Phylogeny of the Saprininae reveals interesting ecological shifts in the history of the subfamily (Coleoptera: Histeridae)

A morphological data set for the histerid beetle subfamily Saprininae comprising 95 adult morphological characters scored (multistate coding) from 72 terminal taxa and four outgroups was developed in order to analyse and determine the relationships amongst the genera and subgenera of the Saprininae subfamily. Cladograms were rooted with exemplars of Dendrophilinae (genus Dendrophilus), Bacaniini (genus Bacanius), Abraeinae (genus Chaetabraeus), and Anapleini (genus Anapleus). Parsimony‐based phylogenetic analyses were performed based on the type species of each genus and subgenus of the Saprininae occurring around the world, with the exception of three taxa: Paramyrmetes foveipennis (type species of the genus Paramyrmetes), Satrapister nitens (type species of the genus Satrapister) and Xerosaprinus (Auchmosaprinus) laciniatus (type species of the subgenus Auchmosaprinus) that were not available. In addition, in order to test the monophyly of several questionable genera, multiple exemplars were added in a few cases. The analysis also included an exemplar of an apparently undescribed genus. The results of the analysis confirm the monophyly of the subfamily supported by two unique synapomorphies: (1) presence of sensory structures of the antenna; and (2) presence of the antennal cavity, as well as several other weaker synapomorphies. However, the phylogeny inferred here shows mostly low support for the deeper branches and consequently no major changes in the Saprininae classification are proposed. The presented cladogram is discussed together with its implications for the evolution of the subfamily. The most informative characters and their respective states are outlined. Multiple shifts in lifestyles have evolved during the evolutionary history of the group. Taxa found near the root of the cladogram are mostly nidicolous or myrmecophilous, and inquiliny is presumed to be the plesiomorphic lifestyle of the subfamily. The nidicolous lifestyle has undergone several transformations to other lifestyles and myrmecophily has evolved three times independently during the evolution of the subfamily. Termitoxeny has evolved two times independently in the group whereas ecological adaptation for life in caves has likewise evolved two times independently. The analyses yielded a large clade of predominantly psammophilous taxa; psammophily is thought to have evolved once and has been subsequently lost several times. © 2014 The Linnean Society of London     

东灵山区阎甲物种多样性研究

本文研究了北京东灵山区阎甲科（Coleptera;Histeridae）特种多样性,野外设样地调查共获标本2224头,分属于6属17种,Saprinus niponicus、Onthophilus foueipennis、Margarinotus niponicus和M.reichardti的个体数量均大于总数的10%, 4种共占总体个数的88.22%,是本地区常见种。本研究采用4种采集方法分别统计,肉诱法捕获的种类和数量最多,共12种占总数的70.19%,杯诱法和堆草诱法都捕到8种,分别占总数的23.61%和4.81%,肉筛法采到的标本仅有2种占总数的1.39%,研究优势物种Onthophilus foueipennis的季节性变化发现,该种集中分布在小龙门林区的阔叶混交林内,个体数量在5月和9月呈现2个季节峰值,其中9月达到全年最高峰。对小龙门林区内肉诱捕获的阎甲类群的物种多样性分析表明,阔叶林中的阎甲多样性指数和均匀度指数较高,混交林的丰富度和个体数量较高,针叶林的多样性系数和个体数量较低,混交林的均匀度指数较低,而相似性分析表明,空间距离是决定小龙门林区内森林生境类型间腐生性阎甲群落相似性程度的因素,生境类型自身对相似性没有显著影响,梨园岭退耕区阎甲的物种多样性指数和均匀度指数高于小龙门林区,但物种个体平均数量低于小龙门林区。以上结果表明,东灵山地区阎甲群落的分布受生境保护程度,干扰程度以及空间距离等因素的影响,保护并提高该地区的森林植被环境对提高阎甲物种多样性有重要意义。     

The function and evolution of male and female genitalia in Phyllophaga Harris scarab beetles (Coleoptera: Scarabaeidae)

Genitalia diversity in insects continues to fuel investigation of the function and evolution of these dynamic structures. Whereas most studies have focused on variation in male genitalia, an increasing number of studies on female genitalia have uncovered comparable diversity among females, but often at a much finer morphological scale. In this study, we analysed the function and evolution of male and female genitalia in Phyllophaga scarab beetles, a group in which both sexes exhibit genitalic diversity. To document the interaction between male and female structures during mating, we dissected flash‐frozen mating pairs from three Phyllophaga species and investigated fine‐scale morphology using SEM. We then reconstructed ancestral character states using a species tree inferred from mitochondrial and nuclear loci to elucidate and compare the evolutionary history of male and female genitalia. Our dissections revealed an interlocking mechanism of the female pubic process and male parameres that appears to improve the mechanical fit of the copulatory position. The comparative analyses, however, did not support coevolution of male and female structures and showed more erratic evolution of the female genitalia relative to males. By studying a group that exhibits obvious female genitalic diversity, we were able to demonstrate the relevance of female reproductive morphology in studies of male genital diversity.     

Morphological association between spermatophores and male genitalia in carabid beetles of the tribe Pterostichini (Coleoptera: Carabidae)

A morphological association between genitalia and ejaculates could provide insight into the function and evolution of genitalia. In this study, the morphologies of the ejaculates and male genitalia of 15 species of Pterostichini and two species of Platynini (Coleoptera: Carabidae) are described. All the species examined formed a spermatophore, the morphology of which could be classified into three types based on its relative volume in the female vaginal cavity and the presence or absence of a pluglike conformation. Male genital morphology could be divided into two types by the direction of the endophallus and gonopore. Species with a strongly bent endophallus invariably formed a pluglike spermatophore. The results suggest that the peculiar shape of endophallus found in some species of Pterostichini may function in forming the pluglike structure of the spermatophore.     

狭胸天牛类雄性外生殖嚣分类研究——附一新种

该文就狭胸天牛类3属6种雄性外生殖器的形态结构,比较研究其系统发育及其归属,并记录太谷芫天牛Mantitheus tazguensts sp. nov.一新种和重新描述细点音天牛的雄性外生殖器。狭胸天牛类属较原始的类群,祖征为中茎突中央加厚,边缘极薄;阳基侧突基部还保留着一条横裂。狭胸天牛属与音天牛属和芫天牛属关系较密切,应为姊妹群,狭胸天牛属较始,其衍征为中茎突较长的分开,内囊具基骨片、骨化细长管和大小较一致骨化微刺。音天牛属和芫天牛属为姊妹群,芫天牛较原始,音天牛较进化,其衍征为阳基侧突背面因基部裂缝消失而完全合并,骨化细长管较细;外部形态表现为触角较短,较圆;中胸发音器中央无缝;而芫天牛内囊基骨片明显易见。     

Differences in genitalia structure and function between subfamilies of longhorn beetles (Coleoptera: Cerambycidae)

Beetle genitalia are usually described only for taxonomic purposes without considering the possible function of structures. Exceptions are sporadic detailed studies on single species. We studied genital structures in the subfamilies of Cerambycidae and outlined assumptions on the function of these structures and the implications for the phylogeny of the Cerambycidae. We found that male genitalia in particular are taxon-specific on a higher taxonomic level; e.g., the parameres are widely variable in Cerambycinae, while in most Lamiinae species they appear relatively uniform and differ from those of the Cerambycinae. Internal sac structures are very different among the various subfamilies. Small backwards-pointing spines are the most common armature of the internal sac. The female genitalia are less variable, although ovipositor morphology may differ among subfamilies. In most species, the connection between the mates during copulation is achieved by the long internal sac and the ovipositor only, whereas the median lobe and parameres are in contact with the female abdomen only at the beginning of copulation. Cerambycinae and Lepturinae have a basal swelling of the endophallus to prevent it from sliding back into the male abdomen during copulation. The long internal sac functions in connecting the mates and guaranteeing the sperm transfer.     

Afroprinus cavicola gen. et sp. n. from the Afrotropical region with notes on cave-dwelling Saprininae (Coleoptera,Histeridae)

A new genus and species from Kenya, Afroprinus cavicola is herein described and illustrated and its systematic position is discussed. By the prosternal pre-apical foveae connected by marginal prosternal stria it resembles most of the Afrotropical species of the genus Chalcionellus Reichardt, 1932, or some species of the genus Pholioxenus Reichardt, 1932 from South Africa and Namibia. Afroprinus can be distinguished from Chalcionellus chiefly by the lack of pronotal depressions and a coarsely sculptured, non-metallic dorsum; from Afrotropical species of Pholioxenus it can be most easily distinguished by the asetose pronotal hypomeron. The new taxon was discovered in a cave, but lacks obvious troglophilic adaptations. Notes on other Saprininae taxa found in caves are given. An identification key to the genera of Afrotropical Saprininae is provided.     

A remarkable new fossil species of Amplectister with peculiar hindleg modifications (Coleoptera: Histeridae): further evidence for myrmecophily in Cretaceous clown beetles

Myrmecophily is a phenomenon of the symbiosis of organisms that depend on various ant (Formicidae) societies. Such interspecies associations are found in several unrelated lineages within the clown beetle family Histeridae. Recent studies have suggested that the origin of myrmecophily can be traced back to mid-Cretaceous based on a few fossil records from Kachin amber from northern Myanmar. Here, we describe a remarkable new species, Amplectister terapoides n. sp., from Kachin amber. This is the second species of the extinct genus Amplectister Caterino and Maddison, which has been found from the same amber deposit and has also been considered to be myrmecophilous. The new species here described has the most heavily modified hindlegs in any fossil histerids or even beetles discovered until now, indicating further evidence for ant colony association. Our discovery demonstrates that significant and diverse morphological adaptations to myrmecophily had already occurred during the Cretaceous.     

First non-feeding Sericini beetles (Coleoptera,Scarabaeidae): new genus from Amazonia and phylogenetic position

Organisms Diversity & Evolution - Two unusual scarab species discovered from two localities in the Amazonian Basin are described. We used comparative morphology and phylogenetic analysis to...     

Phylogenetically informative rearrangements in mitochondrial genomes of Coleoptera, and monophyly of aquatic elateriform beetles (Dryopoidea)

Mitochondrial gene order in Coleoptera has been thought to be conservative but a survey of 60 complete or nearly complete genomes revealed a total of seven different gene rearrangements (deletions, gene order reversals), mainly affecting tRNA genes. All of these were found to be limited to a single taxon or a subclade of Coleoptera. The phylogenetic distribution of a translocation of tRNA(Pro) in three species of elateriform beetles was investigated further by sequencing three nearly complete mitochondrial genomes (Dascillidae, Byrrhidae, Limnichidae) and ten additional individuals for a ～1370 bp diagnostic fragment spanning the relevant region. Phylogenetic analysis consistently recovered the monophyly of families previously grouped in the contentious superfamily Dryopoidea, a group of approximately 10 beetle families with mainly aquatic lifestyles. The Byrrhidae (moss beetles) were not part of this lineage, although they may be its sister group, to recover the widely accepted Byrrhoidea. The tRNA(Pro) translocation was present in all members of Dryopoidea, but not in any other Elateriformia, providing independent support for this lineage and for a single origin of aquatic habits.     

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.     

On the Genus Trypeticus Marseul (Coleoptera: Histeridae: Trypeticinae) in China

This paper describes two new species, Trypeticus fissirostrum n. sp. from Hainan and Trypeticus yunnanensis n. sp. from Yunnan Provinces, China. Three Trypeticus species having been recorded in the territory of China before this study, their total number in this country has now increased to five. A key to the Chinese species is compiled and given in the text. The type specimens are deposited in Institute of Zoology, Chinese Academy of Sciences.     

Carcinops troglodytes (Erichson) (Coleoptera: Histeridae) preying on Alphitobius diaperinus (Panzer) (Coleoptera: Tenebrionidae) in poultry houses

We first record Carcinops troglodytes (Paykull) as a predator of Alphitobius diaperinus (Panzer) larvae in poultry houses in North and Northwest regions of the state of Paraná. Carcinops spp. are commonly recorded as predators of dipterans, and this record in poultry houses indicates the possibility of exploiting such predator for future studies aiming the development of management strategies for A. diaperinus.     

Lamellicorn beetles (Coleoptera: Scarabaeoidea) in Korea and Mongolia

This work was undertaken as an initial part of the collaborative effort between Korean and Mongolian researchers to study the terrestrial and aquatic insects in these countries. An analysis of data on faunal composition, taxonomic diversity and biogeographical structure of lamellicorn beetles in Korea and Mongolia was conducted in order to characterize the faunal relations between two countries. The Korean fauna of lamellicorn beetles is almost two fold greater in both generic diversity and species richness than that of Mongolia. The species compositions of lamellicorn beetles in two countries are substantially different from each other and the relative levels of faunal overlap are insignificant. Not only the faunal diversity and composition, but also the biogeographical structure of lamellicorn beetles in Korea and Mongolia display dissimilar patterns, except for their similar degree of endemism. Additional data on communities of dung beetles in the grasslands of central Mongolia, as well as a brief discussion on conservation issues of the endangered species in Korea, especially Gymnopleurus mopsus, are given.     

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.     

Olfactory receptor neurons detecting plant odours and male volatiles in Anomala cuprea beetles (Coleoptera: Scarabaeidae)

We have identified several types of olfactory receptor neurons in male and female Anomala cuprea beetles. The receptor neurons were sensitive to female sex pheromone components, flower volatiles, green leaf volatiles and unknown volatiles from males. Olfactory sensilla were located on three lamellae forming the antennal club. There was a clear spatial separation between some types of sensilla on each lamella. Receptor neurons for the two sex pheromone components were situated in sensilla placodea covering a specific area on each lamella in both males and females. All sex pheromone receptor neurons were found in these sensilla. Most other receptor neurons were located in a longitudinal, heterogeneous streak formed by various types of sensilla. Receptor neurons for plant-derived compounds appeared to be specialists with a high sensitivity to their respective key compound. The most remarkable among these are the green leaf volatile-specific receptor neurons, which were both sensitive and selective, with the key compound being at least 1000 times as effective as any other compound. These green leaf volatile detectors are apparently homologous to detectors recently found in the scarab Phyllopertha diversa. Our results emphasize the role of single-sensillum recordings as a tool in the identification of biologically active odours.