BackgroundIn the mega-diverse insect order Lepidoptera (butterflies and moths; 165,000 described species), deeper relationships are little understood within the clade Ditrysia, to which 98% of the species belong. To begin addressing this problem, we tested the ability of five protein-coding nuclear genes (6.7 kb total), and character subsets therein, to resolve relationships among 123 species representing 27 (of 33) superfamilies and 55 (of 100) families of Ditrysia under maximum likelihood analysis.ResultsOur trees show broad concordance with previous morphological hypotheses of ditrysian phylogeny, although most relationships among superfamilies are weakly supported. There are also notable surprises, such as a consistently closer relationship of Pyraloidea than of butterflies to most Macrolepidoptera. Monophyly is significantly rejected by one or more character sets for the putative clades Macrolepidoptera as currently defined (P < 0.05) and Macrolepidoptera excluding Noctuoidea and Bombycoidea sensu lato (P ≤ 0.005), and nearly so for the superfamily Drepanoidea as currently defined (P < 0.08). Superfamilies are typically recovered or nearly so, but usually without strong support. Relationships within superfamilies and families, however, are often robustly resolved. We provide some of the first strong molecular evidence on deeper splits within Pyraloidea, Tortricoidea, Geometroidea, Noctuoidea and others.Separate analyses of mostly synonymous versus non-synonymous character sets revealed notable differences (though not strong conflict), including a marked influence of compositional heterogeneity on apparent signal in the third codon position (nt3). As available model partitioning methods cannot correct for this variation, we assessed overall phylogeny resolution through separate examination of trees from each character set. Exploration of "tree space" with GARLI, using grid computing, showed that hundreds of searches are typically needed to find the best-feasible phylogeny estimate for these data.ConclusionOur results (a) corroborate the broad outlines of the current working phylogenetic hypothesis for Ditrysia, (b) demonstrate that some prominent features of that hypothesis, including the position of the butterflies, need revision, and (c) resolve the majority of family and subfamily relationships within superfamilies as thus far sampled. Much further gene and taxon sampling will be needed, however, to strongly resolve individual deeper nodes.

Major progress has been made recently toward resolving the phylogeny of Noctuoidea, the largest superfamily of Lepidoptera. However, numerous questions and weakly supported nodes remain. In this paper we independently check and extend the main findings of multiple recent authors by performing maximum-likelihood analyses of 5-19 genes (6.7-18.6 kb) in 74 noctuoids representing all the families and a majority of the subfamilies. Our results strongly support the six family system of Zahiri et al., with the former Lymantriidae and Arctiidae subsumed within the huge family Erebidae, and Noctuidae restricted largely to the subfamilies with so-called trifine hindwing venation. Our data also strongly corroborate monophyly of the set of four families with quadrifid forewing venation, to the exclusion of Notodontidae, and removal from the latter of Oenosandridae. Other among-family relationships, however, remain unsettled. Our evidence is equivocal on the position of Oenosandridae, which are sister group to either Notodontidae alone or to all other noctuoids. Like other recent nuclear gene studies, our results also provide no strong support for relationships among the four quadrifid forewing families. In contrast, within families our analyses significantly expand the list of robustly resolved relationships, while introducing no strong conflicts with previous molecular studies. Within Notodontidae, for which we present the largest molecular taxon sample to date, we find strong evidence for polyphyly for some, or all, recent definitions of the subfamilies Thaumetopoeinae, Pygaerinae, Notodontinae and Heterocampinae. Deeper divergences are incompletely resolved but there is strong support for multiple 'backbone' nodes subtending most of the subfamilies studied. Within Erebidae, we find much agreement and no strong conflict with a recent previous study regarding relationships among subfamilies, and somewhat stronger support. Although many questions remain, the two studies together firmly resolve positions for over half the subfamilies. Within Noctuidae, we find no strong conflict with previous molecular studies regarding relationships among subfamilies, but much stronger resolution along the 'backbone' of the phylogeny. Combining information from multiple studies yields strongly resolved positions for most of the subfamilies. Finally, our results strongly suggest that the tribes Pseudeustrotiini and Prodeniini, currently assigned to the largest subfamily, Noctuinae, do not belong there. In sum, our results provide additional corroboration for the main outlines of family-level phylogeny in Noctuoidea, and contribute toward resolving relationships within families.

Six new species of the genus Distatrix Mason from Central and South America, D. loretta, D. xanadon, D. vigilis, D. pitillaensis, D. pandora Grinter, n. sp., and D. antirrheae Whitfield & Grinter, n. sp., are described from large-scale caterpillar inventory endeavors, mostly from the larvae of geometrid moths. Biological information and diagnostic features that distinguish these species from other previously described Distatrix, especially those from the Neotropical region, are provided; and the first key to New World species is presented. The new discoveries expand our knowledge of the World's widespread Distatrix fauna by about a third, suggesting that similar survey efforts in other poorly sampled regions will reveal numerous additional undescribed species.

Two new Costa Rican species of the braconid parasitoid wasp subfamily Cardiochilinae, Heteropteron kidonoi Dabek &amp; Whitfield and Heteropteron hasegawai Dabek &amp; Whitfield, are described and illustrated from dry forest in the Area de Conservacion Guanacastae, along with data on rearing from their hosts. Heteropteron kidonoi is a solitary endoparasitoid of Stenoma cathosiota (Lepidoptera: Depressariidae) on Roupala montana (Proteaceae), while H. hasegawai is a solitary endoparasitoid of Carthara abrupta (Lepidoptera: Pyralidae) on the same host plant, but typically at slightly higher elevation localities. Diagnostic characters are provided to distinguish these two new species from each other, and also from the three previously decsribed species of Heteropteron. Heteropteron kidonoi and H. hasegawai are the first species of Heteropteron to have any host data, and also are the first to be reported in Costa Rica.

The descriptive taxonomic study reported here is focused on Glyptapanteles, a species-rich genus of hymenopteran parasitoid wasps. The species were found within the framework of two independent long-term Neotropical caterpillar rearing projects: northwestern Costa Rica (Área de Conservación Guanacaste, ACG) and eastern Andes, Ecuador (centered on Yanayacu Biological Station, YBS). One hundred thirty-six new species of Glyptapanteles Ashmead are described and all of them are authored by Arias-Penna. None of them was recorded in both countries; thus, 78 are from Costa Rica and the remaining 58 from Ecuador. Before this revision, the number of Neotropical described Glyptapanteles did not reach double digits. Reasonable boundaries among species were generated by integrating three datasets: Cytochrome Oxidase I (COI) gene sequencing data, natural history (host records), and external morphological characters. Each species description is accompanied by images and known geographical distribution. Characteristics such as shape, ornamentation, and location of spun Glyptapanteles cocoons were imaged as well. Host-parasitoid associations and food plants are also here published for the first time. A total of 88 species within 84 genera in 15 Lepidoptera families was encountered as hosts in the field. With respect to food plants, these wild-caught parasitized caterpillars were reared on leaves of 147 species within 118 genera in 60 families. The majority of Glyptapanteles species appeared to be relatively specialized on one family of Lepidoptera or even on some much lower level of taxonomic refinement. Those herbivores in turn are highly food-plant specialized, and once caterpillars were collected, early instars (1–3) yielded more parasitoids than later instars. Glyptapanteles jimmilleri Arias-Penna, sp. nov. is the first egg-larval parasitoid recorded within the genus, though there may be many more since such natural history requires a more focused collection of eggs. The rate of hyperparasitoidism within the genus was approximately 4% and was represented by Mesochorus spp. (Ichneumonidae). A single case of multiparasitoidism was reported, Copidosoma floridanum Ashmead (Encyrtidae) and Glyptapanteles ilarisaaksjarvi Arias-Penna, sp. nov. both parasitoid species emerged from the caterpillar of Noctuidae: Condica cupienta (Cramer). Bodyguard behavior was observed in two Glyptapanteles species: G. howelldalyi Arias-Penna, sp. nov. and G. paulhansoni Arias-Penna, sp. nov. A dichotomous key for all the new species is provided. The numerous species described here, and an equal number already reared but not formally described, signal a far greater Glyptapanteles species richness in the Neotropics than suggested by the few described previously.