Review of paleontological data on the evolution of aquatic beetles (Coleoptera)

Paleontological data on the evolution of aquatic beetles are reviewed. A total of 37 beetle families are regarded as aquatic; five of them are extinct; three other extinct families could have had aquatic larvae; seven families are not known from the fossil record. The earliest records of fossil representatives of Recent families of aquatic beetles are given. The geological distribution of extinct families is reviewed, and the data on the modes of life of their representatives and on their evolution are provided. The new name Coleocatiniidae nom. nov. is proposed for Catiniidae Ponomarenko, 1968, a junior homonym of Catiniidae Embelton, 1901 (Crustacaea, Copepoda).     

Morphological evolutionary trends and their implications for the phylogenetic analysis of metendosternite in Chinese dung beetles (Coleoptera, Scarabaeidae)

后胸叉骨是昆虫胸部的内骨骼,是重要的肌肉联结点,在昆虫运中起着不可替代的作用.该结构在鞘翅目高级阶元系统发育关系重建中扮演了重要角色,但由于其在低级阶元间的差异较小以及在形态分析中使用困难等因素限制,致使后胸叉骨的研究并未受到广泛重视.国内学者对于甲虫后胸叉骨的研究非常少,甚至长期以来没有中文名称,相关内容只零星散布于教科书和学位论文中,针对蜣螂后胸叉骨形态学研究尚未见报道.本文对金龟科蜣螂亚科9族65种的后胸叉骨进行了详细的比较形态学研究,归纳了族级形态特征,进而探究蜣螂后胸叉骨形态演化趋势及其在系统发育分析中的意义.此外,还对利用现代形态学和几何形态学等方法研究后胸叉骨功能形态的前景进行了阐述.     

Molecular phylogeny of Omaliinae (Coleoptera: Staphylinidae) and its implications for evolution of atypically long elytra in rove beetles

Staphylinidae, or rove beetles, are a megadiverse family known for their typically very short elytra exposing most of the abdomen, but the putatively early-derived subfamily Omaliinae and its relatives have been known to include multiple taxa with unusually long elytra. The ancestral elytral length of the family and of this subfamily have long been debated. We present a phylogenetic analysis of Omaliinae based on partial mitochondrial COI (1488 bp), COII (366 bp), 12S rDNA (353 bp), nuclear 18S rDNA (1814 bp), 28S rDNA (876 bp) and CAD (869 bp) data. In all, 51 species in 31 genera and four outgroup species were included. The concatenated sequences were analysed by both parsimony- and model-based (Bayesian and maximum likelihood) methods. The subfamily Omaliinae was not supported as a monophyletic group. The model-based analyses (Bayesian and maximum likelihood trees) showed Empelinae nested within Omaliinae (excluding Corneolabiini), whereas parsimony analysis found all three putative ingroup subfamilies, Empelinae, Glypholomatinae and Microsilphinae, grouped within Omaliinae. Within the Omaliinae, the tribes Coryphiini and Eusphalerini were each supported as monophyletic, whereas Anthophagini and Omaliini were each nonmonophyletic. We hypothesize that there have been at least four independent origins of long elytra from short elytra in the omaliine lineage.     

A new genus of Elateriform beetles (Coleoptera,Polyphaga) from the Middle-Late Jurassic of Karatau

A new coleopteran genus, Anacapitis gen. nov., comprising A. karataviensis sp. nov., A. incertus sp. nov., and A. oblongus sp. nov., is described from the Karabastau Formation of the Karatau locality (southern Kazakhstan, Middle-Late Jurassic) in the infraorder Elateriformia. The systematic position of the new genus within the suborder Polyphaga is discussed.     

Repeated losses of TTAGG telomere repeats in evolution of beetles (Coleoptera)

We studied the occurrence of (TTAGG) _n telomere repeats in 12 species of beetles, representing main lineages of the Coleoptera phylogenetic tree, by Southern hybridization and fluorescence in situ hybridization (FISH). In contrast to other insect orders, beetles were heterogeneous with respect to the occurrence of TTAGG repeats. In addition, the presence or absence of (TTAGG) _n motif was irrespective of phylogenetic relationships. In the suborder Polyphaga, six species displayed positive hybridization signals. These were Silpha obscura, Agrilus viridis, Ampedus sanguineus, Stegobium paniceum, Oryzaephilus surinamensis, and Leptinotarsa decemlineata. Whereas negative signals were obtained in three polyphagan species, Geotrupes stercorarius, Thanasimus formicarius, and Sitophilus granarius. In the suborder Adephaga, the TTAGG sequence was present in one species, Graphoderus cinereus, and absent in two species, Orectochilus villosus and Pterostichus oblongopunctatus. We concluded that the telomerase-dependent (TTAGG) _n motif had been repeatedly lost in different phylogenetic branches of Coleoptera and probably replaced with another mechanism of telomere elongation. This had to happen at least 5–6 times. The results suggest a predisposition or a backup mechanism of telomere maintenance in the genome of beetles that enabled them to make frequent evolutionary changes in the telomere composition.     

The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict

A model of evolution based on conflicts of interest between the sexes over mating decisions is examined in relation to diving beetles (Dytiscidae). The model predicts the following evolutionary sequence: (1) cost to females of mating increases, (2) females evolve behavioural resistance to male mating attempts, (3) males evolve devices to overcome female resistance, and (4) females evolve morphological counter-adaptations to the male devices. This model is tested using species of Dytiscidae, in which (1) some species have a very long mating duration while others mate quickly, (2) females of some species resist male mating attempts by swift and erratic swimming when seized by a male, (3) males of some species possess a grasping device in the form of sucker-shaped setae on the legs used to adhere to the pronota or elytra of females prior to mating, and (4) females of some species have a modified dorsal cuticle with irregular sculpturing which appears to interfere with the male adhesive setae. The predicted order of evolution of some of these features was tested in a cladistic analysis of 52 taxa in Dytiscidae and Hygrobiidae using characters from adult and larval morphology and a portion of the gene wingless . The combined analysis resulted in nine most parsimonious cladograms. The consensus cladogram of these indicates that male sucker setae arose a single time in a clade of Dytiscinae. Nested within this clade are five groups with an independently derived, modified dorsal cuticle in females. This pattern of characters in Dytiscinae is consistent with the prediction implied by the model of sexual selection. The utility of wingless as a marker for phylogenetic analysis of diving beetles is discussed, and the resulting phylogeny is compared with previous analyses and current classification.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 359–388.     

Molecules,morphology and minute hooded beetles: a phylogenetic study with implications for the evolution and classification of Corylophidae (Coleoptera: Cucujoidea)

Eight genes (nuclear: 18S, 28S, H3, CAD; mitochondrial: 12S, 16S, COI, COII) and morphology were used to infer the evolutionary history of Corylophidae, some of the smallest free‐living insects. The study included 36 corylophid exemplars, representing approximately 60% of the known generic diversity of the family and 16 cucujoid outgroup taxa. Multiple partitioning strategies, molecular datasets, combined datasets and different taxon sampling regimes using maximum likelihood and mixed‐model Bayesian inference were utilized to analyse these data. Most results were highly concordant across analyses. There was strong agreement across (i) partitioning strategies, (ii) maximum likelihood and Bayesian inference analyses of the molecular data, and (iii) Bayesian inference of the molecular data alone and Bayesian inference of the combined morphological and molecular data when all terminal taxa were included. When a strict taxon sampling protocol was employed so that only single generic exemplars were included, deep relationships were affected in the resulting phylogenetic hypotheses. Under such narrow sampling strategies, deep phylogenetic relationships were also sensitive to the choice of generic exemplars. Although it is often challenging to obtain single representatives for many taxa in higher‐level phylogenetic analyses, these results indicate the importance of using denser taxon sampling approaches even at the specific level for genera included in such studies. Molecular data alone support Anamorphinae (Endomychidae) strongly as the sister group of Corylophidae. In combined data analyses, Coccinellidae is recovered as the sister group to Corylophidae. In all analyses, Corylophidae and the subfamily Corylophinae are recovered as monophyletic. The monophyly of Periptyctinae was untested, as only a single species was included. All included corylophine tribes were recovered as monophyletic with the exception of Aenigmaticini; Aenigmaticum Matthews forms the sister group to Orthoperus Stephens and Stanus?lipiński et al. is recovered as the sister group of Sericoderus Stephens. Stanus tasmanicus?lipiński et al. is transferred to a new genus, Pseudostanus Robertson, ?lipiński & McHugh gen.n. incertae sedis. We propose a new tribe, Stanini Robertson, ?lipiński & McHugh trib.n. for Stanus bowesteadi?lipiński et al. and a new concept of Aenigmaticini sensu.n. to include only the nominate genus. Anatomical transitions associated with corylophid miniaturization are highlighted. Key phenotypic modifications and elevated rates of substitution in nuclear rRNA genes are evident in a subgroup of Corylophinae that includes the most diminutive species. Other taxonomic and evolutionary implications are discussed in light of the results.     

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.     

A new genus of Elateriform beetles (Coleoptera,Polyphaga) from the Jurassic of Daohugou,China

A new coleopteran genus, Parelateriformius, gen. nov., comprising the species P. communis, sp. nov., P. villosus, sp. nov., P. capitifossus, sp. nov., and P. mirabdominis, sp. nov., is described from Daohugou deposits, Inner Mongolia, China, and is assigned to the infraorder Elateriformia. The systematic position of the new genus within the suborder Polyphaga is discussed.     

New mesozoic Ithyceridae beetles (Coleoptera)

A new subfamily, Mongolocarinae subfam. nov. of the family Ithyceridae, from the Middle-Upper Jurassic and Lower Cretaceous of Asia is described. It includes five genera with five species: Palaeocar gen. nov. (with P. princeps sp. nov.), Mongolocar gen. nov. (M. orcinus sp. nov.), Praecar gen. nov. (P. stolidus sp. nov.), Karacar gen. nov. (M. contractus sp. nov.), and Baissacar gen. nov. (B. passarius sp. nov.).     

New species ofElectribius Crowson 1973 (Coleoptera: Artematopodidae) from Baltic amber

Three new species of the genusElectribius Crowson 1973 (Coleoptera: Artematopodidae),E. friederichi, E. balticus, andE. procerus are described from the Baltic amber. The characters separating the extant and fossil species ofElectribius are summarised in a key. A list of the known extant and fossil species of Artematopodidae sensuLawrence (1995) with synonyms and remarks on the geographical distribution is provided. The distribution of the members of the family and the identical morphological specializations of the fossil and extant species ofElectribius suggest that the taxon formerly had a much wider distribution and is relatively old. The possibility that the Upper Jurassic to Lower Cretaceous generaSinoelaterium Ping 1928and Artematopodites Ponomarenko 1990 represent early members of the Artematopodidae is discussed.     

Habitat selection and grouping of beetles (Coleoptera)

Beetles were collected by pitfall trapping for a two-year period in seven adjacent habitats in an upland site in North Wales. Positive correlations were demonstrated between number of beetle species and number and diversity of plant species. Similar correlations were shown between beetle numbers and plant species. However, only 15% of the beetle species were herbivores requiring host plants. The degree of habitat selection by individual beetle species was demonstrated, ranging from habitat specialist, being found in one habitat, lo habitat generalist, being found in most habitats. Herbivores were significantly more habitat specialist than predators or scavengers. The grouping of beetles, demonstrated by ordination analysis, was similar to, but less precise than, the grouping of plant species. The beetle groups reflect habitat selection preferences by individual species rather than a functional relationship between beetle species. They provide an example of the centrifugal structure of habitat selection theory.     

Ground beetles of the Ukraine (Coleoptera, Carabidae)

A review of the ground beetles of the Ukrainian fauna is given. Almost 750 species from 117 genera of Carabidae are known to occur in the Ukraine. Approximately 450 species of ground beetles are registered in the Carpathian region. No less than 300 species of ground beetles are found in the forest zone. Approximately 400 species of Carabidae present in the forest-steppe zone are relatively similar in species composition to those in the forest territories. Some 450 species of Carabidae are inhabitants of the steppe zone. Representatives of many other regions of heterogeneous biotopes such as forest, semi desert, intrazonal, etc. can be found in the steppe areas. The fauna of Carabidae (ca. 100 species) of the lowlands of southern Ukraine (sandy biotopes), situated mostly in the Kherson region, is very peculiar. The fauna of the Crimean mountains contains about 300 species. Conservation measures for the Carabidae are discussed.     

Biogeography of the phoracanthine beetles (Coleoptera: Cerambycidae)

Six biogeographic subregions of the phoracanthine beetles are proposed on the basis of 136 valid species in all eleven genera of this Australasian group of insects: the Kosciuskan, Western, Eyrean, Torresian, Timorian and New Guinean. Both phenetic and cladistic approaches are employed in the biogeographic study of these beetles. Phenetically, the Kosciuskan and Torresian fauna are the most similar and the least faunal similarity occurs between the New Guinean subregion and all others. Cladistically, two distinct peripheral faunas are divided: the southern Western+Kosciuskan and the northern New Guinean+Torresian+Timorian. The Eyrean fauna in the central Australian continent is more distantly related to the remainder than the latter are to each other. Biogeographic history of the phoracanthine beetles is discussed extensively.     

The evolution of mating systems in bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae)

The extraordinary array of mating systems in the Scolytidae and Platypodidae has been largely overlooked by researchers interested in the evolution of sexual behaviour. This paper provides the first overview of reproductive behaviour in this important and widespread group, known to most biologists only by the reputations of tree-killing taxa. Referred to generally as ‘bark beetles’, these insects chew egg tunnels inside a variety of (usually dead) plant tissues, though most species are either phloeophagous (breeding in the inner bark of woody plants) or xylomycetophagous (all stages feeding on mutualistic fungi growing on sapwood or heartwood). In most species, permanent records of many aspects of reproductive behaviour are etched in the host; in many, engravings reveal female fecundity, eggs sired per male, hatching success, and offspring survivorship. Each gallery arm represents a good portion of a given female's lifetime reproduction, but in many species females commonly re-emerge to reproduce in one or two additional sites. In most species of bark beetles, each female initiates her own gallery, to be joined later by a male. These monogynous gallery systems are associated with mating systems defined by how long males stay with females: in a few species, males seldom if ever join females under the bark; in the vast majority of species, males stay for part or all of the oviposition period then leave to seek other mates; and a few groups exhibit permanent monogamy, in that both sexes die in their only gallery system. While these patterns emerge from an overview of the world scolytid fauna, the length of male residency has seldom been quantified, and the costs and benefits associated with male mating strategies have not been measured for any bark beetle. Male-initiated monogyny is uncommon in Scolytidae, though the rule in Platypodidae; all instances of which I am aware are summarized from a phylogenetic perspective. Inbreeding polygyny with highly biased sex ratios has arisen at least seven times in Scolytidae. These taxa are usually characterized by males being dwarfed, flightless, and uncommon. Sex determination is known for only a few examples, but both haplodiploidy and diplodiploidy have been reported. Multiple origins of harem polygyny (otherwise rare in invertebrates) add an exciting dimension to the comparative and experimental study of scolytid mating systems. In harem polygynous taxa, males initiate gallery construction. I summarize what little can be learned from the literature about the fine structure of harem polygynous mating systems in bark beetles, and the problem of measuring reproductive success. Data on the nature of harem polygyny in Pityophthorus lautus are presented, illustrating (a) the fluidity of harems; (b) that average eggs laid per gallery arm is relatively unaffected by harem size, but strongly influenced by resource quality; (c) that male egg-gain is strongly correlated with territory quality (a consequence of (b) above); and (d) the temporal patterning of immigration and emigration and its effects on gallery system sex ratios. The second half of this paper is a discussion of the evolution of bark and ambrosia beetle mating Mating systems, emphasizing sexual selection and the role of resources. Male, residence is interpreted as postcopulalory guarding—preventing sexual liaisons with wandering males. Operational sex ratio, encounter rate, synchrony of breeding, ejaculate competition, and spatiotemporal distribution of resources are discussed as evolutionary forces moulding scolytid and platypodid male postmating behaviour. The nature of male male competition is reviewed. The paucity of information on male behaviour in gallery systems is mentioned; whether or not males significantly aid females is not known. Three hypotheses are presented for why females re-emerge, a feature which strongly affects operational sex ratios. Finally, I summarize features of bark beetle existence predisposing them to the evolution of post-inseminative guarding. Male-initiated monogyny presents a puzzle. I propose that most uncontested examples can be explained by monogyny re-evolving from (male-initiated) harem polygyny, and I present an argument for the evolution of harem polygyny leading to the development of male gallery initiation. The evolution of harem polygyny in birds and mammals has attracted considerable attention. The Verner Willson Orians polygyny threshold model is discussed with respect to bark beetles in general and P. lautus in particular. Resource quality is a major factor in P. lautus harem dynamics: the cost to females of joining harems is apparently slight compared to benefits accrued from moving into sites with higher quality inner bark. Female-biased adult sex ratios have been suggested to lead to harem polygyny, and literature and original data pertinent to this hypothesis are examined. The geometric constraints model, based on the polygyny threshold concept but tailored to bark beetles, is proposed to account for the failure of most species to evolve harem polygyny, and testable predictions are derived that interrelate breeding systems, habitat quality, and progeny size. The evolution of Inbreeding is briefly covered, and two routes to inbreeding polygyny are suggested.     

Anthophagy in the leaf beetles (Coleoptera,Chrysomelidae)

Laboratory experiments were performed on leaf beetles collected on flowers. Flower consumption (anthophagy) was demonstrated for adults of Donacia bicolora, D. brevitarsis, D. obscura, D. thalassina, Plateumaris discolor, Cryptocephalus laetus, C. sericeus, C. solivagus, Labidostomis longimana, Hydrothassa marginella, Phaedon concinnus, Galerucella nymphaeae, Neocrepidodera femorata, Altica oleracea, Aphthona lutescens, Longitarsus pellucidus, and larvae of Entomoscelis adonidis, H. marginella, and G. nymphaeae for the first time for all the species except for P. discolor, L. longimana, and C. sericeus. The feeding behavior was described, and the mouthparts of adults were studied. The mandibles and maxillae of the Donacia, Plateumaris, and Orsodacne species mentioned are adapted to consuming pollen. Some questions of ecology and evolution of anthophagy are discussed. An overview of literature on anthophagy in the leaf beetles is presented.