This paper reports the collecting of adult beetles and third-instar larvae of Coelocorynus desfontainei Antoine, 1999 in Cameroon and provides new data on the biology of this high-altitude Afromontane genus. It also presents the first diagnosis of this genus based on larval characters and examination of its systematic position in a phylogenetic context using 78 parsimony informative larval and adult characters. Based on the results of our analysis we (1) support the hypothesis that the tribe Trichiini is paraphyletic with respect to both Valgini and the rest of the Cetoniinae, and (2) propose that the Trichiini subtribe Cryptodontina, represented by Coelocorynus, is a sister group of the Valgini: Valgina, represented by Valgus. The larvae-only analyses were about twofold better than the adults-only analyses in providing a phylogenetic resolution consistent with the larvae + adults analyses. Only one of the ten clades was consistently supported by the analyses of both the larval and adult datasets, while the remaining nine were invariably strongly supported by one but not the other analysis, thus highlighting the importance of employing different data sources.
Selected representatives of Cucujoidea, Cleroidea, Tenebrionoidea, Chrysomelidae, and Lymexylidae were examined. External and internal head structures of larvae of Sphindus americanus and Ericmodes spp. are described in detail. The data were analyzed cladistically. A sister group relationship between Sphindidae and Protocucujidae is suggested by the vertical position of the labrum. The monophyly of Cucujiformia is supported by the reduced dorsal and anterior tentorial arms, fusion of galea and lacinia, and the presence of tube-like salivary glands. Absence of M. tentoriopraementalis inferior and presence of a short prepharyngeal tube are potential synapomorphies of Cleroidea, Cucujoidea and Tenebrionoidea. The monophyly of Cleroidea and Cucujoidea is suggested by the unusual attachment of the M. tentoriostipitalis to the ventral side of the posterior hypopharynx. Cucujoidea are paraphyletic. The families Endomychidae, Coccinellidae and Nitidulidae are more closely related to the monophyletic Cleroidea, than to other cucujoid groups. Separation of the posterior tentorial arms from the tentorial bridge and presence of a maxillolabial complex are synapomorphic features of Cleroidea and these cucujoid families. For a reliable reconstruction of cucujoid interrelationships, further characters and taxa need to be studied.
External and internal head structures and external structures of the thorax and abdomen of larval representatives of Melandryidae (Orchesia), Ulodidae (Meryx), Oedemeridae (Pseudolycus) and Pythidae (Pytho) are described. The obtained data were compared to characters of other tenebrionoid larvae and to larval characters of other representatives of Cucujiformia. Characters potentially relevant for phylogenetic reconstruction are listed and were analysed cladistically. The data set is characterised by a high degree of homoplasy and the resolution of the strict consensus trees of 2650 or 815 (second analysis) minimal length trees is low. The monophyly of Tenebrionoidea is supported by several larval autapomorphies, e.g. posteriorly diverging gula, anteriorly shifted posterior tentorial arms, asymmetric mandibles and the origin of several bundles of M. tentoriopharyngalis from the well-developed gular ridges. Several features of the larval head are plesiomorphic compared to the cleroid-cucujoid lineage. The interrelationships of most tenebrionoid families not belonging to the pythid-salpingid and anthicid-scraptiid groups were not resolved. Synchroidae were placed as sister group of a clade comprising these two lineages and Prostomidae. A sistergroup relationship between Trictenotomidae and Pythidae seems to be well supported and the monophyly of the anthicid-scraptiid lineage was also confirmed. Another potential clade comprises Prostomidae, Mycteridae and Boridae, and possibly Pyrochroidae (s.str.) and Inopeplinae. The monophyly of Salpingidae (incl. Othniinae and Inopelinae) and Pyrochroidae (incl. Pedilinae) was not supported. Many features such as the shape of the head and body, sutures and ridges of the head capsule, the endocarina, the mandibles, the maxillary apex, and also characters of the terminal abdominal apex are highly variable, even within families. Especially the families Tetratomidae, Melandryidae, Colydiidae and Zopheridae show a high degree of variation in the larval stages. Several taxa appear isolated in terms of larval morphology within the families they are assigned to, e.g. Orchesia within Melandryidae, Sphindocis (Sphindocinae) within Ciidae, Calopus (Calopinae) within Oedemeridae and Penthe (Penthinae) within Tetratomidae. A broader spectrum of characters and a stepwise approach will be needed for a reliable clarification of the relationships within a very complex group like Tenebrionoidea.
Larvae of Rhipsideigma raffrayi are described in detail and those of Distocupes varians are re-examined. Their morphological structures are evaluated with respect to their functional and phylogenetic significance. Larvae of Rhipsideigma are wood-borers with a straight body and a wedge-shaped head capsule. Most of their apomorphic features are correlated with their xylobiontic habits. The strong mandibles, the sclerotized ligula and the wedge-shaped head enable the larvae to penetrate rotting wood. The broadened prothorax, prosternal asperities, tergal ampullae, the short legs, and eversible lobes of segment IX play an important role in locomotion in galleries within rotting wood. Leg muscles are weakly developed, whereas the dorsal, pleural and ventral musculature is complex. The larval features allow Rhipsideigma to be placed in the clades Archostemata, Cupedidae + Micromalthidae, Cupedidae, Cupedidae excl. Priacma, and Cupedidae excl. Priacma and Distocupes. The monophyly of Cupedidae and Cupedidae, excluding Priacma, so far is only supported by apomorphies of the adults. However, the presence of glabrous patches on the prosternum and of a medially divided field of asperities may be larval apomorphies of the family. A clade, which comprises Rhipsideigma, Tenomerga and probably other genera of Cupedidae with hitherto unknown larvae, is well supported by larval apomorphies such as the broadened prothorax, the presence of coxal asperities and the presence of a distinct lateral longitudinal bulge. Increased numbers of antennomeres and labial palpomeres are apomorphies only found in larvae of Distocupes.
Concordant differences in morphology, phenology and RAMS markers, as well as in sequenced mtDNA (COI, COII, cytb) and nuclear DNA (ITS2) fragments, indicate that Dolerus asper Zaddach, 1859 and Dolerus brevicornis Zaddach, 1859 are valid species. On the basis of morphology, molecular markers, and distributional records, both species are distinct from Dolerus gibbosus Hartig, 1837 (= Dolerus planatus Hartig, 1837). Taxonomy of the species is clarified and the neotypes of Dolerus asper Zaddach, 1859 and Dolerus brevicornis Zaddach, 1859 are designated. The synonymies of Dolerus asper Zaddach, 1859, to Dolerus planatus Hartig, 1837 and Dolerus derzavini Malaise, 1931, spec. rev. to D. asper Zaddach, 1859 are abandoned. Dolerus carbonarius Zaddach, 1859 and Dolerus fumosus Zaddach, 1859 are considered to be species inquirendae. Phylogenetic analyses of the ITS2 fragment and fragments of ITS2 + COI and ITS2 + COII yielded the topology [D. asper, (D. brevicornis, D. gibbosus)], while those of all other markers and their combinations resulted in the topology [D. brevicornis, (D. asper, D. gibbosus)]. In the latter hypothesis the clade asper + gibbosus is also supported by structural synapomorphies.
The reservoirs of dorso-abdominal scent glands and the occurrence of the metapleural scent gland evaporatoria in the adults of nine central European and one North American species in the family Rhopalidae (Hemiptera) were studied. All published data about the persistence of the dorso-abdominal scent glands in rhopalid adults are reviewed, and systematic and phylogenetic implications are derived from the patterns of variation.
Ant-like stone beetles (Coleoptera: Scydmaenidae) include more than 4,850 described species in about 90 genera maintained as a separate cosmopolitan family since 1815. Recent authors have hypothesised that Scydmaenidae might be rooted deep inside rove-beetles (Staphylinidae). To test this hypothesis we analysed 206 parsimoniously informative larval and adult morphological characters scored for 38 taxa. Strict consensus topologies from the shortest trees in all 12 analyses consistently placed Scydmaenidae as sister to (Steninae + Euaesthetinae) in a monophyletic Staphylinine Group (with or without Oxyporinae). The single fully resolved and most consistently supported topology maintains a monophyletic Staphylinine Group consisting of Oxyporinae + (Megalopsidiinae + (("Scydmaenidae" + (Steninae + Euaesthetinae)) + (Leptotyphlinae + (Pseudopsinae + (Paederinae + Staphylininae))))); Solierius lacks larval data and is ambiguously placed within the Group. Eight analyses of variably aligned 18S rDNA data for 93 members of Staphylinoidea under parsimony, neighbour-joining and Bayesian approaches were markedly inconsistent, although partly congruent with the Scydmaenidae + (Steninae + Euaesthetinae) hypothesis. Our results strongly suggest that ant-like stone beetles do not form an independent family, but are morphologically modified members of Staphylinidae and, consequently, should be treated as a 32nd recent subfamily within the megadiverse Staphylinidae sensu latissimo. Formal taxonomic acts are: Scydmaeninae Leach, 1815, status novus (= Scydmaenidae Leach, 1815); Scydmaenitae Leach, 1815, status novus (= Scydmaeninae Leach, 1815); Mastigitae Fleming, 1821, status novus (= Mastiginae Fleming, 1821); Hapsomelitae Poinar & Brown, 2004, status novus (= Hapsomelinae Poinar & Brown, 2004). The family Staphylinidae sensu latissimo becomes the largest in Coleoptera and in the whole of the Animal Kingdom, with 55,440 described species (extant plus extinct), thus surpassing Curculionidae with an estimated 51,000 described species.
First instar larvae of Polystoechotidae sp., and first and older instar larvae of Ithone fusca Newman and Oliarces clara Banks are described; those of the latter species for the first time. The family Ithonidae is unique in Neuroptera in having grub-like C-shaped older instar larvae. Potential morphological synapomorphies of mature larvae of Ithonidae and Polystoechotidae are the mandibles with exceptionally broad base and markedly thickened apical part; antennal curvature is fixed and rather characteristic in shape; ocular area reduced or absent; cardo and stipes are markedly enlarged with stipes much larger than the cardo; presence of gula (Polystoechotes) or some traces of gular sclerotisation (Ithone, Oliarces). Larvae of Ithone have numerous larval autapomorphies such as C-shaped first instar larva with reduced abdominal segments IX and X; fused tibia and tarsus on all legs and dorsally directed maxillae. Larvae of Ithonidae and Polystoechotidae have some similarities with those of the family Dilaridae, such as no or one pair of stemmata; body not flattened dorso-laterally; mesothoracic spiracle located on fold between prothorax and mesothorax; short and stout mandibles widened at base and tapered apically; robust and elongated fore legs; tarsi on all legs markedly shortened; more than three larval instars. Older instar larvae of Ithonidae are markedly similar to those of the beetle superfamily Scarabaeoidea in having a C-shaped body, at least in older instars; body round in cross-section; sclerites on thorax and abdomen reduced and body surface membranous; each thoracic and abdominal segment subdivided dorsally into two or three fleshy lobes; ventral surface of abdominal apex bears a field of short and stout setae. Chaetotaxy pattern in first instar Ithonidae and Polystoechotidae larvae suggests that it is possible to homologise the sensilla in different genera and provide a system of sensilla designation for Neuroptera larvae. This study is illustrated with 36 morphological drawings.
A provisional larval groundplan of the family Hygrobiidae is provided through descriptions of internal and external features of three of six extant species, Hygrobia hermanni (Fabricius, 1775), H. wattsi Hendrich 2001 and H. australasiae (Clark, 1862) and phylogenetic interpretations. Hygrobiidae larvae are morphologically differing dramatically from all other known Adephaga by 20 autapomorphies. Structures involved with feeding, i.e., mouthparts, prepharynx and foregut are highly modified as a result of a specialisation on small tubificid worms and chironomid larvae. A placement of Hygrobiidae within Dytiscoidea is well supported by the reduced condition of the terminal abdominal segments, and the presence of 10 ancestral setae on femur and a clade comprising Hygrobiidae, Amphizoidae, and Dytiscidae by the presence of thin and elongate caudal tentorial arms, a very strong musculus verticopharyngalis and a longitudinally divided adductor tendon of the mandible. A highly modified foregut, reduced terminal spiracles VIII and the presence of tubular gills are features which distinguish hygrobiid larvae from those of other groups of Dytiscoidea (i.e, Amphizoidae, Noteridae, Dytiscidae). A sister-group relationship between Hygrobiidae and Dytiscidae is indicated by a distinctly shortened and transverse prepharynx and a cerebrum and suboesophaeal ganglion shifted to the anterior third of the head. Larvae of the Australian species H. wattsi and H. australasiae share the presence of a bluntly rounded mandible and an apical position of the primary pore MNd in instar I as potential synapomorphies.
We present a molecular phylogeny incorporating all five species of the Palaearctic geometrid genus Lythria, based on a 2810-bp combined data matrix comprising the full sequence of the mitochondrial gene cytochrome oxidase subunit one (COI) and fragments of the nuclear genes elongation factor 1 alpha (EF-1α) and wingless (wgl). L. venustata, which was recently rediscovered from Kazakhstan, is shown to be sister taxon to all other members of the genus. The remaining species within the genus form two pairs of sister species: L. purpuraria groups together with L. plumularia, and L. cruentaria with L. sanguinaria. The phylogeny is well supported by characters of the male genitalia of all Lythria species. In addition to the molecular phylogeny of the genus Lythria, we illustrate the external appearance of L. venustata for the first time and describe the anatomy of its male genitalia.