New beetles (Insecta,Coleoptera) from the Lower Triassic of European Russia

Fossil remains of beetles are described from two Lower Triassic localities: Entala (Induan) and Tikhvinskoe (Olenekian). Only one beetle fossil was previously known from the Lower Triassic of Tikhvinskoe. The fossils are rather few and poorly preserved, but they are worth describing as finds rare for the Lower Triassic. Five fossils from Entala most probably belong to beetles of the same species of the formal genus Pseudochrysomelites. Beetles of this genus are especially abundant in deposits close to the Permian–Triassic boundary and can be considered “disaster taxa.” There are no known cases, either in the Permian or in the Middle–Upper Triassic, of a random sample of five specimens belonging to a single species. This suggests that in the Entala oryctocenosis the species diversity of beetles is extremely low. All three beetle fossils found in Tikhvinskoe belong to beetles of different species, showing that diversity had already started to increase. However, it remained low, and all fossils belong to the formal family Schizocoleidae, and two of the three belong to the same genus, Pseudochrysomelites. The Khei-Yaga locality, which immediately follows Tikhvinskoe in time (topmost Olenekian or early Anisian), already contains beetles of the families Asiocoleidae and Permosynidae. In the Lower Anisian of the Buntsandstein, such typical Mesozoic beetles as Cupedidae and Coptoclavidae have been recorded. The appearance of such advanced beetles as early as the Lower Anisian suggests that the famous Permian–Triassic crisis was not as deep as it is usually believed, and many beetles survived it, disappearing, however, from the fossil record in the Early Triassic.     

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.     

A catalogue of Lithuanian beetles (Insecta, Coleoptera)

This paper presents the first complete and updated list of all 3597 species of beetles (Insecta: Coleoptera) belonging to 92 familiesfound and published in Lithuania until 2011, with comments also provided on the main systematic and nomenclatural changes since the last monographic treatment in two volumes (Pileckis and Monsevičius 1995, 1997). The introductory section provides a general overview of the main features of the territory of Lithuania, the origins and formation of the beetle fauna and their conservation, the faunistic investigations in Lithuania to date revealing the most important stages of the faunistic research process with reference to the most prominent scientists, an overview of their work, and their contribution to Lithuanian coleopteran faunal research.Species recorded in Lithuania by some authors without reliable evidence and requiring further confirmation with new data are presented in a separate list, consisting of 183 species. For the first time, analysis of errors in works of Lithuanian authors concerning data on coleopteran fauna has been conducted and these errors have been corrected. All available published and Internet sources on beetles found in Lithuania have been considered in the current study. Over 630 literature sources on species composition of beetles, their distribution in Lithuania and neighbouring countries, and taxonomic revisions and changes are reviewed and cited. An alphabetical list of these literature sources is presented. After revision of public beetle collections in Lithuania, the authors propose to remove 43 species from the beetle species list of the country on the grounds, that they have been wrongly identified or published by mistake. For reasons of clarity, 19 previously noted but later excluded species are included in the current checklist with comments. Based on faunal data from neighbouring countries, species expected to occur in Lithuania are matnioned. In total 1390 species are attributed to this category and data on their distribution in neighbouring countries is presented. Completion of this study provides evidence that the Lithuanian coleopteran fauna has yet to be completely investigated and it is estimated that approximately 28 % of beetle species remain undiscovered in Lithuania. More than 85% of beetle species expected for Lithuania have been found in the following families: Cerylonidae, Geotrupidae, Haliplidae, Kateridae, Lycidae, Lucanidae, Mycetophagidae, Scarabaeidae and Silphidae. In families with few species such as Alexiidae, Boridae, Byturidae, Dascilidae, Drilidae, Eucinetidae, Lampyridae, Lymexilidae, Megalopodidae, Nemonychidae, Nosodendridae, Noteridae, Orsodacnidae, Pyrochroidae, Pythidae, Psephenidae, Rhysodidae, Sphaeritidae, Sphaeriusidae, Sphindidae, Stenotrahelidae and Trogidae, all possible species have already been discovered. However in some beetle families such as Aderidae, Bothrideridae, Eucnemidae, Laemoploeidae, Mordellidae, Ptiliidae, Scraptidae and Throscidae less than 50% of all possible species are known. At present the beetle species recorded in Lithuania belong to 92 families, with species from 9 other families such as Agyrtidae, Biphylidae, Deradontidae, Mycteridae, Ochodaeidae, Phleophilidae, Phloeostichidae, Prostomidae, Trachypachidae are expected to be found.A bibliography and a index of subfamily and genus levels are provided. The information published in the monograph will serve to further faunistic and distribution research of beetles and will help to avoid confusion in the identificatation of coleopteran fauna of Lithuania.     

New Triassic beetles (Coleoptera) from northern European Russia

Beetle remains from the Triassic Khey-Yaga locality, Nenets National District, Korotaikha Basin, Nyadeita Formation, Olenekian-Anisian are described. Only isolated elytra have been found. Tetracoleus tshalyshevi gen. et sp. nov. and T. minimus sp. nov. are described and assigned to Tricoleidae, and six new species are described in formal taxa. Tetracoleus is close to Permian tricoleids just found in the Newcastle Group of Australia and the Vyatkian Aristovo locality of northern European Russia. The composition of elytra in the locality is closer to that of Lower Triassic localities than other Middle Triassic localities and characterizes the beginning of a recovery after the Permian-Triassic ecological crisis. Longxianocupes Hong in Liu, Liu et Hong, 1985 is shown to be a junior synonym of Sogdelytron Ponomarenko, 1969. The species Palademosyne ovum Ponomarenko, 2004, P. elongatum Ponomarenko, 2004, and P. latum Ponomarenko, 2004 are transferred to the genus Pseudochrysomelites Handlirsch, 1906.     

On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera,Insecta)

The relationships of extant and extinct lineages of Adephaga were analysed formally for the first time. Emphasis is placed on the aquatic and semiaquatic groups and their evolution in the Mesozoic. ?Triadogyrus and ?Mesodineutus belong to Gyrinidae, the sister group of the remaining families. ?Triaplidae are the sister group of the following groups (Haliplidae, Geadephaga, Dytiscoidea incl. ?Liadytidae, ?Parahygrobiidae and ?Coptoclavidae [major part]). The lack of a ventral procoxal joint and a very short prosternal process are plesiomorphies of ?Triaplidae. ?Coptoclavidae and ?Timarchopsinae are paraphyletic. ?Timarchopsis is placed in a geadephagan clade. In contrast to other coptoclavids, its metathorax is close to the condition found in Haliplidae, with a complete transverse ridge and coxae with large plates and free mesal walls. ?Coptoclavidae s.str., i.e. excl. ?Timarchopsis, is a dytiscoid subgroup. The mesal metacoxal walls are fused, the coxal plates are reduced, and the transverse ridge is absent. ?Stygeonectes belongs to this dytiscoid coptoclavid unit and is therefore misplaced in ?Timarchopsinae. ?Liadytidae belongs to a dytiscoid subgroup, which also comprises the extant families Aspidytidae, Amphizoidae, Hygrobiidae and Dytiscidae. ?Parahygrobia is the sister group of Hygrobiidae. The larvae are characterized by a broad gula, the absence of the lacinia, retractile maxillary bases and very long urogomphi set with long setae. ?Liadytiscinae is the sister group of extant Dytiscidae. There is no support for a clade ?Eodromeinae and for Trachypachidae incl. ?Eodromeinae. ?Fortiseode is nested within Carabidae. The exclusion of fossil taxa has no effect on the branching pattern. The evolution of Adephaga in the Mesozoic is discussed. Possible reasons for the extinction of ?Coptoclavidae are the rise of teleost fish and the competition of Gyrinidae and Dytiscidae, which possess efficient defensive glands and larval mandibular sucking channels.     

New species of bark beetles from the Rovno amber (Insecta: Coleoptera: Scolytidae)

Two new species of bark beetles are described from the Late Eocene Rovno amber. Xylechinus mozolevskae sp. nov. (Hylesininae: Tomicini) is the first fossil representative of this genus, differing from recent species in having large triangular scales along the suture. Taphramites rovnoensis sp. nov. (Scolytinae: Dryocoetini) differs from the closest species, T. gnathotrichus from Baltic amber, in the presence of short thin hairs on the elytra.     

基于线粒体基因组序列的鞘翅目肉食亚目水生类群系统发育分析

【目的】明确肉食亚目(Adephaga)水生类群线粒体基因组的基本特征,并基于线粒体基因组序列分析肉食亚目水生类群的系统发育关系。【方法】基于Illumina HiSeq X Ten测序技术测定了圆鞘隐盾豉甲Dineutus mellyi和齿缘龙虱Eretes sticticus的线粒体全基因组序列,对其进行了基因注释,并对其tRNA基因二级结构进行了预测分析。加上已公布的鞘翅目(Coleoptera)肉食亚目水生类群17个种的线粒体基因组序列,对该类群共19个种线粒体的蛋白质编码基因(protein-coding genes, PCGs)开展了比较基因组学分析,包括AT含量、密码子偏好性、选择压力等。基于13个PCGs的氨基酸序列和核苷酸序列,利用最大似然法(ML)和贝叶斯法(BI)分别构建鞘翅目肉食亚目水生类群的系统发育关系,并通过FcLM分析进一步评估伪龙虱科(Noteridae)和瀑甲科(Meruidae)的系统发育位置。【结果】圆鞘隐盾豉甲和齿缘龙虱的线粒体基因组全长分别为16 123 bp(GenBank登录号: MN781126)和16 196 bp(GenBank登录号: MN781132),都包含13个PCGs、22个tRNA基因、2个rRNA基因和1个D-loop区(控制区)。19个肉食亚目水生类群线粒体基因组PCGs的碱基组成都呈现A+T偏好性,在密码子使用上也都偏向于使用富含A+T的密码子;在进化过程中13个PCGs的进化模式相同,都受到纯化选择。基于线粒体基因组13个PCGs的氨基酸序列的肉食亚目水生类群的系统发育关系为(豉甲科Gyrinidae+(沼梭甲科Haliplidae+((壁甲科Aspidytidae+(两栖甲科Amphizoidae+龙虱科Dytiscidae))+(水甲科Hygrobiidae+(瀑甲科Meruidae+伪龙虱科Noteridae)))))。【结论】研究结果表明,豉甲科是肉食亚目水生类群的基部类群,接下来是沼梭甲科和龙虱总科;伪龙虱科和瀑甲科互为姐妹群,并一起作为龙虱总科内部的一个分支;两栖甲科与龙虱科具有更近的亲缘关系。     

New beetles (Insecta, Coleoptera) from Vyazniki locality, terminal Permian of European Russia

Recently collected fossil beetles from the families Asiocoleidae, Rhombocoleidae, Schizocoleidae, and Permosynidae are described from the locality Vyazniki, terminal for the Permian of European Russia. The findings are represented by isolated elytra, the position of which in the natural system it is often impossible to establish; they are, therefore, described in a formal system. The oryctocenosis is dominated by schizocoleids and permosynids, which start to play an important role at the very end of the Permian. They probably include members of Adephaga and Polyphaga, the principal groups of Mesozoic-Cenozoic beetles. Each of the other two families is represented by a single specimen.     

Absence asymmetry: the evolution of monorchid beetles (Insecta: Coleoptera: Carabidae)

Asymmetrical monorchy, or the complete absence of one testis coupled with the presence of its bilateral counterpart, is reported for 174 species of the carabid beetle tribes Abacetini, Harpalini, and Platynini (Insecta: Coleoptera: Carabidae) based on a survey of over 820 species from throughout the family. This condition was not found in examined individuals of any other carabid beetle tribes, or of other adephagan beetle families. One monorchid taxon within Platynini exhibits symmetrical vasa deferentia at the beginning of the pupal stadium, suggesting that developmental arrest of the underdeveloped vas deferens takes place in pupation. The point at which development of the testis is interrupted is unknown. Complete absence of one organ of a bilateral pair--absence asymmetry--is rarely found in any animal clade and among insects is otherwise only known for testes in the minute-sized beetles of the family Ptiliidae, ovaries in Scarabaeinae dung beetles, and ovaries of some aphids. Based on current phylogenetic hypotheses for Carabidae, testis loss has occurred independently at least three times, and up to five origins are possible, given the variation within Abacetini. Clear phylogenetic evidence for multiple independent origins suggests an adaptive or functional cause for this asymmetry. A previously posited taxon-specific hypothesis wherein herbivory in the tribe Harpalini led to testis loss is rejected. Optimal visceral packing of the beetle abdomen is suggested as a general explanation. Specifically, based on the function of various organ systems, we hypothesize that interaction of internal organs and pressure to optimize organ size and space usage in each system led to the multiple origins and maintenance of the monorchid condition. Testes are the only redundant and symmetrically paired structures not thought to be developmentally linked to other symmetrical structures in the abdomen. Among all possible organs, they are the most likely--although the observed frequency is very small--to bypass constraints that maintain bilateral symmetry, resulting in absence asymmetry. However, based solely on our observations of gross morphology of internal organs, no function conclusively explains the ontogenetic loss of one testis in these taxa. Unlike the analogous absence asymmetry of organs in other animal groups, no dramatic body-form constraint--e.g., snakes and lung loss, ptiliid beetles' small body-size and relatively giant sperm--or adaptive scenario of improved locomotory performance--e.g., birds and ovary loss due to flight constraints-applies to these carabid beetles. We tentatively suggest that testis loss is driven wholly by an interaction among the internal organs of these beetles, possibly due to selective pressure to maximize the comparatively large accessory glands found in these taxa. However, as the ordering of these evolutionary events of testis loss and accessory gland size increase is not known, large accessory glands might have secondarily evolved to compensate for a decreased testicular output.     

Diversity of litter-dwelling beetles in the Ouachita Highlands of Arkansas, USA (Insecta: Coleoptera)

A survey of forest litter-inhabiting Coleoptera was conducted in deciduous forests of the Ouachita Mountains in western Arkansas during 1991–1992. A total of 102 Berlese samples were collected, weighed, and processed during the 12-month study. From 741kg of sifted forest litter, we counted and sorted 10663 adult beetles representing 46 families and 400 species. The family Staphylinidae was taxonomically and numerically dominant, comprising 46% of species and 63% of individuals. Problems in assessing species richness of forest litter Coleoptera faunas result from a lack of taxonomic revisions, occurrence of sex-limited diagnostic characters, and inadequate information about larval-adult species associations and life histories. A randomized species accumulation curve indicated that species addition ranged from 14 species per sample during the first ten samples to 1.5 species per sample during the final ten. Richness estimates generated from the empirical data ranged from 434 species (Michaelis–Menten and Coleman richness estimators) to 590 species (second order jacknife). The area sampled is biogeographically significant because it harbors numerous habitat- restricted species that are endemic to the Ouachita Mountains or the Interior Highland region, as well as disjunct populations of species that are also found in the southern Appalachian Mountains. Deciduous forest habitats of the Ouachita Mountains, particularly beech-maple riparian forest, should be given special consideration in forest conservation planning because of their significance as refugia for these and other endemic and disjunct arthropods.     

Histological structure of tripartite mushroom bodies in ground beetles (Insecta, Coleoptera: Carabidae)

Contrary to members of the suborder Polyphaga, ground beetles have been found to possess tripartite mushroom bodies, which are poorly developed in members of basal taxa and maximally elaborated in evolutionarily advanced groups. Nevertheless, they do not reach the developmental stage, which has been previously found in particular families of beetles. It has been pointed out that a new formation of the Kenyon cells occurs during at least the first months of adult life, and inactive neuroblasts are found even in one-year-old beetles. It has been suggested that there is a relation between the Kenyon cell number and development of the centers of Kenyon cell new-formation.     

A genus-level supertree of Adephaga (Coleoptera)

A supertree for Adephaga was reconstructed based on 43 independent source trees – including cladograms based on Hennigian and numerical cladistic analyses of morphological and molecular data – and on a backbone taxonomy. To overcome problems associated with both the size of the group and the comparative paucity of available information, our analysis was made at the genus level (requiring synonymizing taxa at different levels across the trees) and used Safe Taxonomic Reduction to remove especially poorly known species. The final supertree contained 401 genera, making it the most comprehensive phylogenetic estimate yet published for the group. Interrelationships among the families are well resolved. Gyrinidae constitute the basal sister group, Haliplidae appear as the sister taxon of Geadephaga+Dytiscoidea, Noteridae are the sister group of the remaining Dytiscoidea, Amphizoidae and Aspidytidae are sister groups, and Hygrobiidae forms a clade with Dytiscidae. Resolution within the species-rich Dytiscidae is generally high, but some relations remain unclear. Trachypachidae are the sister group of Carabidae (including Rhysodidae), in contrast to a proposed sister-group relationship between Trachypachidae and Dytiscoidea. Carabidae are only monophyletic with the inclusion of a non-monophyletic Rhysodidae, but resolution within this megadiverse group is generally low. Non-monophyly of Rhysodidae is extremely unlikely from a morphological point of view, and this group remains the greatest enigma in adephagan systematics. Despite the insights gained, our findings highlight that a combined and coordinated effort of morphologists and molecular systematists is still required to expand the phylogenetic database to enable a solid and comprehensive reconstruction of adephagan phylogeny. See also Supplementary material in the online edition at doi:10.1016/j.ode.2006.05.003     

First record of the family Hygrobiidae (Coleoptera: Adephaga) from Turkey

With the discovery of Hygrobia hermanni, the family Hygrobiidae is reported for the first time from Turkey. An illustration of the habitus and line drawings of diagnostic characters are provided.     

The morphological evolution of the Adephaga (Coleoptera)

The evolution of the coleopteran suborder Adephaga is discussed based on a robust phylogenetic background. Analyses of morphological characters yield results nearly identical to recent molecular phylogenies, with the highly specialized Gyrinidae placed as sister to the remaining families, which form two large, reciprocally monophyletic subunits, the aquatic Haliplidae + Dytiscoidea (Meruidae, Noteridae, Aspidytidae, Amphizoidae, Hygrobiidae, Dytiscidae) on one hand, and the terrestrial Geadephaga (Trachypachidae + Carabidae) on the other. The ancestral habitat of Adephaga, either terrestrial or aquatic, remains ambiguous. The former option would imply two or three independent invasions of aquatic habitats, with very different structural adaptations in larvae of Gyrinidae, Haliplidae and Dytiscoidea.     

Male achiasmatic meiosis in Caraboidea (Coleoptera,Adephaga)

The meiotic process was studied in fifty-three species belonging to the caraboid families Bembidiidae (48). Trechidae (3), Pogonidae (1), and Harpalidae (1). Males show achiasmatic meiosis patterns of the Callimantis type. In females only early prophase stages were observed, that are similar to those of the males. The characteristics of this abnormal meiosis and its evolutionary and cytotaxonomic significance are discussed.     

Comparative histology of mushroom bodies in carnivorous beetles of the suborder polyphaga (Insecta, Coleoptera)

Mushroom bodies in beetles of the families Histeridae, Staphylinidae, Cantharidae, Trogossitidae, Peltidae, Cleridae, Malachiidae, and Coccinellidae are shown to be rather poorly developed. The calyx region of the mushroom bodies in these beetles never forms two separate cups, and the peduncular apparatus includes a unified shaft almost over its entire length. Only the pedunculus contains two separate shafts in a few cases. Two proliferative centers consisting of one to three neuroblasts are often found in each Kenyon cell group. The shift from carnivorous to feeding on pollen or leaves, which has taken place in some taxa, does not visibly affect the degree of mushroom body development.     

Early Cretaceous origin of pollen‐feeding beetles (Insecta: Coleoptera: Oedemeridae)

The taxonomic position of a new pollen‐feeding fossil beetle from Spanish amber (late Albian, 105 Ma) is analysed. A phylogenetic analysis allows me to accommodate Darwinylus marcosi gen. et sp. nov. in the Polyphaga: Oedemeridae within current limits for the family, which clearly belongs in the subfamily Oedemerinae. It corresponds to the oldest definitive record for the family. Some autapomorphies, mainly in antennae, are observable in the fossil compared with extant members of the family. A discussion about these problematic characters and the evolution of the family is proposed.