Brigham Young University BYU ScholarsArchive Theses and Dissertations 2004-03-18 Phylogeny of the Polyneopterous Insects With Emphasis on Plecoptera: Molecular and Morpological Evidence Matthew Dana Terry Brigham Young University - Provo Follow this and additional works at: https://scholarsarchive.byu.edu/etd Part of the Biology Commons BYU ScholarsArchive Citation Terry, Matthew Dana, "Phylogeny of the Polyneopterous Insects With Emphasis on Plecoptera: Molecular and Morpological Evidence" (2004). Theses and Dissertations. 1134. https://scholarsarchive.byu.edu/etd/1134 This Dissertation is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Phylogeny of the Polyneopterous Insects With Emphasis on Plecoptera: Molecular and Morpological Evidence by Matthew D. Terry A dissertation submitted to the faculty of Brigham Young University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Integrative Biology Brigham Young University December 2003 GRADUATE COMMITTEE APPROVAL of a dissertation submitted by Matthew D. Terry This dissertation has been read by each member of the following graduate committee and by majority vote has been found to be satisfactory. ______________ __________________________________________ Date Michael F. Whiting, Chair ______________ __________________________________________ Date Keith A. Crandall ______________ __________________________________________ Date Richard W. Baumann ______________ __________________________________________ Date C. Riley Nelson ______________ __________________________________________ Date Joseph V. McHugh BRIGHAM YOUNG UNIVERSITY As chair of the candidate’s graduate committee, I have read the dissertation of Matthew D. Terry in its final form and have found that (1) its format, citations, and bibliographical style are consistent and acceptable and fulfill university and department style require- ments; (2) its illustrative materials including figures, tables, and charts are in place; and (3) the final manuscript is satisfactory to the graduate committee and is ready for submission to the university library. ___________________ __________________________________________ Date Michael F. Whiting Chair, Graduate Committee Accepted for the Department __________________________________________ Keith A. Crandall Graduate Coordinator Accepted for the College __________________________________________ R. Kent Crookston Dean, College of Integrative Biology ABSTRACT PHYLOGENY OF THE POLYNEOPTEROUS INSECTS WITH EMPHASIS ON PLECOPTERA: MOLECULAR AND MORPHOLOGICAL EVIDENCE Matthew D. Terry Department of Integrative Biology Doctor of Philosophy Polyneoptera is an assemblage of eleven insect orders comprising the “orthopteroid” insects. It includes familiar insects such as grasshoppers, roaches, termites, earwigs and preying mantises; as well as the more obscure web-spinners, angel insects and ice-crawlers. We present a phylogenetic analysis of the polyneopteran orders based on 18S rDNA, 28S rDNA, Histone 3, and a coded morphology matrix for an extensive sampling of taxa. We investigate the use of congruence between separate datasets as an a priori measure of alignment quality. Our results support the paraphyly of Polyneoptera, the monophyly of Dictyoptera, sister taxon relationships between Embiidina + Phasmatodea and Dermaptera + Zoraptera, and a relatively basal placement of Plecoptera. The analyses also support a sister taxon relationship between the newly described Mantophasmatodea and Grylloblattodea, a small order of cryophilic insects confined to the northwestern Americas and northeastern Asia. This placement coupled with the morphological disparity of the two groups validates the creation of a new order for Mantophasmatodea. Our results also suggest the Direct Optimization (formerly Optimization Alignment) produces alignments that are more predictable across the parameter landscape than alignment via CLUSTAL X, as measured by congruence among independent data partitions. Dense taxon sampling and phylogenetic analysis of six molecular markers (12S, 16S, 18S, 28S, COII, and H3) and morphological data for the order Plecoptera demonstrates that the subordinal groups Arctoperlaria and Antarctoperlaria are monophyletic. Euholognatha and Systellognatha are also monophyletic, with the exception of the genus Megaleuctra which is the basal lineage for the order and deserves recognition as a distinct family (Megaleuctridae). Notonemouridae is strongly supported as a monophyletic clade. Within the Systellognatha Styloperlidae is the basal lineage, followed by Peltoperlidae then Pteronarcyidae, and Perloidea is a strongly supported monophyletic group with Chloroperlidae as sister taxon to Perlidae + Perlodidae. The family Gripopterygidae is strongly supported as paraphyletic. Many Plecoptera (stoneflies) exhibit a pre-mating communication known as “drumming.” Species of the genus Isogenoides have complex drumming behavior in which (i) the male calls the female by tapping his abdomen against the substrate, (ii) the female answers with her own distinctive tapping, and (iii) the male responds with a confirmatory series of taps. These drumming patterns are specific to individual species and may vary within a species to form distinct dialects. Phylogenetic analysis for the genus based on six molecular markers (12S, 16S, 18S, 28S, COII, and H3) supports Yugus as its nearest extant relative and I. hansoni as the basal lineage within the genus. Drumming behavioral characters appear to be largely incongruent with the phylogeny. TABLE OF CONTENTS Title Page . i. Graduate Committee Approval . ii. Final Reading Approval . iii. Abstract . .iv-vi. Table of Contents . .vii. Chapter 1. Mantophasmatodea and Phylogeny of the Lower Neopterous Insects . .1-35. Chapter 2. Comparison of Two Alignment Techniques within a Single Data Set: POY v. CLUSTAL. 36-56. Chapter 3. Phylogenetic Systematics of Plecoptera: Evidence from Morphology and Six Genes . .57-91. Chapter 4. Phylogeny of the Genus Isogenoides (Plecoptera: Perlodidae) and the Evolution of Drumming Behavior . 91-118. vii. Mantophasmatodea and Phylogeny of the Lower Neopterous Insects Matthew D. Terry1,2 and Michael F. Whiting1 1Department of Integrative Biology, Brigham Young University, Provo, UT 84602, USA 2Present Address: Department of Molecular and Cellular Biology, University of Arizona, Life Sciences South Building, Room 444, 1007 E. Lowell Street, Tucson, AZ 85721 Corresponding Author: Matthew D. Terry 1007 E. Lowell St. University of Arizona Tucson, AZ 85721 (520) 422-2326 Fax: (520) 621-3709 email: [email protected] 1 Abstract. —Polyneoptera is a name sometimes applied to an assemblage of 11 insect orders comprising the lower neopterous or “orthopteroid” insects. These orders include familiar insects such as Orthoptera (grasshoppers), Blattodea (roaches), Isoptera (termites), (Mantodea) preying mantises, Dermaptera (earwigs), Phasmatodea (stick insects), Plecoptera (stoneflies); as well as the more obscure, Embiidina (web- spinners), Zoraptera (angel insects) and Grylloblattodea (ice-crawlers). Many of these insect orders exhibit a high degree of morphological specialization, a condition that has lead to multiple phylogenetic hypotheses and little consensus among investigators. We present a phylogenetic analysis of the polyneopteran orders, including the recently described Mantophasmatodea, based on 18S rDNA, 28S rDNA, Histone 3, and a coded morphology matrix for an extensive sampling of taxa. Extensive analyses utilizing different alignment methodologies and parameters values across a majority of possible ranges were employed to test for sensitivity of results to ribosomal alignment and to explore patterns across the theoretical alignment landscape. Our results support the paraphyly of Polyneoptera, the monophyly of Dictyoptera, sister taxon relationships between Embiidina + Phasmatodea and Dermaptera + Zoraptera, and a relatively basal placement of Plecoptera. The analyses also support a sister taxon relationship between the newly described Mantophasmatodea, which are endemic to arid portions of southern Africa, and Grylloblattodea, a small order of cryophilic insects confined to the northwestern Americas and northeastern Asia. This placement coupled with the morphological disparity of the two groups validates the creation of a new order for Mantophasmatodea. 2 All neopterous insects, those that can fold their wings, can be placed into one of three groups: Holometabola, insects that have complete metamorphosis; Paraneoptera, true bugs and their allies; or a basal assemblage sometimes called “Polyneoptera” or the “orthopteroid” insects (for the sake of clarity the term Polyneoptera will be used hereafter when referring to these orders collectively). There has been extensive discussion among entomologists regarding the monophyly of the polyneopteran lineages, a group that includes Blattodea (roaches), Isoptera (termites), Mantodea (preying mantises), Dermaptera (earwigs), Embiidina (web- spinners), Grylloblattodea (ice crawlers), Orthoptera (crickets and grasshoppers), Phasmatodea (stick insects), Plecoptera (stoneflies), Zoraptera (angel insects), and the recently described Mantophasmatodea (gladiators). These orders represent one of the largest and earliest insect radiations,