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THE INSECTS OF HAWAII The Insects of Hawaii Series was conceived as a comprehensive taxonomic treatment of the Hawaiian insect fauna. With more than 5,000 native arthropod species already described and many more awaiting discovery, this vast undertaking requires the collaborative efforts of systematists spe cializing in all components of the class. We have created an editorial board that reflects this diver sity ofexpertise. Board members will develop and recruit new projects for the series and will review the scientific content of manuscripts submitted. In view of the efforts required to reach the goal of completely describing Hawaii's entomofauna, the.board invites inquiries from specialists interested in preparing monographs for groups in need of revision. Editor: James K. Liebherr Department of Entomology Comstock Hall Cornell University Ithaca, NY 14853-0901 Taxonomic Editors: Rosemary G. Gillespie-University of California, Berkeley (Arachnida) Dan A. Polhemus-Smithsonian Institution, Washington, D.C. (Hemiptera) Manfred Asche-Humboldt Museum fur Naturkunde, Berlin (Homoptera) James K. Liebherr-Cornell University, Ithaca (Coleoptera) Scott E. Miller-Smithsonian Institution, Washington, D.C. (Lepidoptera) Neal L. Evenhuis-Bernice P. Bishop Museum, Honolulu (Diptera) . David B. Wahl-American Entomological Institute, Gainesville (Hymenoptera) Instructions to Authors Beginning with the publication of Volume 16, the Insects of Hawaii Series is being produced as a camera-ready publication. Authors are responsible for providing final copy to the publisher, following a style and format consistent with Volumes 16 and 17. Prospective authorsshould re quest detailed information on the preparation of camera-ready copy from the editor. For review purposes, contributions should be submitted to the editor in standard double-spaced manuscript format. While there are no page charges for publications in this series, a modest subsidy will be required of each author. The amount of this subsidy will be determined by the size and complexity of the manuscript. Front cover: Hylaeus mana Magnacca and Daly, n. sp. Back cover: Hylaeus difficilis (Perkins) on Dodonaea INSECTS OF HAWAII INSECTS OF HAWAII A Manual of the Insects of the Hawaiian Islands, including an Enumeration of the Species and Notes on Their Origin, Distribution, Hosts, Parasites, etc. VOLUME 17 HAWAIIAN HYLAEUS (NESOPROSOPIS) BEES (HYMENOPTERA: APOIDEA) by HOWELL V. DALY Professor Emeritus of Entomology University of California Berkeley, CA and KARL N. MAGNACCA Department of Entomology Cornell University Ithaca, NY University of Hawai‘i Press Honolulu © 2003 University of Hawai'i Press All rights reserved Printed in the United States ofAmerica 03 04 05 06 07 08 5 4 3 2 1 library of Congress Cataloging-in-Pub1ication Data (Revised for vol. 17) Insects of Hawaii; a manual of the insects of the Hawaiian Islands. Vols. 1-9 by Elwood C. Zimmerman. Vols. 10-14 by D. Elmo Hardy and others. Vol. 15 by Kenneth Christiansen and Peter Bellinger. Vol. 16 by James K. Liebherr and Elwood C. Zimmerman. Includes bibliographies and indexes. Vol. 17 by Howell V. Daly and Karl N. Magnacca 1. Insects-Hawaii. I. Daly, Howell V. II. Magnacca, Karl N. QL489.H3Z5 595.709969 48-45482 ISBN 0-8248-2674-4 (v. 17) Camera-ready text prepared by the authors. University of Hawai'i Press books are printed on acid-free paper and meet the guidelines for permanence and durability of the Council on Library Resources. Printed by The Maple-Vail Book Manufacturing Group CONTENTS ABSTRACT ................................................................................................................... 1 INTRODUCTION ........................................................................................................ 3 THE HAWAIIAN ISLANDS ............................................................................... 3 HISTORY OF COLLECTIONS AND TAXONOMY ........................................ 3 DISPERSAL AND EVOLUTION OF HYLAEUS SPECIES IN HAWAII ........ 7 BIOLOGY OF HYLAEUS ................................................................................... 9 NATIVE AND ADVENTIVE ENEMIES .......................................................... 9 ECOLOGY OF HYLAEUS IN HAWAII ........................................................... 11 NATIVE AND ADVENTIVE BEES OF THE HAWAIIAN ISLANDS .......... 12 DESCRIPTIVE TERMINOLOGY .......................................................................... 14 KEYS TO SPECIES................................................................................................... 24 KEY TO MALES.............................................................................................. 25 KEY TO FEMALES ......................................................................................... 34 TAXONOMY OF HYLAEUS (NESOPROSOPIS) SPECIES ................................ 43 NOMINA DUBIA ..................................................................................................... 209 ACKNOWLEDGMENTS ....................................................................................... 210 BIBLIOGRAPHY .................................................................................................... 211 APPENDIX: RECORDS OF RECENT COLLECTIONS................................... 217 INDEX ....................................................................................................................... 230 HYLAEUS SPECIES-GROUP NAMES ......................................................... 230 ARTHROPOD ENEMIES AND ASSOCIATES ............................................ 231 PLANTS USED AS NESTING SUBSTRATES ............................................ 231 FLOWER VISITATION RECORDS .............................................................. 231 HAWAIIAN HYLAEUS 1 ABSTRACT The native colletid bee species of the genus Hylaeus (Nesoprosopis) are revised for the Hawaiian Islands based on adult morphology. Sixty species are recognized. NEW SPECIES described by Magnacca and Daly are H. akoko and H. gliddenae from Hawaii; H. kukui and H. takumiae from Maui; H. kuakea, H. mana, H. mimicus, and H. nalo from Oahu; and H. kokeensis and H. solaris from Kauai. NEW SYNONYMIES are proposed (senior synonyms first): H. assimulans (Perkins) = H. assimulans oahuensis (Perkins); H. coniceps (Blackburn) = H. comes (Perkins); H. connectens (Perkins) = H. koae (Perkins), H. insulae Meade-Waldo (earlier replacement name for Nesoprosopis vicina Perkins), and Prosopis trigona Warncke (unwarranted replacement name for Nesoprosopis vicina Perkins); H. flavipes (F. Smith) = H. blackburni (F. Smith); H. fuscipennis (F. Smith) = H. fuscipennis obscuripes (Perkins), H. fuscipennis swezeyi (Cockerell), and H. caeruleipennis (Perkins); H. haleakalae (Perkins) = H. neglectus (Perkins); H. nivicola (Meade-Waldo) = Prosopis farinosa Warncke (unwarranted replacement name for Nesoprosopis nivalis Perkins); H. paradoxicus (Perkins) = H. erythrodemas (Perkins); H. rugulosus (Perkins) = Prosopis ovatula Warncke (unwarranted replacement name for Nesoprosopis rugulosa Perkins); H. specularis (Perkins) = H. homoeochromus (Perkins); H. unicus (Perkins) = H. binominatus Meade-Waldo (earlier replacement name for Nesoprosopis laticeps Perkins) and Prosopis avara Warncke (unwarranted replacement name for Nesoprosopis laticeps Perkins). NEW COMBINATIONS are Hylaeus muranus (Warncke) and Hylaeus niloticus (Warncke). The following names are considered nomina dubia: “Nesoprosopis rubrocaudatus Blackburn and Cameron” and “Nesoprosopis attripennis Perkins.” The history of collections of the bees, their taxonomy, attributes for dispersal, and their biology and ecology are discussed. Taxa of adventive bees are characterized as an aid to identification of native species. Terminology used in the treatment is explained, followed by keys to the identification of species based on adult males and females. Treatment of each species includes annotated synonymy and other references; diagnosis of identifying features and general distribution; description of male and female; localities where first collected and recent collections; flower records; remarks on taxonomic problems and other information; and line illustrations. Recent collection records are provided in an appendix. 2 INSECTS OF HAWAII VOL. 17 Fig. 1. Map of major islands in the Hawaiian Island chain (after Zimmerman 1948:10). HAWAIIAN HYLAEUS 3 INTRODUCTION The native bees of the Hawaiian Islands are believed to have evolved from a single lineage of the genus Hylaeus (family Colletidae) and are assigned to the subgenus Nesoprosopis. The genus Hylaeus is worldwide in distribution with 46 subgenera and an estimated 579 species (Michener 2000:62). The subgenus Nesoprosopis is among the largest of the subgenera with 60 of its species found in the Hawaiian Islands and eight species in the Oriental region. The latter includes one species also known from the Palearctic region (Hirashima 1977, Ikudome 1989). About three-quarters of the species described from the Hawaiian Islands by Perkins (1899, 1910, 1911) have been collected recently (Table 1). Some are still as rare or as abundant as he observed, yet others, formerly abundant, have not been collected recently. After more than a century, we have little knowledge of their mating and nesting habits, food preferences, natural enemies, or roles as pollinators of endemic Hawaiian plants. The purpose of this volume is to encourage future research by facilitating identification and by documenting the survival of these interesting