Inventory of the caterpillars, their food plants and parasitoids began in 1978 for today's Area de Conservacion Guanacaste (ACG), in northwestern Costa Rica. This complex mosaic of 120 000 ha of conserved and regenerating dry, cloud and rain forest over 0-2000 m elevation contains at least 10 000 species of non-leaf-mining caterpillars used by more than 5000 species of parasitoids. Several hundred thousand specimens of ACG-reared adult Lepidoptera and parasitoids have been intensively and extensively studied morphologically by many taxonomists, including most of the co-authors. DNA barcoding -the use of a standardized short mitochondrial DNA sequence to identify specimens and flush out undisclosed species -was added to the taxonomic identification process in 2003.
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.
Epstein, Marc E. Revision and Phylogeny of the Limacodid-Group Families, with Evolutionary Studies on Slug Caterpillars (Lepidoptera: Zygaenoidea). Smithsonian Contributions to Zoology, number 582, 102 pages, 409 figures, 7 tables, 1996.-The limacodid group, composed of Limacodidae (+ Chrysopolomidae), Dalceridae, Megalopygidae, Aididae, and Somabrachyidae, is revised, and diagnoses, redescriptions, and keys to larval and adult stages are provided for each family. Comparative morphology and ontogeny of the larval stage are emphasized. Immature stages, including first instars, of a number of taxa are figured for the first time. New character data for adults and immatures are presented, including the presence of larval crochets on abdominal segments A2-A7 and A10 in species of Megalopygidae and Limacodidae. Computer-assisted phylogenetic analyses of limacodid-group families using separate character-data sets from adult and immature stages produces congruent cladograms of equal parsimony, with Zygaenidae selected as the outgroup. Megalopygidae (Megalopyginae + Trosiinae) is found to be the most basal family among the limacodid group, and Aididae, previously a subfamily of Megalopygidae, is the sister group to the Limacodidae + Dalceridae clade. Chrysopolomidae is placed as a subfamily of Limacodidae based on new character evidence from adult and immature stages. Relationships of Somabrachyidae, Epipyropidae, and Cyclotornidae also are discussed. Previous interpretations of character homologies for abdominal proiegs in the larvae are revised based on the phylogenetic analysis, resulting in a novel hypothesis concerning the evolutionary transformation from proiegs to a slug-like ventral surface. New representations of homology for larval body setae and for juxta and valva of male genitalia also are given. Behavioral observations of immatures and adults of species in the limacodid group are presented, along with a review of their biology. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).
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