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Journal: Arthropod structure & development


Mate choice, copulation, genital morphology, and sperm storage are not very well understood in millipedes. The use of three-dimensional x-ray computed tomography (μCT) provides new morphological data regarding millipede reproductive systems in both the female and male, including chitinous sclerites and membranes, muscles, glands, oviducts, and sperm conduits. Here we present a complete integrated account of the morphology and function of the female genital organs in the family Polydesmidae (Diplopoda: Polydesmida) using μCT, UV fluorescence imaging, and scanning electron microscopy. These data allow us to consider competing hypotheses regarding millipede vulva formation. We additionally present the morphology of copulatory interface in Pseudopolydesmus Attems, 1898 using images of a mating pair in copula and by simulating the interface of the organs using 3D models from μCT, allowing us to tentatively identify a lock-and-key-like mechanism. Finally, we use μCT to reveal the topology of the seminal canal in the gonopod of male Pseudopolydesmus, a topic that has remained unresolved for nearly 80 years.


Heteropteran legs are very diverse within and among taxa, and such variation is frequently correlated with life habits. Structural modifications are commonly present in the legs of the Pentatomoidea but are poorly studied. Using scanning electron microscopy, the tibia and pretarsal microstructure of 82 species of Pentatomidae (Heteroptera), three species of Scutelleridae, and ten species of Thyreocoridae were described, focusing on the pretarsal structure, the foretibial apparatus, and the foretibial comb. The Pentatomidae, the Scutelleridae, and the Thyreocoridae have uniform pretarsal structures. Variation can be found in the length of the parempodial setae and in the shape of the parempodial projections. The foretibial combs of the Pentatomidae, the Thyreocoridae, and the Scutelleridae are described for the first time, and we have demonstrated that there is low structural variation in the foretibial comb complex of the studied species. The setae organization and distribution on the foretibial apparatus is uniform in the families studied. However, the Asopinae (Pentatomidae) bear a foretibial apparatus that is uniquely organized. The taxonomic and phylogenetic relevance of the pretarsal traits, the foretibial apparatus, and the foretibial comb are discussed.

Concepts: Electron, Species, Structure, Scanning electron microscope, Organization, Hemiptera, Heteroptera, Pentatomidae


Loss of the flight ability and wing reduction has been reported for many taxa of Coleoptera. If elytra are closed, their roots are clenched between the tergum and the pleuron, forces applied to the elytra can not be transmitted to the field of campaniform sensilla situated on the root. That is why it is plausible to assume that the field becomes redundant in non-flying beetles. We examined the relationships between the hind wing reduction and characters of this mechanosensory field in beetles of six families. We measured the size of the elytron, that of the hind wing and counted the number of sensilla in the sensory field. Mesopterous non-flying beetles retain one half to one third of sensilla present in macropterous species of the same body size. Further reduction of the sensory field in brachypterous species is obvious, but sensilla are still present in insects with strongly reduced wings, as long as their elytra are separable and mesothoracic axillaries are present. Complete loss of sensilla coincides with the existence of a permanent sutural lock. However, some beetles with permanently locked elytra and absence of axillaries still retain few campaniform sensilla. A very special case of an extreme wing modification in feather-wing beetles is considered. No sensilla were revealed either on the root of the elytron or on the basal segment of such fringed wings in flying ptiliid species.

Concepts: Insect, Fungus, Beetle, Field, Wing, Insects, Insect wing, Elytron


In this first of three articles we show construction of the articular part of the elytron, the root. The root bears a conspicuous field of campaniform sensilla. This field was studied using light and scanning electron microscopes. Diversity of shape of the field among beetles, types of orientation of elongated sensilla within the field, individual variability of their number among conspecifics are demonstrated. Elongated sensilla point to the junction of the elytron with the second axillary plate. Presumably, they monitor twist movement in this junction which is possible if the elytron is open. The goal of the whole project is to reveal the effect of both structure and function of the hind wings and elytra on morphology of this mechanosensory field. Our data on allometric relationships between the animal size and quantitative characteristics of the field in normally flying beetles provide an important background for further functional analysis of this sensory organ. We selected 14 series of several species belonging to the same taxon but differing in size from big to small. It is revealed that the area of the sensory field is directly proportional to the elytral area, whereas the number of sensilla is proportional to the square root of the elytral area. Despite the great range in the elytral area (1500 times) in series of selected species the area of an external pit or cap of a single sensillum varies only 25-fold. The density of sensilla per unit area of the sensory field increases with a decrease of the elytral area.

Concepts: Function, Visual system, Vector space, Beetle, Circle, Scanning electron microscope, Complex number, Elytron


The Heteroptera show a diversity of glands associated with the epidermis. They have multiple roles including the production of noxious scents. Here, we examine the cellular arrangement and cytoskeletal components of the scent glands of pentatomoid Heteroptera in three families, Pentatomidae (stink bugs), Tessaratomidae, and Scutelleridae (shield-backed bugs or jewel bugs). The glands are; (1) the dorsal abdominal glands, (2) the tubular glands of the composite metathoracic gland, and (3) the accessory gland component of the composite metathoracic gland. The dorsal abdominal glands are at their largest in nymphs and decrease in size in adults. The metathoracic gland is an adult-specific gland unit with a reservoir and multiple types of gland cells. The accessory gland is composed of many unicellular glands concentrated in a sinuous line across the reservoir wall. The lateral tubular gland is composed of two-cell units. The dorsal abdominal glands of nymphs are composed of three-cell units with a prominent cuticular component derived from the saccule cell sitting between the duct and receiving canal. The cuticular components that channel secretion from the microvilli of the secretory cell to the exterior differ in the three gland types. The significance of the numbers of cells comprising gland units is related to the role of cells in regenerating the cuticular components of the glands at moulting in nymphs.

Concepts: Secretion, Skin, Gland, Exocrine gland, Component video, Pentatomoidea, Tubular gland, Shield bugs


Some flying beetles have peculiar functional properties of their elytra, if compared with the vast majority of beetles. A “typical” beetle covers its pterothorax and the abdomen from above with closed elytra and links closed elytra together along the sutural edges. In the open state during flight, the sutural edges diverge much more than by 90°. Several beetles of unrelated taxa spread wings through lateral incisions on the elytra and turn the elytron during opening about 10-12° (Cetoniini, Scarabaeus, Gymnopleurus) or elevate their elytra without partition (Sisyphus, Tragocerus). The number of campaniform sensilla in their elytral sensory field is diminished in comparison with beetles of closely related taxa lacking that incision. Elytra are very short in rove beetles and in long-horn beetles Necydalini. The abundance of sensilla in brachyelytrous long-horn beetles Necydalini does not decrease in comparison with macroelytrous Cerambycinae. The strong reduction of the sensory field was found in brachyelytrous Staphylinidae. Lastly, there are beetles lacking the linkage of the elytra down the sutural edge (stenoelytry). Effects of stenoelytry were also not uniform: Oedemera and flying Meloidae have the normal amount of sensilla with respect to their body size, whereas the sensory field in the stenoelytrous Eulosia bombyliformis is 5-6 times less than in chafers of the same size but with normally linking broad elytra.

Concepts: Normal distribution, Beetle, Elytron, Polyphaga, Aleochara, Rove beetle, Blister beetle, Staphylinidae


The gill and paranotal lobe theories of insect wing evolution were both proposed in the 1870s. For most of the 20th century, the paranotal lobe theory was more widely accepted, probably due to the fundamentally terrestrial tracheal respiratory system; in the 1970s, some researchers advocated for an elaborated gill (“pleural appendage”) theory. Lacking transition fossils, neither theory could be definitively rejected. Winged insects are abundant in the fossil record from the mid-Carboniferous, but insect fossils are vanishingly rare earlier, and all earlier fossils are from primitively wingless insects. The enigmatic, isolated mandibles of Rhyniognatha (early Devonian) hint that pterygotus may have been present much earlier, but the question remains open. In the late 20th century, researchers used models to study the interaction of body and protowing size on solar warming and gliding abilities, and stability and glide effectiveness of many tiny adjustable winglets versus a single, large pair of immobile winglets. Living stoneflies inspired the surface-skimming theory, which provides a mechanism to bridge between aquatic gills and flapping wings. The serendipitously discovered phenomenon of directed aerial descent suggests a likely route to the early origin of insect flight. It provides a biomechanically feasible sequence from guided falls to fully-powered flight.

Concepts: Evolution, Insect, Respiratory system, Fossil, Wing, Insect wing, Flying and gliding animals, Insect flight


Arthropods operate in an outrageous diversity of environments. From the deep sea to dense tropical forests, to wide open arctic tundra, they have colonized almost every possible habitat. Within these environments, the presence of light is nearly ubiquitous, varying in intensity, wavelength, and polarization. Light provides critical information about the environment, such as time of day or where food sources may be located. Animals take advantage of this prevalent and informative cue to make behavioral choices. However, the types of choices animals face depend greatly on their environments and needs at any given time. In particular, animals that undergo metamorphosis, with arthropods being the prime example, experience dramatic changes in both behavior and ecology, which in turn may require altering the structure and function of sensory systems such as vision. Amphibiotic organisms maintain aquatic lifestyles as juveniles before transitioning to terrestrial lifestyles as adults. However, light behaves differently in water than in air, resulting in distinct aquatic and terrestrial optical environments. Visual changes in response to these optical differences can occur on multiple levels, from corneal structure down to neural organization. In this review, we summarize examples of alterations in the visual systems of amphibiotic larval and adult insects and malacostracan crustaceans, specifically those attributed to environmental differences between metamorphic phases.


The ability of ant colonies to transport, store, and distribute food resources through trophallaxis is a key advantage of social life. Nonetheless, how the structure of the digestive system has adapted across the ant phylogeny to facilitate these abilities is still not well understood. The crop and proventriculus, structures in the ant foregut (stomodeum), have received most attention for their roles in trophallaxis. However, potential roles of the esophagus have not been as well studied. Here, we report for the first time the presence of an auxiliary thoracic crop in Pheidole aberrans and Pheidole deima using X-ray micro-computed tomography and 3D segmentation. Additionally, we describe morphological modifications involving the endo- and exoskeleton that are associated with the presence of the thoracic crop. Our results indicate that the presence of a thoracic crop in major workers suggests their potential role as repletes or live food reservoirs, expanding the possibilities of tasks assumed by these individuals in the colony. Our contribution emphasizes the utility of combining data from external and internal morphology to better understand functional and behavioral mechanisms.


Posture and walking require support of the body weight, which is thought to be detected by sensory receptors in the legs. Specificity in sensory encoding occurs through the numerical distribution, size and response range of sense organs. We have studied campaniform sensilla, receptors that detect forces as strains in the insect exoskeleton. The sites of mechanotransduction (cuticular caps) were imaged by light and confocal microscopy in four species (stick insects, cockroaches, blow flies and Drosophila). The numbers of receptors and cap diameters were determined in projection images. Similar groups of receptors are present in the legs of each species (flies lack Group 2 on the anterior trochanter). The number of receptors is generally related to the body weight but similar numbers are found in blow flies and Drosophila, despite a 30 fold difference in their weight. Imaging data indicate that the gradient (range) of cap sizes may more closely correlate with the body weight: the range of cap sizes is larger in blow flies than in Drosophila but similar to that found in juvenile cockroaches. These studies support the idea that morphological properties of force-detecting sensory receptors in the legs may be tuned to reflect the body weight.