中生代鞘翅目扁甲总科化石的研究进展

扁甲总科隶属于鞘翅目多食亚目扁甲系,目前包含25个现生科和3个化石科,其中10个科仅分布于南半球,特别是澳大利亚、新西兰和南美。当前的分子系统发育学研究显示扁甲总科的起源时间为晚三叠世至早侏罗世,而冈瓦纳古陆与劳亚古陆的分离也发生在早侏罗世约1.8亿年前。因此对中生代扁甲总科化石的研究是解释扁甲总科如今分布格局的重要基础,也有助于研究扁甲总科的演化历史。本文简要回顾了现生扁甲总科的研究历史,并对其中生代的化石研究进行了概述,迄今在中生代地层发现有12科54种扁甲总科化石。     

Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinelloidea (Coleoptera: Cucujiformia)

A large‐scale phylogenetic study is presented for Cucujoidea (Coleoptera), a diverse superfamily of beetles that historically has been taxonomically difficult. This study is the most comprehensive analysis of cucujoid taxa to date, with DNA sequence data sampled from eight genes (four nuclear, four mitochondrial) for 384 coleopteran taxa, including exemplars of 35 (of 37) families and 289 genera of Cucujoidea. Maximum‐likelihood analyses of these data present many significant relationships, some proposed previously and some novel. Tenebrionoidea and Lymexyloidea are recovered together and Cleroidea forms the sister group to this clade. Chrysomeloidea and Curculionoidea are recovered as sister taxa and this clade (Phytophaga) forms the sister group to the core Cucujoidea (Cucujoidea s.n .). The nitidulid series is recovered as the earliest‐diverging core cucujoid lineage, although the earliest divergences among core Cucujoidea are only weakly supported. The cerylonid series (CS) is recovered as monophyletic and is supported as a major Cucujiform clade, sister group to the remaining superfamilies of Cucujiformia. Currently recognized taxa that were not recovered as monophyletic include Cucujoidea, Endomychidae, Cerylonidae and Bothrideridae. Biphyllidae and Byturidae were recovered in Cleroidea. The remaining Cucujoidea were recovered in two disparate major clades: one comprising the nitidulid series + erotylid series + Boganiidae and Hobartiidae + cucujid series, and the other comprising the cerylonid series. Propalticidae are recovered within Laemophloeidae. The cerylonid series includes two major clades, the bothriderid group and the coccinellid group. Akalyptoischiidae are recovered as a separate clade from Latridiidae. Eupsilobiinae are recovered as the sister taxon to Coccinellidae. In light of these findings, many formal changes to cucujiform beetle classification are proposed. Biphyllidae and Byturidae are transferred to Cleroidea. The cerylonid series is formally recognized as a new superfamily, Coccinelloidea stat.n. Current subfamilies elevated (or re‐elevated) to family status include: Murmidiidae stat.n. , Teredidae stat.n. , Euxestidae stat.n. , Anamorphidae stat.rev. , Eupsilobiidae stat.n. , and Mycetaeidae stat.n. The following taxa are redefined and characterized: Cleroidea s.n. , Cucujoidea s.n. , Cerylonidae s.n. , Bothrideridae s.n. , Endomychidae s.n. A new subfamily, Cyclotominae stat.n. , is described. Stenotarsinae syn.n. is formally subsumed within a new concept of Endomychinae s.n.     

Nitidulidae (Coleoptera: Cucujoidea) of Vanuatu

A checklist to the Nitidulidae of Vanuatu is provided based on a combination of historical museum specimens and recent field expeditions. A dichotomous key to the described species recorded from the country is provided. Specific data per island is given where available; in addition, broader distributions are outlined for each described taxon. Several undescribed taxa are listed but not described until further work can be done in the region. A discussion of the movement of invasive Nitidulidae throughout Vanuatu is also included with an emphasis on the relationship between increases in human movement and commerce and the number of invasive species present. This discussion is extended to other islands in Vanuatu that have not yet been sampled in an attempt to predict the presence of invasive species.     

Phylogeny and classification of Corylophidae (Coleoptera: Cucujoidea) with descriptions of new genera and larvae

Abstract Phylogenetic relationships within the family Corylophidae were investigated. Twenty ingroup taxa and six outgroups were included in a cladistic analysis, based on 48 characters derived from adult and larval morphology. Phylogenetic analysis confirms that Corylophidae are monophyletic within the superfamily Cucujoidea and may be subdivided into two subfamilies: the Australian Periptycinae and the cosmopolitan Corylophinae containing 10 tribes: Foadiini trib.n. , Cleidostethini, Aenigmaticini, Parmulini, Sericoderini, Peltinodini, Orthoperini, Corylophini, Teplinini and Rypobiini. All currently recognized family‐group taxa are thoroughly diagnosed, and keys to their identification based on adults and larvae are provided. Two new genera and three species are described: Weirus gen.n ., containing only W. tozer sp . n . (Australia: Queensland), and Stanus gen.n. , with the two species S. bowesteadi sp.n . (New Zealand) and S. tasmanicus sp.n. (Tasmania). The larvae of Pakalukodes bimaculatus?lipiński et al. from Queensland and of Stanus bowesteadi sp.n. from New Zealand are described and illustrated for the first time.     

New record of subfamily Prometopinae (Coleoptera: Cucujoidea: Nitidulidae) in Korea

The subfamily Prometopinae Böving and Craighead, 1931 is recorded from Korea for the first time. Two species from different genera are recognized; Prometopia unidentata Hisamatsu, 1959 and Paramatopia x-rubrum Reitter, 1884. Morphological comments, biology, illustrations of habitus with genitalia of both sexes and a key to species from Korea are provided.     

Phylogeny and placement of Boganiidae (Coleoptera,Cucujoidea) with a review of Australian and New Caledonian taxa

A phylogeny of the cucujoid family Boganiidae (Coleoptera) is inferred for the first time based on a parsimony analysis of 102 morphological characters (70 adult and 32 larval). The analysis resulted in a monophyletic Boganiidae divided into two main clades, Boganiinae and Paracucujinae, each supported by a series of synapomorphies. The Boganiinae genera recovered were Afroboganium Endrödy‐Younga & Crowson and Boganium Sen Gupta & Crowson, whereas Paracucujinae includes Paracucujus Sen Gupta & Crowson, Metacucujus Endrödy‐Younga & Crowson, Athertonium Crowson and the New Caledonian Dzumacium caledonicum gen.n. , sp.n. New specimen data and biological information for the Australian taxa are summarized with overviews on biogeography and comments on the fossil Jurassic Parandrexis Martynov (Parandrexidae). Two new species of Boganium, primarily from the Australian mallee are described: B. malleense sp.n. and B. medioflavum sp.n . Boganium malleense is recorded from flowers of Eucalyptus gracilis F. (Myrtaceae). Adults and larvae of Paracucujus rostratus Sen Gupta & Crowson are redescribed. Athertonium parvum Crowson is redescribed and Athertonium williamsi sp.n. is described from coastal New South Wales. Several host records for Athertonium are also presented.     

First Record of the Family Passandridae in Korea (Coleoptera: Cucujoidea)

A taxonomic study of the Korean Passandridae is presented. The family, genus and species (Ancistria apicalis Reitter) are reported for the first time in Korea. Comparison with a lectotype was made. An illustration of the habitus and line drawings of diagnostic characters are provided.     

Review of the genus Myrabolia Reitter (Coleoptera: Cucujoidea: Myraboliidae)

Abstract  Species of the Australian genus Myrabolia are reviewed. Lectotypes are designated for Silvanus brevicornis Erichson, Myrabolia haroldiana Reitter, M. lindensis Blackburn and M. longicornis Blackburn. The following new species are described and illustrated: M. australis, M. blackburni , M. elongata , M. kioloa , M. lawrencei , M. micra , M. pelion and M. tasmanica. Ocholissa leai Grouvelle, originally described in Colydiidae, is here transferred to Myrabolia as M. leai (Grouvelle) comb. n. The nomenclatural history, a diagnosis and distribution data are provided for each species. A key to the species of the genus is presented.     

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.     

Searching for natural lineages within the Cerylonid Series (Coleoptera: Cucujoidea)

Phylogenetic relationships within the diverse beetle superfamily Cucujoidea are poorly known. The Cerylonid Series (C.S.) is the largest of all proposed superfamilial cucujoid groups, comprising eight families and representing most of the known cucujoid species diversity. The monophyly of the C.S., however, has never been formally tested and the higher-level relationships among and within the constituent families remain equivocal. Here we present a phylogenetic study based on 18S and 28S rDNA for 16 outgroup taxa and 61 C.S. ingroup taxa, representing seven of the eight C.S. families and 20 of 39 subfamilies. We test the monophyly of the C.S., investigate the relationships among the C.S. families, and test the monophyly of the constituent families and subfamilies. Phylogenetic reconstruction of the combined data was achieved via standard static alignment parsimony analyses, Direct Optimization using parsimony, and partitioned Bayesian analysis. All three analyses support the paraphyly of Cucujoidea with respect to Tenebrionoidea and confirm the monophyly of the C.S. The C.S. families Bothrideridae, Cerylonidae, Discolomatidae, Coccinellidae and Corylophidae are supported as monophyletic in all analyses. Only the Bayesian analysis recovers a monophyletic Latridiidae. Endomychidae is recovered as polyphyletic in all analyses. Of the 14 subfamilies with multiple terminals in this study, 11 were supported as monophyletic. The corylophid subfamily Corylophinae and the coccinellid subfamilies Chilocorinae and Scymninae are recovered as paraphyletic. A sister grouping of Anamorphinae+Corylophidae is supported in all analyses. Other taxonomic implications are discussed in light of our results.     

A molecular phylogenetic study of Cucujidae s.l. (Coleoptera: Cucujoidea)

Of all the superfamilies within the megadiverse order Coleoptera (Insecta), Cucujoidea (Cucujiformia) is arguably the most problematic taxonomically. The families comprising Cucujidae s.l. (Silvanidae, Laemophloeidae, Passandridae and Cucujidae s.s. represent a large portion of cucujoid diversity. Herein we present the results of a rigorous molecular phylogenetic analysis of Cucujidae s.l. using maximum‐likelihood and Bayesian analyses of seven genes. Representatives of over half of the families of Cucujoidea (excluding the cerylonid series), as well as a broad sampling of Silvanidae and Laemophloeidae, were analysed. The monophyly of Cucujidae s.l. is rejected but a subgrouping of taxa that may form the core of a natural cucujoid lineage is recovered. This clade consists of two large monophyletic groups including several families each. Relationships among these smaller cucujoid groups are discussed, including several novel phylogenetic hypotheses, whereas morphological characters considered significant for classification in Cucujidae s.l. are evaluated in light of these phylogenetic hypotheses. Silvaninae, Telephanini, Brontini and Brontinae are recovered as monophyletic in the Bayesian analysis, but the former two are recovered as paraphyletic in the maximum‐likelihood analysis. Our results support the placement of Psammoecus Latreille within Telephanini and also recover a paraphyletic Telephanus Erichson. Silvaninae is divided into three lineages, each representing a potential tribal lineage. Laemophloeidae is rendered paraphyletic in all analyses by Propalticidae and the latter is herein formally transferred to Laemophloeidae stat.n . Several suprageneric laemophloeid clades are recovered and discussed as potential higher‐level groups. Laemophloeus Dejean is not recovered as monophyletic.     

Molecular phylogeny of Nitidulidae: assessment of subfamilial and tribal classification and formalization of the family Cybocephalidae (Coleoptera: Cucujoidea)

We present a molecular phylogeny of Nitidulidae based on thirty ingroup taxa representing eight of the ten currently recognized subfamilies. Approximately 10 K base pairs from seven loci (12S, 16S, 18S, 28S, COI, COII and H3) were used for the phylogenetic reconstruction. The phylogeny supports the following main conclusions: (i) Cybocephalidae are formally recognized as a distinct family not closely related to Nitidulidae and its constituent taxa are defined; (ii) Kateretidae are sister to Nitidulidae; (iii) Cryptarchinae are monophyletic and sister to the remaining nitidulid subfamilies; (iv) subfamily Prometopinae stat. res. is reinstated and defined, to accommodate taxa allied to Axyra Erichson, Prometopia Erichson and Megauchenia MacLeay; (v) Amphicrossinae, Carpophilinae and Epuraeinae are shown to be closely related taxa within a well‐supported monophyletic clade; (vi) tribal affinities and respective monophyly within Nitidulinae are poorly resolved by our data and must be more rigorously tested as there was little or no support for prior morphologically based tribes or genus‐level complexes; (vii) Nitidulinae are found to be paraphyletic with respect to Cillaeinae and Meligethinae, suggesting that they should either be subsumed as tribes, or Nitidulinae should be divided into several subfamilies to preserve the status of Cillaeinae and Meligethinae; (viii) Teichostethus Sharp stat. res. is not a synonym of Hebascus Erichson and the former is reinstated as a valid genus. These conclusions and emendations are discussed in detail and presented within a morphological framework.     

Larval morphology and phylogenetic position of Micromalthus debilis LeConte (Coleoptera: Micromalthidae)

Abstract External and internal structures of the cerambycoid and triungulin larvae of Micromalthus debilis are described and compared to features found in larvae of other groups of Coleoptera. The morphological data are evaluated with respect to the systematic position of Micromalthidae. A cladistic analysis was carried out with fifty characters. Micromalthidae are not closely related to Lymexylidae (Polyphaga: Cucujiformia) but belong to Archostemata, which is confirmed as a monophyletic unit. Micromalthidae are specialized in terms of morphology and life cycle and are characterized by a considerable number of larval autapomorphies. Their sister-group relationship with Cupedidae is supported by several apomorphic features, which are probably correlated with xylobiontic habits: head transverse and strongly rounded laterally, absence of stemmata, shortened antennae, presence of sternal asperities and presence of eversible lobes of segment IX. Cupedidae is monophyletic and Priamca is the sister group of the remaining genera of Cupedidae included in the analysis. A closer relationship between Tenomerga and Rhipsideigma is supported by several larval synapomorphies. The ancestral life style of larvae of Archostemata was probably xylobiontic. This is suggested by derived groundplan features of the suborder, which are also found in larvae of non-related, wood-associated families.     

The oldest Cenozoic passandrid (Coleoptera: Cucujoidea: Passandridae) from the Paleocene of Menat (France)

The new oldest representative of the subfamily Passandrinae (Passandridae) was found in materials from Menat (Puy-de-Dôme, France). It was preliminarily assigned to the genus Passandra Dalman in Schönherr, 1817, although the new species (?) Passandra plenaria sp. n., in contrast to other congeners, demonstrates an exposed labrum and an absence of preapical groove or impression on the hypopygidum. Some notes on fossil records and classification of the family Passandridae are made. The distinctiveness of the families Passandridae and Laemophloeidae in light of the new data on fossils is considered, and the attribution of Mesopassandra Jin, Ślipiński, Zhou and Pang, 2019 (type genus of Mesopassandrinae) to the family Passandridae is regarded as problematic.     

Generic revision and phylogeny of Microweiseinae (Coleoptera: Coccinellidae)

The monophyly and phylogenetic relationships of the subfamily Microweiseinae were investigated. Twenty‐three in‐group taxa, representing all known genera of Microweiseinae (except for Microcapillata Gordon) were included in a cladistic analysis, based on 45 adult morphological characters. The parsimony analysis of the resulting data matrix supported the monophyly of Microweiseinae, Carinodulini, Serangiini and Microweiseini (inclusive of Sukunahikonini). The recognition of Sukunahikonini renders Microweiseini paraphyletic, and consequently both tribes were synonymized, retaining Microweiseini as a senior family‐group name ( syn.n. ). Genera and tribes of Microweiseinae are diagnosed thoroughly, illustrated and keys to their identification are provided. The following nomenclatural changes were made: Hikonasukuna Sasaji and Orculus Sicard are synonymized with Scymnomorphus Weise ( syn.n. ); Gnathoweisea Gordon is synonymized with Microweisea Cockerell ( syn.n. ); Hikonasukuna monticola Sasaji and Orculus castaneus Sicard are transferred to Scymnomorphus ( comb.n. ); Smilia planiceps Casey, Gnathoweisea hageni Gordon, Gnathoweisea ferox Gordon, Gnathoweisea micula Gordon and Gnathoweisea texana Gordon are transferred to Microweisea ( comb.n ). Three new genera are described: Allenius gen.n. for Allenius californianus sp.n. (type species) from Mexico and Allenius iviei sp.n. from U.S.A.; Gordoneus gen.n. (type species Gnathoweisea schwarzi Gordon from U.S.A.); and Cathedrana gen.n. (type species Cathedrana natalensis sp.n. from South Africa). The first African member of Carinodulini, Carinodulina ruwenzorii sp.n. is described. The genera Microweisea, Coccidophilus, Serangium and Delphastus are well‐known predators of sternorrynchous Hemiptera, such as scale insects (Diaspididae) and whiteflies (Aleyrodidae), and play a significant role in agricultural ecosystems as biocontrol agents. Host data and biological records are summarized for each genus.