Study on the systematic position of Metriini based on characters of the larval head (Coleoptera: Carabidae)

Abstract. Characters of the head of larvae of Metrius contractus Eschscholtz, Ozaenini and Paussini are interpreted phylogenetically. The monophyly of Metriini + Ozaenini + Paussini is substantiated by several synapomorphies such as hyperprognathism and strong constriction of the neck. Ozaenini and Paussini together form the sister-group of Metriini. Ozaenini are paraphyletic. The monophyly of Paussini + Ozaenini excluding Pachyteles is indicated by two possible synapomorphies. Several synapomorphies are shared by Physea + Paussini. Secondary prognathism, large membranous submento-mental area and other derived features are considered autapomorphies of Paussini. Paussini excluding Platyrhopalopsis are characterized by the loss of the palpifer. The monophyly of a group which comprises Geadephaga excluding Trachypachini is suggested by several synapomorphic features. A very basal position of the metriine—paussine lineage within Carabidae is indicated by several plesiomorphic features. A hypopharyngeal filter apparatus with a dense fringe of well-arranged, long hairs is a possible autapomorphy of Anisochaeta. The results of this study do not indicate a close relationship between the metriine—paussine lineage and the tribes Brachinini and Crepidogastrini as has been suggested in recent works.     

A phylogenetic study of Dermestidae (Coleoptera) based on larval morphology

Abstract.  The results of a phylogenetic analysis of Dermestidae (Coleoptera) based on the morphology of immature stages are presented and implications for the classification of the family are discussed. The monophyly of (( Orphilus ) + (Dermestidae) + (Endecatomidae (Bostrichidae + Anobiidae))) is strongly supported. Dermestidae, excluding Orphilus Erichson, forms a monophyletic group. The phylogenetic position of the Orphilinae remains enigmatic. The hypothesized relationships of the remaining dermestid subfamilies are (Thorictinae (Dermestinae (Attageninae (Megatominae + Trinodinae)))). Phylogenetically informative morphological characters of larvae and pupae are illustrated and described. Character state transformations implied by the phylogenetic hypothesis are discussed. New morphological observations are provided, including the first records of the presence of gastric caeca in the larva of Dermestes L., and the first report of the number of Malpighian tubules and the number of abdominal ganglia in Dermestidae, Bostrichidae, Anobiidae, Nosodendridae and Derodontidae. The form of the mandibular base is described for the first time for Thylodriini Beal and Trinodini Beal. The fossil record and distribution of extant Dermestidae are discussed in the light of the phylogenetic hypothesis. The origin of the family is hypothesized to be in the Late Cretaceous.     

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 Psephenidae (Coleoptera: Byrrhoidea) based on larval, pupal and adult characters

Abstract.  We conducted the first comprehensive phylogenetic analysis of Psephenidae, based on 143 morphological characters of adults, larvae and pupae and coded for 34 taxa, representing three outgroups and 31 psephenid genera, including four undescribed ones. A strict consensus tree calculated (439 steps, consistency index = 0.45, retention index = 0.75) from the two most-parsimonious cladograms indicated that the monophyly of the family and subfamilies is supported, with the exception of Eubriinae, which is paraphyletic when including Afroeubria . Here a new subfamily, Afroeubriinae ( subfam.n. ), is formally established for Afroeubria . The analysis also indicated that the 'streamlined' larva is a derived adaptive radiation. Here, suprageneric taxonomy and the evolution of some significant characters are discussed. Keys are provided to the subfamilies and genera of Psephenidae considering larvae, adults and pupae.     

A preliminary study on the phylogenetic relationships of Copelatus Erichson (Coleoptera: Dytiscidae: Copelatinae) based on larval chaetotaxy and morphology

The phylogenetic relationships of the diving-beetle (Dytiscidae) subfamily Copelatinae are not well known. Some authors postulated a sister-group relationship between Copelatus Erichson and the remaining Dytiscidae, relying mainly on the absence of a mandibular channel in Copelatus. Other authors suggested a more derived position of the genus. Larval characters have been underutilized in phylogenetic studies, mainly because the larvae of many taxa within the family and, in particular, within Copelatinae are unknown. A comprehensive approach aimed to study the phylogenetic placement of a member of this subfamily based on larval characters has not been produced so far. In this study, the phylogenetic relationships of Copelatus are explored based on a cladistic analysis of 24 taxa and 120 chaetotaxic and morphological characters from larvae. For this purpose, larvae of Copelatus longicornis Sharp are described and illustrated in detail for the first time, with particular emphasis on morphometry and chaetotaxy, with the latter being unexplored until now. The results support a derived position of Copelatus within Dytiscidae, with a sister-group relationship between this genus and a clade formed by the subfamilies Lancetinae, Coptotominae, Laccophilinae, Colymbetinae, Matinae, and Dytiscinae, and part of Agabinae. No evidence was found for a sister-group relationship between Copelatus and the remaining Dytiscidae so that the absence of a mandibular channel in this genus is likely a reduction. Copelatus is supported by three apomorphies within Dytiscidae: mandibular channel absent, internal margin of the stipes with three robust spinulae, and seta MX8 inserted subapically on the galea. Handling editor: K. Martens     

Higher‐level phylogeny of diving beetles (Coleoptera: Dytiscidae) based on larval characters

A comprehensive higher‐level phylogeny of diving beetles (Dytiscidae) based on larval characters is presented. Larval morphology and chaetotaxy of a broad range of genera and species was studied, covering all currently recognized subfamilies and tribes except for the small and geographically restricted Hydrodytinae, where the larva is unknown. The results suggest several significant conclusions with respect to the systematics of Dytiscidae including the following: monophyly of all currently recognized subfamilies, although Dytiscinae when considered in a broad context is rendered paraphyletic by Cybistrinae; currently recognized tribes are monophyletic except for Agabini, Hydroporini and Laccornellini; inter‐subfamily and inter‐tribe relationships generally show weak support, except for a few well supported clades; three distinct clades are recognized within Dytiscinae [Dytiscini sensu lato (i.e. including the genera Dytiscus Linnaeus and Hyderodes Hope), Hydaticini sensu lato, and Cybistrini]; and recognition of Pachydrini as a distinct tribe. Other less robust results include: Methlini sister to the rest of Hydroporinae; relative basal position of Laccornini, Hydrovatini and Laccornellini within Hydroporinae; close relationship of Agabinae and Copelatinae; Matinae nested deep within Dytiscidae, as sister to a large clade including Colymbetinae, Coptotominae, Lancetinae and Dytiscinae sensu lato; the sister‐group relationship of Agabetini and Laccophilini is confirmed. The results presented here are discussed and compared with previous phylogenetic hypotheses based on different datasets, and the evolution of some significant morphological features is discussed in light of the proposed phylogeny. All suprageneric taxa are diagnosed, including illustrations of all relevant synapomorphies, and a key to separate subfamilies and tribes is presented, both in traditional (paper) format and as an online Lucid interactive identification key.     

Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae)

Faille, A., Casale, A. & Ribera, I. (2010). Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae). —Zoologica Scripta, 40, 282–295. Carabid beetles of tribe Trechini (Coleoptera) are one of the main groups of insects that colonized the subterranean environment. Many species of this group have developed similar morphological modifications related to the subterranean life, resulting in a characteristic Aphaenops‐like phenotype that obscures their phylogenetic relationships (depigmented, blind, elongated body and appendages, narrow head and pronotum). We present here the result of a molecular study using a combination of nuclear (small ribosomal unit, large ribosomal unit) and mitochondrial (cox1, cyb, rrnL, trnL, nad1) genes to investigate the phylogenetic placement of the highly modified subterranean genera of the tribe Trechini from the west Mediterranean area (France, Spain, Morocco and Sardinia). Our results confirm the multiple independent origin of troglomorphism among these genera, and reveal a pattern largely determined by geographical proximity. We discuss the validity of some groups proposed on the base of morphological features, and provide estimates of divergence between subterranean genera and other groups of Trechini, including epigean species of the same area. We compare the estimated age for the origin of the main groups resulting from two different calibrations, using one the standard mitochondrial mutation rate (2.3% divergence per Myr) and the other the separation between Sardinia and mainland 33 Ma. Under the first scenario, the main groups of genera would have a late Miocene origin, with a subsequent colonization of north Africa at the Pliocene–Pleistocene boundary. The assumption that the main groups originated through vicariance due to the separation of the Sardinian plate in the Oligocene results in a Messinian origin of the north African subterranean taxa, and a global mitochondrial rate reduced to 1% divergence per Myr.     

Phylogenetic analysis of Myxophaga (Coleoptera) using larval characters

A phylogenetic parsimony analysis of fifty-four larval characters of Myxophaga (excluding Lepiceridae) resulted in two minimal length cladograms. The monophyly of Torridincolidae, Hydroscaphidae and Microsporidae is supported by several autapomorphies: miniaturization, flattened body with laterally extended tergites, broadened head, scale-like surface structures, broad tentorial bridge, disc-shaped labral sensilla, spiracular gills and pupation in the last larval exuviae. Hydroscaphidae are the sister group of Microsporidae. Larvae of both families are characterized by semi-entognathous mouthparts, tergites with posterior rows of lancet-shaped setae, claws with flattened basal spines and balloon-shaped spiracular gills. The monophyly of all families is supported by autapomorphies. Torridincolidae excluding Delevea is defined as a monophylum by four derived character states: body ovoid, thorax semicircular and as long as abdomen, labral sensilla fused and abdominal sternite IX distinctly reduced and triangular. The monophyly of Torridincolinae (sensu Endrödy-Younga 1997b) is supported by two autapomorphies. The proposed branching pattern suggests that the early representatives of Myxophaga (excluding Lepiceridae) were living in aquatic conditions with a preference for hygropetric habitats. The tendency to live on rocks in running water and miniaturization have played an important role in myxophagan evolution.     

Higher-level phylogeny of Hydrophilinae (Coleoptera: Hydrophilidae) based on larval, pupal and adult characters

Abstract.  A phylogenetic analysis, at a tribal and subtribal level, of the subfamily Hydrophilinae was conducted. The analysis was based on twenty-nine taxa (twenty-three genera) and 148 characters (fifty-eight from immature stages and bionomics, and ninety from adults). According to the present study, Hydrophilinae is monophyletic, and except for the tribe Hydrophilini which appears as polyphyletic (it includes the subtribes Hydrophilina, Hydrobiina, and Acidocerina), the remaining tribes are monophyletic. The tribes Berosini and Chaetarthriini form the basal clade of Hydrophilinae. One unexpected result is the relationship between the subtribe Hydrobiina and the tribe Sperchopsini, which form a well-supported clade. The final tree has the following structure: (((((Sperchopsini Hydrobiina) (Anacaenini Laccobiini)) Acidocerina) Hydrophilina) (Berosini Chaetarthriini)). The results partially disagree with the phylogeny presented by Hansen, in 1991, which was based mostly on adult characters. Several evolutionary trends are briefly discussed: the types of egg case, the morphology of the clypeolabrum, mouthparts, legs, and breathing adaptations in larvae.     

New morphological aspects and phylogenetic considerations of Cicindis Bruch (Coleoptera: Carabidae: Cicindini)

Cicindis Bruch is a monospecific genus of carabid beetles endemic to Argentina. In this contribution, Cicindis horni Bruch is re-described, with addition of new morphological features of male internal sac, female genital tract and elytral closure. New information on the species' habitat and distribution is also provided. The phylogenetic placement and relationships of Cicindis within the family Carabidae are discussed on the basis of a cladistic analysis. Terminal taxa included representatives of all subfamilies of Carabidae and supertribes of Carabinae, with a major sampling of those taxa considered to be closely related to Cicindini by previous authors. The phylogenetic analysis shows the basal position of Cicindis in a clade that includes Ozaeninae, Omophronini, Scaritinae and Conjuncta. A close relationship of Cicindis with Ozaenini + Metriini is supported by the particular closure of the procoxa and the ventral position of the oviduct with respect to the spermatheca.     

Evaluation of relationships within the endemic Hawaiian Platynini (Coleoptera: Carabidae) based on molecular and morphological evidence

Relationships among 69 species of Hawaiian Platynini, a monophyletic beetle radiation, was investigated based on evidence from five data partitions, comprising mitochondrial and nuclear DNA sequences (cytochrome oxidase II, 624 bp; cytochrome b, 783 bp; 28S rDNA, 668 bp; wingless; 441 bp) and morphology (206 features of external and internal anatomy). Results from individual and combined data analyses generally support the monophyly of three putative divisions within Platynini in Hawaii: Division 0 (Colpocaccus species group), Division 1 (Blackburnia species group), and Division 2 (Metromenus species group). However, relationships within and among these three divisions differ from previous morphological hypotheses. An extensive series of sensitivity analyses was performed to assess robustness of recovered clades under a variety of weighted parsimony conditions. Sensitivity analyses support the monophyly of Divisions 0 and 1, but were equivocal for the monophyly of Division 2. A phylogeny based on combined data suggests at least four independent losses/reductions of platynine flight wings. The combined analysis provides corroboration for biogeographic hypotheses, including (1) colonization of Kauai by Hawaiian Platynini with subsequent dispersal and colonization along the island chain from Oahu to Maui Nui to Hawaii Island and (2) incongruent area relationships among Eastern Molokai, West Maui, and Haleakala for two species triplets.     

Carabidae from Vietnam (Coleoptera)

Members of the Vietnamese carabid fauna are arranged and nomenclaturally revised. As the result, 178 species belonging to 74 genera of 12 subfamilies, Brachininae, Omophroninae, Carabinae, Scaritinae, Bembidiinae, Trechinae, Pterostichinae, Harpalinae, Callistinae, Oodinae, Odacanthinae, and Lebiinae, are recorded from Vietnam. Within these, four species, Abacetus leucocetes Bates, 1873, Ago-nom (Metacolpodes) buchanani Hope, 1831, Steno-lopus (Egadromd) difficilis (Hope, 1845), and Phe-ropsophus {Stenaptinus) beckeri Jedlicka, 1930 are newly recorded. A species checklist with taxonomie information, type localities and worldwide distribution are provided.     

Testing abundance-range size relationships in European carabid beetles (Coleoptera, Carabidae)

Four of the eight hypotheses proposed in the literature for explaining the relationship between abundance and range size (the sampling artifact, phylogenetic non-independence, range position and resource breadth hypotheses) were tested by using atlas data for carabid beetles (Coleoptera, Carabidae) from Belgium, Denmark and the Netherlands. A positive relationship between abundance and partial range size was found in all three countries, and the variation in abundance was lower for widespread species. Analysis of the data did not support three of the proposed hypotheses, but did support the resource breadth hypothesis (species having broader environmental tolerances and being able to use a wider range or resources will have higher local densities and be more widely distributed than more specialised species). Examination of species' characteristics revealed that widespread species are generally large bodied, generalists (species with wide niche breadths occurring in a variety of habitat types) and are little influenced by human-altered landscapes, while species with restricted distributions are smaller bodied, specialists (species with small niche breadths occurring in only one or two habitat types), and favour natural habitat. Landscape alteration may be an important factor influencing carabid abundance and range size in these three countries with a long history of human-induced environmental changes.     

Weighted compromise trees: a method to summarize competing phylogenetic hypotheses

A new consensus method for summarizing competing phylogenetic hypotheses, weighted compromise, is described. The method corrects for a bias inherent in majority‐rule consensus/compromise trees when the source trees exhibit non‐independence due to ambiguity in terminal clades. Suggestions are given for its employment in parsimony analyses and tree resampling strategies such as bootstrapping and jackknifing. An R function is described that can be used with the programming language R to produce the consensus.     

Ground beetles (Coleoptera: Carabidae) as bioindicators

One of the primary goals of research on bioindicators is to identifyspecies or other taxonomic units that would reliably indicate disturbances inthe environment, and reflect the responses of other species or the overallbiodiversity. However, there is no perfect bioindicator and selecting the mostsuitable one depends to a great extent on the goal of the survey. In this paperwe examine the suitability of carabids as bioindicators. Carabids are frequentlyused to indicate habitat alteration. They have been used in grasslands andboreal forests where species number and/or abundances have been noted to changealong a habitat disturbance gradient. A common trend is that large, poorlydispersing specialist species decrease with increased disturbance while smallgeneralist species with good dispersal ability increase. Some species are notaffected by moderate disturbance. There is, however, not enough research todetermine how suitable carabids are for biodiversity studies, or how well theyrepresent the response of other species. We conclude that carabids are usefulbioindicators, but as crucial understanding of their relationship with otherspecies is incomplete, they should be used with caution.