DOI: 10.5061/dryad.772
title: Data from: The Strepsiptera Problem: Phylogeny of the Holometabolous Insect Orders Inferred from 18S and 28S Ribosomal DNA Sequences and Morphology

description: Phylogenetic relationships among the holometabolous insect orders were
 inferred from cladistic analysis of nucleotide sequences of 18S ribosomal
 DNA (rDNA) (85 exemplars) and 28S rDNA (52 exemplars) and morphological
 characters. Exemplar outgroup taxa were Collembola (1 sequence),
 Archaeognatha (1), Ephemerida (1), Odonata (2), Plecoptera (2), Blattodea
 (1), Mantodea (1), Dermaptera (1), Orthoptera (1), Phasmatodea (1),
 Embioptera (1), Psocoptera (1), Phthiraptera (1), Hemiptera (4), and
 Thysanoptera (1). Exemplar ingroup taxa were Coleoptera: Archostemata (1),
 Adephaga (2), and Polyphaga (7); Megaloptera (1); Raphidioptera (1);
 Neuroptera (sensu stricto ;eq Planipennia): Mantispoidea (2),
 Hemerobioidea (2), and Myrmeleontoidea (2); Hymenoptera: Symphyta (4) and
 Apocrita (19); Trichoptera: Hydropsychoidea (1) and Limnephiloidea (2);
 Lepidoptera: Ditrysia (3); Siphonaptera: Pulicoidea (1) and
 Ceratophylloidea (2); Mecoptera: Meropeidae (1), Boreidae (1), Panorpidae
 (1), and Bittacidae (2); Diptera: Nematocera (1), Brachycera (2), and
 Cyclorrhapha (1); and Strepsiptera: Corioxenidae (1), Myrmecolacidae (1),
 Elenchidae (1), and Stylopidae (3). We analyzed ~1 kilobase of 18S rDNA,
 starting 398 nucleotides downstream of the 5' end, and ~400 bp of 28S
 rDNA in expansion segment D3. Multiple alignment of the 18S and 28S
 sequences resulted in 1,116 nucleotide positions with 24 insert regions
 and 398 positions with 14 insert regions, respectively. All Strepsiptera
 and Neuroptera have large insert regions in 18S and 28S. The secondary
 structure of 18S insert 23 is composed of long stems that are GC rich in
 the basal Strepsiptera and AT rich in the more derived Strepsiptera. A
 matrix of 176 morphological characters was analyzed for holometabolous
 orders. Incongruence length difference tests indicate that the 28S +
 morphological data sets are incongruent but that 28S + 18S, 18S +
 morphology, and 28S + 18S + morphology fail to reject the hypothesis of
 congruence. Phylogenetic trees were generated by parsimony analysis, and
 clade robustness was evaluated by branch length, Bremer support,
 percentage of extra steps required to force paraphyly, and sensitivity
 analysis using the following parameters: gap weights, morphological
 character weights, methods of data set combination, removal of key taxa,
 and alignment region. The following are monophyletic under most or all
 combinations of parameter values: Holometabola, Polyphaga, Megaloptera +
 Raphidioptera, Neuroptera, Hymenoptera, Trichoptera, Lepidoptera,
 Amphiesmenoptera (Trichoptera + Lepidoptera), Siphonaptera, Siphonaptera +
 Mecoptera, Strepsiptera, Diptera, and Strepsiptera + Diptera (Halteria).
 Antliophora (Mecoptera + Diptera + Siphonaptera + Strepsiptera),
 Mecopterida (Antliophora + Amphiesmenoptera), and Hymenoptera +
 Mecopterida are supported in the majority of total evidence analyses.
 Mecoptera may be paraphyletic because Boreus is often placed as sister
 group to the fleas; hence, Siphonaptera may be subordinate within
 Mecoptera. The 18S sequences for Priacma (Coleoptera: Archostemata),
 Colpocaccus (Coleoptera: Adephaga), Agulla (Raphidioptera), and Corydalus
 (Megaloptera) are nearly identical, and Neuropterida are monophyletic only
 when those two beetle sequences are removed from the analysis. Coleoptera
 are therefore paraphyletic under almost all combinations of parameter
 values. Halteria and Amphiesmenoptera have high Bremer support values and
 long branch lengths. The data do not support placement of Strepsiptera
 outside of Holometabola nor as sister group to Coleoptera. We reject the
 notion that the monophyly of Halteria is due to long branch attraction
 because Strepsiptera and Diptera do not have the longest branches and
 there is phylogenetic congruence between molecules, across the entire
 parameter space, and between morphological and molecular data.
description: Whiting Data
 Matrixwhiting.txt