On the systematic position of the family Gyrinidae (Coleoptera: Adephaga)

Various characters of adult and larval members of Adephaga and Cupedidae were analyzed, and suggest that Gyrinidae are the sister-group of the remaining Adephaga, and are not closely related to the remaining aquatic Adephaga. The aquatic families Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae seem to form a well founded monophyletic unit. The following characters are considered as synapomorphies of Adephaga excluding Gyrinidae: bifurcate condition of the muscle (= M.) tentoriopraementalis inferior, reduction of hypopharynx, strongly developed prosternal process, reduction in size and specialized modification of the ventral sclerite of the mesothorax, strongly developed mesofurcal arms, a high mesopleural ridge, globular mesocoxae restricted to rotatory movements, invaginated sternum VIII (coxostemum), the strongly curved base of the median lobe of the aedeagus, which articulates with the parameres, the rotated position of the aedeagus in repose, fusion of the larval clypeolabrum with the frons and reduction of the larval lacinia. Mesal shifting of M. episterno-coxalis prothoracis, and the fusion of the apical portions of the malpighian tubules of either side are considered as synapomorphies of Adephaga excluding Rhysodidae and Gyrinidae. Lateral reduction of the meta “sternal” transverse ridge and the presence of the subcubital setal binding patch of the hind wing are considered as synapomorphic characters of Trachypachidae, Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae. We postulate that the metacoxal fusion occurred independently in gyrmids and the common ancestor of Trachypachidae, Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae. Consequently we consider this character state as another synapomorphy of Trachypachidae and Hydradephaga excluding Haliplidae and Gyrinidae. The following characters are considered as synapomorphies of Noteridae, Amphizoidae, Hygrobiidae and Dytiscidae: Loss of tactile setae on the head capsule, metafurcal origin on the intercoxal wall, expansion of the intercoxal wall, elongation of the subcubital setal binding patch, loss of Mm. furca-coxale anterior and posterior, reduction of the larval abdominal segments IX and X, and the shifting of the uropmphi onto the ventral side of segment VIII. Presence of M. tentorio-mandibularis and M. stipitopalpalis intemus are certainly primitive features of adult gyrinids but the distribution of these character states among most members of Adephaga is yet unclear. Chemical defence gland constituents point towards a very isolated position of Gyrinidae. The old age of the group, documented by a larva found in upper Permian deposits, may support the hypothesis of a sister-group relation-ship between Gyrinidae and the remainder of Adephaga.     

Molecular phylogeny of the highly disjunct cliff water beetles from South Africa and China (Coleoptera: Aspidytidae)

The superfamily Dytiscoidea contains six families with an aquatic lifestyle, with most of its extant diversity in two families: the burrowing water beetles (Noteridae) and the diving beetles (Dytiscidae). The other families have few species (up to six) and generally highly disjunct extant distributions. Aspidytidae currently contains one genus with two species, one in China and one in South Africa. Here we provide the first molecular data for the Chinese species, allowing us to explore the phylogenetic relationships and position of both species of this small family for the first time. Based on a matrix of 11 genes we inferred a phylogenetic hypothesis for Dytiscoidea including all extant families. Unexpectedly, Aspidytidae were consistently recovered as paraphyletic relative to Amphizoidae, despite being well characterized by apparently synapomorphic adult features. A re‐examination of larval characters in the two aspidytid species revealed that the larva of the Chinese species is strikingly similar to that of Amphizoidae. Both share a series of plesiomorphic features but also some potential synapomorphies, including a dense vestiture of short setae on the head capsule, anteriorly shifted posterior tentorial grooves and widely separated labial palps. Arguably these features may belong to the groundplan of the clade Aspidytidae + Amphizoidae, with far‐reaching secondary modifications (including reversals) in the South African Aspidytes niobe. At present we retain the family Aspidytidae, however, due to the strong adult morphological synapomorphies of the two extant species, and the fact that the molecular paraphyly of the family may result from the highly divergent nature of the two extant species. This long evolutionary separation and strong divergence, in terms of gene sequences and larval features, is undeniable, substantial levels of saturation in third codon positions of protein‐coding genes being present between the two taxa. We address this issue taxonomically by introducing the new genus S inaspidytes gen. nov. for the Chinese Aspidytes wrasei. The continued contentious relationships amongst Dytiscidae, Hygrobiidae, Aspidytidae and Amphizoidae highlight the need for more data to address dytiscoid phylogenetics, possibly involving a genomic approach. © 2016 The Linnean Society of London     

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.     

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.     

The systematic position of Aspidytidae, the diversification of Dytiscoidea (Coleoptera, Adephaga) and the phylogenetic signal of third codon positions

Characters of the newly discovered larvae of the South African Cliff Water Beetle Aspidytes niobe were examined and integrated into a data matrix including all families of Dytiscoidea as well as Haliplidae. Fifty-three morphological characters of adults and larvae were analysed separately and combined with molecular data from six nuclear and mitochondrial genes. The phylogeny of the group is reconstructed for the study of the evolution of swimming behaviour and larval feeding habits, as well as the shift in diversification rates leading to the two most speciose lineages. The parsimony analysis of all equally weighted morphological and molecular characters combined resulted in a single well supported tree with the topology (Noteridae (Hygrobiidae ((Aspidytidae, Amphizoidae) Dytiscidae))), in agreement with the molecular data alone, but in contradiction to the morphological data, which favoured a topology in which Hygrobiidae is sister to Dytiscidae. The exclusion of third codon positions of the three protein coding genes resulted in a topology identical to that obtained with the morphological data alone, but the use of Bayesian probabilities or the amino acid sequence resulted in the same topology as that of the tree obtained with parsimony using all equally weighted characters. We concluded that interactions of third codon positions with the other data are complex, and their removal is not justified. There was a significant increase in the diversification rate at the base of the richest families (Noteridae and Dytiscidae), which could be associated with the development of simultaneous stroke and higher swimming performance, although data on the swimming behaviour of some basal groups of Noteridae are incomplete. The presence of larval mandibular sucking channels may have contributed to the diversification of Dytiscidae and the species-rich noterid genera Hydrocanthus and Canthydrus .     

Ultraconserved elements show utility in phylogenetic inference of Adephaga (Coleoptera) and suggest paraphyly of ‘Hydradephaga’

The beetle suborder Adephaga has been the subject of many phylogenetic reconstructions utilizing a variety of data sources and inference methods. However, no strong consensus has yet emerged on the relationships among major adephagan lineages. Ultraconserved elements (UCEs) have proved useful for inferring difficult or unresolved phylogenies at varying timescales in vertebrates, arachnids and Hymenoptera. Recently, a UCE bait set was developed for Coleoptera using polyphagan genomes and a member of the order Strepsiptera as an outgroup. Here, we examine the utility of UCEs for reconstructing the phylogeny of adephagan families, in the first in vitro application a UCE bait set in Coleoptera. Our final dataset included 305 UCE loci for 18 representatives of all adephagan families except Aspidytidae, and two polyphagan outgroups, with a total concatenated length of 83 547 bp. We inferred trees using maximum likelihood analyses of the concatenated UCE alignment and coalescent species tree methods (astral ii , ASTRID, svdquartets ). Although the coalescent species tree methods had poor resolution and weak support, concatenated analyses produced well‐resolved, highly supported trees. Hydradephaga was recovered as paraphyletic, with Gyrinidae sister to Geadephaga and all other adephagans. Haliplidae was recovered as sister to Dytiscoidea, with Hygrobiidae and Amphizoidae successive sisters to Dytiscidae. Finally, Noteridae was recovered as monophyletic and sister to Meruidae. Given the success of UCE data for resolving phylogenetic relationships within Adephaga, we suggest the potential for further resolution of relationships within Adephaga using UCEs with improved taxon sampling, and by developing Adephaga‐specific probes.     

The thoracic skeleto-muscular system of Mengenilla (Strepsiptera: Mengenillidae) and its phylogenetic implications

The thorax of Mengenilla was examined using traditional morphological techniques and its features were documented in detail using scanning electron microscopy and computer-based 3D reconstructions. The results were compared to conditions found in other holometabolan insects. The implications for the systematic placement of Strepsiptera are discussed. The observations are interpreted in the light of the recently confirmed sistergroup relationship between Strepsiptera and Coleoptera (Coleopterida). The synapomorphies of the thorax of Strepsiptera and Coleoptera are partly related with posteromotorism (e.g., increased size of the metathorax), partly with a decreased intrathoracic flexibility (e.g., a fused pronotum and propleurum), and partly independent from these two character complexes (e.g., not connected profurca and propleuron). Strepsiptera are more derived than Coleoptera in some thoracic features (e.g., extremely enlarged metathorax) but have also preserved some plesiomorphic conditions (e.g., tegulae in both pterothoracic segments). All potential apomorphies of Mecopterida are missing in Strepsiptera. The last common ancestor of Coleopterida had already acquired posteromotorism but the wings were still largely unmodified. Several reductions in the mesothorax likely occurred independently.     

Phylogenetic analysis of Adephaga (Coleoptera) based on characters of the larval head

Abstract. Characters of the head of adephagan larvae were examined and analysed phylogenetically. A labrum which is completely fused to the clypeofrons and the presence of a closed prepharyngeal tube are autapomorphies of Adephaga. Partial reduction of the fossa maxillaris, cardo and stipes forming a functional unit, the immobilization of the lacinia, attachment of M. craniolacinialis to the lateral stipital wall, and loss of one stipitopalpal muscle, are considered autapomorphies of Adephaga excluding Gyrinidae. Complete reduction of the fossa maxillaris and the presence of M. craniostipitalis medialis are possible autapomorphies of Adephaga excluding Gyrinidae and Haliplidae. The presence of caudal tentorial arms, insertion of the galea on the mesal side of palpomere I, and absence of the lacinia are considered synapomorphies of Trachypachidae and Dytiscoidea (Noteridae, Amphizoidae, Hygrobiidae, Dytiscidae). The presence of a slender, elongated process of the head capsule, which articulates with a corresponding socket of the cardo, is a possible autapomorphy of Dytiscoidea. The sinuate frontal sutures, distinctly protruding prementum, shortened M. craniostipitalis medialis, and absence of M. submentopraementalis are considered autapomorphies of Geadephaga excluding Trachypachidae. The presence of a regular row of hairs along the anterior hypopharyngeal margin is a possible autapomorphy of Geadephaga excluding Trachypachidae and Rhysodidae. Improvement of the hypopharyngeal filter apparatus suggests the monophyly of Anisochaeta. Presence of a penicillum and partial reduction of the lacinia are possible autapomorphies of Anisochaeta excluding Omophronini. Larvae of Cychrini, Carabini, Nebriini and Notiophilini are characterized by a strongly developed, cone-shaped hypodon. Postocular and cervical ridges, crosswise arrangement of antennal muscles, and a completely flattened hypopharynx are considered autapomorphies of Caraboidea Limbata.     

Phylogeny of hydradephagan water beetles inferred from 18S rRNA sequences

Several families in the beetle suborder Adephaga have an aquatic life style and are commonly grouped in the "Hydradephaga," but their monophyly is contentious and relationships between and within these families are poorly understood. Here we present full-length 18S rRNA sequence for 84 species of Hydradephaga, including representatives of most major groups down to the tribal level, and a total of 68 species of the largest family, Dytiscidae. Using a direct optimization method for the alignment of length-variable regions, the preferred tree topology was obtained when the cost of gaps and the cost of nucleotide changes were equal, and three hypervariable regions of 18S rRNA were downweighted by a factor of five. Confirming recent molecular studies, the Hydradephaga were found to be monophyletic, indicating a single colonization of the aquatic medium. The most basal group within Hydradephaga is Gyrinidae, followed in a comb-like arrangement by families Haliplidae, Noteridae, Amphizoidae, and Hygrobiidae plus Dytiscidae. Under most alignment parameters, Hygrobiidae is placed amid Dytiscidae in an unstable position, suggesting a possible data artifact. Basal relationships within Dytiscidae are not well established, nor is the monophyly of subfamilies Hydroporinae and Colymbetinae. In contrast, relationships at the genus level appear generally well supported. Despite the great differences in the rates of change and the significant incongruence of the phylogenetic signal in conserved vs hypervariable regions of the 18S rRNA gene, both contribute to establish relationships at all taxonomic levels.     

The systematic position of Meruidae (Coleoptera, Adephaga) and the phylogeny of the smaller aquatic adephagan beetle families

A phylogenetic analysis of Adephaga is presented. It is based on 148 morphological characters of adults and larvae and focussed on a placement of the recently described Meruidae, and the genus‐level phylogeny of the smaller aquatic families Gyrinidae, Haliplidae and Noteridae. We found a sister group relationship between Gyrinidae and the remaining adephagan families, as was found in previous studies using morphology. Haliplidae are either the sister group of Dytiscoidea or the sister group of a clade comprising Geadephaga and the dytiscoid families. Trachypachidae was placed as the sister group of the rhysodid‐carabid clade or of Dytiscoidea. The monophyly of Dytiscoidea including Meru is well supported. Autapomorphies are the extensive metathoracic intercoxal septum, the origin of the metafurca from this structure, the loss of Mm. furcacoxalis anterior and posterior, and possibly the presence of an elongated subcubital setal binding patch. Meruidae was placed as sister group of the Noteridae. Synapomorphies are the absence of the transverse ridge of the metaventrite, the fusion of abdominal segments III and IV, the shape of the strongly asymmetric parameres, and the enlargement of antennomeres 5, 7 and 9. The Meru‐noterid clade is the sister group of the remaining Dytiscoidea. The exact position of Aspidytes within this clade remains ambiguous: it is either the sister group of Amphizoidae or the sister group of a clade comprising this family and Hygrobiidae + Dytiscidae. The sister group relationship between Spanglerogyrinae and Gyrininae was strongly supported. The two included genera of Gyrinini form a clade, and Enhydrini are the sister group of a monophylum comprising the remaining Enhydrini and Orectochilini. A branching pattern (Peltodytes + (Brychius + Haliplus)) within Haliplidae was confirmed. Algophilus, Apteraliplus and the Haliplus‐subgenus Liaphlus form a clade. The generic status of the two former taxa is unjustified. The Phreatodytinae are the sister group of Noterinae, and Notomicrus (+ Speonoterus), Hydrocoptus, and Pronoterus branch off successively within this subfamily. The search for the larvae of Meru and a combined analysis of morphological and molecular data should have high priority. © The Willi Hennig Society 2006.     

Before the wasp-waist: comparative anatomy and phylogenetic implications of the skeleto-musculature of the thoraco-abdominal boundary region in basal Hymenoptera (Insecta)

The skeleto-musculature of the metathorax and first abdominal segment was studied in representatives from all ’symphytan’ families. Forty-three informative characters were coded and scored. The distribution of character states are discussed with reference to recent cladistic treatments of the Hymenoptera. Previously unreported autapomorphies for the Hymenoptera are the separation of the metathoracic trochantins from the metepisterna and metacoxae, the position of the metafurca anteriorly on the discrimenal lamella of the metathorax and the presence of second abdominal sternum (S2)-metacoxal muscles. The absence of metapleuro-S2 muscles is an autapomorphy for the non-xyelid Hymenoptera. Putative autapomorphies of the Tenthredinoidea are: (1) the presence of transverse metanotal muscles, (2) the subdivision of the second phragmo-third phragmal muscles, part of which arises from the metalaterophragmal lobes, (3) the posterior thoracic spiracle occlusor muscles arising from the mesepisterna, (4) the absence of trochantins and metanoto-trochantinal muscles and (5) the presence of elongate lateral metafurcal arms. Having the paracoxal sulci extending along the anterior margins of the metepisterna and the anterior metafurcal arms reduced are synapomorphies for all tenthredinoid families excluding Blasticotomidae. The presence of transversely extended cenchri with hooks on their entire surface is a putative synapomorphy for Diprionidae + Cimbicidae + Argidae + Pergidae. The clade Cimbicidae + Argidae + Pergidae is supported by the absence of metanoto- metabasalar muscles, the fusion of the first abdominal tergite (T1) with the metepimera and the absence of posterior metapleuro-metafurcal muscles. Autapomorphies of the Cimbicidae are the absence of the metalaterophragmal lobes and the metalaterophragmal-metafurcal muscles. Having the mesoscutello-metanotal muscle inserting on a projection from the anterior margin of the metanotum, surrounding the tendon with sclerotised cuticle, is a synapomorphy for the Argidae and Pergidae. Autapomorphies of the Cephoidea are the absence of cenchri, the presence of distinct articulations between T1 and the metepimera, and having the paracoxal sulci extending subparallel with the metafurcal discrimen. The monophyly of the Siricidae is supported by the absence of the anapleural clefts and the presence of an elongate mesospina projecting posteriorly between the anterior metafurcal arms. The presence of a membranous pouch ventrally of T1 and of large T1-metafurcal muscles is unique to Xiphydria camelus among the taxa examined. The absence of hind wing tegulae, posterior metapleuro-metafurcal, metanoto-trochantinal and anterior metanoto-metacoxal muscles, and the presence of elongate lateral metafurcal arms are synapomorphies for Xiphydriidae + Orussidae + Apocrita. The Orussidae greatly resembles the Apocrita in the region studied, a synapomorphy for the two taxa being the presence of metepisternal depressions. An autapomorphy for the Apocrita is the fusion of T1 with the metapleural arms; these structures closely abut in Orussidae. The fusion of T1 with the metepimera was preceded by the reduction of the posterior parts of the metepimera, as observed in Anaxyelidae, Xiphydriidae, and Orussidae. This makes the lines of fusion between T1 and the metepimera confluent with the metapleural sulci in the Apocrita. There is no compelling evidence for considering the configuration of T1 and the metepimera in Cephoidea to be incipient in the formation of the propodeum in Apocrita. The close association between the meso- and metathorax and the integration of T1 in the metathorax evolved gradually twice within the basal hymenopteran lineages, culminating in the Apocrita and the Cimbicidae + Argidae + Pergidae clade. Accepted: 2 September 1999     

New evidence on the phylogeny and biogeography of the Amphizoidae: Discovery of a new species from China (Coleoptera)

Abstract. A new species, Amphizoa sinica, is described from Chang bei shan in Northern China. A revised key is provided for this and the other four species of this monogeneric and enigmatic group. Phylogcnetic and zoogeographic relationships are discussed.     

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。     

Constancy and variation of the serotonin-like immunoreactive neurons in the metamorphosing ventral nerve cord of the meal beetle,Tenebrio molitor L. (Coleoptera : Tenebrionidae)

Serotonin-like immunoreactive neurons were mapped in the larval, prepupal, pupal, and adult ventral nerve cord (VNC) of the beetle, Tenebrio molitor L. (Coleoptera: Tenebrionidae). The alterations of the shape of these neurons during metamorphosis were analysed. The stage-specific interindividual variability of the examined serotonin-like immunoreactive neurons is low. Serotonin-like immunoreactive neurons of the abdominal and thoracic ganglia behave differently during metamorphosis. Only in thoracic ganglia was an obvious change in the pattern of serotonin-like immunoreactive neurons observed. The shape of the dendritic trees of serotonin-like immunoreactive neurons varies in thoracic., but not in abdominal ganglia. During postlarval development, new emerging neurons that react with the anti-serotonin antibody are found only in the thoracic ganglia. Serotonin-like immunoreactive neurons are serially homologous in the larval ventral nerve cord. The basic organization of the serotonin-like immunoreactive neurons is maintained up to the adult stage. Some aspects of the metamorphosis of the nervous system are discussed with respect to the transformation of the set of immunoreactive neurons from larval to adult stage. The results are compared to those obtained in the study of serotonin-immunoreactive neurons in cockroaches, dipterans and locusts.     

Field observation of predation on a horsehair worm (Gordioida: Chordodidae) by a diving beetle larva Cybister brevis Aubé (Coleoptera: Dytiscidae)

Diving beetles (Coleoptera: Dytiscidae) are carnivorous in both the larval and adult stages; larvae are exclusively predatory, whereas adults also scavenge for food. They are known to prey on zooplankton, insects, gastropods, fish, amphibians and reptiles. However, there have been no previous reports detailing direct predation on adult Gordioida by Dytiscidae in the field. This study represents the first observation of a diving beetle larva, Cybister brevis Aubé (Coleoptera: Dytiscidae), predating on an adult horsehair worm (Gordioida: Chordodidae). This might be the first report of predation on horsehair worms by insects.     

The unique locomotor apparatus of whirligig beetles of the tribe Orectochilini (Gyrinidae,Coleoptera)

Whirligig beetles, which are known for their rapid gliding on the water surface, have evolved a unique locomotor apparatus. External and internal thoracic structures of Orectochilus villosus (Orectochilini) are described in detail and documented with microcomputed tomography, computer‐based 3D reconstructions, and scanning electronic microscopy (SEM). The results are compared with conditions found in other genera of Gyrinidae and other groups of Coleoptera. The focus is on structures linked with locomotion, especially on the unusual flight apparatus, which differs strongly from that of other beetles. As in the other Orectochilini, the prothorax of Orectochilus displays characters typical for Gyrinidae, with triangular procoxae and forelegs transformed into elongated, sexually dimorphic grasping devices. The musculature of this segment is similar to the pattern found in other Coleoptera. Similar to all other extant Gyrinidae, the mesothorax is characterized by an extensive and flat mesoventrite, suitable for gliding on the water surface. As in Heterogyrinae and the other Gyrininae, the pterothoracic legs are transformed into paddle‐like structures, enabling the beetles to move with high speed on the surface film. The musculature of the mesothorax is reduced compared to other Coleoptera, but similar to what is found in the other Gyrininae. The metathoracic skeleton and musculature are simplified in Orectochilini compared to other Gyrininae and other groups of Coleoptera. In O. villosus, only 10 metathoracic muscles are preserved. 36 are present in an archostematan beetle, a condition probably close to the coleopteran ground plan. The metathoracic dorsal longitudinal bundles are absent in Gyrininae, muscles that play a role as indirect flight muscles in most other neopteran insects. The rest of the posteromotoric flight apparatus is distinctly modified, with a limited number of skeletomuscular elements taking over more functions. The large muscle M84 (IIIdvm7) M. noto–trochanteralis, for instance, functions as dominant wing levator, but is also responsible for the powerful and rapid backstroke of the hind legs. The presence of this muscle is a synapomorphy of Heterogyrinae and Gyrininae. The narrow metafurca in the latter group is likely linked to its large size. The elytra likely contribute to the control of the flight of the beetle, whereas they shield and inhibit the flight apparatus during swimming.     

豉甲科及水生肉食亚目的分子系统发育学分析(昆虫纲：鞘翅目)

利用PAUP和MrBayes软件,对线粒体COⅠ基因序列3个密码子位置的数据模块分别进行了豉甲科(Gyrinidae)和水生肉食亚目(Hydradephaga)在亚科或科水平上的系统发育学分析,结果表明第二密码子数据模块获得了理想的分析结果。由PAUP生成的豉甲科最优树来自第二密码子数据模块的分析,而由MrBayes生成的最优树来自全部密码子数据模块的分析。此外,用对应的氨基酸序列生成的ME和MP树与第二密码子数据模块分析的结果也一致。亚科Orectochilinae和Gyrininae以高的支持率形成了单系。然而,来自亚科Enhydrinae的种Porrorhynchus landaisi landaisi呈现了异常的位置。SH-test检验也支持该异常位置,表明这个种可能代表了一个科。在来自第二密码子数据模块的水生肉食亚目最优ML树中,整个Hydradephaga树呈现单系,豉甲科位于树的基部,表明了该科在水生肉食亚目中是一个早期的分支。在树中还产生了一个单系的Dytiscoidea总科,由Dytiscidae、Hygrobiidae、Noteridae和Amphizoidae 4个科组成,单系的Haliplidae与之成为姐妹群。此外线粒体分子钟的结果表明豉甲科的5对相近种间的分化是一个短时期内发生的(0.01～1.81百万年前),这点可能与它们的特殊地理分布有关。     

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     

The structure of the defence glands of the Cicindelidae, Amphizoidae, and Hygrobiidae (Insecta: Coleoptera)

An account is given of the structure of the thoracic and pygidial defence glands of Hygrobia hermanni (F.) and of the pygidial glands of Amphizoa insolens Leconte, and Cicindela campestris L. The phylogenetic significance of differences in structure is discussed.     

A new sensory organ in “primitive” molluscs (Polyplacophora: Lepidopleurida), and its context in the nervous system of chitons

Introduction

Chitons (Polyplacophora) are molluscs considered to have a simple nervous system without cephalisation. The position of the class within Mollusca is the topic of extensive debate and neuroanatomical characters can provide new sources of phylogenetic data as well as insights into the fundamental biology of the organisms. We report a new discrete anterior sensory structure in chitons, occurring throughout Lepidopleurida, the order of living chitons that retains plesiomorphic characteristics.

Results

The novel “Schwabe organ” is clearly visible on living animals as a pair of streaks of brown or purplish pigment on the roof of the pallial cavity, lateral to or partly covered by the mouth lappets. We describe the histology and ultrastructure of the anterior nervous system, including the Schwabe organ, in two lepidopleuran chitons using light and electron microscopy. The oesophageal nerve ring is greatly enlarged and displays ganglionic structure, with the neuropil surrounded by neural somata. The Schwabe organ is innervated by the lateral nerve cord, and dense bundles of nerve fibres running through the Schwabe organ epithelium are frequently surrounded by the pigment granules which characterise the organ. Basal cells projecting to the epithelial surface and cells bearing a large number of ciliary structures may be indicative of sensory function. The Schwabe organ is present in all genera within Lepidopleurida (and absent throughout Chitonida) and represents a novel anatomical synapomorphy of the clade.

Conclusions

The Schwabe organ is a pigmented sensory organ, found on the ventral surface of deep-sea and shallow water chitons; although its anatomy is well understood, its function remains unknown. The anterior commissure of the chiton oesophagial nerve ring can be considered a brain. Our thorough review of the chiton central nervous system, and particularly the sensory organs of the pallial cavity, provides a context to interpret neuroanatomical homology and assess this new sense organ.