to-nocturnal shift in behavior, by recording from photoreceptors of first instar nymphs and adult animals using the patch-clamp method. In the nymphs, ommatidia were smaller and photoreceptors were on average 15-fold less sensitive to light than in adults. The magnitude of A-type K(+) current did not increase but the delayed rectifier doubled in adults compared with nymphs, the K(+) current densities being greater in the nymphs. By contrast, the density of light-induced current did not increase, although its magnitude increased 8.6-fold, probably due to the growth of microvilli. Nymph photoreceptors performed poorly, demonstrating a peak information rate (IR) of 2.9 ± 0.7 bits/s versus 34.1 ± 5.0 bits/s in adults in response to white-noise stimulation. Strong correlations were found between photoreceptor capacitance (a proxy for cell size) and IR, and between light sensitivity and IR, with larger and more sensitive photoreceptors performing better. In adults, IR peaked at light intensities matching irradiation from the evening sky. Our results indicate that biophysical properties of photoreceptors at each age stage and visual behavior are interdependent and that developmental improvement in photoreceptor performance may facilitate the switch from the diurnal to the safer nocturnal lifestyle. This also has implications for how photoreceptors achieve optimal performance.

Concepts: Insect, Developmental biology, Performance, Carausius morosus, Hemimetabolism, Hemiptera, Entomology, Phasmatodea


We report that in a leaf insect, Phyllium westwoodii Wood-Mason (Phasmatodea: Phylliidae), two differing apertures can be used for oviposition, the color of eggs being affected by which aperture is used. Eggs which are forcibly propelled from the internal space within the valvulae of the abdomen are brown, whereas white eggs emerge slowly from the opening between the eighth sternite and the valvulae, and are deposited close to the ventral surface of the female. This unusual oviposition system does not appear to have been previously reported in phasmatids or in other insects.

Concepts: Insect, Thorax, Phasmatodea, Insects, Phylliidae


Many stick insects and mantophasmids possess tarsal ‘heel pads’ (euplantulae) covered by arrays of conical, micrometre-sized hairs (acanthae). These pads are used mainly under compression; they respond to load with increasing shear resistance, and show negligible adhesion. Reflected-light microscopy in stick insects (Carausius morosus) revealed that the contact area of ‘heel pads’ changes with normal load on three hierarchical levels. First, loading brought larger areas of the convex pads into contact. Second, loading increased the density of acanthae in contact. Third, higher loads changed the shape of individual hair contacts gradually from circular (tip contact) to elongated (side contact). The resulting increase in real contact area can explain the load dependence of friction, indicating a constant shear stress between acanthae and substrate. As the euplantula contact area is negligible for small loads (similar to hard materials), but increases sharply with load (resembling soft materials), these pads show high friction coefficients despite little adhesion. This property appears essential for the pads' use in locomotion. Several morphological characteristics of hairy friction pads are in apparent contrast to hairy pads used for adhesion, highlighting key adaptations for both pad types. Our results are relevant for the design of fibrillar structures with high friction coefficients but small adhesion.

Concepts: Force, Classical mechanics, Carausius morosus, Phasmatodea, Hair, Friction, Long hair, Contact area


Stick insects (Carausius morosus) have two distinct types of attachment pad per leg, tarsal “heel” pads (euplantulae) and a pre-tarsal “toe” pad (arolium). Here we show that these two pad types are specialised for fundamentally different functions. When standing upright, stick insects rested on their proximal euplantulae, while arolia were the only pads in surface contact when hanging upside down. Single-pad force measurements showed that the adhesion of euplantulae was extremely small, but friction forces strongly increased with normal load and coefficients of friction were [Formula: see text] 1. The pre-tarsal arolium, in contrast, generated adhesion that strongly increased with pulling forces, allowing adhesion to be activated and deactivated by shear forces, which can be produced actively, or passively as a result of the insects' sprawled posture. The shear-sensitivity of the arolium was present even when corrected for contact area, and was independent of normal preloads covering nearly an order of magnitude. Attachment of both heel and toe pads is thus activated partly by the forces that arise passively in the situations in which they are used by the insects, ensuring safe attachment. Our results suggest that stick insect euplantulae are specialised “friction pads” that produce traction when pressed against the substrate, while arolia are “true” adhesive pads that stick to the substrate when activated by pulling forces.

Concepts: Insect, Force, Shear stress, Carausius morosus, Phasmatodea, Friction


Little is known about the Phasmatodea gut microbial community, including whether phasmids have symbiotic bacteria aiding in their digestion. While symbionts are near ubiquitous in herbivorous insects, the Phasmatodea’s distinctively thin body shape precludes the gut enlargements needed for microbial fermentation. High-throughput sequencing was used to characterize the entire microbiota of the fat bodies, salivary glands, and anterior and posterior midguts of two species of walking stick.

Concepts: Archaea, Bacteria, Gut flora, Metabolism, Microbiology, Insect, Symbiosis, Phasmatodea


Determining the mechanical output of limb joints is critical for understanding the control of complex motor behaviours such as walking. In the case of insect walking, the neural infrastructure for single-joint control is well described. However, a detailed description of the motor output in form of time-varying joint torques is lacking. Here, we determine joint torques in the stick insect to identify leg joint function in the control of body height and propulsion. Torques were determined by measuring whole-body kinematics and ground reaction forces in freely walking animals. We demonstrate that despite strong differences in morphology and posture, stick insects show a functional division of joints similar to other insect model systems. Propulsion was generated by strong depression torques about the coxa-trochanter joint, not by retraction or flexion/extension torques. Torques about the respective thorax-coxa and femur-tibia joints were often directed opposite to fore-aft forces and joint movements. This suggests a posture-dependent mechanism that counteracts collapse of the leg under body load and directs the resultant force vector such that strong depression torques can control both body height and propulsion. Our findings parallel propulsive mechanisms described in other walking, jumping and flying insects, and challenge current control models of insect walking.

Concepts: Insect, Torque, Force, Classical mechanics, Phasmatodea, Reaction, Ground reaction force, Net force


The eggs of stick and leaf insects (Phasmatodea) bear strong resemblance to plant seeds and are commonly dispersed by females dropping them to the litter. Here we report a novel egg-deposition mode for Phasmatodea performed by an undescribed Vietnamese species of the enigmatic subfamily Korinninae that produces a complex egg case (ootheca), containing numerous eggs in a highly ordered arrangement. This novel egg-deposition mode is most reminiscent of egg cases produced by members of unrelated insect orders, e.g. by praying mantises (Mantodea) and tortoise beetles (Coleoptera: Cassidinae). Ootheca production constitutes a striking convergence and major transition in reproductive strategy among stick insects, viz. a shift from dispersal of individual eggs to elaborate egg concentration. Adaptive advantages of ootheca formation on arboreal substrate are likely related to protection against parasitoids and desiccation and to allocation of specific host plants. Our phylogenetic analysis of nuclear (28S, H3) and mitochondrial (COI, COII) genes recovered Korinninae as a subordinate taxon among the species-rich Necrosciinae with Asceles as sister taxon, thus suggesting that placement of single eggs on leaves by host plant specialists might be the evolutionary precursor of ootheca formation within stick insects.

Concepts: Species, Insect, Egg, Reproductive system, Cladistics, Phasmatodea, Taxonomic rank, Insects


Insects have various gaits with specific characteristics and can change their gaits smoothly in accordance with their speed. These gaits emerge from the embodied sensorimotor interactions that occur between the insect’s neural control and body dynamic systems through sensory feedback. Sensory feedback plays a critical role in coordinated movements such as locomotion, particularly in stick insects. While many previously developed insect models can generate different insect gaits, the functional role of embodied sensorimotor interactions in the interlimb coordination of insects remains unclear because of their complexity. In this study, we propose a simple physical model that is amenable to mathematical analysis to explain the functional role of these interactions clearly. We focus on a foot contact sensory feedback called phase resetting, which regulates leg retraction timing based on touchdown information. First, we used a hexapod robot to determine whether the distributed decoupled oscillators used for legs with the sensory feedback generate insect-like gaits through embodied sensorimotor interactions. The robot generated two different gaits and one had similar characteristics to insect gaits. Next, we proposed the simple model as a minimal model that allowed us to analyze and explain the gait mechanism through the embodied sensorimotor interactions. The simple model consists of a rigid body with massless springs acting as legs, where the legs are controlled using oscillator phases with phase resetting, and the governed equations are reduced such that they can be explained using only the oscillator phases with some approximations. This simplicity leads to analytical solutions for the hexapod gaits via perturbation analysis, despite the complexity of the embodied sensorimotor interactions. This is the first study to provide an analytical model for insect gaits under these interaction conditions. Our results clarified how this specific foot contact sensory feedback contributes to generation of insect-like ipsilateral interlimb coordination during hexapod locomotion.

Concepts: Insect, Arthropod, Interaction, Emergence, Oscillation, Feedback, Phasmatodea, Hexapoda