D. Elmo Hardy Memorial Volume Contributions to the Systematics and Evolution of Diptera NEAL L. EVENHUIS is a research entomologist and Chairman of the Department of Natural Sciences, Bishop Museum, Honolulu, Hawai‘i KENNETH Y. KANESHIRO is Director of the Center for Conservation Research and Training at, the University of Hawaii at Mänoa, Hawai‘i D. Elmo Hardy Memorial Volume Contributions to the Systematics and Evolution of Diptera Edited by Neal L. Evenhuis & Kenneth Y. Kaneshiro Bishop Museum Bulletin in Entomology 12 Bishop Museum Press Honolulu, 2004 Published by Bishop Museum Press 1525 Bernice Street Honolulu, Hawai‘i 96817-2704, USA Copyright ©2004 Bishop Museum All Rights Reserved Printed in the United States of America ISBN 1-58178-022-2 ISSN 0893-3146 Evenhuis & Kaneshiro — D. Elmo Hardy Memorial Volume v TABLE OF CONTENTS Preface — Evenhuis, N.L. & Kaneshiro, K.Y. ............................................................................... vii Biography of D. Elmo Hardy (1914–2002) — Evenhuis, N.L. ....................................................... 1 Pipunculidae Research by Elmo Hardy: Another Founding Event on the Hawaiian Islands — Skevington, J.H. & De Meyer, M. .................................................................................... 13 Alishania, a New Genus with Remarkable Female Terminalia from Taiwan, with Notes on Chrysotimus Loew (Diptera: Dolichopodidae) — Bickel, D.J. ............................................ 27 Humongochela, a New Genus of Waterfall-Loving Flies from the Marquesas Islands (Diptera: Dolichopodidae) — Evenhuis, N.L. ....................................................................................... 35 Sex, Flies, and the Genetic Consequences of Small Population Size — Kaneshiro, K.Y. ........... 43 Host Alternation by Gall Midges of the Genus Asphondylia (Diptera: Cecidomyiidae) — Uechi, N., Yukawa, J. & Yamaguchi, D. ........................................................................... 53 New Species and New Records of Tephritidae (Diptera) from New Caledonia — Norrbom, A.L. & Hancock, D.L. ............................................................................................................ 67 Bibionidae (Diptera) of New Caledonia — Fitzgerald, S.J. ........................................................ 79 A Revision of the Shore-fly Genus Trimerogastra Hendel (Diptera: Ephydridae) — Mathis, W.N. & Zatwarnicki, T. ......................................................................................................... 89 Phylogeny and Population Genetics of Hawaiian Drosophilids: Impact on Evolutionary Biology — Carson, H.L. .................................................................................................... 109 Phylogenetic Relationships Within the Drosophila haleakalae Species Group Inferred by Molecular and Morphological Characters (Diptera: Drosophilidae) — O’Grady, P.M. & Zilversmit, M. ....................................................................................................................... 117 Insects Associated with Fruits of the Oleaceae (Asteridae, Lamiales) in Kenya, with Special Reference to the Tephritidae (Diptera) — Copeland, R.S, White, I.M., Olumu, M., Machera, P. & Wharton, R.A. .............................................................................................. 135 Biogeography and Speciation in the Dacini (Diptera: Tephritidae: Dacinae) — Drew, R.A.I. .... 165 Bibliography of and Taxa Described by D. Elmo Hardy — Evenhuis, N.L. & Thompson, F.C. ..... 179 vi BISHOP MUSEUM BULLETIN IN ENTOMOLOGY 12 (2004) PREFACE D. Elmo Hardy, “one of the last polymaths of Diptera Systematics” (see Evenhuis’ “Biography of D. Elmo Hardy”, this volume), died on 17 October 2002. Elmo, as he was known to most who knew him personally, was not only a dipterist extraordinaire, he was a teacher and mentor to students of entomology throughout the Asia-Pacific Region. Over the nearly seven decades of active research, Elmo named and described nearly 2,000 species in 34 different families of Diptera. These were pub- lished in 437 articles and notes (see Evenhuis & Thompson, this volume). Elmo may be considered the “father of Hawaiian Dipterology” and has inspired the research of many students of Diptera sys- tematics. The collection of articles in this volume is a celebration of Elmo’s achievements in the tax- onomy and systematics of flies from the Asia-Pacific Region. Evenhuis’ lead article on the Biography of D. Elmo Hardy is a narrative of Elmo’s personal life as well as his scientific career. The primary reference for the article is an unpublished autobiogra- phy and is full of personal stories about his childhood years, his many travels to exotic parts of the world, and insights into his formative years that led to his interests in insect systematics. The article by Skevington and DeMeyer describes Elmo’s contributions to the systematics of Pipunculidae, or big-headed flies. Bickel, in his article, describes a new genus of Dolichopodidae and names the type species for this genus, Alishania elmohardyi, in recognition of Elmo’s work on the taxonomy of this family of flies. This is followed by Evenhuis’ article describing another new genus, Humongochela, in the family Dolichopodidae. The Kaneshiro paper is a review of his earlier work on the behavioral genetics of the Hawaiian Drosophilidae in understanding the role of sexual selection in the formation of new species in this group. Uechi et al. presents a description of host alternation in species of gall midges in the genus Asphondylia. They present evidence from DNA analyses that A. gennadii can complete its life cycle by utilizing at least four different host plants alternatively. Norrbom and Hancock describe three new species and six new records of Tephritidae increas- ing the total number of species from New Caledonia from 16 to 25. Fitzgerald discusses five species of Bibionidae from New Caledonia, four of which are described as new species. Mathis & Zatwar- nicki present a revision of five species in the shore-fly genus Trimerogastra of which two are described as new species. Carson’s paper is a review of his earlier studies on the use of inversion patterns in the giant polytene chromosomes as “tracers” of phylogenetic relationships among 107 species of picture- winged Hawaiian drosophilids. He also discusses the role of sexual selection in species formation. The O’Grady & Silvestri paper presents some new DNA data and, together with morphologi- cal and behavioral data, the authors infer phylogenetic relationships among the fungus-feeder group of species in the endemic Hawaiian drosophilids. The authors provide evidence that some of the species group relationships based on morphological data alone need to be revised when the molec- ular data are incorporated. Copeland et al. report on insect host associations of Oleaceae in Kenya. They sample four gen- era (13 species) of the olive family and reared five species of tephritids, including one new species, as well as four braconid parasitoids that were reared from the pupae of these tephritids. They also report on the lepidopteran species and their parasites that were reared from the fruit samples. Drew presents a discussion of the biogeography and speciation in the Dacini group of Tephrit- idae, primarily of species in the genera Bactrocera and Dacus. He suggests that the close relation- ship between these fruit fly species and its host plant where courtship and mating occurs has had a significant influence on the speciation process and the evolution of this group of flies. The final chapter is a complete list of Elmo’s publications and of dipteran taxa he described. These include 437 articles and notes where 1,867 species in 34 families of Diptera were treated. Evenhuis & Kaneshiro — D. Elmo Hardy Memorial Volume vii This volume would not have been possible without the willingness and enthusiasm of each of the contributors. We are also indebted to those who reviewed and critiqued the manuscripts; their critical comments and suggestions significantly enhanced the quality of this volume. N.L.E. K.Y.K. Honolulu, Hawai‘i January 2004 viii BISHOP MUSEUM BULLETIN IN ENTOMOLOGY 12 (2004) D. Elmo Hardy Memorial Volume. Contributions to the Systematics 1 and Evolution of Diptera. Edited by N.L. Evenhuis & K.Y. Kaneshiro. Bishop Museum Bulletin in Entomology 12: 1–11 (2004). Biography of D. Elmo Hardy (1914-2002)1 NEAL L. EVENHUIS J. Linsley Gressitt Center for Entomological Research, Bishop Museum, 1525 Bernice Street, Honolulu, Hawai‘i 96817-2704, USA; email: [email protected] Introduction D. Elmo Hardy, “Elmo” to all who knew him personally, was one of the last polymaths in Diptera systematics. His knowledge of Diptera families spanned the order from Nematocera (specializing in Bibionidae), through to the Brachycera (specializing in the families Pipunculidae, Drosophilidae, and Tephritidae). He expanded his knowledge of the systematics to many other Diptera families through his monumental efforts in producing 5 volumes of Diptera for the “Insects of Hawaii” series where he described 581 species in no less than 20 families in the 30 years it took to conduct that research and produce those volumes. He coordinated the cataloging of the Diptera of the Oriental Region in the 1970s, the first and only comprehensive catalog from that region. He helped start the Hawaiian Drosophila project in 1963 which, with the help of numerous collaborators, increased the systematic and genetic knowledge of and stimulated