A phylogeny of the families of Scarabaeoidea (Coleoptera)

Abstract. A study, based on examination of thirteen scarabaeoid families, was made of 134 adult and larval characters from the following character suites: 105 adult characters of the antennae, eye, epipharynx, mandible, maxillae, labium, tentorium, trochantin, procoxae, mesocoxae, mesothoracic spiracles, hind wing articulation, hind wing base, hind wing venation, hind wing folding, abdominal sternites, abdominal spiracles, male genitalia, ovarioles and karyotype; twenty larval characters of the antennae, fronto-clypeal suture, stemmata, labial palpi, maxillae, mandibles, legs, stridulatory apparatus, spiracles and ecdysial process; and nine adult and larval biological characters. In order to assess the reliability of different characters in resolving scarabaeoid family relationships, six data sets were subjected to cladistic analysis: the total evidence character set (134 characters), restricted adult character set (thirty-two characters, not including those of the wings), wing character set (seventy-three characters), larval character set (twenty characters), biological character set (nine characters) and re-coded Howden (1982) character set (thirty-nine characters). The complete character set and wing character set both produced phylograms with all nodes resolved; the restricted adult data set, larval data set, Howden (1982) data set and biological data set produced phylograms with diminishing levels of node resolution. The reconstructed phylogeny, from the preferred phylogram of the total evidence character set, shows that the Scarabaeoidea comprises three major lineages; a glaresid, passalid and scarabaeid lineage. The glaresid lineage consists only of the Glaresidae. The passalid lineage comprises two major lines; a glaphyrid line (containing Glaphyridae, Passalidae, Lucanidae, Diphyllostomatidae, Trogidae, Bolboceratidae and Pleocomidae) and a geotrupid line (containing Geotrupidae, Ochodaeidae, Ceratocanthidae and Hybosoridae). The scarabaeid lineage contains those taxa traditionally included within the Scarabaeidae (Aphodiinae, Scarabaeinae, Orphninae, Melolonthinae, Acoma, Chasmatopterinae, Hopliinae, Oncerinae, Rutelinae, Dynastinae, Trichiinae, Cetoniinae and Valginae).     

Evolution of the scarab hindwing articulation and wing base: a contribution toward the phylogeny of the Scarabaeidae (Scarabaeoidea: Coleoptera)

A study is made of the articulation and base of hindwings of Scarabaeidae. The survey is based on an examination of over 150 genera from sixteen scarabaeid subfamilies and taxa of uncertain phylogenetic status. Relationships among all subfamilies of Scarabaeidae are examined here for the first time. The constructed phylogeny shows that the scarabaeid lineage (Scarabaeidae) is comprised of two major lines: an aphodiine line (containing Aphodiinae, including Aegialiini, Aulonocnemis and Scarabaeinae), and an orphnine line (containing Orphninae, Melolonthinae, Acoma , Chnaunanthus , Hopliini, Oncerus , Rutelinae, Dynastinae, Trichiinae, Cetoniinae, Osmoderma and Valginae).     

Structure of the mushroom bodies in Scarabaeoidea (Insecta: Coleoptera): 1. Basal families and coprophagous scarabaeidae

Mushroom bodies are in general similarly developed in most taxons studied. The calyx region appears as a single structure, and its dual nature is not yet realized. An anterio-posterior asymmetry of the calyx region with Kenyon cell processes running mostly behind the glomerular neuropil of the calyx is characteristic of all the species studied. In this respect, the calyx region of basal Scarabaeoidea resembles greatly the calyx of many dipterans. Lobe compartmentalization occurs at the initial stage. The passalid beetles represent an exception, as their mushroom bodies are much more developed than in related families. This may be connected with the complicated social behavior of Passalidae.     

The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae)

Abstract. To reconstruct the phylogeny of the Sericini and their systematic position among the scarabaeid beetles, cladistic analyses were performed using 107 morphological characters from the adults and larvae of forty‐nine extant scarabaeid genera. Taxa represent most ‘traditional’ subfamilies of coprophagous and phytophagous Scarabaeidae, with emphasis on the Sericini and other melolonthine lineages. Several poorly studied exoskeletal features have been examined, including the elytral base, posterior wing venation, mouth parts, endosternites, coxal articulation, and genitalia. The results of the analysis strongly support the monophyly of the ‘orphnine group’ + ‘melolonthine group’ including phytophagous scarabs such as Dynastinae, Hopliinae, Melolonthinae, Rutelinae, and Cetoniinae. This clade was identified as the sister group to the ‘dung beetle line’ represented by Aphodius + Copris. The ‘melolonthine group’ is comprised in the strict consensus tree by two major clades and two minor lineages, with the included taxa of Euchirinae, Rutelinae, and Dynastinae nested together in one of the major clades (‘melolonthine group I’). Melolonthini, Cetoniinae, and Rutelinae are strongly supported, whereas Melolonthinae and Pachydemini appear to be paraphyletic. Sericini + Ablaberini were identified to be sister taxa nested within the second major melolonthine clade (‘melolonthine group II’). As this clade is distributed primarily in the southern continents, one could assume that Sericini + Ablaberini are derived from a southern lineage. Plausibly, ancestors of Sericini + Ablaberini and Athlia were separated by a vicariance event, such as the separation of the African plate from the rest of Gondwana, whereas Sericini and Ablaberini probably diversified during the early Tertiary, with dispersal of some basal Sericini to South America.     

Wood ingestion by passalid beetles in the presence of xylose-fermenting gut yeasts

During a survey of insect gut micro-organisms, we consistently isolated Pichia stipitis-like yeasts (Fungi: Ascomycota, Saccharomycetes) from the wood-ingesting beetles, Odontotaenius disjunctus and Verres sternbergianus (Coleoptera: Passalidae). The yeasts were isolated from passalid beetles over a wide area, including the eastern and midwestern USA and Panama. Phylogenetic analyses of the nuclear encoded small and large subunit rRNA gene (rDNA) sequences distinguished a well-supported clade consisting of the passalid yeasts and Pichia stipitis, P. segobiensis, Candida shehatae and C. ergatensis. Members of this clade have the ability to ferment and assimilate xylose or to hydrolyse xylan, major components of the polysaccharide, hemicellulose. Sexual reproduction was present in the passalid isolates but was rare among the gut yeasts of other beetles to which they were compared. Minor genetic and phenotypic variation among some of the passalid yeasts was detected using markers from the internal transcribed spacer region of the rDNA repeat unit, morphology, and in vitro metabolic tests. The consistent association of xylose-fermenting yeasts of almost identical genotypes with passalid beetles across a broad geographical distribution, suggests a significant symbiotic association.     

New fossil evidence of the early diversification of scarabs: Alloioscarabaeus cheni (Coleoptera: Scarabaeoidea) from the Middle Jurassic of Inner Mongolia,China

Abstract Scarabaeoidea are known from the Lower Jurassic and may have originated in the Triassic based on fossil evidence and phylogenetic research. However, the early diversification of Scarabaeoidea remains unclear due to the lack of high‐quality fossil evidence. Here we describe an exceptionally well‐preserved new fossil of Scarabaeoidea, Alloioscarabaeus cheni gen. et sp. nov from the Middle Jurassic Jiulongshan Formation of Inner Mongolia, China. Based on a morphometric analysis using 17 landmarks of the hind wing of Alloioscarabaeus and 10 scarabaeoid families, we found that Alloioscarabaeus cheni gen. et sp. nov clearly does not belong to any of the known scarabaeoid families and, consequently, is a new family, Alloioscarabaeidae fam. nov., was erected. The discovery of Alloioscarabaeus brought further evidence for the early diversification of major scarab lineages which could allow more detail in the palaeobiogeography of the Scarabaeoidea and Northeast of China which might be one of the originating places or an important radiation place during the evolution of Scarabaeoidea. Alloioscarabaeidae were very likely not good diggers and might have fed on decaying organic materials. Based on the evidence we have now, we tend to believe that most families and some subfamilies of Scarabaeoidea were present in the Jurassic period.     

The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals

Extant terrestrial biodiversity arguably is driven by the evolutionary success of angiosperm plants, but the evolutionary mechanisms and timescales of angiosperm-dependent radiations remain poorly understood. The Scarabaeoidea is a diverse lineage of predominantly plant- and dung-feeding beetles. Here, we present a phylogenetic analysis of Scarabaeoidea based on four DNA markers for a taxonomically comprehensive set of specimens and link it to recently described fossil evidence. The phylogeny strongly supports multiple origins of coprophagy, phytophagy and anthophagy. The ingroup-based fossil calibration of the tree widely confirmed a Jurassic origin of the Scarabaeoidea crown group. The crown groups of phytophagous lineages began to radiate first (Pleurostict scarabs: 108 Ma; Glaphyridae between 101 Ma), followed by the later diversification of coprophagous lineages (crown-group age Scarabaeinae: 76 Ma; Aphodiinae: 50 Ma). Pollen feeding arose even later, at maximally 62 Ma in the oldest anthophagous lineage. The clear time lag between the origins of herbivores and coprophages suggests an evolutionary path driven by the angiosperms that first favoured the herbivore fauna (mammals and insects) followed by the secondary radiation of the dung feeders. This finding makes it less likely that extant dung beetle lineages initially fed on dinosaur excrements, as often hypothesized.     

基因序列在金龟总科(Scarabaeoidea)分子系统学研究中的应用

基因序列分析是揭示金龟总科系统发育关系的重要工具。统计了应用于金龟总科中13个科的线粒体和核基因序列,综述了COⅠ、16S rRNA、28S rRNA、18S rRNA等基因序列在金龟总科分子系统学研究的新进展,探讨了不同基因序列在分类鉴定、隐存种发现、系统发育关系重建等方面的作用,对未来研究趋势进行了展望,为进一步阐明金龟总科系统发育机制奠定基础。     

Microorganisms in the gut of beetles: evidence from molecular cloning

We have regularly cultured yeasts from the gut of certain beetles in our ongoing research. In this study cloned PCR products amplified from the gut contents of certain mushroom-feeding and wood-ingesting beetles in four families (Erotylidae, Tenebrionidae, Ciidae, and Passalidae) were sequenced and compared with culture results. Cultural techniques detected some yeasts present in the gut of the beetles, including a Pichia stipitis-like yeast associated with wood-ingesting passalid beetles. Clone sequences similar to several ascomycete yeasts and Malassezia restricta, a fastidious basidiomycetous yeast requiring special growth media, however, were not detected by culturing. Unexpectedly, phylogenetic analysis of additional clone sequences discovered from passalid beetles showed similarity to members of the Parabasalia, protists known from other wood-ingesting insects, termites, and wood roaches. Examination of all gut regions of living passalids, however, failed to reveal parabasalids, and it is possible that they were parasites in the gut tissue present in low numbers.     

The morphology and terminology of the hindwing articulation and wing base of the Coleoptera, with specific reference to the Scarabaeoidea

Abstract. Characters of the hindwing articulation and wing base are important for contributing to the solution of phylogenetic and systematic problems in the Coleoptera. In the Scarabaeoidea morphological terms proposed by previous authors do not cover many structures in sufficient detail and additional terms are needed to describe and utilize all characters used in systematic considerations; these can be used for all Coleoptera.
In this paper we identify new structures, the first basal plate and the second basal plate (two subdivisions of the coleopteran wing base), name the various yokes, braces and reinforcements found on them and propose names for various projections, lobes, indentations and embayments on the axillary sclerites (first axillary, head, neck and tail; second axillary, arm and body; third axillary, prong; basalare).     

Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes

Geometridae is one of the most diverse families within the Lepidoptera, comprising nine subfamilies. Winter moths, which have a unique life history, are found in three subfamilies. To examine the phylogeny of the Geometridae at the subfamily level and determine the evolutionary history of winter moths, we constructed phylogenetic trees for all nine geometrid subfamilies using two mitochondrial and two nuclear gene sequences. Specimens of all subfamilies were sampled from Japan. Simultaneous analyses of the combined data from all genes revealed that the Geometridae comprised two major clades: one with subfamilies Larentiinae and Sterrhinae, and the other with the remaining seven subfamilies. The second clade included the largest subfamily, Ennominae, and the subfamily Archiearinae, which is traditionally considered to be an ancestral lineage of the Geometridae. The Larentiinae+Sterrhinae clade contained one winter moth lineage, and the second major clade consisted of three winter moth lineages, including Alsophilinae, which contains winter moths exclusively. Using a Bayesian inference of divergence times, we estimated that geometrids began to diverge 54 Mya (62-48 Mya), whereas winter moth lineages differentiated from non-winter moth lineages 34-12 Mya, during the global cooling events in the Oligocene and the early Miocene. The adaptation to cool climates may have been a preadaptation that facilitated the winter moth life cycle.     

Seasonal and annual dietary changes in Lesser Kestrels Falco naumanni wintering in Lesotho

The diet of Lesser Kestrels Falco naumanni wintering in an agriculturally-transformed area in Lesotho was studied by means of pellet analysis. Orthopterans, beetles and solifugids (sun spiders) comprised the staple food of the wintering Lesser Kestrels. Small vertebrates, termite alates, earwigs and scolopendrans supplemented the diet. The proportion of pellets containing scarabaeid and carabid beetles, as well as those containing locusts and crickets, increased as the wintering season progressed, while the proportion of pellets containing solifugids decreased during the same period. Significant differences in diet composition were recorded between the 1998/1999 and 1999/2000 wintering seasons.     

Habitat Specificity and Variation of Coleopteran Assemblages Between Habitats in a Southern African (Swaziland) Agricultural Landscape

Assessment of Coleopteran diversity and abundance was carried out in the lowveld region of Swaziland by pitfall trapping in five distinct habitats. Additionally, the study was to ascertain if any of the families collected illustrated habitat fidelity. 18 coleopteran families, comprising 2903 individuals were collected, with an additional 29 unidentified specimens, totalling 2932 beetles. Numerically, the top five families were the Scarabaeidae (2425), Carabidae (211), Tenebrionidae (149), Nitidulidae (37) and Curculionidae (22). 66.6% of all specimens collected were harboured within the pine windbreak while the lowest number (2.6%) was collected from the mature citrus orchard. The highest proportion (77%) of scarabaeid beetles collected occurred within the pine windbreak. Habitat specificity was illustrated by one family, Meloidae, which was found only in the young orchard. Community analysis was further carried out to ascertain distribution patterns of the dominant coleopteran families. Three scarabaeid species were found only in the pine windbreak while three carabid species occurred only in the indigenous savanna. Significant differences between habitats were observed in the number of beetles collected while no significant differences were observed in terms of the number of families observed in each habitat (p<0.05). Results indicate that conservation of the various habitat patches within the mosaic studied could facilitate conservation of whole communities rather than individual species thus facilitating effective conservation of the agricultural landscape.     

A comprehensive molecular phylogeny of the starlings (Aves: Sturnidae) and mockingbirds (Aves: Mimidae): congruent mtDNA and nuclear trees for a cosmopolitan avian radiation

We generated a comprehensive phylogeny for the avian families Sturnidae (starlings, mynas, Rhabdornis, oxpeckers, and allies) and Mimidae (mockingbirds, thrashers, and allies) to explore patterns of morphological and behavioral diversification. Reconstructions were based on mitochondrial DNA sequences from five coding genes (4108 bp), and nuclear intron sequences from four loci (2974 bp), for most taxa, supplemented with NDII gene sequences (1041 bp) derived from museum skin specimens from additional taxa; together the 117 sampled taxa comprise 78% of the 151 species in these families and include representatives of all currently or recently recognized genera. Phylogenetic analyses consistently identified nine major clades. The basal lineage is comprised of the two Buphagus oxpeckers, which are presently confined to Africa where they are obligately associated with large mammals. Some species in nearly all of the other major clades also feed on or around large vertebrates, and this association may be an ancestral trait that fostered the world-wide dispersal of this group. The remaining taxa divide into sister clades representing the New-World Mimidae and Old-World Sturnidae. The Mimidae are divided into two subclades, a group of Central American and West Indian catbirds and thrashers, and a pan-American clade of mockingbirds and thrashers. The Sturnidae are subdivided into six clades. The Phillipine endemic Rhabdornis are the sister lineage to a larger and substantially more recent radiation of South Asian and Pacific island starlings and mynas. A clade of largely migratory or nomadic Eurasian starlings (within which the basal lineage is the model taxon Sturnus vulgaris) is allied to three groups of largely African species. These reconstructions confirm that Buphagus should not be included in the Sturnidae, and identify many genera that are not monophyletic. They also highlight the substantial diversity among the major Sturnidae subclades in rates of species accumulation, morphological differentiation, and behavioral variation.     

Phylogenetic analysis of Staphyliniformia (Coleoptera) based on characters of larvae and adults

Abstract. One hundred and twenty-one morphological characters of larvae and adults of the series Staphyliniformia were scored (multistate coding) and analysed to determine the family group relationships of the polyphagan groups Scarabaeoidea, Histeroidea, Hydrophiloidea and Staphylinoidea. Cladograms were rooted with exemplars of Adephaga, Archostemata, Myxophaga and the polyphagan families Dascillidae, Derodontidae, Eucinetidae and Scirtidae. Analyses of the same dataset with multistate characters re-coded as presence/absence (144 characters) produced cladograms that were similar to those produced from analyses of the original characters. Cladograms produced from partitioned larval and adult characters differed strongly, with adult-only trees more similar to those produced by combined data. The results confirm the monophyly of Hydrophiloidea + Histeroidea and of Staphylinoidea (including Hydraenidae). The Epimetopidae + Georissidae are the only strongly supported clade within Hydrophiloidea. A clade comprising Hydrochidae, Spercheidae and Hydrophilidae, and a sister-group relationship between the latter two families were confirmed in analyses of the data with presence/absence coding. Helophoridae, Epimetopidae and Georissidae are probably not a monophyletic unit, and additional evidence is needed for a reliable placement of Helophoridae. Scarabaeoidea are placed as a sister taxon of Hydrophiloidea + Histeroidea, but support for this relationship is weak. The branching pattern ((Hydraenidae + Ptiliidae) + (Leiodidae + Agyrtidae)), and a clade comprising Scydmaenidae, Silphidae and Staphylinidae (= ‘staphylinid group’) are well founded. The branching pattern (Orchymontiinae + (Prosthetopinae + (Ochthebiinae + Hydraeninae))) within Hydraenidae is confirmed. Poor resolution at the base of the trees and the placement of some nonstaphyliniform taxa (Dascillidae, Derodontidae, Scirtidae and Eucinetidae) as a sister group to a clade comprising Scarabaeoidea, Hydrophiloidea and Histeroidea suggests that Staphyliniformia may be paraphyletic. It is recommended that series names are eliminated from the classification of Polyphaga, at least for the more ‘primitive’ groups.     

Beetle evolution in the Asian highlands: insight from a phylogeny of the scarabaeid subgenus Serica (Coleoptera, Scarabaeidae)

Abstract.  The phylogeny of the species of the scarabaeid subgenus Serica (genus Serica MacLeay, 1819) was inferred from a parsimony analysis and the use of 117 morphological characters of adults. Successive weighting was employed to further evaluate the phylogenetic relationships, and the geographical changes in the distribution in certain lineages were explored. Four major lineages may be recognized from the strict consensus tree within the ingroup: (1) Calloserica ; (2) Gastroserica + Neoserica ; (3) Lasioserica ; and (4) Serica including the representatives of Taiwanoserica . Pachyserica proved to be monophyletic only after successive weighting based on the retention index. The taxa of Taiwanoserica are nested within the species of Serica . Consequently, Taiwanoserica ( stat.n. ) is ranked as a subgenus of Serica . The clade Serica (subgenus Serica ) comprised the majority of the species included in this study. Serica ( sensu stricto ) species of the Himalayas are a polyphyletic group. Several major younger clades of Serica ( sensu stricto ) are not restricted to just one geographical region, but cover widely distant ranges (such as the Himalaya and eastern Tibet). Dispersal is hypothesized multiple times from the Himalaya to eastern Tibet/Indochina and vice versa, and also between the Japanese islands and eastern Tibet. Several hypotheses are discussed for an important role of the entire Himalaya–south-eastern Tibet orogenic system for the evolution of the subgenus Serica . More recently, the progressive uplift of the Tibetan Plateau provided an efficient barrier to the westward dispersal of eastern Palaearctic species.     

数学形态学在昆虫分类学上的应用研究.Ⅱ.在总科阶元上的应用研究

从总科角度对鳞翅目（Lepidoptera）和鞘翅目（Coleoptera）5个总科23种昆虫图像中提取的昆虫面积、周长等11项数学形态特征进行了统计分析。结果表明,在总科阶元上,11项特征的可靠性大小为圆形性＞（面积、周长、横轴长、球形性、似圆度、偏心率）＞（纵轴长、叶状性）＞（形状参数、亮斑数）。从数学形态特征角度讲,夜蛾总科等3个总科的亲缘关系远近大小依次为夜蛾总科与凤蝶总科>蚕蛾总科与凤蝶总科>夜蛾总科与蚕蛾总科。     

Hystrignathus dearmasi sp. n. (Oxyurida, Hystrignathidae), first record of a nematode parasitizing a Panamanian Passalidae (Insecta, Coleoptera)

Hystrignathus dearmasi sp. n. (Oxyurida: Hystrignathidae) is described from an unidentified passalid beetle (Coleoptera: Passalidae) from Panama. It resembles Hystrignathus cobbi Travassos & Kloss, 1957 from Brazil, by having a similar form of the cephalic end, extension of cervical spines and absence of lateral alae. It differs from the latter species by having the body shorter, the oesophagus and tail comparatively larger, the vulva situated more posterior and the eggs ridged. This species constitutes the first record of a nematode parasitizing a Panamanian passalid.     

大针茅草原蛴螬群落特征研究

内蒙古典型草原的主体植被类型——大针茅(Stipa grandis)草原中,有蛴螬4科9种．构成大针茅草原蛴螬群落的4个科中,以鳃金龟科种类数、个体数为最多,主要种群的重要值排序为黑皱鳃金龟、东方绢金龟、马铃薯鳃金龟及弓斑常丽金龟．几种蛴螬在发生上大致呈3种类型：春季和秋季大量发生．如东方绢金龟;一年中密度波动较小,如弓斑常丽金龟;秋季大量发生,如黑皱鳃金龟和马铃薯鳃金龟．大针茅草原蛴螬群落的多样性特征在于秋季群落多样性和物种的丰度较高,而夏季相对较低．春季蛴螬群落种群丰度较高,但群落结构相对较单调．     

A new podapolipid species (Acari) on Scarabaeus (Scarabaeus) acuticollis (Insecta: Coleoptera: Scarabaeidae) from Iran

A new species of mites of the genus Tarsopolipus (Acari: Prostigmata: Podapolipidae) is described from southern Iran. Tarsopolipus husbandi Mortazavi and Hajiqanbar n. sp. is a sub-elytral ectoparasite of the scarabaeid beetle, Scarabaeus (Scarabaeus) acuticollis. The new species is closely related to Tarsopolipus corrugatus Berlese 1911, but is distinguished from it by the following characters. Adult female: presence of vestigial setae v(2) and shorter setae sc(2). Adult male: presence of setae v' on tibia IV and shorter setae sc(2) and c(2). Larval female: shorter cheliceral stylets and longer distance between setae v(1) and ch. Species of Tarsopolipus are currently distributed in Afro-tropical, Palaearctic, and Oriental regions and parasitize species in 3 genera of scarabaeid beetles, i.e., Scarabaeus , Kheper, and Drepanopodes, all belonging to the tribe Scarabaeini.