Mechanisms Modeling the Double Rotation of the Elytra in Beetles (Coleoptera)

We recorded transient movements,i.e.opening and closing,of beetle elytra.The beetles were tethered from below and filmed under a skew mirror; two markers were glued on each elytron at the apex and at the base.Body-fixed 3D traces of the apical and basal markers were reconstructed.The trace of the basal marker was,as a rule,non-parallel to the apical trace.The costal edge of the elytron uniformly supinated in the course of adduction of the apical marker.We found two essential attributes of double rotation:(1) the elytron to body articulation is approximately a spherical mechanism; (2) transient opening and closing possess single degree of freedom.The double rotation was modeled with two mechanisms:(1) a flexagon model of the Haas and Wootton's type simulated the elytral movement relative to the movement of one facet of the flexagon; (2) a screw and nut model provided traces as two sectors of a helical thread,one sector was phase shifted with respect to other one.Screw guideways in a spherical mechanism give rise to discrepancies.Exact solution for a spherical mechanism with two guideways was proposed.The modeling revealed the attribute (3):the elytron is actuated by two linked but differently directed drives.Experimental investigations on the elytron to body articulation may be oriented at search of those mechanisms.     

Indirect closing of elytra by the prothorax in beetles (Coleoptera): general observations and exceptions

Voluntary movements of the prothorax and the elytra in tethered flying beetles and manually induced movements of these parts in fresh dead beetles were recorded in 30 species representing 14 families. Participation of prothoracic elevation in the closing of the elytra was demonstrated in three ways. (i) The elevation was always simultaneous with elytral closing, in contrast to depression and elytral opening; a rare exception occurred in Lucanus cervus, whose elytra sometimes started to close before the cessation of wing strokes and the elevation of the prothorax. (ii) The manipulated elevation always induced closing of the spread elytra; the mechanical interaction between the hind edge of the pronotum and the roots of the elytra is a universal mechanism of closing the elytra in beetles. (iii) The prevention of pronoto-elytral contact in live beetles by the excision of the hind edge of the pronotum in front of the root prevented elytral closing after normal flight. Exceptions to this rule included some beetles that were able to close their elytra after such an excision: tiger beetles and diving beetles (seldomly) and rose chafers (always). This ability in Adephaga may be explained by attachments of the muscle actuating the 4th axillary plate, which differ from the attachments in Polyphaga. Cetoniinae open their elytra only by a small amount. It is proposed that their small direct adductors in combination with the elasticity of the sclerites are enough to achieve elytral closing without additional help from the prothorax.     

Lehr's fields of campaniform sensilla in beetles (Coleoptera): Functional morphology. III. Modification of elytral mobility or shape in flying beetles

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. 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.     

Lehr's fields of campaniform sensilla in beetles (Coleoptera): Functional morphology. II. Wing reduction and the sensory field

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.     

Coupling between elytra of some beetles: Mechanism, forces and effect of surface texture

Lightweight materials, structures and coupling mechanisms are very important for realizing advanced flight vehicles. Here, we obtained the geometric structures and morphologies of the elytra of beetles and ascertained its coupling zone by using the histological section technique and SEM. We set up a three-dimensional motion observing system to monitor the opening and closing behaviour of elytra in beetles and to determine the motion mechanism. We constructed a force measuring system to measure the coupling forces between elytra. The results show that elytra open and close by rotating about a single axle, where the coupling forces may be as high as 160 times its own bodyweight, the elytra coupling with the tenon and mortise mechanism, surface texture and opening angle between elytra heavily influence the coupling forces. These results may provide insights into the design mechanism and structure for future vehicles of flight.     

Passive aerodynamic stabilization by beetle elytra (wing covers)

Summary. Most beetles extend their elytra laterally during flight, where they can act as fixed airfoils; typically, such elytra are held at a pronounced dihedral angle. We fixed extended elytra of June beetles (Scarabaeidae: Melolonthinae) and tiger beetles (Cicindelidae) over a range of dihedral angles (-10° to 45°) and towed them in a water tunnel. We also towed them with flexed elytra (i.e. in the resting position). (Reynolds number was used to maintain dynamic similarity with air.) We measured the maximum rolling angle and the lateral oscillation rate as two indices of stability. There was no relationship between oscillation rate and dihedral angle, or between maximum roll angle and dihedral angle. At all dihedral angles tested, however, beetles were much more stable than when they were towed with the elytra flexed. We also included a third beetle type, flower beetle (Scarabaeidae: Cetoniine), which normally flies with flexed elytra. We measured stability indices for these beetles with elytra only in the flexed position. When all beetles were tested with flexed elytra, flower beetles showed no difference in maximum roll angle, but more stability in oscillation rate than the other two beetle types.     

Lehr's fields of campaniform sensilla in beetles (Coleoptera): Functional morphology. I. General part and allometry

In this first of three articles we show the 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. The 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 the 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 decrease of the elytral area.     

Physiological colour change in the elytra of the hercules beetle, Dynastes hercules

The male of the hercules beetle, Dynastes hercules, is able to change the colour of its elytra from yellowish to black and back again to yellowish within a few minutes. The epicuticle of the elytra is transparent and about 3 μm thick. Below it is a yellow spongy layer that is usually about 5 μm thick. The cuticle below the yellow sponge is black. When the layer of yellow sponge is air filled it becomes optically heterogeneous, and the light reflected from the elytra is yellow. When the yellow sponge is liquid filled it becomes optically homogeneous, and the black cuticle below is seen.If a beetle that has yellowish elytra is placed in a saturated atmosphere, the elytra become black. When the relative humidity is appreciably reduced, yellow patches begin to appear on the elytra, usually within 30 sec to 2 min. However, if the beetle is kept at a constant relative humidity that previously caused yellowing, it will become black given enough time. Most colour changes observed were clearly in response to changes in the ambient humidity and were not affected when the beetles were kept in the light or in total darkness nor by blackening their eyes or prodding them or exposing them to sounds of different intensities or frequencies.If an elytron is removed from a live beetle, it changes colour in response to changes in relative humidity exactly like the elytron left attached. When a restricted area of the elytra is subjected to a humidity that normally causes blackening and an adjacent area to a humidity that normally causes yellowing, both change colour in the expected way. This local control of colour change seems to preclude hormonal control. It is suggested that the epidermal cells or both the epidermal and blood cells in the elytra are responsible for the hydration and dehydration of the layer of yellow sponge.     

Differences in loading of the temporomandibular joint during opening and closing of the jaw

Kinematics of the human masticatory system during opening and closing of the jaw have been reported widely. Evidence has been provided that the opening and closing movement of the jaw differ from one another. However, different approaches of movement registration yield divergent expectations with regard to a difference in loading of the temporomandibular joint between these movements. Because of these diverging expectations, it was hypothesized that joint loading is equal during opening and closing. This hypothesis was tested by predicting loading of the temporomandibular joint during an unloaded opening and closing movement of the jaw by means of a three-dimensional biomechanical model of the human masticatory system. Model predictions showed that the joint reaction forces were markedly higher during opening than during closing. The predicted opening trace of the centre of the mandibular condyle was located cranially of the closing trace, with a maximum difference between the traces of 0.45 mm. The hypothesis, postulating similarity of joint loading during unloaded opening and closing of the jaw, therefore, was rejected. Sensitivity analysis showed that the reported differences were not affected in a qualitative sense by muscular activation levels, the thickness of the cartilaginous layers within the temporomandibular joint or the gross morphology of the model. Our predictions indicate that the TMJ is loaded more heavily during unloaded jaw opening than during unloaded jaw closing.     

DISTRIBUTION OF TRABECULAE AND ELYTRAL SURFACE STRUCTURES OF THE HORNED BEETLE, ALLOMYRINA DICHOTOMA (LINNÉ) (COLEOPTERA: SCARABAEIDAE)

Abstract To clarify the dynamic construction of Allomyrina dichotoma (Linné) elytra, the distribution of trabeculae and surface structures has been investigated using scanning electron microscopy and transmitted light. There are solid trabeculae in the elytron and under penetrating light these can be seen as black dots. It is clear that trabeculae arrangement is almost entirely irregular throughout the elytron, except for some approximately straight alignment near some trachea. This irregular arrangement is different from the longitudinal rows of striations that are well known in other species, and there are no hollowed striae (punctures) on the elytral surface of A. dichotoma . Throughout the internal architecture of the elytron, there are mesh-like (honeycomb) structures. Each honeycomb usually has 1–2 trabeculae mainly distributed at the corners of the honeycomb. The number of trabeculae present on each honeycomb is dependent on the size of the honeycomb.     

Surface Chemicals Inform about Sex and Breeding Status in the Biparental Burying Beetle Nicrophorus vespilloides

The multicomponent nature of chemical cues and signals are not very well understood. One reason for the often found complexity of chemical blends might be that they provide multiple messages. Burying beetles which use vertebrate carcasses as food for their larvae and defend these carcasses against inter- and intraspecific competitors are able to recognise the sex and breeding status of conspecifics. Studies have shown that the chemical composition of cuticular lipids is correlated with sex and breeding status, but there is no definitive evidence that these chemicals function in recognition. In the present study, we performed behavioural bioassays to directly asses the role of chemical cues in the recognition system of the burying beetle Nicrophorus vespilloides . After finding a carcass, females were more tolerant of dead males than of females. The behaviour was reversed when a solvent extract from the opposite sex was applied. An earlier experiment had shown that females breeding on a carcass treat non-breeding males more aggressively than breeding ones. In the present study, we could trigger the same dichotomous behaviours by presenting a single elytron from a breeding and a non-breeding beetle. In an additional experiment, females tolerated the elytra of non-breeding beetles if we had first applied an extract from a breeding beetle to these elytra. Our study is the first behavioural proof that female burying beetles obtain multiple information from chemical cues.     

Flow Visualization of Rhinoceros Beetle (Trypoxylus dichotomus) in Free Flight

Aerodynamic characteristics of the beetle,Trypoxylus dichotomus,which has a pair of elytra (forewings) and flexible hind wings,are investigated.Visualization experiments were conducted for various flight conditions of a beetle,Trypoxylus dichotomus:free,tethered,hovering,forward and climbing flights.Leading edge,trailing edge and tip vortices on both wings were observed clearly.The leading edge vortex was stable and remained on the top surface of the elytron for a wide interval during the downstroke of free forward flight.Hence,the elytron may have a considerable role in lift force generation of the beetle.In addition,we reveal a suction phenomenon between the gaps of the hind wing and the elytron in upstroke that may improve the positive lift force on the hind wing.We also found the reverse clap-fling mechanism of the T.dichotomus beetle in hovering flight.The hind wings touch together at the beginning of the upstroke.The vortex generation,shedding and interaction give a better understanding of the detailed aerodynamic mechanism of beetle flight.     

纹吉丁属及其近缘种鞘翅超微结构的比较研究(鞘翅目,吉丁科)

比较研究了8种纹吉丁和7种近缘属种类鞘翅刻点的超微结构,研究表明纹吉丁属鞘翅刻点结构具有属、种特异性.纹吉丁属不同种类鞘翅刻点的形态相似,仅大小、密度和弯曲程度不同.纹吉丁与其近缘属种类鞘翅刻点的形态结构差异显著.同时表明,外部形态相似的类群,鞘翅刻点的形态结构未必相似,体现了吉丁鞘翅形态结构的多样性.     

The Role of Elytra in Beetle Flight: I. Generation of Quasi-Static Aerodynamic Forces

We conducted a comprehensive study to investigate the aerodynamic characteristics and force generation of the elytra of abeetle,Allomyrina dichotoma.Our analysis included wind tunnel experiments and three-dimensional computational fluiddynamics simulations using ANSYS-CFX software.Our first approach was a quasi-static study that considered the effect ofinduced flapping flow due to the flapping motion of the fore-wings (elytra) at a frequency of around 30 Hz to 40 Hz.The dihedralangle was varied to represent flapping motion during the upstroke and downstroke.We found that an elytron producespositive lift at 0° geometric angle of attack,negative lift during the upstroke,and always produces drag during both the upstrokeand downstroke.We also found that the lift coefficient of an elytron does not drop even at a very high geometric angle of attack.For a beetle with a body weight of 5 g,based on the quasi-static method,the fore-wings (elytra) can produce lift of less than 1%of its body weight.     

Righting kinematics in beetles (Insecta: Coleoptera)

Twenty modes of stereotyped righting motions were observed in 116 representative species of coleoptera. Methods included cine and stereocine recording with further frame by frame analysis, stereogrammetry, inverse kinematic reconstruction of joint angles, stroboscopic photography, recording of electromyograms, 3D measurements of the articulations, etc. The basic mode consists of a search phase, ending up with grasping the substrate, and a righting, overturning phase. Leg coordination within the search cycle differs from the walking cycle with respect to phasing of certain muscle groups. Search movements of all legs appear chaotic, but the tendency to move in antiphase is still present in adjacent ipsilateral and contralateral leg pairs. The system of leg coordination might be split: legs of one side might search, while contralateral legs walk, or fore and middle legs walk while hind legs search. Elaborated types of righting include somersaults with the aid of contralateral or diagonal legs, pitch on elytra, jumps with previous energy storage with the aid of unbending between thoracic segments (well-known for Elateridae), or quick folding of elytra (originally described in Histeridae). Righting in beetles is compared with righting modes known in locusts and cockroaches. Search in a righting beetle is directed dorsad, while a walking insect searches for the ground downwards. Main righting modes were schematized for possible application to robotics.     

Die Bedeutung der Elytren bei Vertretern desMelolontha-Flugtyps (Coleoptera)

The function of the flapping elytra was investigated in garden shafers (Melolontha melolontha L.) and rhinoceros beetles (Oryctes boas Fabr.).

FINE STRUCTURE OF TRABECULA IN THE ELYTRA OF ALLOMYRINA DICHOTOMA (LINNE) AND PROSOPOCOILUS INCLINATUS (MOTSCHULSKY) (COLEOPTERA: SCARABAEEDAE)

Abstract To obtain new ideas on optimal composite material structure, the fine structures of the trabeculae of the elytra of Allomyrina dichotoma (Iinné) and Prosopocoilus inclinatus (Motschulsky) were investigated using scanning electron microscope. The shape and size of trabeculae, which form the internal bridges between the upper and lower surfaces of the elytra, depend on the species of beetles. Complex surface structures (stripes or bandings) were found in those of A. dichotoma , but less in P. inclinatus . The trabeculae consisted of three parts: the surface part, the cylindrical layer and the central part. The surface and the central part were mainly protein substances which could be dissolved by KOH. The cylindrical layer had many chitin fibers with different orientation, and they were embedded in a protein matrix, and were connected to endocuticle chitin fibers on the upper and lower parts of the lamination.     

Impact of environmental factors on the body shape variation and sexual shape dimorphism in Carabus granulatus L.(Coleoptera: Carabidae)

Despite plenty of data in insects shape variation,papers on environmental factors effect on such variation are scarce and in ground beetles are practically absent.The aim of this research was to:(i) model the effect of the region of habitation,urbanization and habitat vegetation into the shape variation in widespread carabid species;(ii) describe sexual shape dimorphism in studied species.Samples were pitfall trapped in different regions of its area in the spectrum of anthropogenic influence (cities,suburbs,natural biotopes).One thousand and one hundred sixty-eight specimens were analyzed for six morphometric traits and terminal points of those measurements were used as landmarks for Procrustes analysis.We used linear models to reveal which factor (region,urbanization or vegetation) and in what direction affected beetles shape.Results showed that males in C.granulatus had more convex elytra and head,and the more convex in apical-basal direction pronotum.In its area from the east to the west elytra flattened in medial-distal direction and pronotum flattened in apical-distal direction.In disturbed urban environment beetles pronotum became more convex in both sexes and males elytra became more convex too.In open habitats (meadows,lawns) beetles became more flattened.     

海南皱鞘天牛属一新种（鞘翅目：天牛科，沟胫天牛亚科）

记述采自海南的沟胫天牛亚科皱鞘天牛属Parapolytrechus Breuning一新种：黄跗皱鞘天牛P. flavotarsus Wang et Zheng, sp.nov.。模式标本保存于南开大学生物系昆虫标本室。     

Hearing in tiger beetles (Cicindelidae)

ABSTRACT. Tympanic hearing organs (ears) are reported for several tiger beetle (Cicindelidae) species. The paired ears are positioned bilaterally on the first abdominal tergum and consist of cavities covered by thin tympana. When the beetle is not flying the elytra covers its ears and reduces their sensitivity to sound. However, when the beetle is flying, its exposed ears are capable of detecting ultrasonic pulses. Under a microscope, beetles with their elytra artificially raised contract their abdomens in response to ultrasound. Ultrasonic emissions directed toward flying beetles induce them immediately to fly downward and land, a response which probably aids escape from predators, particularly echolocating bats. Other possible uses for the ears are the avoidance of diurnal insect predators and intraspecific communication.