The insect order Trichoptera (caddisflies) forms the second most species‐rich monophyletic group of animals in freshwater. So far, several attempts have been made to elucidate its evolutionary history with both morphological and molecular data. However, none have attempted to analyse the time frame for its diversification. The order is divided into three suborders – Annulipalpia, Integripalpia and ‘Spicipalpia’. Historically, the most problematic taxon to place within the order is ‘Spicipalpia’, whose larvae do not build traditional cases or filtering nets like the majority of the caddisflies. They have previously been proposed to be the sister group of all other Trichoptera or more advanced within the order, with equivocal monophyly and with different interordinal placements among various studies. In order to resolve the evolutionary history of the caddisflies as well as timing their diversification, we utilized fragments of three nuclear (carbamoylphosphate synthethase, isocitrate dehydrogenase and RNA polymerase II) and one mitochondrial (cytochrome oxidase I) protein coding genes, with 16 fossil trichopteran taxa used for time calibration. The ‘spicipalpian’ families are recovered as ancestral to all other caddisflies, though paraphyletic. We recover stable relationships among most families and superfamilies, resolving many previously unrecognized phylogenetic affinities amongst extant families. The origin of Trichoptera is estimated to be around 234 Ma, i.e. Middle – Late Triassic.

Aim To test whether environmental diversification played a role in the diversification of the New Caledonian Hydropsychinae caddisflies.Location New Caledonia, south-west Pacific.Methods The phylogeny of the New Caledonian Hydropsychinae caddisflies was hypothesized using parsimony and Bayesian methods on molecular characters. The Bayesian analysis was the basis for a comparative analysis of the correlation between phylogeny and three environmental factors: geological substrate (ultrabasic, non-ultrabasic), elevation and precipitation. Phylogenetic divergence times were estimated using a relaxed clock method, and environmental factors were mapped onto a lineage-through-time plot to investigate the timing of environmental diversification in relation to species radiation. The correlation between rainfall and elevation was tested using independent contrasts, and the gamma statistic was calculated to infer the diversification pattern of the group. ResultsThe diversification of extant Orthopsyche-Caledopsyche species began in the Middle-Late Oligocene, when much of the island of New Caledonia was covered by ultrabasic substrate and mountain forming was prevalent. Most lineages originated in the Middle-Late Miocene, a period associated with long-term climate oscillation. Optimization of environmental factors on the phylogeny demonstrated that the New Caledonian Hydropsychinae group adapted to ultrabasic substrate early in its evolutionary history. The clade living mostly on ultrabasic substrate was far more species-rich than the clade living mostly on non-ultrabasic substrate. Elevation and rainfall were significantly correlated with each other. The lineagethrough-time plot revealed that the main environmental diversification preceded species diversification. A constant speciation through time was rejected, and the negative gamma indicates that most of the diversification occurred early in the history of the clade. According to the inferred phylogeny, the genus Orthopsyche McFarlane is a synonym under Caledopsyche Kimmins, and Abacaria caledona Oláh & Barnard should also be included in Caledopsyche. Main conclusionsThe age of the radiation does not support a vicariance origin of New Caledonian Hydropsychinae caddisflies. Environmental diversification pre-dates lineage diversification, and thus environmental heterogeneity potentially played a role in the diversification of the group, by providing a variety of fragmented habitats to disperse into, promoting speciation. The negative gamma indicates that the speciation rate slowed as niches started to fill.

Seven new species of Helicopsyche (Feropsyche) Johanson 2002 (Helicopsychidae) are described from Mexico (H. curvipalpia new species), Panama (H. blantoni new species, H. chiriquensis new species, H. linguata new species, and H. sanblasensis new species), and Brazil (H. paprockii new species and H. cipoensis new species) based on adult material borrowed from the National Museum of Natural History (Smithsonian Institution), Washington, D. C. and the Illinois Natural History Survey, Champaign, Illinois, USA. New records are given for H. sinuata Denning & Blickle from Mexico, and H. incisa Ross and H. woldai Johanson from Panama.

BackgroundLeptoceridae are among the three largest families of Trichoptera (caddisflies). The current classification is founded on a phylogenetic work from the 1980's, based on morphological characters from adult males, i.e. wing venation, tibial spur formula and genital morphology. In order to get a new opinion about the relationships within the family, we undertook a molecular study of the family based on sequences from five genes, mitochondrial COI and the four nuclear genes CAD, EF-1α, IDH and POL.ResultsThe resulting phylogenetic hypotheses are more or less congruent with the morphologically based classification, with most genera and tribes recovered as monophyletic, but with some major differences. For monophyly of the two subfamilies Triplectidinae and Leptocerinae, one tribe of each was removed and elevated to subfamily status; however monophyly of some genera and tribes is in question. All clades except Leptocerinae, were stable across different analysis methods.ConclusionsWe elevate the tribes Grumichellini and Leptorussini to subfamily status, Grumichellinae and Leptorussinae, respectively. We also propose the synonymies of Ptochoecetis with Oecetis and Condocerus with Hudsonema.

Empidoidea represent a large and diverse superfamily of true flies, and to date no stable hypothesis on the phylogeny exists. Previous classifications have been based on morphological data and the relationships among several groups are still unknown. Using the mitochondrial genes cytochrome oxidase c subunit I (COI) and cytochrome (Cyt ) and the nuclear genes carbomoylphosphate synthase domain of rudimentary (CAD), elongation factor-1 (EF-1 ) and isocitrate dehydrogenase (IDH) in a Bayesian analysis, we tested the support of higher taxonomic groups within this large superfamily of flies. We re-evaluated previous hypotheses of evolution within the group and present a highly supported phylogenetic hypothesis. Atelestidae, Dolichopodidae, Empididae and Hybotidae were supported as monophyletic families, with Atelestidae as sister group to the remaining Empidoidea. Within the family Hybotidae, Bicellariinae stat.n. formed the sister group to the other subfamilies. The family Ragadidae stat.n. is established to include the subfamily Ragadinae and the new subfamily Iteaphilinae subfam.n.; Ragadidae was sister group to the Empididae. Dolichopodidae was found to form a sister group to Ragadidae plus Empididae. Within Empididae, Hemerodromiinae was found to be a nonmonophyletic group. The tribes Hilarini and Hemerodromiini stat. rev. were recovered as sister groups, as were Empidini and Chelipodini stat. rev. The former family Brachystomatidae was found to be nested within Empididae. A revised classification and diagnoses of nondolichopodid families, subfamilies and tribes are provided.