Beetle elytra: evolution, modifications and biological functions.

Conversion of forewings into hardened covers, elytra, was a ground-breaking morphological adaptation that has contributed to the extraordinary evolutionary success of beetles. Nevertheless, the knowledge of the functional aspects of these structures is still fragmentary and scattered across a large number of studies. Here, we have synthesized the presently available information on the evolution, development, modifications and biological functions of this crucial evolutionary novelty. The formation of elytra took place in the earliest evolution of Coleoptera, very likely already in the Carboniferous, and was achieved through the gradual process of progressive forewing sclerotization and the formation of inward directed epipleura and a secluded sub-elytral space. In many lineages of modern beetles, the elytra have been distinctly modified. This includes multiple surface modifications, a rigid connection or fusion of the elytra, or partial or complete reduction. Beetle elytra can be involved in a very broad spectrum of functions: mechanical protection of hind wings and body, anti-predator strategies, thermoregulation and water saving, water harvesting, flight, hind wing folding, diving and swimming, self-cleaning and burrow cleaning, phoresy of symbiotic organisms, mating and courtship, and acoustic communication. We postulate that the potential of the elytra to take over multiple tasks has enormously contributed to the unparalleled diversification of beetles.

Within- and Between-Species Variation of Wing Venation in Genus Monochamus (Coleoptera: Cerambycidae).

We have described the morphological variation of five Western Palaearctic species of Monochamus Dejean, 1821. The variation was assessed using wing measurements. Special emphasis was placed on the differences between Monochamus sartor (F., 1787) and Monochamus urussovii (Fischer-Waldheim, 1805). There was an interesting pattern of variation between the two species. Individuals of M. sartor from the Carpathians differed markedly from individuals of M. urussovii from Siberia, but individuals from north-eastern Poland were intermediate between those two populations. The intermediate individuals were more similar to the Siberian M. urussovii than to the Carpathian M. sartor despite the relatively large geographic distance between north-eastern Poland and Siberia. The occurrence of the intermediate individuals in north-eastern Poland may be the effect of hybridization between M. urussovii and M. sartor, which might have occurred after secondary contact between the two species in the Holocene.

Evolution, types, and distribution of flight control devices on wings and elytra in bark beetles.

Gaining the ability to fly actively was a ground-breaking moment in insect evolution, providing an unprecedented advantage over other arthropods. Nevertheless, active flight was a costly innovation, requiring the development of wings and flight muscles, the provision of sufficient energetic resources, and a complex flight control system. Although wings, flight muscles, and the energetic budget of insects have been intensively studied in the last decades, almost nothing is known regarding the flight-control devices of many crucial insect groups, especially beetles (Coleoptera). Here, we conducted a phylogenetic-informed analysis of flight-related mechanosensors in 28 species of bark beetles (Curculionidae: Scolytinae, Platypodinae), an economically and ecologically important group of insects characterized by striking differences in dispersal abilities. The results indicated that beetle flight apparatus is equipped with different functional types of mechanosensors, including strain- and flow-encoding sensilla. We found a strong effect of allometry on the number of mechanosensors, while no effect of relative wing size (a proxy of flight investment) was identified. Our study constitutes the first step to understanding the drivers and constraints of the evolution of flight-control devices in Coleoptera, including bark beetles. More research, including a quantitative neuroanatomical analysis of beetle wings, should be conducted in the future.

Climatic oscillations in Quaternary have shaped the co-evolutionary patterns between the Norway spruce and its host-associated herbivore.

During the Last Glacial Maximum in the Northern Hemisphere, expanding ice sheets forced a large number of plants, including trees, to retreat from their primary distribution areas. Many host-associated herbivores migrated along with their host plants. Long-lasting geographic isolation between glacial refugia could have been led to the allopatric speciation in separated populations. Here, we have studied whether the migration history of the Norway spruce Picea abies in Quaternary has affected its host-associated herbivorous beetle-Monochamus sartor. By using microsatellite markers accompanied by the geometric morphometrics analysis of wing venation, we have revealed the clear geographic structure of M. sartor in Eurasia, encompassing two main clusters: southern (Alpine-Carpathian) and eastern (including northeastern Europe and Asia), which reflects the northern and southern ecotypes of its host. The two beetles' lineages probably diverged during the Pleniglacial (57,000-15,000 BC) when their host tree species was undergoing significant range fragmentation and experienced secondary contact during post-glacial recolonization of spruce in the Holocene. A secondary contact of divergent lineages of M. sartor has resulted in the formation of the hybrid zone in northeastern Europe. Our findings suggest that the climatic oscillations during the Pleistocene have driven an insect-plant co-evolutionary process, and have contributed to the formation of the unique biodiversity of Europe.

Elytra reduction may affect the evolution of beetle hind wings.

Beetles are one of the largest and most diverse groups of animals in the world. Conversion of forewings into hardened shields is perceived as a key adaptation that has greatly supported the evolutionary success of this taxa. Beetle elytra play an essential role: they minimize the influence of unfavorable external factors and protect insects against predators. Therefore, it is particularly interesting why some beetles have reduced their shields. This rare phenomenon is called brachelytry and its evolution and implications remain largely unexplored. In this paper, we focused on rare group of brachelytrous beetles with exposed hind wings. We have investigated whether the elytra loss in different beetle taxa is accompanied with the hind wing shape modification, and whether these changes are similar among unrelated beetle taxa. We found that hind wings shape differ markedly between related brachelytrous and macroelytrous beetles. Moreover, we revealed that modifications of hind wings have followed similar patterns and resulted in homoplasy in this trait among some unrelated groups of wing-exposed brachelytrous beetles. Our results suggest that elytra reduction may affect the evolution of beetle hind wings.

Developmental Costs of Biological Invasion: The Exotic Wood Borer Tetropium fuscum (Coleoptera: Cerambycidae) is More Asymmetric and Smaller in Invaded Area.

Biological invasions provide a unique opportunity to gain insight into basic biological processes occurring under new circumstances. During the process of establishment, exotic species are exposed to various stressors which may affect their development. Presence of the stressors is often detected by measurements of left-right body asymmetry, which consists of two main components: fluctuating asymmetry and directional asymmetry. Fluctuating asymmetry constitutes random differences between the two body sides, whereas directional asymmetry occurs when a particular trait is bigger on one of the sides. The relation between these two asymmetry components is still not fully understood. Our goal was to investigate the potential differences in asymmetry patterns between native and invasive populations of Tetropium fuscum (Fabr. 1787) (Coleoptera: Cerambycidae), a harmful forest pest native to Europe and introduced to North America. Wing asymmetry assessment was based on the geometric morphometrics of hind wings. We found that specimens from invaded area were markedly smaller and have more asymmetric wings than individuals from native population, suggesting some unfavorable conditions in the invaded area. Moreover, we found significant directional asymmetry in the native but not in the invasive population. On the other hand, differences between left and right hind wings were similar in the native and invasive populations, in terms of direction. This suggests that a high level of fluctuating asymmetry in the invasive population may blur the intrinsic directional asymmetry and hinder its detection. Our data show that fluctuating asymmetry has a potential as an indicator of developmental stress in invasive species.

Hind wing morphology facilitates discrimination between two sibling species: Leiopus nebulosus and L. linnei (Coleoptera: Cerambycidae).

The study focused on two sibling beetle species: Leiopus nebulosus (Linnaeus, 1758) and L. linnei Wallin, Nylander & Kvamme, 2009. These species are very similar morphologically and their identification is difficult and possible only by experienced taxonomists. A supporting method for identification of L. nebulosus and L. linnei based on hind wings measurements was developed. The study was based on 115 specimens of L. linnei and 45 specimens of L. nebulosus. The correctness of identification of L. nebulosus amounted to 95.56%, and of L. linnei - 97.39%. The obtained model facilitates reliable identification of L. nebulosus and L. linnei also by less experienced entomologists. Geographical distributions of both species were summarized based on faunistic data from 39 scientific papers. The results show that both species have a Western-Palearctic distribution. Their distribution ranges are markedly overlapping. However, L. linnei is the species reported from larger number of localities, and observed more often.

Hind wing variation in Leptura annularis complex among European and Asiatic populations (Coleoptera, Cerambycidae).

The ability to quantify morphological variation is essential for understanding the processes of species diversification. The geometric morphometrics approach allows reliable description of variation in animals, including insects. Here, this method was used to quantify the morphological variation among European and Asiatic populations of Leptura annularis Fabricius, 1801 and its closely related species L. mimica Bates, 1884, endemic for Japan and Sakhalin islands. Since the taxonomic status of these two taxa is differently interpreted by taxonomists, they are collectively called "Leptura annularis complex" in this paper. The analysis was based on the measurements of hind wings of 269 specimens from six populations from Europe and Asia. The level of morphological divergence between most of continental European and Asiatic populations was relatively small and proportional to the geographic distance between them. However, distinct morphotype was detected in Sakhalin Is. and Japan. These data confirm the morphological divergence of the endemic L. mimica species. Obtained results highlight the potential of the geometric morphometric method in studying morphological variation in beetles.