Tanning of the tarsal and mandibular cuticle in adult Anax imperator (Insecta: Odonata) during the emergence sequence.

The arthropod cuticle offers strength, protection, and lightweight. Due to its limit in expandability, arthropods have to moult periodically to grow. While moulting is beneficial in terms of parasite or toxin control, growth and adaptation to environmental conditions, it costs energy and leaves the soft animal's body vulnerable to injuries and desiccation directly after ecdysis. To investigate the temporal change in sclerotization and pigmentation during and after ecdysis, we combined macrophotography, confocal laser scanning microscopy, scanning electron microscopy and histological sectioning. We analysed the tarsal and mandibular cuticle of the blue emperor dragonfly to compare the progress of tanning for structures that are functionally involved during emergence (tarsus/tarsal claws) with structures whose functionality is required much later (mandibles). Our results show that: (i) the tanning of the tarsal and mandibular cuticle increases during emergence; (ii) the tarsal cuticle tans faster than the mandibular cuticle; (iii) the mandibles tan faster on the aboral than on the oral side; and (iv) both the exo- and the endocuticle are tanned. The change in the cuticle composition of the tarsal and mandibular cuticle reflects the demand for higher mechanical stability of these body parts when holding on to the substrate during emergence and during first walking or hunting attempts.

Extrinsic and intrinsic musculature of the raptorial forelegs in Mantodea (Insecta) in the light of functionality and sexual dimorphism.

Prehensile raptorial forelegs are prey capturing and grasping devices, best known for praying mantises (Mantodea) within insects. They show strong morphological and behavioral adaptations toward a lifestyle as generalist arthropod predators. In the past, few species of Mantodea were investigated, concerning morphological variability of the raptorial forelegs. Especially the knowledge of foreleg anatomy in the light of functional and comparative morphology is scarce. Our comparative approach is based on the, for arthropods very common, "female-biased sexual size dimorphism" (SSD) that occurs in almost every Mantodea species. Within Mantodea, this SSD is likely leading to a shift of the exploited ecological niche between male and female individuals due to changes in, for example, the possible prey size; which might be reflected in the chosen ecomorphs. In this context, we analyzed the musculature of the raptorial forelegs of female and male specimens in five different species with varying SSD, using high-resolution microcomputed tomography and dissection. We were able to confirm the presence of 15 extrinsic and 15 intrinsic muscles-including one previously undescribed muscle present in all species. Thus, presenting a detailed description and illustrative three-dimensional anatomical visualization of the musculature in Mantodea. Interestingly, almost no observable differences were found, neither between species, nor between the sexes. Furthermore, we homologized all described muscles, due to their attachment points, to the comprehensive nomenclature established by Friedrich and Beutel (2008), discussed potential functionality of the muscles and possible homologies to the neuropteran Mantispa styriaca (Büsse et al., 2021) and the newly introduced leg nomenclature by Aibekova et al. (2022). By elucidating the anatomy, particularly in the context of functionality and SSD, our results complement previous knowledge of the raptorial forelegs, and facilitate a better understanding of the underlying biomechanical system of the predatory strike, and ultimately, a future comparison to other insect taxa.

A switchable joint in the head of dragonfly larvae (Insecta: Odonata) as key to the multifunctionality of the prehensile labial mask.

Dragonfly and damselfly larvae (Insecta: Odonata) capture prey by rapid protraction of a raptorial mouthpart, based on a modified labium. Yet, in insects with biting-chewing mouthparts, the labium has an essential role in food handling. These two distinct functions -prey capturing and handling-lead to a mechanical problem in Odonata larvae: while the labium is always protracted in a straight line during prey capture, food handling requires more dexterity. In this study, we investigate the role of the labium in the feeding process and analyse the mechanics of the labial joints in the dragonfly larva Anax imperator. Our results show that the labium features a multiaxial joint connecting the basal segment (postmentum) and the head. During feeding, a combination of rotations around different axes is used to handle and orient prey, which is unique among biting-chewing mouthparts. Furthermore, we identified structures at the joint which likely restrict lateral motion during the predatory strike. Our results provide a further understanding of the unique prey-capturing apparatus of odonate larvae capable of controlling a 'switchable' multiaxial to a restricted monoaxial joint. This concept highlights the evolution of a highly modified raptorial mouthpart appendage where the degrees of freedom can be actively restricted to allow for the respectively needed functionality.

Patients' awareness regarding the quality of their oral hygiene: development and validation of a new measurement instrument.

BACKGROUND: The present research aimed to develop and validate a standardised survey instrument for the assessment of patients' awareness of the quality of their oral hygiene performance. METHODS: A digital questionnaire was developed that assesses both patients' naïve self-perceptions of oral cleanliness (SPOCn) after tooth brushing and patients' perceptions after being informed how oral cleanliness may be captured in dentistry (SPOCd). Three studies (N = 56 adults, N = 66 adolescents and one of their parents, N = 24 university students) assessed the instrument's feasibility (patient reports), reliability (internal consistency), validity (correlation with other constructs; sensitivity to manipulation of actual tooth brushing), and the correlation with actual oral cleanliness after tooth brushing. RESULTS: All study groups accepted the questionnaire well; average answering times were less than 5 min. Cronbach's α exceeds 0.90; correlational analyses support the discriminant validity regarding oral hygiene related self-efficacy expectations and stages of change; manipulation of oral hygiene behaviour results in the expected changes of SPOC scores. Patients' SPOC correlate only moderately with actual oral cleanliness. The comparison between SPOCd scores and actual oral cleanliness indicate that they considerably overestimate their oral hygiene performance. CONCLUSIONS: The SPOC questionnaire is an easy-to-use, well-accepted, reliable and valid instrument for the assessment of patients' awareness of the quality of their oral hygiene for research and clinical purposes. The results of the questionnaire may help to reveal unrealistic self-perceptions of patients regarding their oral hygiene. It can raise their awareness of the need to improve their skills and/or efforts in this regard. Trial registration The third study was an interventional study and was registered in the appropriate national register ( www.drks.de ; ID: DRKS00018781; date of registration: 12/09/2019).

Louse flies holding on mammals' hair: Comparative functional morphology of specialized attachment devices of ectoparasites (Diptera: Hippoboscoidea).

Hippoboscidae and Nycteribiidae of the dipteran superfamily Hippoboscoidea are obligate ectoparasites, which feed on the blood of different mammals. Due to their limited flight capability, the attachment system on all tarsi is of great importance for a secure grasp onto their host and thus for their survival. In this study, the functional morphology of the attachment system of two hippoboscid species and two nycteribiid species was compared in their specificity to the host substrate. Based on data from scanning electron microscopy and confocal laser scanning microscopy, it was shown that the attachment systems of both Hippoboscidae and Nycteribiidae (Hippoboscoidea) differ greatly from that of other calyptrate flies and are uniform within the respective families. All studied species have an attachment system with two monodentate claws and two pulvilli. The claws and pulvilli of the Hippoboscidae are asymmetric, which is an adaptation to the fur of even-toed ungulates (Artiodactyla). The fur of these mammals possesses both, thinner woolen and thicker coat hair; thus, the asymmetry of the attachment system of the hippoboscid species enables a secure attachment to all surfaces of their hosts. The claws and pulvilli of the nyceribiid species do not show an asymmetry, since the fur of their bat (Chiroptera) hosts consists of hairs with the same thickness. The claws are important for the attachment to mammals' fur, because they enable a secure grip by mechanical interlocking of the hairs through the claws. Additionally, well-developed pulvilli are able to attach on thicker hairs of Artiodactyla or on smooth substrates such as the skin.

Functional morphology of cirri in the barnacle Amphibalanus improvisus (crustacea: Balanidae).

Barnacles rely heavily on their mobile cirri for food capture because of the sessile lifestyle. These filamentous food capturing devices are extended into the water current and perform undulating movements. Cuticular structures with corresponding musculature work together, to allow these highly repetitive movements. This paper studies the interplay between structure and function of the cirri using scanning electron microscopy (SEM), microcomputed tomography (µCT) and high-speed video recordings (HSV) in Amphibalanus improvisus. Barnacles use the external cuticle structures (denticles and setae) for efficient grooming, food capturing, and providing support for the muscular system responsible for movement control. Entanglement of the cirri during extension is probably avoided through an interlocking of the serrate setae on cirri IV-VI, creating a "zipper-like" effect, which was recorded here using the HSV. We analyzed the muscular arrangement using µCT and found a new flexor muscle in both the endo- and exopod of cirrus II. Supported by the intrinsic cirral muscles, the new flexor muscles may provide variable movements of the anterior cirri (cirri I-III), which is important for further food handling. Our results provide a foundation for further comparative studies of the feeding apparatus of barnacles and for possible implications in the area of bio-inspired robotics.

The mouthparts of the adult dragonfly Anax imperator (Insecta: Odonata), functional morphology and feeding kinematics.

Insects evolved differently specialized mouthparts. We study the mouthparts of adult Anax imperator, one of the largest odonates found in Central Europe. Like all adult dragonflies, A. imperator possesses carnivorous-type of biting-chewing mouthparts. To gain insights into the feeding process, behavior and kinematics, living specimens were filmed during feeding using synchronized high-speed videography. Additionally, the maximum angles of movement were measured using a measuring microscope and combined with data from micro-computed tomography. The resulting visualizations of the 3D-geometry of each mouthpart were used to study their anatomy and complement the existing descriptive knowledge of muscles in A. imperator to date. Furthermore, confocal laser scanning microscopy-projections allow for estimation of differences in the material composition of the mouthparts' cuticle. By combining all methods, we analyze possible functions and underlying biomechanics of each mouthpart. We also analyzed the concerted movements of the mouthparts; unique behavior of the mouthparts during feeding is active participation by the labrum and distinct movement by the maxillary laciniae. We aim to elucidate the complex movements of the mouthparts and their functioning by combining detailed information on (1) in vivo movement behavior (supplemented with physiological angle approximations), (2) movement ability provided by morphology (morphological movement angles), (3) 3D-anatomy, and (4) cuticle composition estimates.

Taxonomic note on the species status of Epiophlebiadiana (Insecta, Odonata, Epiophlebiidae), including remarks on biogeography and possible species distribution.

The species included in the genus Epiophlebia Calvert, 1903 represent an exception within Recent lineages - they do not belong to either dragonflies (Anisoptera) nor damselflies (Zygoptera). Nowadays, the genus is solely known from the Asian continent. Due to their stenoecious lifestyle, representatives of Epiophlebia are found in often very small relict populations in Nepal, Bhutan, India, Vietnam, China, North Korea, and Japan. We here present a taxonomic re-evaluation on the species status of Epiophlebiadiana Carle, 2012, known from the Sichuan province in China, supplemented with a morphological character mapping on a genetic tree to highlight synapomorphies of E.diana and E.laidlawi Tillyard, 1921. We conclude that E.diana is a junior synonym of E.laidlawi. Furthermore, we discuss the Recent distribution of the group, allowing for predictions of new habitats of representatives of this group.

Sand-throwing behaviour in pit-building antlion larvae: insights from finite-element modelling.

Sandy pitfall traps of antlions are elaborate constructions to capture prey. Antlions exploit the interactions between the particles in their habitat and build a stable trap. This trap is close to the unstable state; prey items will slide towards the centre-where the antlion ambushes-when entering the trap. This is efficient but requires permanent maintenance. According to the present knowledge, antlions throw sand, mainly to cause sandslides towards the centre of the pit. We hypothesized that: (i) sand-throwing causes sandslides towards the centre of the pit and (ii) sand-throwing constantly maintains the pitfall trap and thus keeps its efficiency high. Using laboratory experiments, as well as finite-element analysis, we tested these hypotheses. We show, experimentally and numerically, that sand that accumulates at the centre of the pit will be removed continuously by sand-throwing, this maintenance is leading to slope condition close to an unstable state. This keeps the slope angle steep and the efficiency of the trap constant. Furthermore, the resulting sandslides can relocate the trapped prey towards the centre of the pit. This study adds further insights from specific mechanical properties of a granular medium into the behavioural context of hunting antlion larvae.

A controllable dual-catapult system inspired by the biomechanics of the dragonfly larvae's predatory strike.

The biomechanics underlying the predatory strike of dragonfly larvae is not yet understood. Dragonfly larvae are aquatic ambush predators, capturing their prey with a strongly modified extensible mouthpart. The current theory of hydraulic pressure being the driving force of the predatory strike can be refuted by our manipulation experiments and reinterpretation of former studies. Here, we report evidence for an independently loaded synchronized dual-catapult system. To power the ballistic movement of a single specialized mouthpart, two independently loaded springs simultaneously release and actuate two separate joints in a kinematic chain. Energy for the movement is stored by straining an elastic structure at each joint and, possibly, the surrounding cuticle, which is preloaded by muscle contraction. As a proof of concept, we developed a bioinspired robotic model resembling the morphology and functional principle of the extensible mouthpart. Understanding the biomechanics of the independently loaded synchronized dual-catapult system found in dragonfly larvae can be used to control the extension direction and, thereby, thrust vector of a power-modulated robotic system.

Illuminating nature's beauty: modular, scalable and low-cost LED dome illumination system using 3D-printing technology.

Presenting your research in the proper light can be exceptionally challenging. Meanwhile, dome illumination systems became a standard for micro- and macrophotography in taxonomy, morphology, systematics and especially important in natural history collections. However, proper illumination systems are either expensive and/or laborious to use. Nowadays, 3D-printing technology revolutionizes lab-life and will soon find its way into most people's everyday life. Consequently, fused deposition modelling printers become more and more available, with online services offering personalized printing options. Here, we present a 3D-printed, scalable, low-cost and modular LED illumination dome system for scientific micro- and macrophotography. We provide stereolithography ('.stl') files and print settings, as well as a complete list of necessary components required for the construction of three differently sized domes. Additionally, we included an optional iris diaphragm and a sliding table, to arrange the object of desire inside the dome. The dome can be easily scaled and modified by adding customized parts, allowing you to always present your research object in the best light.

Step by step and frame by frame - Workflow for efficient motion tracking of high-speed movements in animals.

Analysing the motion of animals, especially at high speeds, is often challenging. Motion tracking software needs to deal with a variety of visual contexts, variable lighting conditions, heterogeneous backgrounds and even background movements. Here we present motion tracking via the easy to use and constantly updated Adobe After Effects software - which is often included in software packages most researchers are already using. The provided custom-made Javascript allows for easy exporting of tracking coordinates. Furthermore, some examples for analysing the obtained data in the open source statistical software 'R' will provide reference points, even for an unexperienced user. We present a step-by-step guide of the methodology using high-speed video recordings of locust jumps and additionally validate this method by successful tracking of simulated data under defined subpar filming conditions. This simulated data allows experienced users to compare the tracking software in use with the here presented workflow to weigh the advantages and disadvantages of any motion tracking software on the market.

Pressure-induced silk spinning mechanism in webspinners (Insecta: Embioptera).

The articulated appendages of arthropods are highly adaptable and potentially multifunctional, used for walking, swimming, feeding, prey capture, or other functions. Webspinners (Order Embioptera) are a paragon in this context. In contrast to other arthropods producing silk, they utilize their front feet for silk production. However, employing the same leg for alternative functions rather than for pure locomotion potentially imposes constraints and compromises. We here present morphological and experimental evidence for a "passive" pressure-induced silk spinning mechanism induced by external mechanical stimuli. Furthermore, we demonstrate that, as a consequence of the conflicting functions for their front feet, webspinners have evolved a unique style of walking that reduces the potentially problematic contact between silk ejectors and the substrate. Here we answer for the first time a long-term question within this enigmatic group of insects-how webspinners can use their front feet to spin their nanoscale silk. This knowledge may open the door for experimental studies on an artificial spinning process and for future utilization in applied fields of robotics or chemistry.

Adaptations of dragonfly larvae and their exuviae (Insecta: Odonata), attachment devices and their crucial role during emergence.

Moulting, especially in 'hemimetabolous' insects that emerge upside down, is a crucial moment in their live. Losing their attachment during this situation can be fatal for survival. We here studied the emergence of dragonfly adults, describe structures involved in larval attachment to the substrate, and biomechanically test the pull-off forces of exuviae to natural substrates. Confocal laser scanning microscopy and scanning electron microscopy were used to describe both morphology and material composition of the leg cuticle of Anax imperator larvae. The results show that the combination of morphological and behavioral adaptations provides reliable anchorage of exuviae to the substrates. We determined a safety factor of 14, and demonstrated that this staggered safety system experiencing several unlocking and relocking events withstand multiple disturbances before the entire exuvia is completely detaches. This furthers our understanding of interlocking and anchorage of insects in general and may allow for future applications.

Measurement error in µCT-based three-dimensional geometric morphometrics introduced by surface generation and landmark data acquisition.

Computed-tomography-derived (CT-derived) polymesh surfaces are widely used in geometric morphometric studies. This approach is inevitably associated with decisions on scanning parameters, resolution, and segmentation strategies. Although the underlying processing steps have been shown to potentially contribute artefactual variance to three-dimensional landmark coordinates, their effects on measurement error have rarely been assessed systematically in CT-based geometric morphometric studies. The present study systematically assessed artefactual variance in landmark data introduced by the use of different voxel sizes, segmentation strategies, surface simplification degrees, and by inter- and intra-observer differences, and compared their magnitude to true biological variation. Multiple CT-derived surface variants of the anuran (Amphibia: Anura) pectoral girdle were generated by systematic changes in the factors that potentially influence the surface geometries. Twenty-four landmarks were repeatedly acquired by different observers. The contribution of all factors to the total variance in the landmark data was assessed using random-factor nested permanovas. Selected sets of Euclidean distances between landmark sets served further to compare the variance among factor levels. Landmark precision was assessed by landmark standard deviation and compared among observers and days. Results showed that all factors, except for voxel size, significantly contributed to measurement error in at least some of the analyses performed. In total, 6.75% of the variance in landmark data that mimicked a realistic biological study was caused by measurement error. In this landmark dataset, intra-observer error was the major source of artefactual variance followed by inter-observer error; the factor segmentation contributed < 1% and slight surface simplification had no significant effect. Inter-observer error clearly exceeded intra-observer error in a different landmark dataset acquired by six partly inexperienced observers. The results suggest that intra-observer error can potentially be reduced by including a training period prior to the actual landmark acquisition task and by acquiring landmarks in as few sessions as possible. Additionally, the application of moderate and careful surface simplification and, potentially, also the use of case-specific optimal combinations of automatic local thresholding algorithms and parameters for segmentation can help reduce intra-observer error. If landmark data are to be acquired by several observers, it is important to ensure that all observers are consistent in landmark identification. Despite the significant amount of artefactual variance, we have shown that landmark data acquired from microCT-derived surfaces are precise enough to study the shape of anuran pectoral girdles. Yet, a systematic assessment of measurement error is advisable for all geometric morphometric studies.

Resilin in the flight apparatus of Odonata (Insecta)-cap tendons and their biomechanical importance for flight.

In Odonata, a direct flight mechanism with specialized tendons evolved. One particular adaptation, the implementation of the rubber-like protein resilin in these cap tendons, might be of major importance. Although resilin was first described in one tendon of Odonata, to our knowledge no comprehensive study about the presence of resilin in the thorax exists yet. We investigated various species of Odonata, using µCT, dissection and fluorescence microscopy. Here we show a complete mapping of the odonatan pterothorax, regarding the presence of tendons and their properties. Thus, 20-21 cap tendons in the pterothorax of Odonata show the presence of resilin. While performing outstanding and often-aggressive flight manoeuvres, resilin can provide shock absorption against mechanical damage from strong impacts. It may further improve the wear and fatigue resistance owing to resilin's damping behaviour. Additionally, resilin in tendons can absorb and return kinetic energy to restore muscles to their original shape after contracting and help in maintaining self-oscillation of the flight muscles. Here, the material distribution within the direct flight system of Odonata and the biomechanical importance and possible function of resilin are discussed. These results are an important step towards the understanding of the complex form-material-function interplay of the insect cuticle.

Material composition of the mouthpart cuticle in a damselfly larva (Insecta: Odonata) and its biomechanical significance.

Odonata larvae are key predators in their habitats. They catch prey with a unique and highly efficient apparatus, the prehensile mask. The mandibles and maxillae, however, play the lead in handling and crushing the food. The material composition of the cuticle in the biomechanical system of the larval mouthparts has not been studied so far. We used confocal laser scanning microscopy (CLSM) to detect material gradients in the cuticle by differences in autofluorescence. Our results show variations of materials in different areas of the mouthparts: (i) resilin-dominated pads within the membranous transition between the labrum and the anteclypeus, which support mobility and might provide shock absorption, an adaptation against mechanical damage; (ii) high degrees of sclerotization in the incisivi of the mandibles, where high forces occur when crushing the prey's body wall. The interaction of the cuticle geometry, the material composition and the related musculature determine the complex concerted movements of the mouthparts. The material composition influences the strength, mobility and durability of the cuticular components of the mouthparts. Applying CLSM for extracting information about material composition and material properties of arthropod cuticles will considerably help improve finite-element modelling studies.

The legs of "spider associated" parasitic primary larvae of Mantispa aphavexelte (Mantispidae, Neuroptera) - Attachment devices and phylogenetic implications.

The legs of the primary larva of Mantispa aphavexelte, parasite in egg sacks of spiders, were examined using scanning electron microscopy (SEM), histology and confocal laser scanning microscopy (CLSM). The leg morphology is described in detail, including intrinsic muscles. Functional adaptations of the leg attachment devices are discussed, especially regarding the material composition. For example, a sole-like flexible ventral tarsal surface containing resilin is combined with sclerotized pseudo-claws. This likely enables the larvae to cope with surface structures on the spider's body, with substrates on the ground, and also with various structural elements in the spider's nest. The leg morphology is evaluated with respect to phylogenetic affinities. A trumpet-shaped, elongated empodium has likely evolved early in the evolution of Neuroptera and may consequently belong to the groundplan of a large subgroup of the order. It characterizes most groups of the hemerobiform lineage and is also present in the myrmeleontiform Psychopsidae. The presence of a tarsal protrusion resembling a pretarsus confirms the monophyletic origin of Mantispoidea. A single fixed tooth and a specific surface structure are potential autapomorphies of Mantispidae. A distal tibial subunit partly separated from the main part of the leg segment is an apomorphy only described for larvae of M. aphavexelte.

Holding tight to feathers - structural specializations and attachment properties of the avian ectoparasite Crataerina pallida (Diptera, Hippoboscidae).

The louse fly Crataerina pallida is an obligate blood-sucking ectoparasite of the common swift Apus apus As a result of reduction of the wings, C. pallida is unable to fly; thus, an effective and reliable attachment to their host's plumage is of utmost importance. The attachment system of C. pallida shows several modifications in comparison to that of other calyptrate flies, notably the large tridentate claws and the dichotomously shaped setae located on the pulvilli. Based on data from morphological analysis, confocal laser scanning microscopy, cryo-scanning electron microscopy and attachment force experiments performed on native (feathers) as well as artificial substrates (glass, epoxy resin and silicone rubber), we showed that the entire attachment system is highly adapted to the fly's lifestyle as an ectoparasite. The claws in particular are the main contributor to strong attachment to the host. Resulting attachment forces on feathers make it impossible to detach C. pallida without damage to the feathers or to the legs of the louse fly itself. Well-developed pulvilli are responsible for the attachment to smooth surfaces. Both dichotomously shaped setae and high setal density explain high attachment forces observed on smooth substrates. For the first time, we demonstrate a material gradient within the setae, with soft, resilin-dominated apical tips and stiff, more sclerotized bases in Diptera. The empodium seems not to be directly involved in the attachment process, but it might operate as a cleaning device and may be essential to maintain the functionality of the entire attachment system.

Comparative morphology of the thorax musculature of adult Anisoptera (Insecta: Odonata): Functional aspects of the flight apparatus.

Due to their unique flight mechanism including a direct flight musculature, Odonata show impressive flight skills. Several publications addressed the details of this flight apparatus like: sclerites, wings, musculature, and flight aerodynamics. However, 3D-analysis of the thorax musculature of adult dragonflies was not studied before and this paper allows for a detailed insight. We, therefore, focused on the thorax musculature of adult Anisoptera using micro-computed tomography. Herewith, we present a comparative morphological approach to identify differences within Anisoptera: Aeshnidae, Corduliidae, Gomphidae, and Libellulidae. In total, 54 muscles were identified: 16 prothoracic, 19 mesothoracic, and 19 metathoracic. Recorded differences were for example, the reduction of muscle Idlm4 and an additional muscle IIIdlm1 in Aeshna cyanea, previously described as rudimentary or missing. Muscle Iscm1, which was previously reported missing in all Odonata, was found in all investigated species. The attachment of muscle IIpcm2 in Pantala flavescens is interpreted as a probable adaption to its long-distance migration behaviour. Furthermore, we present a review of functions of the odonatan flight muscles, considering previous publications. The data herein set a basis for functional and biomechanical studies of the flight apparatus and will therefore lay the foundation for a better understanding of the odonatan flight.