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A Systematic Investigation of the Crane Fly Subfamily Limoniinae (Tipuloidea: Limoniidae) Matthew Onj Petersen Iowa State University

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A Systematic Investigation of the Crane Fly Subfamily Limoniinae (Tipuloidea: Limoniidae) Matthew Onj Petersen Iowa State University Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 2008 A systematic investigation of the crane fly subfamily Limoniinae (Tipuloidea: Limoniidae) Matthew onJ Petersen Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Entomology Commons Recommended Citation Petersen, Matthew Jon, "A systematic investigation of the crane fly ubfas mily Limoniinae (Tipuloidea: Limoniidae)" (2008). Retrospective Theses and Dissertations. 15829. https://lib.dr.iastate.edu/rtd/15829 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. A systematic investigation of the crane fly subfamily Limoniinae (Tipuloidea: Limoniidae) by Matthew Jon Petersen A dissertation submitted to the graduate faculty in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Co-Majors: Entomology; Ecology and Evolutionary Biology Program of Study Committee: Gregory W. Courtney, Major Professor Bryony C. Bonning Lynn G. Clark Marlin E. Rice Robert S. Wallace Iowa State University Ames, Iowa 2008 Copyright © Matthew Jon Petersen, 2008. All rights reserved. 3316185 3316185 2008 ii TABLE OF CONTENTS CHAPTER 1. DISSERTATION OVERVIEW 1 1.1 Dissertation Organization 4 1.2 References 6 CHAPTER 2. A COMBINED MORPHOLOGICAL AND MOLECULAR PHYLOGENETIC ANALYSIS OF THE TRUE CRANE FLIES (DIPTERA; TIPULOIDEA) 8 2.1 Abstract 8 2.2 Introduction 9 2.3 Methods and Materials 13 2.3.1 Taxon sampling 13 2.3.2 Outgroup choice 14 2.3.3 DNA isolation, amplification, and sequencing 15 2.3.4 Alignment 16 2.3.5 Morphological characters 16 2.3.6 Phylogenetic analyses and tree support 17 2.4 Results 18 2.4.1 Parsimony analysis of morphological data 18 2.4.2 Parsimony analysis of 28S rDNA data 19 2.4.3 Parsimony analysis of combined data 20 2.4.4 Bayesian analyses 21 2.4.5 Influence of separate and combined datasets 22 2.5 Discussion 23 2.5.1 Pediciidae 25 2.5.2 Tipulidae 26 2.5.3 Cylindrotomidae 26 2.5.4 “Limoniidae” 27 2.5.4.1 Eriopterinae 28 2.5.4.2 Limnophilinae 28 2.5.4.3 Chioneinae 29 2.5.4.4 Limoniinae 30 2.5.4.5 Elephantomyinae 30 2.5.4.6 Lipsothrixinae 30 2.5.4.7 Epiphragminae 31 2.5.4.8 Dactylolabinae 31 2.6 Figures 33 2.7 Tables 41 2.8 References 47 2.9 Appendix 53 iii CHAPTER 3. A CLASSIFICATION OF THE LIMONIINI CRANE FLY GENERA (“LIMONIIDAE”: LIMONIINAE) 59 3.1 Introduction 59 3.2 Methods 60 3.3 Results and Discussion 61 3.3.1 “Limoniidae”: Limoniinae: Limoniini 62 3.3.2 Questionable taxonomic units 109 3.4 Tables 110 3.4 References 115 CHAPTER 4. EVALUATING CRANE FLY (DIPTERA; TIPULOIDEA) TAXONOMIC RICHNESS AND COMMUNITY ASSEMBLAGE IN A BIODIVERSITY HOTSPOT 120 4.1 Abstract 120 4.2 Introduction 121 4.3 Methods and Materials 123 4.3.1 Study area 123 4.3.2 Sampling methods 124 4.3.3 Sample processing 125 4.3.4 Data analysis 125 4.4 Results 127 4.4.1 Observed diversity 127 4.4.2 Estimated diversity 128 4.4.3 Collection methods 128 4.4.4 Regional faunas 129 4.4.5 Biogeographic affinities 129 4.5 Discussion 130 4.6 Figures 135 4.7 Tables 140 4.8 References 145 CHAPTER 5. SYSTEMATICS OF THE CRANE FLY GENUS LIPSOTHRIX LOEW (DIPTERA; TIPULOIDEA) 150 5.1 Abstract 150 5.2 Introduction 151 5.2.1 Taxonomic history 152 5.2.2 Taxonomic placement 152 5.3 Methods and Materials 153 5.3.1 Specimens 153 5.3.2 Measurements 154 5.3.3 Genus and species concept 155 5.3.4 Species delineation 156 5.3.5 Species emergence 158 5.3.6 Species distributions 158 5.3.7 Phylogenetic analysis 159 iv 5.4 Results and Discussion 160 5.4.1 Diagnosis of the genus Lipsothrix 161 5.4.2 Redescription of valid species 163 5.4.3 Species of uncertainty 247 5.4.4 Natural history 253 5.4.4.1 Larvae 253 5.4.4.2 Pupae 253 5.4.4.3 Adult 256 5.4.5 Phylogenetic analyses 257 5.4.6 Biogeography 259 5.5 Conclusions 264 5.6 Figures 266 5.7 Tables 395 5.8 References 300 5.9 Appendix 308 CHAPTER 6. GENERAL CONCLUSIONS 311 1 CHAPTER 1: DISSERTATION OVERVIEW Systematics provides the knowledge base from which all biology is based, and the subdivisions of this science provide many fundamental methodological criteria by which we investigate the natural world. Far from simply a service science that provides a reference system for study by other disciplines (Ebach & Holdrege 2005), systematics is a dynamic science that involves many facets such as the discovery and delineation of species and other taxa (taxonomy), defining current and historical distributional patterns (biogeography), and the study of evolutionary patterns (phylogenetics). Systematics allows researchers to answer the questions of what and how many organisms inhabit the Earth (Agnarsson & Kuntner 2007). The importance of basic taxonomic research is now greater than ever, as deterioration of taxonomic expertise coupled with an increased rate of species extinction, the so called “taxonomic impediment” (Rodman & Cody 2003) or “taxonomic crisis” (Taylor 1976) threatens to eliminate diversity even before it is identified. For the study of insects this is particularly true as taxonomic revisions occur on average once every hundred years (Wheeler, 2004). The impact of a depleting knowledge base has the power to affect multiple aspects of related biological disciplines such as ecological studies (Gotelli 2004) and conservation planning (Golding & Timberlake 2003; Godfrey & Knapp 2004). In the absence of taxonomy, the language by which we interpret and identify life disappears. The acknowledgement of the importance of taxonomic research has led to a recent resurgence of basic taxonomic work that incorporates traditional aspects of systematic investigations, such as the study of morphology, species descriptions, and classification, with new technologies of molecular biology (DNA taxonomy) and electronic resources (“new 2 taxonomy” Wheeler, 2008). With this resurgence comes the opportunity to rapidly disseminate taxonomic resources and world species catalogs, thus greatly decreasing the time spent researching records and searching for taxonomic tools. Within the Diptera (Insecta) the general descriptor of crane fly has been used for the Ptychopteridae (phantom crane flies), Trichoceridae (winter crane flies), Tanyderidae (primitive crane flies), and the Tipuloidea (true crane flies), having been loosely applied to those groups of flies that show a tendency for the elongation of the legs, wings, and abdomen, often in a delicate form. Since the advent of modern cladistic approaches (Henning 1966) and subsequent phylogenetic analyses of the Lower Diptera (Wood & Borkent 1989; Griffith 1990; Oosterbroek & Courtney 1995), a consensus has been reached that the term crane fly in fact represents a paraphyletic or polyphyletic group of independently derived lineages that have convergence on this delicate body form. Of the groups that have had this descriptor attached, the true crane flies of the superfamily Tipuloidea represent the largest and most diverse assemblage of Diptera. The 15,000 described crane fly species account for over 10% of all fly diversity and are placed in 700 genera and subgenera (Oosterbroek 2008). The chapters included in this dissertation examine the systematics of the true crane flies of the superfamily Tipuloidea. This large and diverse group is taxonomically rich in terms of described species, but suffers from an inadequate classification and lack of available and complete taxonomic tools. Although the knowledge of species identity is broad and not limited to any particular region, the effect of inadequate taxonomic tools is largely felt in areas outside of the Nearctic and Western Palearctic where they are limited or often nonexistent. As a result, the biogeographic, phylogenetic, and ecological knowledge is 3 lacking, especially in those groups and regions where basic taxonomy has been insufficiently studied. Some basic biological information can be learned from the few studies on widespread taxa in North America and Europe. The developmental histories of the majority of crane flies are quite similar, with most of the lifecycle spent in the larval stage. Feeding takes place in the larval stage where a variety of feeding types occur, including predation, detritivory, herbivory, and omnivory (Pritchard 1983). Adult feeding is uncommon and rare. While most of the life span is spent in the larval life stage, this is where the least is known. The larval preferences of crane flies are generally thought to be either aquatic, semi-aquatic, or terrestrial (Young & Gelhaus 2000), however the great majority of species occupy habitats that are neither aquatic nor terrestrial but are areas that are either an interface between the two, or occupy both areas in different times of immature development. Examples of intermediate aquatic / terrestrial habitats include algae growing in splash zones, saturated dead wood in aquatic areas, partially saturated soils, and partially aquatic mosses. Species that occupy by terrestrial and aquatic habitats will occupy aquatic habitats in the larval stage but move to terrestrial areas to pupate. While these examples illustrate the diversity of habitats utilized by crane fly larvae, our knowledge of larval behavior and habitat preference is still in its infancy, with life stage associations made for less than 4% of the world species and 16% of the genera and subgenera. The crane fly adult, more specifically restricted to the adult male, is in almost every case the life stage in which the knowledge of species identity is maintained.
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