Summary The phylogeny of carabid tribes is examined with sequences of 18S ribosomal DNA from eighty‐four carabids representing forty‐seven tribes, and fifteen outgroup taxa. Parsimony, distance and maximum likelihood methods are used to infer the phylogeny. Although many clades established with morphological evidence are present in all analyses, many of the basal relationships in carabids vary from analysis to analysis. These deeper relationships are also sensitive to variation in the sequence alignment under different alignment conditions. There is moderate evidence against the monophyly of Migadopini + Amarotypini, Scaritini + Clivinini, Bembidiini and Brachinini. Psydrini are not monophyletic, and consist of three distinct lineages (Psydrus, Laccocenus and a group of austral psydrines, from the Southern Hemisphere consisting of all the subtribes excluding Psydrina). The austral psydrines are related to Harpalinae plus Brachinini. The placements of many lineages, including Gehringia, Apotomus, Omophron, Psydrus and Cymbionotum, are unclear from these data. One unexpected placement, suggested with moderate support, is Loricera as the sister group to Amarotypus. Trechitae plus Patrobini form a monophyletic group. Brachinini probably form the sister group to Harpalinae, with the latter containing Pseudomorpha, Morion and Cnemalobus. The most surprising, well supported result is the placement of four lineages (Cicindelinae, Rhysodinae, Paussinae and Scaritini) as near relatives of Harpalinae + Brachinini. Because these four lineages all have divergent 18S rDNA, and thus have long basal branches, parametric bootstrapping was conducted to determine if their association and placement could be the result of long branch attraction. Simulations on model trees indicate that, although their observed association might be due to long branch attraction, there was no evidence that their placement near Harpalinae could be so explained. These simulations also suggest that 18S rDNA might not be sufficient to infer basal carabid relationships.

Axis patterning and appendage development have been well studied in Drosophila melanogaster, a species in which both limb and segment morphogenesis are derived. In Drosophila, positional information from genes important in anteroposterior and dorsoventral axis formation, including wingless (wg) and decapentaplegic (dpp), is required for allocating and patterning the appendage primordia. We used RNA interference to characterize the functions of wg and dpp in the red flour beetle, Tribolium castaneum, which retains more ancestral modes of limb and segment morphogenesis. We also characterized the expression of potential targets of the WG and DPP signaling pathways in these embryos. Tribolium embryos in which dpp had been downregulated had defects in the dorsalmost body wall, but did not appear to have been globally repatterned and had normal appendages. Downregulation of wg led to the loss of segment boundaries, gnathal and thoracic appendages, and lateral head lobes, and to changes in the expression of dpp, Distal-less, and Engrailed. The functions of wg varied along both the anteroposterior and dorsoventral axes of the embryo. Phylogenetic comparisons indicate that the role of WNT signaling in segment boundary formation is evolutionarily old, but that its role in appendage allocation originated in the common ancestor of holometabolous insects.

The beetle suborder Adephaga is traditionally divided into two sections on the basis of habitat, terrestrial Geadephaga and aquatic Hydradephaga. Monophyly of both groups is uncertain, and the relationship of the two groups has implications for inferring habitat transitions within Adephaga. Here we examine phylogenetic relationships of these groups using evidence provided by DNA sequences from all four suborders of beetles, including 60 species of Adephaga, four Archostemata, three Myxophaga, and ten Polyphaga. We studied 18S ribosomal DNA and 28S ribosomal DNA, aligned with consideration of secondary structure, as well as the nuclear protein-coding gene wingless. Independent and combined Bayesian, likelihood, and parsimony analyses of all three genes supported placement of Trachypachidae in a monophyletic Geadephaga, although for analyses of 28S rDNA and some parsimony analyses only if Coleoptera is constrained to be monophyletic. Most analyses showed limited support for the monophyly of Hydradephaga. Outside of Adephaga, there is support from the ribosomal genes for a sister group relationship between Adephaga and Polyphaga. Within the small number of sampled Polyphaga, analyses of 18S rDNA, wingless, and the combined matrix supports monophyly of Polyphaga exclusive of Scirtoidea. Unconstrained analyses of the evolution of habitat suggest that Adephaga was ancestrally aquatic with one transition to terrestrial. However, in analyses constrained to disallow changes from aquatic to terrestrial habitat, the phylogenies imply two origins of aquatic habit within Adephaga.

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